Wildlife-Habitat Relationships

in Oregon and Washington

 Project Sponsors and Contributing Sponsors
Birds of Oregon Project
Birds of Washington Project
Confederated Tribes of the Warm Springs
Donavin A. Leckenby
Environmental Protection Agency
Federal Highways Administration
Fish and Wildlife Information Exchange
National Fish and Wildlife Foundation
Northwest Power Planning Council
Oregon Chapter of The Wildlife Society
Oregon Cooperative Wildlife Research Unit
Oregon Department of Fish and Wildlife
Oregon Department of Transportation
Oregon Forest Resources Institute
Oregon/Washington Partners In Flight
Pacific States Marine Fish Commission
Quileute Indian Tribe
Rocky Mountain Elk Foundation
Paul F. and Teresa J. Roline
Society for Northwestern Vertebrate Biology
USDA Forest Service
USDI Bureau of Land Management
USDI Fish and Wildlife Service
USGS Biological Resources Division
Washington Chapter of The Wildlife Society
Washington Community, Trade, and Economic Development
Washington Cooperative Fish and Wildlife Research Unit
Washington Department of Natural Resources
Washington Department of Transportation
Washington Forest Protection Association
Weyerhaeuser Company
Wildlife Management Institute

The Washington Department of Fish and Wildlife and the Northwest Habitat Institute
were the lead organizations on this project.

Managing Directors’ Dedication

The ecological face of the Pacific Northwest has been, and continues to be, significantly altered by our
human hands. In the last 200 years, there have been many changes to the wildlife and human populations
that share this most wonderful place on earth. In recognition that to reverse these trends will require
human intervention, we dedicate this book to you, and to those with a profound respect of nature who
have come before us. It will be through your efforts and dedication to restoring and conserving the health
of our natural resources that we may thrive as a people in unison with our environment.
Wildlife-Habitat Relationships
in Oregon and Washington

Managing Directors

David H. Johnson

Thomas A. O’Neil

Oregon State University Press

Corvallis
The paper in this book meets the guidelines for permanence and durability of the
Committee on Production Guidelines for Book Longevity of the Council on Library
Resources and the minimum requirements of the American National Standard for
Permanence of Paper for Printed Library Materials Z39.48-1984.

Library of Congress Cataloging-in-Publication Data

Wildlife-habitat relationships in Oregon and Washington / managing directors,
David H. Johnson, Thomas A. O’Neil.— 1st ed.
p. cm.
Includes bibliographical references and index.
ISBN 0-87071-488-0 (alk. paper)
1. Animal ecology—Oregon. 2. Animal ecology—Washington (State). 3. Habitat
(Ecology)—Oregon. 4. Habitat (Ecology)—Washington (State). I. Johnson, David H.
II. O’Neil, Thomas A.
QH105.O7 W565 2000
591.7’09795—dc21
00-010729

© 2001 Oregon State University Press

All rights reserved. First edition 2001
Printed in the United States of America

Oregon State University Press

101 Waldo Hall
Corvallis OR 97331-6407
541-737-3166 • fax 541-737-3170
http://osu.orst.edu/dept/press
 Table of Contents
Maps of Oregon and Washington Wildlife-Habitat Types ............................................................................ v i i
Managing Directors’ Preface .......................................................................................................................... xii
Foreword ........................................................................................................................................................ xiv
Introduction .................................................................................................................................................... x v
1 Oregon and Washington Wildlife Species and Their Habitats ................................................................ 1
T.A. O’Neil and D.H. Johnson
2 Wildlife Habitats: Descriptions, Status, Trends, and System Dynamics ............................................... 22
C.B. Chappell, R.C. Crawford, C. Barrett, J. Kagan, D.H. Johnson, M. O’Mealy, G.A. Green,
H.L. Ferguson, W.D. Edge, E.L. Greda, and T.A. O’Neil
3 Structural Conditions and Habitat Elements of Oregon and Washington .......................................... 115
T.A. O’Neil, K.A. Bettinger, M. Vander Heyden, B.G. Marcot, C. Barrett, T.K. Mellen,
W. M. Vander Haegen, D.H. Johnson, P.J. Doran, L. Wunder, and K.M. Boula
4 Management of Within-stand Forest Habitat Features ........................................................................ 140
W.C. McComb
5 Conservation of Biodiversity: Considerations and Methods for Identifying and Prioritizing Areas
and Habitats ......................................................................................................................................... 154
M.M. Shaughnessy and T.A. O’Neil
6 Key Ecological Functions of Wildlife Species ..................................................................................... 168
B.G. Marcot and M. Vander Heyden
7 Wildlife of Westside and High Montane Forests ................................................................................. 187
D.H. Olson, J.C. Hagar, A.B. Carey, J.H. Cissel, and F.J. Swanson
8 Wildlife of Eastside (Interior) Forests and Woodlands ........................................................................ 213
R. Sallabanks, B.G. Marcot, R.A. Riggs, C.A. Mehl, and E.B. Arnett
9 Wildlife of Alpine and Subalpine Habitats ......................................................................................... 239
K.M. Martin
10 Wildlife of Westside Grassland and Chaparral Habitats ..................................................................... 261
B. Altman, M. Hayes, S. Janes, and R. Forbes
11 Wildlife of Eastside Shrubland and Grassland Habitats ..................................................................... 292
W.M. Vander Haegen, S.M. McCorquodale, C.R. Peterson, G.A. Green, and E. Yensen
12 Wildlife of Urban Habitats .................................................................................................................. 317
H.L. Ferguson, K. Robinette, and K. Stenberg
13 Wildlife of Agriculture, Pastures, and Mixed Environs ....................................................................... 342
W.D. Edge
14 Wildlife of Riparian Habitats .............................................................................................................. 361
J.B. Kauffman, M. Mahrt, L. Mahrt, and W.D. Edge
15 Wildlife of Coastal and Marine Habitats ............................................................................................. 389
J.B. Buchanan, D.H. Johnson, E.L. Greda, G.A. Green, T.R. Wahl, and S.J. Jeffries
16 Introduced Wildlife of Oregon and Washington ................................................................................. 423
G.W. Witmer and J.C. Lewis
17 Genetic Considerations for Introduced and Augmented Populations ................................................ 444
S.M. Haig and R.S. Wagner
18 Extirpated Species of Oregon and Washington ................................................................................... 452
C. Iten, T.A. O’Neil, K.A. Bettinger, and D.H. Johnson
19 Characterizing Species at Risk ............................................................................................................ 474
J.F. Lehmkuhl, B.G. Marcot, and T. Quinn
20 Terrestrial and Marine Management Activities: Links to Habitat Elements and Ecological Processes ....
............................................................................................................................................................. 501
M. Vander Heyden and B.G. Marcot
21 An Overview of Models and Their Role in Wildlife Management ..................................................... 512
G.J. Roloff, G.F. Wilhere, T. Quinn, and S. Kohlmann
22 Five Case Studies of Wildlife Modeling Applications ........................................................................ 537
T. Quinn and D.H. Johnson, editors
Case Study 1: A Spatially Realistic Population Model for Informing Forest Management Decisions .....
............................................................................................................................................................. 538
G.F. Wilhere, N.H. Schumaker, and S.P. Horton
Case Study 2: A Model to Assess Potential Vertebrate Habitat at Landscape Scales: HABSCAPES ... 544
M. Huff, T.K. Mellen, and R. Hagestedt
Case Study 3: Cross-scale Classification Trees for Assessing Risks of Forest Practices to Headwater
Stream Amphibians ............................................................................................................................. 550
G.D. Sutherland and F.L. Bunnell
Case Study 4: Applying GAP Analysis to County and Regional Land Use Planning ......................... 555
M.R. Stevenson
Case Study 5: A Model to Determine Potential Northern Spotted Owl Nesting Areas ....................... 561
N.W. Darby and T. Young
23 Integrating Wildlife Species Habitat Goals and Quantitative Land Management Planning Processes ...
............................................................................................................................................................. 567
P. Bettinger, K. Boston, J. Sessions, and W.C. McComb
24 Decaying Wood in Pacific Northwest Forests: Concepts and Tools for Habitat Management ............ 580
C.L. Rose, B.G. Marcot, T.K. Mellen, J.L. Ohmann, K.L. Waddell, D.L. Lindley, and B. Schreiber
25 Single Species, Multiple Species, or Ecosystem Management: A Perspective on Approaches to
Wildlife Conservation .......................................................................................................................... 624
H. Salwasser
26 Pacific Salmon and Wildlife: Ecological Contexts, Relationships, and Implications for Management ...
............................................................................................................................................................. 628
C.J. Cederholm, D.H. Johnson, R.E. Bilby, L.G. Dominguez, A.M. Garrett, W.H. Graeber, E.L. Greda,
M.D. Kunze, B.G. Marcot, J.F. Palmisano, R.W. Plotnikoff, W.G. Pearcy, C.A. Simenstad, and P.C. Trotter
27 An Introduction to Wildlife-Habitat Relationships CD-ROM ............................................................ 686
M. Trevithick , T.A. O’Neil, and C. Barrett
Appendix: Scientific and common names for 743 wildlife species found in Oregon and
Washington .......................................................................................................................................... 687
Glossary of Terms ................................................................................................................................ 712
Acknowledgments ............................................................................................................................... 723
Index .................................................................................................................................................... 728

CD-ROM: Matrixes for Wildlife–Habitat Relationships in Oregon and Washington

T.A. O’Neil, D.H. Johnson, C. Barrett, M. Trevithick, K.A. Bettinger, C. Kiilsgaard,
M. Vander Heyden, E.L. Greda, D. Stinson, B.G. Marcot, P.J. Doran, S. Tank, and L. Wunder
WILDLIFE-HABITAT MAPS vii
viii Wildlife–Habitat Relationships in Oregon and Washington
WILDLIFE-HABITAT MAPS ix
x Wildlife–Habitat Relationships in Oregon and Washington
 WILDLIFE-HABITAT MAPS xi

Oregon and Washington Wildlife-Habitat Types

Forest & Woodland Habitats Developed Habitats

Westside Lowland Conifer-Hardwood Forest Agriculture, Pasture & Mixed Environs
Westside Oak & Dry Douglas-fir Forest & Woodlands Urban & Mixed Environs
Southwest Oregon Mixed Conifer-Hardwood Forest
Aquatic & Riparian Habitats
Montane Mixed Conifer Forest
Lakes, Rivers, Ponds & Reservoirs
Eastside (Interior) Mixed Conifer Forest Herbaceous Wetlands
Western Juniper & Mountain Mahogany Woodlands Westside Riparian - Wetlands
Lodgepole Pine Forest & Woodlands Montane Coniferous Wetlands
Ponderosa Pine & Eastside White Oak Forest & Woodlands Eastside (Interior) Riparian - Wetlands
Upland Aspen Forest
Subalpine Parklands Maritime & Coastal Habitats
Coastal Dunes & Beaches
Grassland & Shrubland Habitats Coastal Headlands & Islets
Alpine Grasslands & Shrublands Bays & Estuaries
Westside Grasslands Inland Marine Deeper Waters
Ceanothus-Manzanita Shrublands Marine Nearshore
Eastside (Interior) Canyon Shrublands Marine Shelf
Eastside (Interior) Grasslands Oceanic
Shrub-steppe
Dwarf Shrub-steppe
Desert Playa & Salt Scrub

Oregon: Scale 1:3,500,000

Washington: Scale 1: 3,000,000
Current: circa 1999
Historic: circa 1850 (modeled)
Map compilation and cartography by Chris Kilsgaard and Charley Barrett, Northwest Habitat Institute
Marine contours provided by Terence P. Johnson, Washington Dept. of Fish and Wildlife
Current Eastern Washington Shrub-Steppe data provided by John E. Jacobson, Washington Dept. of Fish and Wildlife
 xii Wildlife–Habitat Relationships in Oregon and Washington

Managing Directors’ Preface

Wildlife–Habitat Relationships in Oregon and Washington and Because so many people and organizations have a stake
accompanying maps and digital information have been in the wildlife heritage of the Pacific Northwest, more than
developed to synthesize and disseminate the current state 600 people have been involved in the creation of this
of knowledge about amphibians, birds, mammals, and document and associated products. Thirty-three Project
reptiles, and their terrestrial, freshwater, and marine Partners and Contributing Sponsors have supported the
habitats of Oregon and Washington. Throughout this project financially or by detailing staff, facilities, and/or
book, we have focused our attention on scientific rigor equipment. The final tally shows that we received input
and an honest evaluation of what we know, and do not from 246 questionnaire respondents, 225 people who
know, about these wildlife species and their habitat attended the landscape modeling workshop in Olympia,
relationships. Under a multiple partnership framework, 73 species specialists who participated in the
we are broadening our understanding of wildlife habitats. fifteenscientific panels, 40 people who accessed and
Our wildlife habitats are made up of three components: contributed to our Internet site, 88 chapter authors, and
wildlife-habitat types, structural conditions, and habitat another 30 people who guided our process by
elements. We have also added exciting new information participating on one of five advisory teams. Please see the
about species’ key ecological functions, management full acknowledgments at the end of the book for details
activity links (with habitat elements), and salmon-wildlife about the many organizations and individuals without
relationships. Throughout this book and the digital whose contribution this project would not have been
information, we have done our best to integrate disciplines completed.
often considered as independent: terrestrial, freshwater, Our work was done in a spirit of cooperation and
and marine systems. We have also designed the collaboration, and relationships were forged that we hope
information within this book to be applicable at the local, will continue into the future as we continue to learn from
watershed, state, and regional levels. Further, we have each other how best to meet our common goals. Our
worked hard to develop the information, and overall travels around Oregon and Washington offered us unique
framework, for easy application to surrounding states and insights into our truly remarkable natural resources and
provinces, the natural extension of our ecological context. people that occupy the region. Our philosophy that lead
We felt this overall approach was fundamentally us to undertake the development of this book is best
important, as we strove to establish a cohesive information reflected within David Suzuki’s Declaration of
source on which to base a common understanding for Interdependence (below). To the many, many people we
management to address ecologically and economically have worked with during the last four years, we thank
important issues. Based on previous works, and the you. It has been an honor and true pleasure to work with
combined strength of the contributions to this book, we so many talented and caring individuals.
anticipate that the information presented here will be used October 12, 2000
in natural resource planning, conservation, and education David H. Johnson and Thomas A. O’Neil
efforts for years to come.
 Declaration of Interdependence
This We Know
We are the earth, through the plants and animals that nourish us.
We are the rains and the oceans that flow through our veins.
We are the breath of the forests of the land, and the plants of the sea.
We are human animals, related to all other life as descendants of the firstborn cell.
We share with these kin a common history, written in our genes.
We share a common present, filled with uncertainty.
And we share a common future, as yet untold.
We humans are but one of thirty million species
weaving the thin layer of life enveloping the world.
The stability of communities of living things depends upon this diversity.
Linked in that web, we are interconnected -
using, cleansing, sharing and replenishing the fundamental elements of life.
Our home, planet Earth, is finite, all life shares its resources and the energy from the sun,
and therefore has limits to growth.
For the first time, we have touched those limits.
When we compromise the air, the water, the soil and the variety of life,
we steal from the endless future to serve the fleeting present.

This We Believe
Humans have become so numerous and our tools so powerful
that we have driven fellow creatures to extinction, dammed the great rivers,
torn down ancient forests, poisoned the earth, rain and wind, and ripped holes in the sky.
Our science has brought pain as well as joy; our comfort is paid for by the suffering of millions.
We are learning from our mistakes, we are mourning our vanished kin,
and we now build a new politics of hope.
We respect and uphold the absolute need for clean air, water and soil.
We see that economic activities that benefit the few while shrinking the inheritance of many are wrong.
And since environmental degradation erodes biological capital forever,
full ecological and social cost must enter all equations of development.
We are one brief generation in the long march of time; the future is not ours to erase.
So where knowledge is limited, we will remember all those who will walk after us,
and err on the side of caution.

This We Resolve
All this that we know and believe must now become the foundation of the way we live.
At this turning point in our relationship with Earth,
we work for an evolution: from dominance to partnership;
from fragmentation to connection; from insecurity, to interdependence.
Declaration of Interdependence published with permission from the David Suzuki Foundation,
219-2211 West 4th Ave, Vancouver, British Columbia, Canada. V6K 4S2.

xiii
 Foreword

The biggest challenge facing natural resource conservation The information reported here is the first
efforts in the next millennium is the maintenance of viable comprehensive effort that we are aware of to compile and
ecosystems and the biological diversity they encompass. synthesize the body of knowledge about Oregon and
Addressing this imbroglio will require us to find solutions Washington wildlife. Concurrent with the advent of
that are efficient yet effective. In meeting this challenge, ecosystem-based management is the need to address
we need to continue to strive and build a common multi-species requirements while addressing single-
foundation of knowledge that will serve to support sound species issues, and to see the land, its habitats and wildlife
and wise decisions concerning our natural resources. This communities as a system. This book is a response to that
book is an attempt to synthesize a substantially large and need and updates our understanding of Oregon’s and
diverse body of wildlife information into an agreed Washington’s forests, shrublands, grasslands, marine and
foundation of knowledge. It began as a grassroots effort aquatic habitats, and describes the wildlife relationships
of the Managing Directors; with a little bit of persuasion, that are believed to exist. To achieve this we have
we were successful in initially convincing the Washington combined the information found in the literature with the
and Oregon Departments of Fish and Wildlife to offer knowledge of individuals expert in wildlife, botany,
support for the project. We realized from the beginning fisheries, conservation biology, the fields of forest,
that simply having these agencies’ endorsement and initial rangeland, and marine ecology, earth-bound and satellite
support was insufficient; if we were to be successful we vegetation mapping technologies, and computer science.
would require participation from numerous professionals The presentation of this information is based on a
and support from a variety of groups and agencies. foundation of sound theoretical and scientific perspectives,
Eventually, 34 entities became project partners or and recognizes that the final products must have practical
contributing sponsors, and provided financial or in-kind applications. After all, the work developed here should
support to the project. As we approached the conclusion allow for new insights, suggest new hypotheses, and offer
of this effort, we found ourselves almost completely mechanisms to address single and multi-species needs in
relying on our project partners’ support. After four years a world that is becoming less and less wild with each
of focused effort, numerous consensus decision-making passing day. Our hope for the future is that people from
work sessions, and involvement of more than 600 people, all walks of life can and will work together to keep those
we present our best effort here in Wildlife—Habitat natural resources we cherish healthy throughout the next
Relationships in Oregon and Washington in hopes of building millennium. For those people who come after us and are
a common understanding for management. willing to continue updating and expanding these
concepts, we hope that you will experience the comradery,
laughter, and passion for the resources that we
experienced; may nature’s forces be with you!

xiv
 Introduction

Effectively managing our natural resources requires the most of it is about fifteen to twenty years old and in need
combination of high-quality science and the availability of updating. Also, there were geographic gaps in the
of data-rich information. A key information need for information, as well as important differences in how the
natural resource managers, scientists, and educators is the habitats were defined and described.
current state of knowledge of wildlife species and their This publication builds on the work of past regional
habitats. In this book, we use the term “wildlife” to refer publications and subscribes to (1) taking a two-state
to terrestrial vertebrates or amphibians, birds, mammals, perspective and making seamless the terrestrial,
marine mammals, and reptiles. While we support the more freshwater, and marine environs of Oregon and
encompassing description of wildlife to include fishes and Washington; (2) providing updated information on
invertebrates, by necessity we had to narrow the focus in wildlife species and their habitat relationships; (3)
this book. reviewing basic concepts and current thinking regarding
Wildlife habitat is a concept related to a particular these relationships; and 4) providing data and approaches
wildlife species.9 More specifically, habitat is an area with that could be used in local, regional, and state planning.
the combination of the necessary resources (e.g., food, Importantly, we have focused a good deal of attention on
cover, water) and environmental conditions (temperature, the integration of terrestrial, freshwater, and marine
precipitation, presence or absence of predators and systems. Also, this project advances the following new
competitors) that promotes occupancy by individuals of concepts: defining the hierarchical nature of wildlife
a given species (or population), and allows those habitats (reflected in our wildlife habitat types, structural
individuals to survive and reproduce.5 The arrangement conditions, and habitat elements) and key ecological
of these habitat resources and features to meet the functions; tying management activities to habitat elements;
biological needs of a species identifies the habitat niche a characterizing salmon—wildlife relationships; assigning
species occupies, and from a systems perspective, provides associations and confidence levels to each species’
a framework for the ecological role or function that an relationship to wildlife habitat types and structural
individual species plays within the environment.6 From a conditions; and developing the first wildlife habitat maps
manager’s perspective, the habitat concept needs to be for each state. The geographic area covered by these
tangible and scientifically supported, inasmuch as habitat systems is substantial: the terrestrial and freshwater areas
features need to be plainly defined, and the wildlife species encompass 75 counties with an area of 105,710,720 acres
associations with them made clear. Addressing this (42,781,128 ha); the surface area of the marine waters
management perspective is the primary focus of this book. include an additional 65,747,200 acres (26,298,880 ha).
Understanding species and their habitat relationships A fundamental paradigm shift has occurred since the
is paramount to predicting species’ responses to past, initial habitat relationship books were published. The shift
present, and future land uses within a managed landscape. reflects the move away from a focus on wildlife habitats
In the past, one approach to habitat management could having value for few individual species, like elk, towards
be considered similar to the motto in the movie Field of wildlife habitats having multiple values for the mix of
Dreams: “Build or protect it (habitat) and they (species) species they may contain. Importantly, the shift
will come.” But in order to build it or conserve it, resource incorporates a view of the land and its value in a systems
managers need to know what the relationships are context, giving rise to an ecosystem-based management
between the individual species and their habitat. philosophy. In 1940, Aldo Leopold3 proposed a “land
Furthermore, because of disturbances, habitats are in a ethic” that put natural resource decisions on a non-special-
constant state of change, and thus wildlife communities interest footing. Leopold stated that a natural resource
constantly change in response. Habitats need to be decision is “right when it tends to preserve the integrity,
evaluated throughout their entire geographic range where stability and beauty of the biotic community,” and “it is
a species occurs to assess the influence of these habitats wrong when it tends otherwise.” Since 1960, an ethic for
on the life history characteristics (such as breeding, our natural resources has been steadily building, and it is
feeding, and wintering) of that wildlife species. influencing our thinking and policies by giving greater
Unfortunately, most wildlife relationship information considerations to species other than our own. This ethical
has generally not been easily accessible. Information perspective is aimed at becoming more holistic with one’s
germane to this topic is often found in largely unread surroundings and recognizes that humans are part of a
journals and transactions of symposia, in researchers’ files, larger system. These broader views are shaping the
and in the minds of knowledgeable individuals.10 Recent principles that underlie the way we think about and value
technological advances in data retrieval (e.g., bibliographic the environment as a whole, as well as how we approach
searches) have been beneficial, but users are still left with natural resource problems.
the daunting task of synthesis. Earlier publications have Over the past forty years, a number of laws have played
forged the basic wildlife relationship information available a part in shaping our landscapes and marine waters.
for Oregon and Washington,1,2,4,7,9 and while important, Among the most influential federal laws are the Mining

xv
 xvi Wildlife–Habitat Relationships in Oregon and Washington

Law of 1872, Multiple Use-Sustained Yield Act of 1960, the segment of society who were dependent on natural
Wilderness Act of 1964, Wild and Scenic Rivers Act of 1968, resources in the Northwest, became the subject of
Clean Air Act of 1970, Clean Water Act of 1972, Marine President Clinton’s Forest Summit hosted in Portland,
Mammal Protection Act (1972), Endangered Species Act Oregon. Hence new ideas and approaches influencing our
of 1973 (ESA), Forest and Rangeland Renewable Resources landscapes were required. Likewise, technological changes
Planning Act of 1974, National Forest Management Act of allowed us to view, and analyze, our landscapes and
1976, and the Federal Land Policy and Management Act marine areas through an increasingly rich array of tools.
of 1976. Likewise, a number of state laws have shaped The growing body of science and technology played a key
our landscape as well. Oregon’s Land Use Planning Act role in shaping the information on which policy decisions
(Oregon Revised Statute #197.005 to .860) requires counties were based.
to prepare comprehensive land-use plans that include More recently, our increasing knowledge about the
consideration of wildlife habitat, open space needs, and environment, and about species such as the northern
the consideration of ecologically significant natural areas. spotted owl and salmon, has led to a growing awareness
The Oregon Endangered Species law has provisions that that human society needs to adapt its activities to protect
protect native vertebrates and plants on state lands only crucial ecological processes. To do so requires an
(Oregon Revised Statute #496.172 to .192; 498.026; 564.100 integration of ecological, economic, and social values to
to .135), and the Oregon Forest Practices Act requires manage biological and physical systems in a way that
consideration of the impacts of forest practices on safeguards their long-term sustainability, natural diversity,
threatened, endangered, and special concern species and productivity. Thus, ecosystem-based management is
(Oregon Revised Statute #527.610). In Washington, there a philosophy that focuses on desired states to provide the
is also an Endangered Species law that covers animals, outputs sought within a framework of sustainable and
but not plants (Revised Code of Washington #77.16.040, viable ecological conditions, rather than management
77.16.120, 77.12.020, 77.08.010, 77.12.055 to 3, 77.21.010, primarily for system outputs. It acknowledges the need
79.08.250). The Washington Growth Management Act to protect or restore critical ecosystem components,
affects land-use activities in counties with populations of functions, and structures in order to sustain ecological
50,000 or more and requires the designation of critical areas systems in perpetuity. A shortcoming of this effort is that
of habitat and open space corridors (Revised Code of we are unable to address the ecosystem processes as a
Washington #36.70A.010 et seq.). Washington also has a whole, and for a good reason: we do not fully understand
Forest Protection Act (Revised Code of Washington #76.09, them. This project only addresses the portion of the
enacted in 1974), which establishes minimum standards ecosystem that is associated with terrestrial, freshwater,
for forest practices, and provides protection for critical and marine vertebrates; users of our information seeking
wildlife habitats of threatened and endangered species. to implement an ecosystem-based management approach
The Washington Hydraulics Code (Revised Code of are urged to seek additional information on the status and
Washington #75.20 enacted in 1949) requires a permit for functions of other ecological components, such as fish and
any activity that will use, divert, obstruct, or change the invertebrates. These other species should not be
bed or flow of state waters. The Washington Shoreline overlooked within the landscapes and aquatic environs
Management Act (Revised Code of Washington #90.58, as they contribute to fundamentally important ecological
enacted in 1972) applies to all marine waters, submerged functions.
tidelands, lakes >20 acres (49.5 ha), and larger streams, Other approaches are also emerging to address an
marshes, bogs, and swamps and adjacent landward area; ecosystem understanding. Organizations are moving
with the intent to ensure that “. . . development of these away from traditional static approaches and towards
shorelines . . . will promote and enhance the public becoming learning institutions. Agencies that are striving
interest.” The Washington State Environmental Policy Act to become learning institutions focus on information
(Revised Code of Washington #43.21C, enacted in 1971), management, training and education, integration and
provides for the review of environmental impacts of cooperation, evaluation and feedback loops, efficient and
activities that may potentially affect Washington’s air, flexible processes, and empowerment of employees and
water, soil, human health, and environment. the public.8 An example of a learning approach is adopting
Whereas these laws and policies help to improve land adaptive management strategies into the management of
and marine conservation, prior to 1990 most natural natural resources. Adaptive management is an approach
resource management was driven by commodity that incorporates uncertainty into managing our natural
production, like the Allowable Sale Quantity (ASQ), the resources. It involves using a systematic process to
amount of board feet of timber from a National Forest; evaluate management actions that subsequently increase
Animal Units per Month (AUMs) for livestock on our level of understanding about our natural systems so
rangelands; and Maximum Sustained Yield (MSY) for that we can make better future decisions. Key to the
fisheries from our marine waters. But in the early 1990s success of an adaptive management strategy is monitoring
these attitudes and policy directions began to change not only the natural communities’ response to a
because several wildlife species became listed as management action, but also future management
threatened or endangered under the federal ESA. In April decisions, and adjusting them in accordance with results
1993, these changes, affecting the economic stability for obtained. Embracing adaptive management would forge
 INTRODUCTION xvii

a closer tie between researchers and managers, blur may agree with the information reported, but we have
boundaries between traditional funding sources, and more reviewed more than 100,000 pieces of literature, impaneled
effectively pool resources and give a higher return on our fifteen groups of expert specialists, cited our sources or
investment. gave comments to support a description or claim where
Managing our natural resources requires the appropriate, and performed quality assurance/quality
combination of science and the availability of information. control checks on the matrixes.
Therefore, a primary emphasis of our endeavor is to Finally, minor changes were made to the matrixes on
develop high-quality data sets on our wildlife habitats and the CD-ROM after the book chapters were finalized.
the species associated with them. We achieved this by first Hence, some query totals found in the chapters may
defining, describing, and depicting various component slightly deviate from the values reported on the CD-ROM.
details about our wildlife habitats that are offered in this Although we would have liked to include a chapter
book, maps, and accompanying CD-ROM. Secondly, we addressing the topic of “cumulative effects”, one is not
brought together as much scientific talent from the Pacific offered in this book. However, components of this book
Northwest as possible to review, determine, comment, and can be arranged into an approach to begin to address these
assign relationships between our wildlife species and their effects, and we propose an illustration (Figure 1) of one
habitats. Our approach moves away from defining what such way of using the matrixes in this book to this end. In
is primary or secondary habitat for a species, and towards the past, most managers have typically thought about
identifying the overall strength and context of the cumulative effects in one or two dimensions, that is, we
relationship between the wildlife species and their have reviewed management activities in terms of direct
habitat(s). As such, the strength of the relationship is impacts to species, groups of species or habitat(s). But here
designated as Closely Associated, Generally Associated, or is where we have stopped. To assess the health of
Present within the wildlife habitat types or structural ecosystems we need to incorporate a third dimension to
conditions. In addition, we also assign a confidence rating our thinking, which in the context of this book is framed
to the relationship and its strength based on what we know
today. This approach allows for an individual species, as
well as multiple species, to be assessed across habitats.
Using all matrixes will allow the reader to begin to acquire
an ecosystem perspective by depicting, for example,
species relationships to: one another, ecological functions,
unique and common life history characteristics (like diets
and reproductive sites), and potential effects from human
activities. These relationships, coupled with the confidence
rating for each association, should allow users to develop
potential species lists with a confidence ranking for each,
along with the ability to model uncertainty of wildlife use
of habitats by applying the confidence rankings as a fuzzy
set logic.11
As part of our updating wildlife-habitat relationships,
we offer seven data sets found on the CD-ROM for the
reader to use. These seven data sets are depicted as
matrixes; three matrixes detail the wildlife species’
associations with Wildlife Habitat Types, Structural
Conditions, and Habitat Elements. Other matrixes are
descriptive and illustrate Key Ecological Functions, Life
History characteristics, Management Activity Links (linkages
between management actions and the Habitat Elements)
and Salmon—Wildlife Relationships. Additionally, where the
literature was used to support life history or management
activity information, it can also be found under a separate
heading called Citations. The matrixes were developed in
a hierarchical manner so that the user can go from a coarse
level (wildlife habitat types) to site-specific, fine-scale
features (habitat elements). The descriptive matrixes are
developed to give the reader an idea of the role that the
species play within the ecosystem and what type of
activities may affect them. Each matrix is offered in a
Schematic of how the Wildlife-Habitat Relationships data
digital format and designed to give the user updated and
accurate information; there are some 60,000 records of data matrixes can be used with 3-dimensional thinking to
in the seven matrixes. We recognize that not all end users addresscumulative impacts.
 xviii Wildlife–Habitat Relationships in Oregon and Washington

by the examination of key ecological functions for each of chapters (7-15) discuss wildlife as part of multi-habitat
the wildlife species. The term “Key Ecological Function” communities; the habitats have been grouped because of
(KEF) refers to the principal set of ecological roles their general ecological similarities: Westside and High
performed by each species in its ecosystem. KEFs directly Montane Forests; Eastside (Interior) Forest and
highlight the influence of organisms on their Woodlands; Alpine and Subalpine; Westside Grasslands
environments, and how the presence (or absence) of a and Chaparral; Eastside (Interior) Shrubland and
species serves to modify its own environment as well as Grasslands; Urban; Agriculture, Pastures, and Mixed
the environments of other organisms (see Chapter 6). By Environs; Riparian; and Coastal and Marine. Chapters 16,
using KEFs, we can begin to assess cumulative effects in a 17, and 18 deal with introduced species, genetic
more consistent context because we can evaluate considerations for introduced and augmented
management actions in terms of their influences on populations, and an examination of the species that have
ecological functions—via habitat elements and species been lost from the terrestrial and marine environs,
relationships, and through the functions provided by these respectively. Chapters 19 through 25 report on some of
species—over time and space. the most recent thinking in regard to characterizing
By design, the text within this book makes limited wildlife species at risk; land-use activities (Chapter 20
management recommendations. Several factors guided directly supports the Management Activity Links matrix);
our rationale for this: (1) the main focus of this project modeling wildlife and their habitats, integrating wildlife
was to offer credible, scientific evidence on wildlife— information into the planning process; assessing snags,
habitat relationships; (2) our area of interest covered a two- green trees, and down wood; and considering multi-
state region (including the marine waters out to the 200- species planning while trying to address single-species
mile exclusive economic zone) and management actions issues. Chapter 26 closely examines salmon and other
are often localized, directed by state and federal policies aquatic resource relationships with wildlife and is linked
or regulations, and rarely does one prescription fit all to the Salmon–Wildlife Relationships matrix. The final
situations; and (3) management recommendations can chapter (27), a summary, gives an introduction to what
become outdated relatively quickly. Our position was that can be found on the CD-ROM. A complete text of the
whereas management recommendations come and go, the metadata is found on the CD-ROM.
scientific information on which management actions are
based (the focus herein) will span a longer time frame. In CD-ROM Organization
summary, our principal intent was to make scientifically- The digital wildlife habitat relationship information that
rigorous and ecosystem-based wildlife information accompanies this book is developed as seven matrixes that
available to resource managers and interested others, are found on the CD-ROM. These data sets comprised the
thereby allowing them to pursue management options that following matrixes: wildlife habitat types, structural
fit their local, watershed, or regional situation. conditions, habitat elements, key ecological functions, life
histories, management activity links, and salmon–wildlife
Chapter Organization relationships. Citations are also given to support the
The chapters that follow are designed to help the reader information in the Life History and Management Activity
become aware of new concepts and approaches as well as Links matrixes.
give concrete examples using the matrixes. The maps on Layout of the information on the CD-ROM is done to
pages vii-xi depict historic (circa 1850) and current (circa report data by wildlife species or by wildlife habitat. There
1999) maps of Oregon and Washington to give the reader are canned queries that allow the user to review
an idea of how and where the wildlife-habitat types have predetermined information, like a list of species that
changed over time. Chapters 1 and 2 describe the process require a snag or are generally associated with lowland
of how the wildlife habitats were derived and contain a mixed conifer forests.
detailed description of each habitat. These chapters The information is accessible on the CD-ROM by using
support and highlight the Wildlife Habitat Types matrix. the Internet Explorer, version 5.0 or greater. By using the
Chapter 3 describes the structural conditions and land Internet browser, the user can print out any of the
use/land cover classes and the importance of habitat information that is displayed. For a description of the
elements in considering management of a site, and is metadata and definition of terms, please see Chapters 1,
supported by the Structural Conditions and Habitat Elements 2, 3, 6, 20, and 26.
matrixes. Chapter 4 offers a discussion of how habitat
elements can be designed into forest management Conclusion
schemes. Chapter 5 provides a review of the differing Because the primary focus of our effort has been wildlife
efforts useful in identifying and prioritizing areas and habitat, we have tried not to lose sight of the population
habitats for conservation, and offers a summary of the dynamics that are associated with habitat changes. With
amounts of each of the 32 wildlife-habitat types in Oregon this in mind, we had an opportunity while convening the
and Washington and the relative amounts of each under panels of species experts to ask them what they thought
current conservation regimes. Chapter 6 is directly linked the population status was for each species by state. As a
to the Key Ecological Functions matrix and gives a rationale result, 558 species were assessed for their population
and examples of how to use this data set. The next nine status, with about 10% decreasing, 10% increasing, 40%
 INTRODUCTION xix

stable, and another 40% unknown. As far as we know this 8. Senge, P. M. 1994. The Fifth Discipline. Doubleday Publications,
is the first time that there has been an attempt to survey New York, NY.
species’ experts to establish a trend estimate for each 9. Thomas, J. W., technical editor. 1979. Wildlife habitats in
species by state. managed forests: the Blue Mountains of Oregon and
As we face the new millennium, we recognize that Washington. U.S. Forest Service, Agriculture Handbook 553.
ecosystems are dynamic and evolutionary, and that Washington, D.C.
steady-state management solutions to environmental 10. ———. 1991. Research on wildlife in old growth forests:
issues aren’t always possible or appropriate. Ecosystems setting the stage. Pages 1-4 in Wildlife and vegetation of
are organized within the hierarchical scales of time and unmanaged Douglas-fir forests. U.S. Forest Service, General
Technical Report PNW-GT-285. Pacific Northwest Forest and
space. They are limited in their ability to adapt to changes,
Range and Experiment Station, Portland, OR.
which may be biophysically or socially defined, although
11. Von Altrock, C. 1995. Fuzzy logic and neurofuzzy applications
some limits may be mitigated with the input of resources
explained. Prentice-Hall, NJ.
and energy. There are also limits to the predictability of
ecosystem patterns and processes; some conditions and
events may be predictable at some temporal and spatial
scales, but not at others. Ecosystem-based management
is an abstract idea, and hence is not an end in itself. Rather,
it is an approach to try and achieve sustainable conditions
and provide wildlife habitat, outdoor recreation,
wilderness, water, wood, mineral resources, and food
while retaining the aesthetic, historical, and spiritual
qualities of the land. It joins the needs of people and
environmental values in such a way that Oregon’s and
Washington’s forests, grasslands, lakes, streams, and
marine environments can continue to represent diverse,
healthy, productive and sustainable ecosystems.
Wildlife-habitat relationship information is
fundamental to our pursuit of ecosystem-based
conservation efforts. We hope this book and its related
products meaningfully assist you in your endeavors, and
provide a springboard for the continued evolution of
conservation, scientific thought, and education.

Literature Cited
1. Brown, E. R., technical editor. 1985. Management of wildlife and
fish habitats in forests of western Oregon and Washington
(Volume 1 & 2). U.S. Forest Service, Publication R6-F&WL-192-
1985. Pacific Northwest Region, Portland, OR.
2. Capp, J., B. Carter, J. Diebert, J. Inman, and E. Styskel. 1976.
Wildlife habitats relations of south central Oregon. U.S. Forest
Service, Bend, OR.
3. Leopold, A. 1976. A Sand County Almanac. Oxford University
Press, New York, NY.
4. Maser, C., J. W. Thomas, and R. G. Anderson. 1984. Wildlife
habitats in managed rangelands—the great basins of
southwestern Oregon. U.S. Forest Service, General Technical
Report PNW-172. Pacific Northwest Forest and Range and
Experiment Station, Portland, OR.
5. Morrison, M., B. Marcot, and R. Mannan. 1992. Wildlife—habitat
Relationships—concepts and applications. University of
Wisconsin Press, Madison, WI.
6. ———, ———, ———. 1998. Wildlife—habitat
relationships—concepts and applications. 2nd Edition.
University of Wisconsin Press, Madison, WI.
7. Proctor, C. M., J. C. Garcia, D.V. Galvin, G. C. Lewis, L. C. Loehr,
and A. M. Massa. 1980. An ecological characterization of the
Pacific Northwest coastal region. 5 volumes. U.S. Fish and
Wildlife Service, Biological Services Program. FWS/OBS-79/11
through 79/15.
Recommended citations:
Book—Johnson, David H., and Thomas A. O’Neil
(managing directors). 2001. Wildlife-Habitat
Relationships in Oregon and Washington. Oregon
State University Press, Corvallis, OR. 768 p.
CD-ROM—O’Neil, Thomas A., David H. Johnson,
Charley Barrett, Marla Trevithick, Kelly A. Bettinger,
Chris Kiilsgaard, Madeleine Vander Heyden, Eva L.
Greda, Derek Stinson, Bruce G. Marcot, Patrick J.
Doran, Susan Tank, and Laurie Wunder. 2001.
Matrixes for Wildlife-Habitat Relationships in
Oregon and Washington. In D.H. Johnson and T.A.
O’Neil (managing directors). 2001. Wildlife-Habitat
Relationships in Oregon and Washington. Oregon
State University Press, Corvallis, OR. 768 p.
 1
Oregon and Washington Wildlife Species
and Their Habitats
Thomas A. O’Neil & David H. Johnson

Introduction
In this chapter, we address the fundamental issues of We used a number of primary sources to establish/
developing a common nomenclature for wildlife species, confirm species occurrences: Atlas of Oregon Wildlife,14 Land
and determining their occurrence and breeding status Mammals of Oregon,45 Mammals of the Pacific States,26 The
within the two-state area. We then describe the methods Mammals of North America,23 The Mammals and Life Zones of
used to identify the 32 wildlife-habitat types. Combined, Oregon,6 Birds of Oregon,20 Oregon Species Information
the species list and habitats form the backbone for the rest System,36 Oregon Wildlife Diversity Plan,38 Amphibians and
of this book, associated maps, and CD-ROM. Reptiles of Washington State: Location Data and Predicted
Distributions,18 Terrestrial Mammals of Washington State:
Developing the Location Data and Predicted Distributions,29 Breeding Birds of
Washington State: Location Data and Predicted Distributions,40
Wildlife Species List and Oregon and Washington Marine Mammal and Seabird
For almost two hundred years, information has been
Survey.10 We also supplemented the above references with
collected about Oregon’s and Washington’s wildlife
Alexander,1 Aubry,4, 5 Best,7 and Bradley.8
species. These data have been obtained from field studies,
Scientific and common names and species occurrence
surveys, and general observations. Most of this
status by state were reviewed by Dick Johnson
information now resides in either museum collections
(Washington State University), B. J. Verts (Oregon State
throughout the United States, or as records in journals,
University), Tom O’Neil (Northwest Habitat Institute),
publications, and public agency files. Over time, each state
Rolf Johnson (Washington Department of Fish and
has developed, mostly independently, its own wildlife
Wildlife [WDFW]), Derek Stinson (WDFW), Kelly
species list based on these data sets. Thus, a first step for
Bettinger (WDFW), Charlie Bruce (Oregon Department of
us was to bring both states’ wildlife lists together into a
Fish and Wildlife [ODFW]), Kelly McAllister (WDFW),
regional data set. As we developed the joint species list,
Bruce Mate (Oregon State University Marine Science Lab),
we simultaneously determined the occurrence and
Steven Jeffries (WDFW), and Robin Brown (ODFW).
breeding status for each of the species. Developing the
Taxonomic order follows regional publications or
combined species list, and status categories, proved
commonly accepted national books so that cross-
somewhat challenging because the common and scientific
referencing could be made easier. The publications that
names for many species have changed over the decades,
guided taxonomic order were Leonard et al. 31 for
and the status of some species have been confusingly
amphibians; Storm and Leonard 42 for reptiles; The
described as “common,” “regular,” “irregular,”
American Ornithologists’ Union Checklist of North
“incidental,” “accidental,” or “vagrant.”
American Birds, Seventh edition2 for birds; and Hall23 and
Once the joint species list was compiled, a variety of
Verts and Carraway45 for mammals and marine mammals.
sources were used to establish current taxonomic
We settled on five occurrence status categories for the
nomenclature (i.e., scientific and common names) for each
species: Occurs, Accidental, Non-native, Reintroduced, and
species. For most species, initial nomenclatures were
Extirpated; the species could be listed as any one of these
obtained from the Oregon Species Information System.36
categories in either state. To list a species as having an
Collins et al.,13 Leonard et al.,31 and Storm and Leonard42
Occurs status within each state required >15 documented
were used to update the amphibians and reptiles. Bird
observations, that is, they are considered to be regularly
nomenclature follows the recent Checklist of North American
occurring species for the area. This figure of 15
Birds, Seventh Edition by the American Ornithologists’
documented observations was derived from its use by the
Union2. Verts and Carraway45 and Wilson and Reeder48
states’ ornithological groups. Accidental denotes those
were used as the primary sources of scientific names for
species with <15 documented occurrences, or >15 records
mammals; Jones et al.30 and Hall23 were used as secondary
but Oregon and Washington are not a regular part of the
sources for scientific and common mammal names. Frost
species’ range; Non-native denotes species that are not
and Timm19 and van Zyll de Jong44 were consulted for bat
native but now are found in Oregon or Washington;
nomenclature.
1
 2 Wildlife–Habitat Relationships in Oregon and Washington

Reintroduced denotes native species that were eliminated are based on the similarity of many wildlife species using
from Oregon or Washington or reduced to such low a suite of vegetation types. However, a wildlife species’
population levels that additional individuals were “habitat” refers to an individual, species-specific use of a
required to supplement or re-establish the species; and wildlife-habitat type. Thus, habitat is fundamentally
Extirpated refers to a native species whose populations linked to the distribution and abundance of species and
have been completely removed from Oregon or underlies explanations of the factors, patterns, and
Washington. Additionally, several categories were used processes that support the fitness of wildlife at the
to describe the breeding status of the species. Breeds is for individual, population, and community levels, as well as
those species with >5 documented breeding records by their continuing evolution.
separate pairs unless professionals familiar with the Too often the term “habitat type” is misused and is
species believed that breeding is probable but has not yet misunderstood. The term was initially used by
been documented. Non-breeder refers to those species that Daubenmire15 as a way to characterize forest vegetation
occur in the state(s) but do not breed, or have <5 associations by describing the potential of that association
documented breeding records. Bred Historically refers to to reach a specific climax stage and was later refined and
those species that used to breed in the state(s) but currently expanded to forest management.16, 17, 37 This early use of
do not. the term “habitat type” described vegetative associations
Comparing the species lists helps us determine, for the in relationship to seral stage, and actually had no wildlife
first time, the degree of commonality and uniqueness of species use clearly identified with it. Further,
wildlife that exists between Oregon and Washington. The Daubenmire’s habitat types classification dealt with the
combined list of 743 wildlife species now allows us to potential of a vegetative association at a site to move
jointly determine common and scientific names for each towards a climax stage, hence, the forest classification
species (Appendix, page 687). The combined species list described future, not current, conditions. Wildlife use an
further allows us to characterize the occurrence and area because of current conditions, not future ones. Thus,
breeding status of each species by state (Table 1, at end of to a large extent, our past thinking revolved around
chapter). These steps towards standardization are vegetation classification schemes developed essentially for
fundamental for building a common understanding about forest production and inventory purposes, and then
our wildlife because species’ names (both common and overlaid with wildlife attributes.
scientific), occurrence, and breeding status have fluctuated Recently, Hall et al.24 reviewed the literature for the
over time. Reaching agreement on these lists not only appropriate use of the term “habitat type” and concluded
brings the information up to date, but also becomes the that based on the Daubenmire definition the term should
basis for identifying which wildlife species are associated not be used when discussing wildlife-habitat relationships.
with the various habitat types. While we agree in part with Hall et al.’s24 statement, we
A total of 743 wildlife species occur in Oregon and recognize that the problem stems not from usage of the
Washington; 700 of these species occur in Oregon and 651 term in wildlife studies, but rather, that the term needs to
species in Washington. Some 593 species (80%) occur in be redefined by recognizing the interrelationships of
both states, whereas 92 species are unique to Oregon and wildlife with different vegetation types. Further, as Hall
43 are unique to Washington. In the following breakdown et al. 24 stated, there is also a need to standardize
we list the species totals for both states, followed by the terminology. Hence, if we are willing to accept the
total for each state. A total of 479 species Occur in both definition of “habitat” as the resources and conditions
states (548 OR, 501 WA), 80 species are Accidental in both present in an area to result in occupancy by a given wildlife
states (120 OR, 116 WA), 22 are Non-native in both states
(24 OR, 28 WA), no similar species have been Reintroduced
in both states (2 OR, 2 WA), and 2 species have been
Extirpated from both states (6 OR, 4 WA). A total of 385
species Breed in both states, 187 are Non-breeders, and 2
Bred Historically. A summary of the breeding status of the
wildlife species in each state is shown in Figure 1.

Determining the Wildlife-Habitats in

Oregon and Washington
In this book, we use the term wildlife-habitat type or habitat
types to mean a group of vegetation cover types (or land
use/land cover types) that were determined based on the
similarity of wildlife use. Our habitat types are based on
actual conditions (e.g., current vegetation), and therefore
can be mapped, and we assume they contain all the
essential needs for a species’ maintenance and viability. Figure 1. The total number of breeding wildlife species in
Wildlife-habitat types are not species-specific because they Oregon and Washington.
 CHAPTER 1: DEFINING HABITATS 3

species, then a redefinition of the term for wildlife is pooling the 287 alliances into 85 vegetative groups12. To
appropriate. Wildlife species affect, and are affected by these groups, we added 5 agricultural land cover types
their habitats, thus we have approached wildlife-habitat and 1 urban classification. Twenty-eight marine
types as a collection of wildlife species interacting with classifications were also identified, based on a series of
different vegetation communities. meetings with a multi-agency team of marine experts.
Several authors have advocated that wildlife-habitat Thus, a total of 119 vegetative/land and aquatic/marine
relationships be placed in the proper spatial and temporal cover types were identified, and used in the subsequent
scales when they are being assessed.34, 47 That is, we need analysis steps.
to recognize that wildlife-habitat relationships are scale We linked the associations of wildlife species with the
dependent. For example, Johnson28 and Hutto25 suggested following three steps:
that animals select habitat in a hierarchical manner: the 1. Existing wildlife-habitat relationships matrixes9, 11, 33, 43
first level is determined by the species’ geographic range; were incorporated into the Oregon Species
the second is at a level where a species conducts its daily/ Information System that supported the Oregon Gap
seasonal activities (i.e., home range); and the third level is Analysis project.
for habitat components that are local or site-specific within 2) The Oregon Gap data were then brought together
their home ranges. In recognizing the scale dependency with other regional projects that classified vegetation
of wildlife-habitat relationships, we have also structured for wildlife purposes, namely the Washington Gap
the wildlife-habitat data associated with this book to go Analysis and the Interior Columbia Basin Ecosystem
from a coarse level or wildlife-habitat type (that includes Management Project. Although these efforts, taken
the essential components for a species to survive and individually, only covered a portion of the two-state
reproduce), to an intermediate spatial level or structural area, taken in composite, these efforts offered
condition, to the finest level of habitat elements that can be information for the entire area, and contained the
found at a specific site. Our hope is that knowing the desired presence/absence information on the wildlife
species’ relationship with its habitat type, structural species for the suite of the vegetation/land use
conditions and habitat elements will help us make better groupings.
predictions for species occurrences and ecological 3) Crosswalks were then developed between the
conditions for an area. Knowing that ecological condition vegetative and land use cover types from each of the
is based on physical parameters should also help us to projects to the appropriate 85 vegetative groups, 5
identify the ecological processes that are operating (as well agricultural land cover, and 1 urban types. Links to
as missing) in an area. the wildlife occurrence data occurred simultaneously.
Presently, our ability to identify floristic types is greater We then reviewed the wildlife occurrence data (from
than our ability to identify associated fauna. Therefore, the combined projects) for inconsistencies.
each identifiable vegetation type probably does not We selected 541 native breeding species to use in
represent a unique wildlife-habitat. 33 We present an conjunction with the 119 classes of vegetative/land cover/
approach to classify vegetation alliances into wildlife- marine types for the subsequent analysis. We used native
habitat types based on the similarity of native breeding breeding species because we believe existing habitat
terrestrial vertebrate species. The basis for this approach information for breeders is more accurate and species
has been discussed in detail by O’Neil et al.35 This approach preferences for breeding habitat are more specific than for
has its advantage because the status of our knowledge of other associations.35 We then used the group of wildlife
wildlife and their habitat relationships has increased species associated with each of the 119 classes as a set of
dramatically, and the process is quantitative and attributes to describe each class. This increased class
repeatable. We summarize the approach as follows. specificity increased our ability to discriminate effectively
Our initial task was to identify plant alliances, as defined among similar vegetative/land cover/marine types.
by dominant plant species that are found in Oregon and We constructed a matrix of 119 rows (vegetation/land
Washington. We used the national vegetation classification cover/marine types) by 541 columns (wildlife species),
system of Grossman et al.22 and Anderson et al.3 as the and populated it with wildlife use data. In the matrix, a
basis for identifying the 287 plant alliances found in “1” indicate that the wildlife species could be found using
Oregon and Washington. Although there are many the vegetation/land cover/marine class, and a “0”
vegetation classification systems available, we chose this indicated absence or limited use of the class. A wildlife
classification system because it addresses current natural species could have an association with any number of the
vegetation cover, allowed both states a unified 119 classes.
classification scheme, and was hierarchical in design, thus Multivariate statistics were then used to undertake a
providing sufficient detail for linking it to wildlife use. cluster analysis process. Consistent patterns of wildlife
Upon review of the 287 plant alliances, we recognized association within groups suggest that wildlife species
that the plant alliances represented a level of detail too perceive the groups as a similar habitat. From this matrix
fine for ascribing wildlife use. To remedy this, we we calculated wildlife species similarity between each pair
contracted with a member(s) from each state’s Natural of types using two different coefficients (Jaccard and
Heritage Program to create alliance groupings based on Ochaia) as a measure of species’ association.21, 27, 32 The
similar vegetative communities; this effort resulted in difference between the two coefficients is that the Jaccard
 4 Wildlife–Habitat Relationships in Oregon and Washington

gives equal weight to the absence or presence of a species used the following categories for characterizing the degree
sharing a vegetation type and the Ochaia gives weight to of association:
species sharing vegetation type(s). In general, the two
Closely Associated. A species is widely known to depend
coefficients gave similar results. We then created a distance
on a habitat or structural condition for part or all of its life
matrix (species’ dissimilarity) for vegetation types (119 x
history requirements. Identifying this association implies
119) by subtracting the Jaccard or Ochaia coefficients from
that the species has an essential need for this habitat or
1. To groups whose vegetation types had high species’
structural condition for its maintenance and viability.
similarities, we performed a cluster analysis using the
Some species may be closely associated with more than
distance matrix and Ward linkage method,46 a sequential,
one habitat or structural condition, others may be closely
agglomerative, hierarchic, nonover-lapping cluster
associated with only one habitat or structural condition.
strategy,41 which minimizes the within-cluster variance to
Examples of species exhibiting close associations are red-
the between-cluster variance at each step. We then
winged blackbirds to wetland habitats, and spotted owls
reviewed each vegetation type’s membership (and the
to mature and giant tree structural conditions.
wildlife species associated with it) with each set of clusters
to see that it was consistent with our ecological Generally Associated. A species exhibits a high degree of
understanding of plant and wildlife associations. If we adaptability and may be supported by a number of
questioned the vegetation membership within a cluster habitats or structural conditions. In other words, the
group then we also reviewed the wildlife species habitats or structural conditions play a supportive role
associated with the particular vegetation type. Experts for its maintenance and viability. Examples include the
then reviewed our initial groups and made suggestions black bear’s association with a variety of forested habitats
for refining them. A total of 32 wildlife-habitats in Oregon or black-tailed deer associated with a number of structural
and Washington were determined from this process. Table conditions.
2 illustrates the results that were achieved using this
Present. A species demonstrates occasional use of a habitat
method. The Frontispiece reflects the mapping of these
or structural condition. The habitat or structural condition
32 wildlife habitats.
provides marginal support to the species for its
maintenance and viability. Examples are the rough-legged
Wildlife-Habitat Data Matrix hawk in desert playa and salt scrub or mink in a montane
To maximize the utility of wildlife-habitat relationship mixed conifer forest. Table 3 depicts the total number of
information, a digital database that links wildlife with its species associated with each wildlife habitat.
habitats can be found on the CD-ROM included with this Finally, the expert panelists assigned an overall
book. Wildlife occurrence with a particular habitat type confidence rating to the occurrence and activity headings
was determined through an expert panel process held for each species within each habitat type. The confidence
during the summer of 1998. The expert panels while ratings were simply high (e.g., many peer or published
cloistered had available to them range maps for each accounts), moderate, and low (e.g., few or no published
species, wildlife-habitat types distribution maps, accounts). By ascribing a confidence rating, our objective
statewide vegetation maps, and a variety of reference was to offer users an evaluation of the overall strength of
materials. These sources of material helped the panelists the scientific evidence. We understand that this is the first
to determine wildlife species occurrence with a particular time a confidence rating system has been applied to
habitat type. The codes used were: Y—Yes the species occurs, wildlife-habitat relationship information of this type.
H—Historically occurred, and U—Unsure. Alongside the
occurrence category, the panelist also identified the types Literature Cited
of activity that the species performs while utilizing the 1. Alexander, L. F. 1996. A morphometric analysis of geographic variation
habitat. The activity codes for the wildlife species within within Sorex monticolus (Insectivora: Soricidae). Miscellaneous
a particular habitat were: B—Both feeds and breeds, F—Feeds Publication. University of Kansas, Natural History Museum 88: 1-54.
2. American Ornithologists’ Union. 1998. Checklist of North American
only, R—Reproduces only, and O—Other. The Other category
birds. Seventh Edition, American Ornithologists’ Union, Washington
reflects activities such as roosting/resting, hibernacula, or D.C.
use of the habitat for cover (thermal and hiding) purposes. 3. Anderson, M. P. Bourgeron, M. Bryer, R. Crawford, L. Engelking, D.
Faber-Langendoen, M. Gallyoun, D. H. Grossman, K. Goodin, S. Landaal,
Defining the Level of Associations K. Metzler, K. P. Patterson, M. Pyne, M. Reid,
L. Sneddon, A. W. Weakley. 1998. International classification of ecological
Between Wildlife and the Habitats communities: terrestrial vegetation of the United States.Volume II: List
The science of wildlife ecology continues to embrace new of vegetation types. The Nature Conservancy, Boulder, CO.
concepts and frameworks. The relatively new paradigm 4. Aubry, K. B. 1982. The recent history and present distribution of the
of ecosystem management challenges us to broaden red fox in Washington. Paper presented at the 63rd annual meeting,
Pacific Northwest Bird and Mammal Society, May 1, 1982.
traditional definitions and incorporate concepts that allow
5. ———, and D. B. Houston. 1992. Distribution and status of the fisher
us to assess the value of landscapes to single as well as (Martes pennanti) in Washington. Northwestern Naturalist 73: 69-79.
multi-species strategies. One such concept that we are 6. Bailey,V. 1936. The mammals and life zones of Oregon. U.S. Department
building on is the degree of association between wildlife of Agriculture, Bureau of Biological Survey, North America Fauna No.
species and their habitats.39 For our purpose, the panelists 55: 1-416.
 CHAPTER 1: DEFINING HABITATS 5

7. Best, T. L. 1988. Morphologic variation in the spotted bat Euderma 28. Johnson, D. 1980. The comparison of usage and availability
maculatum. American Midland Naturalist 119: 244-252. measurements for evaluating resource preference. Ecology 61: 65-71.
8. Bradley, W. P. 1982. Some observations on feral horses on the Yakima 29. Johnson, R. E. and K. M. Cassidy. 1997. Terrestrial mammals of
Reservation. Paper presented at the 63rd annual meeting of the Pacific Washington State: Location data and predicted distributions. Volume 3
Northwest Bird and Mammal Society, May 1, 1982. in: K. M. Cassidy, C. E. Grue, M. R. Smith, and K. M. Dvornich, editors.
9. Brown, E. R., technical editor. 1985. Management of wildlife and fish Washington State Gap Analysis, Final Report. Washington Cooperative
habitats in forests of western Oregon and Washington (Volume 1 & 2). Fish and Wildlife Research Unit, University of Washington, Seattle, WA.
U.S. Forest Service, Publication R6-F&WL-192-1985. Pacific Northwest 30. Jones, J. K., R. S. Hoffman, D. W. Rice, C. Jones, R. J. Baker, and M. D.
Region, Portland, OR. Engstrom. 1992. Revised checklist of North American mammals north
10. Brueggeman, J. J. 1992. Oregon and Washington marine mammal and of Mexico, 1991. Occasional Papers, The Museum Texas Tech University,
seabird surveys. Pacific OCS Region, Mineral Management Service, U.S. Lubbock, TX.
Department of Interior, Los Angeles, CA. 31. Leonard, W. P., H.A. Brown, L. L. C. Jones, K. R. McAllister, and R. M.
11. Capp, J., B. Carter, J. Diebert, J. Inman, and E. Styskel. 1976. Wildlife Storm. 1993. Amphibians of Washington and Oregon. Published by the
habitats relations of south central Oregon. U.S. Forest Service, Bend, Seattle Audubon Society, Seattle, WA.
OR. 32. Ludwig, J. A., and J. F. Reynolds. 1988. Statistical ecology. John Wiley and
12. Chappell, C., R. Crawford, J. Kagen, and P. Doran. 1997. Sons, New York, NY.
A vegetation, land use, and habitat classification system for the 33. Maser. C.J., J. W. Thomas, and R. G. Anderson. 1984. Wildlife habitats in
terrestrial and aquatic systems of Oregon and Washington. Wildlife managed rangelands—the Great Basins of southeastern Oregon. U.S.
habitats and species associations in Oregon and Washington—building Forest Service, Pacific Northwest Forest and Range and Experiment
a common understanding. Progress Report 3. Washington Department Station, General Technical Report PNW-172. Portland, OR.
of Fish and Wildlife, Olympia, WA. 34. Morrison, M., B.G. Marcot, and R. Mannan. 1992. Wildlife-habitat
13. Collins, J. T. 1990. Standard common and current scientific names for relationships: concepts and applications. Univ. Wisconsin Press, Madison,
North American amphibians and reptiles, Third edition. Society for the WI.
Study of Amphibians and Reptiles, University of Kansas, Herpetological 35. O’Neil, T. A., R. J. Steidl, W. D. Edge, and B. Csuti. 1995. Using wildlife
Circular Number 19, Lawerence, KA. communities to improve vegetation classification for conserving
14. Csuti, B., A. J. Kimmerling, T. A. O’Neil, M. M. Shaughnessy, biodiversity. Conservation Biology, Volume 9 No. 6, 1482-1491.
E. P. Gaines, M. M. P. Huso. 1997. Atlas of Oregon wildlife. Oregon State 36. Oregon Department of Fish and Wildlife. 1994. Oregon species
University Press, Corvallis, OR. information system. User manual developed by
15. Daubenmire, R. 1952. Forest vegetation on northern Idaho and W. McKenzie, S. Olson-Edge, and T. O’Neil. Corvallis, OR.
adjacent Washington, and its bearing on concepts of vegetation 37. Pfister, R., B. Kovalchik, S. Arno, and R. Presby. 1977. Forest habitat
classification. Ecological Monographs 22: 301-330. types of Montana. U.S. Forest Service General Technical Report INT-34.
16.———. 1968. Plant communities: a textbook of plant synecology. Intermountain Forest & Range Experiment Station, Ogden, UT.
Harper and Row, New York, NY. 38. Puchy, C., and D. Marshall. 1993. Oregon wildlife diversity plan.
17. ———, and J. B. Daubenmire. 1968. Forest vegetation of eastern Oregon Department of Fish and Wildlife, Portland, OR.
Washington and northern Idaho. Washington Agriculture Experimental 39. Ruggiero, L., L. Jones, and K. Aubry. 1991. Plant and animal habitat
Station, Technical Bulletin 60. associations in Douglas-fir forests of the Pacific Northwest: an
18. Dvornich, K. M., K. R. McAllister, and K. Aubry. 1997. Amphibians and overview. Pp. 447-450 in: Wildlife and vegetation of unmanaged
reptiles of Washington State: Location data and predicted distributions Douglas-fir forests. U.S. Forest Service General Technical Report PNW-
in: K. M. Cassidy, C. E. Grue, M. R. Smith, and K. M. Dvornich, editors. GTR-285. Portland, OR.
Washington Cooperative Fish and Wildlife Research Unit, University of 40. Smith, M. R., P. W. Mattocks Jr., and K. M. Cassidy. 1997. Breeding birds
Washington, Seattle,WA. of Washington State: Location data and predicted distributions. Volume
19. Frost, D. R., and R. M. Timm. 1992. Phylogeny of plecotine bats 4 in: K. M. Cassidy, C. E. Grue,
(Chiropetera: Vespertilionidae): summary of the evidence and proposal M. R. Smith, and K. M. Dvornich, editors. Washington State Gap Analysis,
of a logically consistent taxonomy. American Museum Novitate 3034: 1- Final Report. Seattle Audubon Society Publications in Zoology No. 1,
16. Seattle, WA.
20. Gilligan, J., M. Smith, D. Rogers, and A. Contreras. Birds of Oregon. 41. Sneath, P. H. A., and R. R. Sokal. 1973. Numerical taxonomy.
Cinclus Publications, McMinnville, OR. W. H. Freeman, San Francisco, CA.
21. Goodall, D. W. 1973. Sample similarity and species correlation. Pp. 99- 42. Storm, R. M., and W. P. Leonard. 1995. Reptiles of Oregon and
149 in: R. H. Whittaker, editor. Ordination and classification of Washington. Seattle Audubon Society, Seattle, WA.
communities. W. Junk, The Hague. 43. Thomas, J. W., technical editor. 1979. Wildlife habitats in managed
22. Grossman, D. H., D. Faber-Langendoen, A. W. Weakley, forests: the Blue Mountains of Oregon and Washington. U.S. Forest
M. Anderson, P. Bourgeron, R. Crawford, K. Goodin, S. Landaal, K. Service, Agriculture Handbook 553. Washington, D.C.
Metzler, K. D. Patterson, M. Pyne, M. Reid and L. Sneddon. 1998. 44. van Zyll de Jong, C. 1984. Taxonomic relationships of Nearctic small-
International classification of ecological communities: terrestrial footed bats of the Myotis leibii group (Chiroptera:Vespertilionidae).
vegetation of the United States. Volume I : The National Vegetation Canadian Journal of Zoology 62: 2519-2526.
Classification Standard. The Nature Conservancy, Boulder, CO. 45. Verts, B. J., and L. N. Carraway. 1998. Land mammals of Oregon.
23. Hall, R. E. 1981. The mammals of North America. Second Edition. John University of California Press, Berkeley and Los Angeles, CA.
Wiley & Sons, New York, NY. 46. Ward, J. H., Jr. 1963. Hierarchial grouping to optimize an objective
24. Hall, L. S., P. R. Krausman, and M. L. Morrison. 1997. The habitat function. Journal of the American Statistical Association 58: 236-244.
concept and a plea for standard terminology. Wildlife Society Bulletin 47. Wiens, J. A. 1981. Scale problems in avian censusing. Studies in Avian
25 (1): 173-182. Biology. 6: 513-521.
25. Hutto, R. L. 1985. Habitat selection by nonbreeding, migratory land 48. Wilson, D. E., and D. M. Reeder. 1993. Mammal species of the world,
birds. Pp. 455-476 in: M. L. Cody, editor. Habitat selection in birds. 2nd edition. Smithsonian Institution Press, Washington, D.C.
Academic Press, Orlando, FL.
26. Ingles, L. G. 1965. Mammals of the Pacific States. Stanford University
Press, Stanford, CA.
27. Jaccard, P. 1901. Distribution de la flore alpine dans le Bassin des
Dranes et dans quelques regions voisines. Bulletin Societe Vaudoise des
Sciences Naturelles 37: 241-272.
 6 Wildlife–Habitat Relationships in Oregon and Washington

Table 1. Wildlife species occurrence and breeding status in Oregon and Washington.

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Amphibians
Tiger Salamander Occurs Breeds Occurs Breeds
Northwestern Salamander Occurs Breeds Occurs Breeds
Long-toed Salamander Occurs Breeds Occurs Breeds
Cope’s Giant Salamander Occurs Breeds Occurs Breeds
Pacific Giant Salamander Occurs Breeds Occurs Breeds
Olympic Torrent Salamander Does Not Occur Not Applicable Occurs Breeds
Columbia Torrent Salamander Occurs Breeds Occurs Breeds
Southern Torrent Salamander Occurs Breeds Does Not Occur Not Applicable
Cascade Torrent Salamander Occurs Breeds Occurs Breeds
Rough-skinned Newt Occurs Breeds Occurs Breeds
Dunn’s Salamander Occurs Breeds Occurs Breeds
Larch Mountain Salamander Occurs Breeds Occurs Breeds
Van Dyke’s Salamander Does Not Occur Not Applicable Occurs Breeds
Western Red-backed Salamander Occurs Breeds Occurs Breeds
Del Norte Salamander Occurs Breeds Does Not Occur Not Applicable
Siskiyou Mountains Salamander Occurs Breeds Does Not Occur Not Applicable
Ensatina Occurs Breeds Occurs Breeds
Clouded Salamander Occurs Breeds Does Not Occur Not Applicable
Black Salamander Occurs Breeds Does Not Occur Not Applicable
Oregon Slender Salamander Occurs Breeds Does Not Occur Not Applicable
California Slender Salamander Occurs Breeds Does Not Occur Not Applicable
Tailed Frog Occurs Breeds Occurs Breeds
Great Basin Spadefoot Occurs Breeds Occurs Breeds
Western Toad Occurs Breeds Occurs Breeds
Woodhouse’s Toad Occurs Breeds Occurs Breeds
Pacific Chorus (Tree) Frog Occurs Breeds Occurs Breeds
Red-legged Frog Occurs Breeds Occurs Breeds
Cascades Frog Occurs Breeds Occurs Breeds
Oregon Spotted Frog Occurs Breeds Occurs Breeds
Columbia Spotted Frog Occurs Breeds Occurs Breeds
Foothill Yellow-legged Frog Occurs Breeds Does Not Occur Not Applicable
Northern Leopard Frog Occurs Breeds Occurs Breeds
Bullfrog Non-native Breeds Non-native Breeds
Green Frog Does Not Occur Not Applicable Non-native Breeds
Reptiles
Snapping Turtle Non-native Non-Breeder Non-native Breeds
Painted Turtle Occurs Breeds Occurs Breeds
Western Pond Turtle Occurs Breeds Occurs Breeds
Red-eared Slider Turtle Non-native Breeds Non-native Breeds
Loggerhead Sea Turtle Accidental Non-Breeder Accidental Non-Breeder
Green Sea Turtle Accidental Non-Breeder Accidental Non-Breeder
Pacific Ridley Sea Turtle Accidental Non-Breeder Accidental Non-Breeder
Leatherback Turtle Occurs Non-Breeder Occurs Non-Breeder
Northern Alligator Lizard Occurs Breeds Occurs Breeds
Southern Alligator Lizard Occurs Breeds Occurs Breeds
Mojave Black-collared Lizard Occurs Breeds Does Not Occur Not Applicable
Long-nosed Leopard Lizard Occurs Breeds Does Not Occur Not Applicable
Short-horned Lizard Occurs Breeds Occurs Breeds
Desert Horned Lizard Occurs Breeds Does Not Occur Not Applicable
Sagebrush Lizard Occurs Breeds Occurs Breeds
Western Fence Lizard Occurs Breeds Occurs Breeds
Side-blotched Lizard Occurs Breeds Occurs Breeds
Western Skink Occurs Breeds Occurs Breeds
 CHAPTER 1: DEFINING HABITATS 7

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Western Whiptail Occurs Breeds Does Not Occur Not Applicable

Plateau Striped Whiptail Non-native Breeds Does Not Occur Not Applicable
Rubber Boa Occurs Breeds Occurs Breeds
Racer Occurs Breeds Occurs Breeds
Night Snake Occurs Breeds Occurs Breeds
Sharptail Snake Occurs Breeds Occurs Breeds
Ringneck Snake Occurs Breeds Occurs Breeds
Night Snake Occurs Breeds Occurs Breed
Common Kingsnake Occurs Breeds Does Not Occur Not Applicable
California Mountain Kingsnake Occurs Breeds Occurs Breeds
Striped Whipsnake Occurs Breeds Occurs Breeds
Gopher Snake Occurs Breeds Occurs Breeds
Western Ground Snake Occurs Breeds Does Not Occur Not Applicable
Pacific Coast Aquatic Garter Snake Occurs Breeds Does Not Occur Not Applicable
Western Terrestrial Garter Snake Occurs Breeds Occurs Breeds
Northwestern Garter Snake Occurs Breeds Occurs Breeds
Common Garter Snake Occurs Breeds Occurs Breeds
Western Rattlesnake Occurs Breeds Occurs Breeds
Birds
Red-throated Loon Occurs Non-Breeder Occurs Non-Breeder
Pacific Loon Occurs Non-Breeder Occurs Non-Breeder
Common Loon Occurs Non-Breeder Occurs Breeds
Yellow-billed Loon Accidental Non-Breeder Occurs Non-Breeder
Pied-billed Grebe Occurs Breeds Occurs Breeds
Horned Grebe Occurs Breeds Occurs Breeds
Red-necked Grebe Occurs Breeds Occurs Breeds
Eared Grebe Occurs Breeds Occurs Breeds
Western Grebe Occurs Breeds Occurs Breeds
Clark’s Grebe Occurs Breeds Occurs Breeds
Shy Albatross Does Not Occur Not Applicable Accidental Non-Breeder
Laysan Albatross Occurs Non-Breeder Occurs Non-Breeder
Black-footed Albatross Occurs Non-Breeder Occurs Non-Breeder
Short-tailed Albatross Occurs Non-Breeder Occurs Non-Breeder
Northern Fulmar Occurs Non-Breeder Occurs Non-Breeder
Murphy’s Petrel Accidental Non-Breeder Accidental Non-Breeder
Mottled Petrel Accidental Non-Breeder Accidental Non-Breeder
Cook’s Petrel Does Not Occur Not Applicable Accidental Non-Breeder
Pink-footed Shearwater Occurs Non-Breeder Occurs Non-Breeder
Flesh-footed Shearwater Occurs Non-Breeder Occurs Non-Breeder
Buller’s Shearwater Occurs Non-Breeder Occurs Non-Breeder
Sooty Shearwater Occurs Non-Breeder Occurs Non-Breeder
Short-tailed Shearwater Occurs Non-Breeder Occurs Non-Breeder
Manx Shearwater Does Not Occur Not Applicable Accidental Non-Breeder
Black-vented Shearwater Accidental Non-Breeder Does Not Occur Not Applicable
Wilson’s Storm-petrel Accidental Non-Breeder Accidental Non-Breeder
Fork-tailed Storm-petrel Occurs Breeds Occurs Breeds
Leach’s Storm-petrel Occurs Breeds Occurs Breeds
Black Storm-petrel Accidental Non-Breeder Does Not Occur Not Applicable
Red-billed Tropicbird Does Not Occur Not Applicable Accidental Non-Breeder
Blue-footed Booby Does Not Occur Not Applicable Accidental Non-Breeder
American White Pelican Occurs Breeds Occurs Breeds
Brown Pelican Occurs Non-Breeder Occurs Non-Breeder
Brandt’s Cormorant Occurs Breeds Occurs Breeds
Double-crested Cormorant Occurs Breeds Occurs Breeds
Pelagic Cormorant Occurs Breeds Occurs Breeds
Magnificent Frigatebird Accidental Non-Breeder Accidental Non-Breeder
American Bittern Occurs Breeds Occurs Breeds
 8 Wildlife–Habitat Relationships in Oregon and Washington

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Least Bittern Occurs Breeds Does Not Occur Not Applicable

Great Blue Heron Occurs Breeds Occurs Breeds
Great Egret Occurs Breeds Occurs Breeds
Snowy Egret Occurs Breeds Accidental Non-Breeder
Little Blue Heron Accidental Non-Breeder Accidental Non-Breeder
Tricolored Heron Accidental Non-Breeder Does Not Occur Not Applicable
Cattle Egret Occurs Breeds Occurs Non-Breeder
Green Heron Occurs Breeds Occurs Breeds
Black-crowned Night-heron Occurs Breeds Occurs Breeds
Yellow-crowned Night-heron Does Not Occur Not Applicable Accidental Non-Breeder
White-faced Ibis Occurs Breeds Occurs Non-Breeder
Turkey Vulture Occurs Breeds Occurs Breeds
California Condor Extirpated Bred Historically* Extirpated Bred Historically*
Fulvous Whistling-Duck Accidental Non-Breeder Accidental Non-Breeder
Greater White-fronted Goose Occurs Non-Breeder Occurs Non-Breeder
Emperor Goose Accidental Non-Breeder Accidental Non-Breeder
Snow Goose Occurs Non-Breeder Occurs Non-Breeder
Ross’s Goose Occurs Non-Breeder Occurs Non-Breeder
Canada Goose Occurs Breeds Occurs Breeds
Western Canada Goose Occurs Breeds Occurs Breeds
Giant Canada Goose Non-native Breeds Non-native Breeds
Taverner’s Canada Goose Occurs Non-Breeder Occurs Non-Breeder
Aleutian Canada Goose Occurs Non-Breeder Occurs Non-Breeder
Cackling Canada Goose Occurs Non-Breeder Occurs Non-Breeder
Dusky Canada Goose Occurs Non-Breeder Occurs Non-Breeder
Vancouver Canada Goose Occurs Non-Breeder Occurs Non-Breeder
Lesser Canada Goose Occurs Non-Breeder Occurs Non-Breeder
Brant Occurs Non-Breeder Occurs Non-Breeder
Mute Swan Non-native Breeds Non-native Breeds
Trumpeter Swan Occurs Breeds Occurs Breeds
Tundra Swan Occurs Non-Breeder Occurs Non-Breeder
Whooper Swan Accidental Non-Breeder Does Not Occur Not Applicable
Wood Duck Occurs Breeds Occurs Breeds
Gadwall Occurs Breeds Occurs Breeds
Falcated Duck Does Not Occur Not Applicable Accidental Non-Breeder
Eurasian Wigeon Occurs Non-Breeder Occurs Non-Breeder
American Wigeon Occurs Breeds Occurs Breeds
American Black Duck Accidental Non-Breeder Non-native Breeds
Mallard Occurs Breeds Occurs Breeds
Blue-winged Teal Occurs Breeds Occurs Breeds
Cinnamon Teal Occurs Breeds Occurs Breeds
Northern Shoveler Occurs Breeds Occurs Breeds
Northern Pintail Occurs Breeds Occurs Breeds
Garganey Accidental Non-Breeder Accidental Non-Breeder
Baikal Teal Accidental Non-Breeder Does Not Occur Not Applicable
Green-winged Teal Occurs Breeds Occurs Breeds
Canvasback Occurs Breeds Occurs Breeds
Redhead Occurs Breeds Occurs Breeds
Ring-necked Duck Occurs Breeds Occurs Breeds
Tufted Duck Accidental Non-Breeder Accidental Non-Breeder
Greater Scaup Occurs Non-Breeder Occurs Non-Breeder
Lesser Scaup Occurs Breeds Occurs Breeds
Steller’s Eider Accidental Non-Breeder Accidental Non-Breeder
King Eider Accidental Non-Breeder Accidental Non-Breeder
Harlequin Duck Occurs Breeds Occurs Breeds
Surf Scoter Occurs Non-Breeder Occurs Non-Breeder
White-winged Scoter Occurs Non-Breeder Occurs Non-Breeder
 CHAPTER 1: DEFINING HABITATS 9

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Black Scoter Occurs Non-Breeder Occurs Non-Breeder

Long-tailed Duck (Oldsquaw) Occurs Non-Breeder Occurs Non-Breeder
Bufflehead Occurs Breeds Occurs Breeds
Common Goldeneye Occurs Non-Breeder Occurs Breeds
Barrow’s Goldeneye Occurs Breeds Occurs Breeds
Smew Accidental Non-Breeder Accidental Non-Breeder
Hooded Merganser Occurs Breeds Occurs Breeds
Common Merganser Occurs Breeds Occurs Breeds
Red-breasted Merganser Occurs Non-Breeder Occurs Non-Breeder
Ruddy Duck Occurs Breeds Occurs Breeds
Osprey Occurs Breeds Occurs Breeds
White-tailed Kite Occurs Breeds Occurs Breeds
Bald Eagle Occurs Breeds Occurs Breeds
Northern Harrier Occurs Breeds Occurs Breeds
Sharp-shinned Hawk Occurs Breeds Occurs Breeds
Cooper’s Hawk Occurs Breeds Occurs Breeds
Northern Goshawk Occurs Breeds Occurs Breeds
Red-shouldered Hawk Occurs Breeds Accidental Non-Breeder
Broad-winged Hawk Accidental Non-Breeder Accidental Non-Breeder
Swainson’s Hawk Occurs Breeds Occurs Breeds
Red-tailed Hawk Occurs Breeds Occurs Breeds
Ferruginous Hawk Occurs Breeds Occurs Breeds
Rough-legged Hawk Occurs Non-Breeder Occurs Non-Breeder
Golden Eagle Occurs Breeds Occurs Breeds
American Kestrel Occurs Breeds Occurs Breeds
Merlin Occurs Bred-Historically Occurs Breeds
Gyrfalcon Occurs Non-Breeder Occurs Non-Breeder
Peregrine Falcon Occurs Breeds Occurs Breeds
Prairie Falcon Occurs Breeds Occurs Breeds
Chukar Non-native Breeds Non-native Breeds
Gray Partridge Non-native Breeds Non-native Breeds
Ring-necked Pheasant Non-native Breeds Non-native Breeds
Ruffed Grouse Occurs Breeds Occurs Breeds
Greater Sage-grouse Occurs Breeds Occurs Breeds
Spruce Grouse Occurs Breeds Occurs Breeds
White-tailed Ptarmigan Does Not Occur Not Applicable Occurs Breeds
Blue Grouse Occurs Breeds Occurs Breeds
Sharp-tailed Grouse Reintroduced Breeds Occurs Breeds
Wild Turkey Non-native Breeds Non-native Breeds
Mountain Quail Occurs Breeds Occurs Breeds
Scaled Quail Does Not Occur Not Applicable Non-native Non-Breeder
California Quail Occurs Breeds Non-native Breeds
Northern Bobwhite Non-native Breeds Non-native Breeds
Yellow Rail Occurs Breeds Accidental Non-Breeder
Virginia Rail Occurs Breeds Occurs Breeds
Sora Occurs Breeds Occurs Breeds
Common Moorhen Accidental Non-Breeder Does Not Occur Not Applicable
American Coot Occurs Breeds Occurs Breeds
Sandhill Crane Occurs Breeds Occurs Breeds
Black-bellied Plover Occurs Non-Breeder Occurs Non-Breeder
American Golden-Plover Occurs Non-Breeder Occurs Non-Breeder
Pacific Golden-Plover Occurs Non-Breeder Occurs Non-Breeder
Mongolian Plover Accidental Non-Breeder Does Not Occur Not Applicable
Snowy Plover Occurs Breeds Occurs Breeds
Semipalmated Plover Occurs Non-Breeder Occurs Non-Breeder
Piping Plover Accidental Non-Breeder Accidental Non-Breeder
Killdeer Occurs Breeds Occurs Breeds
Mountain Plover Accidental Non-Breeder Accidental Non-Breeder
 10 Wildlife–Habitat Relationships in Oregon and Washington

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Eurasian Dotterel Does Not Occur Not Applicable Accidental Non-Breeder

Black Oystercatcher Occurs Breeds Occurs Breeds
Black-necked Stilt Occurs Breeds Occurs Breeds
American Avocet Occurs Breeds Occurs Breeds
Greater Yellowlegs Occurs Non-Breeder Occurs Non-Breeder
Lesser Yellowlegs Occurs Non-Breeder Occurs Non-Breeder
Spotted Redshank Accidental Non-Breeder Does Not Occur Not Applicable
Solitary Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Willet Occurs Breeds Occurs Non-Breeder
Wandering Tattler Occurs Non-Breeder Occurs Non-Breeder
Gray-tailed Tattler Does Not Occur Not Applicable Accidental Non-Breeder
Spotted Sandpiper Occurs Breeds Occurs Breeds
Upland Sandpiper Occurs Breeds Extirpated Bred Historically
Whimbrel Occurs Non-Breeder Occurs Non-Breeder
Bristle-thighed Curlew Accidental Non-Breeder Accidental Non-Breeder
Long-billed Curlew Occurs Breeds Occurs Breeds
Hudsonian Godwit Accidental Non-Breeder Accidental Non-Breeder
Bar-tailed Godwit Accidental Non-Breeder Accidental Non-Breeder
Marbled Godwit Occurs Non-Breeder Occurs Non-Breeder
Ruddy Turnstone Occurs Non-Breeder Occurs Non-Breeder
Black Turnstone Occurs Non-Breeder Occurs Non-Breeder
Surfbird Occurs Non-Breeder Occurs Non-Breeder
Great Knot Accidental Non-Breeder Accidental Non-Breeder
Red Knot Occurs Non-Breeder Occurs Non-Breeder
Sanderling Occurs Non-Breeder Occurs Non-Breeder
Semipalmated Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Western Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Red-necked Stint Accidental Non-Breeder Does Not Occur Not Applicable
Little Stint Accidental Non-Breeder Does Not Occur Not Applicable
Long-toed Stint Accidental Non-Breeder Does Not Occur Not Applicable
Least Sandpiper Occurs Non-Breeder Occurs Non-Breeder
White-rumped Sandpiper Does Not Occur Not Applicable Accidental Non-Breeder
Baird’s Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Pectoral Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Sharp-tailed Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Rock Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Dunlin Occurs Non-Breeder Occurs Non-Breeder
Curlew Sandpiper Accidental Non-Breeder Accidental Non-Breeder
Stilt Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Buff-breasted Sandpiper Occurs Non-Breeder Occurs Non-Breeder
Ruff Occurs Non-Breeder Occurs Non-Breeder
Short-billed Dowitcher Occurs Non-Breeder Occurs Non-Breeder
Long-billed Dowitcher Occurs Non-Breeder Occurs Non-Breeder
Common Snipe Occurs Breeds Occurs Breeds
Wilson’s Phalarope Occurs Breeds Occurs Breeds
Red-necked Phalarope Occurs Non-Breeder Occurs Non-Breeder
Red Phalarope Occurs Non-Breeder Occurs Non-Breeder
South Polar Skua Occurs Non-Breeder Occurs Non-Breeder
Pomarine Jaeger Occurs Non-Breeder Occurs Non-Breeder
Parasitic Jaeger Occurs Non-Breeder Occurs Non-Breeder
Long-tailed Jaeger Occurs Non-Breeder Occurs Non-Breeder
Laughing Gull Accidental Non-Breeder Accidental Non-Breeder
Franklin’s Gull Occurs Breeds Occurs Non-Breeder
Little Gull Accidental Non-Breeder Accidental Non-Breeder
Black-headed Gull Accidental Non-Breeder Accidental Non-Breeder
Bonaparte’s Gull Occurs Non-Breeder Occurs Non-Breeder
Heermann’s Gull Occurs Non-Breeder Occurs Non-Breeder
 CHAPTER 1: DEFINING HABITATS 11

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Mew Gull Occurs Non-Breeder Occurs Non-Breeder

Ring-billed Gull Occurs Breeds Occurs Breeds
California Gull Occurs Breeds Occurs Breeds
Herring Gull Occurs Non-Breeder Occurs Non-Breeder
Thayer’s Gull Occurs Non-Breeder Occurs Non-Breeder
Iceland Gull Does Not Occur Not Applicable Accidental Non-Breeder
Slaty-backed Gull Accidental Non-Breeder Accidental Non-Breeder
Western Gull Occurs Breeds Occurs Breeds
Glaucous-winged Gull Occurs Breeds Occurs Breeds
Glaucous Gull Occurs Non-Breeder Occurs Non-Breeder
Sabine’s Gull Occurs Non-Breeder Occurs Non-Breeder
Black-legged Kittiwake Occurs Non-Breeder Occurs Non-Breeder
Red-legged Kittiwake Accidental Non-Breeder Accidental Non-Breeder
Ross’s Gull Accidental Non-Breeder Accidental Non-Breeder
Ivory Gull Does Not Occur Not Applicable Accidental Non-Breeder
Caspian Tern Occurs Breeds Occurs Breeds
Elegant Tern Occurs Non-Breeder Occurs Non-Breeder
Common Tern Occurs Non-Breeder Occurs Non-Breeder
Arctic Tern Occurs Non-Breeder Occurs Breeds
Forster’s Tern Occurs Breeds Occurs Breeds
Least Tern Accidental Non-Breeder Accidental Non-Breeder
Black Tern Occurs Breeds Occurs Breeds
Common Murre Occurs Breeds Occurs Breeds
Thick-billed Murre Accidental Non-Breeder Accidental Non-Breeder
Pigeon Guillemot Occurs Breeds Occurs Breeds
Long-billed Murrelet Accidental Non-Breeder Accidental Non-Breeder
Marbled Murrelet Occurs Breeds Occurs Breeds
Kittlitz’s Murrelet Does Not Occur Not Applicable Accidental Non-Breeder
Xantus’s Murrelet Accidental Non-Breeder Accidental Non-Breeder
Ancient Murrelet Occurs Non-Breeder Occurs Non-Breeder
Cassin’s Auklet Occurs Breeds Occurs Breeds
Parakeet Auklet Accidental Non-Breeder Accidental Non-Breeder
Rhinoceros Auklet Occurs Breeds Occurs Breeds
Horned Puffin Accidental Non-Breeder Accidental Non-Breeder
Tufted Puffin Occurs Breeds Occurs Breeds
Rock Dove Non-native Breeds Non-native Breeds
Band-tailed Pigeon Occurs Breeds Occurs Breeds
White-winged Dove Accidental Non-Breeder Accidental Non-Breeder
Mourning Dove Occurs Breeds Occurs Breeds
Black-billed Cuckoo Does Not Occur Not Applicable Accidental Non-Breeder
Yellow-billed Cuckoo Occurs Bred Historically Extirpated Bred Historically
Barn Owl Occurs Breeds Occurs Breeds
Flammulated Owl Occurs Breeds Occurs Breeds
Western Screech-owl Occurs Breeds Occurs Breeds
Great Horned Owl Occurs Breeds Occurs Breeds
Snowy Owl Occurs Non-Breeder Occurs Non-Breeder
Northern Hawk Owl Accidental Non-Breeder Accidental Non-Breeder
Northern Pygmy-owl Occurs Breeds Occurs Breeds
Burrowing Owl Occurs Breeds Occurs Breeds
Spotted Owl Occurs Breeds Occurs Breeds
Barred Owl Occurs Breeds Occurs Breeds
Great Gray Owl Occurs Breeds Occurs Breeds
Long-eared Owl Occurs Breeds Occurs Breeds
Short-eared Owl Occurs Breeds Occurs Breeds
Boreal Owl Occurs Breeds Occurs Breeds
Northern Saw-whet Owl Occurs Breeds Occurs Breeds
Common Nighthawk Occurs Breeds Occurs Breeds
Common Poorwill Occurs Breeds Occurs Breeds
 12 Wildlife–Habitat Relationships in Oregon and Washington

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Black Swift Occurs Breeds Occurs Breeds

Vaux’s Swift Occurs Breeds Occurs Breeds
White-throated Swift Occurs Breeds Occurs Breeds
Black-chinned Hummingbird Occurs Breeds Occurs Breeds
Anna’s Hummingbird Occurs Breeds Occurs Breeds
Costa’s Hummingbird Accidental Non-Breeder Does Not Occur Not Applicable
Calliope Hummingbird Occurs Breeds Occurs Breeds
Broad-tailed Hummingbird Occurs Breeds Does Not Occur Not Applicable
Rufous Hummingbird Occurs Breeds Occurs Breeds
Allen’s Hummingbird Occurs Breeds Accidental Non-Breeder
Belted Kingfisher Occurs Breeds Occurs Breeds
Lewis’s Woodpecker Occurs Breeds Occurs Breeds
Acorn Woodpecker Occurs Breeds Occurs Non-Breeder
Williamson’s Sapsucker Occurs Breeds Occurs Breeds
Yellow-bellied Sapsucker Accidental Non-Breeder Accidental Non-Breeder
Red-naped Sapsucker Occurs Breeds Occurs Breeds
Red-breasted Sapsucker Occurs Breeds Occurs Breeds
Nuttall’s Woodpecker Accidental Non-Breeder Does Not Occur Not Applicable
Downy Woodpecker Occurs Breeds Occurs Breeds
Hairy Woodpecker Occurs Breeds Occurs Breeds
White-headed Woodpecker Occurs Breeds Occurs Breeds
Three-toed Woodpecker Occurs Breeds Occurs Breeds
Black-backed Woodpecker Occurs Breeds Occurs Breeds
Northern Flicker Occurs Breeds Occurs Breeds
Pileated Woodpecker Occurs Breeds Occurs Breeds
Olive-sided Flycatcher Occurs Breeds Occurs Breeds
Western Wood-pewee Occurs Breeds Occurs Breeds
Eastern Wood-pewee Accidental Non-Breeder Does Not Occur Not Applicable
Willow Flycatcher Occurs Breeds Occurs Breeds
Least Flycatcher Occurs Non-Breeder Occurs Breeds
Hammond’s Flycatcher Occurs Breeds Occurs Breeds
Gray Flycatcher Occurs Breeds Occurs Breeds
Dusky Flycatcher Occurs Breeds Occurs Breeds
Pacific-slope Flycatcher Occurs Breeds Occurs Breeds
Cordilleran Flycatcher Occurs Breeds Occurs Breeds
Black Phoebe Occurs Breeds Accidental Non-Breeder
Eastern Phoebe Accidental Non-Breeder Accidental Non-Breeder
Say’s Phoebe Occurs Breeds Occurs Breeds
Vermilion Flycatcher Accidental Non-Breeder Accidental Non-Breeder
Ash-throated Flycatcher Occurs Breeds Occurs Breeds
Tropical Kingbird Accidental Non-Breeder Accidental Non-Breeder
Western Kingbird Occurs Breeds Occurs Breeds
Eastern Kingbird Occurs Breeds Occurs Breeds
Scissor-tailed Flycatcher Accidental Non-Breeder Accidental Non-Breeder
Fork-tailed Flycatcher Does Not Occur Not Applicable Accidental Non-Breeder
Loggerhead Shrike Occurs Breeds Occurs Breeds
Northern Shrike Occurs Non-Breeder Occurs Non-Breeder
White-eyed Vireo Does Not Occur Not Applicable Accidental Non-Breeder
Bell’s Vireo Accidental Non-Breeder Does Not Occur Not Applicable
Yellow-throated Vireo Does Not Occur Not Applicable Accidental Non-Breeder
Plumbeous Vireo Occurs Non-Breeder Does Not Occur Not Applicable
Cassin’s Vireo Occurs Breeds Occurs Breeds
Hutton’s Vireo Occurs Breeds Occurs Breeds
Warbling Vireo Occurs Breeds Occurs Breeds
Philadelphia Vireo Does Not Occur Not Applicable Accidental Non-Breeder
Red-eyed Vireo Occurs Breeds Occurs Breeds
Gray Jay Occurs Breeds Occurs Breeds
Steller’s Jay Occurs Breeds Occurs Breeds
Blue Jay Accidental Non-Breeder Accidental Non-Breeder
 CHAPTER 1: DEFINING HABITATS 13

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Western Scrub-Jay Occurs Breeds Occurs Breeds

Pinyon Jay Occurs Breeds Accidental Non-Breeder
Clark’s Nutcracker Occurs Breeds Occurs Breeds
Black-billed Magpie Occurs Breeds Occurs Breeds
American Crow Occurs Breeds Occurs Breeds
Northwestern Crow Occurs Non-Breeder Occurs Breeds
Common Raven Occurs Breeds Occurs Breeds
Sky Lark Does Not Occur Not Applicable Non-native Breeds
Horned Lark Occurs Breeds Occurs Breeds
Purple Martin Occurs Breeds Occurs Breeds
Tree Swallow Occurs Breeds Occurs Breeds
Violet-green Swallow Occurs Breeds Occurs Breeds
Northern Rough-winged Swallow Occurs Breeds Occurs Breeds
Bank Swallow Occurs Breeds Occurs Breeds
Cliff Swallow Occurs Breeds Occurs Breeds
Barn Swallow Occurs Breeds Occurs Breeds
Black-capped Chickadee Occurs Breeds Occurs Breeds
Mountain Chickadee Occurs Breeds Occurs Breeds
Chestnut-backed Chickadee Occurs Breeds Occurs Breeds
Boreal Chickadee Does Not Occur Not Applicable Occurs Breeds
Oak Titmouse Occurs Breeds Does Not Occur Not Applicable
Juniper Titmouse Occurs Breeds Does Not Occur Not Applicable
Bushtit Occurs Breeds Occurs Breeds
Red-breasted Nuthatch Occurs Breeds Occurs Breeds
White-breasted Nuthatch Occurs Breeds Occurs Breeds
Pygmy Nuthatch Occurs Breeds Occurs Breeds
Brown Creeper Occurs Breeds Occurs Breeds
Rock Wren Occurs Breeds Occurs Breeds
Canyon Wren Occurs Breeds Occurs Breeds
Bewick’s Wren Occurs Breeds Occurs Breeds
House Wren Occurs Breeds Occurs Breeds
Winter Wren Occurs Breeds Occurs Breeds
Marsh Wren Occurs Breeds Occurs Breeds
American Dipper Occurs Breeds Occurs Breeds
Golden-crowned Kinglet Occurs Breeds Occurs Breeds
Ruby-crowned Kinglet Occurs Breeds Occurs Breeds
Blue-gray Gnatcatcher Occurs Breeds Accidental Non-Breeder
Northern Wheatear Accidental Non-Breeder Does Not Occur Not Applicable
Western Bluebird Occurs Breeds Occurs Breeds
Mountain Bluebird Occurs Breeds Occurs Breeds
Townsend’s Solitaire Occurs Breeds Occurs Breeds
Veery Occurs Breeds Occurs Breeds
Gray-cheeked Thrush Accidental Non-Breeder Accidental Non-Breeder
Swainson’s Thrush Occurs Breeds Occurs Breeds
Hermit Thrush Occurs Breeds Occurs Breeds
Wood Thrush Accidental Non-Breeder Does Not Occur Not Applicable
American Robin Occurs Breeds Occurs Breeds
Varied Thrush Occurs Breeds Occurs Breeds
Wrentit Occurs Breeds Does Not Occur Not Applicable
Gray Catbird Occurs Breeds Occurs Breeds
Northern Mockingbird Occurs Non-Breeder Occurs Breeds
Sage Thrasher Occurs Breeds Occurs Breeds
Brown Thrasher Accidental Non-Breeder Accidental Non-Breeder
California Thrasher Accidental Non-Breeder Does Not Occur Not Applicable
European Starling Non-native Breeds Non-native Breeds
Siberian Accentor Does Not Occur Not Applicable Accidental Non-Breeder
Yellow Wagtail Does Not Occur Not Applicable Accidental Non-Breeder
White Wagtail Does Not Occur Not Applicable Accidental Non-Breeder
 14 Wildlife–Habitat Relationships in Oregon and Washington

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Black-backed Wagtail Accidental Non-Breeder Accidental Non-Breeder

Red-throated Pipit Does Not Occur Not Applicable Accidental Non-Breeder
American Pipit Occurs Breeds Occurs Breeds
Bohemian Waxwing Occurs Non-Breeder Occurs Non-Breeder
Cedar Waxwing Occurs Breeds Occurs Breeds
Phainopepla Accidental Non-Breeder Accidental Non-Breeder
Blue-winged Warbler Accidental Non-Breeder Accidental Non-Breeder
Golden-winged Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Tennessee Warbler Occurs Non-Breeder Accidental Non-Breeder
Orange-crowned Warbler Occurs Breeds Occurs Breeds
Nashville Warbler Occurs Breeds Occurs Breeds
Virginia’s Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Lucy’s Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Northern Parula Accidental Non-Breeder Accidental Non-Breeder
Yellow Warbler Occurs Breeds Occurs Breeds
Chestnut-sided Warbler Accidental Non-Breeder Accidental Non-Breeder
Magnolia Warbler Accidental Non-Breeder Accidental Non-Breeder
Cape May Warbler Accidental Non-Breeder Accidental Non-Breeder
Black-throated Blue Warbler Occurs Non-Breeder Accidental Non-Breeder
Yellow-rumped Warbler Occurs Breeds Occurs Breeds
Black-throated Gray Warbler Occurs Breeds Occurs Breeds
Black-throated Green Warbler Accidental Non-Breeder Accidental Non-Breeder
Townsend’s Warbler Occurs Breeds Occurs Breeds
Hermit Warbler Occurs Breeds Occurs Breeds
Blackburnian Warbler Accidental Non-Breeder Accidental Non-Breeder
Yellow-throated Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Pine Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Prairie Warbler Accidental Non-Breeder Accidental Non-Breeder
Palm Warbler Occurs Non-Breeder Occurs Non-Breeder
Bay-breasted Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Blackpoll Warbler Accidental Non-Breeder Accidental Non-Breeder
Black-and-white Warbler Accidental Non-Breeder Accidental Non-Breeder
American Redstart Occurs Breeds Occurs Breeds
Prothonotary Warbler Accidental Non-Breeder Accidental Non-Breeder
Worm-eating Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Ovenbird Accidental Non-Breeder Accidental Non-Breeder
Northern Waterthrush Occurs Breeds Occurs Breeds
Kentucky Warbler Accidental Non-Breeder Accidental Non-Breeder
Mourning Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Macgillivray’s Warbler Occurs Breeds Occurs Breeds
Common Yellowthroat Occurs Breeds Occurs Breeds
Hooded Warbler Accidental Non-Breeder Accidental Non-Breeder
Wilson’s Warbler Occurs Breeds Occurs Breeds
Canada Warbler Accidental Non-Breeder Does Not Occur Not Applicable
Yellow-breasted Chat Occurs Breeds Occurs Breeds
Summer Tanager Accidental Non-Breeder Does Not Occur Not Applicable
Scarlet Tanager Accidental Non-Breeder Does Not Occur Not Applicable
Western Tanager Occurs Breeds Occurs Breeds
Green-tailed Towhee Occurs Breeds Occurs Breeds
Spotted Towhee Occurs Breeds Occurs Breeds
California Towhee Occurs Breeds Does Not Occur Not Applicable
American Tree Sparrow Occurs Non-Breeder Occurs Non-Breeder
Chipping Sparrow Occurs Breeds Occurs Breeds
Clay-colored Sparrow Occurs Non-Breeder Occurs Breeds
Brewer’s Sparrow Occurs Breeds Occurs Breeds
Black-chinned Sparrow Accidental Non-Breeder Does Not Occur Not Applicable
Vesper Sparrow Occurs Breeds Occurs Breeds
Lark Sparrow Occurs Breeds Occurs Breeds
 CHAPTER 1: DEFINING HABITATS 15

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Black-throated Sparrow Occurs Breeds Occurs Breeds

Sage Sparrow Occurs Breeds Occurs Breeds
Lark Bunting Accidental Non-Breeder Accidental Non-Breeder
Savannah Sparrow Occurs Breeds Occurs Breeds
Grasshopper Sparrow Occurs Breeds Occurs Breeds
LeConte’s Sparrow Accidental Non-Breeder Accidental Non-Breeder
Nelson’s Sharp-tailed Sparrow Does Not Occur Not Applicable Accidental Non-Breeder
Fox Sparrow Occurs Breeds Occurs Breeds
Song Sparrow Occurs Breeds Occurs Breeds
Lincoln’s Sparrow Occurs Breeds Occurs Breeds
Swamp Sparrow Occurs Non-Breeder Occurs Non-Breeder
White-throated Sparrow Occurs Non-Breeder Occurs Non-Breeder
Harris’s Sparrow Occurs Non-Breeder Occurs Non-Breeder
White-crowned Sparrow Occurs Breeds Occurs Breeds
Golden-crowned Sparrow Occurs Non-Breeder Occurs Non-Breeder
Dark-eyed Junco Occurs Breeds Occurs Breeds
McCown’s Longspur Accidental Non-Breeder Does Not Occur Not Applicable
Lapland Longspur Occurs Non-Breeder Occurs Non-Breeder
Chestnut-collared Longspur Accidental Non-Breeder Accidental Non-Breeder
Rustic Bunting Accidental Non-Breeder Accidental Non-Breeder
Snow Bunting Occurs Non-Breeder Occurs Non-Breeder
Mckay’s Bunting Accidental Non-Breeder Accidental Non-Breeder
Rose-breasted Grosbeak Accidental Non-Breeder Accidental Non-Breeder
Black-headed Grosbeak Occurs Breeds Occurs Breeds
Blue Grosbeak Accidental Non-Breeder Does Not Occur Not Applicable
Lazuli Bunting Occurs Breeds Occurs Breeds
Indigo Bunting Accidental Non-Breeder Accidental Non-Breeder
Painted Bunting Accidental Non-Breeder Does Not Occur Not Applicable
Dickcissel Accidental Non-Breeder Accidental Non-Breeder
Bobolink Occurs Breeds Occurs Breeds
Red-winged Blackbird Occurs Breeds Occurs Breeds
Tricolored Blackbird Occurs Breeds Does Not Occur Not Applicable
Western Meadowlark Occurs Breeds Occurs Breeds
Yellow-headed Blackbird Occurs Breeds Occurs Breeds
Rusty Blackbird Accidental Non-Breeder Accidental Non-Breeder
Brewer’s Blackbird Occurs Breeds Occurs Breeds
Common Grackle Accidental Non-Breeder Accidental Non-Breeder
Great-tailed Grackle Accidental Non-Breeder Accidental Non-Breeder
Brown-headed Cowbird Occurs Breeds Occurs Breeds
Orchard Oriole Accidental Non-Breeder Accidental Non-Breeder
Hooded Oriole Accidental Non-Breeder Accidental Non-Breeder
Streak-backed Oriole Accidental Non-Breeder Does Not Occur Not Applicable
Baltimore Oriole Accidental Non-Breeder Accidental Non-Breeder
Bullock’s Oriole Occurs Breeds Occurs Breeds
Scott’s Oriole Accidental Non-Breeder Accidental Non-Breeder
Brambling Accidental Non-Breeder Accidental Non-Breeder
Gray-crowned Rosy-Finch Occurs Breeds Occurs Breeds
Black Rosy-finch Occurs Breeds Does Not Occur Not Applicable
Pine Grosbeak Occurs Breeds Occurs Breeds
Purple Finch Occurs Breeds Occurs Breeds
Cassin’s Finch Occurs Breeds Occurs Breeds
House Finch Occurs Breeds Occurs Breeds
Red Crossbill Occurs Breeds Occurs Breeds
White-winged Crossbill Occurs Non-Breeder Occurs Breeds
Common Redpoll Occurs Non-Breeder Occurs Non-Breeder
Hoary Redpoll Accidental Non-Breeder Accidental Non-Breeder
Pine Siskin Occurs Breeds Occurs Breeds
Lesser Goldfinch Occurs Breeds Occurs Breeds
Lawrence’s Goldfinch Accidental Non-Breeder Does Not Occur Not Applicable
 16 Wildlife–Habitat Relationships in Oregon and Washington

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

American Goldfinch Occurs Breeds Occurs Breeds

Evening Grosbeak Occurs Breeds Occurs Breeds
House Sparrow Non-native Breeds Non-native Breeds
Mammals
Virginia Opossum Non-native Breeds Non-native Breeds
Masked Shrew Does Not Occur Not Applicable Occurs Breeds
Preble’s Shrew Occurs Breeds Occurs Breeds
Vagrant Shrew Occurs Breeds Occurs Breeds
Montane Shrew Occurs Breeds Occurs Breeds
Baird’s Shrew Occurs Breeds Does Not Occur Not Applicable
Fog Shrew Occurs Breeds Does Not Occur Not Applicable
Pacific Shrew Occurs Breeds Does Not Occur Not Applicable
Water Shrew Occurs Breeds Occurs Breeds
Pacific Water Shrew Occurs Breeds Occurs Breeds
Trowbridge’s Shrew Occurs Breeds Occurs Breeds
Merriam’s Shrew Occurs Breeds Occurs Breeds
Pygmy Shrew Does Not Occur Not Applicable Occurs Breeds
Shrew-mole Occurs Breeds Occurs Breeds
Townsend’s Mole Occurs Breeds Occurs Breeds
Coast Mole Occurs Breeds Occurs Breeds
Broad-footed Mole Occurs Breeds Does Not Occur Not Applicable
California Myotis Occurs Breeds Occurs Breeds
Western Small-footed Myotis Occurs Breeds Occurs Breeds
Yuma Myotis Occurs Breeds Occurs Breeds
Little Brown Myotis Occurs Breeds Occurs Breeds
Long-legged Myotis Occurs Breeds Occurs Breeds
Fringed Myotis Occurs Breeds Occurs Breeds
Keen’s Myotis Does Not Occur Not Applicable Occurs Breeds
Long-eared Myotis Occurs Breeds Occurs Breeds
Silver-haired Bat Occurs Breeds Occurs Breeds
Western Pipistrelle Occurs Breeds Occurs Breeds
Big Brown Bat Occurs Breeds Occurs Breeds
Hoary Bat Occurs Non-Breeder Occurs Non-Breeder
Spotted Bat Accidental Non-Breeder Occurs Breeds
Townsend’s Big-eared Bat Occurs Breeds Occurs Breeds
Pallid Bat Occurs Breeds Occurs Breeds
Brazilian Free-tailed Bat Occurs Breeds Does Not Occur Not Applicable
American Pika Occurs Breeds Occurs Breeds
Pygmy Rabbit Occurs Breeds Occurs Breeds
Brush Rabbit Occurs Breeds Does Not Occur Not Applicable
Eastern Cottontail Non-native Breeds Non-native Breeds
Nuttall’s (Mountain) Cottontail Occurs Breeds Occurs Breeds
European Rabbit Does Not Occur Not Applicable Non-native Breeds
Snowshoe Hare Occurs Breeds Occurs Breeds
White-tailed Jackrabbit Occurs Breeds Occurs Breeds
Black-tailed Jackrabbit Occurs Breeds Occurs Breeds
Mountain Beaver Occurs Breeds Occurs Breeds
Least Chipmunk Occurs Breeds Occurs Breeds
Yellow-pine Chipmunk Occurs Breeds Occurs Breeds
Townsend’s Chipmunk Occurs Breeds Occurs Breeds
Allen’s Chipmunk Occurs Breeds Does Not Occur Not Applicable
Siskiyou Chipmunk Occurs Breeds Does Not Occur Not Applicable
Red-tailed Chipmunk Does Not Occur Not Applicable Occurs Breeds
Yellow-bellied Marmot Occurs Breeds Occurs Breeds
Hoary Marmot Does Not Occur Not Applicable Occurs Breeds
Olympic Marmot Does Not Occur Not Applicable Occurs Breeds
 CHAPTER 1: DEFINING HABITATS 17

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

White-tailed Antelope Squirrel Occurs Breeds Does Not Occur Not Applicable
Townsend’s Ground Squirrel Occurs Breeds Occurs Breeds
Merriam’s Ground Squirrel Occurs Breeds Does Not Occur Not Applicable
Piute Ground Squirrel Occurs Breeds Does Not Occur Not Applicable
Washington Ground Squirrel Occurs Breeds Occurs Breeds
Wyoming Ground Squirrel Extirpated Bred Historically Does Not Occur Not Applicable
Belding’s Ground Squirrel Occurs Breeds Does Not Occur Not Applicable
Columbian Ground Squirrel Occurs Breeds Occurs Breeds
California Ground Squirrel Occurs Breeds Occurs Breeds
Golden-mantled Ground Squirrel Occurs Breeds Occurs Breeds
Cascade Golden-mantled Ground Squirrel Does Not Occur Not Applicable Occur Breeds
Eastern Gray Squirrel Non-native Breeds Non-native Breeds
Eastern Fox Squirrel Non-native Breeds Non-native Breeds
Western Gray Squirrel Occurs Breeds Occurs Breeds
Red Squirrel Occurs Breeds Occurs Breeds
Douglas’ Squirrel Occurs Breeds Occurs Breeds
Northern Flying Squirrel Occurs Breeds Occurs Breeds
Northern Pocket Gopher Occurs Breeds Occurs Breeds
Western Pocket Gopher Occurs Breeds Occurs Breeds
Camas Pocket Gopher Occurs Breeds Does Not Occur Not Applicable
Botta’s (Pistol Riv.) Pocket Gopher Occurs Breeds Does Not Occur Not Applicable
Townsend’s Pocket Gopher Occurs Breeds Does Not Occur Not Applicable
Great Basin Pocket Mouse Occurs Breeds Occurs Breeds
Little Pocket Mouse Occurs Breeds Does Not Occur Not Applicable
Dark Kangaroo Mouse Occurs Breeds Does Not Occur Not Applicable
Ord’s Kangaroo Rat Occurs Breeds Occurs Breeds
Chisel-toothed Kangaroo Rat Occurs Breeds Does Not Occur Not Applicable
California Kangaroo Rat Occurs Breeds Does Not Occur Not Applicable
American Beaver Occurs Breeds Occurs Breeds
Western Harvest Mouse Occurs Breeds Occurs Breeds
Deer Mouse Occurs Breeds Occurs Breeds
Columbian Mouse Does Not Occur Not Applicable Occurs Breeds
Canyon Mouse Occurs Breeds Does Not Occur Not Applicable
Pinon Mouse Occurs Breeds Does Not Occur Not Applicable
Northern Grasshopper Mouse Occurs Breeds Occurs Breeds
Desert Woodrat Occurs Breeds Does Not Occur Not Applicable
Dusky-footed Woodrat Occurs Breeds Does Not Occur Not Applicable
Bushy-tailed Woodrat Occurs Breeds Occurs Breeds
Southern Red-backed Vole Occurs Breeds Occurs Breeds
Western Red-backed Vole Occurs Breeds Does Not Occur Not Applicable
Heather Vole Occurs Breeds Occurs Breeds
White-footed Vole Occurs Breeds Does Not Occur Not Applicable
Red Tree Vole Occurs Breeds Does Not Occur Not Applicable
Meadow Vole Does Not Occur Not Applicable Occurs Breeds
Montane Vole Occurs Breeds Occurs Breeds
Gray-tailed Vole Occurs Breeds Occurs Breeds
California Vole Occurs Breeds Does Not Occur Not Applicable
Townsend’s Vole Occurs Breeds Occurs Breeds
Long-tailed Vole Occurs Breeds Occurs Breeds
Creeping Vole Occurs Breeds Occurs Breeds
Water Vole Occurs Breeds Occurs Breeds
Sagebrush Vole Occurs Breeds Occurs Breeds
Muskrat Occurs Breeds Occurs Breeds
Northern Bog Lemming Does Not Occur Not Applicable Occurs Breeds
Black Rat Non-native Breeds Non-native Breeds
Norway Rat Non-native Breeds Non-native Breeds
House Mouse Non-native Breeds Non-native Breeds
Western Jumping Mouse Occurs Breeds Occurs Breeds
Pacific Jumping Mouse Occurs Breeds Occurs Breeds
Common Porcupine Occurs Breeds Occurs Breeds
 18 Wildlife–Habitat Relationships in Oregon and Washington

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

Nutria Non-native Breeds Non-native Breeds

Coyote Occurs Breeds Occurs Breeds
Gray Wolf Extirpated Bred Historically Occurs Breeds
Red Fox Occurs Breeds Occurs Breeds
Kit Fox Occurs Breeds Does Not Occur Not Applicable
Gray Fox Occurs Breeds Does Not Occur Not Applicable
Black Bear Occurs Breeds Occurs Breeds
Grizzly Bear Extirpated Bred Historically Occurs Breeds
Ringtail Occurs Breeds Does Not Occur Not Applicable
Raccoon Occurs Breeds Occurs Breeds
American Marten Occurs Breeds Occurs Breeds
Fisher Occurs Breeds Occurs Breeds
Ermine Occurs Breeds Occurs Breeds
Long-tailed Weasel Occurs Breeds Occurs Breeds
Mink Occurs Breeds Occurs Breeds
Wolverine Occurs Breeds Occurs Breeds
American Badger Occurs Breeds Occurs Breeds
Western Spotted Skunk Occurs Breeds Occurs Breeds
Striped Skunk Occurs Breeds Occurs Breeds
Northern River Otter Occurs Breeds Occurs Breeds
Mountain Lion Occurs Breeds Occurs Breeds
Lynx Occurs Breeds Occurs Breeds
Bobcat Occurs Breeds Occurs Breeds
Wild Burro Non-native Breeds Does Not Occur Not Applicable
Feral Horse Non-native Breeds Non-native Breeds
Feral Pig Non-native Breeds Non-native Non-Breeder
Roosevelt Elk Occurs Breeds Occurs Breeds
Rocky Mountain Elk Occurs Breeds Occurs Breeds
Black-tailed Deer Occurs Breeds Occurs Breeds
Mule Deer Occurs Breeds Occurs Breeds
Columbian White-tailed Deer Occurs Breeds Occurs Breeds
White-tailed Deer (Eastside) Occurs Breeds Occurs Breeds
Moose Accidental Non-Breeder Occurs Breeds
Mountain Caribou Does Not Occur Not Applicable Occurs Breeds
Pronghorn Antelope Occurs Breeds Extirpated Bred Historically
Bison Extirpated Bred Historically Extirpated Bred Historically*
Mountain Goat Reintroduced Breeds Occurs Breeds
Rocky Mountain Bighorn Sheep Reintroduced Breeds Reintroduced Breeds
California Bighorn Sheep Reintroduced Breeds Reintroduced Breeds
Northern Fur Seal Occurs Non-Breeder Occurs Non-Breeder
Northern (Steller) Sea Lion Occurs Breeds Occurs Non-Breeder
California Sea Lion Occurs Non-Breeder Occurs Non-Breeder
Harbor Seal Occurs Breeds Occurs Breeds
Northern Elephant Seal Occurs Breeds Occurs Non-Breeder
Sea Otter Extirpated Bred Historically Reintroduced Breeds
Gray Whale Occurs Non-Breeder Occurs Non-Breeder
Minke Whale Occurs Breeds Occurs Breeds
Sei Whale Accidental Non-Breeder Accidental Non-Breeder
Blue Whale Occurs Non-Breeder Occurs Non-Breeder
Fin Whale Accidental Non-Breeder Accidental Non-Breeder
Humpback Whale Occurs Non-Breeder Occurs Non-Breeder
Northern Right Whale Accidental Non-Breeder Accidental Non-Breeder
Striped Dolphin Accidental Non-Breeder Accidental Non-Breeder
Common Saddle-backed Dolphin Accidental Non-Breeder Accidental Non-Breeder
 CHAPTER 1: DEFINING HABITATS 19

Common Name Oregon Washington

Occurrence Breeding Status Occurrence Breeding Status

False Killer Whale Accidental Non-Breeder Accidental Non-Breeder

Short-finned Pilot Whale Occurs Non-Breeder Occurs Non-Breeder
Killer Whale Occurs Breeds Occurs Breeds
Northern Right-whale Dolphin Occurs Non-Breeder Occurs Non-Breeder
Harbor Porpoise Occurs Breeds Occurs Breeds
Dall’s Porpoise Occurs Breeds Occurs Breeds
North Pacific Bottle-nosed Whale Occurs Non-Breeder Occurs Non-Breeder
Goose-beaked Whale Accidental Non-Breeder Accidental Non-Breeder
Bering Sea Beaked Whale Accidental Non-Breeder Accidental Non-Breeder
Arch-beaked Whale Accidental Non-Breeder Accidental Non-Breeder
Pygmy Sperm Whale Accidental Non-Breeder Accidental Non-Breeder
Sperm Whale Occurs Non-Breeder Occurs Non-Breeder
Beluga Whale Does Not Occur Not Applicable Accidental Non-Breeder
*Historical Breeding Unknown.

Table 2. The 32 wildlife-habitat types as determined from the cluster analysis procedure using 541 native
breeding species and 119 Pacific Northwest vegetation, land use, and marine groupings.

Wildlife-HabitatTypes Wildlife-HabitatTypes
Vegetative/Land Use/Marine Groupings Vegetative/Land Use/Marine Groupings

Westside Lowland Conifer-Hardwood Forest Ponderosa Pine Forest and Woodlands

Alnus rubra-Acer macrophyllum Upland Forests Pinus ponderosa Woodlands
Picea sitchensis-Tsuga heterophylla Forests Eastside Pinus ponderosa-Quercus garryana Forest and
Pseudotsuga menziesii-Alnus rubra-Acer macrophyllus Forests Woodlands
Maritime Tsuga heterophylla-Thuja plicata Forests Eastside Quercus garryana Woodlands
Forested Dunes
Upland Aspen Forest
Westside Oak and Dry Douglas-fir Forest and Populus tremuloides Upland Forests
Woodlands
Subalpine Parklands
Westside Quercus garryana Forests and Woodlands
Subapline and Alpine Wetlands
Westside Quercus garryana-Pseudotsuga menziesii Forests
Pinus albicaulis-Abies lasiocarpa Woodlands and Parklands
Westside Dry Pseudotsuga menziesii Forests
Tsuga mertensiana Parklands
Pseudotsuga menziesii-Arbutus menziesii Forests
Alpine Grasslands and Shrublands
Southwest Oregon Mixed Conifer-Hardwood Forest
Subalpine and Alpine Grasslands
Abies concolor Mixed Conifer Forests
Alpine Dwarf Shrublands-Fellfields and Sedge Turf
Pinus jeffreii Woodlands
Pseudotsuga menziesii-Lithocarpus densiflorus Forests Westside Grasslands
Southwest Oregon Low Elevation Mixed Conifer Forests Westside Festuca idahoensis var. romeri-Danthonia californica
Montane Mixed Conifer Forest Ceanothus-Manzanita Shrublands
Abies amabilis-Tsuga heterophylla Forests Chaparral
Abies lasiocarpa-Picea englemannii Forests
Abies magnifica var. shastensis Forests and Woodlands Western Juniper and Mountain Mahogany Woodlands
Tsuga mertensiana Forests Juniperus occidentalis Scablands
Tsuga mertensiana-Abies amabilis Forests Juniperus occidentalis-Artemisia tridentata Tall Shrublands
Juniperus occidentalis/Bunchgrass
Eastside (Interior) Mixed Conifer Forest Cercocarpus ledifolius
Eastside Abies grandis-Pseudotsuga menziesii Forests
Eastside Pseudotsuga menziesii-Pinus ponderosa Forests Eastside (Interior) Canyon Shrublands
Eastside Tsuga heterophylla-Thuja plicata Forests Eastside Moist Deciduous Shrublands

Lodgepole Pine Forest and Woodlands Eastside (Interior) Grasslands

Pinus contorta Grass understory Pseudoroegneria spicata Grasslands
Pinus contorta Shrub understory Eastside Low-to-Mid-elevation Festuca idahoensis Grasslands
Pinus contorta Subalpine Forests Eastside Modified Grasslands
Pinus contorta Woodlands and Forests on Pumice Sporobolus cryptandrus-Aristida puppurea var. longiseta
Grasslands
 20 Wildlife–Habitat Relationships in Oregon and Washington

Wildlife-HabitatTypes Wildlife-HabitatTypes
Vegetative/Land Use/Marine Groupings Vegetative/Land Use/Marine Groupings

Shrub-steppe Pinus ponderosa Riparian Woodlands

Artemisia tripartita Shrub-steppe Populus tremuloides Riparian/Wetland Forests and Woodlands
Artemisia cana Shrub-steppe
Artemisia tridentata ssp. tridentata and ssp.wyomingensis Shrub- Coastal Dunes and Beaches
steppe Coastal Dune Grasslands
Artemisia tridentata ssp. vaseyana Shrublands Coastal Dune Shrublands
Purshia tridentata Shrub-steppe Coastal Headlands and Islets
Sandy steppe and Shrub-steppe Coastal Headland Shrublands and Grasslands
Dwarf Shrub-steppe Bays and Estuaries*
Artemisia rigida/Eriogonum spp./Poa secunda Dwarf-Shrub Bays and Estuaries (includes Intertidal Marshes)
Scabland
Artemisia arbuscula Dwarf-Shrub-steppe Inland Marine Deeper Waters*
Artemisia nova Dwarf-Shrublands Puget Sound to Strait of Juan de Fuca

Desert Playa and Salt Scrub Marine Nearshore*

Alkali Grasslands and Wetlands Marine environment from shore line to 20 m depth
Atriplex confertifolia Shrublands Marine Shelf*
Mixed Saltdesert Shrub-Non-Playa Marine environment from 20 m to 200 m depth
Mixed Saltdesert Shrub-Playa
Sarcobatus vermiculatus Shrublands Oceanic*
Marine environment greater than 200 m depth
Agriculture, Pasture, and Mixed Environs*
Cultivated Croplands *Wildlife-habitats that were determined by an expert panel
Improved Pasture process rather than through the cluster analysis effort.
Modified Grasslands
Orchard/Vineyard/Nursery
Unimproved Pasture
Urban and Mixed Environs*
High Density
Moderate Density
Low Density
Open Water-Lakes, Rivers, Streams
Riverine
Lacustrine -Open Water
Herbaceous Wetlands
Graminoid Wet Meadow
Freshwater Aquatic Beds
Herbaceous and Sedge Wetlands
Westside Riparian - Wetlands
Alnus viridis ssp. sinuata-Acer circinatum Shrublands
Westside Riparian and Wetland Deciduous Forests
Picea sitchensis Wetland Forests and Woodlands
Tsuga heterophylla-Thuja plicata coniferous wetlands
Westside Riparian/Wetland Shrublands
Shrub/herbaceous Sphagnum Bogs
Wooded Bogs
Montane Coniferous Wetlands
Westside Montane Coniferous Wetlands
Picea engelmannii Forested Wetlands
Eastside (Interior) Riparian - Wetlands
Eastside Midmontane Alnus incana-Salix spp. Riparian
Shrublands
Eastside Lowland Riparian Shrublands
Eastside Populus balsamifera spp. trichocarpa
Alnus rhombifolia Riparian
 CHAPTER 1: DEFINING HABITATS 21

Table 3. Number of wildlife species by habitat and by their association.

Wildlife-HabitatTypes Number of Species*

Total CloselyAssociated GenerallyAssociated Present Unsure

Westside Lowland Conifer-Deciduous Forest 233 48 155 29 1

Westside Oak and Dry Douglas-fir Forest and
Woodlands 229 34 148 40 7
Southwest Oregon Mixed Conifer-Deciduous Forest 236 35 163 33 5
Montane Mixed Conifer Forest 221 35 138 38 10
Eastside (Interior) Mixed Conifer Forest 225 38 146 35 6
Lodgepole Pine Forest and Woodlands 166 15 95 48 8
Ponderosa Pine Forest and Woodlands 247 26 160 46 15
Upland Aspen Forest 144 4 85 42 13
Subalpine Parkland 184 19 92 69 4
Alpine Grasslands and Shrublands 132 19 35 71 7
Westside Grasslands 157 12 105 39 1
Ceanothus-Manzanita Shrublands 138 4 89 39 6
Western Juniper and Mountain Mahogany Woodlands 173 17 113 36 7
Eastside (Interior) Canyon Shrubland 147 13 84 38 12
Eastside (Interior) Grasslands 174 33 100 38 3
Shrub-steppe 184 47 100 34 3
Dwarf Shrub-steppe 149 23 86 32 8
Desert Playa and Salt Scrub 143 27 78 33 5
Agriculture and Pastures Mixed Environs 346 68 174 99 5
Urban and Mixed Environs 266 18 144 102 2
Open Water - Lakes, Rivers, Streams 162 96 53 13 0
Herbaceous Wetlands 228 105 90 31 2
Westside Riparian/Wetlands 256 74 145 35 2
Montane Coniferous Wetlands 148 17 101 28 2
Eastside (Interior) Riparian/Wetlands 271 81 149 36 5
Coastal Dunes and Beaches 155 31 79 41 4
Coastal Headlands and Islets 118 28 63 24 3
Bays and Estuaries 175 66 74 34 1
Inland Marine Deeper Waters 61 26 20 15 0
Marine Nearshore 86 38 33 14 1
Marine Shelf 72 41 20 11 0
Oceanic 44 31 8 5 0
* Historical associations are not shown.
 2
Wildlife Habitats: Descriptions, Status,
Trends, and System Dynamics
Christopher B. Chappell, Rex C. Crawford, Charley Barrett,
Jimmy Kagan, David H. Johnson, Mikell O’Mealy,
Greg A. Green, Howard L. Ferguson,W. Daniel Edge,
Eva L. Greda, & Thomas A. O’Neil

Introduction
In the previous chapter, the authors described the process (horizontal dimension) of vegetation; layering (vertical
by which the 32 wildlife-habitat types of Oregon and dimension) of vegetation; dominant growth forms, leaf
Washington (Table 1, Frontispiece) were defined. In this phenologies (evergreen or deciduous), leaf characters
chapter, we offer detailed descriptions of each wildlife- (conifer or broadleaf), and vegetation persistence (annual
habitat type to support a common understanding for their or perennial) represented in different structural layers; and
delineation, inventory, and management across the region. significant structural components of dead and decaying
Each wildlife habitat below is described as to its vegetation. Growth forms include trees, shrubs (>1.6 ft
geographic distribution, physical setting, landscape [0.5 m] tall), dwarf-shrubs (<1.6 ft [0.5 m] tall), graminoids
setting, structure, and composition. Additionally, we (grasses, sedges, rushes), forbs, ferns, mosses, lichens, and
include other information that might help managers, algae. Vegetation cover categories frequently referred to
researchers, and others gain further insight into each include forest (>60% cover of trees), woodland (25-60%
habitat, such as listing other classification systems and key cover of trees), shrubland (>25% cover of shrubs),
references, natural disturbance regimes, succession and dwarf-shrubland (>25% cover of dwarf-shrubs), and
stand dynamics, effects of management and grassland (graminoids dominant). Water-dominated
anthropogenic impacts, and status and trends. We have habitats (e.g., marine and open water) may be described
also included photographs of each wildlife-habitat type in terms of the physical aspects of the water column and
to give the reader an idea of what each habitat type looks the bottom substrate of the habitat.
like. Multiple photographs are offered for most habitats
Composition. Describes the species composition of the
in order to depict some of the variability that exists within
vegetation that creates structure. Composition is described
each type. Each of the habitats below is numbered (Table
as dominant, co-dominant (shares dominance with •1
1); the descriptions in this chapter reflect the same
species), or important indicator species by structural layer.
numbering sequence.
English names for all vertebrates are used in the text and
The following are definitions of each category used to
corresponding standard names are in Appendix I. The
characterize the wildlife-habitat types:
geographic distribution or physical setting is noted for
Geographic Distribution. Describes the broad geographic those dominant species that occur only in particular
range within which the habitat is located, both within physical settings or specific geographic areas of the overall
Oregon and Washington and elsewhere. Major variations habitat’s range of occurrence.
in dominance type are noted either here or under
Other Classifications and Key References. Notes other
Composition.
names that have been applied to this habitat by other
Physical Setting. Describes physical features of the classifications or major summary publications, and
environment on sites where the habitat is found in Oregon important references that describe the habitat or parts of
and Washington. These typically include climate, the habitat in greater detail.
elevation, soils, hydrology, geology, and topography.
Natural Disturbance Regime. Describes the major natural
Landscape Setting. Describes the landscape pattern and disturbances that are important in the habitat. The regime
distribution of the habitat in relation to other habitats. includes the disturbance type, severity, frequency, extent,
Primary land use is also noted. and range of variation in these characteristics.
Structure. Describes the physical structure of the habitat, Succession and Stand Dynamics. Describes the way in
both its typical aspect and the range of variation in which structure and composition change over time in
structure present within the habitat. Aspects of physical relation to natural disturbances.
structure include some description of cover or density

22
 CHAPTER 2: HABITAT DESCRIPTIONS 23

Table 1. The 32 wildlife habitats and their total acreage in Oregon and Washington.
The marine waters extend out to the 200-mile Exclusive Economic Zone.

Wildlife Habitat Oregon Washington Page Number

Total Acreage Total Acreage

1. Westside Lowlands Conifer-Hardwood Forest 9,349,756 9,064,128 24

2. Westside Oak and Dry Douglas-fir Forest and Woodlands 433,132 425,038 26
3. Southwest Oregon Mixed Conifer-Hardwood Forest 4,020,320 Does Not Occur 28
4. Montane Mixed Conifer Forest 2,949,586 4,653,306 30
5. Eastside (Interior) Mixed Conifer Forest 4,126,957 4,662,101 31
6. Lodgepole Pine Forest and Woodlands 532,587 119,201 33
7. Ponderosa Pine Forest and Woodlands (includes Eastside Oak) 6,226,351 1,927,176 35
8. Upland Aspen Forest 19,685 100,621 36
9. Subalpine Parkland 84,240 327,442 37
10. Alpine Grassland and Shrublands 291,494 1,591,115 39
11. Westside Grasslands 1331 22,491 40
12. Ceanothus-Manzanita Shrublands 52,104 Does Not Occur 42
13.Western Juniper and Mountain Mahogany Woodlands 4,037,221 Does Not Occur 43
14. Eastside (Interior) Canyon Shrublands 358,250 Not Mapped3 44
15. Eastside (Interior) Grasslands 1,935,794 1,002,076 46
16. Shrub-steppe 17,420,753 7,144,6973 47
17. Dwarf Shrub-steppe 514,066 Not Mapped3 49
18. Desert Playa and Salt Scrub Shrublands 719,503 Not Mapped3 50
19. Agriculture, Pasture and Mixed Environs 6,197,887 9,251,107 52
20. Urban and Mixed Environs 575,087 1,204,680 53
21. Open Water - Lakes, Rivers, Streams 780,901 761,360 56
22. Herbaceous Wetlands 1,031,343 210,451 58
23. Westside Riparian-Wetlands 168,872 347,653 90
24. Montane Coniferous Wetlands 56,099 241,450 92
25. Eastside (Interior) Riparian-Wetlands 31,121 100,763 93
26. Coastal Dunes and Beaches 52,451 Not Mapped2 95
27. Coastal Headlands and Islets 9,137 7,776 96
28. Bays and Estuaries 172,748 226,336 97
29. Inland Marine Deeper Water Does Not Occur 1,855,780 99
30. Marine Nearshore 223,371 750,329 100
31. Marine Shelf 3,905,164 4,780,625 101
32. Oceanic 33,987,189 19,845,660 102

Totals 100,263,303 70, 532,093

1
Because of difficulty in classifying this type using remote sensing (i.e., discerning native grasslands from pasture lands) native westside
grasslands have inadvertently been classified within the agriculture habitat type. Nonetheless, there are few areas known to be native
westside grasslands in Oregon.
2
This type was not part of the vegetation classification when the Washington Gap Project mapped the state of Washington. Thus, no
wildlife habitat area was determined.
3
In Washington, Eastside Canyon Shrublands, Dwarf Shrub-steppe, and Desert Playa and Salt Scrub Shrublands were mapped as part
of Shrub-steppe for the Washington GAP Project. Thus, no wildlife habitat area was determined.

Effects of Management and Anthropogenic Impacts. Status and Trends. Describes the general extent of the type
Describes typical changes in structure and composition in Oregon and Washington, its current ecological
observed after typical management activities (human condition, and historical and current trends in extent and
disturbances) and widespread changes in the habitat that condition. Ecological condition refers primarily to how
have occurred since Euro-American settlement. similar the current structure, composition, and disturbance
Disturbances addressed include land uses that do not regime is to natural or presettlement conditions. The total
necessarily convert the habitat to urban or agriculture, but number of plant associations recognized in the habitat and
have a significant influence on structure or composition, the number of those that are considered globally imperiled
e.g., hydrologic alterations, logging, and grazing. Exotic provide some idea of the degree of loss, degradation, and
species that have become abundant in the habitat are threat that is associated with the habitat.
noted.
 24 Wildlife–Habitat Relationships in Oregon and Washington

1. Westside Lowlands Conifer- large logs on the ground. Early seral stands typically have
smaller trees, single-storied canopies, and may be
Hardwood Forest dominated by conifers, broadleaf trees, or both. Coarse
Christopher B. Chappell & Jimmy Kagan woody debris is abundant in early seral stands after
natural disturbances but much less so after clearcutting.
Forest understories are structurally diverse: evergreen
shrubs tend to dominate on nutrient-poor or drier sites;
deciduous shrubs, ferns, and/or forbs tend to dominate
on relatively nutrient-rich or moist sites. Shrubs may be
low (1.6 ft [0.5 m] tall), medium-tall (3.3-6.6 ft [1-2 m]), or
tall (6.6-13.1 ft [2-4 m]). Almost all structural stages are
represented in the successional sequence within this
habitat. Mosses are often a major ground cover. Lichens
are abundant in the canopy of old stands.
Composition. Western hemlock (Tsuga heterophylla) and
Douglas-fir (Pseudotsuga menziesii) are the most
characteristic species and one or both are typically present.
Most stands are dominated by one or more of the
following: Douglas-fir, western hemlock, western redcedar
Geographic Distribution. This forest habitat occurs (Thuja plicata), Sitka spruce (Picea sitchensis), red alder
throughout low-elevation western Washington, except on (Alnus rubra), or bigleaf maple (Acer macrophyllum). Trees
extremely dry or wet sites. In Oregon it occurs on the of local importance that may be dominant include Port-
western slopes of the Cascades, around the margins of Orford cedar (Chamaecyparis lawsoniana) in the south, shore
the Willamette Valley, in the Coast Range, and along the pine (Pinus contorta var. contorta) on stabilized dunes, and
outer coast. The global distribution extends from grand fir (Abies grandis) in drier climates. Western white
southeastern Alaska south to southwestern Oregon. pine (Pinus monticola) is frequent but subordinate in
Physical Setting. Climate is relatively mild and moist to importance through portions of this habitat. Pacific silver
wet. Mean annual precipitation is mostly 35-100 inches fir (Abies amabilis) is largely absent except on the wettest
(90-254 cm), but can vary locally. Snowfall ranges from portion of the western Olympic Peninsula, where it is
rare to regular, but is transitory. Summers are relatively common and sometimes co-dominant. Common small
dry. Summer fog is a major factor on the outer coast in the subcanopy trees are cascara buckthorn (Rhamnus
Sitka spruce zone. Elevation ranges from sea level to a purshiana) in more moist climates and Pacific yew (Taxus
maximum of about 2,000 ft (610 m) in much of northern brevifolia) in somewhat drier climates or sites.
Washington and 3,500 ft (1,067 m) in central Oregon. Soils Sitka spruce is found as a major species only in the outer
and geology are very diverse. Topography ranges from coastal area at low elevations where summer fog is a
relatively flat glacial till plains to steep mountainous significant factor. Bigleaf maple is most abundant in the
terrain. Puget Lowland, around the Willamette Valley, and in the
central Oregon Cascades, but occurs elsewhere also.
Landscape Setting. This is the most extensive habitat in Douglas-fir is absent to uncommon as a native species in
the lowlands on the westside of the Cascades, except in the very wet maritime outer coastal area of Washington,
southwestern Oregon, and forms the matrix within which including the coastal plain on the west side of the Olympic
other habitats occur as patches, especially Westside Peninsula. However, it has been extensively planted in
Riparian-Wetlands and less commonly Herbaceous that area. Port-Orford cedar occurs only in southern
Wetlands or Open Water. It also occurs adjacent to or in a Oregon. Paper birch (Betula papyrifera) occurs as a co-
mosaic with Urban and Mixed Environs (hereafter Urban) dominant only in Whatcom County, Washington. Grand
or Agriculture, Pasture and Mixed Environs (hereafter fir occurs as an occasional co-dominant only in the Puget
Agriculture) habitats. In the driest areas, it occurs adjacent Lowland and Willamette Valley.
to or in a mosaic with Westside Oak and Dry Douglas-fir Dominant or co-dominant understory shrub species of
Forest and Woodlands. Bordering this habitat at upper more than local importance include salal (Gaultheria
elevations is Montane Mixed Conifer Forest. Along the shallon), dwarf Oregongrape (Mahonia nervosa), vine maple
coastline, it often occurs adjacent to Coastal Dunes and (Acer circinatum), Pacific rhododendron (Rhododendron
Beaches. In southwestern Oregon, it may border macrophyllum), salmonberry (Rubus spectabilis), trailing
Southwest Oregon Mixed Conifer-Hardwood Forest. The blackberry (R. ursinus), red elderberry (Sambucus racemosa),
primary land use for this habitat is forestry. fools huckleberry (Menziesia ferruginea), beargrass
Structure. This habitat is forest, or rarely woodland, (Xerophyllum tenax), oval-leaf huckleberry (Vaccinium
dominated by evergreen conifers, deciduous broadleaf ovalifolium), evergreen huckleberry (V. ovatum), and red
trees, or both. Late seral stands typically have an huckleberry (V. parvifolium). Salal, rhododendron, and
abundance of large (>164 ft [50 m] tall) coniferous trees, a beargrass are particularly associated with low nutrient or
multi-layered canopy structure, large snags, and many dry sites.
 CHAPTER 2: HABITAT DESCRIPTIONS 25

Swordfern (Polystichum munitum) is the most common not establish early, the site may remain as a shrubland for
herbaceous species and is often dominant on nitrogen- an indeterminate period. Early seral tree species can be
rich or moist sites. Other forbs and ferns that frequently any of the potential dominants for the habitat, depending
dominate the understory are Oregon oxalis (Oxalis on environment, type of disturbance, and seed source. All
oregana), deerfern (Blechnum spicant), bracken fern of these species except the short-lived red alder are capable
(Pteridium aquilinum), vanillaleaf (Achlys triphylla), of persisting for at least a few hundred years. Douglas-fir
twinflower (Linnaea borealis), false lily-of-the-valley is the most common dominant after fire, but is uncommon
(Maianthemum dilatatum), western springbeauty (Claytonia in the wettest zones. It is also the most fire resistant of the
siberica), foamflower (Tiarella trifoliata), inside-out flower trees in this habitat and survives moderate-severity fires
(Vancouveria hexandra), and common whipplea (Whipplea well. After the tree canopy closes, the understory may
modesta). become sparse, corresponding with the stem-exclusion
stage.168 Eventually tree density will decrease and the
Other Classifications and Key References. This habitat
understory will begin to flourish again, typically at stand
includes most of the forests and their successional seres
age 60-100 years. As trees grow larger and a new
within the Tsuga heterophylla and Picea sitchensis zones.88
generation of shade-tolerant understory trees (usually
This habitat is also referred to as Douglas-fir-western
western hemlock, less commonly western redcedar) grows
hemlock and Sitka spruce-western hemlock forests,87
up, a multi-layered canopy will gradually develop and
spruce-cedar-hemlock forest (Picea-Thuja-Tsuga, No. 1) and
be well expressed by stand age 200-400 years.89 Another
cedar-hemlock-Douglas-fir forest (Thuja-Tsuga-
fire is likely to return before the loss of shade-intolerant
Pseudotsuga, No. 2).136 The Oregon Gap II Project126 and
Douglas-fir from the canopy at stand age 800-1,000 years,
Oregon Vegetation Landscape-Level Cover Types127 would
unless the stand is located in the wet maritime zone.
crosswalk with Sitka spruce-western hemlock maritime
Throughout this habitat, western hemlock tends to
forest, Douglas-fir-western hemlock-redcedar forest, red
increase in importance as stand development proceeds.
alder forest, red alder-bigleaf maple forest, mixed conifer/
Coarse woody debris peaks in abundance in the first 50
mixed deciduous forest, south coast mixed-deciduous
years after a fire and is least abundant at about stand age
forest, and coastal lodgepole forest. The Washington Gap
100-200 years.193
Vegetation map includes this vegetation as conifer forest,
mixed hardwood/conifer forest, and hardwood forest in Effects of Management and Anthropogenic Impacts. Red
the Sitka spruce, western hemlock, Olympic Douglas-fir, alder is more successful after typical logging disturbance
Puget Sound Douglas-fir, Cowlitz River and Willamette than after fire alone on moist, nutrient-rich sites, perhaps
Valley zones.37 A number of other references describe because of the species’ ability to establish abundantly on
elements of this habitat.13, 25, 26, 40, 42, 66, 90, 104, 110, 111, 114, 115, 210 scarified soils. 100 Alder is much more common now
because of large-scale logging activities.87 Alder grows
Natural Disturbance Regime. Fire is the major natural
more quickly in height early in succession than the
disturbance in all but the wettest climatic area (Sitka spruce
conifers, thereby prompting many forest managers to
zone), where wind becomes the major source of natural
apply herbicides for alder control. If alder is allowed to
disturbance. Natural fire-return intervals generally range
grow and dominate early successional stands, it will
from about 100 years or less in the driest areas to several
decline in importance after about 70 years and die out
hundred years.1, 115, 160 Mean fire-return interval for the
completely by age 100. Often there are suppressed conifers
western hemlock zone as a whole is 250 years, but may
in the subcanopy that potentially can respond to the death
vary greatly. Major natural fires are associated with
of the alder canopy. However, salmonberry sometimes
occasional extreme weather conditions.1 Fires are typically
forms a dense shrub layer under the alder, which can
high-severity, with few trees surviving. However, low- and
exclude conifer regeneration.88 Salmonberry responds
moderate-severity fires that leave partial to complete live
positively to soil disturbance, such as that associated with
canopies are not uncommon, especially in drier climatic
logging.19 Bigleaf maple sprouts readily after logging and
areas. Occasional major windstorms hit outer coastal
is therefore well adapted to increase after disturbance as
forests most intensely, where fires are rare. Severity of
well. Clearcut logging and plantation forestry have
wind disturbance varies greatly, with minor events being
resulted in less diverse tree canopies, and have focused
extremely frequent and major events occurring once every
mainly on Douglas-fir, with reductions in coarse woody
few decades. Bark beetles and fungi are significant causes
debris over natural levels, a shortened stand initiation
of mortality that typically operate on a small scale.
phase, and succession truncated well before late-seral
Landslides are another natural disturbance that occur in
characteristics are expressed. Douglas-fir has been almost
some areas.
universally planted, even in wet coastal areas of
Succession and Stand Dynamics. After a severe fire or Washington, where it is rare in natural stands.
blowdown, a typical stand will be briefly occupied by
Status and Trends. Extremely large areas of this habitat
annual and perennial ruderal forbs and grasses as well as
remain. Some loss has occurred, primarily to development
predisturbance understory shrubs and herbs that
in the Puget Lowland. Condition of what remains has been
resprout.102 Herbaceous species generally give way to
degraded by industrial forest practices at both the stand
dominance by shrubs or a mixture of shrubs and young
and landscape scale. Most of the habitat is probably now
trees within a few years. If shrubs are dense and trees did
 26 Wildlife–Habitat Relationships in Oregon and Washington

in Douglas-fir plantations. Only a fraction of the original in the Olympic Mountains, but is mainly below 1,500 ft
old-growth forest remains, mostly in national forests in (457 m). Topography ranges from nearly level to very steep
the Cascade and Olympic mountains. Areal extent slopes, where aspect tends to be southern or western. Soils
continues to be reduced gradually, especially in the Puget on dry sites are typically shallow over bedrock, very stony,
Lowland. An increase in alternative silviculture practices or very deep and excessively drained. Willamette Valley
may be improving structural and species diversity in some soils are typically much older and have more moderate
areas. However, intensive logging of natural-origin mature drainage and water availability. Parent materials include
and young stands and even small areas of old growth various types of bedrock, shallow or very coarse glacial
continues. Of the 62 plant associations representing this till, alluvium, and glacial outwash.
habitat listed in the National Vegetation Classification, 27
Landscape Setting. This habitat is found in a mosaic with,
% are globally imperiled or critically imperiled.10
or adjacent to, Westside Grasslands, Westside Lowlands
Conifer-Hardwood Forest, Westside Riparian-Wetlands,
2. Westside Oak and Dry Douglas-fir Southwest Oregon Mixed Conifer-Hardwood Forest,
Forest and Woodlands Urban, and Agriculture. Inclusions of Open Water or
Christopher B. Chappell & Jimmy Kagan Herbaceous Wetlands sometimes occur. In the Puget
Lowland, this habitat is sometimes found adjacent to Puget
Sound (Nearshore Marine). Land use of this habitat
includes forestry (generally small scale), livestock grazing,
and low-density rural residential.
Structure. This is a forest or woodland dominated by
evergreen conifers, deciduous broadleaf trees, evergreen
broadleaf trees, or some mixture of conifers and broadleaf
trees. Canopy structure varies from single- to multi-
storied. Large conifers, when present, typically emerge
above broadleaf trees in mixed canopy stands. Large snags
and logs are less abundant than in other westside forest
habitats, but can be prominent, especially in unlogged old
stands. Understories vary in structure: grasses, shrubs,
ferns, or some combination will typically dominate.
Deciduous broadleaf shrubs are perhaps most typical as
understory dominants in the existing landscape. Early
Geographic Distribution. This habitat is primarily found successional stand structure varies depending on
in the Willamette Valley, Puget Lowlands, and Klamath understory species present and if initiated following
Mountains ecoregions. In the Puget Lowlands, it is logging or fire.
common in and around the San Juan Islands and in parts
of Thurston, Pierce, and Mason counties. In southwestern Composition. The canopy is typically dominated by one
Oregon, it is now restricted mainly to the valleys of the or more of the following species: Douglas-fir (Pseudotsuga
Rogue and Umpqua rivers. Minor occurrences can also menziesii), Oregon white oak (Quercus garryana), Pacific
be found in the northeastern Olympic Mountains and madrone (Arbutus menziesii), shore (lodgepole) pine (Pinus
western Cascades. contorta var. contorta), or California black oak (Q. kelloggii).
This habitat is comprised of several geographic Ponderosa pine (Pinus ponderosa) is important in
variants: California black oak and ponderosa pine are southwestern Oregon and the southern Willamette Valley
important only in southwestern Oregon and the southern as a subordinate or co-dominant with oak. Grand fir (Abies
Willamette Valley. The latter is also found in a small area grandis) is occasionally co-dominant with Douglas-fir in
of Pierce County, Washington. Shore (lodgepole) pine is the northern Puget Lowland or in the Willamette Valley.
only important in the Puget Lowland, mainly in San Juan Oregon ash (Fraxinus latifolia) is occasionally co-dominant
and Mason counties. Dry Douglas-fir forests (without oak with white oak in riparian oak stands. Several other tree
or madrone) are mainly in the Puget Lowland and rarely species may be present, but western hemlock (Tsuga
in the Olympic Mountains, west Cascades, and Willamette heterophylla) and western redcedar (Thuja plicata) generally
Valley. Pacific madrone and Douglas-fir/Pacific madrone cannot regenerate successfully because of dry conditions.
stands without oak are limited to the Puget Lowland and This lack of shade-tolerant tree regeneration, along with
the southern Willamette Valley foothills. Mixed oak- understory indicators like tall Oregongrape (Mahonia
madrone stands occur primarily in Oregon, especially aquifolium), and blue wildrye (Elymus glaucus), help
southwestern Oregon. distinguish dry Douglas-fir forests from mid-seral
Douglas-fir stands on more mesic sites, which are part of
Physical Setting. This habitat typically occupies dry sites the Westside Lowlands Conifer-Hardwood Forest. Tree
west of the Cascades. Annual mean precipitation ranges regeneration, when present, is typically Douglas-fir, less
from 17 to 60 inches (43 to 152 cm), occasionally higher. commonly grand fir. Sweet cherry (Prunus avium) and/or
Elevation ranges from sea level to about 3,500 ft (1,069 m) English hawthorn (Crataegus monogyna) have invaded and
 CHAPTER 2: HABITAT DESCRIPTIONS 27

now dominate a subcanopy layer in many oak forests of Succession and Stand Dynamics. Many of these forests
the Willamette Valley. and woodlands were formerly either grasslands or
Deciduous shrubs that commonly dominate or co- savannas that probably burned frequently, thus preventing
dominate the understory are oceanspray (Holodiscus dominance by trees.41, 54 Some portions of this habitat in
discolor), baldhip rose (Rosa gymnocarpa), poison-oak the central Puget Lowlands may have formerly been
(Toxicodendron diversiloba), serviceberry (Amelanchier dominated by shrubs (salal, beaked hazel, evergreen
alnifolia), beaked hazel (Corylus cornuta), trailing blackberry huckleberry, hairy manzanita [Arctostaphylos columbiana])
(Rubus ursinus), Indian plum (Oemleria cerasiformis), for lengthy periods, probably also because of the particular
snowberries (Symphoricarpos albus and S. mollis), wedge- combination of fire frequency and intensity. Other areas
leaf ceanothus (Ceanothus cuneatus), and oval-leaf were woodlands to semi-open forests that burned
viburnum (Viburnum ellipticum). Evergreen shrubs or vines moderately frequently, as evidenced by the relict stands
that sometimes are dominant where conifers are important of old-growth Douglas-fir. The dominant trees in this
in the canopy include salal (Gaultheria shallon), dwarf habitat establish most abundantly after fire. Moderate-
Oregongrape (Mahonia nervosa), Pacific rhododendron severity fires kill many trees but also leave many alive,
(Rhododendron macrophyllum), hairy honeysuckle (Lonicera creating opportunities for establishment of new cohorts
hispidula), evergreen huckleberry (Vaccinium ovatum), and of trees and increasing structural complexity.1 Oaks and
Piper’s barberry (Mahonia piperiana). madrone resprout after fire if they are top-killed. Without
Native graminoids that commonly dominate or co- periodic fire, most oak-dominated stands will eventually
dominate the understory are western fescue (Festuca convert to Douglas-fir forests.1 Animal dissemination of
occidentalis), Alaska oniongrass (Melica subulata), blue acorns may be important in dispersal of oaks. Shore pine,
wildrye, and long-stolon sedge (Carex inops). Kentucky where present, is an early-seral upper canopy species that
bluegrass (Poa pratensis) is a major non-native dominant grows quickly and dies out after about 100-150 years,
in oak woodland understories. Swordfern (Polystichum yielding to a mature Douglas-fir stand unless another fire
munitum) or, less commonly, bracken fern (Pteridium intervenes before the death of the pine.
aquilinum) sometimes co-dominates the understory,
Effects of Management and Anthropogenic Impacts.
especially on sites that formerly supported grasslands and
Clearcut or similar logging reduces canopy structural
savannas. Forbs, many of which are characteristic of these
complexity and abundance of large woody debris. Dry
dry sites, are often abundant and diverse, but typically
Douglas-fir stands are well suited to alternative
do not dominate. Common camas (Camassia quamash),
silvicultural practices, such as uneven-aged management
cleavers (Galium aparine), or other forbs are occasionally
or maintaining two-storied stands. Oaks and madrone will
co-dominant with graminoids.
typically resprout after logging and thus can increase in
Other Classifications and Key References. This habitat importance relative to conifers in mixed canopy stands.
has been described as oak groves and dry site Douglas-fir Selective logging of Douglas-fir in oak stands can prevent
forest in the Tsuga heterophylla zone of western Washington long-term loss of oak dominance. With fire exclusion,
and northwestern Oregon as well as oak woodland in the stands have probably increased in tree density and grassy
interior valleys of western Oregon.88 It is also referred to understories have been replaced by deciduous shrubs.41
as Oregon oakwoods No. 22 and a minor part of Cedar- Moderate to heavy grazing or other significant ground
hemlock-Douglas-fir forest No. 2;136 The Oregon Gap II disturbance, especially in grassy understories, leads to
Project126 and Oregon Vegetation Landscape-Level Cover increases in non-native invader species, many of which
Types127 that would represent this type are Oregon white are now abundant in stands with grassy or formerly grassy
oak forest and Douglas-fir/white oak forest. The understories. Scot’s broom (Cytisus scoparius) is an exotic
Washington Gap Project represents this habitat as part of shrub particularly invasive and persistent in oak
hardwood forest, mixed hardwood/conifer forest, and woodlands. Exotic herbaceous invaders include colonial
conifer forest in the Woodland/Prairie Mosaic, Puget bentgrass (Agrostis capillaris), common velvetgrass (Holcus
Sound Douglas-fir, and, to a minor degree, Cowlitz River, lanatus), Kentucky bluegrass, tall oatgrass (Arrhenatherum
and Willamette Valley zones of Washington.37 Other elatius), rigid brome (Bromus rigidus), orchardgrass
references also describe elements of this habitat.13, 17, 40, 41, 86, (Dactylis glomerata), hedgehog dogtail (Cynosurus
111, 115, 202, 210
echinatus), tall fescue (Festuca arundinacea), and common
St. Johnswort (Hypericum perforatum).
Natural Disturbance Regime. Fire is the major natural
disturbance in this habitat. In presettlement times, fire Status and Trends. This habitat is relatively limited in area
frequency probably ranged from frequent (every few and is currently declining in extent and condition. With
years) to moderately frequent (once every 50-100 years), the cessation of regular burning 100-130 years ago, many
and reflected low-severity and moderate-severity fire grasslands and savannas were invaded by a greater
regimes.1 Fire frequency has been much lower in the last density of trees and thus converted to a different habitat.
100 years. Windstorms are an occasional disturbance, most Conversely, large areas of this habitat have been converted
important in the San Juan Islands and vicinity. to Urban or Agriculture habitats. Most of what remains
Understories are sometimes browsed heavily by deer in has been considerably degraded by invasion of exotic
the San Juan Islands, thus preventing dominance by species or by logging and consequent loss of structural
deciduous shrubs and favoring grasses and forbs.
 28 Wildlife–Habitat Relationships in Oregon and Washington

diversity. Ongoing threats include residential Grazing occurs on some areas, especially at lower
development, increase and spread of exotic species, and elevations.
fire suppression effects (the latter especially in
Structure. Conifer trees typically dominate this forest or
oak-dominated stands). Thirteen of 27 plant associations
woodland habitat. In some generally more coastal areas,
listed in the National Vegetation Classification are
a well developed subcanopy layer of smaller evergreen
considered globally imperiled or critically imperiled.10
broadleaf trees is present. Occasionally, deciduous
broadleaf trees are co-dominant. Complex multi-layered
3. Southwest Oregon Mixed Conifer- canopies are typical, though single-layered canopies also
Hardwood Forest occur, especially in areas of intensive forest management.
Christopher B. Chappell & Jimmy Kagan Dominant canopy trees vary from 60 to >300 ft (18 to >91
m) tall at maturity. Large woody debris (snags and logs)
is typically common, although variable. Understories are
mostly dominated by shrubs, but can be dominated by
forbs, graminoids, or may be largely depauperate.
Composition. The tree canopy is often diverse. Douglas-
fir (Pseudotsuga menziesii), white fir (Abies concolor), sugar
pine (Pinus lambertiana), ponderosa pine (P. ponderosa), or
incense cedar (Calocedrus decurrens) are typically dominant
or co-dominant. Port-Orford cedar (Chamaecyparis
lawsoniana), tanoak (Lithocarpus densiflorus), canyon live
oak (Quercus chrysolepis), and Pacific madrone (Arbutus
menziesii) are locally important. Jeffrey pine (Pinus jeffreyi)
is dominant on serpentine parent materials in the Siskiyou
Mountains, and to a lesser degree in the southwestern
Cascades.
Douglas-fir is found in almost every area; ponderosa
Geographic Distribution. This upland forest and pine is also found in most stands, although it has been
woodland habitat occurs in southwestern Oregon, declining with fire suppression. White fir, incense cedar,
northwestern California, and the Sierra Nevada. In and sugar pine are common in mixed stands in the
southern Oregon, it is found at low and middle elevations Cascades and central and eastern Siskiyous on all but the
in the Klamath Mountains, Cascades, Coast Range, and driest sites. White fir dominates the canopy in only the
Eastern Cascade Slopes and Foothills ecoregions. Portions moist, cool sites at higher elevations, although it is the
of Curry, Josephine, Jackson, Douglas, Lane, and Klamath major tree regeneration in most areas. Jeffrey pine and
counties are included in the range of this habitat. knobcone pine (Pinus attenuata) are limited primarily to
Physical Setting. The climate varies from relatively dry serpentine soils, which they dominate. Port-Orford cedar
and very warm to moderately moist and cool to slightly dominates some more moist sites near the coast and
warm and very moist. Mean annual precipitation ranges riparian and wetland habitats inland. Brewer’s spruce
from 20 to 140 inches (51 to 356 cm). Snow is uncommon (Picea breweri) is an uncommon dominant at high
except at the highest elevations, where a winter snow pack elevations in the Siskiyous. The broadleaf subcanopy is
occurs for a few months. Summers are hot and dry. most prominent on the western sides of the Coast Range
Elevation ranges from near sea level to 6,000 ft (1,829 m). and Siskiyous, where tanoak is most abundant, with
Topography is mostly mountainous but also includes two Pacific madrone, golden chinquapin (Castanopsis
fairly large valleys, and a corresponding variety of terrain. chrysophylla), or canyon live oak also sometimes
Soils are diverse as is the bedrock geology. Serpentine soils dominating the subcanopy. Coast redwood (Sequoia
are common in portions of the Siskiyou Mountains, where sempervirens) occurs only in a very small area near the coast
they have a major effect on vegetation. in far southern Oregon.
Dominant or co-dominant evergreen shrubs include
Landscape Setting. This habitat is typically bounded at pinemat manzanita (Arctostaphylos nevadensis), green-leaf
its upper elevation limits by Montane Mixed Conifer manzanita (A. patula), white-leaf manzanita (A. viscida),
Forest and at its lower limits, along the coast, by Westside kinnikinnick (A. uva-ursi), Piper’s barberry (Mahonia
Lowlands Conifer-Hardwood Forest. At lower elevations piperiana), dwarf Oregongrape (M. nervosa), tobacco brush
in the Rogue and Umpqua valleys it can be found in a (Ceanothus velutinus), squawcarpet (C. prostratus), salal
mosaic with Westside Oak and Dry Douglas-fir Forest and (Gaultheria shallon), deer oak (Quercus sadleriana),
Woodland, Ceanothus-Manzanita Shrublands, Urban, and huckleberry oak (Q. vacciniifolia), snow bramble (Rubus
Agriculture. Small inclusions of Open Water, Herbaceous nivalis), Pacific rhododendron (Rhododendron
Wetlands, Westside Riparian-Wetlands, and Ceanothus- macrophyllum), and evergreen huckleberry (Vaccinium
Manzanita Shrublands occur scattered throughout this ovatum). Major deciduous shrubs are serviceberry
habitat. The predominant land use is forestry. Low-density (Amelanchier alnifolia), sticky currant (Ribes viscosissimum),
residential is prominent in the Rogue and Umpqua valleys.
 CHAPTER 2: HABITAT DESCRIPTIONS 29

oceanspray (Holodiscus discolor), creeping snowberry and canyon live oak can survive fairly hot fires if the fuels
(Symphoricarpos mollis), baldhip rose (Rosa gymnocarpa), do not extend into the canopy. Conifers are at a
beaked hazel (Corylus cornuta), Rocky Mountain maple disadvantage in regeneration following stand replacement
(Acer glabrum), vine maple (A. circinatum), poison-oak fires because of dependence on local seed-fall. Many
(Toxicodendron diversiloba), big huckleberry (Vaccinium conifers of this habitat are able to survive moderate-
membranaceum), deerbrush (Ceanothus integerrimus), and severity fire well, including, in decreasing order of fire
trailing blackberry (Rubus ursinus). Early seral shrublands, resistance, Ponderosa pine, Jeffrey pine, Douglas-fir, sugar
part of this habitat, can be difficult to distinguish from pine, coast redwood, incense cedar, and Port-Orford cedar.
Ceanothus-Manzanita Shrublands. They are best These species are fairly well represented throughout the
separated by their different species composition, especially successional sequence, unless a high-severity fire was
the predominance in this habitat of Ceanothus velutinus, closely followed by another, in which case the subcanopy
Arctostaphylos patula, and A. nevadensis. broadleaf species are likely to dominate.1 Development
Graminoids that are most prominent are long-stolon of complex multi-layered canopies of conifers and
sedge (Carex inops), Idaho fescue (Festuca idahoensis), and broadleaf evergreens are typical under a moderate-
California fescue (F. californica). Forbs that are indicative severity fire regime.
of site conditions or dominate understories include Where hardwoods are absent and white fir is
common whipplea (Whipplea modesta), twinflower (Linnaea prominent, succession differs from that described above.
borealis), sidebells (Orthilia secunda), rattlesnake plantain Under a low-severity fire regime with frequent fires, white
(Goodyera oblongifolia), vanillaleaf (Achlys triphylla), fir is relatively unimportant and fire-resistant conifers,
beargrass (Xerophyllum tenax), and starry false solomonseal especially Douglas-fir and ponderosa pine, dominate.
(Maianthemum stellata). White fir increases in the absence of fire. 171 With a
moderate-severity fire regime, i.e., less frequent fires, white
Other Classifications and Key References. This habitat
fir can dominate or co-dominate, especially on cooler sites.1
includes the conifer-dominated forests and their
Small gaps created by moderate-severity fires, blowdown,
successional seres within the Interior Valley, Mixed-
or disease afford opportunities for regeneration of less
Conifer, Mixed-Evergreen, and Abies concolor zones of
shade-tolerant tree species, thus maintaining a diverse tree
southwestern Oregon, plus Redwood forests in the Picea
canopy for lengthy periods. Evergreen shrubs, especially
sitchensis Zone. 88 It is also referred to as Klamath
tobacco brush, often dominate after high-severity fire and
Mountains mixed evergreen forests and Sierran-type
may persist as a cover type for decades, especially if they
mixed conifer forests,87 Pseudotsuga menziesii/hardwood
are reburned.51, 88 On the driest, hottest sites in this habitat,
forests and Abies concolor forests,1 Mixed conifer forest No.
white fir does not grow and tree regeneration is limited to
5, Redwood forest No. 6, California mixed evergreen forest
Douglas-fir and ponderosa pine, with the former tending
No. 29, and Montane chaparral No. 34.136 The Oregon Gap
to increase in the absence of fire.
II Project126 and Oregon Vegetation Landscape-Level Cover
Types 127 that would represent this type are the Effects of Management and Anthropogenic Impacts.
southwestern portion of the Douglas-fir dominant-mixed Clearcut logging where hardwoods are present favors
conifer forest, Jeffrey pine forest and woodland, serpentine post-disturbance dominance of tanoak or madrone.
conifer woodland, Douglas-fir-Port Orford cedar forest, Control of this competing vegetation has been a major
Douglas-fir mixed deciduous forest, Douglas-fir-white fir/ focus of timber management in this habitat. Fire control
tanoak-madrone mixed forest, and Siskiyou Mountains over the last 100 years has decreased fire frequencies and
mixed deciduous forest. Other references also describe this altered stand structure through increases in small tree
habitat.13, 15, 17, 111, 117 density and heavy fuels, especially where low-severity fire
regimes were prevalent. As a result, most of these areas
Natural Disturbance Regime. Fire is the predominant
are more susceptible to stand-replacement fires. White fir
natural disturbance. Fire regime varies depending on
has increased dramatically on drier sites where it occurs,
environmental conditions Drier, hotter sites within this
creating dense subcanopy thickets.1, 128 Evergreen shrubs
area have a low-severity fire regime. Cooler and/or
often dominate after clearcut logging and in some cases
moister sites typically have a moderate-severity fire
hinder the establishment of conifers.140 Clearcut logging
regime. Presettlement mean fire return intervals vary from
tends to decrease tree species diversity, coarse woody
10 years to about 80 years.1, 98, 154 Lightning ignitions are
debris loads, and structural diversity. The non-native
more frequent here than anywhere else in the region and
species white pine blister rust (Cronartium ribicola) and
Native Americans probably burned some areas
Phytophthora lateralis, a root rot disease, have had
intentionally.1 Wind is a somewhat important disturbance
significant negative impacts on the abundance of sugar
at higher elevations. Root rot fungi and insects are other
pine and Port-Orford cedar, respectively.
important disturbances in some forests, mostly operating
at small-scales. Status and Trends.. This habitat covers most of
southwestern Oregon and has declined little in areal
Succession and Stand Dynamics. Most evergreen
extent. Conditions of most communities and stands have
broadleaf trees, when present, are top-killed by moderate-
been degraded by forestry practices and by fire
severity fires but resprout vigorously to dominate or co-
suppression. The low-elevation, driest communities have
dominate after most fires.14, 152 Mature Oregon white oak
 30 Wildlife–Habitat Relationships in Oregon and Washington

been altered by grazing and invasion of exotic species. timber values and much of it has been harvested in a
Port-Orford cedar has declined dramatically in extent from dispersed-patch pattern.
logging and Phytophthora lateralis.230 Effects of fire
Structure. This is a forest, or rarely woodland, dominated
suppression and logging-related impacts continue to be
by evergreen conifers. Canopy structure varies from
threats. Twenty-one % of 68 plant associations representing
single- to multi-storied. Tree size also varies from small to
this habitat listed in the National Vegetation Classification
very large. Large snags and logs vary from abundant to
are listed as globally imperiled.10
uncommon. Understories vary in structure: shrubs, forbs,
ferns, graminoids or some combination of these usually
4. Montane Mixed Conifer Forest dominate, but they can be depauperate as well. Deciduous
Christopher B.Chappell broadleaf shrubs are most typical as understory
dominants. Early successional structure after logging or
fire varies depending on understory species present.
Mosses are a major ground cover and epiphytic lichens
are typically abundant in the canopy.
Composition. This forest habitat is recognized by the
dominance or prominence of one of the following species:
Pacific silver fir (Abies amabilis), mountain hemlock (Tsuga
mertensiana), subalpine fir (A. lasiocarpa), Shasta red fir (A.
magnifica var. shastensis), Engelmann spruce (Picea
engelmannii), noble fir (A. procera), or Alaska yellow-cedar
(Chamaecyparis nootkatensis). Several other trees may co-
dominate: Douglas-fir (Pseudotsuga menziesii), lodgepole
pine (Pinus contorta), western hemlock (Tsuga heterophylla),
western redcedar (Thuja plicata), or white fir (A. concolor).
Tree regeneration is typically dominated by Pacific silver
fir in moist westside middle-elevation zones; by mountain
Geographic Distribution. These forests occur in
hemlock, sometimes with silver fir, in cool, very snowy
mountains throughout Washington and Oregon, excepting
zones on the westside and along the Cascade Crest; by
the Basin and Range of southeastern Oregon. These
subalpine fir in cold, drier eastside zones; and by Shasta
include the Cascade Range, Olympic Mountains,
red fir in the snowy mid- to upper-elevation zone of
Okanogan Highlands, Coast Range (rarely), Blue and
southwestern and south-central Oregon.
Wallowa mountains, and Siskiyou Mountains.
Subalpine fir and Engelmann spruce are major species
Physical Setting. This habitat is typified by a moderate to only east of the Cascade Crest in Washington, in the Blue
deep winter snow pack that persists for three to nine and Wallowa mountains, and in the northeastern Olympic
months. The climate is moderately cool and wet to Mountains (spruce is largely absent in the Olympic
moderately dry and very cold. Mean annual precipitation Mountains). Lodgepole pine is important east of the
ranges from about 40 inches (102 cm) to >200 inches (508 Cascade Crest throughout and in central and southern
cm). Elevation is mid- to upper montane, as low as 2,000 Oregon. Douglas-fir is important east of the Cascade Crest
ft (610 m) in northern Washington, to as high as 7,500 ft and at lower elevations on the westside. Pacific silver fir
(2,287 m) in southern Oregon. On the westside, it occupies is a major species on the westside as far south as central
an elevational zone of about 2,500 to 3,000 vertical feet Oregon. Noble fir, as a native species, is found primarily
(762 to 914 m), and on the eastside it occupies a narrower in the western Cascades from central Washington to
zone of about 1,500 vertical feet (457 m). Topography is central Oregon. Mountain hemlock is a common dominant
generally mountainous. Soils are typically not well at higher elevations along the Cascade Crest and to the
developed, but varied in their parent material: glacial till, west. Western hemlock, and to a lesser degree western
volcanic ash, residuum, or colluvium. Spodosols are redcedar, occur as dominants primarily with silver fir at
common. lower elevations on the westside. Alaska yellow-cedar
occurs as a co-dominant west of the Cascade Crest in
Landscape Setting. This habitat is found adjacent to
Washington, rarely in northern Oregon. Shasta red fir and
Westside Lowlands Conifer-Hardwood Forest, Eastside
white fir occur only from central Oregon south, the latter
Mixed Conifer Forest, or Southwest Oregon Mixed
mainly at lower elevations.
Conifer-Hardwood Forest at its lower elevation limits and
Deciduous shrubs that commonly dominate or co-
to Subalpine Parkland at its upper elevation limits.
dominate the understory are oval-leaf huckleberry
Inclusions of Montane Forested Wetlands, Westside
(Vaccinium ovalifolium), big huckleberry (V. membranaceum),
Riparian-Wetlands, and less commonly Open Water or
grouseberry (V. scoparium), dwarf huckleberry (V.
Herbaceous Wetlands occur within the matrix of montane
cespitosum), fools huckleberry (Menziesia ferruginea),
forest habitat. The typical land use is forestry or recreation.
Cascade azalea (Rhododendron albiflorum), copperbush
Most of this type is found on public lands managed for
(Elliottia pyroliflorus), devil’s-club (Oplopanax horridus), and,
 CHAPTER 2: HABITAT DESCRIPTIONS 31

in the far south only, baldhip rose (Rosa gymnocarpa), large areas of mountain hemlock forests in the Oregon
currants (Ribes spp.), and creeping snowberry Cascades.72
(Symphoricarpos mollis). Important evergreen shrubs
Succession and Stand Dynamics. After fire, a typical
include salal (Gaultheria shallon), dwarf Oregongrape
stand will briefly be occupied by annual and perennial
(Mahonia nervosa), Pacific rhododendron (Rhododendron
ruderal forbs and grasses, as well as predisturbance
macrophyllum), deer oak (Quercus sadleriana), pinemat
understory shrubs and herbs that resprout. Stand initiation
manzanita (Arctostaphylos nevadensis), beargrass
can take a long time, especially at higher elevations,
(Xerophyllum tenax), and Oregon boxwood (Paxistima
resulting in shrub/herb dominance (with or without a
myrsinites).
scattered tree layer) for extended periods.3, 109 Early seral
Graminoid dominants are found primarily just along
tree species can be any of the potential dominants for the
the Cascade Crest and to the east and include pinegrass
habitat, or lodgepole pine, depending on the environment,
(Calamagrostis rubescens), Geyer’s sedge (Carex geyeri),
type of disturbance, and seed source. Fires tend to favor
smooth woodrush (Luzula glabrata var. hitchcockii), and
early seral dominance of lodgepole pine, Douglas-fir, noble
long-stolon sedge (Carex inops). Deerfern (Blechnum
fir, or Shasta red fir, if their seeds are present.1 In some
spicant) and western oakfern (Gymnocarpium dryopteris) are
areas, large stand-replacement fires will result in
commonly co-dominant. The most abundant forbs include
conversion of this habitat to the Lodgepole Pine Forest
Oregon oxalis (Oxalis oregana), single-leaf foamflower
and Woodland habitat, distinguished by dominance of
(Tiarella trifoliata var. unifoliata), rosy twisted-stalk
lodgepole. After the tree canopy closes, the understory
(Streptopus roseus), queen’s cup (Clintonia uniflora), western
typically becomes sparse for a time. Eventually tree
bunchberry (Cornus unalaschkensis), twinflower (Linnaea
density will decrease and the understory will begin to
borealis), prince’s pine (Chimaphila umbellata), five-leaved
flourish again, but this process takes longer than in lower
bramble (Rubus pedatus), dwarf bramble (R. lasiococcus),
elevation forests, generally at least 100 years after the
sidebells (Orthilia secunda), avalanche lily (Erythronium
disturbance, sometimes much longer. 1 As stand
montanum), Sitka valerian (Valeriana sitchensis), false lily-
development proceeds, relatively shade-intolerant trees
of-the-valley (Maianthemum dilatatum), and Idaho
(lodgepole pine, Douglas-fir, western hemlock, noble fir,
goldthread (Coptis occidentalis).
Engelmann spruce) typically decrease in importance and
Other Classifications and Key References. This habitat more shade-tolerant species (Pacific silver fir, subalpine
includes most of the upland forests and their successional fir, Shasta red fir, mountain hemlock) increase. Complex
stages, except lodgepole pine dominated forests, in the multi-layered canopies with large trees will typically take
Tsuga mertensiana, Abies amabilis, A. magnifica var. shastensis, at least 300 years to develop, often much longer, and on
A. lasiocarpa zones of Franklin and Dyrness.88 Portions of some sites may never develop. Tree growth rates, and
this habitat have also been referred to as A. amabilis-Tsuga therefore the potential to develop these structural features,
heterophylla forests, A. magnifica var. shastensis forests, and tend to decrease with increasing elevation.
Tsuga mertensiana forests.87 It is equivalent to Silver fir-
Effects of Management and Anthropogenic Impacts.
Douglas-fir forest No. 3, closed portion of Fir-hemlock
Forest management practices, such as clearcutting and
forest No. 4, Red fir forest No. 7, and closed portion of
plantations, have in many cases resulted in less diverse
Western spruce-fir forest No. 15.136 The Oregon Gap II
tree canopies with an emphasis on Douglas-fir. They also
Project126 and Oregon Vegetation Landscape-Level Cover
reduce coarse woody debris compared to natural levels,
Types127 that would represent this type are mountain
and truncate succession well before late-seral
hemlock montane forest, true fir-hemlock montane forest,
characteristics are expressed. Post-harvest regeneration of
montane mixed conifer forest, Shasta red fir-mountain
trees has been a perpetual problem for forest managers in
hemlock forest, and subalpine fir-lodgepole pine montane
much of this habitat.16, 97 Planting of Douglas-fir has often
conifer; also most of the conifer forest in the Silver Fir,
failed at higher elevations, even where old Douglas-fir
Mountain Hemlock, and Subalpine Fir Zones of
were present in the unmanaged stand.115 Slash burning
Washington Gap.37 A number of other references describe
often has negative impacts on productivity and
this habitat.13, 15, 17, 25, 26, 36, 38, 90, 108, 111, 114, 115, 118, 144, 148, 158, 212, 221
regeneration.186 Management has since shifted away from
Natural Disturbance Regime. Fire is the major natural burning and toward planting noble fir or native species,
disturbance in this habitat. Fire regimes are primarily of natural regeneration, and advance regeneration.16, 103 Noble
the high-severity type,1 but also include the moderate- fir plantations are now fairly common in managed
severity regime (moderately frequent and highly variable) landscapes, even outside the natural range of the species.
for Shasta red fir forests.39 Mean fire-return intervals vary Advance regeneration management tends to simulate
greatly, from •800 years for some mountain hemlock-silver wind disturbance but without the abundant downed
fir forests to about 40 years for red fir forests. Windstorms wood component. Shelterwood cuts are a common
are a common small-scale disturbance and occasionally management strategy in Engelmann spruce or subalpine
result in stand replacement. Insects and fungi are often fir stands.221
important small-scale disturbances. However, they may
Status and Trends. This habitat occupies large areas of
affect larger areas also, for example, laminated root rot
the region. There has probably been little or no decline in
(Phellinus weirii) is a major natural disturbance, affecting
the extent of this type over time. Large areas of this habitat
 32 Wildlife–Habitat Relationships in Oregon and Washington

are relatively undisturbed by human impacts and include Northwest. It is located between the subalpine portions
significant old-growth stands. Other areas have been of the Montane Mixed Conifer Forest habitat in eastern
extensively affected by logging, especially dispersed patch Oregon and Washington and lower tree line Ponderosa
clearcuts. The habitat is stable in area, but is probably still Pine and Forest and Woodlands.
declining in condition because of continued logging. This
Structure. Eastside Mixed Conifer habitats are montane
habitat is one of the best protected, with large areas
forests and woodlands. Stand canopy structure is
represented in national parks and wilderness areas. The
generally diverse, although single-layer forest canopies
only threat is continued road building and clearcutting in
are currently more common than multilayered forests with
unprotected areas. None of the 81 plant associations
snags and large woody debris. The tree layer varies from
representing this habitat listed in the National Vegetation
closed forests to more open-canopy forests or woodlands.
Classification are considered imperiled.10
This habitat may include very open stands. The
undergrowth is complex and diverse. Tall shrubs, low
5. Eastside Mixed Conifer Forest shrubs, forbs or any combination may dominate stands.
Rex C.Crawford Deciduous shrubs typify shrub layers. Prolonged canopy
closure may lead to development of a sparsely vegetated
undergrowth.
Composition. This habitat contains a wide array of tree
species (nine) and stand dominance patterns. Douglas-fir
(Pseudotsuga menziesii) is the most common tree species in
this habitat. It is almost always present and dominates or
co-dominates most overstories. Lower elevations or drier
sites may have ponderosa pine (Pinus ponderosa) as a co-
dominant with Douglas-fir in the overstory and often have
other shade-tolerant tree species growing in the
undergrowth. On moist sites, grand fir (Abies grandis),
western redcedar (Thuja plicata) and/or western hemlock
(Tsuga heterophylla) are dominant or co-dominant with
Douglas-fir. Other conifers include western larch (Larix
occidentalis) and western white pine (Pinus monticola) on
mesic sites, Engelmann spruce (Picea engelmannii),
Geographic Distribution. The Eastside Mixed Conifer
lodgepole pine (Pinus contorta), and subalpine fir (Abies
Forest habitat appears primarily in the Blue Mountains,
lasiocarpa) on colder sites. Rarely, Pacific yew (Taxus
East Cascades, and Okanogan Highland Ecoregions of
brevifolia) may be an abundant undergrowth tree or tall
Oregon, Washington, adjacent Idaho, and western
shrub.
Montana. It also extends north into British Columbia.
Undergrowth vegetation varies from open to nearly
Douglas-fir-ponderosa pine forests occur along the
closed shrub thickets with one to many layers. Throughout
eastern slope of the Oregon and Washington Cascades,
the eastside conifer habitat, tall deciduous shrubs include
the Blue Mountains, and the Okanogan Highlands of
vine maple (Acer circinatum) in the Cascades, Rocky
Washington. Grand fir-Douglas-fir forests and western
Mountain maple (A. glabrum), serviceberry (Amelanchier
larch forests are widely distributed throughout the Blue
alnifolia), oceanspray (Holodiscus discolor), mallowleaf
Mountains and, less so, along the east slope of the
ninebark (Physocarpus malvaceus), and Scouler’s willow
Cascades south of Lake Chelan and in the eastern
(Salix scouleriana) at mid- to lower elevations. Medium-
Okanogan Highlands. Western hemlock-western
tall deciduous shrubs at higher elevations include fools
redcedar-Douglas-fir forests are found in the Selkirk
huckleberry (Menziesia ferruginea), Cascade azalea
Mountains of eastern Washington, and on the east slope
(Rhododendron albiflorum), and big huckleberry (Vaccinium
of the Cascades south of Lake Chelan to the Columbia
membranaceum). Widely distributed, generally drier site
River Gorge.
mid-height to short deciduous shrubs include baldhip rose
Physical Setting. The Eastside Mixed Conifer Forest (Rosa gymnocarpa), shiny-leaf spirea (Spiraea betulifolia), and
habitat is primarily mid-montane with an elevation range snowberry (Symphoricarpos albus, S. mollis, and S.
of between 1,000 and 7,000 ft (305-2,137 m), mostly oreophilus). Low shrubs of higher elevations include low
between 3,000 and 5,500 ft (914-1,676 m). Parent materials huckleberries (Vaccinium cespitosum, and V. scoparium) and
for soil development vary. This habitat receives some of five-leaved bramble (Rubus pedatus). Evergreen shrubs
the greatest amounts of precipitation in the inland represented in this habitat are chinquapin (Castanopsis
northwest, 30-80 inches (76-203 cm)/year. Elevation of this chrysophylla), a tall shrub in southeastern Cascades, low
habitat varies geographically, with generally higher to mid-height dwarf Oregongrape (Mahonia nervosa in the
elevations to the east. east Cascades and M. repens elsewhere), tobacco brush
(Ceanothus velutinus), an increaser with fire, Oregon
Landscape Setting. This habitat makes up most of the
boxwood (Paxistima myrsinites) generally at mid- to lower
continuous montane forests of the inland Pacific
 CHAPTER 2: HABITAT DESCRIPTIONS 33

elevations, beargrass (Xerophyllum tenax), pinemat Generally, wetter sites burn less frequently and stands are
manzanita (Arctostaphylos nevadensis) and kinnikinnick (A. older with more western hemlock and western redcedar
uva-ursi). than drier sites. Many sites dominated by Douglas-fir and
Herbaceous broadleaf plants are important indicators ponderosa pine, which were formerly maintained by
of site productivity and disturbance. Species generally wildfire, may now be dominated by grand fir (a fire
indicating productive sites include western oakfern sensitive, shade-tolerant species).
(Gymnocarpium dryopteris), vanillaleaf (Achlys triphylla),
Succession and Stand Dynamics. Successional
wild sarsparilla (Aralia nudicaulis), wild ginger (Asarum
relationships of this type reflect complex interrelationships
caudatum), queen’s cup (Clintonia uniflora), goldthread
between site potential, plant species characteristics, and
(Coptis occidentalis), false bugbane (Trautvetteria
disturbance regime.228 Generally, early seral forests of
caroliniensis), windflower (Anemone oregana, A. piperi, A.
shade-intolerant trees (western larch, western white pine,
lyallii), fairybells (Disporum hookeri), Sitka valerian
ponderosa pine, Douglas-fir) or tolerant trees (grand fir,
(Valeriana sitchensis), and pioneer violet (Viola glabella).
western redcedar, western hemlock) develop some 50
Other indicator forbs are dogbane (Apocynum
years following disturbance. This stage is preceded by
androsaemifolium), false solomonseal (Maianthemum
forb- or shrub- dominated communities. These early stage
stellata), heartleaf arnica (Arnica cordifolia), several lupines
mosaics are maintained on ridges and drier topographic
(Lupinus caudatus, L. latifolius, L. argenteus ssp. argenteus
positions by frequent fires. Early seral forest develops into
var laxiflorus), western meadowrue (Thalictrum occidentale),
mid-seral habitat of large trees during the next 50-100
rattlesnake plantain (Goodyera oblongifolia), skunkleaf
years. Stand replacing fires recycle this stage back to early
polemonium (Polemonium pulcherrimum), trailplant
seral stages over most of the landscape. Without high-
(Adenocaulon bicolor), twinflower (Linnaea borealis), western
severity fires, a late-seral condition develops either single-
starflower (Trientalis latifolia), and several wintergreens
layer or multilayer structure during the next 100-200 years.
(Pyrola asarifolia, P. picta, Orthilia secunda).
These structures are typical of cool bottomlands that
Graminoids are common in this forest habitat.
usually only experience low-intensity fires.
Columbia brome (Bromus vulgaris), oniongrass (Melica
bulbosa), northwestern sedge (Carex concinnoides) and Effects of Management and Anthropogenic Impacts. This
western fescue (Festuca occidentalis) are found mostly in habitat has been most affected by timber harvesting and
mesic forests with shrubs or mixed with forb species. fire suppression. Timber harvesting has focused on large
Bluebunch wheatgrass (Pseudoroegneria spicata), Idaho shade-intolerant species in mid- and late-seral forests,
fescue (Festuca idahoensis), and junegrass (Koeleria leaving shade-tolerant species. Fire suppression enforces
macrantha) are found in drier more open forests or those logging priorities by promoting less fire-resistant,
woodlands. Pinegrass (Calamagrostis rubescens) and shade-tolerant trees. The resultant stands at all seral stages
Geyer’s sedge (C. geyeri) can form a dense layer under tend to lack snags, have high tree density, and are
Douglas-fir or grand fir trees. composed of smaller and more shade-tolerant trees. Mid-
seral forest structure is currently 70% more abundant than
Other Classifications and Key References. This habitat
in historical, native systems.181 Late-seral forests of shade-
includes the moist portions of the Pseudotsuga menziesii,
intolerant species are now essentially absent. Early-seral
the Abies grandis, and the Tsuga heterophylla zones of eastern
forest abundance is similar to that found historically but
Oregon and Washington.88 This habitat is called Douglas-
lacks snags and other legacy features.
fir (No. 12), Cedar-Hemlock-Pine (No. 13), and Grand fir-
Douglas-fir (No. 14) forests in Kuchler.136 The Oregon Gap Status and Trends. Quigley and Arbelbide181 concluded
II Project126 and Oregon Vegetation Landscape-Level Cover that the Interior Douglas-fir, Grand fir, and Western
Types127 that would represent this type are the eastside redcedar/Western hemlock cover types are more
Douglas-fir dominant-mixed conifer forest, ponderosa abundant now than before 1900, whereas the Western larch
pine dominant mixed conifer forest, and the northeast and Western white pine types are significantly less
Oregon mixed conifer forest. Quigley and Arbelbide181 abundant. Twenty percent of Pacific Northwest Douglas-
referred to this habitat as Grand fir/White fir, the Interior fir, grand fir, western redcedar, western hemlock, and
Douglas-fir, Western larch, Western redcedar/Western western white pine associations listed in the National
hemlock, and Western white pine cover types and the Vegetation Classification are considered imperiled or
Moist Forest potential vegetation group. Other references critically imperiled.10 Roads, timber harvest, periodic
detail forest associations for this habitat.45, 59, 117, 118, 123, 122, 144, grazing, and altered fire regimes have compromised these
148, 208, 209, 212, 221, 228
forests. Even though this habitat is more extensive than
pre-1900, natural processes and functions have been
Natural Disturbance Regime. Fires were probably of
modified enough to alter its natural status as functional
moderate frequency (30-100 years) in presettlement times.
habitat for many species.
Inland Pacific Northwest Douglas-fir and western larch
forests have a mean fire interval of 52 years.22 Typically,
stand-replacement fire-return intervals are 150-500 years
with moderate severity-fire intervals of 50-100 years.
Specific fire influences vary with site characteristics.
 34 Wildlife–Habitat Relationships in Oregon and Washington

6. Lodgepole Pine Forest and graminoid undergrowth appears with open to closed tree
canopies.
Woodlands
Rex C.Crawford Composition. The tree layer of this habitat is dominated
by lodgepole pine (Pinus contorta var. latifolia and P. c. var.
murrayana), but it is usually associated with other montane
conifers (Abies concolor, A. grandis, A. magnifica var.
shastensis, Larix occidentalis, Calocedrus decurrens, Pinus
lambertiana, P. monticola, P. ponderosa, Pseudotsuga menziesii).
Subalpine fir (Abies lasiocarpa), mountain hemlock (Tsuga
mertensiana), Engelmann spruce (Picea engelmannii), and
whitebark pine (Pinus albicaulis), indicators of subalpine
environments, are present in colder or higher sites.
Quaking aspen (Populus tremuloides) may occur in small
numbers.
Shrubs can dominate the undergrowth. Tall deciduous
shrubs include Rocky Mountain maple (Acer glabrum),
serviceberry (Amelanchier alnifolia), oceanspray (Holodiscus
discolor), or Scouler’s willow (Salix scouleriana). These tall
shrubs often occur over a layer of mid-height deciduous
Geographic Distribution. This habitat is found along the shrubs such as baldhip rose (Rosa gymnocarpa), russet
eastside of the Cascade Range, in the Blue Mountains, the buffaloberry (Shepherdia canadensis), shiny-leaf spirea
Okanogan Highlands and ranges north into British (Spiraea betulifolia), and snowberry (Symphoricarpos albus
Columbia and south to Colorado and California. and/or S. mollis). At higher elevations, big huckleberry
With grassy undergrowth, this habitat appears (Vaccinium membranaceum) can be locally important,
primarily along the eastern slope of the Cascade Range particularly following fire. Mid-tall evergreen shrubs can
and occasionally in the Blue Mountains and Okanogan be abundant in some stands, for example, creeping
Highlands. Subalpine lodgepole pine habitat occurs on Oregongrape (Mahonia repens), tobacco brush (Ceanothus
the broad plateau areas along the crest of the Cascade velutinus), and Oregon boxwood (Paxistima myrsinites).
Range and the Blue Mountains, and in the higher Colder and drier sites support low-growing evergreen
elevations in the Okanogan Highlands. On pumice soils shrubs, such as kinnikinnick (Arctostaphylos uva-ursi) or
this habitat is confined to the eastern slope of the Cascade pinemat manzanita (A. nevadensis). Grouseberry (V.
Range from near Mt. Jefferson south to the vicinity of scoparium) and beargrass (Xerophyllum tenax) are consistent
Crater Lake. evergreen low shrub dominants in the subalpine part of
this habitat. Manzanita (Arctostaphylos patula),
Physical Setting. This habitat is located mostly at mid- to kinnikinnick, tobacco brush, antelope bitterbrush (Purshia
higher elevations (3,000-9,000 ft [914-2,743 m]). These tridentata), and wax current (Ribes cereum) are part of this
environments can be cold and relatively dry, usually with habitat on pumice soil.
persistent winter snowpack. A few of these forests occur Some undergrowth is dominated by graminoids with
in low-lying frost pockets, wet areas, or under edaphic few shrubs. Pinegrass (Calamagrostis rubescens) and/or
control (usually pumice) and are relatively long-lasting Geyer’s sedge (Carex geyeri) can appear with grouseberry
features of the landscape. Lodgepole pine is maintained in the subalpine zone. Pumice soils support grassy
as a dominant by the well-drained, deep Mazama pumice undergrowth of long-stolon sedge (C. inops), Idaho fescue
in eastern Oregon. (Festuca idahoensis) or western needlegrass (Stipa
Landscape Setting. This habitat appears within Montane occidentalis). The latter two species may occur with
Mixed Conifer Forest east of the Cascade crest and the bitterbrush or big sagebrush and other bunchgrass steppe
cooler Eastside Mixed Conifer Forest habitats. Most species. Other nondominant indicator graminoids
pumice soil lodgepole pine habitat is intermixed with frequently encountered in this habitat are California
Ponderosa Pine Forest and Woodland habitats and is oatgrass (Danthonia californica), blue wildrye (Elymus
located between Eastside Mixed Conifer Forest habitat and glaucus), Columbia brome (Bromus vulgaris) and
either Western Juniper Woodland or Shrub-steppe habitat. oniongrass (Melica bulbosa). Kentucky bluegrass (Poa
pratensis), and bottlebrush squirreltail (Elymus elymoides)
Structure. The lodgepole pine habitat is composed of open can be locally abundant where livestock grazing has
to closed evergreen conifer tree canopies. Vertical structure persisted.
is typically a single tree layer. Reproduction of other more The forb component of this habitat is diverse and varies
shade-tolerant conifers can be abundant in the with environmental conditions. A partial forb list includes
undergrowth. Several distinct undergrowth types develop goldthread (Coptis occidentalis), false solomonseal
under the tree layer: evergreen or deciduous medium-tall (Maianthemum stellata), heartleaf arnica (Arnica cordifolia),
shrubs, evergreen low shrub, or graminoids with few several lupines (Lupinus caudatus, L. latifolius, L. argenteus
shrubs. On pumice soils, a sparsely developed shrub and ssp. argenteus var. laxiflorus), meadowrue (Thalictrum
 CHAPTER 2: HABITAT DESCRIPTIONS 35

occidentale), queen’s cup (Clintonia uniflora), rattlesnake Effects of Management and Anthropogenic Impacts. Fire
plantain (Goodyera oblongifolia), skunkleaf polemonium suppression has left many single-canopy lodgepole pine
(Polemonium pulcherrimum), trailplant (Adenocaulon bicolor), habitats unburned to develop into more multilayered
twinflower (Linnaea borealis), Sitka valerian (Valeriana stands. Thinning of serotinous lodgepole pine forests with
sitchensis), western starflower (Trientalis latifolia), and fire intervals <20 years can reduce their importance over
several wintergreens (Pyrola asarifolia, P. picta, Orthilia time. In pumice-soil lodgepole stands, lack of natural
secunda). regeneration in harvest units has lead to creation of
“pumice deserts” within otherwise forested habitats.47
Other Classifications and Key References. The
Lodgepole Pine Forest and Woodland habitat includes the Status and Trends. Quigley and Arbelbide181 concluded
Pinus contorta zone of eastern Oregon and Washington.88 that the extent of the lodgepole pine cover type in Oregon
The Oregon Gap II Project126 and Oregon Vegetation and Washington is the same as before 1900 and in regions
Landscape-Level Cover Type127 that would represent this may exceed its historical extent. Five percent of Pacific
type is lodgepole pine forest and woodlands. Quigley and Northwest lodgepole pine associations listed in the
Arbelbide181 referred to this habitat as Lodgepole pine National Vegetation Classification are considered
cover type and as a part of the Dry Forest potential imperiled.10 At a finer scale, these forests have been
vegetation group. Other references detail plant fragmented by roads, timber harvest, and influenced by
associations with this habitat.117, 118, 122, 123, 144, 212, 221 periodic livestock grazing and altered fire regimes.
Natural Disturbance Regime. This habitat typically
reflects early successional forest vegetation that originated 7. Ponderosa Pine Forest and
with fires. Inland Pacific Northwest lodgepole pine has a Woodlands (includes Eastside Oak)
mean fire interval of 112 years.22 Summer drought areas Rex C. Crawford & Jimmy Kagan
generally have low to medium-intensity ground fires
occurring at intervals of 25-50 years, whereas areas with
more moisture have a sparse undergrowth and slow fuel
build-up that results in less frequent, more intense fire.
With time, lodgepole pine stands increase in fuel loads.
Woody fuels accumulate on the forest floor from insect
(mountain pine beetle) and disease outbreaks and residual
wood from past fires. Mountain pine beetle outbreaks thin
stands that add fuel and create a drier environment for
fire or open canopies and create gaps for other conifer
regeneration. High-severity crown fires are likely in young
stands, when the tree crowns are near deadwood on the
ground. After the stand opens up, shade-tolerant trees
increase in number.
Succession and Stand Dynamics. Most Lodgepole Pine
Forest and Woodlands are early- to mid seral stages
initiated by fire. Typically, lodgepole pine establishes Geographic Distribution. This habitat occurs in much of
within 10-20 years after fire. This can be a gap phase eastern Washington and eastern Oregon, including the
process where seed sources are scarce. Lodgepole stands eastern slopes of the Cascades, the Blue Mountains and
break up after 100-200 years. Without fires and insects, foothills, and the Okanogan Highlands. Variants of it also
stands become more closed-canopy forest with sparse occur in the Rocky Mountains, the eastern Sierra Nevada,
undergrowth. Because lodgepole pine cannot reproduce and mountains within the Great Basin. It extends into
under its own canopy, old unburned stands are replaced south-central British Columbia as well.
by shade-tolerant conifers. Lodgepole pine on pumice soils In the Pacific Northwest, ponderosa pine-Douglas-fir
is not seral to other tree species; these extensive stands, if woodland habitats occur along the eastern slope of the
not burned, thin naturally, with lodgepole pine Cascades, the Okanogan Highlands, and in the Blue
regenerating in patches. On poorly drained pumice soils, Mountains. Ponderosa pine woodland and savanna
quaking aspen sometimes plays a mid-seral role and is habitats occur in the foothills of the Blue Mountains, along
displaced by lodgepole when aspen clones die. Serotinous the eastern base of the Cascade Range, the Okanogan
cones (cones releasing seeds after fire) are uncommon in Highlands, and in the Columbia Basin in northeastern
eastern Oregon lodgepole pine (P. c. var. murrayana). On Washington. Ponderosa pine is widespread in the pumice
the Colville National Forest in Washington, only 10% of zone of south-central Oregon between Bend and Crater
lodgepole pine (P. c. var. latifolia) trees in low-elevation Lake east of the Cascade Crest. Ponderosa pine-Oregon
Douglas-fir habitats had serotinous cones, whereas 82% white oak habitat appears east of the Cascades in the
of cones in high-elevation subalpine fir habitats were vicinity of Mt. Hood near the Columbia River Gorge north
serotinous.4 to the Yakama Nation and south to the Warm Springs
Nation. Oak-dominated woodlands follow a similar
 36 Wildlife–Habitat Relationships in Oregon and Washington

distribution as Ponderosa Pine-White Oak but are more a tall to medium-tall deciduous shrub layer of mallowleaf
restricted and less common. ninebark (Physocarpus malvaceus) or common snowberry
(Symphoricarpos albus). Grand fir seedlings or saplings may
Physical Setting. This habitat generally occurs on the
be present in the undergrowth. Pumice soils support a
driest sites supporting conifers in the Pacific Northwest.
shrub layer represented by green-leaf or white-leaf
It is widespread and variable, appearing on moderate to
manzanita (Arctostaphylos patula or A. viscida). Short
steep slopes in canyons, foothills, and on plateaus or plains
shrubs, pinemat manzanita (Arctostaphylos nevadensis) and
near mountains. In Oregon, this habitat can be maintained
kinnikinnick (A. uva-ursi) are found across the range of
by the dry pumice soils, and in Washington it can be
this habitat. Antelope bitterbrush (Purshia tridentata), big
associated with serpentine soils. Average annual
sagebrush (Artemisia tridentata), black sagebrush (A. nova),
precipitation ranges from about 14 to 30 inches (36 to 76
green rabbitbrush (Chrysothamnus viscidiflorus), and in
cm) on ponderosa pine sites in Oregon and Washington,
southern Oregon, curl-leaf mountain mahogany
and often as snow. This habitat can be found at elevations
(Cercocarpus ledifolius) often grow with Douglas-fir,
of 100 ft (30m) in the Columbia River Gorge to dry, warm
ponderosa pine and/or Oregon white oak, which typically
areas over 6,000 ft (1,829 m). Timber harvest, livestock
have a bunchgrass and shrub-steppe ground cover.
grazing, and pockets of urban development are major land
Undergrowth is generally dominated by herbaceous
uses.
species, especially graminoids. Within a forest matrix,
Landscape Setting. This woodland habitat typifies the these woodland habitats have an open to closed sodgrass
lower tree line zone forming transitions with Eastside undergrowth dominated by pinegrass (Calamagrostis
Mixed Conifer Forest and Western Juniper and Mountain rubescens), Geyer’s sedge (Carex geyeri), Ross’ sedge (C.
Mahogany Woodland, Shrub-steppe, Eastside Grasslands, rossii), long-stolon sedge (C. inops), or blue wildrye (Elymus
or Agriculture habitats. Douglas-fir-ponderosa pine glaucus). Drier savanna and woodland undergrowth
woodlands are found near or within the Eastside Mixed typically contains bunchgrass steppe species, such as
Conifer Forest habitat. Oak woodlands appear in the Idaho fescue (Festuca idahoensis), rough fescue (F.
driest, most restricted, landscapes in transition to Eastside campestris), bluebunch wheatgrass (Pseudoroegneria spicata),
Grasslands or Shrub-steppe. Indian ricegrass (Oryzopsis hymenoides), or needlegrasses
(Stipa comata, S. occidentalis). Common exotic grasses that
Structure. This habitat is typically a woodland or savanna
may appear in abundance are cheatgrass (Bromus
with tree canopy coverage of 10-60%, although closed-
tectorum), and bulbous bluegrass (Poa bulbosa). Forbs are
canopy stands are possible. The tree layer is usually
common associates in this habitat and are too numerous
composed of widely spaced large conifer trees. Many
to be listed.
stands tend towards a multilayered condition with
encroaching conifer regeneration. Isolated taller conifers Other Classifications and Key References. This habitat
above broadleaf deciduous trees characterize part of this is referred to as Merriam’s Arid Transition Zone, Western
habitat. Deciduous woodlands or forests are an important ponderosa forest (Pinus), and Oregon Oak wood (Quercus)
part of the structural variety of this habitat. Clonal in Kuchler,136 and as Pacific ponderosa pine-Douglas-fir,
deciduous trees can create dense patches across a grassy Pacific ponderosa pine, and Oregon white oak by the
landscape rather than scattered individual trees. The Society of American Foresters. The Oregon Gap II Project126
undergrowth may include dense stands of shrubs or, more and Oregon Vegetation Landscape-Level Cover Types127
often, be dominated by grasses, sedges, or forbs. Shrub- that would represent this type are ponderosa pine forest
steppe shrubs may be prominent in some stands and create and woodland, ponderosa pine-white oak forest and
a distinct tree-shrub-sparse-grassland habitat. woodland, and ponderosa pine-lodgepole pine on pumice.
Other references describe elements of this habitat.45, 62, 88,
Composition. Ponderosa pine (Pinus ponderosa) and 117, 118, 121, 122, 123, 144, 148, 209, 212, 221, 222
Douglas-fir (Pseudotsuga menziesii) are the most common
evergreen trees in this habitat. The deciduous conifer, Natural Disturbance Regime. Fire plays an important role
western larch (Larix occidentalis), can be a co-dominant in creating vegetation structure and composition in this
with the evergreen conifers in the Blue Mountains of habitat. Most of the habitat has experienced frequent low-
Oregon, but seldom as a canopy dominant. Grand fir (Abies severity fires that maintained woodland or savanna
grandis) may be frequent in the undergrowth on more conditions. A mean fire interval of 20 years for ponderosa
productive sites, giving stands a multilayer structure. In pine is the shortest of the vegetation types listed by Barrett
rare instances, grand fir can be co-dominant in the upper et al.22 Soil drought plays a role in maintaining an open
canopy. Tall ponderosa pine over Oregon white oak tree canopy in part of this dry woodland habitat.
(Quercus garryana) trees form stands along part of the east
Succession and Stand Dynamics. This habitat is climax
Cascades. These stands usually have younger cohorts of
on sites near the dry limits of each of the dominant conifer
pines. Oregon white oak dominates open woodlands or
species and is more seral as the environment becomes
savannas in limited areas.
more favorable for tree growth. Open seral stands are
The undergrowth can include dense stands of shrubs
gradually replaced by more closed shade-tolerant climax
or, more often, be dominated by grasses, sedges, and/or
stands. Oregon white oak can reproduce under its own
forbs. Some Douglas-fir and ponderosa pine stands have
shade but is intolerant of overtopping by conifers. Oregon
 CHAPTER 2: HABITAT DESCRIPTIONS 37

white oak woodlands are considered fire climax and are 8. Upland Aspen Forest
seral to conifers. In drier conditions, unfavorable to Rex C. Crawford & Jimmy Kagan
conifers, oak is climax. Oregon white oak sprouts from
the trunk and root crown following cutting or burning
and form clonal patches of trees.
Effects of Management and Anthropogenic Impacts.
Before 1900, this habitat was mostly open and park like
with relatively few undergrowth trees. Currently, much
of this habitat has a younger tree cohort of more shade-
tolerant species that gives the habitat a more closed,
multilayered canopy. For example, this habitat includes
previously natural fire-maintained stands in which grand
fir can eventually become the canopy dominant. Fire
suppression has lead to a buildup of fuels that in turn
increase the likelihood of stand-replacing fires. Heavy
grazing, in contrast to fire, removes the grass cover and
tends to favor shrub and conifer species. Fire suppression
combined with grazing creates conditions that support Geographic Distribution. Quaking aspen groves are the
cloning of oak and invasion by conifers. Large late-seral most widespread habitat in North America, but are a
ponderosa pine, Douglas-fir, and Oregon white oak are minor type throughout eastern Washington and Oregon.
harvested in much of this habitat. Under most Upland Aspen habitat is found in the isolated mountain
management regimes, typical tree size decreases and tree ranges of southeastern Oregon, e.g., Steens Mountain, and
density increases in this habitat. Ponderosa pine-Oregon in the northeastern Cascades of Washington. Aspen stands
white oak habitat is now denser than in the past and may are much more common in the Rocky Mountain states.
contain more shrubs than in presettlement habitats. In
Physical Setting. This habitat generally occurs on well-
some areas, new woodlands have been created by patchy
drained mountain slopes or canyon walls that have some
tree establishment at the forest-steppe boundary.
moisture. Rockfalls, talus, or stony north slopes are often
Status and Trends. Quigley and Arbelbide181 concluded typical sites. It may occur in steppe on moist microsites,
that the Interior Ponderosa Pine cover type is significantly but is not associated with streams, ponds, or wetlands.
less in extent than pre-1900 and that the Oregon White This habitat is found from 2,000 to 9,500 ft (610 to 2,896
Oak cover type is greater in extent than pre-1900. They m) elevation.
included much of this habitat in their Dry Forest potential
Landscape Setting. Aspen forms a “subalpine belt” above
vegetation group,181 which they concluded has departed
the Western Juniper and Mountain Mahogany Woodland
from natural succession and disturbance conditions. The
habitat and below montane Shrub-steppe Habitat and
greatest structural change in this habitat is the reduced
Alpine Grasslands on Steens Mountain in southern
extent of the late-seral, single-layer condition. This habitat
Oregon. It can occur in seral stands in the lower Eastside
is generally degraded because of increased exotic plants
Mixed Conifer Forest and Ponderosa Pine Forest and
and decreased native bunchgrasses. One third of Pacific
Woodland habitats. Primary land use is livestock grazing.
Northwest Oregon white oak, ponderosa pine, and dry
Douglas-fir or grand fir community types listed in the Structure. Deciduous trees usually <48 ft (15 m) tall
National Vegetation Classification are considered dominate this woodland or forest habitat. The tree layer
imperiled or critically imperiled.10 grows over a forb-, grass-, or low-shrub-dominated
undergrowth. Relatively simple two-tiered stands
characterize the typical vertical structure of woody plants
in this habitat. This habitat is composed of one to many
clones of trees with larger trees toward the center of each
clone. Conifers invade and create mixed evergreen-
deciduous woodland or forest habitats.
Composition. Quaking aspen (Populus tremuloides) is the
characteristic and dominant tree in this habitat. It is the
sole dominant in many stands although scattered
ponderosa pine (Pinus ponderosa) or Douglas-fir
(Pseudotsuga menziesii) may be present. Snowberry
(Symphoricarpos oreophilus and less frequently S. albus) is
the most common dominant shrub. Tall shrubs, Scouler’s
willow (Salix scouleriana) and serviceberry (Amelanchier
alnifolia) may be abundant. On mountain or canyon slopes,
antelope bitterbrush (Purshia tridentata), mountain big
 38 Wildlife–Habitat Relationships in Oregon and Washington

sagebrush (Artemisia tridentata ssp. vaseyana), low 9. Subalpine Parkland

sagebrush (A. arbuscula), and curl-leaf mountain Rex C. Crawford & Christopher B. Chappell
mahogany (Cercocarpus ledifolius) often occur in and
adjacent to this woodland habitat.
In some stands, pinegrass (Calamagrostis rubescens) may
dominate the ground cover without shrubs. Other
common grasses are Idaho fescue (Festuca idahoensis),
California brome (Bromus carinatus), or blue wildrye
(Elymus glaucus). Characteristic tall forbs include
horsemint (Agastache spp.), aster (Aster spp.), senecio
(Senecio spp.), coneflower (Rudbeckia spp.). Low forbs
include meadowrue (Thalictrum spp.), bedstraw (Galium
spp.), sweetcicely (Osmorhiza spp.), and valerian (Valeriana
spp.).
Other Classifications and Key References. This habitat
is called “Aspen” by the Society of American Foresters
and “Aspen woodland” by the Society of Range
Management. The Oregon Gap II Project126 and Oregon Geographic Distribution. The Subalpine Parkland habitat
Vegetation Landscape-Level Cover Type127 that would occurs throughout the high mountain ranges of
represent this type is aspen groves. Other references Washington and Oregon (e.g., Cascade crest, Olympic
describe this habitat2, 88, 119, 161, 222. Mountains, Wallowa and Blue mountains, and Okanogan
Highlands), extends into mountains of Canada and
Natural Disturbance Regime. Fire plays an important role
Alaska, and to the Sierra Nevadas and Rocky Mountains.
in maintenance of this habitat. Quaking aspen will
colonize sites after fire or other stand disturbances through Physical Setting. Climate is characterized by cool
root sprouting. Research on fire scars in aspen stands in summers and cold winters with deep snowpack, although
central Utah119 indicated that most fires occurred before much variation exists among specific vegetation types.
1885, and concluded that the natural fire return interval Mountain hemlock sites receive an average precipitation
was 7-10 years. Ungulate browsing plays a variable role of >50 inches (127 cm) in 6 months and several feet of snow
in aspen habitat; ungulates may slow tree regeneration often accumulates. Whitebark pine sites receive 24-70
by consuming aspen sprouts on some sites, and may have inches (61-178 cm) per year and some sites only rarely
little influence in other stands. accumulate a significant snowpack. Summer soil drought
is possible in eastside parklands but rare in westside areas.
Succession and Stand Dynamics. There is no generalized
Elevation varies from 4,500 to 6,000 ft (1,371 to 1,829 m) in
successional pattern across the range of this habitat. Aspen
the western Cascades and Olympic Mountains and from
sprouts after fire and spreads vegetatively into large clonal
5,000 to 8,000 ft (1,524 to 2,438 m) in the eastern Cascades
or multiclonal stands. Because aspen is shade intolerant
and Wallowa mountains.
and cannot reproduce under its own canopy, conifers can
invade most aspen habitat. In central Utah, quaking aspen Landscape Setting. The Subalpine Parkland habitat lies
was invaded by conifers in 75-140 years. Apparently, some above the Mixed Montane Conifer Forest or Lodgepole
aspen habitat is not invaded by conifers, but eventually Pine Forest habitat and below the Alpine Grassland and
clones deteriorate and succeed to shrubs, grasses, and/or Shrubland habitat. Associated wetlands in subalpine
forbs. This transition to grasses and forbs occurs more parklands extend up a short distance into the alpine zone.
likely on dry sites. Primary land use is recreation, watershed protection, and
grazing.
Effects of Management and Anthropogenic Impacts.
Domestic sheep reportedly consume four times more Structure. Subalpine Parkland habitat has a tree layer
aspen sprouts than do cattle. Heavy livestock browsing typically between 10 and 30% canopy cover. Openings
can adversely impact aspen growth and regeneration. among trees are highly variable. The habitat appears either
With fire suppression and alteration of fine fuels, fire as parkland, that is, a mosaic of treeless openings and small
rejuvenation of aspen habitat has been greatly reduced patches of trees often with closed canopies, or as
since about 1900. Conifers now dominate many seral aspen woodlands or savanna-like stands of scattered trees. The
stands and extensive stands of young aspen are ground layer can be composed of (1) low to matted dwarf-
uncommon. shrubs (<1 ft [0.3 m] tall) that are evergreen or deciduous
and often small-leaved; (2) sod grasses, bunchgrasses, or
Status and Trends. With fire suppression and change in
sedges; (3) forbs; or (4) moss- or lichen-covered soils. Herb
fire regimes, the Aspen Forest habitat is less common than
or shrub-dominated wetlands appear within the parkland
before 1900. None of the 5 Pacific Northwest upland
areas and are considered part of this habitat; wetlands can
quaking aspen community types in the National
occur as deciduous shrub thickets up to 6.6 ft (2 m) tall, as
Vegetation Classification are considered imperiled.10
scattered tall shrubs, as dwarf shrub thickets, or as short
 CHAPTER 2: HABITAT DESCRIPTIONS 39

herbaceous plants <1.6 ft (0.5 m) tall. In general, western lupine (Lupinus arcticus ssp. subalpinus), and alpine aster
Cascades and Olympic areas are mostly parklands (Aster alpigenus). Showy sedge (Carex spectabilis) is also
composed of a mosaic of patches of trees interspersed with locally abundant.
heather shrublands or wetlands, whereas, eastern
Other Classifications and Key References. This habitat
Cascades and Rocky mountain areas are parklands and
is called the Hudsonian Zone,155 Parkland subzone,134
woodlands typically dominated by grasses or sedges, with
meadow-forest mosaic, 74 upper subalpine zone, 88
fewer heathers.
Meadows and Park, and Subalpine Parkland.20 Quigley
Composition. Species composition in this habitat varies and Arbelbide181 called this habitat Whitebark pine and
with geography or local site conditions. The tree layer can Whitebark pine-Subalpine larch cover types. Kuchler136
be composed of one or several tree species. Subalpine fir included this within the subalpine fir-mountain hemlock
(Abies lasiocarpa), Engelmann spruce (Picea engelmannii) forest. The Oregon Gap II Project126 and Oregon Vegetation
and lodgepole pine (Pinus contorta) are found throughout Landscape-Level Cover Types127 that would represent this
the Pacific Northwest, whereas limber pine (P. flexilis) is type are whitebark-lodgepole pine montane forest and
restricted to southeastern Oregon. Alaska yellowcedar subalpine parkland. Additional references describe this
(Chamaecyparis nootkatensis), Pacific silver fir (A. amabilis), habitat.11, 49, 75, 105, 112, 114, 115, 139, 144, 221
and mountain hemlock (Tsuga mertensiana) are most
Natural Disturbance Regime. Although fire is rare to
common in the Olympics and western Cascades.
infrequent in this habitat, it plays an important role,
Whitebark pine (P. albicaulis) is found primarily in the
particularly in drier environments. Whitebark pine
eastern Cascades mountains, Okanogan Highlands, and
woodland fire intervals varied from 50 to 300 years before
Blue Mountains. Subalpine larch (Laryx lyallii) occurs only
1900. Mountain hemlock parkland fire reccurrence is 400-
in the northern Cascade Mountains, primarily east of the
800 years. Wind blasting by ice and snow crystals is a
crest.
critical factor in these woodlands and establishes the
West Cascades and Olympic areas generally are
higher limits of the habitat. Periodic shifts in climatic
parklands. Tree islands often have big huckleberry
factors, such as drought, snowpack depth, or snow
(Vaccinium membranaceum) in the undergrowth
duration either allow tree invasions into meadows and
interspersed with heather shrublands between. Openings
shrublands or eliminate or retard tree growth. Volcanic
are composed of pink mountain-heather (Phyllodoce
activity plays a long-term role in establishing this habitat.
empetriformis), and white mountain-heather (Cassiope
Wetlands are usually seasonally or perennially flooded
mertensiana) and Cascade blueberry (Vaccinium deliciosum).
by snowmelt and springs, or by subirrigation.
Drier areas are more woodland or savanna-like, often with
low shrubs, such as common juniper (Juniperus communis), Succession and Stand Dynamics. Succession in this
kinnikinnick (Arctostaphylos uva-ursi), low whortleberries habitat occurs through a complex set of relationships
or grouseberries (Vaccinium myrtillus or V. scoparium) or between vegetation response to climatic shifts and
beargrass (Xerophyllum tenax) dominating the ground catastrophic disturbance, and plant species interactions
cover. Wetland shrubs in the Subalpine Parkland habitat and site modification that create microsites. A typical
include bog-laurel (Kalmia microphylla), Booth’s willow succession of subalpine trees into meadows or shrublands
(Salix boothii), undergreen willow (S. commutata), Sierran begins with the invasion of a single tree, subalpine fir and
willow (S. eastwoodiae), and blueberries (Vaccinium mountain hemlock in the wetter climates and whitebark
uliginosum or V. deliciosum) pine and subalpine larch in drier climates. If the
Undergrowth in drier areas may be dominated by environment allows, tree density slowly increases (over
pinegrass (Calamagrostis rubescens), Geyer’s sedge (Carex decades to centuries) through seedlings or branch layering
geyeri), Ross’ sedge (C. rossii), smooth woodrush (Luzula by subalpine fir. The tree patches or individual trees
glabrata var. hitchcockii), Drummond’s rush (Juncus change the local environment and create microsites for
drummondii), or short fescues (Festuca viridula, F. shade-tolerant trees, Pacific silver fir in wetter areas, and
brachyphylla, F. saximontana). Various sedges are subalpine fir and Engelmann spruce in drier areas.
characteristic of wetland graminoid-dominated habitats: Whitebark pine, an early invading tree, is dispersed long
black (Carex nigricans), Holm’s Rocky Mountain (C. distances by Clark’s nutcrackers and shorter distances by
scopulorum), Sitka (C. aquatilis var. dives) and Northwest mammals. Most other tree species are wind dispersed.
Territory (C. utriculatia) sedges. Tufted hairgrass
Effects of Management and Anthropogenic Impacts. Fire
(Deschampsia caespitosa) is characteristic of subalpine
suppression has contributed to change in habitat structure
wetlands.
and functions. For example, the current “average”
The remaining flora of this habitat is diverse and
whitebark pine stand will burn every 3,000 years or longer
complex. The following herbaceous broadleaf plants are
because of fire suppression. Blister rust, an introduced
important indicators of differences in the habitat:
pathogen, is increasing whitebark pine mortality in these
American bistort (Polygonum bistortoides), American false
woodlands.4 Even limited logging can have prolonged
hellebore (Veratrum viride), fringe leaf cinquefoil (Potentilla
effects because of slow invasion rates of trees. This is
flabellifolia), marsh marigolds (Caltha leptosepala), avalanche
particularly important on drier sites and in subalpine larch
lily (Erythronium montanum), partridgefoot (Luetkea
stands. During wet cycles, fire suppression can lead to
pectinata), Sitka valerian (Valeriana sitchensis), subalpine
 40 Wildlife–Habitat Relationships in Oregon and Washington

tree islands coalescing and the conversion of parklands are generally poorly developed and shallow, though in
into a more closed forest habitat. Parkland conditions can subalpine grasslands they may be somewhat deeper or
displace alpine conditions through tree invasions. better developed. Geologic parent material varies with
Livestock use and heavy horse or foot traffic can lead to local geologic history.
trampling and soil compaction. Slow growth in this habitat
Landscape Setting. This habitat always occurs above
prevents rapid recovery.
upper treeline in the mountains or a short distance below
Status and Trends. This habitat is generally stable with it (grasslands in the subalpine parkland zone). Typically,
local changes to particular tree variants. Whitebark pine it occurs adjacent to, or in a mosaic with, Subalpine
is maybe declining because of the effects of blister rust or Parkland. Occasionally, it may grade quickly from this
fire suppression that leads to conversion of parklands to habitat down into Montane Mixed Conifer Forest without
more closed forest. Global climate warming will likely intervening Subalpine Parkland. In southeastern Oregon,
have an amplified effect throughout this habitat. Less than this habitat occurs adjacent to and above Upland Aspen
10% of Pacific Northwest subalpine parkland community Forest and Shrub-steppe habitats. Small areas of Open
types listed in the National Vegetation Classification are Water, Herbaceous Wetlands, and Subalpine Parkland
considered imperiled.10 habitats sometimes occur within a matrix of this habitat.
Cliffs, talus, and other barren areas are common features
10. Alpine Grasslands and Shrublands within or adjacent to this habitat. Land use is primarily
recreation, but in some areas east of the Cascade Crest, it
Christopher B. Chappell & Jimmy Kagan
is grazing, especially by sheep.
Structure. This habitat is dominated by grassland, dwarf-
shrubland (mostly evergreen microphyllous), or forbs.
Cover of the various life forms is extremely variable, and
total cover of vascular plants can range from sparse to
complete. Patches of krummholz (coniferous tree species
maintained in shrub form by extreme environmental
conditions) are a common component of this habitat,
especially just above upper treeline. In subalpine
grasslands, which are considered part of this habitat,
widely scattered coniferous trees sometimes occur. Five
major structural types can be distinguished: (1) subalpine
and alpine bunchgrass grasslands, (2) alpine sedge turf,
(3) alpine heath or dwarf-shrubland, (4) fellfield and
boulderfield, and (5) snowbed forb community. Fellfields
have a large amount of bare ground or rocks with a diverse
Geographic Distribution. This habitat occurs in high and variable open layer of forbs, graminoids, and less
mountains throughout the region, including the Cascades, commonly dwarf-shrubs. Snowbed forb communities
Olympic Mountains, Okanogan Highlands, Wallowa have relatively sparse cover of few species of mainly forbs.
Mountains, Blue Mountains, Steens Mountain in In the alpine zone, these types often occur in a complex
southeastern Oregon, and, rarely, the Siskiyous. It is most fine-scale mosaic with each other.
extensive in the Cascades from Mount Rainier north and
Composition. Most subalpine or alpine bunchgrass
in the Wallowa Mountains. Similar habitats occur
grasslands are dominated by Idaho fescue (Festuca
throughout mountains of northwestern North America.
idahoensis), alpine fescue (F. brachyphylla), green fescue (F.
Physical Setting. The climate is the coldest of any habitat viridula), Rocky Mountain fescue (F. saximontana), or timber
in the region. Winters are characterized by moderate to oatgrass (Danthonia intermedia), and to a lesser degree,
deep snow accumulations, very cold temperatures, and purple reedgrass (Calamagrostis purpurascens), downy oat-
high winds. Summers are relatively cool. Growing seasons grass (Trisetum spicatum) or muttongrass (Poa fendleriana).
are short because of persistent snow pack or frost. Blowing Forbs are diverse and sometimes abundant in the
snow and ice crystals on top of the snow pack at and above grasslands. Alpine sedge turfs may be moist or dry and
treeline prevent vegetation such as trees from growing are dominated by showy sedge (Carex spectabilis), black
above the depth of the snow pack. Snow pack protects alpine sedge (C. nigricans), Brewer’s sedge (C. breweri),
vegetation from the effects of this winter wind-related capitate sedge (C. capitata), nard sedge (C. nardina),
disturbance and from excessive frost heaving. Community dunhead sedge (C. phaeocephala), or western single-spike
composition is much influenced by relative duration of sedge (C. pseudoscirpoidea).
snow burial and exposure to wind and frost heaving.75 One or more of the following species dominates alpine
Elevation ranges from a minimum of 5,000 ft (1,524 m) in heaths: pink mountain-heather (Phyllodoce empetriformis),
parts of the Olympics to •10,000 ft (3,048 m). The green mountain-heather (P. glanduliflora), white mountain-
topography varies from gently sloping broad ridgetops, heather (Cassiope mertensiana), or black crowberry
to glacial cirque basins, to steep slopes of all aspects. Soils (Empetrum nigrum). Other less extensive dwarf-shrublands
 CHAPTER 2: HABITAT DESCRIPTIONS 41

may be dominated by the evergreen coniferous common Effects of Management and Anthropogenic Impacts. The
juniper (Juniperus communis), the evergreen broadleaf major human impacts on this habitat are trampling and
kinnikinnick (Arctostaphylos uva-ursi), the deciduous associated recreational impacts, e.g., tent sites. Resistance
shrubby cinquefoil (Pentaphylloides floribunda) or willows and resilience of vegetation to impacts varies by life form.48
(Salix cascadensis and S. reticulata ssp. nivalis). Tree species Sedge turfs are perhaps most resilient to trampling and
occurring as shrubby krummholz in the alpine are heaths are least resilient. Trampling to the point of
subalpine fir (Abies lasiocarpa), whitebark pine (Pinus significantly opening an alpine heath canopy will initiate
albicaulis), mountain hemlock (Tsuga mertensiana), a degradation and erosion phase that results in continuous
Engelmann spruce (Picea engelmannii), and subalpine larch bare ground, largely unsuitable for vascular plant
(Larix lyallii). growth.80 Bare ground in the alpine zone left alone after
Fellfields and similar communities are typified by recreational disturbance will typically not revegetate in a
variable species assemblages and co-dominance of time frame that humans can appreciate. Introduction of
multiple species, including any of the previously exotic ungulates can have noticeable impacts (e.g.,
mentioned species, especially the sedges, as well as golden mountain goats in the Olympic Mountains). Domestic
fleabane (Erigeron aureus), Lobb’s lupine (Lupinus sellulus sheep grazing has also had dramatic impacts,196 especially
var. lobbii), spreading phlox (Phlox diffusa), eight-petal in the bunchgrass habitats east of the Cascades.
mountain-avens (Dryas octopetala), louseworts (Pedicularis
Status and Trends. This habitat is naturally very limited
contorta, P. ornithorhyncha) and many others. Snowbed forb
in extent in the region. There has been little to no change
communities are dominated by Tolmie’s saxifrage
in abundance over the last 150 years. Most of this habitat
(Saxifraga tolmiei), Shasta buckwheat (Eriogonum
is still in good condition and dominated by native species.
pyrolifolium), or Piper’s woodrush (Luzula piperi).
Some areas east of the Cascade Crest have been degraded
Other Classifications and Key References. This habitat by livestock use. Recreational impacts are noticeable in
is equivalent to the alpine communities and the subalpine some national parks and wilderness areas. Current trends
Festuca communities of Franklin and Dyrness.88 It is also seem to be largely stable, though there may be some slow
referred to as Alpine meadows and barren No. 52.136 The loss of subalpine grassland to recent tree invasion. Threats
Oregon Gap II Project 126 and Oregon Vegetation include increasing recreational pressures, continued
Landscape-Level Cover Types127 that would represent this grazing at some sites, and, possibly, global climate change
type are subalpine grassland and alpine fell-snowfields; resulting in expansion of trees into this habitat. Only 1
represented by nonforest in the alpine/parkland zone of out of 40 plant associations listed in the National
Washington Gap.37 Other references describe this habitat.61, Vegetation Classification is considered imperiled.10
65, 75, 80, 94, 105, 112, 123, 139, 195, 207

Natural Disturbance Regime. Most natural disturbances 11. Westside Grasslands

seem to be small scale in their effects or very infrequent. Christopher B. Chappell & Jimmy Kagan
Herbivory and associated trampling disturbance by elk,
mountain goats, and occasionally bighorn sheep seems
to be an important disturbance in some areas, creating
patches of open ground, though the current distribution
and abundance of these ungulates is in part a result of
introductions. Small mammals can also have significant
effects on vegetation: e.g., the heather vole occasionally
overgrazes heather communities.80 Frost heaving is a
climatically related small-scale disturbance that is
extremely important in structuring the vegetation 80
Extreme variation from the norm in snow pack depth and
duration can act as a disturbance, exposing plants to
winter dessication,80 shortening the growing season, or
facilitating summer drought. Subalpine grasslands
probably burn on occasion and can be formed or expanded
in area by fires in subalpine parkland.139 Geographic Distribution. This habitat is restricted
primarily to the Puget Lowland and Willamette Valley
Succession and Stand Dynamics. Little is known about
ecoregions, with most now occurring in Pierce, Thurston,
vegetation changes in these communities, in part because
and San Juan counties, Washington. It also occurs in
changes are relatively slow. Tree invasion rates into
scattered small outliers in the Coast Range of Oregon, the
subalpine grasslands are relatively slow compared to other
eastern Olympic Mountains and the Western Cascades of
subalpine communities.139 Seedling establishment for
southern Washington and Oregon, and in adjacent
many plant species in the alpine zone is poor. Heath
southwestern British Columbia.
communities take about 200 years to mature after initial
establishment and may occupy the same site for thousands Physical Setting. The climate is mild and moderately dry
of years.139 (17-55 inches [43-140 cm] mean annual precipitation), with
 42 Wildlife–Habitat Relationships in Oregon and Washington

moist winters and dry summers. Elevation is mostly low Puget Lowland and in montane balds of the Oregon Coast
and ranges up to a maximum of about 3,500 ft (1,067 m). Range and the Columbia Gorge. Non-native varieties of
Topography varies from flat, to mounded or rolling, to red fescue can occur throughout the area, especially in
steep slopes. Most sites are topoedaphically dry and degraded habitats. California oatgrass communities are
experience extreme soil drought in the summer. Much of found in the San Juan Islands and in the Willamette Valley.
what currently remains of this habitat is found on the Junegrass is a co-dominant in some montane balds and
South Puget prairies, which are underlain by very deep the Willamette Valley prairies; it occurs less abundantly
gravelly/sandy glacial outwash that is excessively well throughout the area. Lemmon’s needlegrass is primarily
drained. Many other small sites, often called “balds”, have found on shallow-soiled balds of Willamette Valley fringes
shallow soils overlying bedrock and typically are on south- and the San Juan Islands.
or west-facing slopes. The habitat also occurs rarely in The most common savanna tree is Douglas-fir
Oregon on deeper soils that are not excessively drained. (Pseudotsuga menziesii). Oregon white oak (Quercus
garryana) formerly was part of extensive savannas, but is
Landscape Setting. This habitat occurs adjacent to or in a
now rare in that structural condition. Ponderosa pine
mosaic with Westside Riparian-Wetlands, Westside Oak
(Pinus ponderosa) is very local. The most common shrub is
and Dry Douglas-fir Forests and Woodlands, Agriculture,
the exotic species Scot’s broom (Cytisus scoparius), which
or Urban habitats. Westside grassland habitat occurs less
frequently forms open stands over the grass. Common
commonly in a matrix of Westside Lowland Conifer-
snowberry (Symphoricarpos albus), Nootka rose (Rosa
Hardwood Forest. In the San Juan Islands, the habitat
nutkana), poison-oak (Toxicodendron diversilobum), and
sometimes occurs on bluffs or slopes adjacent to marine
serviceberry (Amelanchier alnifolia) are other common
habitats. Currently this habitat is used for grazing,
shrubs. The dwarf shrubs kinnikinnick (Arctostaphylos uva-
recreation, and, in the southern Puget Sound area, for
ursi) and common juniper (Juniperus communis) sometimes
military training.
dominate small areas in montane balds, and the former
Structure. This habitat is grassland or, less commonly, sometimes on South Puget prairies. Racomitrium canescens
savanna, with <30% tree or shrub cover. Bunchgrasses is the most common ground moss.
predominate in native-dominated sites, with space
Other Classifications and Key References. Portions of
between the vascular plants typically covered by mosses,
this habitat have been referred to as prairies by many
fruticose lichens, or native forbs. Montane balds are
authors. Franklin and Dyrness88 described this habitat as
sometimes dominated in part by short forbs (<1.6ft. [0.6
prairie in the Puget Sound area, grassland in the San Juan
m]) or dwarf shrubs. Degraded sites are dominated by
Islands and Interior Valley zone of Oregon, and grass balds
rhizomatous exotic grasses with some native herbaceous
in the Oregon Coast Range. The Oregon Gap II Project126
component still present. Scattered trees are either
and Oregon Vegetation Landscape-Level Cover Types127
evergreen conifers or deciduous broadleaves. Shrubs may
effort did not map this type; it was inadvertently
be absent, scattered, or very prominent, and include
aggregated with the agriculture classification. The
evergreen and deciduous broadleaf physiognomy.
Washington Gap project mapped this habitat as part of
Composition. The major native dominant bunchgrass is nonforested in the Woodland/Prairie Mosaic Zone. Other
Roemer’s fescue (Festuca idahoensis var. roemeri). Red fescue references describe elements of this habitat.7, 40, 41, 54, 99, 142, 211
(F. rubra) and California oatgrass (Danthonia californica) are
Natural Disturbance Regime. Historically, fire was a
frequently dominant or co-dominant on a local basis.
major component of this habitat. In addition to occasional
Long-stolon sedge (Carex inops) is occasionally co-
lightning strikes, fires were intentionally set by indigenous
dominant, especially in savannas and in the Columbia
inhabitants to maintain food staples such as camas and
Gorge. Slender wheatgrass (Elymus trachycaulus), blue
bracken fern.165 Although there is no definitive fire history
wildrye (Elymus glaucus), prairie junegrass (Koeleria
information, evidence suggests that many, if not most, of
macrantha), and Lemmon’s needlegrass (Stipa lemmonii)
these grasslands burned every few years. Annual soil
can be important locally. Major exotic dominant species
drought naturally eliminated or thinned invading trees
are colonial bentgrass (Agrostis capillaris), sweet
and promoted higher frequency fire regimes in the past.
vernalgrass (Anthoxanthum odoratum), Kentucky bluegrass
(Poa pratensis), tall oatgrass (Arrhenatherum elatius), Succession and Stand Dynamics. Historically, regular
medusahead (Taeniatherum caput-medusae), tall fescue (F. fires or extreme environmental conditions on the most
arundinacea), and soft brome (Bromus mollis). Common xeric sites prevented the establishment and continued
camas (Camassia quamash) is probably the most important growth of most woody vegetation, thereby maintaining
forb in terms of cover, but it rarely dominates. The bracken the grasslands and oak savannas. In some patches,
fern (Pteridium aquilinum) is sometimes co-dominant. A scattered oaks or even Douglas-fir survived long enough
rich diversity of native forbs is typical of sites in good to obtain some fire resistance and the frequent light fires
condition. then helped to maintain savannas.1, 41 Oaks were also able
Roemer’s fescue is distributed throughout the Puget to resprout if the above-ground stem was killed. High fire
Lowland and the Willamette Valley and in montane balds frequencies combined with digging of roots by Native
of the eastern and northeastern Olympics. Native red Americans could have favored the abundance of forbs
fescue is a major component neat saltwater in the northern over that of grasses in many areas of the pre-European
landscape.
 CHAPTER 2: HABITAT DESCRIPTIONS 43

Effects of Management and Anthropogenic Impacts. The 12. Ceanothus-Manzanita Shrublands

exclusion of fire from most of this habitat over the last Christopher B. Chappell & Jimmy Kagan
100+ years has resulted in profound changes. Oak savanna
has, for all practical purposes, disappeared from the
landscape.41 Douglas-fir encroachment, in the absence of
fire, is a “natural” process that occurs eventually on the
vast majority of westside grasslands, except perhaps on
the very driest sites. This encroachment leads to the
conversion of grasslands to forests. Fire exclusion has also
resulted in increases in shrub cover and the conversion of
some grasslands to shrublands. Exotic species are
prominent in this habitat and generally increase after
ground-disturbing activities like grazing or off-road
vehicle use. Scot’s broom, tall oatgrass, colonial bentgrass,
sweet vernalgrass, tall fescue, common velvetgrass (Holcus
lanatus), Kentucky bluegrass, soft brome, common St.
Johnswort (Hypericum perforatum), and hairy catsear
(Hypochaeris radicata) are among the most troublesome
Geographic Distribution. This habitat ranges from
species. The dominant native grass, Roemer’s fescue, can
southwestern Oregon south through much of California.
be eliminated with heavy grazing. Prescribed fire and
Within Oregon, it is primarily located in the Rogue and
other management tools have been used recently to control
Illinois valleys in Curry, Josephine, and Jackson counties;
Scot’s broom, Douglas-fir encroachment, and to attempt
it is also found scattered in the Siskiyou Mountains of the
to mimic historical conditions in some areas.78
same counties, in Douglas County, and in the southern
Status and Trends. This habitat is very rare and limited Cascades of Jackson and western Klamath counties.
in areal extent. In the southern Puget Sound area, only
Physical Setting. Climate is mostly very warm and
about 10% of the original area of the habitat is extant, and
relatively dry (about 17-30 inches [43-76 cm] mean annual
only 3% is dominated by native species.54 Overall decline
precipitation), but extends less commonly on serpentine
is significantly greater than these figures suggest because
or extremely dry sites into somewhat cooler and moister
the habitat is even more decimated and degraded
climates (up to 50 inches [127 cm] annual precipitation).
elsewhere. Causes of the decline are fire suppression,
Summers are very dry; winters are only slightly cool, much
conversion to agriculture and urban, and invasion of exotic
warmer than shrublands on the eastside. Primary
species. Most of what remains is dominated or co-
elevation range is about 1,000-2,000 ft (305-610 m), but
dominated by exotic species. Current trends are continued
extends up to a maximum of 5,000 ft (1,524 m).
decline both in area and condition. Ongoing threats
Topography is typified by mainly lower valley slopes and
include urban conversion, increase of exotic species,
foothills, but extends to nearly flat valley bottoms (where
ground disturbance via tracked vehicle use for military
before European settlement this habitat was a major type)
training, and effects of fire suppression. Eleven out of 12
and sporadically onto mountain slopes. This habitat tends
native plant associations representing this habitat listed
to occur on southern aspects when it does occur outside
for the National Vegetation Classification are considered
of low valleys. Soils are typically shallow to bedrock or
imperiled or critically imperiled.10
are derived from coarse alluvial deposits. Ultramatic
bedrock is a major parent material in the western Siskiyou
Mountains, whereas the eastern Siskiyou Mountains and
adjacent valleys are largely volcanic.
Landscape Setting. This habitat occurs adjacent to or in a
mosaic with Southwest Oregon Mixed Conifer-Hardwood
Forest, Westside Oak and Dry Douglas-fir Forest and
Woodlands, Agriculture, and rarely, Westside Grassland.
Urban is also adjacent in a few areas. Westside Riparian-
Wetlands habitat occurs as small inclusions within this
habitat. This habitat covers large areas only in lower
elevation valleys or on extensive areas of serpentine soils.
At moderate to high elevations it is mainly small patches
within a forest mosaic. Major land use of this habitat is
grazing and low-density residential development.
Structure. This habitat consists mainly of shrubland
dominated by sclerophyllous evergreen broadleaf shrubs,
but can also include grasslands with scattered tall shrubs.
 44 Wildlife–Habitat Relationships in Oregon and Washington

Deciduous broadleaf shrubs are less important, but are in species regenerate abundantly after fire from a long-lived
some cases dominant. The shrubs are mostly 3.3-13 ft (1-4 seedbank that is scarified by fire.187 Many seedlings die in
m) high. Shrub canopy ranges from very open to the first 3 years after fire.50 Wedge-leaf ceanothus can
completely closed. Herbaceous cover varies inversely with maintain prominence for >100 years.124 Shrub canopy cover
shrub canopy cover. Perennial bunchgrasses are the generally increases up to 30 years or so after the last fire,
dominant understory at sites in good condition, whereas and in the absence of another fire, the herbaceous
annual grasses dominate at sites in poorer condition. If understory can be reduced under dense late-successional
shrubs are not too dense, forbs are abundant. Historically, shrub canopies. Wedge-leaf ceanothus is considered a
many of these shrublands were probably grasslands with climax, or late-successional, dominant species on low-
scattered shrubs. Occasional conifers or broadleaf trees elevation dry sites in the Rogue Valley and on some
are sometimes scattered in the habitat. serpentine sites.14, 69 On many other sites, this habitat seems
to be maintained by occasional fires, and trees, especially
Composition. Sclerophyllous and hemi-sclerophyllous
oaks and ponderosa pine, will gradually increase in the
shrubs that dominate are, most commonly, wedge-leaf
absence of fire.
ceanothus (Ceanothus cuneatus) and white-leaf manzanita
(Arctostaphylos viscida), and less commonly, chaparral Effects of Management and Anthropogenic Impacts. Fire
whitethorn (Ceanothus leucodermis), blueblossom (C. suppression has probably increased the predominance of
thyrsiflorus), deerbrush (C. integerrimus), and deer oak dense, tall shrub stands versus a more open, patchy
(Quercus sadleriana). Wedge-leaf ceanothus is the most structure. It also seems to reduce the cover of bunchgrasses
abundant species at low elevations in the major valleys and forbs as stands become old. Grazing reduces native
and is the shrub most tolerant of xeric conditions. Other bunchgrasses in favor of exotics and/or the native
common, but not typically dominant shrubs include hairy rhizomatous California brome (Bromus carinatus). Exotic
manzanita (Arctostaphylos columbiana), pinemat manzanita species that have successfully invaded the understory of
(A. nevadensis), birchleaf mountain-mahogany (Cercocarpus this habitat are soft brome (Bromus mollis), medusahead
montanus var. glaber), Klamath plum (Prunus subcordata), (Taeniatherum caput-medusae), hedgehog dogtail (Cynosurus
bitter cherry (P. emarginata), chokecherry (P. virginiana), echinatus), and yellow star-thistle (Centaurea solstitialis).
Brewer ’s oak (Quercus garryana var. breweriana),
Status and Trends. This habitat occupies a small area
huckleberry oak (Q. vacciniifolia), California yerba-santa
within this region; it has declined considerably because
(Eriodictyon californicum), and bearbrush (Garrya fremontii).
of conversion to agriculture, residential development, and
The native bunchgrasses are Idaho fescue (Festuca
fire suppression. Most of this habitat has been degraded
idahoensis), California fescue (F. californica), California
by fire suppression, grazing, and exotic species invasions.
oatgrass (Danthonia californica), Lemmon’s needlegrass
This habitat is still declining in extent from development
(Stipa lemmonii), western needlegrass (S. occidentalis), and
pressures. One out of 7 Oregon plant associations listed
bluegrass (Poa secunda). Forb diversity is often high and
in the National Vegetation Classification is considered
common genera include Lilium, Calochortus, Fritillaria,
imperiled globally,10 but this type of vegetation has been
Microseris, Monardella, and Erigeron. One of several species
poorly studied in Oregon and there may be other
of oak (Quercus) or pine (Pinus), Douglas-fir (Pseudotsuga
associations.
menziesii), or incense cedar (Calocedrus decurrens) are
sometimes present as scattered individuals, especially on
less xeric sites.
Other Classifications and Key References. Franklin and
Dyrness88 referred to this habitat as sclerophyllous shrub
communities in the interior valleys of Oregon. It is also
called chaparral.125, 130, 141 The Oregon Gap II Project126 and
Oregon Vegetation Landscape-Level Cover Type127 that
would represent this type is Siskiyou Mountain serpentine
shrubland. Other references include describe aspects of
this habitat.69, 188
Natural Disturbance Regime. Fire is the major natural
disturbance. Fire regimes have not been studied in Oregon,
but in central California the fire-return interval has been
estimated at 30-60 years.84 Fire frequency may have been
greater during historic times based on chaparral fire
regimes in southern California.157 High-severity fires are
typical, with most shrubs being top-killed.
Succession and Stand Dynamics. Wedge-leaf ceanothus
and white-leaf manzanita are killed by fire. Some less
common shrubs sprout after fire. The dominant shrub
 CHAPTER 2: HABITAT DESCRIPTIONS 45

13. Western Juniper and Mountain communities are more common in depressions or steep
slopes with bunchgrass undergrowth. In the Great Basin,
Mahogany Woodlands mountain mahogany may form a distinct belt on mountain
Rex. C. Crawford & Jimmy Kagan slopes and ridgetops above pinyon-juniper woodland.
Mountain mahogany can occur in isolated, pure patches
that are often very dense. The primary land use is livestock
grazing.
Structure. This habitat is made up of savannas,
woodlands, or open forests with 10-60% canopy cover. The
tallest layer is composed of short (6.6-40 ft [2-12 m] tall)
evergreen trees. Dominant plants may assume a tall-shrub
growth form on some sites. The short trees appear in a
mosaic pattern with areas of low or medium-tall (usually
evergreen) shrubs alternating with areas of tree layers and
widely spaced low or medium-tall shrubs. The herbaceous
layer is usually composed of short or medium tall
bunchgrass or, rarely, a rhizomatous grass-forb
undergrowth. These vegetated areas can be interspersed
with rimrock or scree. A well-developed cryptogam layer
Geographic Distribution. This habitat is distributed from often covers the ground, although bare rock can make up
the Pacific Northwest south into southern California and much of the ground cover.
east to western Montana and Utah, where it often occurs Composition. Western juniper and/or mountain
with pinyon-juniper habitat. In Oregon and Washington, mahogany dominate these woodlands either with
this dry woodland habitat appears primarily in the bunchgrass or shrub-steppe undergrowth. Western juniper
Owyhee Uplands, High Lava Plains, and northern Basin (Juniperus occidentalis) is the most common dominant tree
and Range ecoregions. Secondarily, it develops in the in these woodlands. Part of this habitat will have curl-leaf
foothills of the Blue Mountains and East Cascades mountain mahogany (Cercocarpus ledifolius) as the only
ecoregions, and seems to be expanding into the southern dominant tall shrub or small tree. Mahogany may be co-
Columbia Basin ecoregion, where it was naturally found dominant with western juniper. Ponderosa pine (Pinus
in outlier stands. ponderosa) can grow in this habitat and in some rare
Western juniper woodlands with shrub-steppe species instances may be an important part of the canopy.
appear throughout the range of the habitat primarily in The most common shrubs in this habitat are basin,
central and southern Oregon. Many isolated mahogany Wyoming, or mountain big sagebrush (Artemisia tridentata
communities occur throughout canyons and mountains ssp. tridentata, ssp. wyomingensis, and ssp. vaseyana) and/
of eastern Oregon. Juniper-mountain mahogany or bitterbrush (Purshia tridentata). They usually provide
communities are found in the Ochoco and Blue Mountains. significant cover in juniper stands. Low or stiff sagebrush
Physical Setting. This habitat is widespread and variable, (Artemisia arbuscula or A. rigida) are dominant dwarf
occurring in basins and canyons, and on slopes and valley shrubs in some juniper stands. Mountain big sagebrush
margins in the southern Columbia Plateau, and on fire- appears most commonly with mountain mahogany and
protected sites in the northern Basin and Range province. mountain mahogany mixed with juniper. Snowbank
It may be found on benches and foothills. Western juniper shrubland patches in mountain mahogany woodlands are
and/or mountain mahogany woodlands are often found composed of mountain big sagebrush with bitter cherry
on shallow soils, on flats at mid- to high elevations, usually (Prunus emarginata), quaking aspen (Populus tremuloides),
on basalts. Other sites range from deep, loess soils and and serviceberry (Amelanchier alnifolia). Shorter shrubs
sandy slopes to very stony canyon slopes. At lower such as mountain snowberry (Symphoricarpos oreophilus)
elevations, or in areas outside of shrub-steppe, this habitat or creeping Oregongrape (Mahonia repens) can be
occurs on slopes and in areas with shallow soils. Mountain dominant in the undergrowth. Rabbitbrush
mahogany can occur on steep rimrock slopes, usually in (Chrysothamnus nauseosus and C. viscidiflorus) will increase
areas of shallow soils or protected slopes. This habitat can with grazing.
be found at elevations of 1,500- 8,000 ft (457-2,438 m), Part of this woodland habitat lacks a shrub layer.
mostly between 4,000-6,000 ft (1,220-1,830 m). Average Various native bunchgrasses dominate different aspects
annual precipitation ranges from approximately 10 to 13 of this habitat. Sandberg bluegrass (Poa sandbergii), a short
inches (25 to 33 cm), with most occurring as winter snow. bunchgrass, is the dominant and most common grass
throughout many juniper sites. Medium-tall bunchgrasses
Landscape Setting. This habitat reflects a transition such as Idaho fescue (Festuca idahoensis), bluebunch
between Ponderosa Pine Forest and Woodlands and wheatgrass (Pseudoroegneria spicata), needlegrasses (Stipa
Shrub-steppe, Eastside Grasslands, and rarely Desert occidentalis, S. thurberiana, S. lemmonii), bottlebrush
Playa and Salt Desert Scrub habitats. Western juniper squirreltail (Elymus elymoides) can dominate undergrowth.
generally occurs on higher topography, whereas the shrub
 46 Wildlife–Habitat Relationships in Oregon and Washington

Threadleaf sedge (Carex filifolia) and basin wildrye (Leymus Animals seeking shade under trees decrease or eliminate
cinereus) are found in lowlands and Geyer’s and Ross’ bunchgrasses and contribute to increasing cheatgrass
sedge (Carex geyeri, C. rossii), pinegrass (Calamagrostis cover.
rubescens), and blue wildrye (E. glaucus) appear on
Status and Trends. This habitat is dominated by fire-
mountain foothills. Sandy sites typically have needle-and-
sensitive species, and therefore, the range of western
thread (Stipa comata) and Indian ricegrass (Oryzopsis
juniper and mountain mahogany has expanded because
hymenoides). Cheatgrass (Bromus tectorum) or bulbous
of an interaction of livestock grazing and fire suppression.
bluegrass (Poa bulbosa) often dominate overgrazed or
Quigley and Arbelbide181 concluded that in the Inland
disturbed sites. In good condition this habitat may have
Pacific Northwest, Juniper/Sagebrush, Juniper
mosses growing under the trees.
Woodlands, and Mountain Mahogany cover types now
Other Classifications and Key References. This habitat are significantly greater in extent than before 1900.
is also called Juniper Steppe Woodland.136 The Oregon Gap Although it covers more area, this habitat is generally in
II Project126 and Oregon Vegetation Landscape-Level Cover degraded condition because of increased exotic plants and
Types127 that would represent this type are ponderosa pine- decreased native bunchgrasses. One third of Pacific
western juniper woodland, western juniper woodland, Northwest juniper and mountain mahogany community
and mountain mahogany shrubland. Other references types listed in the National Vegetation Classification are
describe this habitat.64, 79, 122, 207 considered imperiled or critically imperiled.10
Natural Disturbance Regime. Both mountain mahogany
and western juniper are fire intolerant. Under natural high- 14 Eastside Canyon Shrublands
frequency fire regimes both species formed savannas or Rex. C. Crawford & Jimmy Kagan
occurred as isolated patches on fire-resistant sites in shrub-
steppe or steppe habitat. Western juniper is considered a
topoedaphic climax tree in a number of
sagebrush-grassland, shrub-steppe, and drier conifer sites.
It is an increaser in many earlier seral communities in these
zones and invades without fires. Most trees >13 ft (4 m)
tall can survive low-intensity fires. The historic fire regime
of mountain mahogany communities varies with
community type and structure. The fire-return interval for
mountain mahogany (along the Salmon River in Idaho)
was 13-22 years until the early 1900’s and has increased
ever since. Mountain mahogany can live to 1,350 years in
western and central Nevada. Some old-growth mountain
mahogany stands avoid fire by growing on extremely
rocky sites.
Succession and Stand Dynamics. Juniper invades shrub-
Geographic Distribution. This habitat occurs primarily
steppe and steppe and reduces undergrowth productivity.
on steep canyon slopes in the Blue Mountains and the
Although slow seed dispersal delays recovery time,
margins of the Columbia Basin in Idaho, Oregon, and
western juniper can regain dominance in 30-50 years
Washington. This habitat also appears as isolated patches
following fire. A fire-return interval of 30-50 years typically
across Washington’s Columbia Basin.
arrests juniper invasion. The successional role of curl-leaf
mountain mahogany varies with community type. Physical Setting. This habitat develops in hot dry climates
Mountain brush communities where curl-leaf mountain in the Pacific Northwest. Annual precipitation totals 12-
mahogany is either dominant or co-dominant are 20 inches (31-51 cm); only 10% falls in the hottest months,
generally stable and successional rates are slow. July through September. Snow accumulation is low (1-6
inches [3-15 cm]), persisting only a few weeks. Sites are
Effects of Management and Anthropogenic Impacts.
generally steep (>60%) on all aspects but most common
Over the past 150 years, with fire suppression,
on northerly aspects in deep, dry canyons. Columbia River
overgrazing, and changing climatic factors, western
basalt is the major geologic substrate although many sites
juniper has increased its range into adjacent shrub-steppe,
are underlain with loess deposits mixed with colluvium.
grasslands, and savannas. Increased density of juniper and
Steep northerly aspects in the Palouse Hills can also
reduced fine fuels from an interaction of grazing and
support this habitat. This habitat is found from 500 to 5,000
shading result in high severity fires that eliminate woody
ft (152 to 1,524 m) in elevation.
plants and promote herbaceous cover, primarily annual
grasses. Diverse mosses and lichens occur on the ground Landscape Setting. This habitat is generally found in steep
in this type if it has not been too disturbed by grazing. canyons surrounded by the Eastside Grassland Habitat
Excessive grazing will decrease bunchgrasses and increase and below or in a mosaic with the Ponderosa Pine Forest
exotic annual grasses plus various native and exotic forbs. and Woodland habitat. This habitat can develop near talus
 CHAPTER 2: HABITAT DESCRIPTIONS 47

slopes, at the heads of dry drainages, and toe slopes in Natural Disturbance Regime. This habitat is within the
moist shrub-steppe and steppe zones. At lower elevation sagebrush and bunchgrass vegetation type of Barrett et
sites, these are more often in a mix with bluebunch al.22 who concluded it had a fire-return interval of 25 years.
wheatgrass, dry rocky grasslands, and low-elevation Canyon shrublands associated with talus burn less
riparian habitats. The primary surrounding land use is frequently but are subject to talus movement. Similar
livestock grazing. shrubfields are associated with forest landscapes and are
early seral stages of the Eastside Mixed Conifer Forest
Structure. The Eastside Canyon Shrubland habitat is
Habitat.
generally a mix of tall (5 ft [1.5 m]) to medium (1.6 ft [0.5
m]) deciduous shrublands in a mosaic with bunchgrass Succession and Stand Dynamics. Many of the major
or annual grasslands. Shrub canopies are almost always shrubs sprout following fire and will be maintained with
closed (>60% cover), forming a thicket of interwoven stems moderate fire frequency. Most thickets will increase in size
and branches. Shrub layers can be one or two-tiered but without fire. This habitat has increased primarily in moist
often are so dense they restrict the herbaceous layer to steppe and shrub-steppe habitat with fire suppression and
shade-tolerant rhizomatous species. restricted grazing. Prolonged fire suppression may lead
to invasions by tree species. Apparently some
Composition. Mallowleaf ninebark (Physocarpus
representatives of this habitat could potentially support
malvaceus), a major dominant, bitter cherry (Prunus
Douglas-fir or ponderosa pine woodlands after a long fire-
emarginata), chokecherry (Prunus virginiana), oceanspray
free period.
(Holodiscus discolor) or Rocky Mountain maple (Acer
glabrum) are the most common tall shrubs in this habitat. Effects of Management and Anthropogenic Impacts.
In moist areas, black hawthorn (Crataegus douglasii) may Livestock grazing in adjacent grassland or shrub-steppe
appear and can dominate some sites as a tall shrub or small habitat changes the surrounding fine-fuel matrix for fire.
tree. Other tall shrubs such as syringa (Philadelphus lewisii) That, combined with fire suppression, leads to a change
or serviceberry (Amelanchier alnifolia) often dominate sites in habitat patch size, structure, and composition. In
associated with talus. Common medium-tall shrubs are response to fire suppression, shrub thickets on northerly
common snowberry (Symphoricarpos albus), rose (Rosa aspects near lower treeline tend to increase in patch size
nutkana, R. woodsii), smooth sumac (Rhus glabra), and and height and are invaded by tree species. With heavy
currants (Ribes spp.). Basin or Wyoming big sagebrush livestock grazing, shrubs are browsed, broken, and
(Artemisia tridentata ssp. tridentata or A. t. ssp. trampled, which eventually creates a more open shrubland
wyomingensis), along with rabbitbrush (Chrysothamnus with a more abundant herbaceous layer.
spp.), may be important members of these thickets in
Status and Trends. The Eastside Canyon Shrubland
weedy sites, dry areas, or transitions with grasslands.
habitat is restricted in range and probably has increased
Scattered ponderosa pine (Pinus ponderosa), black
locally in area. Johnson and Simon123 reported increases
cottonwood (Populus balsamifera ssp. trichocarpa) and rarely
in common snowberry-rose communities as a response to
Douglas-fir (Pseudotsuga menziesii) trees may be found in
fire suppression and heavy grazing that depleted
and adjacent to this habitat.
bunchgrass cover. One of the three Eastside Canyon
Idaho fescue (Festuca idahoensis), bluebunch wheatgrass
Shrubland community types in the National Vegetation
(Pseudoroegneria spicata), Thurber’s needlegrass (Stipa
Classification is considered imperiled.10
thurberiana), and Sandberg’s bluegrass (Poa sandbergii)
found in the surrounding steppe or shrub-steppe are
common but never abundant in these thickets. Basin
wildrye (Leymus cinereus) can be locally important.
Kentucky bluegrass (Poa pratensis) is a common introduced
grass and, where grazed by livestock, is a dominant
undergrowth species. Annual grasses (Bromus tectorum,
B. briziformis) can be abundant especially on disturbed dry
sites. Cleavers (Galium aparine) is a frequent member of
the herbaceous component of this habitat. Other common
forbs include red avens (Geum triflorum), horsemint
(Agastache urticifolia), sticky cinquefoil (Potentilla gracilis),
balsamroots (Balsamorhiza spp.), and fleabanes (Erigeron
spp.).
Other Classifications and Key References. This habitat
is called shrub garland88 or talus thickets. The Oregon Gap
II Project126 and Oregon Vegetation Landscape-Level Cover
Type127 that would represent this type is eastside big
sagebrush shrubland. Other references describe this
habitat.60, 122, 123, 207
 48 Wildlife–Habitat Relationships in Oregon and Washington

15. Eastside Grassslands riparian bottomlands dominated by non-native grasses.

Rex. C. Crawford & Jimmy Kagan This habitat is found from 500 to 6,000 ft (152-1,830 m) in
elevation.
Landscape Setting. Eastside grassland habitats appear
well below and in a matrix with lower treeline Ponderosa
Pine Forest and Woodlands or Western Juniper and
Mountain Mahogany Woodlands. It can also be part of
the lower elevation forest matrix. Most grassland habitat
occurs in two distinct large landscapes: plateau and
canyon grasslands. Several rivers flow through narrow
basalt canyons below plateaus supporting prairies or
shrub-steppe. The canyons can be some 2,132 ft (650 m)
deep below the plateau. The plateau above is composed
of gentle slopes with deep silty loess soils in an expansive
rolling dune-like landscape. Grasslands may occur in a
patchwork with shallow soil scablands or within biscuit
scablands or mounded topography. Naturally occurring
grasslands are beyond the range of bitterbrush and
Geographic Distribution. This habitat is found primarily sagebrush species. This habitat exists today in the shrub-
in the Columbia Basin of Idaho, Oregon, and Washington, steppe landscape where grasslands are created by brush
at mid- to low elevations and on plateaus in the Blue removal, chaining or spraying, or by fire. Agricultural uses
Mountains, usually within the ponderosa pine zone in and introduced perennial plants on abandoned or planted
Oregon. fields are common throughout the current distribution of
Idaho fescue grassland habitats were formerly eastside grassland habitats.
widespread in the Palouse region of southeastern
Structure. This habitat is dominated by short to medium-
Washington and adjacent Idaho; most of this habitat has
tall grasses (<3.3 ft [1 m]). Total herbaceous cover can be
been converted to agriculture. Idaho fescue grasslands still
closed to only sparsely vegetated. In general, this habitat
occur in isolated, moist sites near lower treeline in the
is an open and irregular arrangement of grass clumps
foothills of the Blue Mountains, the Northern Rockies, and
rather than a continuous sod cover. These medium-tall
east Cascades near the Columbia River Gorge. Bluebunch
grasslands often have scattered and diverse patches of low
wheatgrass grassland habitats are common throughout
shrubs, but few or no medium-tall shrubs (<10% cover of
the Columbia Basin, both as modified native grasslands
shrubs are taller than the grass layer). Native forbs may
in deep canyons and the dry Palouse and as fire-induced
contribute significant cover or they may be absent.
representatives in the shrub-steppe. Similar grasslands
Grasslands in canyons are dominated by bunchgrasses
appear on the High Lava Plains ecoregion, where they
growing in lower densities than on deep-soil prairie sites.
occur in a matrix with big sagebrush or juniper woodlands.
The soil surface between perennial plants can be covered
In Oregon they are also found in burned shrub-steppe and
with a diverse cryptogamic or microbiotic layer of mosses,
canyons in the Basin and Range and Owyhee Uplands.
lichens, and various soil bacteria and algae. Moister
Sand dropseed and three-awn needlegrass grassland
environments can support a dense sod of rhizomatous
habitats are restricted to river terraces in the Columbia
perennial grasses. Annual plants are a common spring and
Basin, Blue Mountains, and Owyhee Uplands of Oregon
early summer feature of this habitat.
and Washington. Primary location of this habitat extends
along the Snake River from Lewiston south to the Owyhee Composition. Bluebunch wheatgrass (Pseudoroegneria
River. spicata) and Idaho fescue (Festuca idahoensis) are the
characteristic native bunchgrasses of this habitat and either
Physical Setting. This habitat develops in hot, dry climates
or both can be dominant. Idaho fescue is common in more
in the Pacific Northwest. Annual precipitation totals 8-20
moist areas and bluebunch wheatgrass more abundant in
inches (20-51 cm); only 10% falls in the hottest months,
drier areas. Rough fescue (F. campestris) is a characteristic
July through September. Snow accumulation is low (1-6
dominant on moist sites in northeastern Washington. Sand
inches [3-15 cm]) and occurs only in January and February
dropseed (Sporobolus cryptandrus) or three-awn (Aristida
in eastern portions of its range and November through
longiseta) are native dominant grasses on hot dry sites in
March in the west. More snow accumulates in grasslands
deep canyons. Sandberg bluegrass (Poa sandbergii) is
within the forest matrix. Soils are variable: (1) highly
usually present, and occasionally codominant in drier
productive loess soils up to 51 inches (130 cm) deep, (2)
areas. Bottlebrush squirreltail (Elymus elymoides) and
rocky flats, (3) steep slopes, and (4) sandy, gravel or cobble
Thurber needlegrass (Stipa thurberiana) can be locally
soils. An important variant of this habitat occurs on sandy,
dominant. Annual grasses are usually present; cheatgrass
gravelly, or silty river terraces or seasonally exposed river
(Bromus tectorum) is the most widespread. In addition,
gravel or Spokane flood deposits. The grassland habitat
medusahead (Taeniatherum caput-medusae), and other
is typically upland vegetation but it may also include
 CHAPTER 2: HABITAT DESCRIPTIONS 49

annual bromes (Bromus commutatus, B. mollis, B. japonicus) grazing or repeated early season fires can result in annual
may be present to co-dominant. Moist environments, grasslands of cheatgrass, medusahead, knapweed, or
including riparian bottomlands, are often co-dominated yellow star-thistle. Annual exotic grasslands are common
by Kentucky bluegrass (Poa pratensis). in dry grasslands and are included in modified grasslands
A dense and diverse forb layer can be present or entirely as part of the Agriculture habitat.
absent; >40 species of native forbs can grow in this habitat
Effects of Management and Anthropogenic Impacts.
including balsamroots (Balsamorhiza spp.), biscuitroots
Large expanses of grasslands are currently used for
(Lomatium spp.), buckwheat (Eriogonum spp.), fleabane
livestock ranching. Deep soil Palouse sites are mostly
(Erigeron spp.), lupines (Lupinus spp.), and milkvetches
converted to agriculture. Drier grasslands and canyon
(Astragalus spp.). Common exotic forbs that can grow in
grasslands, those with shallower soils, steeper topography,
this habitat are knapweeds (Centaurea solstitialis, C. diffusa,
or hotter, drier environments, were more intensively
C. maculosa), tall tumblemustard (Sisymbrium altissimum),
grazed and for longer periods than were deep-soil
and Russian thistle (Salsola kali).
grasslands.207 Evidently, these drier native bunchgrass
Smooth sumac (Rhus glabra) is a deciduous shrub locally
grasslands changed irreversibly to persistent annual grass
found in combination with these grassland species.
and forblands. Some annual grassland, native bunchgrass,
Rabbitbrushes (Chrysothamnus nauseosus, C. viscidiflorus)
and shrub-steppe habitats were converted to intermediate
can occur in this habitat in small amounts, especially where
wheatgrass, or more commonly, crested wheatgrass
grazed by livestock. In moist Palouse regions, common
(Agropyron cristatum)-dominated areas. Apparently, these
snowberry (Symphoricarpos albus) or Nootka rose (Rosa
form persistent grasslands and are included as modified
nutkana) may be present, but is shorter than the
grasslands in the Agriculture habitat. With intense
bunchgrasses. Dry sites contain low succulent pricklypear
livestock use, some riparian bottomlands become
(Opuntia polyacantha). Big sagebrush (Artemisia tridentata)
dominated by non-native grasses. Many native dropseed
is occasional and may be increasing in grasslands on
grasslands have been submerged by dam reservoirs.
former shrub-steppe sites. Black hawthorn (Crataegus
douglasii) and other tall shrubs can form dense thickets Status and Trends. Most of the Palouse prairie of
near Idaho fescue grasslands. Rarely, ponderosa pine southeastern Washington and adjacent Idaho and Oregon
(Pinus ponderosa) or western juniper (Juniperus occidentalis) has been converted to agriculture. Remnants still occur in
can occur as isolated trees. the foothills of the Blue Mountains and in isolated, moist
Columbia Basin sites. The Palouse is one of the most
Other Classifications and Key References. This habitat
endangered ecosystems in the U.S.166 with only 1% of the
is called Palouse Prairie, Pacific Northwest grassland,
original habitat remaining; it is highly fragmented with
steppe vegetation, or bunchgrass prairie in general
most sites <10 acres. All these areas are subject to weed
ecological literature. Quigley and Arbelbide181 called this
invasions and drift of aerial biocides. Since 1900, 94% of
habitat Fescue-Bunchgrass and Wheatgrass Bunchgrass
the Palouse grasslands have been converted to crop, hay,
and the dry Grass cover type. The Oregon Gap II Project126
or pasture lands. Quigley and Arbelbide181 concluded that
and Oregon Vegetation Landscape-Level Cover Types127
Fescue-Bunchgrass and Wheatgrass bunchgrass cover
that would represent this type are northeast Oregon
types have significantly decreased in area since before1900,
canyon grassland, forest-grassland mosaic, and modified
while exotic forbs and annual grasses have significantly
grassland; Washington Gap37 types 13, 21, 22, 24, 29-31,
increased since pre-1900. Fifty percent of the plant
82, and 99 map this habitat. Kuchler136 includes this within
associations recognized as components of Eastside
Fescue-wheatgrass and wheatgrass-bluegrass. Franklin
Grasslands habitat listed in the National Vegetation
and Dyrness88 include this habitat in steppe zones of
Classification are considered imperiled or critically
Washington and Oregon. Other references describe this
imperiled.10
habitat.1, 28, 60, 159, 166, 206, 207
Natural Disturbance Regime. The fire-return interval for
sagebrush and bunchgrass is estimated at 25 years.22 The
native bunchgrass habitat apparently lacked extensive
herds of large grazing and browsing animals until the late
1800s. Burrowing animals and their predators likely
played important roles in creating small-scale patch
patterns.
Succession and Stand Dynamics. Currently fires burn less
frequently in the Palouse grasslands than historically
because of fire suppression, roads, and conversions to
cropland.159 Without fire, black hawthorn shrubland
patches expand on slopes along with common snowberry
and rose. Fires covering large areas of Shrub-steppe habitat
can eliminate shrubs and their seed sources and create
Eastside Grasslands habitat. Fires that follow heavy
 50 Wildlife–Habitat Relationships in Oregon and Washington

16. Shrub-steppe Juniper and Mountain Mahogany Woodland habitats. It

Rex. C. Crawford & Jimmy Kagan forms mosaic landscapes with these woodland habitats
and Eastside Grasslands, Dwarf Shrub-steppe, and Desert
Playa and Salt Scrub habitats. Mountain sagebrush shrub-
steppe occurs at high elevations occasionally within the
dry Eastside Mixed Conifer Forest and Montane Mixed
Conifer Forest habitats. Shrub-steppe habitat can appear
in large landscape patches. Livestock grazing is the
primary land use in the shrub-steppe although much has
been converted to irrigation or dry land agriculture. Large
areas occur in military training areas and wildlife refuges.
Structure. This habitat is a shrub savanna or shrubland
with shrub coverage of 10-60%. In an undisturbed
condition, shrub cover varies between 10 and 30%. Shrubs
are generally evergreen although deciduous shrubs are
prominent in many habitats. Shrub height typically is
medium-tall (1.6-3.3 ft [0.5-1.0 m]) although some sites
Geographic Distribution. Shrub-steppe habitats are support shrubs approaching 9 ft (2.7 m) tall. Vegetation
common across the Columbia Plateau of Washington, structure in this habitat is characteristically an open shrub
Oregon, Idaho, and adjacent Wyoming, Utah, and Nevada. layer over a moderately open to closed bunchgrass layer.
They extend up into the cold, dry environments of The more northern or productive sites generally have a
surrounding mountains. denser grass layer and sparser shrub layer than southern
Basin big sagebrush shrub-steppe occurs along stream or more xeric sites. In fact, the rare good-condition site is
channels, in valley bottoms and flats throughout eastern better characterized as grassland with shrubs than a
Oregon and Washington. Wyoming sagebrush shrub- shrubland. The bunchgrass layer may contain a variety of
steppe is the most widespread habitat in eastern Oregon forbs. Good-condition habitat has very little exposed bare
and Washington, occurring throughout the Columbia ground, and has mosses and lichens carpeting the area
Plateau and the northern Great Basin. Mountain big between taller plants. However, heavily grazed sites have
sagebrush shrub-steppe occurs throughout the mountains dense shrubs making up >40% cover, with introduced
of the eastern Oregon and Washington. Bitterbrush shrub- annual grasses and little or no moss or lichen cover. Moist
steppe appears primarily along the eastern slope of the sites may support tall bunchgrasses (>3.3 ft [1 m]) or
Cascades, from north-central Washington to California rhizomatous grasses. More southern shrub-steppe may
and occasionally in the Blue Mountains. Three-tip have native low shrubs dominating with bunchgrasses.
sagebrush shrub-steppe occurs mostly along the northern
Composition. Characteristic and dominant mid-tall
and western Columbia Basin in Washington and
shrubs in the shrub-steppe habitat include all three
occasionally appears in the lower valleys of the Blue
subspecies of big sagebrush, basin (Artemisia tridentata ssp.
Mountains and in the Owyhee Upland ecoregions of
tridentata), Wyoming (A. t. ssp. wyomingensis) or mountain
Oregon. Interior shrub dunes and sandy steppe and shrub-
(A. t. ssp. vaseyana), antelope bitterbrush (Purshia
steppe is concentrated at low elevations near the Columbia
tridentata), and two shorter sagebrushes, silver (A. cana)
River and in isolated pockets in the Northern Basin and
and three-tip (A. tripartita). Each of these species can be
Range and Owyhee Uplands. Bolander silver sagebrush
the only shrub or appear in complex seral conditions with
shrub-steppe is common in southeastern Oregon.
other shrubs. Common shrub complexes are bitterbrush
Mountain silver sagebrush is more prevalent in the Oregon
and Wyoming big sagebrush, bitterbrush and three-tip
East Cascades and in montane meadows in the southern
sagebrush, Wyoming big sagebrush and three-tip
Ochoco and Blue Mountains.
sagebrush, and mountain big sagebrush and silver
Physical Setting. Generally, this habitat is associated with sagebrush. Wyoming and mountain big sagebrush can
dry, hot environments in the Pacific Northwest although codominate areas with tobacco brush (Ceanothus velutinus).
variants are in cool, moist areas with some snow Rabbitbrush (Chrysothamnus viscidiflorus) and short-spine
accumulation in climatically dry mountains. Elevation horsebrush (Tetradymia spinosa) are common associates and
range is wide (300-9,000 ft [91-2,743 m]) with most habitat often dominate sites after disturbance. Big sagebrush
occurring between 2,000 and 6,000 ft (610-1,830 m). Habitat occurs with the shorter stiff sagebrush (A. rigida) or low
occurs on deep alluvial, loess, silty or sandy-silty soils, sagebrush (A. arbuscula) on shallow soils or high elevation
stony flats, ridges, mountain slopes, and slopes of lake sites. Many sandy areas are shrub-free or are open to
beds with ash or pumice soils. patchy shrublands of bitterbrush and/or rabbitbrush.
Silver sagebrush is the dominant and characteristic shrub
Landscape Setting. Shrub-steppe habitat defines a
along the edges of stream courses, moist meadows, and
biogeographic region and is the major vegetation on
ponds. Silver sagebrush and rabbitbrush are associates in
average sites in the Columbia Plateau, usually below
disturbed areas.
Ponderosa Pine Forest and Woodland, and Western
 CHAPTER 2: HABITAT DESCRIPTIONS 51

When this habitat is in good or better ecological postfire moisture, and plant competition. Three-tip
condition a bunchgrass steppe layer is characteristic. sagebrush is a climax species that reestablishes (from seeds
Diagnostic native bunchgrasses that often dominate or commonly from sprouts) within 5-10 years following a
different shrub-steppe habitats are (1) mid-grasses: disturbance. Certain disturbance regimes promote three-
bluebunch wheatgrass (Pseudoroegneria spicata), Idaho tip sagebrush and it can out-compete herbaceous species.
fescue (Festuca idahoensis), bottlebrush squirreltail (Elymus Bitterbrush is a climax species that plays a seral role
elymoides), and Thurber needlegrass (Stipa thurberiana); (2) colonizing by seed onto rocky and/or pumice soils.
short grasses: threadleaf sedge (Carex filifolia) and Bitterbrush may be declining and may be replaced by
Sandberg bluegrass (Poa sandbergii); and (3) the tall grass, woodlands in the absence of fire. Silver sagebrush is a
basin wildrye (Leymus cinereus). Idaho fescue is climax species that establishes during early seral stages
characteristic of the most productive shrub-steppe and coexists with later arriving species. Big sagebrush,
vegetation. Bluebunch wheatgrass is codominant at xeric rabbitbrush, and short-spine horsebrush invade and can
locations, whereas western needlegrass (Stipa occidentalis), form dense stands after fire or livestock grazing. Frequent
long-stolon (Carex inops) or Geyer’s sedge (C. geyeri) or high-intensity fire can create a patchy shrub cover or
increase in abundance in higher elevation shrub-steppe can eliminate shrub cover and create Eastside Grasslands
habitats. Needle-and-thread (Stipa comata) is the habitat.
characteristic native bunchgrass on stabilized sandy soils.
Effects of Management and Anthropogenic Impacts.
Indian ricegrass (Oryzopsis hymenoides) characterizes
Shrub density and annual cover increase, whereas
dunes. Grass layers on montane sites contain slender
bunchgrass density decreases, with livestock use.
wheatgrass (Elymus trachycaulus), mountain fescue (F.
Repeated or intense disturbance, particularly on drier sites,
brachyphylla), green fescue (F. viridula), Geyer’s sedge, or
leads to cheatgrass dominance and replacement of native
tall bluegrasses (Poa spp.). Bottlebrush squirreltail can be
bunchgrasses. Dry and sandy soils are sensitive to grazing,
locally important in the Columbia Basin, sand dropseed
with needle-and-thread replaced by cheatgrass at most
(Sporobolus cryptandrus) is important in the Basin and
sites. These disturbed sites can be converted to modified
Range and basin wildrye is common in the more alkaline
grasslands in the Agriculture habitat.
areas. Nevada bluegrass (Poa secunda), Richardson muhly
(Muhlenbergia richardsonis), or alkali grass (Puccinella spp.) Status and Trends. Shrub-steppe habitat still dominates
can dominate silver sagebrush flats. Many sites support most of southeastern Oregon although half of its original
non-native plants, primarily cheatgrass (Bromus tectorum) distribution in the Columbia Basin has been converted to
or crested wheatgrass (Agropyron cristatum) with or agriculture. Alteration of fire regimes, fragmentation,
without native grasses. Shrub-steppe habitat, depending livestock grazing, and the addition of >800 exotic plant
on site potential and disturbance history, can be rich in species have changed the character of shrub-steppe
forbs or have little forb cover. Trees may be present in some habitat. Quigley and Arbelbide181 concluded that Big
shrub-steppe habitats, usually as isolated individuals from Sagebrush and Mountain Sagebrush cover types are
adjacent forest or woodland habitats. significantly smaller in area than before 1900, and that
Bitterbrush/Bluebunch Wheatgrass cover type is similar
Other Classifications and Key References. This habitat
to the pre-1900 extent. They concluded that Basin Big
is called Sagebrush steppe and Great Basin sagebrush by
Sagebrush and Big Sagebrush-Warm potential vegetation
Kuchler.136 The Oregon Gap II Project 126 and Oregon
type’s successional pathways are altered, that some
Vegetation Landscape-Level Cover Types127 that would
pathways of Antelope Bitterbrush are altered and that
represent this type are big sagebrush shrubland, sagebrush
most pathways for Big Sagebrush-Cool are unaltered.
steppe, and bitterbrush-big sagebrush shrubland. Franklin
Overall this habitat has seen an increase in exotic plant
and Dyrness88 discussed this habitat in shrub-steppe zones
importance and a decrease in native bunchgrasses. More
of Washington and Oregon. Other references describe this
than half of the Pacific Northwest shrub-steppe habitat
habitat. 60, 116, 122, 123, 212, 224, 225
community types listed in the National Vegetation
Natural Disturbance Regime. Barrett et al.22 concluded Classification are considered imperiled or critically
that the fire-return interval for this habitat is 25 years. The imperiled.10
native shrub-steppe habitat apparently lacked extensive
herds of large grazing and browsing animals until the late
1800s. Burrowing animals and their predators likely
played important roles in creating small-scale patch
patterns.
Succession and Stand Dynamics. With disturbance,
mature stands of big sagebrush are reinvaded through
soil-stored or windborne seeds. Invasion can be slow
because sagebrush is not disseminated over long distances.
Site dominance by big sagebrush usually takes a decade
or more depending on fire severity and season, seed rain,
 52 Wildlife–Habitat Relationships in Oregon and Washington

17. Dwarf Shrub-steppe occasionally occur in a mosaic with big sagebrush, stiff
Rex C. Crawford & Jimmy Kagan sagebrush, or black sagebrush steppe or within lower
treeline woodlands. Stiff sagebrush stands may also be
extensive, but usually occur in a mosaic with grassland,
big sagebrush or occasionally in juniper (Juniperus
occidentalis) or Ponderosa pine (Pinus ponderosa)
woodlands. Black sagebrush stands are extensive and may
occur in a mosaic with low sagebrush or mountain or
Wyoming big sagebrush.
Structure. These low shrub (<1.6 ft [0.5 m] high)
communities have an undergrowth of short grasses and
forbs with extensive exposed rock and cryptogamic crusts.
More productive sites have an open, native medium-tall
bunchgrass layer with scattered low shrubs. Some
scablands in the Columbia Basin have few to no dwarf
shrubs and the habitat is entirely dominated by grasses
and forbs. Total vegetation cover is open to sparse.
Individual trees can appear among the low shrubs when
Geographic Distribution. Dwarf-shrub and related this habitat appears in the forest matrix.
scabland habitats are located throughout the Columbia
Composition. Several dwarf-shrub species characterize
Plateau and in adjacent woodland and forest habitats.
this habitat: low sagebrush (Artemisia arbuscula), black
They are more common in southern Oregon than in
sagebrush (A. nova), stiff sagebrush (A. rigida), or several
Washington.
shrubby buckwheat species (Eriogonum douglasii, E.
Low sagebrush steppe is common in the Basin and
sphaerocephalum, E. strictum, E. thymoides, E. niveum, E.
Range and the Owyhee Uplands in eastern Lake, Harney,
compositum). These dwarf-shrub species can be found as
and Malheur counties and is a minor type in eastern
the sole shrub species or in combination with these or other
Washington and northeastern Oregon. It usually occurs
low shrubs. Purple sage (Saliva dorrii) can dominate
on low, scabby plateaus above lake basins. Stiff sagebrush/
scablands on steep sites with rock outcrops.
Sandberg bluegrass is a major type widely distributed in
Sandberg bluegrass (Poa sandbergii) is the characteristic
the Columbia Basin, particularly associated with the
and sometimes the dominant grass making up most of
channeled scablands, High Lava Plains, and in isolated
this habitat’s sparse vegetative cover. Taller bluebunch
spots throughout the Blue Mountains and the Palouse.
wheatgrass (Pseudoroegneria spicata) or Idaho fescue
Black sagebrush steppe is not found in Washington and is
(Festuca idahoensis) grasses may occur on the most
rare in Oregon, occurring along the Nevada border in
productive sites with Sandberg bluegrass. Bottlebrush
southern Lake, Harney, and Malheur counties, in the
squirreltail (Elymus elymoides) and Thurber needlegrass
southern Basin and Range and Owyhee Uplands
(Stipa thurberiana) are typically found in low cover areas,
Physiographic Province.
although they can dominate some sites. One-spike
Physical Setting. This habitat appears on sites with little oatgrass (Danthonia unispicata), prairie junegrass (Koeleria
soil development that often have extensive areas of macrantha), and Henderson ricegrass (Achnatherum
exposed rock, gravel, or compacted soil. The habitat is hendersonii) are occasionally important. Exotic annual
characteristically associated with flats, plateaus, or gentle grasses, commonly cheatgrass (Bromus tectorum), increase
slopes although steep slopes with rock outcrops are with heavy disturbance and can be locally abundant.
common. Scabland types within the shrub-steppe area Common forbs include serrate balsamroot (Balsamorhiza
occur on barren, usually fairly young basalts or shallow serrata), Oregon twinpod (Physaria oregana), Oregon
loam over basalt <12 inches (30 cm) deep. In woodland or bitterroot (Lewisia rediviva), big-head clover (Trifolium
forest mosaics, scabland soils are deeper (still <26 inches macrocephalum), and Rainier violet (Viola trinervata). Several
[65 cm]) but too droughty or extreme soils for tree growth. other forbs (Arenaria, Collomia, Erigeron, Lomatium, and
Topoedaphic drought is the major process influencing Phlox spp.) are characteristic, early blooming species. A
these communities on ridge tops and gentle slopes around diverse lichen and moss layer is a prominent component
ridgetops. Spring flooding is characteristic of scablands of these communities.
in concave topographic positions. This habitat is found Medium-tall shrubs, such as big sagebrush (Artemisia
across a wide range of elevations from 500 to 7,000 ft (152 tridentata), Silver sagebrush (A. cana), antelope bitterbrush
to 2,134 m). (Purshia tridentata), and rabbitbrush (Chrysothamnus spp.)
occasionally appear in these scablands.
Landscape Setting. These scabland habitats form a mosaic
with Shrub-steppe habitat, Eastside Grasslands habitat, Other Classifications and Key References. This habitat
and with Western Juniper and Mountain Mahogany is called scabland, biscuit-swale topography, lithosolic
Woodland or Ponderosa Pine Forest and Woodland steppe, or low shrub-steppe. Quigley and Arbelbide181
habitats. Low sagebrush stands are often extensive and called this habitat low sagebrush cover type and “Low
 CHAPTER 2: HABITAT DESCRIPTIONS 53

Sagebrush-Xeric” and “Low Sagebrush-Mesic” potential 18. Desert Playa and Salt Scrub
vegetation groups. The Oregon Gap II Project126 and
Oregon Vegetation Landscape-Level Cover Type127 that
Shrublands
would represent this type is low-dwarf sagebrush. Rex C. Crawford & Jimmy Kagan
Kuchler136 did not distinguish this habitat but included it
within Sagebrush Steppe. Franklin and Dyrness 88
discussed this habitat as lithosolic sites in steppe and
shrub-steppe zones of Washington and as plant
associations in steppe and shrub-steppe zones of central
and southern Oregon. Other references describe this
habitat.60, 64, 122, 123, 207
Natural Disturbance Regime. Scabland habitats often do
not have enough vegetation cover to support wildfires.
Bunchgrass sites with black or low sagebrush may burn
enough to damage shrubs and decrease shrub cover with
repetitive burns. Many scabland sites have poorly drained
soil and because of shallow soil are prone to winter
flooding. Freezing of saturated soil results in “frost-
heaving” that churns the soil and is a major disturbance
factor in vegetation patterns. Stiff sagebrush is a preferred Geographic Distribution. The desert playa and salt scrub
browse for elk as well as livestock. Native ungulates use habitat centers on the Great Basin of Nevada and Utah. In
scablands in early spring and contribute to churning of the Pacific Northwest, it is most common and abundant
the soil surface. in the larger, alkaline lake basins in southeastern Oregon,
Succession and Stand Dynamics. Grazing reduces the although it is represented throughout the Columbia
cover of bunchgrasses and increases the abundance of Plateau, Basin and Range, and Owyhee Provinces.
common yarrow (Achillea millefolium), phlox species, Shadscale salt desert shrub and mixed salt desert shrub
bighead clover, serrate balsamroot, bottlebrush squirreltail range from southeastern Oregon south into Utah and
and annual bromes on dwarf shrublands. Increased Nevada. Black greasewood salt desert scrub and alkaline/
ground disturbing activities increases exotic plant saline bottomland grasslands and wetlands appear
abundance, particularly on deeper soil sites. All dwarf- throughout the Columbia Plateau of Washington and
shrub species are intolerant of fire and do not sprout. Oregon.
Consequently, redevelopment of dwarf shrub-steppe Physical Setting. This habitat typically occupies the lowest
habitat is slow following fire or any disturbance that elevations in hydrologic basins in the driest regions of the
removes shrubs. Pacific Northwest. Elevation range is highly variable, from
Effects of Management and Anthropogenic Impacts. 3,000 to 7,500 ft (914 to 2.286 m) in southeastern Oregon
Scabland habitats provide little forage and consequently to 500 to 5,500 ft (152-1,676 m) in central Washington.
are used only as a final resort by livestock. Heavy use by Structural and compositional variation in this habitat are
livestock or vehicles disrupts the moss/lichen layer and related to changes in salinity and fluctuations in the water
increases exposed rock and bare ground that create habitat table. Areas with little or no vegetative cover have highly
for exotic plant invasion. Exotic annual bromes have alkaline and saline soils and are poorly drained or
become part of these habitats with natural soil churning irregularly flooded. Other arid soil types include desert
disturbance. pavement and barren ash. The wettest variants of the
habitat are usually found at the mouth of stream drainages
Status and Trends. Quigley and Arbelbide181 concluded or in areas with some freshwater input into a playa. These
that the low sagebrush cover type is as abundant as it was have finer, deeper alluvial soils that occur in low alkaline
before 1900. They concluded that “Low Sagebrush-Xeric” dunes, around playas, on slopes above alkaline basins or
successional pathways have experienced a high level of in small, poorly drained basins in sagebrush steppe.
change from exotic invasions and that some pathways of Topographically, this habitat occurs on playas or desert
“Low Sagebrush-Mesic” are unaltered. Twenty percent of pavement, or on low benches above playas with occasional
Pacific Northwest dwarf shrub-steppe community types low alkaline dune ridges.
listed in the National Vegetation Classification are
considered imperiled or critically imperiled.10 Landscape Setting. This habitat is typically surrounded
by shrub-steppe habitat. It forms a habitat mosaic of
playas, salt grass meadows, salt desert shrublands and
sagebrush shrublands. This habitat may be associated with
Herbaceous Wetland habitat. Local land use can result in
juxtaposition with Agriculture or Eastside Grasslands
habitat. Most of this habitat provides rangeland for
 54 Wildlife–Habitat Relationships in Oregon and Washington

livestock, particularly as winter range. Portions of this maritimus), and creeping spikerush (Eleocharis palustris) are
habitat associated with water are most attractive to diagnostic of the wettest parts of this habitat.
livestock. Other portions of this type are designated
Other Classifications and Key References. Popular
wildlife refuges.
literature refers to this habitat as shadscale, salt desert
Structure. This habitat is structurally diverse, ranging scrub, and saltflat desert. This habitat encompasses the
from dense shrublands to sparse grasslands to “Desert or Salt Desert Shrub” and “Distichlis stricta
unvegetated flats. Generally, low to medium-tall alkali or Associations on Saline-Alkali Soils” in Franklin and
saline tolerant shrubs form an open layer over a grass and Dyrness88 and Saltbush-greasewood in Kuchler.136 The
annual, often succulent, forb undergrowth. Deciduous Oregon Gap II Project 126 and Oregon Vegetation
shrubs, when present, usually create <50% cover but Landscape-Level Cover Types127 that would represent this
occasionally can exceed 70% on previously disturbed type are salt desert scrub shrubland and alkali playa. Other
ground. Ground cover between shrubs is variable, ranging references describe this habitat. 29, 30, 52, 60, 123, 131, 147, 175, 184
from widely spaced tall, medium-tall, or short
Natural Disturbance Regime. Fire plays a minor role over
bunchgrasses to dense, short rhizomatous grasses. Other
much of the distribution of the type because of sparse
areas have no shrubs and support a fairly continuous cover
vegetation and lack of fuel. Many of these areas are prone
of graminoids, occasionally with widely spaced
to irregular flooding and prolonged droughts; both factors
bunchgrasses. Sites can have extensive bare ground,
lead to a redistribution of component species and creation
usually gravelly flats, ash, desert pavement, or low
of sparsely or unvegetated areas.
alkaline dune ridges. Typically, this habitat is a mosaic of
open medium-tall to low shrubland communities, patchy Succession and Stand Dynamics. Many of the dominant
grasslands or herb lands, and sparsely to unvegetated shrub species sprout following fire, herbicide treatments,
areas. or heavy grazing.4 The characteristic shrubs of this habitat
increase with grazing and can invade adjacent big
Composition. Characteristic medium-tall shrubs that
sagebrush communities with intense grazing.
dominate well-drained sites are shadscale (Atriplex
confertifolia), bud sagebrush (Artemisia spinescens), and Effects of Management and Anthropogenic Impacts.
hopsage (Grayia spinosa). Characteristic low shrubs are Several exotic species invade this habitat with grazing.
greenmolly (Kochia americana), saltbush (Atriplex gardneri Halogeton, a toxic exotic plant, is found most commonly
or A. nuttallii), and winter fat (Krascheninnikovia lanata). in this habitat. Other noxious but nontoxic exotics that
Other medium-tall shrubs, big sagebrush (Artemisia increase with grazing are Russian thistle (Salsola kali), tall
tridentata), horsebrush (Tetradymia nuttallii or T. glabrata), tumblemustard (Sisymbrium altissimum), and cheatgrass.
Mormon tea (Ephedra viridis), or rabbitbrush These can replace native grasses and change the structure
(Chrysothamnus nauseosus or C. viscidiflorus) can be co- of the native habitat.
dominant. The medium-tall shrub black greasewood
Status and Trends. Agricultural development is generally
(Sarcobatus vermiculatus) or low shrubs, iodinebush
not feasible; consequently, little of this habitat is converted
(Allenrolfea occidentalis) or Mojave seablite (Suaeda moquinii)
to other uses. Most of this habitat is used for livestock
can be dominant or co-dominant on less well drained,
grazing, which overall has increased shrub and annual
generally more saline parts of this habitat.
cover and decreased bunchgrass cover. Quigley and
Herbaceous indicators of salt desert habitats appear in
Arbelbide181 concluded that the Salt Desert Shrub cover
various habitats. On densely vegetated sites, native
type is less abundant now than before 1900. They further
bunchgrasses, basin wildrye (Leymus cinereus), curly
noted that the cover type has undergone a moderate level
bluegrass (Poa secunda), and needle-and-threadgrass (Stipa
of change, so that some successional pathways have been
comata) are important, usually with shrubs in this habitat.
unaltered. Approximately one third of Pacific Northwest
Basin wildrye is also a common and diagnostic grass in
salt desert and related community types listed in the
sites with less alkaline, deeper soils and some movement
National Vegetation Classification are considered
of water. Indian ricegrass (Oryzopsis hymenoides) and
imperiled or critically imperiled.10
bottlebrush squirreltail (Elymus elymoides) are dominant
grasses on the alkaline dunes. Introduced plants,
particularly cheatgrass (Bromus tectorum) or halogeton
(Halogeton glomeratus), often dominate overgrazed sites.
Saltgrass (Distichlis spicata) is a common, diagnostic native
sod-forming grass on more saline sites that often
dominates large areas with and without shrubs.
Pickleweed (Salicornia virginica) is found in wetter saline
areas. Alkaline sites have mat muhly (Muhlenbergia
richardsonis), alkali bluegrass (Poa secunda ssp. juncifolia),
beardless wildrye (Leymus triticoides), and Lemmon’s
alkaligrass (Puccinella lemmonii). Common reedgrass
(Phragmites australis), bulrush (Scirpus americanus, S.
 CHAPTER 2: HABITAT DESCRIPTIONS 55

19. Agriculture, Pastures, and Mixed grasslands are typified by periods of bare soil and harvest
whereas pastures are mowed, hayed, or grazed one or
Environs more times during the growing season. Structural
W. Daniel Edge, Rex C. Crawford, & David H. Johnson diversity of agricultural habitat is increased at local scales
by the presence of noncultivated or less intensively
managed vegetation such as fencerows, roadsides, field
borders, and shelterbelts.
Composition. Agricultural habitat varies substantially in
composition among the cover types it includes. Cultivated
cropland includes >50 species of annual and perennial
plants in Oregon and Washington, and hundreds of
varieties ranging from vegetables such as carrots, onions,
and peas to annual grains such as wheat, oats, barley, and
rye. Row crops of vegetables and herbs are characterized
by bare soil, plants, and plant debris along bottomland
areas of streams and rivers and areas having sufficient
water for irrigation. Annual grains, such as barley, oats,
and wheat are typically produced in almost continuous
stands of vegetation on upland and rolling hill terrain
Geographic Distribution. Agricultural habitat is widely without irrigation.
distributed at low to mid-elevations (<6,000 ft [1,830 m]) The orchard/vineyard/nursery cover type is composed
throughout both states. This habitat is most abundant in of fruit and nut (apples, peaches, pears, and hazelnuts)
broad river valleys throughout both states and on gentle trees, vineyards (grapes, Kiwi), berries (strawberries,
rolling terrain east of the Cascades. blueberries, blackberries, and raspberries), Christmas
trees, and nursery operations (ornamental container and
Physical Setting. This habitat is maintained across a range greenhouses). This cover type is generally located on
of climatic conditions typical of both states. Climate upland sites with access to abundant irrigation.
constrains agricultural production at upper elevations Cultivation for most orchards, vineyards and Christmas
where there are <90 frost-free days. Agricultural habitat tree farms includes an undergrowth of short-stature
in arid regions east of the Cascades with <10 inches (25 perennial grasses between the rows of trees, vines, or
cm) of rainfall require supplemental irrigation or fallow bushes. Christmas trees are typically produced without
fields for 1-2 years to accumulate sufficient soil moisture. irrigation on upland sites with poorer soils.
Soils types are variable, but usually have a well developed Improved pastures are used to produce perennial
A horizon. This habitat is found from 0 to 6,000 ft (0 to herbaceous plants for grass seed and hay. Alfalfa and
1,830 m) elevation. several species of fescue (Festuca spp.) and bluegrass (Poa
Landscape Setting. Agricultural habitat occurs within a spp.), orchardgrass (Dactylis glomerata), and timothy
matrix of other habitat types at low to mid-elevations, (Phleum pratensis) are commonly seeded in improved
including Eastside grasslands, Shrub-steppe, Westside pastures. Grass seed fields are single-species stands,
Lowlands Conifer-Deciduous Forest and other low- to whereas pastures maintained for haying are typically
mid-elevation forest and woodland habitats. This habitat composed of two to several species. The improved pasture
often dominates the landscape in flat or gently rolling cover type is one of the most common agricultural uses in
terrain, on well-developed soils, broad river valleys, and both states and produced with and without irrigation.
areas with access to abundant irrigation water. Unlike Unimproved pastures are predominately grassland
other habitat types, agricultural habitat is often sites, often abandoned fields that have little or no active
characterized by regular landscape patterns (squares, management such as irrigation, fertilization, or herbicide
rectangles, and circles) and straight borders because of applications. These sites may or may not be grazed by
ownership boundaries and multiple crops within a region. livestock. Unimproved pastures include rangelands
Edges can be abrupt along the habitat borders within planted to exotic grasses that are found on private land,
agricultural habitat and with other adjacent habitats. state wildlife areas, federal wildlife refuges and U.S.
Department of Agriculture Conservation Reserve Program
Structure. This habitat is structurally diverse because it (CRP) sites. Grasses commonly planted on CRP sites are
includes several cover types ranging from low-stature crested wheatgrass (Agropyron cristatum), tall fescue (F.
annual grasses and row crops (<3.3 ft [1 m]) to mature arundinacea), perennial bromes (Bromus spp.) and
orchards (>66 ft [20 m]). However, within any cover type, wheatgrasses (Elytrigia spp.). Intensively grazed
structural diversity is typically low because usually only rangelands, which have been seeded to intermediate
one to a few species of similar height are cultivated. wheatgrass (Elytrigia intermedia), crested wheatgrass, or
Depending on management intensity or cultivation are dominated by increaser exotics such as Kentucky
method, agricultural habitat may vary substantially in wheatgrass (Poa pratensis) or tall oatgrass (Arrhenatherum
structure annually; cultivated cropland and modified elatius) are unimproved pastures. Other unimproved
 56 Wildlife–Habitat Relationships in Oregon and Washington

pastures have been cleared and intensively farmed in the oak. Douglas-fir or other trees can be primary invaders in
past, but are allowed to convert to other vegetation. These some environments.
sites may be composed of uncut hay, litter from previous
Effects of Management and Anthropogenic Impacts. The
seasons, standing dead grass and herbaceous material,
dominant characteristic of agricultural habitat is a regular
invasive exotic plants (tansy ragwort [Senecio jacobea],
pattern of management and vegetation disturbance. With
thistle [Cirsium spp.], Himalaya blackberry [Rubus
the exception of the unimproved pasture cover type, most
discolor], and Scot’s broom [Cytisus scoparius]) with patches
areas classified as agricultural habitat receive regular
of native black hawthorn (Crataegus douglasii), snowberry
inputs of fertilizer and pesticides and have some form of
(Symphoricarpos spp.), spirea (Spirea spp.), poison oak
vegetation harvest and manipulation. Management
(Toxicodendron diversilobum), and encroachment of various
practices in cultivated cropland include different tillage
tree species, depending on seed source and environment.
systems, resulting in vegetation residues during the non-
Modified grasslands are generally overgrazed habitats
growing season that range from bare soil to 100% litter.
that formerly were native eastside grasslands or shrub-
Cultivation of some crops, especially in the arid eastern
steppe but are now dominated by annual plants with only
portions of both states, may require the land to remain
remnant individual plants of the native vegetation.
fallow for 1-2 growing seasons in order to store sufficient
Cheatgrass (Bromus tectorum), other annual bromes,
soil moisture to grow another crop. Harvest in cultivated
medusahead (Taeniatherum caput-medusae), bulbous
cropland, Christmas tree plantations, and nurseries, and
bluegrass (Poa bulbosa), and knapweeds (Centaurea spp.)
mowing or haying in improved pasture cover types
are common increasers that form modified grasslands.
substantially change the structure of vegetation. Harvest
Fire, following heavy grazing or repeated early season fires
in orchards and vineyards is typically less intrusive, but
can create modified grassland monocultures of cheatgrass.
these crops as well as Christmas trees and some
Agricultural habitat also contains scattered dwellings
ornamental nurseries are regularly pruned. Improved
and outbuildings such as barns and silos, rural cemeteries,
pastures are often grazed after haying or during the
ditchbanks, windbreaks, and small inclusions of remnant
nongrowing season. Livestock grazing is the dominant
native vegetation. These sites typically have a
use of unimproved pastures. All of these practices prevent
discontinuous tree layer or one to a few trees over a ground
agricultural areas from reverting to native vegetation.
cover similar to improved or unimproved pastures.
Excessive grazing in unimproved pastures may increase
Other Classifications and Key References. Quigley and the prevalence of weedy or exotic species.
Arbelbide181 referred to this as agricultural and exotic
Status and Trends. Agricultural habitat has steadily
forbs-annual grasses cover types. Csuti et al.58 referred to
increased in amount and size in both states since Eurasian
this habitat as agricultural. The Oregon Gap II Project126
settlement of the region. Conversion to agricultural habitat
and Oregon Vegetation Landscape-Level Cover Type127
threatens several native habitat types.166 The greatest
that would represent this type is agriculture. U.S.
conversion of native habitats to agricultural production
Department of Agriculture Conservation Reserve Program
occurred between 1950 and 1985, primarily as a function
lands are included in this habitat.
of U.S. agricultural policy.96 Since the 1985 Farm Bill and
Natural Disturbance Regime. Natural fires are almost the economic downturn of the early to mid 1980s, the
totally suppressed in this habitat, except for unimproved amount of land in agricultural habitat has stabilized and
pastures and modified grasslands, where fire-return begun to decline.164 The 1985 and subsequent Farm Bills
intervals can resemble those of native grassland habitats. contained conservation provisions encouraging farmers
Fires are generally less frequent today than in the past, to convert agricultural land to native habitats.96, 153 Clean
primarily because of fire suppression, construction of farming practices and single-product farms have become
roads, and conversion of grass and forests to cropland.159 prevalent since the 1960s, resulting in larger farms and
Bottomland areas along streams and rivers are subject to widespread removal of fencerows, field borders,
periodic floods, which may remove or deposit large roadsides, and shelterbelts.96, 153, 164 In Oregon, land-use
amounts of soil. planning laws prevent or slow urban encroachment and
subdivisions into areas zoned as agriculture. Washington’s
Succession and Stand Dynamics. Management practices
growth management is currently controlled by counties
disrupt natural succession and stand dynamics in most
and agricultural land conversion to urban development
of the agricultural habitats. Abandoned eastside
is much less regulated.
agricultural habitats may convert to other habitats, mostly
grassland and shrub habitats from the surrounding native
habitats. Some agricultural habitats that occur on highly
erodible soils, especially east of the Cascades, have been
enrolled in the U.S. Department of Agriculture
Conservation Reserve Program. In the absence of fire or
mowing, westside unimproved pastures have increasing
amounts of hawthorn, snowberry, rose (Rosa spp.),
Himalaya blackberry, spirea, Scot’s broom, and poison
 CHAPTER 2: HABITAT DESCRIPTIONS 57

20. Urban and Mixed Environs for the construction of buildings and associated
Howard L.Ferguson infrastructure. Many structural features typical of the
historical vegetation, such as snags, dead and downed
wood, and brush piles, are often completely removed from
the landscape. Understory vegetation may be completely
absent, or if present, is diminutive and single-layered.
Typically, three zones are characteristic of urban habitat.

High-density Zone
The high-density zone is the downtown area of the inner
city. It also encompasses the heavy industrial and large
commercial interests of the city in addition to high-density
housing areas such as apartment buildings or high-rise
condominiums. This zone has >60% of its total surface
area covered by impervious surfaces. This zone has the
smallest lot size, the tallest buildings, the least amount of
total tree canopy cover, the lowest tree density, the highest
percentage of exotics, the poorest understory and
Geographic Distribution. Urban habitat occurs subcanopy, and the poorest vegetative structure.4a, 116a, 185a
throughout Oregon and Washington. Most urban Human structures have replaced almost all vegetation.23b,
development is located west of the Cascades of both 148a
Road density is the highest of all zones. An example of
Oregon and Washington, with the exception of Spokane, road density can be seen from Washington’s Growth
Washington. However, urban growth is being felt in almost Management Plan requiring Master Comprehensive Plans
every small town throughout the Pacific Northwest. to set aside 20% of the identified urban growth area for
Physical Setting. Urban development occurs in a variety roads and road rights-of-way. For example, Spokane’s
of sites in the Pacific Northwest. It creates a physical setting urban growth area is approximately 57,000 acres (23,077
unique to itself: temperatures are elevated and background ha); therefore >11,000 acres (4,453 ha) were set aside for
lighting is increased; wind velocities are altered by the road surfaces.
urban landscape, often reduced except around the tallest In the high-density zone, land-use practices have
structures downtown, where high-velocity winds are removed most of the native vegetation. Patch sizes of
funneled around the skyscrapers. Urban development remaining natural areas often are so small that native
often occurs in areas with little or no slope and frequently interior species cannot be supported. Not only are
includes wetland habitats. Many of these wetlands have remaining patches of native vegetation typically
been filled in and eliminated. Today, ironically, many disconnected, but also they are frequently missing the full
artificial “wetland” impoundments are being created for complement of vertical strata.149 Stream corridors become
stormwater management, whose function is the same as heavily impacted and discontinuous. Most, if not all,
the original wetland that was destroyed. wetlands have been filled or removed. Large buildings
dominate the landscape and determine the placement of
Landscape Setting. Urban development occurs within or vegetation in this zone.30a This zone has the most street
adjacent to nearly every habitat type in Oregon and tree strips or sidewalk trees, most of which are exotics.
Washington, and often replaces habitats that are valuable There is virtually no natural tree replacement, and new
for wildlife. The highest urban densities normally occur trees are planted only when old ones die or are removed.
in lower elevations along natural or human-made Replacement trees are chosen for their small root systems
transportation corridors, such as rivers, railroad lines, and are generally short in stature with small diameters.
coastlines, or interstate highways. These areas often Ground cover in this zone, if not synthetic or impervious,
contain good soils with little or no slope and lush is typically exotic grasses or exotic annuals, most of which
vegetation. Once level areas become crowded, growth are rarely allowed to go to seed. Snags, woody debris, rock
continues along rivers or shores of lakes or oceans, and piles, and any other natural structures are essentially
eventually up elevated sites with steep slopes or rocky nonexistent. There are few tree cavities because of cosmetic
outcrops. Because early settlers often modified the original pruning, cavity filling, snag removal, and tree thinning.149
landscape for agricultural purposes, many of our urban
areas are surrounded by agricultural and grazing lands. Medium-density Zone
This zone, continuing out from the center of the
Structure. The original habitat is drastically altered in
continuum, is composed of light industry mixed with
urban environments and is replaced by buildings,
high-density residential areas. Housing density of 3-6
impermeable surfaces, bridges, dams, and planting of non-
single-family homes per acre (7-15 per ha) is typical.
native species. Some human-made structures provide
Compared with the high-density zone, this zone has more
habitats similar to those of cavities, caves, fissures, cliffs,
potential wildlife habitat. With 30-59% impervious soil
and ledges. With the onset of urban development, total
cover, this zone has 41-70% of the ground available for
crown cover and tree density are reduced to make way
plants. Road density is less than the high-density zone.
 58 Wildlife–Habitat Relationships in Oregon and Washington

Vegetation in this mid-zone is typically composed of areas around houses, is relatively intact. Lawns are fewer
non-native plant species. Native plants, when present, and native ground covers are more common than in the
represent only a limited range of the natural diversity for previous two zones.
the area. The shrub layer is typically clipped or minimal,
Composition. Remnant isolated blocks of native
even in heavily vegetated areas. Characteristic of this zone
vegetation may be found scattered throughout a town or
are manicured lawns, trimmed hedges, and topped trees.
city mixed with a multitude of introduced exotic
Lawns can be highly productive.82a, 97a Tree canopy is still
vegetation. As urban development increases, these
discontinuous and consists of 1-2 levels, if present at all.
remnant native stands become fragmented and isolated.
Consequently, vertical vegetative diversity and total
The dominant species in an urban setting may be exotic
amount of understory are still low. Coarse and fine woody
or native; for example, in Seattle, the dominant species in
debris is minimal or absent; most snags and diseased live
one area may be Douglas-fir (Pseudotsuga menziesii),
trees are still removed as hazards in this zone.119a, 119b
whereas a few blocks away it may be the exotic silver
Isolated wetlands, stream corridors, open spaces, and
maple (Acer saccharinum). Dominant species will not only
greenbelts are more frequently retained in this zone than
vary from city to city but also within each city and within
in the high-density zone. However, remnant wetland and
each of the three urban zones. Nowack167 found that in
upland areas are often widely separated by urban
the high-density urban zone, species richness is low, and
development.
in one case, four species made up almost 50% of the cover.
Low-density Zone In the same study, exotics made up 69% of the total species.
The low-density zone is the outer zone of the urban-rural In urban and suburban areas, species richness is often
continuum. This zone contains only 10-29% impervious increased because of the introduction of exotics. The
ground cover and normally contains only single-family juxtaposition of exotics interspersed with native
homes. It has more natural ground cover than artificial vegetation produces a diverse mosaic with areas of
surfaces. Vegetation is denser and more abundant than in extensive edge. Also, because of irrigation and the addition
the previous two zones. Typical housing densities are 0.4- of fertilizers, the biomass in the urban communities is often
1.6 single-family homes per acre (1-4 per ha). Road density increased. 149 Interest in the use of native plants for
is lowest of all three zones and consists of many secondary landscaping is rapidly expanding,135, 172 particularly in the
and tertiary roads. Although this zone may have large more arid sites where drought-resistant natives are the
areas of native vegetation and is generally the least only plants able to survive without water.
impacted of all three zones; it still has been significantly Across the U.S., urban tree cover ranges from 1 to
altered by human activities and associated disturbances. 55%.167 As expected, tree cover tends to be highest in cities
Roads, fences, livestock paddocks, and pets are more developed in naturally forested areas with an average of
abundant than in neighboring rural areas. With many 32% cover in forested areas, 28% in grasslands, and 10%
animals and limited acreage, pasture conditions may be in arid areas. Yakima, Washington, has an overall city tree
more overgrazed in this zone than in the rural zone; cover of 18%, ranging from 10% to 12% in the industrial/
overgrazing can significantly affect shrub layers as well. commercial area to 23% in the low-density residential
Areas around home sites are often cleared for fire zone.167 Remnant blocks of native vegetation or native trees
protection. Dogs are more likely to be loose and allowed left standing in yards and parks will compositionally be
to run free, increasing disturbance levels and wildlife related to whatever native habitat was on site prior to
harassment in this zone. Vegetable and flower gardens are development. In the Puget Sound and Willamette Valley
widespread; fencing is prevalent. areas, Douglas-fir is a major constituent, whereas the
Many wetlands remain and are less impacted. Water Spokane area has a lot of ponderosa pina (Pinus ponderosa).
levels are more stable and peak flows are more typical of Other Classifications and Key References. Many
historical flows. Watertables are less impacted and vernal attempts have been made to classify or describe the
wetlands are more frequent; stream corridors are less complex urban environment. The Washington GAP
impacted and more continuous. Analysis37 classified urban environments as “developed”
This zone has the most vertical and horizontal structure land cover using the same three zones as described above:
and diversity of any of the three urban zones.30a, 80a, 140a, 187a (1) high density (>60% impervious surface); (2) medium
In forested areas, tree conditions are semi-natural, density (30-60% impervious surface); and (3) low density
although stand characteristics vary from parcel to parcel. (10-30% impervious surface). The Oregon Gap II Project126
The tree canopy is more continuous and may include and Oregon Vegetation Landscape-Level Cover Types127
multiple levels. Patch sizes are large enough to support represented this type as an urban class. Several other
native interior species. Large blocks of native vegetation relevant strudies characterizing the urban environment
may still be found, and some of these may be connected have been reported. 182, 129, 34, 70, 151
to large areas of native undeveloped land.220a In this zone,
snags, diseased trees, coarse and fine woody debris, brush Natural Disturbance Regime. In many instances, natural
piles, and rock piles are widespread. Structural diversity disturbances are modified or prevented from occurring
approaches historical levels. Non-native hedges are nearly by humans over the landscape and this is particularly true
nonexistent and the native shrub layer, except for small of urban areas. However, disturbances such as ice, wind,
 CHAPTER 2: HABITAT DESCRIPTIONS 59

or firestorms still occur. The severity of these intermittent structures, large-scale storm and wastewater management,
disturbances varies greatly in magnitude and their impact large-scale sewage treatment, water and air pollution, toxic
on the landscape varies accordingly. One of the differences chemicals, toxic chemical use on urban lawns and gardens,
between urban and nonurban landscapes is the removal of species considered to be pests, predation and
lengthening of the disturbance cycles. Another is found disturbance by pets and feral cats and dogs, and the
in the aftermath of these disturbances. In urban areas, extensive and continual removal of habitat due to
damaged trees are often entirely removed and if they are expanding urbanization, and in some cases, uncontrolled
replaced, a shorter, smaller tree, often non-native, is development. Another significant impact is the
selected. The natural fire disturbance interval is highly introduction and cultivation of exotics in urban areas.
modified in the urban environment. Fire (mostly Native vegetation is often completely replaced by exotics,
accidental or arson) still occurs, and is quickly suppressed. leaving little trace of the native vegetative cover.
Another natural disturbance in many of our Pacific
Status and Trends. From 1970 to 1990, >30,000 mile2
Northwest towns is flooding, which historically altered
(77,700 km2) of rural lands in the U.S. became urban, as
and rerouted many of our rivers and streams, and still
classified by the U.S. Census Bureau. That amount of land
scarifies fields and deposits soil on flood plains and
equals about one third of Oregon’s total land area.12 From
potentially recharges local aquifers. Floods now are more
1940 to 1970, the population of the Portland urban region
frequent and more violent than in the past because of the
doubled and the amount of land occupied by that
many modifications made to our watersheds. Attempts
population quadrupled.201 More than 300 new residents
to lessen flooding in urban areas often lead to
arrive in Washington each day, and each day, Washington
channelization, paving, or diking of our waterways, most
loses 100 acres (41 ha) of forest to development.215 Using
of which fail in their attempt to stem the flooding and
satellite photos and GIS software, American Forests9
usually result in increased flooding for the communities
discovered that nearly one third of Puget Sound’s most
farther downstream.
heavily timbered land has disappeared since the early
Succession and Stand Dynamics. Due to anthropogenic 1970s. The amount of land with few or no trees more than
influences found in the urban environment, succession doubled, from 25% to 57%, an increase of >1 million acres
differs in the urban area from that expected for a native (404,858 ha). Development and associated urban growth
stand. Rowntree185 emphasized that urbanization is not was blamed as the single biggest factor affecting the area’s
in the same category as natural disturbance in affecting environment. This urban growth is predicted to continue
succession. He points out that urbanization is to increase at an accelerated pace, at the expense of native
anthropogenic and acts to remove complete vegetation habitat.
associations and creates new ones made of mixes of native
residual vegetation and introduced vegetation. Much 21. Open Water—
human effort in the city goes toward either completely
removing native vegetation or sustaining or maintaining
Lakes, Rivers, and Streams
a specific vegetative type, e.g., lawns or hedges. Much of Eva L. Greda, David H. Johnson, & Thomas A. O’Neil
the vegetative community remains static. Understory and
ground covers are constantly pruned or removed,
seedlings are pulled and lawns are planted, fertilized,
mowed, and meticulously maintained. Trees may be
protected to maturity or even senescence, yet communities
are so fragmented or modified that a genuine old-growth
community never exists. However, a type of “urban
succession” occurs across the urban landscape. The older
neighborhoods with their mature stands are at a later seral
stage than new developments; species diversity is
characteristically higher in older neighborhoods as well.
An oddity of the urban environment is the absence of
typical structure generally found within the various seral
stages. For example, the understory is often removed in a
typical mid-seral stand to give it a “park-like” look. Or if
the understory is allowed to remain, it is kept pruned to a
consistent height. Lawns are the ever-present substitute Lakes, Ponds, and Reservoirs
for native ground covers. Multilayered habitat is often
Geographic Distribution. Lakes in Oregon and
reduced to one or two heights. Vertical and horizontal
Washington occur statewide and are found from near sea
structural diversity is drastically reduced.
level to about 10,200 ft (3,110 m) above sea level. There
Effects of Management and Anthropogenic Impacts. are 3,887 lakes and reservoirs in western Washington and
These additional, often irreversible, impacts include more
Text continues on page 91
impervious surfaces, more and larger human-made
 60 Wildlife–Habitat Relationships in Oregon and Washington

Westside Lowlands Conifer-Hardwood Forest

1. Goodman Creek, Oregon 2. H.J. Andrews Experimental Forest, Oregon

3. Deception Pass State Park, Island County, Washington 4. Capitol Forest, Thurston County, Washington

5. Dungeness River Valley, Clallam County, Washington 6. Lake Quinalt, Grays Harbor County, Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 61

Westside Oak and Dry Douglas-fir Forest and Woodlands

1. Orcas Island, San Juan County, Washington 2. James Island, San Juan County, Washington

3. San Juan Island, San Juan County, Washington 4. Fort Lewis, Pierce County, Washington
 62 Wildlife–Habitat Relationships in Oregon and Washington

Southwest Oregon Mixed Conifer-Hardwood Forest

1. Southwestern Oregon

2. Jackson County, Oregon 3. Ruch, Oregon

 CHAPTER 2: HABITAT DESCRIPTIONS 63

Montane Mixed Conifer Forest

1. Mt. Pilchuck Conservation Area, Snohomish Co., WA 2. Pend Oreille County, Washington

3. Mt. Pilchuck Conservation Area, Snohomish County, 4. Arlecho Creek, Whatcom County, Washington
Washington
 64 Wildlife–Habitat Relationships in Oregon and Washington

Eastside (Interior) Mixed Conifer Forest

1. Wenatchee Mountains, Kittitas County, Washington 2. Alice Mae Mountain, Stevens County, Washington

2. Alice Mae Mountain, Stevens County, Washington 4. Rainbow Creek Research Natural Area, Blue
Mountains, Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 65

Lodgepole Pine Forest and Woodlands

1. Loomis State Forest, Okanogan County, Washington

2. Loomis State Forest, Okanogan County, Washington 3. Loomis State Forest, Okanogan County, Washington

4. Loomis State Forest, Okanogan County, Washington 5. Loomis State Forest, Okanogan County, Washington
 66 Wildlife–Habitat Relationships in Oregon and Washington

Ponderosa Pine Forest and Woodlands (includes Eastside Oak)

1. Barker Mountain, Okanogan County, Washington 2 Near Sisters, Oregon

3. Turnbull National Wildlife Refuge, Spokane County, 4. Indian Ford, north of Sisters, Oregon
Washington

5. Briske Canyon, Washington 6. Badger Gulch Natural Area Preserve, Klickitat

County, Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 67

Upland Aspen Forest

1. Winthrop, Okanogan County, Washington

2. Hart Mountain National Wildlife Refuge, Oregon

3. Hart Mountain Mountain National Wildlife Refuge, 4. Steens Mountain, Oregon

Oregon
 68 Wildlife–Habitat Relationships in Oregon and Washington

Subalpine Parkland

1. Mt. Rainier National Park, Washington

4. Eagle Cap Wilderness Area, Wallowa Mts., Oregon

2. Strawberry Mountain, Oregon

3. Mt. Pilchuck Conservation Area, Snohomish County, 5. Goat Rocks Wilderness Area, Lewis County,
Washington Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 69

Alpine Grasslands and Shrublands

1. Mt. Rainier National Park, Washington

2. Steens Mountain, Oregon

3. Buckhorn Wilderness Area, Clallam County, Washington 4. Mt. Rainier National Park, Washington
 70 Wildlife–Habitat Relationships in Oregon and Washington

Westside Grasslands

1. Scatter Creek Wildlife Area, Thurston County, Washington

2. Lane County, Oregon 3. Near Stayton, Linn County, Oregon

4. Burrows Island, Skagit County, Washington 5. Mima Mounds Natural Area Preserve, Thurston
County, Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 71

Ceanothus – Manzanita Shrublands

1. Southwestern Oregon 2. South of Shady Cove, Oregon

3. Southwestern Oregon 4. South of Shady Cove, Oregon

 72 Wildlife–Habitat Relationships in Oregon and Washington

Western Juniper and Mountain Mahogany Woodlands

1. Sheldon National Wildlife Refuge, Nevada 2. Sheldon National Wildlife Refuge, Nevada

3. Jefferson County, Oregon 4. Klamath County, Oregon

 CHAPTER 2: HABITAT DESCRIPTIONS 73

Eastside (Interior) Canyon Shrublands

1. Campus Prairie Biological Study Area, Whitman County, Washington

2. Douglas Creek, Douglas County, Washington 3. Asotin County, Washington

4. Kramer Palouse Biological Study Area, Whitman 5. Grand Ronde River, Asotin County, Washington
County, Washington
 74 Wildlife–Habitat Relationships in Oregon and Washington

Eastside (Interior) Grasslands

1. Gibraltar Mountain, Ferry County, Washington 2. Palouse River, Franklin County, Washington

3. Columbia Hills Natural Area Preserve, Klickitat 4. Arid Lands Ecology Reserve, Hanford, Washington
County, Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 75

Shrub-Steppe

1. Horse Heaven Hills, Benton County, Washington

2. Yakima Firing Range, Washington 3. Yakima Firing Range, Washington

3. Steens Mountain, Oregon 4. Vernita, Grant County, Washington

 76 Wildlife–Habitat Relationships in Oregon and Washington

Dwarf Shrub-Steppe

1. Castle Rock, Grand Coulee, Washington 2. Umtanum Ridge, Kittitas County, Washington

3. Saddle Mountains, Hanford, Washington 4. Sheldon National Wildlife Refuge, Nevada

 CHAPTER 2: HABITAT DESCRIPTIONS 77

Desert Playa and Salt Scrub Shrublands

1. Alvord Desert, Oregon

2. Alvord Desert, Oregon 3. Alvord Desert, Oregon

4. Harney Basin, Oregon 5. Alvord Desert, Oregon

 78 Wildlife–Habitat Relationships in Oregon and Washington

Agriculture, Pasture, and Mixed Environs

1. Near Samish Bay, Skagit County, Washington 2. Sinlahekin Valley, Okanogan County, Washington

3. Ellensburg area, Kittitas County, Washington 4. Southeast Washington (wheat field)

5. Benton County, Oregon 6. Scotch Creek Wildlife Area, Okanogan County,

Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 79

Urban and Mixed Environs

1. Example of high density urban, Seattle, Washington 2. Example of medium density urban, Washington

3. Example of low density urban, Benton County, Oregon 4. Example of low density urban, Wenatchee,
Washington
 80 Wildlife–Habitat Relationships in Oregon and Washington

Open Water—Lakes, Rivers, Streams

1. Lake Ozette, Washington 2. Lower Soleduck River, Washington

3. Grant County, Washington

4. Snake River, Washington 5. Willamette River, Linn County, Oregon

 CHAPTER 2: HABITAT DESCRIPTIONS 81

Herbaceous Wetlands

1. Okanogan County, Washington

2. Methow Valley, Okanogan County, Washington 3. Pacific County, Washington

4. Lincoln County, Washington 5. Linn County, Oregon

 82 Wildlife–Habitat Relationships in Oregon and Washington

Westside Riparian - Wetlands

1. Stequaleho Creek, Jefferson County, Washington

2. Cranberry Creek, Grays Harbor County, Washington 3. Maxfield Creek, Clallam County, Washington

4. Cranberry Creek, Grays Harbor, Washington 5. Quinalt River, Grays Harbor County, Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 83

Montane Coniferous Wetlands

1. Arlecho Creek, Whatcom County, Washington 2. Roger Lake, Okanogan County, Washington

3. Mt. Rainier National Park, Washington 4. Mt. Pilchuck Conservation Area, Snohomish
County, Washington
 84 Wildlife–Habitat Relationships in Oregon and Washington

Eastside (Interior) Riparian-Wetlands

1. Northrup Canyon, Grant County, Washington 2. Little Pend Oreille River, Stevens County,
Washington

3. Douglas Creek, Douglas County, Washington 4. Myers Creek, Okanogan County, Washington

5. Eastern Klickitat County, Washington 6. Crimm’s Creek, Washington

 CHAPTER 2: HABITAT DESCRIPTIONS 85

Coastal Dunes and Beaches

1. Florence, Oregon

2. Whidbey Island, Washington 3. Ocean Shores, Grays Harbor County, Washington

4. Whidbey Island, Washington 5. Whidbey Island, Washington

 86 Wildlife–Habitat Relationships in Oregon and Washington

Coastal Headlands and Islets

1. Near Cape Perpetua, Oregon 2. Heceta Head, Oregon

3. Near Yaquina Head, Oregon

4. Near Cape Perpetua, Oregon 5. Coastline near Quinalt River, Grays Harbor
County, Washington
 CHAPTER 2: HABITAT DESCRIPTIONS 87

Bays and Estuaries

1. Estuary, Washington 2. Dogfish Point, Skagit County, Washington

3. Niawiakum River Natural Area Preserve, Willapa Bay, 4. Nisqually Delta, Thurston County, Washington
Washington

5. Niawiakum River, Willapa Bay, Washington 6. Newport, Oregon

 88 Wildlife–Habitat Relationships in Oregon and Washington

Marine Nearshore

1. Ben Ure Island, Island County, Washington

2. Cypress Island, Skagit County, Washington 3. Guemes Island, Skagit County, Washington

4. Near Biz Point, Skagit County, Washington 5. Near Cascade Head, Lincoln County, Oregon
 CHAPTER 2: HABITAT DESCRIPTIONS 89

Inland Marine Deeper Water

1. Squaxin Island, South Puget Sound, Washington

2. San Juan Islands, Washington

 90 Wildlife–Habitat Relationships in Oregon and Washington

Marine Habitats

Bays and Esturaries Marine Shelf

Inland Marine Deeper Water Oceanic

Marine Nearshore
 CHAPTER 2: HABITAT DESCRIPTIONS 91

they total 176,920 acres (71,628 ha)226. In contrast, there river. Washington has more streams than any other state
are 4,073 lakes and reservoirs in eastern Washington that except Alaska. In Washington, the coastal region has 3,783
total 436,843 acres (176,860 ha).227 There are 6,000 lakes, rivers and streams totaling 8,176 miles (13,155 km)174. The
ponds, and reservoirs in Oregon including almost 1,800 Puget Sound Region has 10,217 rivers and streams, which
named lakes and over 3,800 named reservoirs, all add to 16,600 miles (26,709 km) in length.223 The rivers
amounting to 270,641 acres (109,571 ha). Oregon has the and streams range from cold, fast-moving high-elevation
deepest lake in the nation, Crater Lake, at 1,932 ft (589 streams to warmer lowland valley rivers.223 In all, there
m).23 are 13,955 rivers and streams that add up to 24,774 miles
(39,861 km). 174 There are many more streams in
Physical Setting. Continental glaciers melted and left
Washington yet to be catalogued.174 Streams reflect flowing
depressions, where water accumulated and formed many
water •6 feet (2 m) wied; narrower water bodies are
lakes in the region. These kinds of lakes are predominantly
considered within their respective habitats.
found in Lower Puget Sound. Landslides that blocked
natural valleys also allowed water to fill in behind them Physical Setting. Climate of the area’s coastal region is
to form lakes, like Crescent Lake, Washington. The lakes very wet. The northern region in Washington is volcanic
in the Cascades and Olympic ranges were formed through and bordered to the east by the Olympic Mountain Range,
glaciation and range in elevation from 2,500 to 5,000 ft on the north by the Strait of Juan de Fuca, and on the west
(762 to 1,524 m). Beavers create many ponds and marshes by the Pacific Ocean. In contrast, the southern portion in
in Oregon and Washington. Craters created by extinct Washington is characterized by low-lying, rolling hills.174
volcanoes, like Battleground Lake, Washington, also The Puget Sound Region has a wet climate. Most of the
formed lakes. Human-made reservoirs created by dams streams entering Puget Sound have originated in glacier
impound water that creates lakes behind them, like fields high in the mountains. Water from melting
Bonneville Dam on the main stem of the Columbia River. snowpacks and glaciers provide flow during the spring
In the lower Columbia Basin, many lakes formed in and winter. Annual rainfall in the lowlands ranges from
depressions and rocky coulees through the process of 35 to 50 inches (89-127 cm), from 75 to 100 inches (191 to
seepage from irrigation waters.226 254 cm) in the foothills, and from 100 to >200 inches (254
to 508 cm) in the mountains (mostly in the form of snow).174
Structure. There are four distinct zones within this aquatic
Rivers and streams in southwestern Oregon are fed by
system: (1) the littoral zone at the edge of lakes is the most
rain and are located in an area composed of sheared
productive with diverse aquatic beds and emergent
bedrock and thus an unstable terrain. Streams in that area
wetlands (part of Herbaceous Wetlands habitat); (2) the
have high suspended-sediment loads. Beds composed of
limnetic zone is deep open water, dominated by
gravel and sand are easily transported during floods. The
phytoplankton and freshwater fish, and extends down to
western Cascades in Washington and Oregon are
the limits of light penetration; (3) the profundal zone below
composed of volcanically derived rocks and are more
the limnetic zone, devoid of plant life and dominated with
stable. They have low sediment-transport rates and stable
detritivores; (4) and the benthic zone reflecting bottom soil
beds composed largely of cobbles and boulders, which
and sediments. Nutrients from the profundal zone are
move only during extreme events.81 Velocities of river flow
recycled back to upper layers by the spring and fall
ranges from as little as 0.2 to 12 mph (0.3 to19.3 km/hr)
turnover of the water. Water in temperate climates
while large streams have an average annual flow of 10
stratifies because of the changes in water density. The
cubic feet (0.3 m3) per second or greater.23, 169 Rivers and
uppermost layer, the epilimnion, is where water is warmer
streams in the Willamette Valley are warm, productive,
(less dense). Next, the metalimnion or thermocline, is a
turbid, and have high ionic strength. They are
narrow layer that prevents the mixing of the upper and
characterized by deep pools, and highly embedded stream
lowermost layers. The lowest layer is the hypolimnion,
bottoms with claypan and muddy substrates, and the
with colder and most dense waters. During the fall
greatest fish species diversity. High desert streams of the
turnover, the cooled upper layers are mixed with other
interior are similar to those of the Willamette Valley but
layers through wind action.
are shallower, with fewer pools, and more runs, glides,
cobbles, boulders, and sand. The Cascades and Blue
Rivers and Streams mountains are similar in that they have more runs and
glides and fewer pools, similar fish assemblages, and
Geographic Distribution. Streams and rivers are similar water quality.218
distributed statewide in Oregon and Washington, forming
a continuous network connecting high mountain areas to Lakes, Rivers and Streams
lowlands and the Pacific coast. There are >12,000 named
Landscape setting. This habitat occurs throughout
rivers and streams in Oregon, totalling 112,640 miles
Washington and Oregon. Ponds, lakes, and reservoirs are
(181,238 km) 23 in length. Oregon’s longest stretch of river
typically adjacent to Herbaceous Wetlands, while rivers
is the Columbia (309 miles [497 km]) that borders Oregon
and streams typicaslly adjoin the Westside Riparian-
and Washington. The longest river in Oregon is the John
Wetlands, Eastside Riparian-Wetlands, Herbaceous
Day (284 miles [457 km]) and the shortest river is the D
Wetlands, and Bays and Estuaries habitats.
River (440 ft [134 m]) that is the world’s second shortest
 92 Wildlife–Habitat Relationships in Oregon and Washington

Other Classifications and Key References. This habitat passage, competition between species, loss of spawning
is called riverine and lacustrine in Anderson et al.,10 areas because of flooding, and declines in native fish
Cowardin et al.,53 Washington Gap Analysis Project,37 populations.146 Historically, the region’s rivers contained
Mayer and Laudenslayer,150 and Wetzel.217 However, this more braided multi-channels. Flood control measures such
habitat is referred to as Open Water in the Oregon Gap II as channel straightening, diking, or removal of streambed
Project126 and Oregon Vegetation Landscape-Level Cover material along with urban and agriculture development
Types.127 have all contributed to a loss of oxbows, river meanders,
and flood plains. Unauthorized or over-appropriated
Natural Disturbance Regime. There are seasonal and
withdrawals of water from the natural drainages also has
decadal variations in the patterns of precipitation. In the
caused a loss of open water habitat that has been
Coast Range, there is usually 1 month of drought per year
detrimental to fish and wildlife production, particularly
(usually July or August) and 2 months of drought once in
in the summer.174
a decade. The Willamette Valley and the Cascades
Agricultural, industrial, and sewage runoff such as
experience 1 month with no rain every year and a 2-month
salts, sediments, fertilizers, pesticides, and bacteria harm
dry period every third year. In eastern Oregon, dry periods
aquatic species.146 Sludge and heavy waste buildup in
last 2 or 3 months every year, with dry spells as long as 4-
estuaries is harmful to fish and shellfish. Unregulated
6 months occurring once every 4 years. Dry years, with
aerial spraying of pesticides over agricultural areas also
<33% of normal precipitation, occur once every 30 years
poses a threat to aquatic and terrestrial life.174 Direct loss
along the coast, every 20 years in the Willamette Valley,
of habitat and water quality occurs through irrigation.130
every 30 years in the Cascades, and every 15 years in most
The Oregon Department of Environmental Quality, after
of eastern Oregon.23
a study of water quality of the Willamette River,
Floods occur in Oregon and Washington every year.
determined that up to 80% of water pollution enters the
Flooding season west of the Cascades occurs from October
river from nonpoint sources and especially agricultural
through April, with more than half of the floods occurring
activity.23 Very large floods (e.g., Oregon Flood of 1964)
during December and January. Floods are the result of
may change the channels permanently through the settling
precipitation and snow melts. Floods west of the Cascades
of large amounts of sediments from hillslopes, through
are influenced by precipitation mostly and thus are short-
debris flow, and through movement of large boulders,
lived, while east of the Cascades floods are caused by
particularly in the montane areas. The width of the channel
melting snow, and the amount of flooding depends on
along the main middle fork of the Willamette increased
how fast the snow melts. High water levels frequently last
over a period of 8 years. Clearcutting creates excessive
up to 60 days. In 1984, heavy precipitation flooded
intermittent runoff conditions and increases erosion and
Malheur and Harney lakes to the point where the two lakes
siltation of streams as well as diminishes shade, and
were joined together for several years. The worst floods
therefore causes higher water temperatures, fewer
have resulted from cloudbursts caused by thunderstorms,
terrestrial and aquatic food organisms, and increased
like Heppner, Oregon’s 1903 flood. Other “flash floods”
predation. Landslides, which contributed to the widening
in the region were among the largest floods in the U.S.
of the channel, were a direct result of clearcutting. Clearcut
and occurred in the John Day Basin’s Meyers Canyon in
logging can alter snow accumulation and increase the size
1956 and the Umatilla Basin’s Lane Canyon in 1965.23
of peak flows during times of snowmelt.197 Clearcutting
Effects of Management and Anthropogenic Impacts. and vegetation removal affects the temperatures of
Sewage effluents caused eutrophication of Lake streams, increasing them in the summer and decreasing
Washington in Seattle, where plants increased in biomass in winter, especially in eastern parts of the Oregon and
and caused decreased light transmission. The situation Washington.24 Building of roads, especially those of poor
was corrected, however, before it became serious as a result quality, can be a major contributor to sedimentation in
of a campaign of public education, and timely cleanup of the streams.82
the lake.146 Irrigation projects aimed at watering drier
Status and Trends. The principal trend has been in
portions of the landscape may pose flooding dangers, as
relationship to dam building or channelization for
was the case with Soap Lake and Lake Leonore in eastern
hydroelectric power, flood control, or irrigation purposes.
Washington. Finally, natural salinity of lakes can decrease
As an example, in 1994, there were >900 dams in
as a result of irrigation withdrawal and can change the
Washington alone. The dams vary according to size,
biota associated with them.92
primary purpose, and ownership (state, federal, private,
Removal of gravel results in reduction of spawning
local).214 The first dam and reservoir in Washington was
areas for anadromous fish. Overgrazing, and loss of
the Monroe Street Dam and Reservoir, built in 1890 at
vegetation caused by logging produces increased water
Spokane Falls. Since then the engineering and equipment
temperatures and excessive siltation, harming the
necessary for dam building developed substantially,
invertebrate communities such as that reported in the John
culminating in such projects as the Grand Coulee Dam on
Day River Basin, Oregon.146 Incorrectly installed culverts
the Columbia River.214 In response to the damaging effects
may act as barriers to migrating fish and may contribute
of dams on the indigenous biota and alteration and
to erosion and siltation downstream.174 Construction of
destruction of freshwater aquatic habitats, Oregon and
dams is associated with changes in water quality, fish
Washington state governments questioned the benefits of
 CHAPTER 2: HABITAT DESCRIPTIONS 93

dams, especially in light of the federal listing of several temporarily to seasonally flooded meadows and generally
salmon species. There are now talks of possibly removing occur on clay, pluvial, or alluvial deposits within montane
small dams, like the Savage Rapids Dam in Oregon, to meadows, or along stream channels in shrubland or
removing large federal dams like those on the lower Snake woodland riparian vegetation. In general, this habitat is
River.23 flat, usually with stream or river channels or open water
present. Elevation varies between sea level to 10,000 ft
22. Herbaceous Wetlands (3,048 m), although infrequently above 6,000 ft (1,830 m).
Rex C. Crawford, Jimmy Kagan, and Christopher B. Chappell Landscape Setting. Herbaceous wetlands are found in all
terrestrial habitats except Subalpine Parkland and Alpine
Grasslands and Shrublands habitats. Herbaceous
wetlands commonly form a pattern with Westside and
Eastside Riparian-Wetlands and Montane Coniferous
Wetlands habitats along stream corridors. These marshes
and wetlands also occur in closed basins in a mosaic with
open water by lakeshores or ponds. Extensive deflation
plain wetlands have developed between Coastal Dunes
and Beaches habitat and the Pacific Ocean. Herbaceous
wetlands are found in a mosaic with alkali grasslands in
the Desert Playa and Salt Scrub habitat.
Structure. The herbaceous wetland habitat is generally a
mix of emergent herbaceous plants with a grass-like life
form (graminoids). These meadows often occur with deep
or shallow water habitats with floating or rooting aquatic
forbs. Various wetland communities are found in mosaics
or in nearly pure stands of single species. Herbaceous
cover is open to dense. The habitat can be comprised of
tule marshes >6.6 ft (2 m) tall or sedge meadows and
wetlands <3.3 ft (1 m) tall. It can be a dense, rhizomatous
sward or a tufted graminoid wetland. Graminoid wetland
vegetation generally lacks many forbs, although the open
Geographic Distribution. Herbaceous wetlands are found extreme of this type contains a diverse forb component
throughout the world and are represented in Oregon and between widely spaced tall tufted grasses.
Washington wherever local hydrologic conditions Composition. Various grasses or grass-like plants
promote their development. This habitat includes all those dominate or co-dominate these habitats. Cattails (Typha
except bogs and those within Subalpine Parkland and latifolia) occur widely, sometimes adjacent to open water
Alpine habitats. with aquatic bed plants. Several bulrush species (Scirpus
Freshwater aquatic bed habitats are found throughout acutus, S. tabernaemontani, S. maritimus, S. americanus, S.
the Pacific Northwest, usually in isolated sites. They are nevadensis) occur in nearly pure stands or in mosaics with
more widespread in valley bottoms and high rainfall areas cattails or sedges (Carex spp.). Burreed (Sparganium
(e.g., Willamette Valley, Puget Trough, coastal terraces, angustifolium, S. eurycarpum) are the most important
coastal dunes), but are present in montane and arid graminoids in areas with up to 3.3 ft (1m) of deep standing
climates as well. Hardstem bulrush-cattail-burreed water. A variety of sedges characterize this habitat. Some
marshes occur in wet areas throughout Oregon and sedges (Carex aquatilis, C. lasiocarpa, C. scopulorum, C.
Washington. Large marshes are common in the lake basins simulata, C. utriculata, C. vesicaria) tend to occur in cold to
of Klamath, Lake, and Harney counties, Oregon. Sedge cool environments. Other sedges (C. aquatilis var. dives, C.
meadows and montane meadows are common in the Blue angustata, C. interior, C. microptera, C. nebrascensis) tend to
and Ochoco mountains of central and northeastern be at lower elevations in milder or warmer environments.
Oregon, and in the valleys of the Olympic and Cascade Slough sedge (C. obnupta), and several rush species (Juncus
mountains and Okanogan Highlands. Extensive wet falcatus, J. effusus, J. balticus) are characteristic of coastal
meadow habitats occur in Klamath, Deschutes, and dune wetlands that are included in this habitat. Several
western Lake counties in Oregon. spike rush species (Eleocharis spp.) and rush species can
Physical Setting. This habitat is found on permanently be important. Common grasses that can be local
flooded sites that are usually associated with oxbow lakes, dominants and indicators of this habitat are American
dune lakes, or potholes. Seasonally to semi-permanently sloughgrass (Beckmannia syzigachne), bluejoint reedgrass
flooded wetlands are found where standing freshwater is (Calamagrostis canadensis), mannagrass (Glyceria spp.), and
present through part of the growing season and the soils tufted hairgrass (Deschampsia caespitosa). Important
stay saturated throughout the season. Some sites are introduced grasses that increase and can dominate with
 94 Wildlife–Habitat Relationships in Oregon and Washington

disturbance in this wetland habitat include reed canary invaded by Oregon ash (Fraxinus latifolia) with fire
grass (Phalaris arundinacea), tall fescue (Festuca arundinacea) suppression.
and Kentucky bluegrass (Poa pratensis).
Effects of Management and Anthropogenic Impacts.
Aquatic beds are part of this habitat and support a
Direct alteration of hydrology (i.e., channeling, draining,
number of rooted aquatic plants, such as yellow pond lily
damming) or indirect alteration (i.e., roading or removing
(Nuphar lutea) and unrooted, floating plants such as
vegetation on adjacent slopes) results in changes in
pondweeds (Potamogeton spp.), duckweed (Lemna minor),
amount and pattern of herbaceous wetland habitat. If the
or water-meals (Wolffia spp.). Emergent herbaceous
alteration is long term, wetland systems may reestablish
broadleaf plants, such as Pacific water parsley (Oenanthe
to reflect new hydrology, e.g., cattail is an aggressive
sarmentosa), buckbean (Menyanthes trifoliata), water star-
invader in roadside ditches. Severe livestock grazing and
warts (Callitriche spp.), or bladderworts (Utricularia spp.)
trampling decreases aquatic sedge, Northwest Territory
grow in permanent and semi-permanent standing water.
sedge (Carex utriculata), bluejoint reedgrass, and tufted
Pacific silverweed (Argentina egedii) is common in coastal
hairgrass. Native species, however, such as Nebraska
dune wetlands. Montane meadows occasionally are forb-
sedge, Baltic and jointed rush (Juncus nodosus), marsh
dominated with plants such as arrowleaf groundsel
cinquefoil (Comarum palustris), and introduced species
(Senecio triangularis) or ladyfern (Athyrium filix-femina).
dandelion (Taraxacum officinale), Kentucky bluegrass,
Climbing nightshade (Solanum dulcamara), purple
spreading bentgrass (Agrostis stolonifera), and fowl
loosestrife (Lythrum salicaria), and poison hemlock (Conium
bluegrass (Poa palustris) generally increase with grazing.
maculatum) are common non-native forbs in wetland
habitats. Status and Trends. Nationally, herbaceous wetlands have
Shrubs or trees are not a common part of this declined and the Pacific Northwest is no exception. These
herbaceous habitat although willow (Salix spp.) or other wetlands receive regulatory protection at the national,
woody plants occasionally occur along margins, in patches state, and county level; still, herbaceous wetlands have
or along streams running through these meadows. been filled, drained, grazed, and farmed extensively in
the lowlands of Oregon and Washington. Montane
Other Classifications and Key References. This habitat
wetland habitats are less altered than lowland habitats
is called Palustrine emergent wetlands in Cowardin et al.53
even though they have undergone modification as well.
Other references describe this habitat.43, 44, 57, 71, 131, 132, 138, 147,
219 A keystone species, the beaver, has been trapped to near
This habitat occurs in both lotic and lentic systems. The
extirpation in parts of the Pacific Northwest and its
Oregon Gap II Project 126 and Oregon Vegetation
population has been regulated in others. Herbaceous
Landscape-Level Cover Types127 that would represent this
wetlands have decreased along with the diminished
type are wet meadow, palustrine emergent, and National
influence of beavers on the landscape. Quigley and
Wetland Inventory (NWI) palustrine shrubland.
Arbelbide181 concluded that herbaceous wetlands are
Natural Disturbance Regime. This habitat is maintained susceptible to exotic, noxious plant invasions.
through a variety of hydrologic regimes that limit or
exclude invasion by large woody plants. Habitats are 23. Westside Riparian-Wetlands
permanently flooded, semipermanently flooded, or
Christopher B. Chappell & Jimmy Kagan
flooded seasonally and may remain saturated through
most of the growing season. Most wetlands are resistant
to fire and those that are dry enough to burn usually burn
in the fall. Most plants are sprouting species and recover
quickly. Beavers play an important role in creating ponds
and other impoundments in this habitat. Trampling and
grazing by large native mammals is a natural process that
creates habitat patches and influences tree invasion and
success.
Succession and Stand Dynamics. Herbaceous wetlands
are often in a mosaic with shrub- or tree-dominated
wetland habitat. Woody species can successfully invade
emergent wetlands when this herbaceous habitat dries.
Emergent wetland plants invade open-water habitat as
soil substrate is exposed; e.g., aquatic sedge and Northwest
Territory sedge (Carex utriculata) are pioneers following
beaver dam breaks. As habitats flood, woody species Geographic Distribution. This habitat is patchily
decrease to patches on higher substrate (soil, organic distributed in the lowlands throughout the area west of
matter, large woody debris) and emergent plants increase the Cascade Crest south into northwestern California and
unless the flooding is permanent. Fire suppression can north into British Columbia. It also occurs less extensively
lead to woody species invasion in drier herbaceous at mid- to higher elevations in the Cascade and Olympic
wetland habitats; e.g., Willamette Valley wet prairies are
 CHAPTER 2: HABITAT DESCRIPTIONS 95

mountains, where it is limited to more specific ssp. lasiandra) can form woodlands on major floodplains
environments. or co-dominate with other willows in tall shrublands.
Oregon white oak (Quercus garryana) and California black
Physical Setting. This habitat is characterized by wetland
oak (Q. kelloggii) can be important in the interior valleys
hydrology or soils, periodic riverine flooding, or perennial
of western Oregon. Conifers that frequently dominate or
flowing freshwater. The climate varies from very wet to
co-dominate include western redcedar (Thuja plicata),
moderately dry and from mild to cold. Mean annual
western hemlock (Tsuga heterophylla), and Sitka spruce
precipitation ranges from 20 to >150 inches (51 to >381
(Picea sitchensis). Grand fir (Abies grandis) sometimes co-
cm) per year. This habitat is found at elevations mostly
dominates, especially in drier climates and riverine flood
below 3,000 ft (914 m), but it does extend up to 5,500 ft
plains. Douglas-fir (Pseudotsuga menziesii) is relatively
(1,676 m) in Washington and 6,500 ft (1,981 m) in Oregon
uncommon. Shore pine (Pinus contorta var. contorta) is
in the form of Sitka alder communities. Wetlands above
common in bogs and in deflation plain wetlands along
these elevations are generally considered part of the
the outer coast. Dominant species in tall shrublands
Subalpine Parkland habitat and are not included here.
include Sitka willow (Salix sitchensis), Hooker’s willow (S.
Topography is typically flat to gently sloping or
hookeriana), Douglas’ spirea (Spirea douglasii), red-osier
undulating, but can include moderate to steep slopes in
dogwood (Cornus sericea), western crabapple (Malus fusca),
the mountains. Geology is extremely variable. Gleyed or
salmonberry (Rubus spectabilis), stink currant (Ribes
mottled mineral soils, organic soils, or alluvial soils are
bracteosum), devil’s-club (Oplopanax horridum), and sweet
typical. Flooding regimes include permanently flooded
gale (Myrica gale). Labrador-tea (Ledum groenlandicum, L.
(aquatic portions of small streams), seasonally flooded,
glandulosum), western swamp-laurel (Kalmia microphylla),
saturated, and temporarily flooded. Nutrient-poor acidic
sweet gale, and salal (Gaultheria shallon) often dominate
bogs, except those higb in the mountains, are considered
sphagnum bogs. Vine maple (Acer circinatum) or Sitka
part of this habitat.
alder (Alnus viridis ssp. sinuata) dominate tall shrublands
Landscape Setting. This habitat typically occupies patches in the mountains that are located on moist talus or in snow
or linear strips within a matrix of forest or regrowing avalanche tracks.
forest. The most frequent matrix habitat is Westside Forests and willow, spirea, and dogwood shrublands
Lowlands Conifer-Hardwood Forest. If not forest, the within this habitat are limited to the area west of the
matrix can be Agriculture, Urban, or Coastal Dunes and Cascade Crest. Oregon ash communities occur primarily
Beaches habitats, or rarely Westside Grasslands or in the southern Puget Lowland (King County south),
Ceanothus-Manzanita Shrublands. This habitat also forms Willamette Valley, and Klamath Mountains ecoregions.
mosaics with or includes small patches of Herbaceous White alder occurs only in the Willamette Valley and
Wetlands. Open Water habitat is often adjacent to Westside southwestern Oregon. Sitka spruce communities are
Riparian-Wetlands. The major land use of the forested mainly found in the Coast Range area and western
portions of this habitat is timber harvest. Livestock grazing Olympic Peninsula in areas of coastal fog influence.
occurs in some areas. Peat mining occurs in some bogs. Western hemlock and western redcedar riparian and
wetland habitats are largely absent from the southern
Structure. Most often this habitat is either a tall (6-30 ft [2-
Oregon Cascades and the Klamath Mountains. Sitka alder
9 m]) deciduous broadleaf shrubland, woodland or forest,
and vine maple communities are located in the mountains,
or some mosaic of these. Short to medium-tall evergreen
mainly in western Washington but to a lesser degree on
shrubs or graminoids and mosses dominate portions of
the east slope of the Cascades and in the Oregon Cascades.
bogs. Trees are evergreen conifers or deciduous broadleaf
Sweet gale communities are found primarily at low
or a mixture of both. Conifer-dominated wetlands in the
elevations on the western Olympic Peninsula. Lodgepole
lowlands are included here, whereas mid-elevation conifer
pine-dominated communities are found as bogs in western
sites are part of Montane Coniferous Wetlands habitat.
Washington and along the outer coast of Oregon. Most
Height of the dominant vegetation can be >200 ft (62 m).
sphagnum bogs are found in low elevation western
Canopy height and structure vary greatly. Typical
Washington.
understories are composed of shrubs, forbs, and/or
Shrubs that commonly dominate underneath a tree
graminoids. Water is sometimes present on the surface
layer include salmonberry, salal, vine maple, red-osier
for a portion of the year. Large woody debris is abundant
dogwood, stink currant, Labrador-tea, devil’s-club,
in late seral forests and adjacent stream channels. Small
thimbleberry (Rubus parviflorus), common snowberry
stream channels and small backwater channels on larger
(Symphoricarpos albus), beaked hazel (Corylus cornuta), and
streams are included in this habitat.
Pacific ninebark (Physocarpus capitatus). Understory
Composition. Red alder (Alnus rubra) is the most dominant herbs include slough sedge (Carex obnupta),
widespread tree species, but is absent from sphagnum Dewey sedge (C. deweyana), Sitka sedge (C. aquatilis var.
bogs. Other deciduous broadleaf trees that commonly dives), skunk-cabbage (Lysichiton americanus), coltsfoot
dominate or co-dominate include black cottonwood (Petasites frigidus), great hedge-nettle (Stachys ciliata),
(Populus balsamifera ssp. trichocarpa), bigleaf maple (Acer youth-on-age (Tolmiea menziesii), ladyfern (Athyrium filix-
macrophyllum), Oregon ash (Fraxinus latifolia), and, locally, femina), oxalis (Oxalis oregana, O. trillifolia), stinging nettle
white alder (Alnus rhombifolia). Pacific willow (Salix lucida (Urtica dioica), swordfern (Polystichum munitum), great
 96 Wildlife–Habitat Relationships in Oregon and Washington

burnet (Sanguisorba officinalis), scouring-rush (Equisetum ungulates, e.g., elk, can have a major effect on vegetation.
hyemale), blue wildrye (Elymus glaucus), Pacific golden-
Succession and Stand Dynamics. Riparian, i.e.,
saxifrage (Chrysosplenium glechomifolium), and field
streamside, habitats are extremely dynamic.162 Succession
horsetail (Equisetum arvense). Bogs often have areas
varies greatly depending on the hydro-geomorphic
dominated by •1 species of sedge (Carex spp.) or beakrush
environment. A typical sequence on a riparian terrace on
(Rhynchospora alba) and sphagnum moss (Sphagnum spp.)
a large stream involves early dominance by Sitka willow,
that are included within this habitat, despite their lack of
mid-seral dominance by red alder or cottonwood, with a
woody vegetation. Sphagnum moss is a major ground
gradual increase in conifers, and eventual late-seral
cover in most bogs.
dominance of spruce, redcedar, and/or hemlock. Such a
Other Classifications and Key References. This habitat sequence corresponds with increasing terrace height above
includes all palustrine, forested wetlands and scrub-shrub the bankfull stream stage.85 Some communities in bogs or
wetlands at lower elevations on the westside as well as a depressional wetlands, as opposed to riverine, seem to be
small subset of persistent emergent wetlands, those within relatively stable given a particular flooding regime and
sphagnum bogs.53 However, drier portions of this habitat environment. Successional sequences are not completely
in riparian flood plains may not qualify as wetlands understood and can be complex. Beaver dams or other
according to Cowardin’s definition.53 They are associated alterations of flood regime often result in vegetation
with both lentic and lotic systems. Much of this habitat is changes.
probably not mapped as distinct types by the Gap projects
Effects of Management and Anthropogenic Impacts.
because of its relatively small scale on the landscape and
Intense logging disturbance in conifer or mixed riparian
the difficulty of distinguishing forested wetlands. A
or wetland forests, except bogs, often results in
portion of this habitat is mapped as the Oregon Gap II
establishment of red alder, and its ensuing long-term
Project126 and Oregon Vegetation Landscape-Level Cover
dominance. Salmonberry responds similarly to this
Types127 westside cottonwood riparian gallery, palustrine
disturbance and tends to dominate the understory.
forest, palustrine shrubland, NWI (National Wetland
Logging activities reduce amounts of large woody debris
Inventory) palustrine emergent, NWI estuarine emergent,
in streams and remove sources of that debris.27 Timber
and alder/cottonwood riparian gallery. In the Washington
harvest can also alter hydrology, most often resulting in
Gap project, this habitat occupies portions of open water/
post-harvest increases in peak flows.107 Mass wasting and
wetlands (especially riparian), hardwood forest, and
related disturbances (stream sedimentation, debris
mixed hardwood/conifer forest, and to a minor degree,
torrents) in steep topography increase in frequency with
conifer forest in the following zones: Western hemlock,
road building and timber harvest.198 Roads and other water
Sitka spruce, Olympic Douglas-fir, Puget Sound Douglas-
diversion/retention structures change watershed
fir, Cowlitz River, Willamette Valley, and Woodland/
hydrology with wide-ranging and diverse effects, 93
prairie mosaic.37 This habitat also occupies much of
including major vegetation changes. The most significant
hardwood forest in the Silver fir, Mountain hemlock,
of these are the major flood controlling dams, which have
portions of Subalpine fir, Interior western hemlock/
greatly altered the frequency and intensity of bottomland
redcedar, and Grand fir zones.37 Other references describe
flooding. Increases in nutrients and pollutants are other
this habitat.41, 71, 85, 88, 90, 91, 104, 113, 114, 115, 138, 210, 220
common anthropogenic impacts, the former with
Natural Disturbance Regime. The primary natural particularly acute effects in bogs. Reed canarygrass
disturbance is flooding. Flooding frequency and intensity (Phalaris arundinacea) is an abundant non-native species
vary greatly with hydro-geomorphic setting. Floods can in low-elevation, disturbed settings dominated by shrubs
create new surfaces for primary succession, erode existing or deciduous trees. Many other exotic species also occur.
streambank communities, deposit sediment and nutrients
Status and Trends. This habitat occupies relatively small
on existing communities, and selectively kill species not
areas and has declined greatly in extent with conversion
adapted to a particular duration or intensity of flood. Most
to urban development and agriculture. What remains is
plant communities are more or less adapted to a particular
mostly in poor condition, having experienced any of
flooding regime,138 or they occupy a specific time in a
various anthropogenic impacts that have degraded the
successional sequence after a major disturbance.85 Debris
functionality of these ecosystems: channeling, diking,
flows/torrents are also an important, typically infrequent,
dams, logging, road-building, invasion of exotic species,
and severe disturbance where topography is
changes in hydrology and nutrients, and livestock grazing.
mountainous.200 Fires were probably infrequent or absent
Current threats include all of the above as well as
because of the combination of landscape position and site
development. Some protection has been afforded to this
moisture, although fires within the watershed would
habitat through government regulations that vary in their
usually have effects on the habitat through impacts on
scope and enforcement with jurisdiction. Of the 77 plant
flooding, sedimentation, and large woody debris inputs.
associations representing this habitat in the National
Windthrow of trees can also be significant, especially near
Vegetation Classification, almost half are considered
the outer coast or on saturated soils. Beavers act as
imperiled or critically imperiled.10
important disturbances by changing the hydrology of a
stream system through dams. Grazing by native
 CHAPTER 2: HABITAT DESCRIPTIONS 97

24. Montane Coniferous Wetlands is often a prominent feature, although it can be lacking on
Christopher B. Chappell & Jimmy Kagan less productive sites.
Composition. Indicator tree species for this habitat, any
of which can be dominant or co-dominant, are Pacific
silver fir (Abies amabilis), mountain hemlock (Tsuga
mertensiana), and Alaska yellow-cedar (Chamaecyparis
nootkatensis) on the westside, and Engelmann spruce (Picea
engelmannii), subalpine fir (Abies lasiocarpa), lodgepole pine
(Pinus contorta), western hemlock (T. heterophylla), or
western redcedar (Thuja plicata) on the eastside. Lodgepole
pine is prevalent only in wetlands of eastern Oregon.
Western hemlock and redcedar are common associates
with silver fir on the westside. They are diagnostoc of this
habitat on the east slope of the central Washington
Cascades, and in the Okanogan Highlands. Douglas-fir
(Pseudotsuga menziesii) and grand fir (Abies grandis) are
sometimes prominent on the eastside. Quaking aspen
Geographic Distribution. This habitat occurs in (Populus tremuloides) and black cottonwood (P. balsamifera
mountains throughout much of Washington and Oregon, ssp. trichocarpa) are in certain instances important to co-
except the Basin and Range of southeastern Oregon, the dominant, mainly on the eastside.
Klamath Mountains of southwestern Oregon, and the Dominant or co-dominant shrubs include devil’s-club
Coast Range of Oregon. This includes the Cascade Range, (Oplopanax horridus), stink currant (Ribes bracteosum), black
Olympic Mountains, Okanogan Highlands, Blue and currant (R. hudsonianum), swamp gooseberry (R. lacustre),
Wallowa mountains. salmonberry (Rubus spectabilis), red-osier dogwood
(Cornus sericea), Douglas’ spirea (Spirea douglasii), common
Physical Setting. This habitat is typified as forested
snowberry (Symphoricarpos albus), mountain alder (Alnus
wetlands or floodplains with a persistent winter snow
incana), Sitka alder (Alnus viridis ssp. sinuata), Cascade
pack, ranging from moderately to very deep. The climate
azalea (Rhododendron albiflorum), and glandular Labrador-
varies from moderately cool and wet to moderately dry
tea (Ledum glandulosum). The dwarf shrub bog blueberry
and very cold. Mean annual precipitation ranges from
(Vaccinium uliginosum) is an occasional understory
about 35 to >200 inches (89 to >508 cm). Elevation is mid-
dominant. Shrubs more typical of adjacent uplands are
to upper montane, as low as 2,000 ft (610 m) in northern
sometimes co-dominant, especially big huckleberry (V.
Washington, to as high as 9,500 ft (2,896 m) in eastern
membranaceum), oval-leaf huckleberry (V. ovalifolium),
Oregon. Topography is generally mountainous and
grouseberry (V. scoparium), and fools huckleberry
includes everything from steep mountain slopes to nearly
(Menziesia ferruginea).
flat valley bottoms. Gleyed or mottled mineral soils,
Graminoids that may dominate the understory include
organic soils, or alluvial soils are typical. Subsurface water
bluejoint reedgrass (Calamagrostis canadensis), Holm’s
flow within the rooting zone is common on slopes with
Rocky Mountain sedge (Carex scopulorum), widefruit sedge
impermeable soil layers. Flooding regimes include
(C. angustata), and fewflower spikerush (Eleocharis
saturated, seasonally flooded, and temporarily flooded.
quinquiflora). Some of the most abundant forbs and ferns
Seeps and springs are common in this habitat.
are ladyfern (Athyrium filix-femina), western oakfern
Landscape Setting. This habitat occurs along stream (Gymnocarpium dryopteris), field horsetail (Equisetum
courses or as patches, typically small, within a matrix of arvense), arrowleaf groundsel (Senecio triangularis), two-
Montane Mixed Conifer Forest, or less commonly, Eastside flowered marshmarigold (Caltha leptosepala ssp. howellii),
Mixed Conifer Forest or Lodgepole Pine Forest and false bugbane (Trautvetteria carolinensis), skunk-cabbage
Woodlands. It also can occur adjacent to other wetland (Lysichiton americanus), twinflower (Linnaea borealis),
habitats: Eastside Riparian-Wetlands, Westside Riparian- western bunchberry (Cornus unalaschkensis), clasping-
Wetlands, or Herbaceous Wetlands. The primary land uses leaved twisted-stalk (Streptopus amplexifolius), singleleaf
are forestry and watershed protection. foamflower (Tiarella trifoliata var. unifoliata), and five-
leaved bramble (Rubus pedatus).
Structure. This is a forest or woodland (>30% tree canopy
cover) dominated by evergreen conifer trees. Deciduous Other Classifications and Key References. This habitat
broadleaf trees are occasionally co-dominant. The includes nearly all of the wettest forests within the Abies
understory is dominated by shrubs (most often deciduous amabilis and Tsuga mertensiana zones of western
and relatively tall), forbs, or graminoids. The forb layer is Washington and northwestern Oregon and most of the
usually well developed even where a shrub layer is wet forests in the Tsuga heterophylla and Abies lasiocarpa
dominant. Canopy structure includes single-storied zones of eastern Oregon and Washington.88 On the
canopies and complex multi-layered ones. Typical tree eastside, they may extend down into the Abies grandis zone
sizes range from small to very large. Large woody debris also. This habitat is not well represented by the Gap
 98 Wildlife–Habitat Relationships in Oregon and Washington

projects because of its relatively limited acreage and the reduces large woody debris and canopy structural
difficulty of identification from satellite images. But in the complexity. Timber harvest on some sites can cause the
Oregon Gap II Project 126 and Oregon Vegetation water table to rise and subsequently prevent trees from
Landscape-Level Cover Types127 the vegetation types that establishing.221 Wind disturbance can be greatly increased
include this type would be higher elevation palustrine by timber harvest in or adjacent to this habitat.
forest, palustrine shrubland, and NWI palustrine
Status and Trends. This habitat is naturally limited in its
emergent. These are primarily palustrine forested
extent and has probably declined little in area over time.
wetlands with a seasonally flooded, temporarily flooded,
Portions of this habitat have been degraded by the effects
or saturated flooding regime.54 They occur in both lotic
of logging, either directly on site or through geohydrologic
and lentic systems. Other references describe this habitat.36,
57, 90, 101, 108, 111, 114, 115, 118, 123, 132, 221 modifications. This type is probably relatively stable in
extent and condition, although it may be locally declining
Natural Disturbance Regime. Flooding, debris flow, fire, in condition because of logging and road building. Five
and wind are the major natural disturbances. Many of of 32 plant associations representing this habitat listed in
these sites are seasonally or temporarily flooded. Floods the National Vegetation Classification are considered
vary greatly in frequency depending on fluvial position. imperiled or critically imperiled.10
Floods can deposit new sediments or create new surfaces
for primary succession. Debris flows/torrents are major 25. Eastside Riparian-Wetlands
scouring events that reshape stream channels and riparian
Rex C. Crawford & Jimmy Kagan
surfaces, and create opportunities for primary succession
and redistribution of woody debris. Fire is more prevalent
east of the Cascade Crest. Fires are typically high in
severity and can replace entire stands, as these tree species
have low fire resistance. Although fires have not been
studied specifically in these wetlands, fire frequency is
probably low. These wetland areas are less likely to burn
than surrounding uplands, and so may sometimes escape
extensive burns as old forest refugia.1 Shallow rooting and
wet soils are conducive to windthrow, which is a common
small-scale disturbance that influences forest patterns.
Snow avalanches probably disturb portions of this habitat
in the northwestern Cascades and Olympic mountains.
Fungal pathogens and insects also act as important small-
scale natural disturbances.
Succession and Stand Dynamics. Succession has not been
well studied in this habitat. Following disturbance, tall Geographic Distribution. Riparian and wetland habitats
shrubs may dominate for some time, especially mountain dominated by woody plants are found throughout eastern
alder, stink currant, salmonberry, willows (Salix spp.), or Oregon and eastern Washington.
Sitka alder. Quaking aspen and black cottonwood in these Mountain alder-willow riparian shrublands are major
habitats probably regenerate primarily after floods or fires, habitats in the forested zones of eastern Oregon and
and decrease in importance as succession progresses. eastern Washington. Eastside lowland willow and other
Lodgepole pine is often associated with post-fire riparian shrublands are the major riparian types
conditions in eastern Oregon, 131 although in some throughout eastern Oregon and Washington at lower
wetlands it can be an edaphic climax species. Pacific silver elevations. Black cottonwood riparian habitats occur
fir, subalpine fir, or Engelmann spruce would be expected throughout eastern Oregon and Washington, at low to
to increase in importance with time since the last major middle elevations. White alder riparian habitats are
disturbance. Western hemlock, western redcedar, and restricted to perennial streams at low elevations, in drier
Alaska yellow-cedar typically maintain co-dominance as climatic zones in Hells Canyon at the border of Oregon,
stand development progresses because of the frequency Washington, and Idaho, in the Malheur River drainage
of small-scale disturbances and the longevity of these and in western Klickitat and southcentral Yakima counties,
species. Tree size, large woody debris, and canopy layer Washington. Quaking aspen wetlands and riparian
complexity all increase for at least a few hundred years habitats are widespread but rarely a major component
after fire or other major disturbance. throughout eastern Washington and Oregon. Ponderosa
pine-Douglas-fir riparian habitat occurs only around the
Effects of Management and Anthropogenic Impacts.
periphery of the Columbia Basin in Washington and up
Roads and clearcut logging practices can increase the
into lower montane forests.
frequency of landslides and resultant debris flows/
torrents, as well as sediment loads in streams.198, 199, 229 This Physical Setting. Riparian habitats appear along perennial
in turn alters hydrologic patterns and the composition and and intermittent rivers and streams. This habitat also
structure of montane riparian habitats. Logging typically appears in impounded wetlands and along lakes and
 CHAPTER 2: HABITAT DESCRIPTIONS 99

ponds. Their associated streams flow along low to high (B. nana) and Douglas spiraea (Spiraea douglasii) can occur
gradients. The riparian and wetland forests are usually in in wetter stands. Red-osier dogwood and common
fairly narrow bands along the moving water that follows snowberry are shade-tolerant and dominate stand
a corridor along montane or valley streams. The most interiors, while these and other shrubs occur along forest
typical stand is limited to 100-200 ft (31-61 m) from or woodland edges and openings. Mountain alder is
streams. Riparian forests also appear on sites subject to frequently a prominent shrub, especially at middle
temporary flooding during spring runoff. Irrigation of elevations. Tall shrubs (or small trees) often growing under
streamsides and toeslopes provides more water than or with white alder include chokecherry (Prunus
precipitation and is important in the development of this virginiana), water birch, shining willow, and netleaf
habitat, particularly in drier climatic regions. hackberry (Celtis reticulata).
Hydrogeomorphic surfaces along streams supporting this Shrub-dominated communities contain most of the
habitat have seasonally- to temporarily-flooded species associated with tree communities. Willow species
hydrologic regimes. Eastside riparian and wetland (Salix bebbiana, S. boothii, S. exigua, S geyeriana, or S.
habitats are found from 100- 9,500 ft (31-2,896 m) in lemmonii) dominate many sites. Mountain alder can be
elevation. dominant and is at least codominant at many sites.
Chokecherry, water birch, serviceberry (Amelanchier
Landscape Setting. Eastside riparian habitats occur along
alnifolia), black hawthorn (Crataegus douglasii), and red-
streams, seeps, and lakes within the Eastside Mixed
osier dogwood can also be codominant to dominant.
Conifer Forest, Ponderosa Pine Forest and Woodlands,
Shorter shrubs, Woods rose, spiraea, snowberry and
Western Juniper and Mountain Mahogany Woodlands,
gooseberry are usually present in the undergrowth.
and part of the Shrub-steppe habitat. This habitat may be
The herb layer is highly variable and is composed of
described as occupying warm montane and adjacent
an assortment of graminoids and broadleaf herbs. Native
valley and plain riparian environments.
grasses (Calamagrostis canadensis, Elymus glaucus, Glyceria
Structure. The Eastside riparian and wetland habitat spp., and Agrostis spp.) and sedges (Carex aquatilis, C.
contains shrublands, woodlands, and forest communities. angustata, C. lanuginosa, C. lasiocarpa, C. nebrascensis, C.
Stands are closed to open canopies and often multilayered. microptera, and C. utriculata) are significant in many
A typical riparian habitat would be a mosaic of forest, habitats. Kentucky bluegrass (Poa pratensis) can be
woodland, and shrubland patches along a stream course. abundant where heavily grazed in the past. Other weedy
The tree layer can be dominated by broadleaf, conifer, or grasses, such as orchard grass (Dactylis glomerata), reed
mixed canopies. Tall shrub layers, with and without trees, canarygrass (Phalaris arundinacea), timothy (Phleum
are deciduous and often nearly completely closed thickets. pratense), bluegrass (Poa bulbosa, P. compressa), and tall
These woody riparian habitats have an undergrowth of fescue (Festuca arundinacea) often dominate disturbed
low shrubs or dense patches of grasses, sedges, or forbs. areas. A short list of the great variety of forbs that grow in
Tall shrub communities (20-98 ft [6-30 m], occasionally tall this habitat includes Columbian monkshood (Aconitum
enough to be considered woodlands or forests) can be columbianum), alpine leafybract aster (Aster foliaceus),
interspersed with sedge meadows or moist, forb-rich ladyfern (Athyrium filix-femina), field horsetail (Equisetum
grasslands. Intermittently flooded riparian habitat has arvense), cow parsnip (Heracleum maximum), skunkcabbage
ground cover composed of steppe grasses and forbs. Rocks (Lysichiton americanus), arrowleaf groundsel (Senecio
and boulders may be a prominent feature in this habitat. triangularis), stinging nettle (Urtica dioica), California false
hellebore (Veratrum californicum), American speedwell
Composition. Black cottonwood (Populus balsamifera ssp.
(Veronica americana), and pioneer violet (Viola glabella).
trichocarpa), quaking aspen (P. tremuloides), white alder
(Alnus rhombifolia), peachleaf willow (Salix amygdaloides), Other Classifications and Key References. This habitat
and, in northeast Washington, paper birch (Betula is called Palustrine scrub-shrub and forest in Cowardin et
papyrifera) are dominant and characteristic tall deciduous al.53 Other references describe this habitat.44, 57, 60, 131, 132, 147, 156
trees. Water birch (B. occidentalis), shining willow (Salix This habitat occurs in both lotic and lentic systems. The
lucida ssp. caudata), and, rarely, mountain alder (Alnus Oregon Gap II Project 126 and Oregon Vegetation
incana) are co-dominant to dominant mid-size deciduous Landscape-Level Cover Types127 that would represent this
trees. Each can be the sole dominant in stands. Conifers type are eastside cottonwood riparian gallery, palustrine
can occur in this habitat, rarely in abundance, more often forest, palustrine shrubland, and National Wetland
as individual trees. The exceptions are ponderosa pine Inventory (NWI) palustrine emergent.
(Pinus ponderosa) and Douglas-fir (Pseudotsuga menziesii)
Natural Disturbance Regime. This habitat is tightly
that characterize a conifer-riparian habitat in portions of
associated with stream dynamics and hydrology. Flood
the shrub-steppe zones.
cycles occur within 20-30 years in most riparian
A wide variety of shrubs are found in association with
shrublands although flood regimes vary among stream
forest/woodland versions of this habitat. Red-osier
types. Fires recur typically every 25-50 years but fire can
dogwood (Cornus sericea), mountain alder, gooseberry
be nearly absent in colder regions or on topographically
(Ribes spp.), rose (Rosa spp.), common snowberry
protected streams. Rafted ice and logs in freshets may
(Symphoricarpos albus) and Drummonds willow (Salix
cause considerable damage to tree boles in mountain
drummondii) are important shrubs in this habitat. Bog birch
 100 Wildlife–Habitat Relationships in Oregon and Washington

habitats. Beavers crop younger cottonwood and willows 26. Coastal Dunes and Beaches
and frequently dam side channels in these stands. These Christopher B. Chappell, David H. Johnson, & Jimmy Kagan
forests and woodlands require various flooding regimes
and specific substrate conditions for reestablishment.
Grazing and trampling is a major influence in altering
structure, composition, and function of this habitat; some
portions are very sensitive to heavy grazing.
Succession and Stand Dynamics. Riparian vegetation
undergoes “typical” stand development that is strongly
controlled by the site’s initial conditions following
flooding and shifts in hydrology. The initial condition of
any hydrogeomorphic surface is a sum of the plants that
survived the disturbance, plants that can get to the site,
and the amount of unoccupied habitat available for
invasions. Subsequent or repeated floods or other
influences on the initial vegetation selects species that can
survive or grow in particular life forms. A typical woody
riparian habitat dynamic is the invasion of woody and
herbaceous plants onto a new alluvial bar away from the Geographic Distribution. This habitat occurs primarily
main channel. If the bar is not scoured in 20 years, a tall along the outer coast from southern Washington (Grays
shrub and small deciduous tree stand will develop. Harbor County) south to northern California. It occurs in
Approximately 30 years without disturbance or change all coastal Oregon counties, most abundantly in Tillamook
in hydrology will allow trees to overtop shrubs and form County and between Florence and Reedsport. In
woodland. Another 50 years without disturbance will Washington it occurs mainly in Grays Harbor and Pacific
allow conifers to invade and in another 50 years a mixed counties, and sporadically along the inland marine waters
hardwood-conifer stand will develop. Many deciduous of Clallam, San Juan, Skagit, Jefferson, Whatcom, King,
tall shrubs and trees cannot be invaded by conifers. Each Pierce, Kitsap, Snohomish, and Island counties. It also
stage can be reinitiated, held in place, or shunted into occurs in British Columbia.
different vegetation by changes in stream or wetland
Physical Setting. This habitat occurs primarily in wet,
hydrology, fire, grazing, or an interaction of those factors.
mild outer coastal climates. Precipitation, almost always
Effects of Management and Anthropogenic Impacts. rain, typically averages >80 inches (203 cm) annually.
Management effects on woody riparian vegetation can be Summers are relatively dry, but fog is common. Elevation
obvious, e.g., removal of vegetation by dam construction, is at and very near sea level, only extending as high as the
roads, logging, or they can be subtle, e.g., removing highest dunes. Topography is mildly to strongly
beavers from a watershed, removing large woody debris, undulating in the form of mostly north-south trending
or construction of a weir dam for fish habitat. In general, dune ridges and troughs. Soils, when present, are always
excessive livestock or native ungulate use leads to less sandy and are underlain by deep deposits of sand, thereby
woody cover and an increase in sod-forming grasses creating edaphically dry sites. Soils are also very poor in
particularly on fine-textured soils. Undesirable forb nutrients and organic matter. These dunes, spits, and
species, such as stinging nettle and horsetail, increase with berms are derived from sand carried by longshore drift
livestock use. and wind erosion. Dunes consist of several types that differ
in their physical form, including foredunes, transverse
Status and Trends. Quigley and Arbelbide181 concluded
dunes, parabola dunes, and retention ridges.220 Outlier
that the Cottonwood-Willow cover type covers
examples away from the outer coast in the Puget Trough
significantly less in area now than before 1900 in the Inland
are small in extent, occur in a drier climate, and mainly
Pacific Northwest. The authors concluded that although
occur in the form of sand spits and berms as opposed to
riparian shrubland was a minor part of the landscape,
dunes.
occupying 2%, they estimated it to have declined to 0.5%
of the landscape. Approximately 40% of riparian Landscape Setting. This habitat occurs in a natural mosaic
shrublands occurred above 3,280 ft (1,000 m) in elevation with Westside Lowland Conifer-Hardwood Forest,
before1900; now nearly 80% is found above that elevation. Westside Riparian-Wetlands, and Herbaceous Wetlands.
This change reflects losses to agricultural development, Forests adjacent to this habitat are found on stabilized
roading, dams and other flood-control activities. The dunes and are dominated by shore pine (Pinus contorta
current riparian shrublands contain many exotic plant var. contorta) and Sitka spruce (Picea sitchensis). Wooded,
species and generally are less productive than historically. shrubby, and herbaceous wetlands occur in seasonally
Quigley and Arbelbide181 found that riparian woodland flooded deflation plains or dune troughs. Hooker’s willow
was always rare and the change in extent from the past is (Salix hookeriana) and slough sedge (Carex obnupta) are the
substantial. two most characteristic species in these wetlands. This
habitat is in a mosaic with the Urban habitat, as coastal
 CHAPTER 2: HABITAT DESCRIPTIONS 101

areas have been developed extensively for tourism and completely stabilized shrub-dominated dunes. 220
low-density residential uses. Recreation is a major land Unstabilized sand, such as foredunes with little European
use and includes the use of off-road vehicles. In southern beachgrass, has the most open and herbaceous vegetation.
Washington and northern Oregon, the wetlands are often Closing of the vegetation typically results in stabilization
converted to agriculture for cranberries. of the sand. Recently stabilized dunes are now primarily
dominated by European beachgrass. Given more time
Structure. This habitat consists of a variable mosaic of
without a major disturbance, shrubs and/or trees colonize
structures ranging from open sand with sparse herbaceous
the grasslands. Shrublands are sometimes an intermediate
vegetation to dense shrublands. Trees are typically absent
stage in succession toward forests. Pine woodlands are
but may be scattered. Unstabilized sand may have very
another very common intermediate stage. Eventually, pine
little vegetation or open short grasslands or forb-
woodlands are colonized by Sitka spruce or Douglas-fir
dominated communities, though these are now relatively
and become mixed pine-spruce or pine-Douglas-fir forests.
uncommon and local. Medium-tall grasslands, typically
Any one of these stages can be set back to sand by a
closed, are a major component in the current landscape.
blowout or reburial by dunes, and a cyclic successional
Tall broadleaf evergreen shrubs, typically dense, are also
sequence is common in many areas.42
a significant component of the mosaic.
Effects of Management and Anthropogenic Impacts.
Composition. Where they are vegetated, unstabilized
European beachgrass has been extensively planted for
dunes or strand are typically dominated or co-dominated
stabilization purposes and has also spread widely on its
by American dunegrass (Leymus mollis), dune bluegrass
own. Unstabilized sand is now a relatively rare condition
(Poa macrantha), or Chinook lupine (Lupinus littoralis). Red
primarily because of the introduction of this species. The
fescue (Festuca rubra) was once a major dominant on more
physical forms of dunes also have been altered by
stabilized dunes but has been largely replaced by
beachgrass.55, 56 Forests are probably forming at a greater
European beachgrass (Ammophila arenaria), an introduced
rate than they did in the past because of increased
species that is now the most common dune grass. Many
stabilization. Exotic species, especially sweet vernalgrass
forb species are largely confined to herb-dominated dunes
(Anthoxanthum odoratum) and common velvetgrass (Holcus
or strand and may take on local importance.
lanatus), are now a nearly ubiquitous component of herb-
Tall shrublands are dominated primarily by salal
dominated communities. The spread of such species may
(Gaultheria shallon) and evergreen huckleberry (Vaccinium
be related to past livestock grazing in many areas.42 Scot’s
ovatum), but may also have prominent amounts of hairy
broom and gorse are aggressive exotic shrub invaders that
manzanita (Arctostaphylos columbiana), kinnikinnick
were planted for stabilization and have spread widely.
(Arctostaphylos uva-ursi), bush lupine (Lupinus arboreus),
Since both are legumes, they result in major nitrogen
or California wax-myrtle (Myrica califorica). Coyotebrush
increases where they establish. Off-road vehicle use has
(Baccharis pilularis) is abundant in southern Oregon. Both
resulted in complete destruction of native herbaceous
Scot’s broom (Cytisus scoparius) and gorse (Ulex europaeus)
communities in some areas220 Trampling is a potential
are exotic shrubs that dominate disturbed areas. Scattered
threat in herbaceous communities.42, 220
trees are mainly shore pine (Pinus contorta var. contorta),
or, less commonly, Sitka spruce (Picea sitchensis). Status and Trends. This habitat covers a relatively limited
area and major expanses of it have been converted to other
Other Classifications and Key References. Franklin and
uses. The vast majority of herbaceous vegetation that
Dryness 88 called this habitat sand dune and strand
remains is in poor condition, being dominated by exotic
communities. The Oregon Gap II Project126 and Oregon
species. Current trends are probably decreasing in both
Vegetation Landscape-Level Cover Types 127 would
extent and condition because of continued development
crosswalk with coastal dunes habitat. This habitat is not
in coastal areas and continuing expansion of exotic species
well represented by the Washington Gap project: it takes
into the few remaining native-dominated areas. Six of 11
up small percentages of several types in the Sitka spruce
plant associations currently listed in the National
zone, including conifer forest, hardwood forests, and
Vegetation Classification representing this habitat are
coastline, sandy beaches, and rocky islands. Other
considered imperiled or critically imperiled.10
references describe this habitat.8, 42, 137, 219, 220
Natural Disturbance Regime. Erosion and deposition of
sand are the primary natural processes controlling this
habitat. Sand is deposited initially on beaches, and then
moved into dunes through wind erosion.220 Wind also
maintains unstabilized dune areas. Major winter storm
events may result in blowouts that create holes in existing
stabilized or unstabilized dunes, creating new areas of
sand deposition.
Succession and Stand Dynamics. The different structural
variants of the mosaic within this habitat are primarily
stages in succession from freshly deposited sand to
 102 Wildlife–Habitat Relationships in Oregon and Washington

27. Coastal Headlands and Islets Composition. Shrublands are dominated by salal
Christopher B. Chappell & David H. Johnson (Gaultheria shallon), evergreen huckleberry (Vaccinium
ovatum), salmonberry (Rubus spectabilis), black twinberry
(Lonicera involucrata), California wax-myrtle (Myrica
californica), thimbleberry (Rubus parviflorus), or the dwarf
shrub, crowberry (Empetrum nigrum). Deer brush
(Ceanothus integerrimus), and hairy manzanita
(Arctostaphylos columbiana) become important on the
southern Oregon coast, as does the non-native gorse (Ulex
europaeus). Sitka spruce (Picea sitchensis) is the most
common tree, although western hemlock (Tsuga
heterophylla), Douglas-fir (Pseudotsuga menziesii), or red
alder (Alnus rubra) also may occur. Native dominant
grasses are red fescue (Festuca rubra) or Nootka reedgrass
(Calamagrostis nutkaensis). Blue wildrye (Elymus glaucus),
California danthonia (Danthonia californica), and Sitka
brome (Bromus sitchensis) can also be important. A diversity
of forbs occurs, with some of the most prominent being
Canada goldenrod (Solidago canadensis), Martindale’s
Geographic Distribution. This shrubland, grassland, and
lomatium (Lomatium martindalei), giant vetch (Vicia
nearshore rocky island habitat occurs along slopes and
gigantea), giant horsetail (Equisetum telmateia), and coastal
exposed headlands along the outer coast, from Cape
wormwood (Artemisia suksdorfii). Bracken (Pteridium
Flattery, Clallam County, Washington, southward to (and
aquilinum) is a fern that often co-dominates. Southern
beyond) California. On small islands it also extends into
Oregon has a number of unique herbaceous species.
the Strait of Juan de Fuca and the inland marine waters of
Puget Sound and Hood Canal, Washington. Sporadic Other Classifications and Key References. Franklin and
along the Washington coast (absent between Point Dyrness88 described portions of this habitat as oceanfront
Grenville and Cape Disappointment), this habitat becomes communities on northern Oregon headlands and the
most extensive on the southern Oregon coast. southern Oregon coast. The Oregon Gap II Project126 and
Oregon Vegetation Landscape-Level Cover Type127 that
Physical Setting. Wind is extreme in this environment and,
would represent this type is coastal strand. The
in combination with abundant salt-spray, limits tree
Washington Gap project mapped parts of this habitat as
growth. Fog is common in the summer. Climate is
coastline, sandy beaches, and rocky islands. Other
generally mild and moist to wet, with mean annual
important references describe elements of the habitat.7, 63,
precipitation ranging from about 70 to 120 inches (178 to 177, 183
305 cm). Elevation is sea level to about 500 ft (152 m). This
habitat occurs mainly on coastal headlands, bluffs, and Natural Disturbance Regime. Wind may topple trees if
islands with steep slopes or cliffs. Soils are typically they do attain upright stature. Charcoal in the soil at some
shallow to bedrock or consist of exposed glacial deposits sites in Oregon suggests that this habitat may have had
on steep erodable bluffs. Slopes range from gentle to very occasional fires in the past.183
steep. In some areas, seeps create moist to wet microsites.
Succession and Stand Dynamics. Little is known about
Landscape Setting. This habitat is always located adjacent the dynamics of this habitat. Trees slowly invade some
to, or in the case of the rock islets (“sea stacks”), within areas of this habitat. As they do so, herbaceous or shrub-
the Marine Nearshore habitat. It is found mainly along dominated vegetation declines. Fires would favor
the outer coastline where it typically occupies small areas maintenance of grasslands or forblands.
between the Marine Nearshore and Westside Lowland
Effects of Management and Anthropogenic Impacts.
Conifer-Hardwood Forest or on small islands. Cliffs are a
Livestock grazing of the grasslands results in decreasing
common feature. In far southern Oregon (Curry County),
importance of native grasses, especially bunchgrasses, and
it occupies continuous ocean-facing slopes for many miles.
increasing importance of exotic species. Sweet vernalgrass
Land use is recreation or low-density residential.
(Anthoxanthum oderatum), common velvetgrass (Holcus
Structure. This habitat is a shrubland, grassland, forbland, lanatus), and orchardgrass (Dactylus glomerata) are major
rocky island, or often a mosaic of these. The dominant exotic grass species that dominate significant areas. Gorse
shrubs may be tall or short and composed of evergreen or has invaded large areas on the southern Oregon coast.
deciduous broadleaf shrubs. Native grasses can be short
Status and Trends. This habitat occupies a very small area
or up to 5 ft (1.5 m) in height and rhizomatous or cespitose.
relative to other habitats in the Pacific Northwest.
Forbs or ferns dominate some patches. Coniferous trees
Condition of the grasslands is generally poor, with an
are sometimes scattered, occur in small clumps, or form
abundance of non-native species. Grasslands continue to
dominant patches of short wind-blasted individuals.
decline in condition and extent over time. Shrublands are
probably more stable. Three of 5 plant associations listed
 CHAPTER 2: HABITAT DESCRIPTIONS 103

in the National Vegetation Classification are considered and Islets, Marine Nearshore, and Inland Marine Deeper
imperiled,10 but portions of this habitat have not been Waters habitats. Major uses of bays and estuaries are
described at the association level. recreation, tourism, the shellfish industry, and navigation.
The terrestrial interface portions of this habitat have been
28. Bays and Estuaries extensively converted for agricultural crop production,
livestock grazing, and residential and commercial
Mikell O’Mealy & David H. Johnson
development. Water channels of many areas have been
dredged for ship navigation.
Structure. At the most seaward extent (e.g., river mouths),
water depths are shallow (mostly <20 ft [6 m]) except for
dredged channels. This habitat is strongly influenced by
the daily tides and currents. Depending on location, mean
higher high water to mean lower low water ranges from
6.1 to 10.2 ft (1.9 to 3.1 m). Tidal currents in channels of
the principal estuaries typically range from 1 to 5 knots
(0.5 to 2.6 m/sec) 176.
Diverse habitats result from riverine discharges and
tidal fluxes, salinity, mixing, sedimentation, discharge, and
insolation. Unconsolidated or consolidated tideflats are
composed of rocks, gravel, sand, silt and clay as well as
abundant organic material.68, 204 Inundated by daily tidal
Geographic Distribution. This habitat reflects areas with flows, tideflats may support eelgrass, various algal species,
significant mixing of salt and freshwater, including lower and invertebrate communities.5 Eelgrass meadows create
reaches of rivers, intertidal sand and mud flats, saltwater protected environments and structured habitats for
and brackish marshes, and open-water portions of various wildlife species.173 Salt marshes form at the upper
associated bays. The habitat is distributed along the marine tidal boundary above tideflats.120 Salt marshes are usually
coast and shoreline of Washington and Oregon. There are open to closed graminoid or forb communities. Highly
some 21 principal bays and estuaries on the Oregon coast, branched estuarine channels drain across salt marshes and
and 34 in Washington.5, 95, 178 Willapa Bay and Grays Harbor tideflats, creating a diverse mix of structures.192 At the most
(both in Washington) are expansive and have the largest inland extent of this habitat, transitional marsh forms
and second largest intertidal areas of the two states. The between salt marshes and bordering upland vegetation
Columbia River Estuary is the largest estuary in the Pacific dominated by grass or woody vegetation.6
Northwest. This habitat does not include open water areas The Columbia River estuary is characterized as a
of Puget Sound (see Inland Marine Deeper Waters). Similar partially mixed estuary and can be divided into three
bay and estuarine habitats exist on the coasts of California sections along the salinity gradient: from the mouth to
and British Columbia. about river mile 7 it is basically marine; from river mile 7
The greater Puget Sound at times is considered a very to mile 23 it is transitional (mixing); and above river mile
large estuary;180 for purposes of this project Puget Sound 23 it is fluvial (fresh water). Pruter and Alverson 179
is comprised of three wildlife habitats: Bays and Estuaries, compiled available physical and biological studies at the
Marine Nearshore, and Inland Marine Deeper Waters. interface between riverine and marine waters in the
nearshore aspects of the Columbia River estuary and
Physical Setting. Climate is moderated by the Pacific
adjacent waters.
Ocean and is usually mild. Mean temperatures at coastal
stations generally range from 40 to 70½F (4-21½C) year- Composition. Eelgrass meadows stabilize submerged
round with little north-south variation within Washington tideflats and are co-dominated by surfgrass and eelgrass
and Oregon. Annual rainfall along the coastal zone species. Three diagnostic surfgrass species (Phyllospadix
averages 80-90 inches (203-229 cm) and is concentrated in scouleri, P. torreyi, and P. serrulatus) occur on rocky
winter months, producing correspondingly high river substrates in exposed waters, whereas two species of
runoff to bays and estuaries.173, 176. Elevation is at sea level eelgrasses (Zostera marina, Z. japonica) are characteristic
to a few feet above. Coastal zone topography is of mud or mixed mud-sand substrates in areas sheltered
characterized by long stretches of sandy beaches broken from turbulent waters.68, 173 Highly productive macroalgae
by steep rocky cliffs, rocky headlands, and the mouths of that dominate estuarine channels include various blue-
bays and estuaries. Organics, silt, and sand are the primary green algae, green algae (Enteromorpha spp.) and rockweed
substrate components of this habitat and vary in specific (Fucus spp.).192 Tideflats bordering salt marshes often are
composition and distribution with variable physical co-dominated by pickleweed (Salicornia virginica),
factors.120 arrowgrass (Triglochin maritima) and three-square rush
(Scirpus americanus).5 The transition to higher areas of the
Landscape Setting. This habitat is adjacent to Westside
low-marsh zone is indicated by the dominance of jaumea
Riparian-Wetlands, Coastal Dunes and Beaches, Westside
(Jaumea carnosa), saltgrass (Distichlis spicata), and Lyngby’s
Lowland Conifer-Hardwood Forest, Coastal Headlands
 104 Wildlife–Habitat Relationships in Oregon and Washington

sedge (Carex lyngbyei).68 Major components of mid and sediment accretion and gradual rise in land elevation,
high salt marsh areas are alkaligrass (Puccinellia pumila) changes that shift environmental conditions and permit
and Canadian sand spurry (Spergularia canadensis).120 Salt other plants to establish. Arrowgrass, pickleweed, sand
rush (Juncus lesueurii), tufted hairgrass (Deschampsia spurry, and spike rush can invade the emerging marsh,
caespitosa), Pacific silverweed (Argentina egedii) and further increasing and stabilizing substrates. Saltgrass and
spreading bentgrass (Agrostis stolonifera) are salt-tolerant sedge establish on higher areas of the marsh. When initial
upland species diagnostic of high salt marshes that colonizers die back, tufted hairgrass and salt rush may
experience freshwater runoff or riverine discharge.6 establish.5 Various exotic species have become naturalized
in Oregon and Washington, including spreading bentgrass
Other Classifications and Key References. Cowardin et
(Agrostis stolonifera) and sand spurry (Spergularia marina)
al. 53 included marine and estuarine systems of the
introduced from Europe, brass buttons (Cotula
Columbian Province. Dethier68 described a classification
coronopifolia) introduced from South Africa, and marsh
for marine and estuarine habitat types in Washington
cordgrass (Spartina alterniflora) introduced from the
State. Habitat types are defined by depth, substratum type,
Atlantic Coast of North America.6, 120 These successional
energy level, and a few modifiers. Species (plants and
stages can be disrupted by riverine or tidal scouring and
animals) are described for combinations of these physical
succession can be reinitiated at any point.
variables. The Oregon Gap II Project126 and Oregon
Vegetation Landscape-Level Cover Types127 that would Effects of Management and Anthropogenic Impacts.
represent this type are exposed tidal flats and estuarine Management, water quality, contaminants, and land-use
emergent. Harper et al. 106 described a shore-zone practices have altered significant portions of this habitat
sensitivity mapping system. Proctor et al.176 described an and continue to impact remaining areas.216 The dredging
ecological characterization of the Pacific Northwest and filling of marshes and tideflats to serve various human
Coastal Region, including physical and chemical needs remove estuarine vegetation. Channel flow, tidal
environments as well as socioeconomic aspects of inundation, and fresh water discharges are disrupted by
watersheds of the region. Schoch and Dethier189 provided construction of seawalls, jetties, dikes, and dams. The
high-resolution data on the physical features and physical and chemical conditions of these habitats are
associated biota of Puget Sound’s shorelines using the degraded by the discharge of municipal, industrial, and
SCALE model (Shoreline Classification and Landscape agricultural effluents. Functional plant and animal
Extrapolation). Downing76 offered a detailed review of the communities are altered by domestic and agricultural
geological and broad ecological development of Puget runoff of pesticides, herbicides, and fertilizers. Invasions
Sound. of exotic plants (e.g., Spartina) and invertebrates (e.g., green
crabs) pose significant, long-term ecological and economic
Natural Disturbance Regime. Natural disturbance
threats to this habitat. Large tracts of habitat have been
perpetuates the dynamic, transitional nature of this
lost and converted for coastal development. Additionally,
habitat. Tides, seasonal riverine discharges, winds, storm
upland activities occurring throughout the watershed,
events, erosion, and accretion are the primary natural
including logging, mining, and hydroelectric power
processes that shape this habitat. Tides are mixed,
development, can have destructive impacts downstream
characterized by two unequal high and low tides daily,
in estuarine and bay environments.192, 205
with varying intrusion into estuaries and bays at different
locations along the coast. Tides and winds push salt-water Status and Trends. Significant quantitative and qualitative
wedges up through the system, causing varying degrees alterations of this habitat have occurred with
of mixing with incoming riverine waters and significant Euro-American settlement. Although natural erosion and
vertical stratification.5 Riverine discharges and fresh-water accretion processes continue, most habitat modification
runoff vary seasonally with precipitation and freshet can be attributed to anthropogenic impacts.192 Because of
regimes. Generally, a large range in annual discharge exists original diking for crop production and flood control,
with high volumes of fresh water entering the system in almost no areas of natural high marsh remain in Oregon.120
winter and significantly reduced flows in summer.192 Short- These dikes, and other more recent barriers, prevent
term storm events produce dramatic variations in physical natural recovery and re-establishment of this habitat.
habitat conditions. Sudden erosion or accretion may result Remaining examples of the bay and estuarine habitat exist
from strong oceanic currents at the mouth of the system in various conditions, from the more natural areas, areas
or from increased fresh water discharges at the head of undergoing active restoration, to the more prevalent
the system. For a detailed conceptual model of disturbance polluted, degraded, or overused areas throughout Oregon
regimes in estuary zones, see Proctor et al.176 and Washington. With increasing population pressures in
coastal areas and the corresponding threats of habitat use
Succession and Stand Dynamics. General successional
and conversion, future trends will likely be continued
stages reflect unconsolidated barren tideflats to stabilized
degradation and reduction of remaining bay and estuarine
high salt marshes and salt meadows. Unvegetated tideflats
areas.
are colonized by pioneer plants, commonly eelgrass, that
are tolerant of extended tidal inundation and vary
depending on sediment type.173 Initial colonization causes
 CHAPTER 2: HABITAT DESCRIPTIONS 105

29. Inland Marine Deeper Water Lake Washington, Lake Sammamish, Hood Canal, and the
David H. Johnson major basins of Puget Sound. The arrangement of the
present shorelines was established 13,000 years ago when
glacial ice retreated from the Puget Lowland.76 Organics,
silt, and sand are the primary substrate components of
this habitat and vary in specific composition and
distribution with fluctuating physical factors.120 Through
deposition of sediments, major river deltas have advanced
substantial distances into the deep basins of Puget Sound.76
Composition. Marine waters dominate fresh water
influences in areas away from riverine discharges or from
the shoreline. Because of the water depths involved,
sunlight is diffused, and few if any plants attached to the
benthic substrates are capable of growing.
Other Classifications and Key References. Cowardin et
al.53 included this region in the Columbia Province and
described a hierarchical classification for wetlands and
deepwater habitats in the U.S. Dethier68 described a
Geographic Distribution. This habitat is located in the
classification for marine and estuarine habitat types in
northwestern portion of Washington and adjacent areas
Washington State. Habitat types were defined by depth,
of southwest British Columbia. It includes the open waters
substratum type, energy level, and a few modifiers. Harper
of the Strait of Georgia, Puget Sound, Hood Canal, and
et al.106 described a shore-zone mapping system for use in
the Strait of Juan de Fuca. More specifically, this habitat
sensitivity mapping and shoreline countermeasures.
reflects waters >66 ft (20 m) deep, found inland from a
Proctor et al.176 described an ecological characterization
line between the Elwha River (just west of Port Angeles)
of the Pacific Northwest Coastal Region, including
on the Washington side of the Strait of Juan de Fuca,
physical and chemical environments as well as
northward to Race Rocks on the southeastern tip of
socioeconomic aspects of watershed units and of the
Vancouver Island, British Columbia. This line was
region. Schoch and Dethier189 provided high-resolution
independently determined based on (1) kelp distribution,
data on the physical features and associated biota of Puget
(2) marine bird distribution, and (3) fish species and
Sound’s shorelines using the SCALE model (Shoreline
abundance data. With the exception of Marine Nearshore
Classification and Landscape Extrapolation).
areas, waters west of this line are considered Marine Shelf.
The Inland Marine Deeper Water habitat is not found in Natural Disturbance Regime. Seasonal and larger,
Oregon. periodically occurring disturbances shape this habitat.
Seasonal variation in tidal regimes, precipitation and
Physical Setting. This habitat lies largely within the Puget
riverine discharges (winter highs), as well as periodic
Lowland and northward in Georgia Strait on the east side
storm events cause changes in temperature, salinity,
of Vancouver Island, British Columbia. Mean air
energy level, and gradual or sudden erosion and accretion
temperatures generally range between 40 and 70½F (4-
in localized areas.
21½C) year round, with little north-south variation. Rainfall
averages 20 to 80 inches (50 to 200 cm) annually and is Successional and Community Dynamics. Diverse plant
concentrated in winter months, producing and invertebrate communities compete for a variety of
correspondingly high river runoff to bays, estuaries, and habitats in this region. Succession occurs in each habitat
inland marine waters. area as disturbances create temporary vacancies, allowing
opportunistic species to become established.
Landscape Setting. This habitat is commonly adjacent to
Bays and Estuaries, Coastal Headlands and Islets, and Effects of Management and Anthropogenic Impacts.
Marine Nearshore habitats and merges with the Marine Land conversion, use, and management have altered
Shelf habitat in the Strait of Juan de Fuca. Inland marine significant portions of this habitat. The physical, chemical,
waters are used extensively for navigation, commercial and biological condition of some habitats are degraded
transport of goods, recreation, tourism, and fishery by both point and nonpoint discharges from municipal
operations. and industrial effluents. Functional plant and animal
communities are altered by domestic and agricultural
Structure. A diversity of underwater structures are created
runoff of pesticides, herbicides, and fertilizers. Large
as swift tidal currents circulate waters of the Pacific Ocean
portions of shoreline have been converted for residential,
through the reaches of Strait of Georgia, Puget Sound,
commercial, and port development, affecting inputs into
Hood Canal and the Strait of Juan de Fuca. Aspects of
the adjacent deeper waters. Benthic communities are
geology are particularly important in understanding the
significantly impacted by maintenance dredging done to
structure and dynamics of this habitat. Glacial ice initially
support navigation and commerce. The transport of oil
excavated several long, narrow valleys that today form
and chemical substances creates the potential for harmful
 106 Wildlife–Habitat Relationships in Oregon and Washington

spills that can affect these areas for extended periods of Hardwood Forest, and Urban). It occurs in a mosaic with
time. Passage of vessels from other regions increases the Coastal Headlands and Islets.
introduction rate of exotic species which, once established,
Structure. Fresh waters drain from lands surrounding
can effectively out-compete native species.
these inland marine waters to create estuarine
Status and Trends. With the important exceptions of environments nearshore 133 (see Bays and Estuaries
locally increased sedimentation rates and contaminant habitat). Nearshore subtidal habitats are diversified by
deposition/retention, the status and trends in the physical degree of wave and current action, availability of sunlight,
and biological aspects of this habitat are poorly known. and presence of vegetation. Submerged unvegetated
habitats cover a greater area than do vegetated nearshore
30. Marine Nearshore habitats, such as salt marshes and eelgrass beds. Various
combinations of water depth, character of substrates, and
David H. Johnson
exposure to tidal action create a wide range of benthic
habitats. Sand, cobble, boulders, and hardpan are
commonly found in areas of moderate to strong currents,
whereas silt and clay settle out in protected inlets and
bays.67, 145
Composition. This habitat supports marine organisms
capable of withstanding short-term exposure to air. Bottom
substrates in exposed areas are generally rock or sand,
but can include cobble or gravel. The subtidal photic zone
includes the region from mean low low water (MLLW or
the 0 ft depth) to about -50 ft (-15 m) where water is deep
enough to prevent sufficient light penetration to the
marine floor for primary productivity of kelp and other
marine plants. The rocky-bottom intertidal habitats
support kelps (Laminaria spp., Lessoniopsis spp.,
Hedophyllum sessile), brown rockweed (Pelvetiopsis limitata),
Geographic Setting. This habitat reflects marine water red algae (Iridaea spp.), and surfgrass (Phyllospadix scouleri),
areas (high tide line to depth of 66 ft [20 m]) along as well as an abundance and variety of sessile benthic
shorelines not significantly affected by freshwater inputs invertebrates. The larger kelps, such as Macrocystis
(i.e., excludes Bays and Estuaries). This includes all marine integrifolia and Nereocystis leutkeana, are found in the rocky-
shorelines of Puget Sound, Hood Canal, San Juan Islands, bottom subtidal areas. Because of constant wave action,
Strait of Georgia, Strait of Juan de Fuca, and along the the sandy-bottom areas of the intertidal and subtidal zones
outer coastlines of Washington and Oregon. In support few or no plants. The moderate to low energy
Washington, there are 3,100 miles (4,990 km) of this intertidal and subtidal areas where sand, mud, and gravel
nearshore habitat85 (H. D. Berry, Department of Natural accumulate support eelgrass (Zostera marina and Z.
Resources, Aquatic Resources Division, Olympia, pers. japonica), and the red alga (Gracilaria pacifica).
comm.); in Oregon, there are 377 miles (607 km) of this
Other Classifications and Key References. Dethier 68
nearshore habitat (C. Barrett, Northwest Habitat Institute,
provided a detailed classification scheme for the estuary,
Corvallis, Oregon, pers. comm.). For mapping and
intertidal, and shallow subtidal areas of Washington. The
classification purposes, this habitat does not extend into,
Cowardin et al. 53 classification scheme has several
or overlap with, shallow or intertidal areas found within
limitations with regards to adopting it for marine and
Bays and Estuaries.
estuarine systems. Levings and Thom143 described 9
Physical Setting. The outer coastline of Washington and categories of nearshore habitat in Puget Sound and
Oregon can be characterized as a series of sandy beaches Georgia Basin.
interspersed with rocky headlands. This coastline is
Natural Disturbance Regimes. This habitat is strongly
oriented in a north-south direction and is subjected to long-
influenced by tidal rhythms, wave action, storm events,
fetch, high-energy waves. Nearshore areas within Puget
light penetration, and bottom substrate. Because of these
Sound, Hood Canal, and elsewhere landward from the
factors, this habitat is characterized by a high degree of
Strait of Juan de Fuca, are more protected. With the
patchiness; this patchiness leads to differences in its faunal
exception of the far-reaching Columbia River plume,194 the
makeup and use. Herbivory by marine invertebrates also
effects of coastal streams are generally local and
causes significant disturbance in plant communities, as
seasonal.170
evidenced by the direct control of kelp beds by urchin
Landscape Setting. This habitat is adjacent to the Marine populations.
Shelf, Inland Marine Deeper Water, Bays and Estuaries,
Succession and Stand Dynamics. The primary natural
and a number of terrestrial-based habitats (e.g., Coastal
processes that shape the nearshore habitats include tides,
Dunes and Beaches, Westside Lowland Conifer-
erosion, accretion, and storm events. The rocky surf zone
 CHAPTER 2: HABITAT DESCRIPTIONS 107

of the outer coast of the Olympic Peninsula includes some submarine canyons. Beyond the shelf and slope are deep
of the most complex and diverse shores in the United abyssal oceanic waters.
States.67 Here, high wave energy provides space for
Landscape Setting. This habitat is located between the
habitation for species as materials are eroded away, and
Nearshore Marine and Oceanic habitats; at about one third
by increasing the capacity of algae to acquire nutrients
of the way into the Strait of Juan de Fuca, this habitat
and use sunlight. Examples of succession can be found
adjoins the Inland Marine Deeper Water habitat.
on rocky intertidal shores where wave energy periodically
disturbs established communities, or in kelp forests where Structure. The marine shelf extends seaward from the 60-
herbivory or the scouring action of swift tidal currents ft (20 m) to the 656-ft (200 m) isobath. It is occasionally
removes vegetation. divided into the inner (0-328 ft [100 m] deep) and outer
(328-656 ft [100-200 m] deep) shelf areas, reflective of
Effects of Management and Anthropogenic Impacts. This
differing oceanographic influences. The bottom substrate
habitat reflects the interface between land and sea, and is
of the shelf is mostly sand, giving way to silt on the outer
the site of intense commercial and navigational activities,
edges, and is described as smooth as a result of sediment
such as seaports, marinas, ferry docks, and log booms. A
accumulation. Currents over the shelf tend to follow the
significant concern identified by Broadhurst35 is the site-
seasonal pattern of the oceanic currents (i.e., northward
by-site consideration of projects with no ability to account
during winter [Davidson current]). The mean surface
for and assess the cumulative environmental effects of
temperatures in summer were >5½C lower, and mean
various development activities (from small residential
salinities were 0.1-0.3% higher in upwelling areas than
projects to large commercial and industrial development
farther offshore. These conditions reflect an active
projects). Without the ability to measure or understand
upwelling process.83 The shelf areas off Oregon and
cumulative effects, managers are permitting individual
Washington are known for their heavy surface waves;
activities that may result in dramatic resource losses over
extremes of wave heights ranging from 49 to 95 ft (15 to
time. Making high-quality nearshore vegetation and
29 m) have been recorded. More typical are waves of 20 to
shoreline characteristics inventory mapping available to
33 ft (6 to 10 m) during storm events.
land-use planners, natural resource scientists, and the
public will increase opportunities to protect this habitat. Composition. The Marine Shelf, as with other deep-water
regions, does not support rooted plant life. Strickland and
Status and Trends. Shoreline modification such as
Chasan194 offered a synthesis of phytoplankton; the
bulkheading, filling, and dredging can lead to direct
Washington shelf has been ranked in the highest
habitat loss. Indirectly, it can lead to changes in the
productivity category of U.S. continental shelves.
sediment and wave energy on a beach and in adjacent
subtidal areas. One third of Puget Sound’s shorelines, Other Classifications and Key References. Strickland and
approximately 800 miles (1,287 km), has been modified.180, Chasan194 offered a synthesis of information on Coastal
190, 191
The Central Puget Sound region, with high human Washington, as related to understanding impacts of
population levels, shows the highest level of modification offshore oil and gas exploration. The National Oceanic and
overall (52%).180 In Washington there are 26 species of kelp, Atmospheric Administration 163 reflects the Final
more than any other area worldwide.77 Data on floating Environmental Impact Statement/Management Plan for
kelp along the Strait of Juan de Fuca suggest that while the Olympic Coast National Marine Sanctuary. Bottom et
kelp areas are dynamic, the overall extent of kelp has al.,33 Dodimead et al.,73 Favorite et al.,83 Thomson,203 and
remained stable during 1993-1997.180 Ware and McFarlane213 offered classification schemes and
terminology for understanding the oceanic systems in the
31. Marine Shelf Northeastern Pacific.
David H. Johnson Natural Disturbance Regimes. Currents over the shelf
tend to follow the seasonal pattern of the oceanic currents,
Geographic Setting. This habitat consists of marine waters but also are strongly influenced by local winds, bottom
along the outer coast of Washington and Oregon that are and shoreline configuration, and freshwater input. On
66-656 ft (20-200 m) deep. This also includes the western average, water flows southward in the upper 328 ft (100
portion of the Strait of Juan de Fuca (excluding the Marine m) during summer, and northward below that. Water over
Nearshore areas), west of a line from the Elwha River on the shelf flows generally northward at all depths during
the Washington side of the Strait of Juan de Fuca to Race the winter.194 In addition to ocean currents, this habitat is
Rocks on the southeastern tip of Vancouver Island. heavily influenced by freshwater infusion from the
Columbia River, the largest river on the Pacific west coast.
Physical Setting. Along the coasts of Oregon and
The Columbia River effluent amounts to approximately
Washington, the Marine Shelf (also called the Continental
60% of the freshwater entering the Pacific Ocean between
Shelf) habitat exists as a relatively shallow, flat, submerged
San Francisco and the Strait of Juan de Fuca in winter,
area, which varies from about 9 to 40 miles (14 to 64 km)
and >90% the rest of the year.21 Furthermore, the 12- to 16-
in width. At about the 656-ft (200-m) isobath, roughly the
mile (20 to 25 km)-wide Strait of Juan de Fuca, separating
edge of the marine shelf, the bottom drops off more steeply
Vancouver Island from mainland Washington, is a glacially
to the continental slope, which is indented by several major
excavated channel that is the primary avenue for vigorous
 108 Wildlife–Habitat Relationships in Oregon and Washington

estuarine exchange between the shelf and the inland and summer results in episodic upwelling of cold nutrient-
marine waters of Washington and British Columbia. rich water; the poleward surface flow in the autumn and
winter is when the downwelling prevails. The transition
Effects of Management and Anthropogenic Impacts. The
from poleward to equatorward flow occurs abruptly in
chief human influence on the ecology of Oregon and
the spring and the reverse transition occurs somewhat less
Washington marine waters is fishing, especially bottom
abruptly in the autumn.
and mid-water trawl fishing in shelf waters for flatfish
and Pacific hake. Other significant fisheries include salmon Other Classifications and Key References. A number of
(inner shelf), shrimp (outer shelf), and albacore (slope). oceanographic studies have attempted to identify distinct
The effects of overfishing have been documented. Other features or parameters to define the geographical extent
risks to the shelf environment include pollution, of the Subarctic and Subtropical Pacific waters. These
contaminants, and oil spills. studies were based mostly on temperature-salinity
properties and oxygen-salinity curves.73, 213 Favorite et al.83
Status and Trends. Until the early 1990s, the Minerals
and Thomson 203 discussed new information and
Management Service of the U.S. Department of the Interior
terminology for the oceanic systems in the Northeast
had planned to conduct Lease Sales for offshore oil and
Pacific. A recent review by Bottom et al.33 synthesized
gas exploration in federal waters on the outer marine shelf
existing information on the characteristics of the ocean
of Oregon and Washington.194 A moratorium on these
environment and the influence of ocean variability on the
leases is now in place. Designated in 1994, the Olympic
capacity of the Northeast Pacific to produce salmon.
Coast National Marine Sanctuary covers 2.11 million acres
Bottom et al.31, 32 provided information on classifications
(854,251 ha) and is managed to protect its natural resources
of subsystems of the Coastal Upwelling Domain.
while encouraging compatible commercial and
recreational uses. Natural Disturbance Regime. This deeper water habitat
is influenced by seasonal water temperatures, winds,
32. Oceanic currents, and upwelling. The shelf slope essentially defines
the pathway of the large-scale California Current, which
David H. Johnson & Greg Green
transports water eastward from mid-ocean, then
southward to California. The current is essentially driven
Geographic Setting. Deep water (>656 ft [200 m] deep),
by seasonal northwesterly winds. The relatively wide shelf
open areas of the northeast Pacific Ocean extending
off Oregon and Washington tends to isolate the California
seaward from the 656-ft (200-m) isobath along the outer
Current and its infusion of nutrients from nearshore flow.
coast of Washington and Oregon.
It is pushed farther offshore during the winter when
Physical Setting. This habitat includes the continental southeasterly winds generate the seasonal northern flow
slope, which is generally found at depths between the 656- of the more coastal Davidson Current.141
and 6,560-ft (200- and 2,000-m) isobath. In general, the 656- In addition to driving currents, seasonal winds also
ft (200-m) isobath follows a north-south line. promote upwelling of colder, nutrient-rich waters. Coastal
upwelling occurs most frequently in summer and fall,
Landscape Setting. The Oceanic habitat adjoins the
promoted by northerly and northwesterly winds. The
shallower marine waters of the Marine Shelf habitat.
upwelling season runs from April to October, with
Structure. The oceanic area off Oregon and Washington maximum intensity in July and August.18 Upwelling
includes marine waters >656 ft (200 m) deep. This region intensity is typically greatest along the southern Oregon
can be divided into two general habitats reflecting coast (Cape Blanco upwelling zone) and diminishes
geomorphic features and water depth. northward. However, it can occur anywhere along the
The continental shelf along Oregon is characterized by Oregon-Washington coast under favorable winds. Shelf
a series of oceanic banks including Daisy, Stonewall, promontories such as Heceta Bank, the submarine canyons
Perpetua, Heceta, and Silcoos banks, and a major off Washington, and the shelf edge and slope create
promontory, Cape Blanco. In contrast, the Washington biologically productive mixing zones influenced both by
shelf is furrowed by Nitinat, Juan de Fuca, Quinault, large-scale currents and upwelling. Sporadic events
Grays, Guide, Willapa, and Astoria submarine canyons, causing large-scale changes in both nearshore and offshore
remnants of the last glacial period. habitats include extreme La Niña (cold water) and El Niño
The continental slope is usually defined as depths (warm water) events, and toxic red tide blooms.
between 656 and 6,560 ft (200 and 2,000 m) deep. Beyond
Effects of Management and Anthropogenic Impacts. The
the slope is a more gradual sloping area known as the
main activities in this habitat are fishing and commercial
continental rise. The rise eventually terminates at the
transport; other than regulations to support these actions,
abyssal sea floor (approximately 13,120 ft [4,000 m] deep).
little active management for biological resources takes
Boundaries between the rise and abyssal plain are not
place in this habitat.
clearly distinct. Both the rise and abyssal sea floor are
composed largely of mud.
Marine currents define important aspects of this habitat,
as wind-driven equatorward surface flow in the spring
 CHAPTER 2: HABITAT DESCRIPTIONS 109

Literature Cited 22. Barrett, S. W., S. F. Arno, and J. P. Menakis. 1997. Fire episodes in the
1. Agee, J. K. 1993. Fire ecology of Pacific Northwest forests. Island Press, inland Northwest (1540-1940) based on fire history data. U.S. Forest
Washington, DC. Service, Intermountain Research Station. General Technical Report INT-
2. ———. 1994. Fire and weather disturbances in terrestrial ecosystems GTR-370.
of the eastern Cascades. U.S. Forest Service, Pacific Northwest 23. Bastasch, R. 1998. Waters of Oregon. A source book on Oregon’s
Research Station. General Technical Report PNW-GTR-320. water and water management. Oregon State University Press, Corvallis,
3. ———, and L. Smith. 1984. Subalpine tree establishment after fire in OR.
the Olympic Mountains, Washington. Ecology 65:810-819. 23b. Beisiinger, S. R. and D. R. Osborne. 1982. Effects of urbanization on
4. Ahlenslager, K. E. 1987. Pinus albicaulis. In W.C. Fischer, compiler.The Fire avian community organization. Condor 84: 75-83.
Effects Information System (Data base). Missoula, Montana. U.S. Forest 24. Beschta, R. L., R. E. Bilby, G. W. Brown, L. B. Holtby, and T. J. D. Hofstra.
service, Intermountain Research Station, Intermountain Fire Sciences 1987. Pages 191-232 in E. O. Salo and T. W. Cundy, editors. Streamside
Laboratory. http://www.fs.fed.us/database/feis/plants/tree/pinalb. management: forestry and fishery interactions. College of Forest
4a. Airola, T. M., and K. Buchholz. 1984. Species structure and soil Resources, University of Washington, Seattle,WA.
characteristics of five urban sites along the New Jersey Palisades. 25. Bigley, R., and S. Hull. 1992. Siouxan guide to site interpretation and
Urban Ecology 8: 149-164. forest management. Washington Department of Natural Resources,
5. Akins, G. J., and C. A. Jefferson. 1973. Coastal wetlands of Oregon. Olympia,WA.
Oregon Conservation and Development Commission, Portland, OR. 26. ———, and ———. 1995. Draft guide to plant associations on the
6. Albright, R., R. Hirschi, R. Vanbianchi, and C.Vita. 1980. Pages 449-887 in Olympic Experimental Forest. Washington Department of Natural
Coastal zone atlas of Washington, land cover/land use narratives, Resources, Olympia WA.
Volume 2. Washington State Department of Ecology, Olympia, WA. 27. Bilby, R. E., and J. W. Ward. 1991. Large woody debris characteristics
7. Aldrich, F. T. 1972. A chorological analysis of the grass balds in the and function in streams draining old growth, clear-cut, and second-
Oregon Coast Range. Ph.D. Dissertation. Oregon State University, growth forests in southwestern Washington. Canadian Journal of
Corvallis, OR. Fisheries and Aquatic Sciences 48:2499-2508.
8. Alpert, P. 1984. Inventory and analysis of Oregon coastal dunes. 28. Black, A. E., J. M. Scott, E. Strand, R.G.Wright, P. Morgan, and C. Watson.
Unpublished manuscript prepared for the Oregon Natural Heritage 1998. Biodiversity and land-use history of the Palouse Region: pre-
Program, Portland, OR. European to present. Chapter 10 in Perspectives on the land use
9. American Forests. 1998. Study documents dramatic tree loss in Puget history of North America: a context for understanding our changing
Sound area. American Forest Press Release July 14, 1998. environment. USDI/USGS. Biological Resources Division, Biological
10. Anderson, M., P. Bourgeron, M. T. Bryer, R. Crawford, L. Engelking, D. Science Report USGS/BRD-1998-003.
Faber-Langendoen, M. Gallyoun, K. Goodin, D. H. Grossman, S. Landaal, 29. Blackburn, W. H., P. T. Tueller, and R. E. Eckert Jr. 1969. Vegetation and
K. Metzler, K. D. Patterson, M. Pyne, M. Reid, L. Sneddon, and A. S. soils of the Coils Creek Watershed. Nevada Agricultural Experiment
Weakley. 1998. International classification of ecological communities: Station Bulletin R-48. Reno, NV.
terrestrial vegetation of the United States. Volume II. The National 30. ———, ———, and ———. 1969. Vegetation and soils of the Cow
Vegetation Classification System: list of types. The Nature Conservancy, Creek Watershed. Nevada Agricultural Experiment Station Bulletin
Arlington,VA. R-49. Reno, NV.
11. Arno, S. F. 1970. Ecology of alpine larch (Larix lyallii Parl.) in the Pacific 30a. Blair, R. B. 1996. Land use and avian species diversity along an urban
Northwest. Ph.D. Dissertation. University of Montana, Missoula. gradient. Ecological Applications 6: 506-519.
12. Associated Press. 1991. Census: cities takeover U.S., Statesman Journal, 31. Bottom, D. K., K. K. Jones, J. D. Rodgers, and R. F. Brown. 1989.
December 18, 1991. Management of living marine resources: a research plan for the
13. Atzet, T., and L. A. McCrimmon. 1990. Preliminary plant associations of Washington and Oregon continental margin. National Coastal
the southern Oregon Cascade Mountain Province. U.S. Forest Service, Resources Research and Development Institute, Publication No. NCRI-
PNW Region, Siskiyou National Forest, Grants Pass, OR. T-89-004.
14. ———, and D. L. Wheeler. 1982. Historical and ecological 32. ———,———, ———, and ———. 1993. Research and
perspectives on fire activity in the Klamath Geological Province of the management in the Northern California Current ecosystem. Pages
Rogue River and Siskiyou National Forests. U.S. Forest Service, Pacific 259-271 in K. Sherman, L. M. Alexander, and B. D. Gold, editors. Large
Northwest Region, Portland, OR. marine ecosystems: stress, mitigation, and sustainability. AAAS Press,
15. ———, and ———. 1984. Preliminary plant associations of the Washington DC
Siskiyou Mountains Province, Siskiyou National Forest. U.S. Forest 33. ———, J. A. Lichatowich, and C. A. Frissell. 1998. Variability of Pacific
Service, Pacific Northwest Region, Portland, OR. Northwest marine ecosystems and relation to salmon production.
16. ———, ———, G. Riegel, and others. 1984. The mountain hemlock Pages 181-252 in B. R. McMurray and R. J. Bailey, editors. Change in
and Shasta red fir series of the Siskiyou Region of southwest Oregon. Pacific coastal ecosystems. National Oceanic and Atmospheric
FIR Report 6(1): 4-7. Administration Coastal Ocean Program Decision Analysis Series No.
17. ———, D.E. White, L.A. McCrimmon, P.A. Martinez, P.R. Fong, and V.D. 11. NOAA Coastal Ocean Office, Silver Spring, MD.
Randall. 1996. Field guide to the forested plant associations of 34. Brady, R. F., T. Tobius, P. F. J. Eagles, R. Ohrner, J. Micak, B.Veale, and R. S.
southwestern Oregon. U.S. Forest Service, Pacific Northwest Research Dorney. 1979. A typology for the urban ecosystem and its relationship
Paper R6-NR-ECOL-TP-17-96. to large biogeographical landscape units. Urban Ecology. 4:11-28.
18. Bakun, A. 1973. Coastal upwelling indices, west coast of North 35. Broadhurst, G. 1998. Puget Sound nearshore habitat regulatory
America, 1946-71. U.S. Department of Commerce, National Oceanic perspective: a review of issues and obstacles. Puget Sound Water
and Atmospheric Administration, National Marine Fisheries Service. Quality Action Team. Olympia, WA.
19. Barber, W. H., Jr. 1976. An autecological study of salmonberry (Rubus 36. Brockway, D. G., C. Topik, M. A. Hemstrom, and W. H. Emmingham.
spectabilis, Pursh) in western Washington. M.S.Thesis. University of 1983. Plant association and management guide for the Pacific silver fir
Washington, Seattle,WA. zone, Gifford Pinchot National Forest. U.S. Forest Service. R6-Ecol-
20. Barbour, M. G., and W. D. Billings, editors. 1988. North American 130a.
terrestrial vegetation. Cambridge University Press, New York, NY. 37. Cassidy, K. M. 1997. Land cover of Washington state: description and
21. Barnes, C. A., A. C. Duxbury, and B. A. Morse. 1972. Circulation and management.Volume 1 in K. M. Cassidy, C. E. Grue, M. R. Smith, and K.
selected properties of the Columbia River effluent at sea. Pages 41-80 M. Dvornich, editors. Washington State Gap Analysis Project Final
in A. T. Pruter and D. L. Alverson, editors. The Columbia River Estuary Report. Washington Cooperative Fish and Wildlife Research Unit,
and adjacent ocean waters, bioenvironmental studies. University of University of Washington, Seattle,WA.
Washington Press, Seattle,WA.
 110 Wildlife–Habitat Relationships in Oregon and Washington

38. Chappell, C. B. 1991. Fire ecology and seedling establishment in Shasta 58. Csuti, B., A. J. Kimerling, T. A. O’Neil, M. M. Shaughnessy, E. P. Gaines,
red fir forests of Crater Lake National Park, Oregon. M.S. Thesis. and M. M. P. Huso. 1997. Atlas of Oregon wildlife. Oregon State
University of Washington, Seattle, WA. University Press, Corvallis, OR.
39. ———, and J. K. Agee. 1996. Fire severity and tree seedling 59. Daniels, J. D. 1969. Variation and integration in the grand fir-white fir
establishment in Abies magnifica forests, southern Cascades, Oregon. complex. Ph.D. Dissertation, University of Idaho, Moscow.
Ecological Applications 6:628-640. 60. Daubenmire, R. F. 1970. Steppe vegetation of Washington. Washington
40. ———, R. Bigley, R. Crawford, and D. F. Giglio. In prep. Field guide to State University Agricultural Experiment Station Technical Bulletin No.
terrestrial plant associations of the Puget Lowland, Washington. 62.
Washington Natural Heritage Program, Department of Natural 61. ———. 1981. Subalpine parks associated with snow transfer in the
Resources, Olympia,WA. mountains of Idaho and eastern Washington. Northwest Science
41. ———, and R. C. Crawford. 1997. Native vegetation of the South 55(2):124-135.
Puget Sound prairie landscape. Pages 107-122 in P. Dunn and K. Ewing, 62. ———, and J. B. Daubenmire. 1968. Forest vegetation of eastern
editors. Ecology and conservation of the South Puget Sound prairie Washington and northern Idaho. Technical Bulletin 60. Washington
landscape. The Nature Conservancy of Washington, Seattle, WA. Agricultural Experiment Station, College of Agriculture, Washington
42. Christy, J.A ., J. S. Kagan, and A. M. Wiedemann. 1998. Plant associations State University, Pullman,WA.
of the Oregon Dunes National Recreation Area, Siuslaw National 63. Davidson, E. D. 1967. Synecological features of a natural headland
Forest, Oregon. Technical paper R6-NR-ECOL-TP-09-98. U.S. Forest prairie on the Oregon coast. M.S. Thesis. Oregon State University,
Service, Pacific Northwest Region, Portland, OR. Corvallis, OR.
43. ____, and J. A. Putera. 1993. Lower Columbia River natural area 64. Dealy, J. E. 1971. Habitat characteristics of the Silver Lake mule deer
inventory, 1992. Unpublished report to the Washington Field Office of range. U.S. Forest Service Research Paper PNW-125.
The Nature Conservancy, Seattle,Washington. Oregon Natural 65. del Moral, R. 1979. High elevation vegetation of the Enchantment
Heritage Program, Portland, OR. Lakes Basin, Washington. Canadian Journal of Botany 57(10):1111-1130.
44. ———, and J. H. Titus. 1996. Draft, wetland plant communities of 66. ———, and J. N. Long. 1977. Classification of montane forest
Oregon. Unpublished manuscript, Oregon Natural Heritage Program, community types in the Cedar River drainage of western Washington,
Portland, OR. U.S.A. Canadian Journal of Forest Research 7(2):217-225.
45. Clausnitzer, R. R., and B. A. Zamora. 1987. Forest habitat types of the 67. Dethier, M. N. 1988. A survey of intertidal communities of the Pacific
Colville Indian Reservation. Unpublished report prepared for the coastal area of Olympic National Park, Washington. Prepared for the
Department of Forest and Range Management, Washington State National Park Service and cooperating agencies.
University, Pullman,WA. 68. ———. 1990. A marine and estuarine habitat classification system for
46. Clemens, J., C. Bradley, and O. L. Gilbert. 1984. Early development of Washington State. Washington Natural Heritage Program, Department
vegetation on urban demolition sites in Sheffield, England. Urban of Natural Resources, Olympia, WA.
Ecology. 8:139-148. 69. Detling, L. E. 1961. The chaparral formation of southwestern Oregon,
47. Cochran, P. H. 1985. Soils and productivity of lodgepole pine. in D. M. with considerations of its postglacial history. Ecology 42:348-357.
Baumgartner, R. G. Krebill, J. T. Arnott, and G. F. Gordon, editors. 70. Detwyler, T. R. 1972. Urbanization and environment. Duxbury Press,
Lodgepole pine: the species and its management: symposium Belmont, CA.
proceedings, Washington State University, Cooperative Extension, 71. Diaz, N. M., and T. K. Mellen. 1996. Riparian ecological types, Gifford
Pullman,WA. Pinchot and Mt. Hood National Forests, Columbia River Gorge
48. Cole, D. N. 1977. Man’s impact on wilderness vegetation: an example National Scenic Area. U.S. Forest Service, Pacific Northwest Region,
from Eagle Cap Wilderness, NE Oregon. Ph.D. Dissertation. University R6-NR-TP-10-96.
of Oregon, Eugene, OR. 72. Dickman, A., and S. Cook. 1989. Fire and fungus in a mountain
49. ———. 1982.Vegetation of two drainages in Eagle Cap Wilderness, hemlock forest. Canadian Journal of Botany 67(7):2005-2016.
Wallowa Mountains, Oregon. U.S. Forest Service Research Paper 73. Dodimead, A. J., F. Favorite, and T. Hirano. 1963. Salmon of the North
INT-288. Pacific Ocean— Part II. Review of oceanography of the subarctic Pacific
50. Conard, S. G., A. E. Jaramillo, K. Cromack, Jr., and S. Rose, compilers. region. International Commission Bulletin No. 13.
1985.The role of the genus Ceanothus in western forest ecosystems. 74. Douglas, G. W. 1970. A vegetation study in the subalpine zone of the
General Technical Report PNW-182. U.S. Forest Service, Pacific western North Cascades, Washington. M.S. Thesis, University of
Northwest Forest and Range Experiment Station, Portland, OR. Washington, Seattle,WA.
51. ———, and S. R. Radosevich. 1981. Photosynthesis, xylem pressure 75. ———, and L. C. Bliss. 1977. Alpine and high subalpine plant
potential, and leaf conductance of three montane chaparral species in communities of the North Cascades Range, Washington and British
California. Forest Science 27(4):627-639. Columbia. Ecological Monographs 47:113-150.
52. Copeland, W. N. 1979. Harney Lake RNA Guidebook, Supplement No. 76. Downing, J. P. 1983. The coast of Puget Sound: its process and
9. U.S. Forest Service Experiment Station, Portland, OR. development. Washington Sea Grant Publication, University of
53. Cowardin, L. M., V. Carter, F. C. Golet, and E. T. LaRoe. 1979. Washington. Seattle,WA.
Classification of wetlands and deepwater habitats of the United States. 77. Druehl, L. D. 1969. The northeast Pacific rim distribution of the
U.S. Fish and Wildlife Service. FWS/OBS-79.31. Laminariales. Proceedings of the International Seaweed Symposium
54. Crawford, R. C., and H. Hall. 1997. Changes in the South Puget Sound 6:161-170.
prairie landscape. Pages 11-15 in P. Dunn and K. Ewing, editors. Ecology 78. Dunn, P.V., and K. Ewing, editors. 1997. Ecology and conservation of
and conservation of the South Puget Sound prairie landscape. The the South Puget Sound Prairie Landscape. The Nature Conservancy,
Nature Conservancy of Washington, Seattle, WA. Seattle,WA.
55. Crook, C. S. 1979. An introduction to beach and dune physical and 79. Eddleman, L. E. 1984. Ecological studies on western juniper in central
biological processes. In K. B. Fitzpatrick, editor. Articles of the Oregon Oregon. In Proceedings western juniper management short course,
Coastal Zone Management Association, Inc., Newport, OR. 1984 October 15-16. Oregon State University, Extension Service and
56. ———. 1979. A system of classifying and identifying Oregon’s coastal Department of Rangeland Resources, Corvallis, OR.
beaches and dunes. In K. B. Fitzpatrick, editor. Articles of the Oregon 80. Edwards, O. M. 1980. The alpine vegetation of Mount Rainier National
Coastal Zone Management Association, Inc., Newport, OR. Park: structure, development, and constraints. Ph.D. Dissertation.
57. Crowe, E. A., and R. R. Clausnitzer. 1997. Mid-montane wetland plant University of Washington, Seattle, WA.
associations of the Malheur, Umatilla and Wallowa-Whitman National 80a. Emlen, J. T. 1974. An urban bird community of Tucson, Arizona:
Forests. U.S. PNW Technical Paper, R6-NR-ECOL-TP-22-97. derivation, structure, regulation. The Condor 76: 184-197.
 CHAPTER 2: HABITAT DESCRIPTIONS 111

81. Everest, F. H. 1987. Salmonids of western forested watersheds. Pages 101. Hall, F. C. 1973. Plant communities of the Blue Mountains in eastern
3-38 in E. O. Salo and T. W. Cundy, editors. Streamside management: Oregon and southeastern Washington. U.S. Forest Service, R-6 Area
forestry and fishery interactions. College of Forest Resources, Guide 3-1.
University of Washington, Seattle,WA. 102. Halpern, C. B. 1989. Early successional patterns of forest species:
82. ———, R. L. Beschta, J. C. Scrivener, K. V. Koski, J. R. Sedell, and C. J. interactions of life history traits and disturbance. Ecology 70:704-720.
Cederholm. 1987. Fine sediments and salmonid production: a paradox. 103. Halverson, N. M., and W. H. Emmingham. 1982. Reforestation in the
Pages 98-142 in E. O. Salo and T. W. Cundy, editors. Streamside Cascades Pacific silver fir zone: a survey of sites and management
management: forestry and fishery interactions. College of Forest experiences on the Gifford Pinchot, Mt. Hood and Willamette National
Resources, University of Washington, Seattle. Forests. U.S. Forest Service. R6-ECOL-091-1982.
82a. Falk, J. H. 1976. Energetics of a suburban lawn ecosystem. Ecology 57: 104. ———, C. Topik, and R. van Vickle. 1986. Plant associations and
141-150. management guide for the western hemlock zone, Mt. Hood National
83. Favorite, F., A. J. Dodimead, and K. Nasu. 1976. Oceaonography of the Forest. U.S. Forest Service, R6-ECOL-232A-1986.
subarctic Pacific region, 1960-71. International North Pacific Fisheries 105. Hamann, M. J. 1972. Vegetation of alpine and subalpine meadows of
Commission Bulletin No. 33. Mount Rainier National Park, Washington. M.S. Thesis. Washington State
84. Florence, M. 1987. Plant succession on prescribed burn sites in University, Pullman.
chamise chaparral. Rangelands 9(3):119-122. 106. Harper, J. R., D. E. Howes, and P. D. Reimer. 1991. Shore-zone mapping
85. Fonda, R. W. 1974. Forest succession in relation to river terrace system for use in sensitivity mapping and shoreline countermeasures.
development in Olympic National Park, Washington. Ecology Proceedings of the 14th Arctic and Marine Oil spill Program (AMOP),
55:927-942. Environment Canada.
86. ———, and J. A. Bernardi. 1976.Vegetation of Sucia Island in Puget 107. Harr, R. D., and B. A. Coffin. 1992. Influence of timber harvest on
Sound, Washington. Bulletin of the Torrey Botanical Club 103(3):99-109. rain-on-snow runoff: a mechanism for cumulative watershed effects.
87. Franklin, J. F. 1988. Pacific Northwest forests. Pages 104-130 in M. G. Pages 455-469 in M.. E. Jones and A. Laemon, editors. Interdisciplinary
Barbour and W. D. Billings, editors. North American terrestrial approaches in hydrology and hydrogeology. American Institute of
vegetation. Cambridge University Press, New York, NY. Hydrology. Minneapolis, MN.
88. ———, and C.T. Dyrness. 1973. Natural vegetation of Oregon and 108. Hemstrom, M. A., W. H. Emmingham, N. M. Halverson, S. E. Logan, and
Washington. U.S. Pacific Northwest Forest and Range Experiment C. Topik. 1982. Plant association and management guide for the Pacific
Station, General Technical Report. PNW-8, Portland, OR. silver fir zone, Mt. Hood and Willamette National Forests. U.S. Forest
89. ———, K. Cromack, Jr., W. Denison, A. McKee, C. Maser, J. Sedell, F. Service R6-Ecol 100-1982a.
Swanson, and G. Juday. 1981. Ecological characteristics of old-growth 109. ———, and J. F. Franklin. 1982. Fire and other disturbances of the
Douglas-fir forests. U.S. Forest Service, Pacific Northwest Forest and forests in Mount Rainier National Park. Quaternary Research 18:32-51.
Range Experiment Station. General Technical Report PNW-118. 110. ———, and S.E. Logan. 1986. Plant association and management
Portland, OR. guide, Siuslaw National Forest. U.S. Forest Service Report R6-Ecol
90. ———, W. H. Moir, M. A. Hemstrom, S. E. Greene, and B. G. Smith. 220-1986a. Portland, OR.
1988. The forest communities of Mount Rainier National Park. U.S. 111. ———, ———, and W. Pavlat. 1987. Plant association and
National Park Service, Scientific Monograph Series 19, Washington, D.C. management guide, Willamette National Forest. U.S. Forest Service.
91. Frenkel, R. E., and E. F. Hieinitz. 1987. Composition and structure of R6-ECOL 257-B-86.
Oregon ash (Fraxinus latifolia) forest in William L. Finley National 112. Henderson, J. A. 1973. Composition, distribution, and succession of
Wildlife Refuge, Oregon. Northwest Science 61:203-212. subalpine meadows in Mount Rainier National Park, Washington. Ph.D.
92. Frey, D. G., editor. 1966. Limnology in North America. The University Dissertation. Oregon State University, Corvallis, OR.
of Wisconsin Press, Madison,WI. 113. ———. 1978. Plant succession on the Alnus rubra/Rubus spectabilis
93. Furniss, M. J., T. D. Roeloggs, and C. S.Yee. 1991. Road construction and habitat type in western Oregon. Northwest Science 52(3):156-167.
maintenance. Pages 297-323 in W. R. Meehan, editor. Influences of forest 114. ———, D. A. Peter, and R. Lesher. 1992. Field guide to the Forested
and rangeland management on salmonid fishes and their habitats. Plant Associations of the Mt. Baker-Snoqualmie National Forest. U.S.
American Fisheries Society Special Publication No. 19, Bethesda, MD. Forest Service Technical Paper R6-ECOL 028-91.
94. Ganskopp, D. C. 1979. Plant communities and habitat types of the 115. ———, ———, ———, and D.C. Shaw. 1989. Forested Plant
Meadow Creek Experimental Watershed. M.S. Thesis. Oregon State Associations of the Olympic National Forest. U.S. Forest Service
University, Corvallis, OR. Publication R6-ECOL-TP 001-88.
95. Gaumer, T. F., S. L. Benson, L. W. Brewer, L. Osis, D. G. Skeesick, R. M. 116. Hironaka, M., M. A. Fosberg, and A. H. Winward. 1983.
Starr, and J. F. Watson. 1985. Estuaries. In E. R. Brown, editor. Sagebrush-grass habitat types of southern Idaho. Forestry, Wildlife, and
Management of wildlife and fish habitats in forests of western Oregon Range Experiment Station Bulletin No. 15,University of Idaho, Moscow.
and Washington. U.S. Forest Service, Pacific Northwest Region, 116a. Hobbs, E. 1988. Using ordination to analyze the composition and
Portland, OR. structure of urban forest islands. Forest Ecology and Management 23:
96. Gerard, P. W. 1995. Agricultural practices, farm policy, and the 139-158.
conservation of biological diversity. USDI, National Biological Service, 117. Hopkins, W. E. 1979. Plant associations of the Fremont National
Biological Science Report 4. Forest. U.S. Forest Service Publication R6-ECOL-79-004.
97. Gordon, D. T. 1970. Natural regeneration of white and red fir: 118. ———. 1979. Plant associations of South Chiloquin and Klamath
influence of several factors. U.S. Forest Service, Research Paper PSW- Ranger Districts— Winema National Forest. U.S. Forest Service
90. Publication R6-ECOL-79-005.
97a. Green, R. J. 1984. Native and exotic birds in a suburban habitat. 119. Howard, J. L. 1996. Populus tremuloides. In D. G. Simmerman, compiler.
Australian Wildlife Research 11: 181-190. The Fire Effects Information System [Data base]. U.S. Forest Service,
98. Greenlee, J. M., and J. H. Langenheim. 1990. Historic fire regimes and Intermountain Research Station, Intermountain Fire Sciences
their relation to vegetation patterns in the Monterey Bay area of Laboratory. Missoula, MT. http://www.fs.fed.us/database/feis/plants/tree/
California. American Midland Naturalist 124(2):239-253. poptre.
99. Habeck, J. R. 1961. Original vegetation of the mid-Willamette Valley, 119a.Ingold, D. J. 1996. Delayed nesting decreased reproductive success in
Oregon. Northwest Science 35:65-77. northern flickers: implications for competition with European
100. Haeussler, S., and D. Coates. 1986. Autecological characteristics of starlings. Journal of Field Ornithology 67: 321-326.
selected species that compete with conifers in British Columbia: a 119b. Ingold, D. J. and R. J. Densmore. 1992. Competition between
literature review. Land Management Report No. 33. Ministry of Forests, European starlings and native woodpeckers for nest cavities in Ohio.
Information Services Branch,Victoria, BC, Canada. Sialia 14: 43-48.
 112 Wildlife–Habitat Relationships in Oregon and Washington

120. Jefferson, C. A. 1975. Plant communities and succession in Oregon 143. Levings, C. D., and R. M. Thom. 1994. Habitat changes in Georgia
coastal salt marshes. Ph.D. Dissertation. Oregon State University, Basin: implications for resource management and restoration. Pages
Corvallis, OR. 330-351 in R. C. H. Wilson, R. J. Beamish, F. Aitkins, and J. Bell, editors.
121. John, T., and D. Tart. 1986. Forested plant associations of the Yakima Review of the marine environment and biota of Strait of Georgia, Puget
Drainage within the Yakima Indian Reservation. Review copy prepared Sound and Juan de Fuca Strait. Canadian Technical Report of Fisheries
for the Yakima Indian Nation—Bureau of Indian Affairs Soil and Aquatic Sciences. No. 1948.
Conservation Service. 144. Lillybridge, T. R., B. L. Kovalchik, C. K. Williams, and B. G. Smith. 1995.
122. Johnson, C. G., and R. R. Clausnitzer. 1992. Plant associations of the Field guide for forested plant association of the Wenatchee National
Blue and Ochoco mountains. U.S. Forest Service, Pacific Northwest Forest. U.S. Forest Service General Technical Report PNW-GTR-359,
Region, Wallowa-Whitman National Forest R6-ERW-TP-036-92. Portland, OR.
123. ———, and S.A. Simon. 1987. Plant associations of the 145. Little, C., and J. A. Kitching. 1996. The biology of rocky shores.
Wallowa-Snake Province. U.S. Forest Service R6-ECOL-TP-255A-86. Oxford University Press, New York, NY.
124. Keeley, J. E. 1975. Longevity of nonsprouting Ceanothus.American 146. Mac, M. J., P. A. Opler, C. E. Puckett Haecker, and P. D. Doran. 1998.
Midland Naturalist 93(2):504-507. Status and trends of the nation’s biological resources.Volume 1. U.S.
125. ———, and S. C. Keeley. 1988. Chaparral. Pages 165-208 in M. G. Department of the Interior, U.S. Geological Survey, Reston,VI.
Barbour and W. D. Billings, editors. North American terrestrial 147. Manning, M. E., and W. G. Padgett. 1992. Riparian community type
vegetation. Cambridge University Press, New York, NY. classification for the Humboldt and Toiyabe national forests, Nevada
126. Kiilsgaard, C. 1999. Oregon vegetation: mapping and classification of and eastern California. Unpublished Draft Report prepared for U.S.
landscape level cover types. Final Report. U.S. Geological Survey- Forest Service, Intermountain Region Ecology and Classification
Biological Resources Division: Gap Analysis Program. Moscow, ID Program, Ogden, UT.
127. ———, and C. Barrett. 1998. Oregon vegetation landscape-level 148. Marsh, F., R. Helliwell, and J. Rodgers. 1987. Plant association guide
cover types 127. Northwest Habitat Institute, Corvallis, OR. for the commercial forest of the Warm Springs Indian Reservation.
128. Kilgore, B. M. 1973. The ecological role of fire in Sierran conifer Confederated Tribes of the Warm Springs Indians, Warm Springs, OR.
forests—its application to National Park management. Quaternary 148a. Marzluff, J. M. 1997. Effects of urbanization and recreation on
Research 3:496-513. songbirds. Pages 89-102 in W. M. Block and D. M. Finch, editors.
129. King County Park, Planning and Resource Department. 1987. Songbird ecology in southwestern ponderosa pine forests: a literature
Wildlife habitat profile—King County Open Space Program, Seattle, review. U.S. Forest Service General Technical Report RM-292, Fort
WA. Collins, CO.
130. Knutson, K. L., and V. L. Naef. 1997. Priority habitat management 149. Marzluff, J. M., F. R. Gehlbach, and D. A. Manuwal. 1998. Urban
recommendations: riparian. Washington Department of Fish and environments: influences on avifauna and challenges for the avian
Wildlife, Olympia,WA. conservationist. Pages 283-299 in J. M. Marzluff and R. Sallabanks,
131. Kovalchik, B. L. 1987. Riparian zone associations—Deschutes, editors. Avian conservation, research, and management. Island Press,
Ochoco, Fremont, and Winema national forests. U.S. Forest Service R6 Washington D.C.
ECOL TP-279-87. 150. Mayer, K. E., and W. F. Laudenslayer, Jr., editors. 1988. A guide to
132. ———. 1993. Riparian plant associations of the National Forests of wildlife habitats of California. State of California, the Resources Agency,
eastern Washington. A partial draft version 1. U.S. Forest Service, Department of Fish and Game, Wildlife Management Division, CWHR
Colville National Forest. Program, Sacramento, CA.
133. Kozloff, E. N. 1973. Seashore life of Puget Sound, the Strait of 151. McBride, J. R., and C. Reid. 1988. Urban. Pages 142-144 in K. E. Mayer
Georgia, and the San Juan Archipelago. University of Washington Press, and W. F. Laudenslayer, Jr., editors. A guide to wildlife habitats of
Seattle,WA. California. California Department of Forestry and Fire Protection,
134. Krajina, V. J. 1965. Bioclimatic zones and classification of British Sacramento, CA.
Columbia. Pages 1-17 in V. J. Krajina, editor. Ecology of western North 152. McDonald, P. M., and J.C. Tappeiner, II. 1987. Silviculture, ecology, and
America. Volume 1. University of British Columbia, Vancouver, British management of tanoak in northern California. Pages 64-70 in T. R.
Columbia, Canada. Plumb and N. H. Pillsbury, technical coordinators. Proceedings of the
135. Kruckeberg, A. R. 1996. Gardening with native plants of the Pacific symposium on multiple-use management of California’s hardwood
Northwest: an illustrated guide. University of Washington Press, Seattle. resources; 12-14 November 1986; San Luis Obispo, California. U.S.
ISBN 0-295-97476-1. Forest Service General Technical Report PSW-100.
136. Kuchler, A.W. 1964. Manual to accompany the map: potential natural 153. McKenzie, D. F., and T. Z. Riley, editors. 1995. How much is enough? A
vegetation of the conterminous United States. Special Publication. 36, regional wildlife habitat needs assessment for the 1995 Farm Bill.
American Geographic Society, New York, NY. Wildlife Management Institute, Washington, D.C.
137. Kumler, M. L. 1969. Plant succession on the sand dunes of the 154. McNeil, R. C., and D. B. Zobel. 1980.Vegetation and fire history of a
Oregon coast. Ecology 50(4):695-704. ponderosa pine-white fir forest in Crater Lake National Park.
138. Kunze, L. M. 1994. Preliminary classification of native, low elevation, Northwest Science 54(1):30-46.
freshwater wetland vegetation in western Washington. Washington 155. Merriam, C. H. 1898. Life zones and crop zones of the United States.
Natural Heritage Program, Department of Natural Resources, Olympia, U.S. Department of Agriculture, Division of Biological Survey, Bulletin
Washington. 10.
139. Kuramoto, R. T., and L. C. Bliss. 1970. Ecology of subalpine meadows 156. Miller, T. B. 1976. Ecology of riparian communities dominated by
in the Olympic Mountains, Washington. Ecological Monograph white alder in western Idaho. M.S. Thesis. University of Idaho, Moscow.
40:317-347. 157. Minnich, R. A. 1983. Fire mosaics in southern California and north
140. Laacke, R .J., and J. N. Fiske. 1983. Red fir and white fir. Pages 41-43 in Baja California. Science 219:1287-1294.
R. M. Burns, compiler. Silvicultural systems for the major forest types of 158. Mitchell, R., and W. Moir. 1976.Vegetation of the Abbott Creek
the United States. U.S. Forest Service Agriculture Handbook No. 44. Research Natural Area, Oregon. Northwest Science 50:42-57.
Washington, D.C. 159. Morgan, P., S. C. Bunting, A. E. Black, T. Merrill, and S. Barrett. 1996.
141. Landry, M. R., and B. M. Hickey, editors. 1989. Coastal oceanography Fire regimes in the interior Columbia River Basin: past and present.
of Washington and Oregon. Elsevier Science Publishing Company, New Final Report RJVA-INT-94913. U.S. Forest Service, Intermountain
York, NY. Research Station, Intermountain Fire Sciences Lab, Missoula, MT.
142. Lang, F. A. 1961. A study of vegetation change on the gravelly prairies 160. Morrison, P., and F. J. Swanson. 1990. Fire history and pattern in a
of Pierce and Thurston counties, western Washington. M.S. Thesis. Cascade Range landscape. U.S. Forest Service General Technical Report
University of Washington, Seattle, WA. PNW-GTR-254.
 CHAPTER 2: HABITAT DESCRIPTIONS 113

161. Mueggler, W. F. 1988. Aspen community types of the Intermountain 184. Roberts, K., L. Bischoff, K. Brodersen, G. Green, D. Gritten, S.
Region. U.S. Forest Service, General Technical Report INT-250. Hamilton, J. Kierstead, M. Benham, E. Perkins, T. Pogson, S. Reed, and D.E.
Intermountain Research Station, Ogden, UT. Kerley. 1976. A preliminary ecology survey of the Alvord Basin, Oregon.
162. Naiman, R. J., H. Decamps, and M. Pollock. 1993. The role of riparian Unpublished, Final Technical Report, Eastern Oregon State College, La
corridors in maintaining regional biodiversity. Ecological Applications Grande. NSF Grant 76-08175.
3:209-212. 185. Rowntree, R. A. 1986. Ecology of the urban forest—introduction to
163. National Oceanic and Atmospheric Administration. 1993. Olympic part II. Urban Ecology 9(3/4):229-243.
Coast National Marine Sanctuary, Final Environmental Impact 185a. Rudnicky, J. L., and M. J. McDonnell. 1989. Forty-eight years of
Statement/Management Plan, November 1993 . NOAA, Sanctuaries and canopy change in a hardwood-hemlock forest in New York City.
Reservoirs Division, Washington D.C. Bulletin of the Torrey Botanical Club 116: 52-64.
164. National Research Council. 1989. Alternative agriculture. National 186. Ruth, R. H. 1974. Regeneration and growth of west-side mixed
Academy Press,Washington, D.C. conifers. In O. P. Camer, editor. Environmental effects of forest residues
165. Norton, H. H. 1979. The association between anthropogenic prairies in the Pacific Northwest: a state-of-knowledge compendium. U.S.
and important food plants in western Washington. Northwest Forest Service General Technical Report PNW-24.
Anthropological Research Notes 13:199-219. 187. Sampson, A. W., and B. S. Jespersen. 1963. California range brushlands
166. Noss, R. F., E. T. LaRoe, and J. M. Scott. 1995. Endangered ecosystems and browse plants. University of California, Division of Agricultural
of the United States: a preliminary assessment of loss and degradation. Sciences, California Agricultural Experiment Station, Extension Service,
U.S. National Biological Service, Biological Report 28. Berkeley, CA.
167. Nowak, D. J. 1994. Understanding of the structure of urban forests. 188. Sawyer, J. O., and T. Keeler-Wolf. 1995. A manual of California
Journal of Forestry October: 42-46. vegetation. Native Plant Society of California, Sacramento, CA.
168. Oliver, C. D. 1981. Forest development in North America following 189. Schoch, G. C., and M. N. Dethier. 1997. Analysis of shoreline
major disturbances. Forest Ecology and Management 3:153-168. classification and biophysical data for Carr Inlet. Washington State
169. Oregon Department of Forestry. 1994. Water protection rules: Department of Natural Resources. Olympia, WA.
purpose, goals, classification, and riparian management. OAR No.629- 190. Shipman, H. 1997. Shoreline armoring on Puget Sound. In T. Ransom,
635-200-Water classification. Oregon Department of Forestry, Salem, editor. Puget Sound Notes No. 40. Puget Sound Water Quality Action
OR. Team, Olympia,WA.
170. Oregon State University. 1971. Oceanography of the nearshore 191. Shreffler, D. K., R. M. Thom, and K. B. MacDonald. 1995. Shoreline
coastal waters of the Pacific Northwest relating to possible pollution. armoring effects on biological resources and coastal ecology in Puget
Volume 1. Corvallis, OR. Sound. In E. Robichaud, editor. Puget Sound Research 1995:
171. Parsons, D. J., and S. H. DeBenedetti. 1979. Impact of fire suppression Proceedings. Puget Sound Water Quality Action Team, Olympia, WA.
on a mixed-conifer forest. Forest Ecology and Management 2:21-33. 192. Simenstad, C. A. 1983. The ecology of estuarine channels of the
172. Pettinger, A. 1996. Native plants in the coastal garden: a guide for Pacific Northwest coast: a community profile. U.S. Fish and Wildlife
gardeners in British Columbia and the Pacific Northwest. Whitecap Services. FWS/OBS-83/05.
Books 1-55110-405-9.Vancouver, BC. 193. Spies, T. A., J. F. Franklin, and T. B. Thomas. 1988. Coarse woody debris
173. Phillips, R. C. 1984. The ecology of eelgrass meadows in the Pacific in Douglas-fir forests of western Oregon and Washington. Ecology
Northwest: a community profile. U.S. Fish and Wildlife Service, FWS/ 69:1689-1702.
OBS-84/24. 194. Strickland, R., and D. J. Chasan. 1989. Coastal Washington, a synthesis
174. Phinney, L. A., and P. Bucknell. 1975. A catalog of Washington streams of information. Washington State and Offshore Oil and Gas,
and salmon utilization. Washington Department of Fisheries.Volume 2: Washington Sea Grant, University of Washington, Seattle, WA.
coastal region. 195. Strickler, G. S. 1961.Vegetation and soil condition changes on a
175. Poulton, C. E. 1955. Ecology of the non-forested vegetation in subalpine grassland in eastern Oregon. U.S. Forest Service Research
Umatilla and Morrow counties, Oregon. Ph.D. Dissertation. State Paper PNW-40, Portland, OR.
College of Washington, Pullman, WA. 196. ———, and W. B. Hall. 1980. The Standley allotment: a history of
176. Proctor, C. M., J. C. Garcia, D.V. Galvin, G. B. Lewis, L. C. Loehr, and A. range recovery. U.S. Forest Service, Forest and Range Experiment
M. Massa. 1980. An ecological characterization of the Pacific Northwest Station Research Paper, PNW-278.
coastal region.Volume 2. U.S. Fish and Wildlife Service, Biological 197. Sullivan, K., T. E. Lidle, C. A. Dolloff, G. E. Grant, and L. M. Reid. 1987.
Services Program. FWS/OBS-79/14. Stream channels: the link between forest and fishes. Pages 39-97 in E. O.
177. ———, ———, ———, ———, ———, and ———. 1980. An Salo and T. W. Cundy, editors. Streamside management: forestry and
ecological characterization of the Pacific Northwest coastal region. fishery interactions. College of Forest Resources. Contribution No. 57.
Volume 3. U.S. Fish and Wildlife Service, Biological Services Program. University of Washington, Seattle,WA.
FWS/OBS-79/14. 198. Swanson, F. J., L. E. Benda, S. H. Duncan, G. E. Grant, W. F. Megaham, L.
178. ———, ———, ———, ———, ———, and ———. 1980. An M. Reid, and R. R. Zeimer. 1987. Mass failures and other processes of
ecological characterization of the Pacific Northwest coastal region. sediment production in Pacific Northwest forest landscapes. Pages 9-38
Volume 4. U.S. Fish and Wildlife Service, Biological Services Program. in E. O. Salo and T. W. Cundy, editors. Streamside management: forestry
FWS/OBS-79/14. and fisheries interactions. College of Forest Resources Contribution
179. Pruter, A. T., and D. L. Alverson, editors. 1972. The Columbia River No. 57, University of Washington, Seattle, WA.
estuary and adjacent waters: bioenvironmental studies. University of 199. ———, and C. T. Dyrness. 1975. Impact of clearcutting and road
Washington Press, Seattle. construction on soil erosion by landslides in the western Cascade
180. Puget Sound Water Quality Authority. 1997. 1997 Puget Sound Range, Oregon. Geology 3:393-396.
update. Seventh annual report of the Puget Sound Ambient Monitoring 200. ———, R. L. Fredriksen, and F. M. McCorison. 1982. Material transfer
Program. Puget Sound Water Quality Authority, Olympia, Washington. in a western Oregon forested watershed. Pages 223-266 in R. L.
181. Quigley, T. M., and S. J. Arbelbide, technical editors. 1997. An Edmonds, editor. Analysis of coniferous forest ecosystems in the
assessment of ecosystem components in the interior Columbia basin western United States. Hutchinson Ross, Stroudsburg, PA.
and portions of the Klamath and Great Basins.Volume 2. U.S. Forest 201. The University of Oregon’s Atlas of Oregon. 1976.
Service General Technical Report PNW-GTR-405. 202.Thilenius, J. F. 1968.The Quercus garryana forests of the Willamette
182. Quinn, T. 1997. Coyote (Canis latrans) food habits in three urban Valley, Oregon. Ecology 49:1124-1133.
habitat types of western Washington. Northwest Science 71(1):1-5. 203. Thomson, R. E. 1981. Oceanography of the British Columbia coast.
183. Ripley, J. D. 1983. Description of the plant communities and
succession of the Oregon coast grasslands. M.S. Thesis. Oregon State
University, Corvallis, OR.
 114 Wildlife–Habitat Relationships in Oregon and Washington

Canadian Special Publication, Fisheries and Aquatic Sciences 56:1-292. 223. Williams, R. W., R. M. Laramie, and J. J. Ames. 1975. A catalog of
204. Thompson, K., and D. Snow. 1974. Fish and Wildlife Resources: Washington streams and salmon utilization. Washington Department of
Oregon coastal zone. Oregon Coastal Conservation and Development Fisheries.Volume 1: Puget Sound Region.
Commission, Portland, OR. 224. Winward, A. H. 1970. Taxonomic and ecological relationships of the
205. Tiner, R. W. 1984. Wetlands of the United States: current status and big sagebrush complex in Idaho. Ph.D. Dissertation. University of Idaho,
recent trends. National Wetlands Inventory. U.S. Fish and Wildlife Moscow.
Service. 225. ———. 1980. Taxonomy and ecology of sagebrush in Oregon.
206. Tisdale, E. W. 1983. Grasslands of western North America: the Pacific Oregon State University Agricultural Experiment Station Bulletin
Northwest bunchgrass type. Pages 223-245 in A. C. Nicholson, A. 642:1-15.
McLean and T. E. Baker, editors. Grassland ecology and classification 226. Wolcott, E. E. 1973. Lakes of Washington. Water Supply. State of
symposium proceedings. British Columbia Ministry of Forests, Victoria, Washington, Department of Conservation, Bulletin No. 14.Volume 1:
BC. Western Washington. Olympia,WA.
207. ———. 1986. Canyon grasslands and associated shrublands of 227. ———. 1973. Lakes of Washington. Water Supply. State of
west-central Idaho and adjacent areas. Bulletin No. 40. Forest, Wildlife Washington, Department of Conservation, Bulletin No. 14.Volume 2:
and Range Experiment Station, University of Idaho, Moscow, ID. Eastern Washington. Olympia,WA.
208. Topik, C. 1989. Plant association and management guide for the 228. Zack, A. C., and P. Morgan. 1994. Early succession on hemlock habitat
Grand Fir Zone, Gifford Pinchot National Forest. U.S. Forest Service, types in northern Idaho. Pages 71-84 in D. M. Baumgartner, J. E. Lotan,
R6-ECOL-006-88. and J. R. Tonn, editors. Interior cedar-hemlock-white pine forests:
209. ———, N. M. Halverson, and T. High. 1988. Plant association and ecology and management. Cooperative Extension Program, Washington
management guide for the Ponderosa Pine, Douglas-fir, and Grand Fir State University, Seattle, WA.
Zones, Mount Hood National Forest. U.S. Forest Service, R6-ECOL- 229. Ziemer, R. R. 1981. Roots and the stability of forested slopes. Pages
TP-004-88. 343-361 in Proceedings of a symposium on erosion and sediment
210. ———, ———, and D. G. Brockway. 1986. Plant association and transport in Pacific Rim steeplands. Publication 132. International
management guide for the Western Hemlock Zone, Gifford Pinchot Association of Hydrological Scientists. Washington, D.C.
National Forest. U.S. Forest Service, R6-ECOL-230A-1986. 230. Zobel, D. B., L. F. Roth, and G. L. Hawk. 1985. Ecology, pathology and
211. Turner, R. B. 1969.Vegetation changes of communities containing management of Port-Orford cedar (Chamaecyparis lawsoniana). U.S.
medusahead (Taeniatherum asperum [Sim.] Nevski) following herbicide, Forest Service General Technical Report PNW-184.
grazing and mowing treatments. Ph.D. Dissertation. Oregon State
University, Corvallis, OR.
212.Volland, L. A. 1976. Plant communities of the central Oregon pumice
zone. U.S. Forest Service R-6 Area Guide 4-2. Pacific Northwest
Region, Portland, OR.
212a. Walcott, C. F. 1974. Changes in bird life in Cambridge,
Massachusetts from 1960 to 1964. The Auk 91: 151-160.
213. Ware, D. M., and G. A. McFarlane. 1989. Fisheries production domains
in the Northeast Pacific Ocean. Pages 359-379 in R. J. Beamish and G. A.
McFarlane, editors. Effects of ocean variability on recruitment and
evaluation of parameters used in stock assessment models. Canadian
Special Publication, Fisheries and Aquatic Sciences 108.
214. Washington Department of Ecology. 1994. Inventory of dams.
Washington Department of Ecology, Water Resources Program, Dam
Safety Section. Publication No.9
215. Washington Department of Natural Resources. 1998. Our changing
nature—natural resource trends in Washington State. Washington
Department of Natural Resources, Olympia, WA.
216. West, J. E. 1997. Protection and restoration of marine life in the
inland waters of Washington State. Puget Sound/Georgia Basin
Environmental Report Series: No. 6. Puget Sound Water Quality Action
Team, Olympia,WA.
217. Wetzel, R. G. 1983. Limnology. Saunders College Publishing. New
York, NY.
218. Whittier, T. R., R. M. Hughes, and D. P. Larsen. 1988. Correspondence
between ecoregions and spatial patterns in stream ecosystems in
Oregon. Canadian Journal of Fisheries and Aquatic Sciences 45:1264-
1278.
219. Wiedemann, A. M. 1966. Contributions to the plant ecology of the
Oregon Coastal Sand Dunes. Ph.D. Dissertation. Oregon State
University, Corvallis, OR.
220. ———. 1984. The ecology of Pacific Northwest coastal sand dunes:
a community profile. U.S. Fish and Wildlife Service, FWS/OBS-84/04.
221. Williams, C. K., B. F. Kelley, B. G. Smith, and T. R. Lillybridge. 1995.
Forested plant associations of the Colville National Forest. U.S. Forest
Service General Technical Report PNW-GTR-360. Portland, OR.
222. ———, and T.R. Lillybridge. 1983. Forested plant associations of the
Okanogan National Forest. U.S. Forest Service, R6-Ecol-132b. Portland,
OR.
 3
Structural Conditions and Habitat
Elements of Oregon and Washington
Thomas A. O’Neil, Kelly A. Bettinger, MadeleineVander
Heyden, Bruce G. Marcot, Charley Barrett,T. Kim Mellen,
W. MatthewVanderhaegen, David H. Johnson,
Patrick J. Doran, Laurie Wunder, & Kathryn M. Boula

Introduction
Wildlife habitat is a concept that describes attributes in or site. We achieve this knowledge by interpolating
the environment that serve as life requisites for wildlife ecological condition from assessing wildlife habitats
allowing them to survive and reproduce. An overt hierarchically, that is, from knowing something about the
assumption is that wildlife use habitats that are arranged wildlife habitats, structural conditions, and habitat
or comprised of vital components; these arrangements elements from area or site. Thus, current and desired
result in healthy and viable wildlife populations. However, ecological conditions can be assessed for wildlife and
the wildlife habitat concept does not address demographic written:
(survival and reproduction of individuals), environmental
(food supply, predators, weather), natural catastrophe Wildlife Habitats = wildlife cover type(s) + structural
(flood, drought, fire), or genetic (genetic drift or condition(s) + habitat element(s)
inbreeding) uncertainties. These uncertainties, though
important, are difficult to determine, predict, and manage. Structural Conditions
Nonetheless, we have studied and assessed wildlife life Structure is what a natural resource manager can
history and how species use their habitat(s) for quite some manipulate to achieve various objectives. After all,
time. Thus, the wildlife habitat concept does address manipulating the structural features of a forest stand is
features in the environment that we can manipulate to what silviculture is all about.10 Structural conditions are
enhance or discourage wildlife use of an area. often thought to follow the plant succession series. That
The principal purpose of this chapter is to report what is, if a stand, say in a forest, is left alone and given enough
we know or surmise about wildlife and its use of the time it will achieve a climax or old-growth state. So the
landscape into features that land managers can continuum starts with the earliest stages, and given a forest
understand, recognize, and strive to achieve. Describing example, this would be at grass/forb, and then
and defining these features allows us to begin to establish progressing through shrub/seedling; sapling/pole; small,
a common understanding for management so that we can medium, and large trees; and eventually achieving giant
affect and influence the landscape to meet wildlife needs. trees. Catastrophic events along with some management
What follows is an expansion of our wildlife habitats; two prescriptions can reset the succession stages (fire burning
major features that also service wildlife species needs are a tract of land mimics in many ways the effect of a clearcut).
structural conditions and specific habitat elements of a So several key structural elements for forest and
site. These two features should be viewed in a hierarchical shrubland/grasslands habitats are a wide range in tree
manner, with structural conditions occurring on tracts of and shrub sizes, a wide range in tree diameters and tree
land such as at a forest stand or watershed level, whereby and shrub heights, and varying amounts of tree and shrub
habitat elements are described at a site-specific or local canopies. Multiple canopy layers are also considered to
level, like within a forest stand. Knowing something about be significant and this includes the continuous distribution
structural conditions of an area along with the habitat of foliar surface from the top of the crown to the ground.
elements occurring at a site will allow us to better predict Such canopy distributions are thought to create greater
what kinds of wildlife may use the site, as well as predict, quantities and greater diversity of animal habitats.10, 13 That
if enhancements or modifications were made what wildlife is, canopies are important not only for their physical
might continue to use the site or area. characteristics, such as thermal qualities that influence
The collective set of environmental conditions provided stream temperatures and ground conditions (i.e. shade,
by wildlife habitats, structural conditions, and habitat shelter from precipitation) but also for their abiotic abilities
elements constitutes a species’ overall habitat. Hence, from such as fixating CO2 or nitrogen from the atmosphere.2, 11,
17
an ecological perspective, we are striving to determine the Finally, plant and tree understories may or may not be
current and potential ecological conditions of a landscape present depending on the structural condition of the stand;
115
 116 Wildlife–Habitat Relationships in Oregon and Washington

however, their presence can influence the kind and 1. Grass/Forb—Open

number of species that use the stand. Grass/Forb dominated with <70% coverage by grasses
and forbs. Shrubs and small seedlings may be present,
Defining Structural Conditions but do not dominate stand, (seedlings <10% canopy cover),
Knowing the wildlife-habitat type(s) of an area will allow and there may be remnant trees (trees remaining from the
a person to predict a list of species that may be found at previous stand) that provide <10% canopy cover (Figure
that site. To refine the prediction, however, requires further 1).
information, like knowing a species range extent and
condition of the habitat in the area of concern. The term 2. Grass/Forb—Closed
condition implies, knowing the types and amounts of Grass/forb dominated with >70% coverage by grasses and
different structural stages and habitat elements. The forbs. Shrubs and small seedlings may be present, but do
condition of the habitat can predispose a species to use an not dominate stand, (seedlings <10% canopy cover), and
area, and thus can serve as a driver for its occurrence. there may be remnant trees (trees remaining from the
Definitions of structural conditions were determined previous stand) that provide <10% canopy cover (Figure
by the Species Habitat Team that was formed from 2).
multiple organizations to help clarify terms, represent field
3. Shrub/Seedling—Open
ecologist and biologist needs, and to help ensure a usable
Seedlings are large enough to add structure to the stand
end-product. The definitions are divided into four major
but are small enough that the structure is similar to shrubs
categories: Forest, Shrubland/Grassland, Agriculture, and
and may have remnant trees (trees remaining from the
Urban. The definitions for the first two categories are based
previous stand) that provide <10% canopy cover. There is
on describing the characteristics of trees, shrubs, and
<70% cover of shrubs or seedlings. Tree size is <1 inch
grasses. The last two categories are based on describing
dbh, and there is 1 canopy stratum (Figure 3).
land use/land cover types. This was done because of the
difficulty in describing these categories from a plant 4. Shrub/Seedling—Closed
community perspective and because there is very little Seedlings are large enough to add structure to the stand
literature available that would allow a more detailed but are small enough that the structure is similar to shrubs.
breakdown. Table 1 was created to depict those wildlife Remnant trees (trees remaining from the previous stand)
habitats that are associated with each structural condition may provide <10% canopy cover. There is >70% cover of
or land use/land cover type. A computer simulation that shrubs or seedlings. Tree size is <1 inch dbh, and there is
represents each Forest and Shrubland/Grassland 1 canopy stratum (Figure 4).
structural condition can be found by reviewing the color
photographs that accompany this chapter. The following 5. Sapling/Pole—Open
are the definitions for structural conditions: The canopy is open enough that understory vegetation
may be abundant. Remnant trees (trees remaining from
Forest Structural Conditions the previous stand) may provide <10% canopy cover.
The forest structural conditions are based on the following There is 10-39% cover of sapling and pole-sized trees. Tree
attributes: (1) tree size diameter at breast height (dbh); (2) size is 1-9 inches dbh, and there is 1 canopy stratum (Figure
percent canopy cover (or percent grass/forb cover); and, 5).
(3) number of canopy layers. These attributes have the
following values: 6. Sapling/Pole—Moderate
Understory development is hampered by available light
Tree size (dbh) and moisture. Remnant trees (trees remaining from the
shrub/seedling <1 inch <2.5 cm previous stand) may provide <10% canopy cover. There
sapling/pole 1-9 inches 2.5-24 cm is 40-69% cover of sapling and pole-sized trees. Tree size
small tree 10-14 inches 25-37 cm is 1-9 inches dbh, and there is 1 canopy stratum (Figure
medium tree 15-19 inches 38-49 cm 6).
large tree 20-29 inches 50-75 cm
giant tree >30 inches >76 cm 7. Sapling/Pole—Closed
The understory is depauperate or absent. Remnant trees
Percent canopy cover (trees remaining from the previous stand) may provide
open 10-39% <10% canopy cover. There is >70% cover of sapling and
moderate 40-69%
pole-sized trees. Tree size is 1- 9 inches dbh and there is
closed 70-100%
1canopy stratum (Figure 7).
Number of canopy layers
8. Small Tree—Single Story—Open
single story 1 stratum
A grass/forb or shrub understory may be present.
multi-story >2 strata
Remnant trees (trees remaining from the previous stand)
The above attributes have been combined into the may provide <10% canopy cover. There is 10-39% cover
following structural conditions: of small trees, with <10% cover of other tree sizes. Tree
size is 10-14 inches dbh, and there is 1 canopy stratum
(Figure 8).
 CHAPTER 3: STRUCTURES AND ELEMENTS 117

Table 1. Matrix for wildlife habitats that are associated with structural conditions
or land use/land cover types.

Wildlife habitats Forest Grassland/Shrubland Structural Conditions Land Use/Land Cover

Structural Conditions
Conditions1
Grass/ Low Medium Tall Urban2 Agriculture3
Forb shrub shrub shrub

Westside lowlands conifer-hardwood forest X — — — — — —

Westside oak & Douglas-fir forest X — — — — — —
& woodlands
Southwest Oregon mixed conifer- X — — — — — —
hardwood forest
Montane mixed conifer forest X — — — — — —
Eastside (Interior) mixed conifer forest X — — — — — —
Lodgepole pine forest & woodlands X — — — — — —
Ponderosa pine forests & woodlands X — — — — — —
Upland aspen forest X — — — — — —
Subalpine parklands X — — — — — —
Alpine grasslands & shrublands — X X — — — —
Westside grasslands — X — — — — —
Ceanothus/manzanita shrublands — — — — X — —
Western juniper & mountain X — — — — — —
mahogany woodlands
Eastside (Interior) canyon shrublands — — — X X — —
Eastside (Interior) grasslands — X — — — — —
Shrub-steppe — — — X — — —
Dwarf shrub-steppe — — X — — — —
Desert playa & salt scrub — — — X — — —
Agriculture, pasture, and mixed environs — — — — — — X
Urban and mixed environs — — — — — X —
Open water—lakes, rivers, streams — — — — — — —
Herbaceous wetlands — — — — — — —
Westside riparian/wetlands X — — — X — —
Montane coniferous wetlands X — — — — — —
Eastside (Interior) riparian/wetlands X — — — X — —
Coastal dunes & beaches — X — X X — —
Coastal headlands & islets — X — X X — —
Bays & estuaries — — — — — — —
Inland marine deeper waters — — — — — — —
Marine nearshore — — — — — — —
Marine shelf — — — — — — —
Oceanic — — — — — — —
1
Includes a variety of tree sizes and canopies
2
Includes low, medium, and high densities of impervious areas.
3
Includes areas like croplands, orchards, and pastures
 118 Wildlife–Habitat Relationships in Oregon and Washington

9. Small Tree—Single Story—Moderate 17. Small Tree—Multi-story—Open

Some grass/forb or shrub understory may be present. These stands have an overstory of small trees with a
Remnant trees (green trees remaining from the previous distinct subcanopy of saplings and/or poles. Scattered
stand) may provide <10% canopy cover. There is 40-69% larger trees may be present but make up <10% canopy
cover of small trees with <10% cover of other sized trees. cover. Grass/forb or shrub understory may be present.
Tree size is 10-14 inches dbh, and there is 1 canopy stratum There is 10-39% total canopy cover dominated by small
(Figure 9). trees, >10% canopy cover of >1 other smaller tree sizes.
Tree size is 10-14 inches dbh, and there are >2 canopy strata
10. Small Tree—Single Story—Closed (Figure 17).
Grass/Forb or shrub understory minor or absent.
Remnant trees (trees remaining from the previous stand) 18. Small Tree—Multi-story—Moderate
may provide <10% canopy cover. There is >70% cover of These stands have an overstory of small trees with a
small trees, with <10% cover of other-sized trees. Tree size distinct subcanopy of saplings and/or poles. Scattered
is 10-14 inches dbh, and there is 1 canopy stratum (Figure larger trees may be present but make up <10% canopy
10). cover. Grass/forb or shrub understory may be present,
but is probably limited. There is 40-69% total canopy cover
11. Medium Tree—Single Story—Open dominated by small trees, >10% canopy cover of >1 or
A grass/forb or shrub understory may be present. more other smaller tree sizes. Tree size is 10-14 inches dbh,
Remnant trees (trees remaining from the previous stand) and there are >2 canopy strata (Figure 18).
may provide <10% canopy cover. There is 10-39% cover
of medium trees, with <10% cover of other-sized trees. 19. Small Tree—Multi-story—Closed
Tree size is 15-19 inches dbh, and there is 1 canopy stratum These stands have an overstory of small trees with a
(Figure 11). distinct subcanopy of saplings and/or poles. Scattered
larger trees may be present but make up <10% canopy
12. Medium Tree—Single Story—Moderate cover. Grass/forb or shrub understory extremely limited
Grass/Forb or shrub understory may be present. Remnant or absent. There is >70% total canopy cover dominated
trees (trees remaining from the previous stand) may by small trees, >10% canopy cover of >1 other smaller tree
provide <10% canopy cover. There is 40-69% cover of sizes. Tree size is 10-14 inches dbh, and there are >2 canopy
medium trees with <10% cover of other-sized trees. Tree strata (Figure 19).
size is 15-19 inches dbh, and there is 1 canopy stratum
(Figure 12). 20. Medium Tree—Multi-story—Open
These stands have an overstory of medium trees with a
13. Medium Tree—Single Story—Closed distinct subcanopy of smaller trees. Scattered larger trees
A grass/forb or shrub understory may be present. may be present but make up <10% canopy cover. Grass/
Remnant trees (trees remaining from the previous stand) forb or shrub understory may be present, but is probably
may provide <10% canopy cover. There is >70% cover of limited. There is 10-39% total canopy cover dominated by
medium trees with <10% cover of other-sized trees. Tree medium trees, >10% or more canopy cover of >1 other
size is 15-19 inches dbh, and there is 1 canopy stratum smaller tree sizes. Tree size is 15-19 inches dbh, and there
(Figure 13). are >2 canopy strata (Figure 20).
14. Large Tree—Single Story—Open 21. Medium Tree—Multi-story—Moderate
Grasses, shrubs, and/or seedlings may occur in the These stands have an overstory of medium trees with a
understory. There is 10-39% cover of large and/or giant distinct subcanopy of smaller trees. Scattered larger trees
size trees with <10% cover of other-sized trees. Tree size may be present but make up <10% canopy cover. Grass/
is 20-29 inches dbh, and there is 1 canopy stratum (Figure forb or shrub understory may be present, but is probably
14). limited. There is 40-69% total canopy cover dominated by
medium trees, >10% or more canopy cover of 1 or more
15. Large Tree—Single Story—Moderate
smaller tree sizes. Tree size is 15-19 inches dbh, and there
Some grass/forb or shrub understory may be present.
are >2 canopy strata (Figure 21).
There is 40-69% cover of large and/or giant trees with
<10% cover of other-sized trees. Tree size is 20-29 inches 22. Medium Tree—Multi-story—Closed
dbh, and there is 1 canopy stratum (Figure 15). These stands have an overstory of medium trees with a
distinct subcanopy of smaller trees. Scattered larger trees
16. Large Tree—Single Story—Closed
may be present but make up <10% canopy cover. Grass/
Grasses, shrubs, and/or seedlings may occur in the
forb understory may be present, but is probably limited.
understory. There is >70% cover of large and/or giant trees
There is >70% total canopy cover dominated by medium
with <10% cover of other-sized trees. Tree size is 20-29
trees, >10% or more canopy cover of >1 smaller tree sizes.
inches dbh, and there is 1 canopy stratum (Figure 16).
Tree size is 15-19 inches dbh, and there are >2 canopy strata
(Figure 22).
 CHAPTER 3: STRUCTURES AND ELEMENTS 119

23. Large Tree—Multi-story—Open Percent shrub cover

These stands have an overstory of large or giant-sized trees open 10-69% shrub cover
with one or more distinct canopy layers of smaller trees. closed 70-100% shrub cover
Stands with >40% cover of giant trees are classified in the
Giant, multi-storied stage. In westside forests, stands Shrub age class
dominated by large trees usually have giant trees scattered seedling/young negligible crown decadence
mature < 25% crown decadence
in the stand, with fewer in eastside forests. A grass/forb
old 26-100% crown decadence
or shrub understory is often present, especially in canopy
gaps. There is 10-39 % total canopy cover, with >10% or The above attributes have been combined into the
more canopy cover from large and/or giant trees and following structural conditions:
another >10% canopy cover from >1 or more smaller tree
size classes. Tree size is 20-29 inches dbh, and there are >2 1. Grass/Forb—Open
canopy strata (Figure 23) Grasslands that have <10% shrub cover and <10% tree
canopy cover. Grasses and forbs cover are <70% of the
24. Large Tree—Multi-story—Moderate ground, and bare ground is evident (Figure 27).
These stands have an overstory of large or giant-sized trees
with >1 distinct canopy layers of smaller trees. Stands with 2. Grass/Forb—Closed
>40% cover of giant trees are classified in the giant, Grasslands that have <10% shrub cover and <10% tree
multi-storied stage. In westside forests, stands dominated canopy cover. Grasses and forbs cover >70% of the ground
by large trees usually have giant trees scattered in the (Figure 28).
stand, with fewer in eastside forests. Grass/Forb or shrub
3. Low Shrub—Open Shrub Overstory—Seedling/
understory is often present, especially in canopy gaps.
Young
There is 40-69% total canopy cover, >10% canopy cover
Shrublands with shrubs <1.6 ft tall, shrub canopy cover
from large trees with another >10% canopy cover from >1
10-69% and may have <10% tree canopy cover. Areas with
smaller tree size classes. Tree size is 20-29 inches dbh, and
<10% shrub cover are categorized as Grass/Forb. These
there are >2 canopy strata (Figure 24).
are post-disturbance regenerating shrublands dominated
25. Large Tree—Multi-story—Closed by seedlings or young shrubs. Mature, legacy shrubs may
These stands have an overstory of large or giant-sized trees persist from before the disturbance, but occur as scattered
with >1 distinct canopy layers of smaller trees. Stands with singles or widely scattered clumps. Crown decadence is
> 40% cover of giant trees are classified in the giant, negligible (Figure 29).
multi-storied stage. In westside forests, stands dominated
4. Low Shrub—Open Shrub Overstory—Mature
by large trees usually have giant trees scattered in the
Shrublands with shrubs <1.6 ft tall, shrub canopy cover
stand, with fewer in eastside forests. A grass/forb or shrub
10-69% and may have <10% tree canopy cover. Areas with
understory is often present, especially in canopy gaps.
<10% shrub cover are categorized as Grass/Forb. Crown
There is >70% total canopy cover, >10% or more canopy
decadence is <25% (Figure 30).
cover from large trees with another >10% canopy cover
from >1 smaller tree size classes. Tree size is 20- 29 inches 5. Low Shrub—Open Shrub Overstory—Old
dbh, and there are >2 canopy strata (Figure 25). Shrublands with shrubs <1.6 ft tall, shrub canopy cover
10-69% and may have <10% tree canopy cover. Areas with
26. Giant Tree—Multi-story
<10% shrub cover are categorized as Grass/Forb. Crown
These stands have an overstory of giant-sized trees with
decadence is >25% (Figure 31).
>1 distinct canopy layers of smaller trees. Stands with
<40% canopy cover are classified in the large tree 6. Low Shrub—Closed Shrub Overstory—Seedling/
multi-story open stage. There is >40% canopy cover. Tree Young
size is >30 inches dbh, and there are >2 canopy strata Shrublands with shrubs <1.6 ft tall, shrub canopy cover
(Figure 26a and b). >70%, and may have <10% tree canopy cover. These are
post-disturbance regenerating shrublands dominated by
Shrubland and Grassland Structural seedlings or young shrubs. Mature, legacy shrubs may
Conditions persist from before the disturbance, but occur as scattered
The shrubland and grassland structural conditions are singles or widely scattered clumps. Crown decadence is
based on the following attributes: (1) shrub height, (2) negligible (Figure 32).
percent shrub cover (or percent grass/forb cover), and (3)
shrub age class. These attributes have the following values: 7. Low Shrub—Closed Shrub Overstory—Mature
Shrublands with shrubs <1.6 ft tall and shrub canopy cover
Shrub height >70%, and may have <10% tree canopy cover. Crown
low <1.6 ft <0.5 m decadence is <25% (Figure 33).
medium 1.6-6.5 ft 0.5-2.0 m
tall >6.5 ft-16.5 ft >2.0 m-5.0 m
 120 Wildlife–Habitat Relationships in Oregon and Washington

8. Low Shrub—Closed Shrub Overstory—Old 16. Tall Shrub—Open Shrub Overstory—Mature

Shrublands with shrubs <1.6 ft tall, shrub canopy cover Shrublands with shrubs >6.5 ft tall, shrub canopy cover
>70%, and may have <10% tree canopy cover. Crown >10% and <70% and may have <10% tree canopy cover.
decadence is >25% (Figure 34). Areas with <10% shrub cover are categorized as Grass/
Forb. Crown decadence is <25% (Figure 42).
9. Medium Shrub—Open Shrub Overstory—
Seedling/Young 17. Tall Shrub—Open Shrub Overstory—Old
Shrublands with shrubs 1.6 - 6.5 ft tall, shrub canopy cover Shrublands with shrubs >6.5 ft tall, shrub canopy cover
>10% and <70%, and may have <10% tree canopy cover. >10% and <70%, and may have tree canopy cover <10%.
Areas with <10% shrub cover are categorized as Grass/ Areas with <10% shrub cover are categorized as Grass/
Forb. These are post- disturbance regenerating shrublands Forb. Crown decadence is >25% (Figure 43).
dominated by seedlings or young shrubs. Mature, legacy
shrubs may persist from before the disturbance, but occur 18. Tall Shrub—Closed Shrub Overstory—Seedling/
as scattered singles or widely scattered clumps. Crown Young
decadence is negligible (Figure 35). Shrublands with shrubs >6.5 ft tall, shrub canopy cover
>70%, and may have tree canopy cover <10%. These are
10. Medium Shrub—Open Shrub Overstory— post-disturbance regenerating shrublands dominated by
Mature seedlings or young shrubs. Mature, legacy shrubs may
Shrublands with shrubs 1.6 - 6.5 ft tall, shrub canopy cover persist from before the disturbance, but occur as scattered
>10% and <70%, and may have <10% tree canopy cover. singles or widely scattered clumps. Crown decadence is
Areas with <10% shrub cover are categorized as Grass/ negligible (Figure 44).
Forb. Crown decadence is <25% (Figure 36).
19. Tall Shrub—Closed Shrub Overstory—Mature
11. Medium Shrub—Open Shrub Overstory—Old Shrublands with shrubs >6.5 ft tall, shrub canopy cover
Shrublands with shrubs 1.6 - 6.5 ft tall, shrub canopy cover >70%, and may have tree canopy cover <10%. Crown
>10% and <70%, and may have <10% tree canopy cover. decadence is <25% (Figure 45).
Areas with <10% shrub cover are categorized as Grass/
Forb. Crown decadence is >25% (Figure 37). 20. Tall Shrub—Closed Shrub Overstory—Old
Shrublands with shrubs >6.5 ft tall, shrub canopy cover
12. Medium Shrub—Closed Shrub Overstory— >70%, and may have <10% tree canopy cover. Crown
Seedling/Young decadence is >25% (Figure 46).
Shrublands with shrubs 1.6 - 6.5 ft tall, shrub canopy cover
>70%, and may have <10% tree canopy cover. These are Urban Land Use/Land Cover Conditions
post-disturbance regenerating shrublands dominated by The Urban land use/land cover conditions are based on
seedlings or young shrubs. Mature, legacy shrubs may the level of urban development as determined by the
persist from before the disturbance, but occur as scattered percent of land surface covered by impervious materials.
singles or widely scattered clumps. Crown decadence is The Urban land use/land cover classification consists
negligible (Figure 38). of the following conditions:

13. Medium Shrub—Closed Shrub Overstory— 1. Low Density

Mature Surfaces that are covered with 10-29% of impervious
Shrublands with shrubs 1.6 - 6.5 ft tall, shrub canopy cover material. Examples would include rural residential areas,
>70%, and may have <10% tree canopy cover. Crown suburban housing with large lots (>1 acre [2.4 ha]).
decadence is <25% (Figure 39).
2. Medium Density
14. Medium Shrub—Closed Shrub Overstory—Old Surfaces that are covered by 30-59% of impervious
Shrublands with shrubs 1.6 - 6.5 ft tall, shrub canopy cover material. Examples would include single family housing
>70%, and may have <10% tree canopy cover. Crown areas (lot size >1 acre [2.4 ha]), suburban development.
decadence is >25% (Figure 40).
3. High Density
15. Tall Shrub—Open Shrub Overstory—Seedling/ Surfaces that are covered by 60-100% of impervious
Young material. Examples would include core downtown areas
Shrublands with shrubs >6.5ft tall, shrub canopy cover within cities (e.g., Seattle, Portland), commercial areas (e.g.,
>10% and <70%, and may have <10% tree canopy cover. shopping malls), industrial areas, high density housing
Areas with <10% shrub cover are categorized as Grass/ (e.g., apartment buildings), and transportation corridors
Forb. These are post- disturbance regenerating shrublands (e.g., highways).
dominated by seedlings or young shrubs. Mature, legacy
shrubs may persist from before the disturbance, but occur
as scattered singles or widely scattered clumps. Crown
decadence is negligible (Figure 41).
 CHAPTER 3: STRUCTURES AND ELEMENTS 121

Agricultural Land Use/Land Cover Conditions land might be grazed by livestock, but shows no evidence
of irrigation. It can also be characterized by uncut hay,
1. Cultivated Cropland organic debris from the previous season, uncut standing
Farmland used for production of annual crops such as dead grass, exotic plants like tansy ragwort (Senecio
vegetables and herbs is characterized by bare soil, and jacobea), thistle (Cirsium spp.), Himalaya blackberry (Rubus
plant debris either in the field or along the periphery. The discolor) and their debris, patches of shrubs such as
location tends to be along bottomland areas of streams hawthorn (Crataegus spp.), snowberry (Symphoricarpos
and rivers and areas with a sufficient source of irrigation. spp.), spirea (Spirea spp.), poison oak (Rhus diversiloba), and
Farmland used for production of annual grasses such as encroachment by various tree species. This land has
wheat, oats, barley, and rye is characterized by upland usually been cleared and farmed intensively in the past.
and rolling hill terrain, generally without irrigation. This This category also includes lands that are designated
agricultural division has similar pesticide use and/or within the Conservation Reserve Program (CRP) and areas
irrigation requirements. That is, row crops are treated the planted with crested wheatgrass (Agropyron cristatum).
same way in regard to the general application of pesticides Land owners use unimproved pasture for grazing
and cultural techniques in land preparation and harvest. livestock, otherwise it lies dormant. Thus, those lands that
There is a wide range of soil conservation practices in this are not grazed either revert to brushy field or volunteer
category. forest.
2. Improved Pasture
Farmland used for the production of perennial grass such Structural Conditions Data Matrix
as grass seed and hay. Perennial grass is generally grown To maximize the utility of wildlife-habitat relationship
without irrigation. Perennial crops are treated the same information, a digital database that links wildlife with its
way in regard to the general application of pesticides and structural conditions can be found on the CD-ROM
cultural techniques. included with this book. Wildlife occurrence within a
particular structural condition type was determined
3. Orchards/Vineyards/Nursery through an expert panel process held during the fall of
Farmland used for production of tree fruits (apples, 1998. Table 1 was created to assist the expert panel in
peaches, pears, hazelnuts), vineyards (grapes), berries identifying what wildlife habitats are associated with what
(strawberries, raspberries, blueberries, blackberries), structural conditions. The categories that depict a wildlife
Christmas trees, and nursery stock (ornamental container species occurrence with a particular structural condition
and greenhouse operations). This cover type is generally are Y—Yes the species occurs, H—Historically occurred, and
located in upland areas with access to a high volume of U—Unsure. Alongside the occurrence category, we
irrigation. Christmas trees are characterized by upland identify the types of activity that the species does while
areas, poorer soils and no irrigation. The use of chemicals using the structural condition. The activity codes for the
in non-food crops, such as Christmas trees and nursery wildlife species within a particular condition are: B—Both
stock, is considerably different both in materials and time feeds and breeds, F/R-HE—Feeds and Reproduces when a
of applications. specific habitat element is present, F—Feeds only, R—
Reproduces only, and O—Other. The Other category reflects
4. Modified Grasslands activities such as roosting/resting, hibernating, or using
Annual or introduced perennial grasslands, including the habitat for cover (thermal and hiding) purposes.
cheatgrass (Bromus tectorum), medusahead (Taeniatherum
caput-medusae), and other annual bromes; moist Defining the Level of Associations Between
environments, including riparian bottomlands, are often Wildlife and Structural Conditions
dominated by Kentucky bluegrass. Annual grasslands As mentioned in Chapter 1, we continue to embrace the
(and areas of introduced forbs) are usually dominated by new concept of degree of association between wildlife species
one or two introduced annuals which comprise most of and their habitats.19 For the purposes of this project, we
the vegetation cover. Perennial grasslands are usually used the following categories for characterizing the degree
dominated by a single planted bunchgrass with of association.
introduced annuals and weedy forbs between the bunches.
Some environments support rhizomatous perennial Closely Associated. A species is widely known to depend
grasses. These areas occur mostly on uplands but also on a habitat or structural condition for part or all of its life
includes riparian bottomlands that are dominated by history requirements. Identifying this association implies
non-native grasses. Modified grasslands can be found that the species has an essential need for this habitat or
throughout the steppe and grasslands areas of eastern structural condition for its maintenance and viability.
Oregon and Washington and at low elevation sites in Some species may be closely associated with more than
southwestern Oregon. one habitat or structural condition, others may be closely
associated with only one habitat or structural condition.
5. Unimproved Pasture Examples of species exhibiting close associations are red-
Farmland that seems to have no active management such winged blackbirds to wetland habitats, and spotted owls
as fertilizer application, irrigation or weed control. This to mature and giant tree structural conditions.
 122 Wildlife–Habitat Relationships in Oregon and Washington

Generally Associated. A species exhibits a high degree of expanding the definition of habitat to encompass all
adaptability and may be supported by a number of environmental correlates, naming the entirety of these
habitats or structural conditions. In other words, the attributes Key Ecological Correlates or KECs. All
habitats or structural conditions play a supportive role environmental scales, from broad floristic communities
for its maintenance and viability. Examples include the to fine-scale within-stand features, were included in their
black bear’s association with a variety of forested habitats definition of a KEC. The word Key in Key Environmental
or the black-tailed deer associated with a number of Correlate refers to the high degree of influence (either
structural conditions. positive or negative) the environmental correlates exert
on the realized fitness of a given species. Nonetheless,
Present. A species demonstrates occasional use of a habitat
when this information was determined, only direct
or structural condition. The habitat or structural condition
relationships between an HE and a species were identified.
provides marginal support to the species for its
Most of the HE-species associations refer to on the CD-
maintenance and viability. Examples are the rough-legged
ROM are mostly a positive influence between the HE and
hawk in desert playa and salt scrub shrubland or the mink
the species. Negative influence between HE and the
in a montane mixed conifer forest.
species may be viewed as environmental stressors;
Finally, the expert panelists assigned an overall
however, a comprehensive list of negative influences is
confidence rating to the occurrence and activity headings
not presented here. The following are HE definitions.
for each species within each structural condition. The
confidence ratings were simply high (e.g., many peer or 1. Forest, Shrubland and Grassland Habitat
published accounts), moderate, and low (e.g., few or no Elements
published accounts). By assigning a confidence rating, our Biotic, naturally occurring attributes of forest and
objective was to offer users an evaluation of the overall shrubland communities; the information that follows is
strength of the scientific evidence. for mostly positive relationships.
1.1 Forest/woodland vegetative elements or substrates.
Habitat Elements Biotic components found within a forested context.
Site-specific habitat elements (HEs) are those components 1.1.1 Down wood. Includes downed logs, branches,
of the environment believed to most influence wildlife and rootwads.
species’ distribution, abundance, fitness, and viability 1.1.1.1 Decay class. A system by which down
(definition adapted from Marcot et al.14 and Mayer and wood is classified based on its deterioration.
Laudenslayer.)16 In this context, HEs include natural 1.1.1.1.1 hard (class 1, 2). Little wood decay
attributes, both biological and physical (e.g., large trees, evident; bark and branches present; log
woody debris, cliffs, and soil characteristics) as well as resting on branches, not fully in contact with
anthropogenic features and their effects such as roads, ground; includes classes 1 and 2 as described
buildings, and pollution. Including these fine-scale in Thomas22.
attributes of an animal’s environment when describing 1.1.1.1.2 moderate (class 3). Moderate decay
the habitat associations for a particular species expands present; some branches and bark missing or
the concept and definition of habitat, a term widely used loose; most of log in contact with ground;
only to characterize the vegetative community or includes class 3 as described in Thomas22.
structural condition occupied by a species. Failing to assess 1.1.1.1.3 soft (class 4, 5). Well decayed logs;
and inventory HEs within these communities and bark and branches missing; fully in contact
conditions may lead to errors of commission; species may with ground; includes classes 4 and 5 as
be presumed to occur when in actuality they do not. described in Thomas.22
Habitat elements that influence a species negatively may 1.1.1.2 Down wood in riparian areas. Includes
preclude occupancy or breeding despite adequate floristic down wood in the terrestrial portion of riparian
or structural conditions. zones in forest habitats. Does not refer to in-
stream woody debris.
Defining Habitat Elements 1.1.1.3 Down wood in upland areas. Includes
Traditionally defined, the term habitat is that set of downed wood in upland areas of forest
environmental conditions, usually depicted as food, water, habitats.
and cover, used and selected for by a given organism. 1.1.2 Litter. The upper layer of loose, organic
Despite this broad definition, many land management (primarily vegetative) debris on the forest floor.
agencies use the term habitat to denote merely the Decomposition may have begun, but components
vegetation conditions and/or structural or seral stages still recognizable.
used by a particular species. However, many other 1.1.3 Duff. The matted layer of organic debris
environmental attributes or features influence and affect beneath the litter layer. Decomposition more
the population viability of wildlife species. Marcot et al.14 advanced than in litter layer; intergrades with
in their assessment of the terrestrial species of the uppermost humus layer of soil.
Columbia River Basin emphasized the importance of 1.1.4 Shrub layer. Refers to the shrub strata within
examining all features that exert influence on wildlife by forest stands.
 CHAPTER 3: STRUCTURES AND ELEMENTS 123

1.1.4.1 Shrub size. Refers to shrub height. 1.1.14.2.5 large tree 20-29 inches dbh
1.1.4.2 Percent shrub canopy cover. Percent of 1.1.14.2.6 giant tree >30 inches dbh
ground covered by vertical projection of shrub 1.1.14.3 Tree size. Measured in dbh, same as
crown diameter. 1.1.14.2 above.
1.1.4.3 Shrub canopy layers. Within a shrub 1.1.14.3.1 seedling <1 inch dbh
community, differences in shrub height and 1.1.14.3.2 sapling/pole 1-9 inches dbh
growth form produce multi-layered shrub 1.1.14.3.3 small tree 10-14 inches dbh
canopies in the forest understory. 1.1.14.3.4 medium tree 15-19 inches dbh
1.1.5 Moss. Large group of green plants without 1.1.14.3.5 large tree 20-29 inches dbh
flowers but with small leafy stems growing in 1.1.14.3.6 giant tree >30 inches dbh
clumps. 1.1.14.4 Mistletoe brooms/witches brooms. Dense
1.1.6 Flowers. A modified plant branch for the masses of deformed branches caused by any
production of seeds and bearing leaves specialized type of broom-forming parasite (fungal or
into floral organs. plant).
1.1.7 Lichens. Any of various lower plants made up 1.1.14.5 Dead parts of live tree. Portions of live
of an alga and a fungus growing as a unit on a trees with rot; can include broken tops;
solid surface. branches with decay; tree base with rot.
1.1.8 Forbs. Broad-leaved herbaceous plants. Does 1.1.14.6 Hollow living trees (chimney trees). Tree
not include grasses, sedges, or rushes. bole with large hollow chambers.
1.1.9 Cactus. Any of a large group of drought- 1.1.14.7 Tree cavities. Smaller chamber in a tree;
resistant plants with fleshy, usually jointed stems can be in bole, limbs, or forks of live or dead
and leaves replaced by scales or spines. trees. May be excavated or result from decay or
1.1.10 Fungi. Mushrooms, molds, yeasts, rusts, etc. damage.
1.1.11 Roots, tubers, underground plant parts. Any 1.1.14.8 Bark. Includes crevices or fissures, and
underground part of a plant that functions in loose or exfoliating bark.
nutrient absorbtion, aeration, storage, 1.1.14.9 Live remnant/legacy trees. A live mature
reproduction and/or anchorage. or old-growth tree remaining from the previous
1.1.12 Ferns. Any of a group of flowerless, seedless stand. Context is remnant trees in recently
vascular green plants. harvested or burned stands up through young
1.1.13 Herbaceous layer. Understory non-woody forested stands. See dead parts of live trees,
vegetation layer beneath shrub layer (forest hollow living trees, tree cavities, and bark to
context). May include forbs, grasses, ferns. see which species benefit from remnant trees
1.1.14 Trees. Includes both coniferous and with these attributes.
hardwood species. 1.1.14.10 Large live tree branches. Large branches
1.1.14.1 Snags. Standing dead trees. often growing horizontally out from the tree
1.1.14.1.1 Decay class. A system by which bole.
snags are classified based on their 1.1.14.11 Tree canopy layer. Refers to the strata
deterioration. occupied by tree crowns.
1.1.14.1.1.1 hard. Little wood decay 1.1.14.11.1 Sub-canopy. The space below the
evident; bark, branches, top, present; predominant tree crowns.
recently dead; includes class 1 as 1.1.14.11.2 Above canopy. The space above the
described in Brown3. predominant tree crowns.
1.1.14.1.1.2 moderate. Moderately decayed 1.1.14.11.3 Tree bole. The tree trunk.
wood; some branches and bark missing 1.1.14.11.4 Canopy. The more or less
and/or loose; top broken; includes classes continuous cover of branches and foliage
2 and 3 as described in Brown.3 formed collectively by the crowns of
1.1.14.1.1.3 soft. Well decayed wood; bark adjacent trees and other woody growth.
and branches generally absent; top 1.1.15 Fruits/seeds/nuts. Plant reproductive bodies
broken; includes classes 4 and 5 as that are used by animals.
described in Brown.3 1.1.16 Edges. The place where plant communities
1.1.14.2 Snag size. Measured in diameter at meet or where successional stages or vegetative
breast height (dbh) the standard measurement conditions within plant communities come
for standing trees taken at 4.5 feet above the together.
ground. 1.2 Shrubland/grassland vegetative elements or substrates.
1.1.14.2.1 seedling <1 inch dbh
1.1.14.2.2 sapling/pole 1-9 inches dbh
1.1.14.2.3 small tree 10-14 inches dbh
1.1.14.2.4 medium tree 15-19 inches dbh
Text continues on page 132
 124 Wildlife–Habitat Relationships in Oregon and Washington

Figure 1. Grass/Forb—Open

Figure 5. Sapling/Pole—Open

Figure 2. Grass/Forb—Closed

Figure 6. Sapling/Pole—Moderate

Figure 3. Shrub/Seedling—Open

Figure 7. Sapling/Pole—Closed
Figure 4. Shrub/Seedling—Closed
 CHAPTER 3: STRUCTURES AND ELEMENTS 125

Figure 8. Small Tree—Single Story—Open Figure 11. Medium Tree—Single Story—Open

Figure 9. Small Tree—Single Story—Moderate Figure 12. Medium Tree—Single Story—Moderate

Figure 10. Small Tree—Single Story—Closed Figure 13. Medium Tree—Single Story—Closed
 126 Wildlife–Habitat Relationships in Oregon and Washington

Figure 14. Large Tree—Single Story—Open Figure 17. Small Tree—Multi-story—Open

Figure 15. Large Tree—Single Story—Moderate Figure 18. Small Tree—Multi-story—Moderate

Figure 16. Large Tree—Single Story—Closed Figure 19. Small Tree—Multi-story—Closed

 CHAPTER 3: STRUCTURES AND ELEMENTS 127

Figure 20. Medium Tree—Multi-story—Open Figure 23. Large Tree—Multi-story—Open

Figure 21. Medium Tree—Multi-story—Moderate Figure 24. Large Tree—Multi-story—Moderate

Figure 22. Medium Tree—Multi-story—Closed Figure 25. Large Tree—Multi-story—Closed

 128 Wildlife–Habitat Relationships in Oregon and Washington

Figure 26a. Giant Tree—Multi-story Figure 27. Grass/Forb—Open

Figure 28. Grass/Forb—Closed

Figure 26b. Giant Tree—Multi-story. This photograph has
been included due to the limitations of the computer model.
 CHAPTER 3: STRUCTURES AND ELEMENTS 129

Figure 29. Low Shrub—Open Shrub Overstory—Seedling/ Figure 32. Low Shrub—Closed Shrub Overstory—Seedling/
Young Young

Figure 30. Low Shrub—Open Shrub Overstory—Mature Figure 33. Low Shrub—Closed Shrub Overstory—Mature

Figure 31. Low Shrub—Open Shrub Overstory—Old Figure 34. Low Shrub—Closed Shrub Overstory—Old
 130 Wildlife–Habitat Relationships in Oregon and Washington

Figure 35. Medium Shrub—Open Shrub Overstory— Figure 38. Medium Shrub—Closed Shrub Overstory—
Seedling/Young Seedling/Young

Figure 36. Medium Shrub—Open Shrub Overstory—Mature Figure 39. Medium Shrub—Closed Shrub Overstory—
Mature

Figure 37. Medium Shrub—Open Shrub Overstory—Old Figure 40. Medium Shrub—Closed Shrub Overstory—Old
 CHAPTER 3: STRUCTURES AND ELEMENTS 131

Figure 41. Tall Shrub—Open Shrub Overstory—Seedling/ Figure 44. Tall Shrub—Closed Shrub Overstory—Seedling/
Young Young

Figure 42. Tall Shrub—Open Shrub Overstory—Mature Figure 45. Tall Shrub—Closed Shrub Overstory—Mature

Figure 43. Tall Shrub—Open Shrub Overstory—Old Figure 46. Tall Shrub—Closed Shrub Overstory—Old
 132 Wildlife–Habitat Relationships in Oregon and Washington

Biotic components found within a shrubland or 1.2.12.1.1.3 soft. Well decayed wood; bark
grassland context (these are positive influences only). and branches generally absent; top
1.2.1 Herbaceous layer. Zone of understory broken; includes classes 4 and 5 as
nonwoody vegetation beneath shrub layer described in Brown.3
(nonforest context). May include forbs, grasses. 1.2.12.2 Snag size. Measured in dbh, as
1.2.2 Fruits/seeds/nuts. Plant reproductive bodies previously defined.
that are used by animals. 1.2.12.2.1 shrub/seedling <1 inch dbh
1.2.3 Moss. Large group of green plants without 1.2.12.2.2 sapling/pole 1-9 inches dbh
flowers but with small leafy stems growing in 1.2.12.2.3 small tree 10-14 inches dbh
clumps. 1.2.12.2.4 medium tree 15-19 inches dbh
1.2.4 Cactus. Any of a large group of drought- 1.2.12.2.5 large tree 20-29 inches dbh
resistant plants with fleshy, usually jointed stems 1.2.12.2.6 giant tree >30 inches dbh
and leaves replaced by scales or spines. 1.2.12.3 Tree size. Measured in dbh, as
1.2.5 Flowers. A modified plant branch for the previously defined.
production of seeds and bearing leaves specialized 1.2.12.3.1 shrub/seedling <1 inch dbh
into floral organs. 1.2.12.3.2 sapling/pole 1-9 inches dbh
1.2.6 Shrubs. Plant with persistent woody stems 1.2.12.3.3 small tree 10-14 inches dbh
and <16.5 feet tall; usually produces several basal 1.2.12.3.4 medium tree 15-19 inches dbh
shoots as opposed to a single bole. 1.2.12.3.5 large tree 20-29 inches dbh
1.2.6.1 Shrub size. Refers to shrub height. 1.2.12.3.6 giant tree >30 inches dbh
1.2.6.1.1 small <2.0 feet 1.2.13 Edges The place where plant communities
1.2.6.1.2 medium 2.0 - 6.5 feet meet or where successional stages or vegetative
1.2.6.1.3 large 6.5 - 16.5 feet conditions within plant communities come
1.2.6.2 Percent shrub canopy cover. Percent of together.
ground covered by vertical projection of shrub
crown diameter. 2. Ecological Habitat Elements
1.2.6.3 Shrub canopy layer. Within a shrub Selected interspecies relationships within the biotic
community, differences in shrub height and community; they include both positive and negative
growth form produce multi-layered shrub influences.
canopies. 2.1 Exotic species. Any non-native plant or animal,
1.2.6.3.1 Sub-canopy. The space below the including cats, dogs, and cattle.
predominant shrub crowns. 2.1.1 Plants. This field refers to the relationship
1.2.6.3.2 Above canopy. The space above the between an exotic plant species and animal
predominant shrub crowns. species.
1.2.7 Fungi. Mushrooms, molds, yeasts, rusts, etc. 2.1.2 Animals. This field refers to the relationship
1.2.8 Forbs. Broad-leaved herbaceous plants. Does between an exotic animal species and the animal
not include grasses, sedges, or rushes. species.
1.2.9 Bulbs/tubers. Any underground part of a plant 2.1.2.1 Predation. The species queried is preyed
that functions in nutrient absorbtion, aeration, upon by or preys upon an exotic species.
storage, reproduction and/or anchorage. 2.1.2.2 Direct displacement. The species queried
1.2.10 Grasses. Members of the Graminae family. is physically displaced by an exotic species,
1.2.11Cryptogamic crusts. Non-vascular plants that either by competition or actual disturbance.
grow on the soil surface. Primarily lichens, 2.1.2.3 Habitat structure change. The species
mosses, and algae. Often found in arid or semi- queried is affected by habitat structural
arid regions. May form soil surface pinnacles. changes caused by an exotic species, for
1.2.12 Trees (located in a shrubland/grassland example, cattle grazing.
context). Small groups of trees or isolated 2.1.2.4 Other. Any other effects of an exotic
individuals. species on a native species.
1.2.12.1 Snags. Standing dead trees. 2.2 Insect population irruptions. The species directly
1.2.12.1.1 Decay class. System by which snags benefits from insect population irruptions (i.e.,
are classified based on their deterioration. benefits from the insects themselves, not the resulting
1.2.12.1.1.1 hard. Little wood decay tree mortality or loss of foliage).
evident; bark, branches, top, present; 2.2.1 Mountain pine beetle. The species directly
recently dead; includes class 1 as benefits from mountain pine beetle eruptions.
described in Brown.3 2.2.2 Spruce budworm. The species directly benefits
1.2.12.1.1.2 moderate. Moderately decayed from spruce budworm irruptions.
wood; some branches and bark missing 2.2.3 Gypsy moth. The species directly benefits
and/or loose; top broken; includes classes from gypsy moth irruptions.
2 and 3 as described in Brown.3
 CHAPTER 3: STRUCTURES AND ELEMENTS 133

2.3 Beaver/muskrat activity. The results of beaver bare agricultural fields; natural bare exposed rock
activity including dams, lodges, and ponds, that are is under the rocky outcrop category.
beneficial to other species. 3.3.7 Playa (alkaline, saline). Shallow desert basins
2.4 Burrows. Aquatic or terrestrial cavities produced that are without natural drainage-ways where
by burrowing animals that are beneficial to other water accumulates and evaporates seasonally.
species. 3.4 Snow. Selected features of snow.
3.4.1 Snow depth. Any measure of the distance
3. Non-Vegetative, Abiotic, Terrestrial Habitat between the top layer of snow and the ground
Elements below.
Nonliving components found within any ecosystem. 3.4.2 Glaciers, snow field. Areas of permanent snow
Primarily positive influences with a few exceptions as and ice.
indicated.
3.1 Rocks. Solid mineral deposits. 4. Freshwater Riparian and Aquatic Bodies Habitat
3.1.1 Gravel. Particle size from 0.1-3.0 inches (0.2- Elements
7.6 cm) in diameter; gravel bars associated with Includes selected forms and characteristics of any body
streams and rivers are a separate category. of freshwater.
3.1.2 Talus. Accumulations of rocks at the base of 4.1Water characteristics. Includes various freshwater
cliffs or steep slopes; rock/boulder sizes varied attributes. Ranges of continuous attributes that are
and determine what species can inhabit the spaces key to the queried species, if known, will be in the
between them. comments.
3.1.3 Talus-like habitats. Refers to areas that contain 4.1.1 Dissolved oxygen. Amount of oxygen passed
many rocks and boulders but are not associated into solution.
with cliffs or steep slopes. 4.1.2 Water depth. Distance from the surface of the
3.2 Soils. Various soil characteristics. water to the bottom substrate.
3.2.1 Soil depth. The distance from the top layer of 4.1.3 Dissolved solids. A measure of dissolved
the soil to the bedrock or hardpan below. minerals in water
3.2.2 Soil temperature. Any measure of soil 4.1.4 Water pH. A measure of water acidity or
temperature or range of temperatures that are key alkalinity.
to the queried species. 4.1.5 Water temperature. Water temperature range
3.2.3 Soil moisture. The amount of water contained that is key to the queried species; if known, it is in
within the soil. the comments field.
3.2.4 Soil organic matter.The accumulation of 4.1.6 Water velocity. Speed or momentum of water
decomposing plant and animal materials found flow.
within the soil. 4.1.7 Water turbidity. Amount of roiled sediment
3.2.5 Soil texture. Refers to size distribution and within the water.
amount of mineral particles (sand, silt, and clay) 4.1.8 Free water. Water derived from any source.
in the soil; examples are sandy clay, sandy loam, 4.1.9 Salinity and alkalinity. The presence of salts.
silty clay, etc. 4.2 Rivers and streams. Various characteristics of
3.3 Rock substrates. Various rock formations. streams and rivers.
3.3.1 Avalanche chute. An area where periodic snow 4.2.1 Oxbows. A pond or wetland created when a
or rock slides prevent the establishment of forest river bend is cut off from the main channel of the
conditions; typically shrub and herb dominated river.
(sitka alder [Alnus sinuata] and/or vine maple 4.2.2 Order and class. Systems of stream
[Acer circinatum]). classification.
3.3.2 Cliffs. A high, steep formation, usually of 4.2.2.1 Intermittent. Streams/rivers that contain
rock. Coastal cliffs are a separate category under nontidal flowing water for only part of the
Marine Habitat Elements. year; water may remain in isolated pools.
3.3.3 Caves. An underground chamber open to the 4.2.2.2 Upper perennial. Streams/rivers with a
surface with varied opening diameters and high gradient, fast water velocity, no tidal
depths; includes cliff-face caves, intact lava tubes, influence; some water flowing throughout the
coastal caves, and mine shafts. year, substrate consists of rock, cobbles, or
3.3.4 Rocky outcrops and ridges. Areas of exposed gravel with occasional patches of sand; little
rock. floodplain development.
3.3.5 Rock crevices. Refers to the joint spaces in 4.2.2.3 Lower perennial. Streams/rivers with a
cliffs, and fissures and openings between slab low gradient, slow water velocity, no tidal
rock; crevices among rocks and boulders in talus influence; some water flowing throughout the
fields are a separate category (talus). year, substrate consists mainly of sand and
3.3.6 Barren ground. Bare exposed soil with >40% mud; floodplain is well developed.
of area not vegetated; includes mineral licks and
 134 Wildlife–Habitat Relationships in Oregon and Washington

4.2.3 Zone. System of water body classification 4.3 Ephemeral pools. Pools that contain water for only
based on the horizontal strata of the water brief periods of time usually associated with periods
column. of high precipitation.
4.2.3.1 Open water. Open water areas not closely 4.4 Sand bars. Exposed areas of sand or mud
associated with the shoreline or bottom. substrate.
4.2.3.2 Submerged/benthic. Relating to the bottom 4.5 Gravel bars. Exposed areas of gravel substrate.
of a body of water, includes the substrate and 4.6 Lakes/ponds/reservoirs. Various characteristics of
the overlaying body of water within 3.2 feet (1 lakes, ponds, and reservoirs.
m) of the substrate. 4.6.1 Zone. System of water body classification
4.2.3.3 Shoreline. Continually exposed substrate based on the horizontal strata of the water
that is subject to splash, waves, and/or periodic column.
flooding. Includes gravel bars, islands, and 4.6.1.1 Open water. Open water areas not closely
immediate nearshore areas. associated with the shoreline or bottom
4.2.4 In-stream substrate. The bottom materials in a substrates.
body of water. 4.6.1.2 Submerged/benthic. Relating to the bottom
4.2.4.1 Rocks. Rocks >10 inches (256 mm ) in of a body of water, includes the substrate and
diameter. the overlaying body of water within one meter
4.2.4.2 Cobble/gravel. Rocks or pebbles, .1-10 of the substrate.
inches (2.5-256 mm) in diameter, substrata may 4.6.1.3 Shoreline. Continually exposed substrate
consist of cobbles, gravel, shell, and sand with that is subject to splash, waves, and/or periodic
no substratum type >70% cover. flooding. Includes gravel bars, islands, and
4.2.4.3 Sand/mud. Fine substrata <.01 inch immediate nearshore areas.
(1mm) in diameter, little gravel present, may be 4.6.2 In-water substrate. The bottom materials in a
mixed with organics. body of water.
4.2.5 Vegetation. Herbaceous plants. 4.6.2.1 Rock. Rocks >10 inches (256 mm) in
4.2.5.1 Submergent vegetation. Rooted aquatic diameter.
plants that do not emerge above the water 4.6.2.2 Cobble/gravel. Rocks or pebbles, .1-10
surface. inches (2.5-256 mm) in diameter, substrata may
4.2.5.2 Emergent vegetation. Rooted aquatic consist of cobbles, gravel, shell, and sand with
plants that emerge above the water surface. no substratum type exceeding 70% cover.
4.2.5.3 Floating mats. Unrooted plants that form 4.6.2.3 Sand/mud. Fine substrata <.1 inch (2.5
vegetative masses on the surface of the water. mm) in diameter, little gravel present, may be
4.2.6 Coarse woody debris in streams and rivers. Any mixed with organics.
piece of woody material (debris piles, stumps, root 4.6.3 Vegetation. Herbaceous plants.
wads, fallen trees) that intrudes into or lies within 4.6.3.1 Submergent vegetation. Rooted aquatic
a river or stream. plants that do not emerge above the water
4.2.7 Pools. Portions of the stream with reduced surface.
current velocity, often with water deeper than 4.6.3.2 Emergent vegetation. Rooted aquatic
surrounding areas. plants that emerge above the water surface.
4.2.8 Riffles. Shallow rapids where the water flows 4.6.3.3 Floating mats. Unrooted plants that form
swiftly over completely or partially submerged vegetative masses on the surface of the water.
obstructions to produce surface agitation, but 4.6.4 Size. Refers to whether or not the species is
where standing waves are absent. differentially associated with water bodies based
4.2.9 Runs/glides. Areas of swiftly flowing water, on their size.
without surface agitation or waves, which 4.6.4.1 Ponds. Bodies of water <5 acre (2 ha).
approximates uniform flow and in which the 4.6.4.2 Lakes. Bodies of water >5 acre (2 ha).
slope of the water surface is roughly parallel to the 4.7Wetlands/marshes/wet meadows/bogs and swamps.
overall gradient of the stream reach. Various components and characteristics related to
4.2.10 Overhanging vegetation. Herbaceous plants any of these systems.
that cascade over stream and river banks and are 4.7.1 Riverine wetlands. Wetlands found in
<3.2 feet (1m) above the water surface. association with rivers.
4.2.11 Waterfalls. Steep descent of water within a 4.7.2 Context When checked, indicates that the
stream or river. setting of the wetland, marsh, wet meadow, bog,
4.2.12 Banks. Rising ground that borders a body of or swamp is key to the queried species.
water. 4.7.2.1 Forest. Wetlands within a forest.
4.2.13 Seeps or springs. A concentrated flow of 4.7.2.2 Nonforest. Wetlands that are not
ground water issuing from openings in the surrounded by forest.
ground.
 CHAPTER 3: STRUCTURES AND ELEMENTS 135

4.7.3 Size. When checked, indicates that the 5.2.8 Mixed-fine. Mixture of sand and mud
queried species is differentially associated with a particles <0.2 inch (4 mm) in diameter, little gravel
wetland, marsh, wet meadow, bog, or swamp present.
based on the size of the water body. 5.2.9 Mud. Fine substrata <.002 inch (0.06 mm) in
4.7.4 Marshes. Frequently or continually inundated diameter, little gravel present, usually mixed with
wetlands characterized by emergent herbaceous organics.
vegetation (grasses,sedges, reeds) adapted to 5.2.10 Organic. Substrata composed primarily of
saturated soil conditions. organic matter such as wood chips, leaf litter, or
4.7.5 Wet meadows. Grasslands with waterlogged other detritus.
soil near the surface but without standing water 5.3 Energy. Degree of exposure to oceanic swell,
for most of the year. currents, and wind waves.
4.8 Islands. A piece of land made up of either rock 5.3.1 Protected. No sea swells, little or no current,
and/or unconsolidated material that projects above and restricted wind fetch.
and is completely surrounded by water. 5.3.2 Semi-protected. Shorelines protected from sea
4.9 Seasonal flooding. Flooding that occurs periodically swell, but may receive waves generated by
due to precipitation patterns. moderate wind fetch, and/or moderate-to-weak
tidal currents.
5. Marine Habitat Elements 5.3.3 Partially exposed. Oceanic swell attenuated by
Selected biotic and abiotic components and characteristics offshore reefs, islands, or headlands, but shoreline
of marine systems. substantially exposed to wind waves, and/or
5.1 Zone. System of marine classification based on strong-to-moderate tidal currents.
water depth, and relationship to substrate. 5.3.4 Exposed. Highly exposed to oceanic swell,
5.1.1 Supratidal. The zone that extends landward wind waves, and/or very strong currents.
from the higher high water line up to either the 5.4 Vegetation. Includes herbaceous plants and plants
top of a coastal cliff or the landward limit of lacking vascular systems.
marine process (i.e., storm surge limit). 5.4.1 Mixed macro algae. Includes brown, green,
5.1.2 Intertidal. The zone between the higher high and red algae.
water line and the lower low water line. 5.4.2 Kelp. Subaquatic rooted vegetation found in
5.1.3 Nearshore subtidal. The zone that extends from the nearshore marine environment
the lower low water line seaward to the 65 foot (20 5.4.3 Eelgrass. Subaquatic rooted vegetation found
m) isobath, typically within .6 miles (1 km) of in an estuarine environment
shore. 5.5 Water depth. Refers to the vertical layering of the
5.1.4 Shelf. The area between the 65-650 feet (20- water column.
200 m) isobath, typically within 36 miles (60 km) 5.5.1 Surface layer. The uppermost part of the water
of shore. column.
5.1.5 Oceanic. The zone that extends seaward from 5.5.1.1 Tide rip. A current of water disturbed by
the 650 feet (200 m) isobath. an opposing current, especially in tidal water or
5.2 Substrates. The bottom materials of a body of by passage over an irregular bottom.
water. 5.5.1.2 Surface microlayer (neuston). The thin
5.2.1 Bedrock. The solid rock underlying surface uppermost layer of the water’s surface.
materials. 5.5.2 Euphotic. Upper layer of a water body that
5.2.2 Boulders. Large, worn, rocks >10 inches (256 receives sufficient sunlight for the photosynthesis
mm) in diameter. of plants.
5.2.3 Hardpan. Consolidated clays forming a 5.5.3 Disphotic. Area below the euphotic zone
substratum firm enough to support an epibenthos where photosynthesis ceases.
and too firm to support a normal infauna (clams, 5.5.4 Demersal/benthic. Submerged lands including
worms, etc.), but with an unstable surface that vegetated and unvegetated areas.
sloughs frequently. 5.6 Water temperature. Measure of ocean water
5.2.4 Cobble. Rocks or pebbles, 2.5-10 inches (64- temperature.
256 mm) in diameter, may be a mix of cobbles, 5.7 Salinity. The presence and concentration of salts;
gravel, shells, and sand, with no type exceeding salinity range that is key to the species, if it is known,
70% cover. will be in the comments field.
5.2.5 Mixed-coarse. Substrata consisting of cobbles, 5.8 Forms. Morphological elements within marine
gravel, shell, and sand with no substratum type areas.
exceeding 70% cover. 5.8.1 Beach. An accumulation of unconsolidated
5.2.6 Gravel. Small rocks or pebbles, 0.2-2.5 inches material (sand, gravel, angular fragments) formed
(4-64 mm) in diameter. by waves and wave-induced currents in the
5.2.7 Sand. Fine substrata <0.2 inch (4 mm) in intertidal and subtidal zones.
diameter, little gravel present, may be mixed with
organics.
 136 Wildlife–Habitat Relationships in Oregon and Washington

5.8.2 Off-shore islands/rocks/sea stacks/off-shore cliffs. harvest/poaching, incidental take (resulting from
A piece of land made up of either rock and/or fishing net by-catch, or by hay mowing, for example),
unconsolidated material that projects above and is and targeted removal for pest control.
completely surrounded by water at higher high 8.7 Fences/corrals. Wood, barbed wire, or electric
water for large (spring) tide. Includes off-shore fences.
marine cliffs. 8.8 Supplemental food. Food deliberately provided for
5.8.3 Marine cliffs (mainland). A sloping face steeper wildlife (e.g., bird feeders, ungulate feeding
than 20½ usually formed by erosion and composed programs, etc.) as well as spilled or waste grain along
of either bedrock and/or unconsolidated railroads and cattle feedlots.
materials. 8.9 Refuse. Any source of human-derived garbage
5.8.4 Delta. An accumulation of sand, silt, and (includes landfills).
gravel deposited at the mouth of a stream where it 8.10.Supplemental boxes, structures and platforms.
discharges into the sea. Includes bird houses, bat boxes, raptor and
5.8.5 Dune. In a marine context; a mound or ridge waterfowl nesting platforms.
formed by the transportation and deposition of 8.11 Guzzlers and waterholes. Water sources typically
wind-blown material (sand and occasionally silt). built for domestic animal use.
5.8.6 Lagoon. Shallow depression within the shore 8.12 Toxic chemical use. Proper use of regulated
zone continuously occupied by salt or brackish chemicals; documented effects only.
water lying roughly parallel to the shoreline and 8.12.1 Herbicides/fungicides. Chemicals used to kill
separated from the open sea by a barrier. vegetation and fungi.
5.8.7 Salt marsh. A coastal wetland area that is 8.12.2 Insecticides. Chemicals used to kill insects.
periodically inundated by tidal brackish or salt 8.12.3 Pesticides. Chemicals used to kill vertebrate
water and that supports significant (15% cover) species.
nonwoody vascular vegetation (e.g., grasses, 8.12.4 Fertilizers. Chemicals used to enhance
rushes, sedges) for at least part of the year. vegetative growth.
5.8.8 Reef. A rock outcrop, detached from the 8.13 Hedgerows/windbreaks. Woody and/or shrubby
shore, with maximum elevations below the high- vegetation either planted or that develops naturally
water line. along fencelines and field borders.
5.8.9 Tidal flat. A level or gently sloping (<5½) 8.14 Sewage treatment ponds. Settling ponds associated
constructional surface exposed at low tide, usually with sewage treatment plants.
consisting primarily of sand or mud with or 8.15 Repellents. Various methods used to repel or
without detritus, and resulting from tidal deter wildlife species that damage crops or property
processes. (excluding pesticides and insecticides).
5.9 Water clarity. As influenced by sediment load. 8.15.1 Chemical (taste, smell, or tactile). Chemical
substances that repel wildlife.
6. (No Data) 8.15.2 Noise or visual disturbance. Nonchemical
Formerly contained topographic information, such as methods to deter wildlife.
elevation, that has been moved to the life history matrix. 8.16 Culverts. Drain crossings under roads or
railroads.
7. Fire as a Habitat Element
8.17 Irrigation ditches/canals. Ditches built to transport
Refers to species that benefit from fire. The time frame
water to agricultural crops or to handle runoff.
after which the habitat is suitable for the species, if known,
8.18 Powerlines/corridors. Utility lines, poles, and
will be found in the comments field.
rights-of-way associated with transmission,
8. Anthropogenic Related Habitat Elements telephone, and gas lines.
This section contains selected examples of human-related 8.19 Pollution. Human-caused environmental
Habitat Elements that may be a key part of the contamination.
environment for many species. These Habitat Elements 8.19.1 Chemical. Contamination caused by
may have either a negative or positive influence on the chemicals.
queried species. 8.19.2 Sewage. Contamination caused by human
8.1 Campgrounds/picnic areas.Sites developed and waste.
maintained for camping and picnicking. 8.19.3 Water. Aquatic contamination from any
8.2 Roads. Either paved or unpaved. source.
8.3 Buildings. Permanent structures. 8.20. Piers. Structures built out over water.
8.4 Bridges. Permanent structures typically over water 8.21 Mooring piles, dolphins, buoys. Floating objects
or ravines. anchored out in the water for nautical purposes.
8.5 Diseases transmitted by domestic animals. Some 8.22 Bulkheads, seawalls, revetment. Retaining
domestic animal diseases may be a source of structures built to protect the shoreline from wave
mortality or reduced vigor for wild species. action.
8.6 Animal harvest or persecution. Includes illegal 8.23 Jetties, groins, breakwaters. Structures built to
influence the current or protect harbors.
 CHAPTER 3: STRUCTURES AND ELEMENTS 137

8.24 Water diversion structures. Structures built to attributes found in the category HEs; other HEs may play
funnel or direct water, including dams, dikes and a larger role in driving the species’ occurrence.
levies. Salamanders and other amphibians, for example, may
8.25 Log boom. A raft of logs lashed together either to occur in forested habitats but are most influenced by water
transport the logs or as barriers to boat traffic near characteristics, found in the category Freshwater Riparian
marinas or dams. and Aquatic Bodies discussed below. Thus, while the
8.26 Boats/ships. Watercraft, either motorized or broader Wildlife-habitat determines which HEs (either
nonmotorized. forested or nonforested) a species may be associated with,
8.27 Dredge spoil islands. Sediment deposited from however, it does not mandate an association. That is, when
dredging operations. a HE from either or both of the two subclasses Forest/
8.28 Hatchery facilities and fish. Fish that are hatched in Woodland Vegetative Elements or Shrubland/Grassland
captivity and later released into the wild. For Vegetative Elements is associated with a species, the
simplicity this refers to freshwater areas, though association(s) imply that the HE is key to the species well
marine birds and mammals likely feed on hatchery- being because it promotes the species’ distribution,
released fish too. This also includes the facilities and abundance, fitness, and viability
their operation. Category 2, Ecological Habitat Elements, contains HEs
that describe interspecific relationships. Namely, these
Habitat Elements Data Matrix elements depict the influence of exotic species, insect
Based on Marcot’s work, the Habitat Elements or HEs population irruptions, beaver/muskrat activities, and
Digital Matrix (found on the CD-ROM) focuses on those burrows on a given species. The first two, Exotic Species
correlates that consist of fine-scale or within-stand and Insect Population Control may exert either positive
features. In keeping with the initial intent, HEs do not or negative influences on a given species, whereas the
include those elements that may be used, but are latter two depict positive relationships only. For example,
nonessential to the success or viability of a population, only species that are positively influenced by another
thus they may also be thought of as the drivers of habitat species’ burrow (e.g., secondary burrow users) will be
selection. The HEs matrix depicts associations with all associated with burrows. Species that are negatively
common species, including both residents and migrants, influenced, such as animals that may break their legs
in Oregon and Washington. stepping into badger burrows, are not.
The list of HEs and their definitions was derived from Nonvegetative, Abiotic Terrestrial Habitat Elements,
Marcot et al.14 and was refined and edited based on the Category 3, contains a variety of abiotic HEs such as Rocks,
published literature (in Literature Cited, these are Soils, Rock Substrates, and Snow; each has >2 subclasses.
identified with an asterisk) and expert review. The final Habitat Elements under the classes Soils and Snow may
list comprises 287 HEs, including naturally occurring depict either negative or positive influences. Freshwater
biological and physical elements as well as elements Riparian and Aquatic Bodies Habitat Elements, Category
created or caused by human actions. Definitions are 4 characterizes many aquatic attributes, including those
provided to characterize each element, and clarify the associated with rivers and streams, lakes, ponds and
nature of its influence on wildlife species. The HEs are reservoirs, and wetlands/marshes/wet meadows/bogs/
grouped into seven major categories, six of which have swamps. These HEs depict primarily positive influences,
related subclasses (Table 2). with the exception of the class Water Characteristics,which
Category 1, entitled Habitat Elements, contains two includes Habitat Elements such as Water Temperature,
broad subclasses: Forest/Woodland Vegetative Elements Water Velocity and Water Depth, all of which could either
or Substrates and Shrubland/grassland Vegetative promote or inhibit species’ viability.
Elements or Substrates. These subclasses denote the Marine Habitat Elements are found in Category 5. Only
floristic components and attributes that are found within species that occur in the Marine Wildlife-habitats are
forested and nonforested (shrubland/grassland) associated with Marine Habitat Elements. Examples
communities. Floristic components include plants (lichens, include the classes Zone, Substrates, Energy Vegetation,
fungi, and ferns), and plant parts (roots, flowers, and bark); Water Depth, Water Temperature Salinity, Forms, and
attributes include the herbaceous and shrub layers. The Water Clarity. The only HEs that may exhibit either
subclasses are nested within the broad wildlife habitats positive or negative influences are Water Temperature and
(see Chapter 1). That is, only species that occur within Salinity.
Forested Wildlife-habitats may be associated with Forest/ Category 7 consists of only one Habitat Element, Fire.
Woodland Vegetative Elements or Substrates, and only This element, when associated with a given species,
species that occur within Shrubland/Grassland Wildlife- indicates that fire has a positive effect on the species
habitats may be associated with Shrubland/Grassland distribution, occurrence, and viability. Some species that
Vegetative Elements or Substrates. Some species that occur benefit from fire include grazers, which feed on the lush
within both forested and nonforested habitats may be new growth initiated after burns, cavity and snag users,
associated with HEs in both of the subclasses. Conversely, and species that use down wood. Panelists indicated in
some forest or shrubland/grassland species may not be the comments the period of time after which the burned
associated with any of the floristic components and stand becomes beneficial to the species.
 138 Wildlife–Habitat Relationships in Oregon and Washington

Lastly, Category 8, Anthropogenic-related Habitat example, there are 130 species closely associated with the
Elements, contains human-made features, such as roads, forest structural conditions, but 40 (38%) of these species
buildings, and fences; activities such as toxic chemical use, require a specific Habitat Element to reproduce in the
animal harvest and hatchery fish releases; and outcomes, forest environment. Thus, when classifying what is or is
such as pollution and refuse. We included these HEs to not habitat for a particular species it is imperative to look
acknowledge that most animals encounter the effects of across scales and ask, What is important at the landscape
humans in their environment, some of which may benefit or regional scale? This information is provided in the broad
certain species, many of which do not. Because of the dual Wildlife-habitat Types Matrix that depicts habitat
nature of these anthropogenic elements, all of them may relationships based on floristic communities (see Chapter
exert positive or negative influences depending on a 1). Next, one must assess the particular structures and/or
variety of factors, including the species involved, the seral stages within those communities to derive
intensity and scale of the activity or effect, and the duration information on how species occurrence is influenced by
of the activity. vegetation of varying age, size class, and density. The
On completion of the HEs list, we used expert panels Structural Conditions Matrix, defined in this chapter,
to indicate which HEs are associated with each species. elucidates wildlife-habitat relationships at what can be
Specifically, we posed the question, Which HEs, based on thought of as the “stand-level.” Lastly, the fine-scale
our list, most strongly influence this species’ distribution, features, both physical and biological, that are essential
abundance, fitness, and viability? As a group, the panelists for a species occurrence or precludes it, must be identified.
reviewed the HEs list, and indicated those elements they The Habitat Elements Matrix, described in this chapter,
believed to be most relevant. When necessary, the panelists delineates the with in-stand attributes that most influence
formulated comments to elucidate the relationship each species.
between the Habitat Element and the species. Clarification
was particularly important for Habitat Elements that may Literature Cited
exert both positive and negative influences. Other types 1. *American Fisheries Society. 1985. Aquatic habitat inventory, glossary
of information that may be found in the comments include and standard methods. Habitat Inventory Committee, American
the following: (1) the range of values necessary for use of Fisheries Society, Western Division.
the HE. For example, if the element Water Depth is listed 2. Bilby, R. 1988. Interactions between aquatic and terrestrial systems.
as being important to the animal, the specific water Pages 13-29 in: K. Raedeke, editor. Streamside management-riparian
wildlife and forestry interactions. University of Washington, Institute of
depth(s) may be described; (2) the spatial, temporal, or
Forest Resources, Seattle, WA. Contribution 59.
geographical/topographical contexts necessary for use of 3. *Brown, E. R., editor. 1985. Management of wildlife and fish habitats in
the HE. That is, whether the HE is used only in certain forests of western Oregon and Washington. U.S. Forest Service,
parts of the animal’s range, or in specific seasons of the Publication R6-F&WL-192-1985, Portland, OR.
year, or on particular slopes, etc.; (3) whether the 4. *Buckman, H. O., and N. C. Brady. 1969. The nature and properties of
importance of the HE is dependent on relationships with soils, 7th edition. Macmillan Co., New York, NY.
other HEs, Structural Conditions, and/or Wildlife 5. *Bull, E. L., C. G. Parks, and T. R. Torgersen. 1997. Trees and logs
important to wildlife in the interior Columbia River basin. U.S. Forest
Habitats. If so, the relevant habitat components are listed;
Service General Technical Report 391. Portland, OR.
and (4) the activity (e.g., reproduction, foraging, etc.) 6. *Cooperrider, A. Y., R. J. Boyd, and H. R. Stuart. 1986. Inventory and
associated with the HE. Some HEs may only be used for monitoring of wildlife habitat. U.S. Bureau of Land Management Service
specific purposes, for example, snags during the nesting Center, BLM-YA-PT-87/001. Denver, CO.
season. 7. *Cowardin, L. M., V. Carter, F. C. Golet, and E. T. LaRoe. 1979.
Classification of wetland and deepwater habitats of the United States.
U.S. Fish and Wildlife Service, FWS/OBS-79/31.
Conclusion 8. *Dethier, M. N. 1990. A marine and estuarine habitat classification
The final Structural Conditions and Habitat Elements system for Washington state. Washington Natural Heritage Program,
Matrices are the most complete characterization of Department of Natural Resources, Olympia, WA.
intermediate and fine-scale features to date; however, 9. *Forest Ecosystem Management Assessment Team. 1993. Forest
ecosystem management: an ecological, economic and social assessment.
because of the sheer magnitude of the exercise, some
U.S. Forest Service, Portland, OR.
species associations may have been mistakenly omitted. 10. Franklin, J., and T. Spies. 1991. Composition, function, and structure of
We expect that as our collective knowledge about wildlife- old-growth Douglas-fir forests. Pages 71-80 in: L. F. Ruggiero, K. B. Aubry,
habitat relationships evolves, additions and corrections A. B. Carey, and M. H. Huff, editors. Wildlife and vegetation of
to all three habitat matrices (Wildlife-Habitat Types, unmanaged Douglas-fir forests. U.S. Forest Service, Pacific Northwest
Structural Conditions, and Habitat Elements) may be Research Station, Portland, OR. General Technical Report PNW-GTR-
identified. The matrices associated with this book are 285.
11. Hattenschwiler, S., F. Miglietta, A. Raschi, and C. Korner. 1997. Thirty
relationally tied to illustrate the fact that wildlife exist in
years of in situ tree growth under elevated CO2, a model for future
a multi-dimensional environment. For this reason, we forest responses? Global Change Biology 2: 377-387.
stress the importance of getting the big picture and using 12. *Howes, D. E., J. R. Harper, and E. H. Owens. 1994. Physical shore-zone
the matrices to their best advantage, that is, as inter-related mapping system for British Columbia. British Columbia Ministry of
databases, each providing a part of the equation that Environment, Lands and Parks, Victoria, BC, Canada.
depicts in its entirety the wildlife environment. For
 CHAPTER 3: STRUCTURES AND ELEMENTS 139

13. Kauffman, B. 1988. The Status of Riparian Habitats in Pacific

Northwest Forests. Pages 45-55 in: K. Raedeke, editor. Streamside
management!riparian wildlife and forestry interactions. University of
Washington, Institute of Forest Resources, Seattle, WA. Contribution
59.
14. Marcot, B. G., M. A. Castellano, J. A. Christy, L. K. Croft, J. F. Lehmkuhl,
R. H. Naney, K. Nelson, C. G. Niwa, R. E. Rosentreter, R. E. Sandquist, B.
C.Wales, and E. Zieroth. 1997. Pages 1497-1713 in: T. M. Quigley and S. J.
Arbelbide, editors. Terrestrial ecology assessment: an assessment of
ecosystem components in the interior Columbia Basin and portions of
the Klamath and Great Basins. Volume 3. US Forest Service General
Technical Report PNW-GTR-405. U.S. Forest Service Pacific
Northwest Research Station, Portland, OR.
15. *Maser, C., J. M. Geist, D. M. Concannon, R. Anderson, and B. Lovell.
1979. Wildlife-habitats in managed rangelands. The Great Basin of
southeastern Oregon: geomorphic and edaphic habitats. U.S. Bureau of
Land Management, General Technical Report PNW-99.
16. *Mayer, K. E., and W. F. Laudenslayer, Jr., editors. 1988. A guide to
wildlife-habitats of California. California Department of Forestry and
Fire Protection, Sacramento, CA.
17. Norby, R.J., C. A. Gunderson, S. D. Wullschleger, E. G. O’Neill, and M. K.
McCracken. 1992. Productivity and compensatory responses of
yellow-poplar trees in elevated CO2, Nature 357: 322-324.
18. *Parks, C. G., E. L. Bull and T. R. Torgerson. 1997. Field guide for the
identification of snags and logs in the interior Columbia River basin.
U.S. Forest Service General Technical Report 390. Portland, OR.
19. Ruggiero, L. F., L. C. Jones, K. B. Aubry. 1991. Plant and animal habitat
associations in Douglas-fir forests of the Pacific Northwest: an
overview. Pages 447-462 in: L. F. Ruggerio, K. B. Aubry, A. B. Carey, and M.
H. Huff, editors. Wildlife and vegetation of unmanaged Douglas-fir
forests. U.S. Forest Service Pacific Northwest Research Station,
Portland, OR. General Technical Report PNW-GTR-285.
20. *Smith, R. L. 1966. Ecology and field biology. Harper and Row, New
York, NY.
21. *Stevens, V. 1995. Wildlife diversity in British Columbia: distribution
and habitat use of amphibians, reptiles, birds, and mammals in
biogeoclimatic zones. Research Branch, British Columbia Ministry of
Forestry, Wildlife Branch, British Columbia Ministry of the Environment,
Lands and Parks,Victoria, BC, Canada. Work Paper 04/1995.
22. *Thomas, J. W. 1979. Wildlife-habitats in managed forests—the Blue
Mountains of Oregon and Washington. U.S. Forest Service Agricultural
Handbook No. 553, Portland, OR.
23. *Washington Department of Fish and Wildlife. 1993. Guidelines for
using lake and stream survey forms. Washington Department of Fish
and Wildlife, Fisheries Management Division. Olympia, WA.
24. *West, N. R. 1990. Structure and function of microphytic soil crusts in
wildland ecosystems of arid to semi-arid regions. In M. Begon, A. H.
Fitter and A. Macfadyen, editor. Advances in Ecological Research,
Academic Press, Volume 20. San Diego, CA.
 4
Management of Within-stand Forest
Habitat Features
William C. McComb

Introduction
Managing forests to produce a desirable mix of forest distribution of animals in our forests. I will introduce
resources, including timber and wildlife, requires an concepts of habitat function, population change, and
understanding of how animals respond to habitat within habitat patterning. These concepts are used to illustrate
and among stands over a landscape. Management how silvicultural systems and habitat patterning can
strategies aimed at long-term population change are most influence demographic patterns of animals over large
likely to succeed if they alter habitat quantity, quality, and/ forest areas. The result, finally, is a framework for meeting
or distribution. In this chapter, I focus on characteristics forest-wildlife goals through habitat manipulation.
of habitat that could be manipulated through silviculture
or vegetation management within stands. Habitat also Habitat Function
includes abiotic factors such as soils, geology and climate, A population gains energy from food resources and
but vegetative characteristics represent the greatest conserves energy by exploiting cover resources (Figure
opportunity for manipulation. 1). The rate of net primary production is fixed over large
I provide a conceptual framework for wildlife habitat areas and time, because there is a solar constant and
management at the stand scale that silviculturists can because climate changes are relatively slow. Herbivores
consider when taking actions to manage wildlife habitat in forests exist in a sea of energy, but food quantity may
and, ultimately, wildlife populations and communities. not be as important as food quality.36 Animals require the
That framework builds on a common understanding of digestible energy in food, but indigestible portions of food
several terms: habitat is the place where a species lives. It (e.g., cellulose, lignin, chiton, or bones) or compounds in
includes the physical and biological resources necessary the plants that inhibit digestion (tannins and other
to support a viable population over space and through phenols) reduce food quality. Net energy available after
time. Each species and each population has its own habitat metabolism increases the animal’s fitness depending on
requirements. Populations are self-sustaining assemblages the quality of the cover available to the animal.
of individuals of one species over space and through time. Many browse species contain high levels of phenols
Communities, by contrast, are assemblages of populations that reduce their digestibility for many herbivores,19 but
over space and through time. mule deer saliva contains a substance (prolene) that binds
There are two common approaches to management of with the phenols and reduces their effectiveness.49 Some
forest habitat for wildlife: management of individual plants, therefore, may have high levels of compounds that
species and management of communities. Occasionally
managers choose to focus management on indicator Solar Energy
species or guilds of species in order to meet the needs for
many other species. This approach should be avoided, Photosynthesis
Fitness: Reproduction,
because each species has its own habitat requirements and Movement, Basal
each will respond differently to management activities.31 Food: Energy Metabolism
Forest disturbances change the abundance of individuals Inputs
in most populations, and those changes also affect the
composition of wildlife communities. Management of Digestible Energy Available Energy
individual species, therefore, has consequences for other Heat Loss
species in the community occupied by the species that is
being managed. Forest disturbances that benefit black- Feces, Net Energy
tailed deer, for example, probably would benefit creeping Urine Cover: Energy
voles and orange-crowned warblers but not Douglas Conservation
squirrels or pileated woodpeckers.
Vegetation management by forest-land managers is Figure 1. Conceptual diagram of acquisition and conservation
probably the greatest factor influencing the abundance and of energy by an animal and the effect on individual fitness.

140
 CHAPTER 4: MANAGING FOREST ELEMENTS 141

Lower critical temperature pocket gophers). Some species use water as a form of cover
to enhance evaporative cooling (e.g., elk) or to escape
predators (e.g., white-tailed deer). Still others such as
amphibians require free water or moist environments for
reproduction.
Metabolic rate

The size of habitat is an important determinant of its

Upper critical suitability for a species. A patch of habitat must be
temperature sufficiently large to provide energy inputs and energy
Zone of thermal conservation features to sustain a population. Habitat may
neutrality occur in one large unit, but more commonly it is
distributed in patches through other less suitable habitat.
If these habitat patches are too widely distributed, then
the animal expends more energy moving among patches
than it receives from those patches. The amount of habitat,
Ambient temperature
therefore, and its quality and distribution are interrelated.
Figure 2. The zone of thermal neutrality is the ambient Increasing any or all of these attributes of habitat increases
temperature where a homeothermic organism expends the the net energy available to animals that use this energy to
least amount of energy maintaining its body temperature. maintain body temperature, move to food and cover, and
reproduce.

reduce digestibility for some wildlife species, but the co- Population Growth
evolution of plants and herbivores has resulted in plant Populations can be manipulated by modifying habitat and
defense mechanisms that are less effective for herbivores thereby influencing animal fitness. The linkages between
such as deer. animal demography and habitat are complex, but some
It is advantageous for an animal to conserve any energy understanding of these relations is necessary for successful
that it acquires. Mammals and birds that maintain a habitat management. Each species has its own potential
constant body temperature expend a large amount of for population increase, and this potential is described as
energy to maintain that temperature. Energy expenditures the intrinsic rate of natural increase. There is a solar constant,
to maintain body temperature are minimized in an so energy available to animals is limited (Figure 1). Food,
animal’s thermal neutral zone (Figure 2). Any departure therefore, becomes scarcer or of poorer quality as the
from the thermal neutral zone results in increased population grows. As populations grow, cover is occupied
expenditure of energy, so animals often select habitat that by other individuals and the risk of disease and parasitism
reduces climatic extremes. There are upper and lower increases. Interspecific competition for resources causes
critical temperatures beyond which is lethal. Cover from some subordinate individuals to use suboptimal cover.18
overheating is especially important to large animals such As food, cover, or other resources become limiting, the
as elk that may find it particularly difficult to release excess population growth rate decreases, because mortality
heat unless water is available to aid in evaporative cooling. increases or because reproduction decreases. This process
Cover from severe cold is especially important to a species is termed logistic growth (Figure 3). If we assume that
with a high surface area to mass ratio (e.g., small bird). resources are constant, then the population reaches a point
Cover that allows an animal to stay within an acceptable where births equal deaths and growth becomes 0. This
range of temperatures (particularly those that approach point is termed the carrying capacity of the habitat for the
the thermal neutral zone) is important to maintaining a population.
positive balance of net energy and hence influences animal But resources are not constant; they change daily,
fitness. seasonally, and annually. Birth rates, death rates, and
Cover also can refer to the portion of habitat that an movement rates are variable over both space and time as
animal uses for nesting and escape from predators. The habitat changes through disturbances and succession.
effectiveness of nest box programs for wood ducks, Carrying capacity, consequently, is always changing. The
bluebirds and other secondary cavity-using species, concept of a dynamic carrying capacity is useful to land
demonstrates that manipulation of the quantity, quality, managers, because it provides the link between habitat
and availability of cover resources can be an effective quality and population growth. Manipulating habitat to
management technique.39 Land managers can manage change carrying capacity is a particularly effective
habitat for a species by altering food quality, quantity, and/ approach to long-term manipulation of wildlife
or availability while also altering the quality, quantity, populations.
and/or availability of cover. This strategy can lead to Populations do not always reach carrying capacity in
drastic changes in habitat quality for the species. relation to habitat quality. Some species, such as voles,
Water is differentially important to animal species. snowshoe hares, and ruffed grouse, follow a “boom and
Some require free water or high humidity (mountain bust” population pattern. Populations grow for about 3-6
beaver, for example, have a primitive uretic system). Other years and then rapidly decline for another 3-6 years. High-
species obtain most of their water from their food (e.g., quality habitat usually increases the highs and decreases
 142 Wildlife–Habitat Relationships in Oregon and Washington

Food scarcity The following section describes natural disturbances

Disease common in the Pacific Northwest and the habitat features
Competition that seem to change dramatically depending on the type
Carrying
Cover quality capacity of disturbance.

Natural Disturbance in Western

Population size

Coniferous Forests
Species inhabiting forests often are adapted to the natural
forest disturbance regimes in the region. Some species
colonize after large, recent disturbances; others recolonize
after the forest develops; and still others occupy late
successional stages. Dead wood can be an important
component of each of these seral stages (Table 1). Some
colonizers of early seral stages are more abundant now
than ever before. They include many herbivores that can
Time influence the trajectory of ecological succession within a
stand,45 particularly where the size and distribution of
Figure 3. Logistic population growth occurs as resources various seral stages optimize rates of population growth
become more limited with increasing populations. and colonization for each species.
Knowledge of natural disturbances in western
the lows of a population cycle, but habitat probably does
coniferous forests can help when developing silvicultural
not directly mediate these cycles, because they occur
systems that might meet the needs of forest-associated
throughout much of the geographic range of the species.26
wildlife.40 Coarse-scale disturbances typically occur over
tens or hundreds of acres (hectares) both within stands as
Movement and Dispersal well as among them. Fine-scale disturbances occur within
Animals move for various reasons, but movement to and stands at a scale of <1 tree height in width. Coarse- and
from food and cover is the most obvious daily requirement fine-scale disturbances have affected the establishment,
of any species. This distance is very short for species such development, and destruction of unmanaged western
as clouded salamanders, while others such as American coniferous forests.
marten may move among stands over a landscape. The The term patch is used to describe a homogeneous unit
arrangement of food and cover in an area can affect energy of resources that is different from the matrix in which it
costs associated with movements from one resource to occurs.42 Stands and patches can be synonymous or
another. Another type of movement, dispersal, may result patches can occur within stands, depending on the
from overcrowding. As a population approaches carrying resources under consideration. Patches may occur at any
capacity, it can be energetically advantageous for some scale; a canopy gap may be a patch within a stand and a
individuals search for more acceptable habitat.18 Dispersal stand may be a patch within a landscape.
can be influenced by habitat quality. The rate of movement
and the time- and habitat-specific probability of survival Coarse-scale disturbances
in various habitat types during dispersal influence the Except in coastal forests, fire was the most frequent and
probability that an organism will survive its dispersal to widespread coarse-scale disturbance occurring at scales
a good patch of habitat.62 up to 250,000 acres (100,000 ha). The fire regime of the
If we assume that species have evolved and persisted Douglas-fir (Pseudotsuga menziesii) region in western
within a range of natural variability from natural Oregon and Washington is complex and includes fires of
disturbances in forest systems, then we might better be varying frequencies, intensities, sizes, and patterns. For
able to understand how human-induced disturbances may instance, return intervals for stand-replacement fires at
influence habitat availability and population dynamics. Mount Rainier National Park in the Washington Cascades

Table 1. Examples of animal species that would use four types of dead or dying trees in four stages of stand
development in Pacific Northwest forests.

Tree decay stage

Seral stage Live cavity-tree Hard snag Soft snag Log

Stand reinitiation House wren American kestrel Western bluebird Alligator lizard
Stem exclusion Winter wren Hairy woodpecker Chestnut-backed chickadee Ensatina salamander
Transition Red-breasted nuthatch Northern flicker Red-breasted sapsucker Clouded salamander
Shifting-gap phase Spotted owl Pileated woodpecker Northern flying squirrel American marten
 CHAPTER 4: MANAGING FOREST ELEMENTS 143

are 400-500 years.24 Stand-replacement fires in the central

Oregon Cascades have return intervals of >200 years,
while smaller ground fires that kill some green trees have
return intervals of about 100-150 years.44, 58
Currently several different scenarios of forest
development and succession are recognized for Oregon
and Washington.1, 44, 57 If a fire kills most of the trees in a
stand and initiates establishment of Douglas-fir and
western hemlock (Tsuga heterophylla) seedlings, then the
stand will probably develop old-growth structure at a
stand age of 150-250 years in the absence of management
(depending on site quality). Snags and logs in such stands
would probably follow a pattern of increase, decrease, and
increase over a 400- to 800-year cycle55 (Figure 4).
Lower intensity fires kill fewer trees in a patchy pattern
(2-100 acres [1-40 ha]). A stand of various tree sizes and Figure 4. A stand replacement fire kills most trees in a stand,
large dead trees produced a multi-layered canopy sooner but leaves considerable amounts of dead wood, which is used
than in the first scenario. Inputs of dead wood are more by many species of cavity-nesting wildlife. (Photograph by
constant compared to the first scenario,55 although fires W.C. McComb)
may recur and delay forest development (Figure 5).
Fire return intervals in ponderosa pine (Pinus ponderosa)
forests are much shorter (every 5-15 years), but return
intervals increase with increasing elevation in eastside
forests.1 Fire intensity also often increases with elevation
in these eastside areas.
Wind may cause either coarse- or fine-scale
disturbances. Areas of high wind, such as the Pacific coast
and the Columbia River Gorge may produce large windfall
areas following the opening of a small part of a stand as
the stand progressively blows down, in a domino effect.
Elsewhere, wind disturbance usually consists of single
trees or small patches (<10 acres [<4 ha]) of trees that are
broken off or blown over. Rarely are all trees in a patch
blown over, and large amounts of dead wood remain after
the event (Figure 6). Figure 5. A partially burned stand leaves many live green
trees to grow after the disturbance as well as many dead trees
Fine-scale disturbances that provide habitat for cavity nesters. A two-story stand is
Fine-scale (<2 acres [<1 ha]) disturbances influence stand likely to develop after this disturbance. (Photograph by J.
dynamics in natural stands. Fine-scale disturbances such Tappeiner)
as suppression mortality, root rots (e.g., Phellinus spp. and
Armillaria spp.), localized windthrow, and light ground
fires can lead to the death of individual or small groups
of trees (Figure 7). Formation rates of these gaps are
relatively low in old-growth Douglas-fir stands56 (0.1-0.8%
of the stand/year) compared to other old-growth forest
types.51 However, these canopy gaps are the sites of tree
regeneration in mature stands,56, 58 particularly for shade
tolerant species. Considerable mortality and replacement
of dominant trees occurred during 36 years in 500-year-
old Douglas-fir/western hemlock forest. 16 Rates of
dominant tree mortality were 0.75%/year, but the
mortality was balanced by recruitment of shade-tolerant
tree species. Gaps may be slow to fill because >70% of
old-growth canopy trees die without disrupting the forest
floor.16, 56 In these cases, existing shade-tolerant shrubs
dominate gaps.
Figure 6. Windthrow leads to a dramatic increase in coarse
In eastern Oregon and Washington, fire control has led
woody debris in a stand and these structures are used by
to homogenization of large landscapes.1 Especially in
many species of mammals and amphibians in northwest
ponderosa pine forests, fire exclusion has caused many forests. (Photograph by J. Tappeiner)
 144 Wildlife–Habitat Relationships in Oregon and Washington

Mycorrhizal fungi aid plants in the uptake of water and

nutrients, and they can be particularly important to early
plant growth and survival on harsh sites.47 These fungi
produce fruits underground, unlike most other fungi, and
they do not, therefore, rely on aerial spore dispersal, as
do other fungi. They seem, instead, to be well adapted to
animal dispersal. Some fungi known as truffles are
important components of the diets of some small
mammals, particularly red-backed voles. These animals
eat fruits and ingest spores, which then pass through the
digestive system in a few days and are deposited at a new
site. A new fungal mat may then grow from this site and
ensure the presence and widespread distribution of
mycorrhizae in the soil.11 Mixing organic matter in the soil
Figure 7. A canopy gap provides a site for tree and shrub by these burrowing animals also is likely to influence
regeneration and can aid in development of vertical structure decomposition rates and soil function.
in the stand. (Photograph by J. Tappeiner) The activities of some herbivores can have tremendous
impact on habitat for other species.45 The activities of
stands to develop outside of the range of natural variability American beaver, for example, create early seral-stage
in terms of species composition and structure. Clearly fire riparian forest and pool habitat that can be important to
or some other disturbance will need to be reintroduced to other species. For example, Suzuki59 found very different
recover these stands. amphibian and mammal communities associated with
All of these scenarios have occurred in the region. The areas in the Oregon Coast Range that were impacted by
landscape patterns prior to timber management were a beavers compared to similar areas where beavers did not
combination of stands created by disturbances of various build dams. Other examples include black bears that kill
frequencies, intensities, and sizes. Since timber patches of trees in plantations, gophers that eliminate
management began in Pacific Northwest forests, the regeneration in patches, or elk herds that browse heavily
managed stands represent a much reduced range of next to riparian zones. All these activities create patchiness
variability in frequency, intensity, and size of disturbance or heterogeneity in affected sites, and such patches can be
from the natural disturbance regime. important resource areas for other species.

Plant-Animal Interactions Spatial and Temporal

Biotic interactions also can affect forest development and Scales of Habitat
consequently the structure and composition of habitat for Habitat typically is characterized by the physical setting
a variety of species. Certainly herbivory can play a (soils, aspect, elevation, and climate) and by the structure
significant role in directing successional pathways. Some and composition of vegetation occupying the site. One
plants have chemical and physical defenses against attribute of habitat is its size. A 2-acre (1-ha) patch of
herbivory.14 Plants also respond to animal herbivory by habitat may be more than sufficient for a California red-
altering growth rates. Among many grasses, forbs, and backed vole but clearly is inadequate for its predator, the
some shrubs, moderate levels of herbivory can actually northern spotted owl. Habitat is also dynamic. It is affected
stimulate growth above the levels of either undisturbed by the geomorphic setting that slowly changes, by the
or heavily grazed or browsed plants.2, 13 It is widely weather that changes daily, seasonally, yearly, and over
assumed that this compensatory growth occurs at the centuries, and by plant-community succession, which is
expense of reproduction and that herbivory, therefore, rapid in the early stages and slower in the later stages.
results in decreased seed production or smaller seed sizes. Vegetation management can alter the composition and
Herbivores affect forest systems in ways other than structure of habitat and the direction of plant succession.
consumption. They aid in the dissemination of seeds, and Forest managers, therefore, must think of habitat over a
they may help maintain site quality. Some plants are well range of spatial and temporal scales. One convenient way
adapted to dispersal on animals (for example, bedstraws of dealing with these scales is to visualize a hierarchy of
[Gallium spp.]). Other plant species (e.g., dogwoods small patches of habitat for a species that serve as building
[Cornus spp.] and cherries [Prunus spp.]) are well adapted blocks within the species’ home range, which in turn serve
to scarification that results from passing through animal as building blocks for populations (Figure 8). I discuss how
digestive systems and “direct-seeding” in a packet of habitat for a species might function at the stand level (a
fertilizer. Many fencerows, consequently, are dominated unit of forest land that will receive a silvicultural
by cherries, hawthorns (Crataegus spp.), and dogwoods treatment), and then how habitat functions in aggregates
because birds often perch on fences after eating the fruits of stands at the forest or landscape scales.
of these plant species.
 CHAPTER 4: MANAGING FOREST ELEMENTS 145

Figure 8. Adequate foraging and

cover patches must be present within
a species home range for it to be
occupied, and sufficient home ranges
must be available to support a
population.

Spatial Scales of Habitat Natural Levels of Variability in

Stands do not exist in isolation. Each stand is surrounded Managed Forests
by other stands, corridors of vegetation, water courses,
Estimating parameters characteristic of natural
and other features (the context for the stand). All these
disturbance regimes can facilitate prediction of forest
features comprise all or a portion of a landscape. A
recovery through the direction of succession and the
landscape typically is considered the mosaic of vegetation
subsequent development of vegetation structure.
and geomorphic features that occur across an area.15 The
elements of the landscape pattern that are important to Disturbance size and shape can influence the animal
some species of wildlife include the types of patches that species that either remain in or recolonize a disturbed
make up the landscape, the sizes of those patches, the area.50, 60 Organisms would likely be displaced by severe
length of surrounding edge, and the nature of corridors disturbances of a size larger than a home range, but they
or barriers between patches. Many of these elements may not be displaced if the disturbance is sufficiently small
interact to determine the suitability of habitat over the relative to the home range size. I suspect that species
landscape for various plant and animal species. Large inhabiting mature forests have been able to persist (1) by
carnivores use very large areas for foraging, necessitating including fine-scale disturbances within their home
a landscape perspective in habitat management.29 The total ranges, or (2) by recolonizing stands of sufficient size that
amount of habitat patches across a landscape, accordingly, regrow to maturity following coarse-scale disturbances.
strongly influences the occurrence or abundance of a Disturbance shape influences the amount of edge and hence
species on that landscape.37 Managers can manipulate microclimatic characteristics of the stand. The
stand structure and distribution across the landscape to characteristics of stand edges can have profound effects
influence the abundance and distribution of species within on a number of bird and mammal species.24, 42
a planning area.
Disturbance intensity influences the amount of organic
Species may be influenced by stand arrangement on
material destroyed and redistributed by the disturbance
the landscape.28 Species that require forest-interior (core)
and hence the amount and form (living or dead) of
patches may be adversely impacted by the proliferation
material that remains after the disturbance. These residual
of edges within the landscape that may result from
structures may directly or indirectly be habitat for mature
disturbance.34, 41 As more of the landscape is affected by
forest species. Gaps in mature and old-growth forests
disturbance, animals that move among remaining patches
produce the snags and logs in the stand, and enhance its
of habitat expend increasing amounts of energy, and
vertical complexity.40 The residual structures following
availability of forest-interior habitat decreases.17, 60 This
coarse-scale disturbances may persist into the next stand
process of habitat loss and fragmentation can be mediated
and provide the large trees and snags used by some
by managing stands within the landscape to meet the
species.38, 46, 53 The residual organic material that remains
needs of managed species.
after disturbance can influence the direction of succession
and the rate of subsequent development following the
disturbance.57
 146 Wildlife–Habitat Relationships in Oregon and Washington

Disturbance frequency will influence the tree species and noncommercial plant species than natural
composition and the amount of living and dead organic disturbances,23 so it may not imitate natural disturbances
material present on the site over time. Frequent coarse- for other forest resources as well as it does for timber.
scale disturbances can delay the onset of forest
development, or may preclude it. Infrequent fine-scale Habitat Characteristics Associated with
disturbance may delay the development of vertical
structure in a stand.56
Forest Wildlife
I assume that animals in forests use certain key structural
Disturbance density is the percentage of the stand or and compositional features over home range-sized areas
landscape occupied by a patch type resulting from a or larger. However, each species differs in its requirements
disturbance. The density of fine-scale disturbances also for the magnitude and condition of these structures. To
influences the abundance of structural features produced meet the needs of forest wildlife species, these habitat
by the disturbance. A stand in which 1% of the stand is in features should be considered as part of silvicultural
canopy gaps <30 years old will have a less well developed systems employed in stands over areas at least as large as
vertical structure and fewer hard snags than one in which the largest home range size among the species under
15% is in gaps. Finally, the disturbance pattern may also consideration multiplied by the desired population (Figure
influence habitat quality for some species. Clumped 8).
distributions of fine-scale disturbances may result in a
cumulative decrease in habitat availability within an Large Trees
individual’s home range. A more random or uniform Some trees within a stand should be designated as legacy
distribution of disturbances may allow that individual to trees and left to grow to maturity and die through natural
tolerate the same disturbance density because only a small processes. Douglas-firs >50 inches dbh (>125 cm) are used
portion of any individual’s home range in the stand would by marbled murrelets,53 red tree voles, and northern
be affected. spotted owls. Large trees add to the vertical structure
No single stand management system will precisely within old forests. Vertical structure may be associated
match the variability inherent in natural forests that with bird community structure in some forest types,30 but
resulted from a variety of disturbance regimes. But some it is unclear how important vertical structure is in western
of the variation can be incorporated into the managed coniferous forests. Large trees also add large surfaces of
forest landscapes of the region by using a variety of deeply-fissured or scaly bark that is used by bark-foraging
silvicultural systems. The choice of these systems will birds such as brown creepers32 and they support lichens,
depend on the biological, social, and economic objectives an important food source for northern flying squirrels.33
for the stand and the landscape, and they will imitate Designating Douglas-fir, western redcedar (Thuja plicata),
natural disturbances to varying degrees (Figure 9). Indeed, bigleaf maple (Acer macrophyllum), Pacific madrone
the basis for development of existing silvicultural systems (Arbutus menziesii), red alder (Alnus rubra) and western
for timber objectives was that these systems reflect the hemlock as legacy trees in coastal and Cascades forests
regeneration and growth strategies of the commercially would provide a range of growth rates and bark surfaces
important tree species in a region. Intensive timber and contribute to complexity in the stand. See chapter 24
management as currently practiced leaves less dead wood for information on guidelines for numbers of living trees
per acre (ha).
Some hardwoods, shrubs, and noncommercial conifers
Range of Managed should be allowed to grow to maturity and die within
acceptable stands these stands. Hardwoods often are considered competing
conditions vegetation in conifer stands. However, species such as
Functional
managed Oregon oak (Quercus garryanna), Pacific madrone, and
stands bigleaf maple (especially trees >20 inches [> 50 cm] dbh)
produce many natural cavities and dead limbs (effectively,
Frequency

elevated snags), that are used by cavity nesting animals.21,

48
Hardwoods and shrubs also provide forage, fruits, and
foliar insects that are food resources for many forest
vertebrates40 and hardwood composition in unmanaged
stands in western Oregon influenced the abundance and
occurrence of several bird species.25

Dead Wood
Habitat characteristic Species vary in their use of dead wood size and decay
Figure 9. Species require certain habitat features over a range classes.38 Although some quantitative information is
of sizes or amounts. The degree to which managed stands can available on snag use by some species, such as red-
provide these conditions will influence the proportion of the breasted sapsuckers, pileated woodpeckers, and hairy
landscape that can be used by each species of wildlife. woodpeckers,3, 46, 52 we have little quantitative information
for other species, especially log-users.38 Hence, setting
 CHAPTER 4: MANAGING FOREST ELEMENTS 147

stand goals for dead wood for species such as bats, shrew- legacy, plant Douglas-fir and western hemlock, seedling
moles, tailed frogs, and northwestern salamanders is release, and thin to 300 trees/acre (750 trees/ha). The result
difficult, but must be done to develop management would probably be a conifer-dominated stand condition
prescriptions. McComb and Lindenmayer38 offer one with a grass-forb understory (sparse shrubs) during the
approach to addressing this problem. early stages of development. Each condition is habitat for
a different suite of species on the same site managed in
Forest Floor Litter one of two ways. Many possible decisions could be made
Forest floor and below-ground conditions influence early in stand development that could produce a wide
habitat quality for ground-foraging and burrowing range of stand conditions (Figure 10).
species. Some terrestrial mammals and amphibians remain
active below-ground during the summer. For instance, Traditional Management Approaches
rough-skinned newts use logs and burrow systems of
voles and shrews as summer daytime refugia (W.C. Even-age Management
McComb and C.L. Chambers, Northern Arizona Clearcut and plant regeneration has been the system most
University, unpubl. data). Burrow systems of mountain frequently used in the westside forests of the Pacific
beavers and pocket gophers are used by other species.35 Northwest, especially on private lands, but there are other
Except for anecdotal descriptions, we have no quantitative options. Seed-tree and shelterwood systems are useful on
information that could be used to set goals for species certain sites and with certain tree species. The deferred
associated with forest floor litter and burrows (e.g., rotation system described by Smith et al.54 is an alternative
northwestern salamanders, Pacific shrews, and ensatina that benefits certain species of animals by leaving large
salamanders). trees to grow through >2 rotations. Whichever system is
chosen, the stand usually proceeds through site
Identifying Desired Future Conditions preparation, stand re-establishment, vegetation
Because habitat is dynamic, it is important to plan for a management, and stand-density management before it is
series of desired future conditions over space and through ready for harvest at the end of the rotation. Decisions at
time for a stand. The first step is to develop diameter each stage influence stand character and affect habitat
distributions (and associated levels of variance) for living quality for the wildlife species present at various stages
and dead wood from stands that are functioning for the of stand development (Figure 10).
species of wildlife that are desired. The goal of the
Regeneration System. The selection of a regeneration
silvicultural system would be to provide the structure,
system affects stand structure during the early stages of
composition, and as nearly as possible, the dead wood
stand development. Seed-tree and shelterwood systems
distribution in the managed stand to fall within the range
leave vertical structures (important to some birds and
of variability acceptable to the species. Models that can
arboreal mammals) that remain in the stand until the seed
help the manager understand the dynamics of trees and
trees and overwood are removed. Duration of the grass-
dead wood should be used when setting goals (see chapter
forb stand condition is usually shorter and shrub
24).
conditions in the stand are more predominant with these
systems than with traditional clearcutting, because of the
Effects of Silvicultural Treatments difficulties associated with vegetation management when
on Habitat residual trees and snags are present. Deferred rotation
Silvicultural activities have partially replaced natural systems offer the advantage of some vertical structure
disturbances in managed forests. Forest managers can remaining in the stand throughout the rotation. Even-age
select the types and rates of disturbances that will meet management, however, often results in the creation of
specific resource objectives. Some habitat management sharp, induced edges (depending on the stature of adjacent
issues that foresters and wildlife biologists face in the stands). These high contrast edges are beneficial to some
Pacific Northwest may result from insufficient species,41 but not to others, especially some species of
consideration of the size, frequency, intensity and amphibians. 34 Plant species associated with these
patterning of silvicultural disturbances on the landscape. regeneration systems are often shade intolerant, although
A range of management decisions can be made on any this can be adjusted with shelterwood systems and
given site that will result in stand conditions and plant artificial regeneration. Partial-cut and clearcut stands
communities that support only certain species (Figure 10). function similarly when <10 trees/acre (<25/ha) are
Consider, for example, an Oregon Coast Range site retained in partial cut stands.7 Nonetheless, partial cut
managed with the following combination of decisions: stands seem to provide habitat for more species of birds
clearcut, no legacy (retention of logs, snags or green trees), than clearcut stands, at least in the breeding season.7, 61
no site preparation, rely only on natural regeneration, no More species were detected in uncut or patch-cut stands
vegetation management, and no precommercial thinning. than in clearcut or partial cut stands during the winter,
The result would probably be a hardwood-shrub plant- however.6
community of small stature. Now consider the same site Legacy. At harvest and before site preparation, the land
managed with the following decisions: clearcut, snag manager may decide to leave certain structural
 148 Wildlife–Habitat Relationships in Oregon and Washington

components of the previous stand on the site and into the Stand re-establishment. Land managers can determine
next rotation. Snags are the most visible type of legacy the composition of developing stands by deciding which
left or created on many sites and they can significantly plant species will be re-established after site preparation.
influence occurrence of a number of species.4, 5, 38, 52 Logs With natural advance regeneration, that decision may be
and living conifers and hardwoods (and the lichens, unnecessary. Planting seedlings, however, can greatly
bryophytes, and other less vagile species associated with affect the structure and composition of stands. A stand
them) left on the site can provide structural and that includes several tree and shrub species, with different
compositional features that create conditions in the new growth rates, will be more diverse than a stand planted
stand more typical of those found after natural with a single tree species. Density and spacing of the
disturbances. Animal communities associated with stands regeneration also could be affected. The size distribution
that include these features would be more complex than of trees in the stand can be narrow (uniform spacing) or
communities in stands that lack similar components. broad (irregular spacing) in an even-aged stand. Mixed-
species planting or variable-density planting can provide
Site preparation. Site preparation ranges from very
heterogeneity in an otherwise homogenous system.
intense (mechanical scarification) to none. Mechanical
scarification may significantly affect the below ground Vegetation Management. Management of competing
structure of the stand by temporarily removing burrows vegetation can significantly affect the availability of certain
and compacting soils. It also may affect plant communities plant species as food and cover. Herbicide applications
that develop after the disturbance. Intense scarification, that release conifers can temporarily decrease the
burning, or some herbicides may reduce shrub availability of shrubs for shrub-nesting birds.43 For small
development in subsequent stages of stand development. mammals and amphibians, site preparation seems to have
Light fires may proliferate the sprouting of shrubs and a greater effect on animal abundance than spraying of
reduce the presence of grasses and forbs in an early stand. glyphosate.9, 10 Either chemical or manual vegetation
The choice of site preparation influences not only the control can influence the heterogeneity of developing
trajectory of the plant community that develops on the stands. Spot control of competing vegetation could lead
site, but also the level of residual “legacy” that remains to a more heterogeneous stand than possible with
after the treatment. An intense burn or mechanical broadcast application of a treatment. Manual control of
scarification, for example, reduces levels of dead wood many shrub species can lead to a proliferation of sprouts
on the site. This may be a desirable method for that increase amounts of available browse.
manipulating the habitat of some species that may Perhaps the most profound effect of vegetation
influence regeneration success (such as mountain beaver), management is the influence that such activities have on
but it also may have adverse consequences for other the composition and structure of developing stands. Lack
species. Nonetheless, species such as Oregon voles and of any vegetation management in many Coast Range sites
vagrant shrews increase after intense site preparation, but may lead to stands with a large component of shrubs or
others such as Pacific and Trowbridge’s shrews, ensatina hardwoods that will benefit some species of wildlife.25
salamanders, and Pacific giant salamanders decrease after Intensive vegetation management may lead to a conifer-
site preparation. 9, 10 dominated stand with little shrub development unless
stand density is manipulated as the stand develops.

Site
preparation Regeneration Vegetation
Legacy artificial management
burn
Regeneration leave tree species manual
season
system conifer fuels density season
Figure 10. A wide range of herbicide
Thinning
clearcut hardwood brown & burn size
vegetative conditions can be seedtree shrubs herbicide survival season density
created on any one site depending shelterwood snags season natural chemical spacing
rate uniform
on the management choices made deferred rotation logs scalp
group selection none variable
early in stand development. none none species
single-tree
selection crown class
high grade none
Define the site
Choices made from each list will
size
shape
combine to describe a stand capable
plant association of supporting a certain assemblage
elevation of species
natural disturbances
fire
wind
insects
disease
 CHAPTER 4: MANAGING FOREST ELEMENTS 149

Intermediate Treatments. Precommercial and commercial intensity all affect the structure and composition of
thinning influences the structure and composition of the uneven-aged stands.
understory and may influence, consequently, the vertical The choice of which silvicultural system to use should
complexity and quantity of browse in a stand. The plant be determined by the plant community, site conditions,
composition of the stand is unlikely to be influenced to logging constraints, and species. Activities in even-aged
any great degree at this point in stand development. stands that could enhance conditions for species typical
Opening the crowns by thinning also influences habitat of late seral stages include:
quality for those species that find cover and food (cones) • allocation of dead and living trees as structural legacy
in tree crowns. Also, those species that feed among tree after harvest,
crowns, such as Hammond’s flycatchers may benefit from
• variable-density planting and thinning, and
thinning.22 Variable-intensity thinning can produce a wide
range of tree diameters and greatly influence the • harvesting techniques that minimize soil disturbance.
production of small snags early in stand development. It Uneven-age management strategies that could enhance
also can influence the distribution of food and cover in habitat quality for many species that inhabit late seral-
young, pole-stage stands. Small openings <0.2 acre (<(0.1 stage conditions include:
ha) in second-growth stands of spruce-hemlock (Picea- • establishing a large target tree size,
Tsuga) in southeast Alaska allowed access to food close to • lengthening cutting cycles,
cover from snow for Sitka black-tailed deer and allowed
• minimizing disturbance to the stand during logging
certain bird species to occur in otherwise unsuitable
with designated skid trails,
habitat.12
• harvesting with small-group or single-tree selection
Uneven-age Management systems where they are appropriate,
Uneven-age management usually involves group selection • managing for shade-tolerant tree species,
or individual tree selection. These regeneration systems
• maintaining high-density groups of regeneration, and
cause a fine-scale disturbance, so stand-level vertical
structure is usually high, edge and fragmentation effects • allocate dead or large, living trees.
are usually low, and stand heterogeneity is usually high
compared to even-aged systems. The decision to leave a Non-traditional Management
legacy of dead wood depends on the objectives of the land Approaches
manager; retention of green trees normally is part of the The following systems represent management approaches
silvicultural system, but the allocation of large reserve trees that differ from those traditionally used for timber
in the stand is also feasible. management. They are only a few of the possible strategies
Site preparation is usually minimal in these systems, to produce structural characteristics typical of those
because they usually rely on natural advance regeneration. following natural disturbances within western Oregon
Artificial regeneration is feasible, however, and especially and Washington.40 The first system, modified single-
within the group selection system. Chemical or mechanical storied stands, represents a slight modification of current
site preparation, therefore, may help ensure establishment stand management practices that are designed to produce
and survival of tree seedlings in these harvest groups. relatively uniform stands for timber production using the
Mechanical scarification of the harvest groups and clearcut regeneration system. Rather than complete
compaction of the soils along designated skid trails can removal of all trees from the unit, 3-10 live trees/ acre (8-
significantly affect below-ground habitat by reducing the 25/ha) and large dead wood are left on the unit to provide
availability of burrow systems and restricting the ability some of the organic carry-over important to organisms
of animals to burrow in the surface soil. Careful harvest that use these features in early seral stages (Figure 11).
planning and use of designated skid trails is essential.27 The second system, few-storied stands, is the creation of
Artificial regeneration in harvest patches with a group stands with 2 or 3 layers of canopy trees as well as retention
selection system provides a significant opportunity to of large dead wood (Figure 12). The third option, many-
influence the composition and structure of the stand. More storied stands, uses small group selection cutting to create
shade-intolerant species can be regenerated with group a stand that is composed of >3 layers of canopy trees in a
selection than with single-tree selection. Small patch, mosaic of gaps (Figure 13). Again, large legacy trees and
group-selection systems or single-tree selection systems snags are maintained in the stand. Each of these systems
that rely on existing advance-regeneration or large represents elements found in natural disturbance regimes,
planting stock, however, may eliminate grass-forb stand and each is applicable to many forest types throughout
conditions and reduce the length of time that a stand the Pacific Northwest. The first two systems imitate coarse-
remains in a shrub stage. Indeed, Chambers et al.7 reported scale disturbances. Stands that develop from these systems
that many bird species found in clearcut and green-tree may be recolonized by species associated with mature
retention stands did not occur in stands managed using forests as the stand develops, but would not provide
small group selection. Many of the species using the group adequate habitat for mature forest species until the
selection system were also found in uncut mature forest regeneration develops to a sufficient size. Disturbance
stands.6, 8 Cutting cycle length, target tree size, and thinning associated with the single-story system is more intense
 150 Wildlife–Habitat Relationships in Oregon and Washington

Figure 11. A managed stand with a legacy of living and dead Figure 13. An uneven-aged stand managed by group selection
trees retained to provide habitat for species that are adapted to cutting. Species typical of mature forest conditions continue
post-fire conditions. (Photograph by W.C. McComb) to use this stand despite repeated entries. (Photograph by
W.C. McComb)

and leaves fewer large living and dead trees than most
natural disturbances, except when fires have occurred
within a stand at high frequency. This system represents
the low end of within-stand natural variation in patch size,
canopy cover, and large live tree survival. The few-storied
system also has elements of coarse-scale disturbances, but
produces greater variability in vertical structure and tree
diameter classes than the single-story system. Both
systems may be smaller in size and more frequent in
occurrence than some naturally occurring coarse-scale
disturbances on the landscape. Long rotations and large
harvest units would more nearly imitate natural
Figure 12. This two-story stand can provide habitat for disturbances in western Oregon and Washington.
species adapted to stands that develop after partial burns. The many-storied system is patterned after fine-scale
(Photograph by W.C. McComb) natural disturbances. Cut gaps may have to be larger than
most natural canopy gaps to allow natural regeneration
of shade intolerant species and to make harvesting more
efficient. On gentle terrain, harvesting costs are not
significantly higher than for clearcutting.27 This system
would have high within-stand variability in tree size and
vertical complexity. This system might provide acceptable
habitat for mature forest species while allowing some
small but regular timber removal.
 CHAPTER 4: MANAGING FOREST ELEMENTS 151

Natural disturbances occur in a range of sizes, shapes, habitat for some mature-forest species and provide higher
frequencies, intensities, and patterns across landscapes. connectivity among mature forest patches than young
These parameters can be varied within both coarse- and plantations. Modified single-storied and few-storied
fine-scale silvicultural systems by varying managed stand stands could be used on steep side slopes where harvesting
stand sizes and shapes, the frequency of entry, the levels costs for uneven-aged management may be prohibitive.27
of residual living and dead wood, and the arrangement The few-storied stands may be most appropriate on steep
of stands on the landscape, respectively. Diversity can be slopes adjacent to riparian buffer strips because they
retained within landscapes by adopting a broader range would provide at least a low contiguous canopy cover
of silvicultural systems and applying them to reflect the and a source of large logs for the riparian system. The
variety in size, intensity, and frequency of the natural resulting landscape would be a mosaic of many-storied
disturbance regimes. stands interspersed with single- and few-storied stands
Density management during stand development might in various seral stages. To be effective, managers would
enhance structural complexity in these stands as well as have to be committed to seeing the landscape designed to
in existing plantations. Typically a stand is meet the needs of various wildlife species over time. This
precommercially thinned and then may receive 1 or 2 landscape design may not meet management objectives
commercial thins prior to harvest. Usually thinning leaves for other resources. Considerations for aesthetics, water
the remaining trees uniformly spaced with adequate room and fisheries resources should be included. Although early
among crowns for continued volume growth.22 Thinnings seral stage habitats will be created during initial stages of
that vary in intensity within the stand provide the development following single-storied and few-storied
opportunity for rapid growth by some trees in the stand system harvests, species associated with young stands
and reduced growth or death (snags) by others within the may not be as abundant as with traditional even-aged
stand. These patterns imposed over several thinnings can management systems.
result in a range of tree sizes within the stand. If some of Regardless of the landscape design, there would be
these trees were designated at final harvest for retention continual (though low) volume removal until the areas
into the next rotation, and variable density planting and came into a fully managed condition, and then there
thinning were used, then an even more diverse forest would be an increase in timber production after this time.20
structure could result and the stand would become two- Volume removals would be less than from stands managed
storied. These stands might provide habitat for most solely for timber production. The level of volume
species typical of mature forest within 80-120 years. reduction will depend on the commitment made by the
forest managers to providing habitat for selected species.
Landscapes by Design
Currently we are inheriting landscapes created by past Summary
disturbances and timber-driven management objectives Habitat is a species-specific concept. Habitat represents
on mixed ownerships that do not consider large-scale the resources necessary to support a population over space
habitat patterning. Combining different silvicultural and through time. Habitat manipulation represents the
systems on the landscape in a manner that considers size best opportunity for long-term management of
and connectedness of mature forest habitat over large areas populations. Within forests, key structures such as large
through time would be one step toward designing forest trees, species composition of trees and shrubs, dead wood,
patterns. The silvicultural systems proposed above should and forest litter layers should be given particular attention
be considered only examples; they will not apply in all when designing stand prescriptions. Each species has its
situations nor will they meet all wildlife objectives. own habitat requirements and these should be estimated
Silviculturists will have to work with wildlife biologists, as well as possible when developing goals for stand
forest ecologists, harvesting specialists, and other resource prescriptions.
managers to: Structure and composition of a developing stand can
1. identify clear objectives for habitat (for 1 or many be dramatically influenced by a wide range of
species) and timber, management decisions. If managers wish to provide
2. identify existing examples of conditions that meet habitat for selected species, decisions should be made
the objectives, based on the potential of the site to develop the magnitude
3. design prescriptions specific to local conditions that and composition of required habitat features over time. It
represent desired future conditions to meet the objectives, is important to recognize that even currently common
and species should be considered during management to
4. plan the location of the stands on the landscape to ensure that both common and rare species continue to be
create landscape patterns that meet the objectives.7 represented across managed landscapes over time.
As an example, consider how several of the approaches Further, these habitat features should be distributed over
described above might be distributed on a landscape to space so that the species’ ability to use resources
meet a management objective of maintaining mature forest throughout its home range is considered. It is this planning
species while allowing some timber removal. Areas known over space and time at biologically meaningful scales that
to support mature-forest associated species could be will be required to meet the needs for wildlife in managed
managed using the many-storied management to provide forests.
 152 Wildlife–Habitat Relationships in Oregon and Washington

Literature Cited 22. Hagar, J., W. C. McComb and W. H. Emmingham. 1996. Bird
communities in commercially thinned and unthinned Douglas-fir stands
1. Agee, J. 1999. Disturbance effects on landscape fragmentation in
of western Oregon. Wildlife Society Bulletin 24:353-366.
Interior West Forests. Chapter 3 in Forest Fragmentation: Wildlife and
23. Hansen, A. J., T. A. Spies, F. J. Swanson and J. L. Ohmann. 1991. Lessons
Management Implications. J. P. Rochelle, L. A. Lehman, and J. Wisniewski,
from natural forests. BioScience 41:382-392.
editors. Forest fragmentation: Wildlife and Management Implications.
24. Hemstrom, M. A., and J. F. Franklin. 1982. Fire and other disturbances
Brill Press, Netherlands.
of the forests in Mount Rainier National Park. Quaternary Research
2. Belsky, J. 1986. Does herbivory benefit plants? A review of the evidence.
18:32-51.
American Naturalist 127: 870-892.
25. Huff, M. H., and C. M. Raley. 1991. Regional patterns of diurnal
3. Brown, E. R., technical coordinator. 1985. Management of wildlife and
breeding bird communities in Oregon and Washington, pages 177-206
fish habitats in western Oregon and Washington. Publication R6-
in L.F. Ruggiero, K. B. Aubrey, A. B. Carey, and M. H. Huff, editors. Wildlife
F&WL-192-1985. U.S. Forest Service. 2 volumes.
and vegetation of unmanaged Douglas-fir forests. U.S. Forest Service
4. Bull, E. L., and A. D. Partridge. 1986. Methods of killing trees for use by
General Technical Report 285.
cavity-nesters. Wildlife Society Bulletin 14:142-146.
26. Keith, L., and L. A. Windberg. 1978. A demographic analysis of the
5. Chambers, C. L., T. Carrigan, T. Sabin, J. Tappeiner, and W. C. McComb.
snowshoe hare cycle. Wildlife Monographs 58.
1997. Use of artificially created Douglas-fir snags by cavity-nesting
27. Kellogg, L. D., P. Bettinger, and R. M Edwards. 1996. A comparison of
birds. Western Journal of Applied Forestry 12(3):93-97.
logging planning, felling, and skyline yarding between clearcutting and
6. Chambers, C. L., and W. C. McComb. 1997. Effects of silvicultural
five group-selection harvesting methods. Western Journal of Applied
treatments on wintering bird communities in the Oregon Coast Range.
Forestry 11:90-96.
Northwest Science 71(4):298-304.
28. Leopold, A.S. 1933. Game management. New York: Charles Scribner’s
7. Chambers, C. L., W. C. McComb, J. C. Tappeiner II, L. D. Kellogg, R. L.
Sons, New York, NY.
Johnson, and G. Spycher. 1999. CFIRP: What we learned in the first ten
29. Lindstedt, S. L., B. J. Miller, and S. W. Buskirk. 1986. Home range, time,
years. The Forestry Chronicle 74:431-434.
and body size in mammals. Ecology 67:413-418.
8. Chambers, C. L., W. C. McComb, and J. C. Tappeiner. 1999. Breeding bird
30. MacArthur, R. H., and J. W. MacArthur. 1961. On bird species diversity.
responses to three silvicultural treatments in the Oregon Coast Range.
Ecology 42:594-598.
Ecological Applications 9(1):171-185.
31. Mannan, R. W., M. L. Morrison, and E. C. Meslow. 1984. Comment: the
9. Cole, E. C., W.C. McComb, M. Newton, C. L. Chambers, and J. P.
use of guilds in forest bird management. Wildlife Society Bulletin
Leeming. 1997. Response of amphibians to clearcutting, burning, and
12:426-430.
glyphosate application in the Oregon Coast Range. Journal of Wildlife
32. Mariani, J. M. 1987. Brown creeper (Certhia americana) abundance
Management 61:656-664.
patterns and habitat use in the southern Washington Cascades. Thesis,
10. Cole, E. C., W. C. McComb, M. Newton, J. P. Leeming, and C. L.
University of Washington, Seattle, WA.
Chambers. 1998. Response of small mammals to clearcutting, burning,
33. Martin, K. J. 1994. Movements and habitat associations of northern
and glyphosate application in the Oregon Coast Range. Journal of
flying squirrels in the central Oregon Cascades. Thesis, Oregon State
Wildlife Management 62:1207-1216.
University, Corvallis, OR.
11. Cork, S. J. and G. J. Kenagy. 1989. Rates of gut passage and retention of
34. Martin, K. J. 1999. Habitat associations of small mammals and
hypogeous fungal spores in two forest-dwelling rodents. Journal of
amphibians in the central Oregon Coast Range. Dissertation, Oregon
Mammalogy 70:512-519.
State University, Corvallis, OR. .
12. DellaSala, D. A., J. C. Hagar, K. A. Engel, W. C. McComb, R. L. Fairbanks,
35. Maser, C., B. R. Mate, J. F. Franklin, and C. T. Dyrness. 1981. Natural
and E. G. Campbell. 1996. Effects of silvicultural modifications of
history of Oregon Coast mammals. U.S. Forest Service General
temperate rainforest on breeding and wintering bird communities,
Technical Report PNW-133.
Prince of Wales Island, Southeast Alaska. Condor 98:706-721.
36. Mautz, W.W. 1978. Nutrition and carrying capacity. Pages 321-348 in J.
13. du Toit, J. T., J. P. Bryant, and K. Frisby. 1990. Regrowth and palatability
L. Schmidt and D. L. Gilbert, D.L., editors. Big game of North America.
of acacia shoots following pruning by African savanna browsers.
Stackpole Books, Harrisburg, PE.
Ecology 71:149-154.
37. McComb, W. C. 1999. Forest fragmentation: wildlife and management
14. Farentinos, R. C., P. J. Capretta, R. E. Kapner, and V. M. Littlefield. 1981.
implications – synthesis of the conference. Chapter 17 in J. P. Rochelle,
Selective herbivory in tassel-eared squirrels: role of monoterpenes in
L. A. Lehman, and J. Wisniewski, editors. Forest fragmentation: Wildlife
ponderosa pines chosen as feeding trees. Science 213:1273-1275.
and Management Implications. Brill Press, Netherlands.
15. Forman, R. T.T. and M. Godron. 1986. Landscape ecology. New York:
38. McComb, W. C., and D. Lindenmayer. 1999. Dying, Dead, and Down
John Wiley & Sons, New York, NY.
Trees, Chapter 10, in Hunter, M. L., Jr., Maintaining Biodiversity in Forest
16. Franklin, J. F. and D. S. DeBell. 1988. Thirty-six years of tree population
Ecosystems. Cambridge University Press, Cambridge, England.
change in an old-growth Pseudotsuga-Tsuga forest. Canadian Journal of
39. McComb, W. C., and R. E. Noble. 1981. Nest box and natural cavity
Forest Research 18:633-639.
use in three mid-South forest habitats. Journal of Wildlife Management
17. Franklin, J. F. and R. T. T. Forman. 1987. Creating landscape patterns by
45:92-101.
forest cutting: ecological consequences and principles. Landscape
40. McComb, W. C., T. A. Spies, and W. H. Emmingham. 1993. Stand
Ecology 1:3-18.
management for timber and mature-forest wildlife in Douglas-fir
18. Fretwell, S. D., and H. L. Lucas, Jr. 1969. On territorial behavior and
forests. Journal of Forestry 91(12):31-42.
other factors influencing habitat distribution in birds. I. Theoretical
41. McGarigal, K., and W. C. McComb. 1995. Relationships between
development. Acta Biotheoroligica 19:16-36.
landscape structure and breeding birds in the Oregon Coast Range.
19. Friesen, C. A. 1991. The effect of broadcast burning on the quality of
Ecological Monographs 65:235-260.
winter forage for elk, western Oregon. Thesis, Oregon State University,
42. McGarigal, K., and W. C. McComb. 1999. Forest fragmentation and
Corvallis, OR.
breeding bird communities in the Oregon Coast Range. Chapter 12 in J.
20. Gagliuso, R. and W. C. McComb. 1992. Designing landscapes for
P. Rochelle, L. A. Lehman, and J. Wisniewski, editors. Forest
wildlife and timber, pages 379-388 in G. Wood and B. Turner, editors.
fragmentation: Wildlife and Management Implications. Brill Press,
Integrating forest information over space and time. ANUTECH Pty Ltd,
Netherlands.
Canberra, Australia.
43. Morrison, M. L., and E. C. Meslow. 1984. Effects of the herbicide
21. Gumtow-Farrior, D. L. 1991. Cavity resources in Oregon white oak
glyphosate on bird community structure, western Oregon. Forest
and Douglas-fir stands in the mid-Willamette valley, Oregon. Thesis,
Science 30:95-106.
Oregon State University, Corvallis, OR.
 CHAPTER 4: MANAGING FOREST ELEMENTS 153

44. Morrison, P. H. and F. J. Swanson. 1990. Fire history and pattern in a

Cascade Range landscape. U.S. Forest Service General Technical Report
PNW-GTR-254.
45. Naiman, R. J. 1988. Animal influences on ecosystem dynamics.
BioScience. 38(11):750-752.
46. Nelson, S. K. 1989. Habitat use and densities of cavity-nesting birds in
the Oregon Coast Range. Thesis, Oregon State University, Corvallis,
OR.
47. Perry, D. A., M. P. Amaranthus, J. G. Borchers, [and others]. 1989.
Bootstrapping in ecosystems. BioScience. 39:230-237.
48. Raphael, M. G. 1987. Use of Pacific madrone by cavity-nesting birds.
Pages 198-202 in Symposium on multiple-use management of
California’s hardwood resources. U.S. Forest Service General Technical
Report PSW-100.
49. Robbins, C. T., S. Mole, A. E. Hagerman, and T. A. Hanley. 1987. Role of
tannins in defending plants against ruminants: Reduction in dry matter
digestion? Ecology 68:1606-1615.
50. Rosenberg, K.V., and M. G. Raphael. 1986. Effects of forest
fragmentation on vertebrates in Douglas-fir forests. Pages 263-272 in J.
Verner, M. L. Morrison, and C. J. Ralph, editors. Wildlife 2000: Modeling
habitat relationships of terrestrial vertebrates. The University of
Wisconsin Press, Madison, WI.
51. Runkle, J. R. 1985. Disturbance regimes in temperate forests. Pages
17-34 in S. T. A. Pickett and P. S. White, editors. The ecology of natural
disturbance and patch dynamics. Academic Press, New York, NY.
52. Schreiber, B. 1988. Diurnal bird use of snags on clearcuts in central
coastal Oregon. Thesis. Oregon State University, Corvallis, OR.
53. Singer, S. W., N. L. Naslund, S. A. Singer, and C. J. Ralph. 1991. Discovery
and observations of two tree nests of the marbled murrelet. Condor
93:330-339.
54. Smith, H. C., N. I. Lamson, and G. W. Miller. 1986. An esthetic
alternative to clearcutting. Journal of Forestry 87:14-18.
55. Spies, T. A., and J. F. Franklin. 1988. Old-growth and forest dynamics in
the Douglas-fir region of western Oregon and Washington. Natural
Areas Journal 8:190-201.
56. Spies T. A., J. F. Franklin, and M. Klopsch. 1990. Canopy gaps in Douglas-
fir forests of the Cascade Mountains. Canadian Journal of Forest
Research 20:649-658.
57. Spies, T. A., J. F. Franklin, and T. B. Thomas. 1988. Coarse woody debris
in Douglas-fir forests of western Oregon and Washington. Ecology
69:1689-1702.
58. Stewart, G. H. 1986. Forest development in canopy openings in old-
growth Pseudotsuga forests of the western Oregon Cascade Range,
Oregon. Canadian Journal of Forest Research 16:558-568.
59. Suzuki, N. 1992. Habitat classification and characteristics of small
mammal and amphibian communities in beaver-pond habitats of the
Oregon Coast Range. Thesis, Oregon State University, Corvallis, OR.
60. Temple, S. A. 1986. Predicting impacts of habitat fragmentation on
forest birds: a comparison of two models. Pages 301-303 in J.Verner, M.
L. Morrison, and C. J. Ralph, editors. Wildlife 2000: modeling habitat
relationships of terrestrial vertebrates. University of Wisconsin Press,
Madison,WI.
61. Vega, R. M. S. 1993. Bird communities in managed conifer stands in the
Oregon Cascades: habitat associations and nest predation. Thesis,
Oregon State University, Corvallis, OR.
62. With, K. 1999. Is landscape Connectivity necessary for wildlife
management? Chapter 7 in J. P. Rochelle, L. A. Lehman, and J. Wisniewski,
editors. Forest fragmentation: Wildlife and Management Implications.
Brill Press, Netherlands.
 5
Conservation of Biodiversity:
Considerations and Methods for Identifying
and Prioritizing Areas and Habitats
Margaret M. Shaughnessy & Thomas A. O’Neil

Introduction
Conservationists have advocated two approaches to developing methods to identify and conserve areas that
counter the accelerating loss of biodiversity. One operates will complement that existing conservation network of
in a crisis mode, rescuing species in danger of extinction. private and public lands; and (3) developing easily
The other focuses on protecting communities of plants and accessible information sources that can be used as tools
animals not yet in serious jeopardy, but which are likely by natural resource managers and the public to identify
to be driven towards extinction with increased habitat loss. priority habitats, conserve native species, and increase
These two complementary approaches have been awareness and understanding of (a) habitats at risk; (b)
described as “fine filter” and “coarse filter” strategies for function, distribution, and abundance of habitats; and (c)
maintaining biodiversity.26 Implementation of the “fine effects of land management activities.4, 7, 48
filter” approach is exemplified in actions taken in support This chapter will review the methods currently used
of the Endangered Species Act. In implementing the to prioritize terrestrial habitats and areas, focusing on the
“coarse filter” approach, conservationists and natural data sets used, important considerations and inclusions,
resource organizations often include wildlife-habitat and qualitative and quantitative aspects of each. In
associations in their assessment of biological diversity.37, addition, several statewide strategies for identifying and
43
In doing so, these organizations often look to conserve prioritizing conservation areas are summarized to provide
vegetation types in hopes that they will also protect the examples of strategies for addressing this complex issue.
plant, invertebrate, and vertebrate species associated with Finally, total and protected acreages for each of the 32
them. This premise implies that vegetation serves as a habitats found in Oregon and Washington are provided.
satisfactory indicator of the environmental variables that
interact on a particular site41, 43 and as such, assumes that
plant communities can serve as substitutes for ecosystems Methods for Identifying and
and the elements of biodiversity.27 Given the complex ways Prioritizing Sites
in which species interact with their ecosystem, the Methods for identifying and prioritizing areas important
combination of proactive fine and coarse filter strategies to conservation values range from subjective techniques
present a greater hope for conserving biodiversity.37 based on the knowledge of experts to more quantitative
Until recently, many conservation strategies have methodologies. Generally, to achieve the desired
focused on identifying areas that will function as reserves, conservation goal, most processes rely on both subjective
or areas that could be set aside to conserve some aspect of and quantitative techniques, with the latter often used as
biodiversity. However, reserves are vulnerable to a tool to aid in the decision-making process. 4, 7 An
evolutionary processes, catastrophic events, and global advantage of using quantitative techniques is that the
warming. In addition, no area or group of areas likely will methods and data sets can be made available to
be sufficient to support the full range of native species individuals and organizations who, in turn, can learn how
and ecological processes that contribute to biodiversity at decisions were made and gain understanding about how
ecoregional, state, and continental scales.14, 35 Therefore, their values and land-management decisions affect large-
the identification and establishment of reserves represents scale conservation decisions.4, 52 Important components of
a relatively short-term solution, and the focus of quantitative techniques include well-defined goals, an
conservation efforts has expanded to include managing appropriate spatial scale for the analyses, analyses
multiple-use public and private lands for biodiversity.27 conducted at several scales (e.g., local, ecoregion, and
Several recent conservation efforts have demonstrated this state), an understanding of the limitations of the maps
broader focus by (1) identifying and mapping locations that are used (e.g., resolution, accuracy, and habitats and
of priority habitats, species, and areas; (2) identifying lands habitat elements that may be excluded), appropriate units
currently managed for biodiversity values and then for the analyses, and indices of viability and threat. The

154
 CHAPTER 5: PRIORITY HABITATS 155

goals of the conservation program will determine the type Historic Vegetation Maps. Maps depicting the historic
and quality of the data required (e.g., resolution of distribution of vegetation are useful tools that allow the
vegetation map) and the spatial scale for the analysis.42, 52 user to gain understanding of the changes that have
occurred in the relative amounts and general distribution
Maps and Units of Analysis of vegetation types, and thereby help identify changes
Critical to most conservation efforts is our ability to (1) relevant to conservation efforts. 7 For example,
map habitats and species distributions, (2) identify the comparisons between maps of current and historic
habitat associations of our wildlife species, and (3) vegetation provide a context in which changes in
understand the changes that have occurred in the vegetation from native species to exotic vegetation types
landscape over time. Therefore, several types of maps and or those associated with human uses (e.g., urban and
digital information are useful to conservation efforts, agricultural land cover types) can be evaluated.7
including maps of vegetation, species distribution, and
historic vegetation. Units of Analysis. Conservation areas have been
identified and prioritized using various units of analysis
Vegetation Maps. Vegetation maps developed though including (1) sampling grids or hexagons of uniform size
interpretation of satellite imagery provide a good source that have been superimposed over the landscape;5, 16, 49, 52
of information about vegetation and land forms currently (2) mapped vegetation or habitat polygons; and (3) existing
existing on the landscape. Generally, these maps do not conservation reserves or areas known to be important,
adequately represent small features or linear features such rare, or limiting. All of these methods have benefits as well
as wetlands, riparian areas, and small areas of specific as limitations, and the goals of the project will help
vegetation types such as Romer’s fescue grasslands also determine which method is most appropriate. For
are not represented well.4, 6, 7, 37, 39 In addition, although they example, sampling grids represent a convenient way to
are important biological characteristics for predicting divide the landscape and are useful tools for large-scale
distributions of vertebrate and invertebrate species, many regional or national assessments because attributes such
fine-scale features such as structure, habitat elements, and as species presence or absence can be assigned easily to
presence of individual plant species generally are not each cell of the grid. However, because the cells or
included in maps developed from remotely sensed data.37, hexagons defined by the sampling grids represent
39
Nevertheless, maps developed from remotely sensed arbitrarily defined areas that bear no relationship to
data provide some of our best and most cohesive sources features on the ground and often encompass a wide range
of information with which to conduct analyses and of elevations and a variety of vegetation and habitat types,
identify conservation priorities. the cell may provide only a small piece of the habitat
needed by many of the species attributed to it.7 Therefore,
Species Distribution Maps. Maps of species distributions
the cell identified through an analysis may not contain
for plants, vertebrates, and invertebrates also are
the characteristics or considerations that are important or
important components of most fine and coarse filter
necessary to achieve the desired conservation goal.4 Field
strategies for conservation. For rare, threatened, and
surveys may be conducted to confirm that the cell contains
endangered species, species distribution maps generally
the desired biological characteristics; 6 however, the
can be developed from databases that track occurrences
biological characteristics of the cells not chosen through
of those species. For more common species, distributions
the analysis would remain unverified and therefore
can be modeled using information such as museum
unknown even though they may contain characteristics
records, verified sighting records, maps of current
that would be considered more desirable to the
vegetation, and information on habitat associations and
conservation goal.
elevation.6, 38 Accuracy of distribution maps is difficult to
Vegetation patches or polygons identified through
determine; however, the accuracy will be related to the
various mapping methodologies with remotely sensed
resolution, vegetation classification, and attributes of the
data also have been used as units of analysis for
vegetation map as well as the predictive capabilities of
identification of conservation priorities.4, 6, 7 An assumption
the habitat associations.
of these analyses is that the vegetation polygons represent
Because our knowledge of species and habitat
a single vegetation type, although the degree to which that
associations is limited, O’Neil et al.28 developed a method
is true is a function of the vegetation classifications, map
for grouping vegetation types into more generalized
accuracy, mapability, minimum mapping unit (mmu, the
habitat types based on use by similar groups of wildlife
smallest polygon or unit of classification), and scale. Until
species. The generalized habitat classifications allowed
recently, most statewide vegetation maps produced from
investigators to predict with greater certainty the
satellite imagery were created with a relatively large mmu
associations between species and habitats.28 However,
(e.g., 350 acres or 140 ha). However, advances in
databases that provide only general associations between
technology have enabled investigators to more easily
species and generalized habitat features provide weak
create maps with smaller mmu, making these vegetation
inferences for predicting the presence and distribution of
maps more useful for natural resource managers and local-
species,39 and may not be the best choice for the units of
scale conservation assessments. Other limitations of
biodiversity analyses since information about vegetation
vegetation maps were discussed previously.
type, structure, successional stage, and habitat elements
will be lacking.
 156 Wildlife–Habitat Relationships in Oregon and Washington

Quantitative methods to the general public. Therefore, some authors recommend

Generally, most quantitative methodologies for identifying the use of single criterion rankings that then can be used
and prioritizing conservation sites can be categorized into to subjectively evaluate and rank sites for conservation
three types, (1) species richness, (2) scoring techniques and value.45
indices, and (3) complementarity or iterative analyses. Complementarity or Iterative Analyses. A more recently
Species Richness. Species richness analyses are simple developed method for identifying and prioritizing areas
approaches used by many investigators to evaluate the involves an iterative process in which an algorithm is used
relative value of sites as wildlife habitat. In its most basic to identify sites with the largest number of species from a
form, species richness methodologies rank the value of a target group (e.g., threatened species, amphibians, species
site based on the total number of species that are predicted with restricted ranges) and then iteratively identifies
to occur in that area.5, 15, 16, 36, 45, 52 Several investigators have additional sites that contribute the most species not
expressed concern over the use of species richness already represented in the site selected previously.6, 16, 19, 22,
25, 31, 32
methodologies to identify or prioritize sites for The iterative process continues until a subset of sites
conservation values because areas of species richness for is identified in which all species of the target group are
a particular group (e.g., mammals, threatened and represented. An advantage of this approach is that, unlike
endangered species, plants) rarely overlap with areas of many scoring and ranking methodologies that simply
species richness for other groups.6, 20, 30, 36 However, other select the highest ranking sites, replicating many species
investigators have found that richness analyses for species and excluding other species,19 the iterative selection
with restricted ranges may identify areas that are more process takes into consideration the attributes of other
biologically diverse and include areas important to a sites, and then identifies and selects complementary
variety of taxa.12, 44 areas.32 As with other prioritization methodologies, it is
Additional concern about species richness important that these analyses include information on the
methodologies has arisen because often the analyses are total number of individuals of a species or an index of
based on predicted distributions of species or do not abundance to ensure that viable populations are identified
include an estimate or index of abundance for each and conserved.45
species.6, 16, 45, 52 Therefore, a site that contains only 1 Factors that should be considered for inclusion in any
individual of each of 30 species would have a ranking of the subjective or quantitative analyses include the
similar to a site with 100 individuals of each of 30 species. following:
Similarly, a site ranked as poor in terms of species richness 1. management needs of a habitat or area (e.g., fire in
may contain many individuals of each species. In addition, Oregon White Oak communities);
habitats of intermediate quality frequently are ranked as 2. abundance or total numbers of each species;
good in terms of species richness, but may contain many 3. important breeding habitat, seasonal range,
urban or non-native species.33 To address these concerns movement corridors;
and ensure that viable populations are identified and 4. limited abundance or distribution of habitats;
conserved, several investigators have included estimates 5. vulnerability to disease, habitat alteration,
of population density or indices of abundance into the proximity to threat, development, and land-use
analyses.45, 52 In addition, the use of species richness activities;
analyses now usually is limited to identification of areas 6. unique or dependent species;
important for specific species or groups of species rather 7. uniqueness of plant or wildlife community;
than being used for more general coarse-filter approaches 8. function of plant, invertebrate, or vertebrate
to identification and prioritization of conservation areas. species;
9. function of habitat or some component of habitat
Scoring Techniques and Indices. A widely used approach for plant, invertebrate, or vertebrate species;
to evaluate areas in terms of their contribution to 10. status of plant, vertebrate, and invertebrate
conservation goals involves scoring techniques and species;
indices that enable investigators to rank or prioritize areas 11. endemism or species with restricted ranges;
based on various selection criteria.1, 3, 4, 24, 51, 53 Areas may be 12. land ownership;
ranked for criteria such as species richness, species 13. protection status or management goals for site;
diversity, species rarity, species priority, endemism, total and
number of individuals of each species or an index of 14. species richness, species rarity, and species
abundance, naturalness, size, or economic and scientific priority.
value.4, 9, 23, 31, 45, 46, 53 Selection criteria often are weighted in
complex multi-criteria indices that are used to evaluate
and prioritize the potential conservation areas.3, 4, 24, 51 Current Approaches for Evaluating and
However, the relationships between criteria often are too Identifying Priority Areas and Habitats
complex to allow for the development of a meaningful At the local, state, regional, and national levels, the task
index,9, 21, 40, 46 and the factors contributing to a site’s of identifying and conserving priority areas and habitats
importance become hidden in a complex formula and are is daunting. Until recently, few states have had the data
not easily identifiable to the investigators themselves or sets needed to develop quantitative approaches to this
 CHAPTER 5: PRIORITY HABITATS 157

conservation challenge. However, the National Gap distribution of the species. Finally, the National Gap
Analysis Program has provided vision and start-up Analysis Program also relies on maps depicting land
funding for most states to begin development of these ownership and land management. These maps identify
basic data sets. With the completion of some of these data the major patterns of land ownership and management,
sets, several states have begun to develop approaches for aiding in the identification of lands managed for
identifying and conserving priority habitats and areas. In conservation values and in identification of “gaps” in the
addition to the nation-wide approach developed by the current network of conservation lands.
National Gap Analysis Program, three statewide
approaches are reviewed in this section. All three Washington: Priority Habitats and
approaches are quite different, reflecting differing goals,
confidence or availability of data sets, and available
Species Program
technology. Of the three statewide approaches, The Washington Department of Fish and Wildlife Habitat
Washington’s program was developed the earliest and Program (WDFW) has developed a Priority Habitats and
consequently has the simplest design. The approaches for Species list that identifies habitats and species considered
Oregon and Colorado were developed in 1997 and 1998, priorities for conservation and management.48 The WDFW
respectively. Both of theses approaches are complex, using program has a broad focus and includes information for
recently developed data sets and technology and terrestrial, aquatic, and marine species and habitats. The
incorporating quantitative analyses as well as subjective program identifies (1) priority habitats, (2) priority species,
evaluations. All three approaches take important steps to and (3) priority areas. Habitats, species, or areas may be
begin addressing conservation needs at local, ecoregion, considered priorities throughout the state or that
and statewide scales. designation may be restricted to specific geographic areas.
Biologists use the criteria described below to identify
priority habitats, species, and areas; these occurrences are
National Gap Analysis Program then mapped, and locational and descriptive data are
The National Gap Analysis Program is a nationwide recorded in a GIS. WDFW then develops management
program managed by the United States Geological Survey, recommendations for Washington’s priority habitats,
Biological Resources Division. The program was species, and areas. These documents can be obtained from
developed in the mid-1980s and uses a coarse-filter the WDFW Habitat Program Internet site at www.wa.gov/
approach to assess conservation needs at state, regional, wdfw/hab/phspage.htm. The Priority Habitats and
and national scales. The program focuses on working with Species program currently has identified approximately
each state in the United States and developing digital data 160 priority species and 20 priority habitats.48
layers that can be used with GIS to identify “gaps” or
natural land cover types and native vertebrate species not Priority Habitats. Priority habitats are defined as habitat
adequately represented in the existing network of types or elements with unique or significant value to a
conservation lands. Specific objectives of the National Gap diverse assemblage of species. Priority habitats may
Program include conducting statewide, regional, and consist of a unique vegetation type or dominant plant
national level analyses to: species, a specific successional stage, or specific habitat
1. Determine the area occupied by vertebrate species elements (e.g., talus, caves). For a habitat type or element
and land cover/vegetation types that are within to be considered a priority habitat, it must have at least
lands managed for conservation values. oneof the following characteristics:
2. Identify public lands having vertebrate species and 1. relatively high fish and wildlife density or species
land cover/vegetation types that are not well diversity;
represented in the existing conservation network. 2. important fish and wildlife breeding habitat,
3. Identify gaps in the natural reserve system of the seasonal range, or movement corridor;
United States. 3. rare or of limited availability;
4. high vulnerability to habitat alteration; or unique
The National Gap Analysis Program uses several data or dependent species.
layers to meet their program goals including maps
depicting (1) current vegetation and land cover in each Priority Species. Priority Species are defined as species
state, (2) distribution of terrestrial vertebrates, and (3) land that require protective measures for their perpetuation
ownership and management. For each state, vegetation is because of their (1) population status; (2) sensitivity to
mapped from satellite imagery to the alliance level habitat alteration; or (3) recreational, commercial, or tribal
(dominant/co-dominant cover types) using the National importance. Priority species include:
Vegetation Classification System.8 The mmu for each of 1. all state listed (threatened, endangered and
the statewide maps is <250 acres (100 ha). Using a variety sensitive) and candidate species;
of techniques and data sets, maps of vertebrate distribution 2. vulnerable aggregations: species or groups of
also are developed for each state. Because few states have species susceptible to population decline because
been systematically surveyed for all species, vertebrate of their tendency to aggregate (e.g., heron
distribution maps generally identify the predicted rookeries, sea bird concentrations, marine
distribution of each species rather than the known mammal haul outs, shellfish beds, fish spawning
and rearing areas); and
 158 Wildlife–Habitat Relationships in Oregon and Washington

3. native and non-native species with recreational, Habitat Institute and has a mmu of 10 acres (4 ha). Both
commercial, or tribal importance that are at risk maps have 62 vegetation and land cover types and were
due to habitat loss or degradation. developed by Killsgaard and Barrett.18
Priority Areas. Priority areas are defined as specific areas Historic Vegetation Map. This map was created by the
or locations that are a priority because they support OBP, and represents a blending of several regional data
relatively high numbers of individuals (e.g., heron sets including those developed by the Interior Columbia
rookeries, locations of rare species) or are important to River Basin Ecosystem Management Project, U.S. Forest
the life history and ecology of the species. Examples of Service vegetation maps, and General Land Office maps.
priority areas include the following: The historic vegetation map was not intended to identify
1. breeding, rearing, and hibernation sites; the vegetation types that existed at a specific place and
2. leks; time; rather it was developed as an indicator of the relative
3. areas commonly or traditionally used by amount and general distribution of vegetation, enabling
individuals of a species or a group of animals; investigators to identify historic vegetation patterns and
4. migration corridors; or land-cover changes that have occurred over the past
5. foraging areas. century. As with the map of current vegetation, the scale
of the historic vegetation map limits the detail that could
Oregon: be shown, under-representing or omitting several
vegetation and land-cover types.
The Oregon Biodiversity Project
Currently in Oregon, there is not a state-sponsored At-risk Species. At-risk species are defined as plant and
program to identify or prioritize habitats or areas animal (aquatic and terrestrial vertebrates and
important to conservation values. However, a nonprofit invertebrates) species considered highly vulnerable to
organization, The Defenders of Wildlife, has begun to try extinction. The OBP used the Natural Heritage Program’s
to address these concerns for Oregon.7 The Oregon database and ranking system to identify approximately
Biodiversity Project (OBP), a project overseen by 200 species considered to be at risk in Oregon. The
Defenders of Wildlife, recently completed an analysis of Heritage Program’s database incorporates information on
Oregon’s biodiversity and identified processes and factors such as known occurrences, threats, sensitivity, and
opportunities for conserving the State’s native species and area occupied, as well as species status at state, national,
habitats.7 Both statewide and ecoregional analyses were and global levels.
conducted, resulting in the identification of the network Land Ownership, Administration, and Management.
of conservation lands existing currently in Oregon and The OBP refined the existing data layer that identified
the development of a process for identifying conservation current land ownership and administration for Oregon.
needs and strategies for addressing those needs. In addition, the OBP developed a ranking system, the
Biodiversity Management Rating System, to identify the
Data Layers
degree to which lands were managed for biodiversity
The OBP relied on both quantitative and qualitative
values. Lands managed by state and federal agencies and
processes to meet its goals. Data layers and digital data
those managed by The Nature Conservancy were rated
sets used for both the statewide and ecoregional analyses
subjectively on a 10-point scale to indicate their
included the following:
contribution to long-term conservation of native
Statewide Vegetation Map. This map was developed in biodiversity. Land managers assessed and rated their lands
cooperation with the Oregon Gap Analysis Program13 and for the following criteria:
depicts the current vegetation and land cover of Oregon. 1.Management Objectives: is biodiversity
The map was created through visual interpretation of conservation or protection of natural values the
satellite photos (1:250,000 scale) taken in the late 1980s primary objective for the area?
and has a mmu of 350 acres (140 ha). One hundred thirty- 2. Security: is the area formally designated for long-
three vegetation and land cover types were identified term protection of natural values?
based on dominant species (e.g., Oregon White Oak, 3. Biodiversity Values: does the area include land that
Ponderosa Pine) and basic structure (e.g., woodland, has high-quality habitat or does it make an
grassland, forest). This large mmu resulted in several important contribution to ecosystem function?
vegetation and land cover types not being represented 4. Size: is the land a large block, and is it managed as
(e.g., small stands of native forest types) or being under- a unit?
represented on the map (e.g., wetlands, riparian areas).
Current Conservation Network. The land ownership data
The map was used for “coarse filter” analyses to assess
layer and the ratings developed though the Biodiversity
the distribution, abundance, and status of Oregon’s
Management Rating System were used to identify lands
biodiversity. Two statewide vegetation maps of Oregon
in Oregon that are devoted to the long-term protection of
have been completed since the ORB completed its
biodiversity, or a “Network of Conservation Lands.” Only
assessment. One map was developed by the Oregon Gap
lands that were rated at the high end of the scale (i.e., 8-
Analysis Program and has a mmu of 250 acres (100 ha).
10) by the Biodiversity Management Rating System were
The second map was developed in 1999 by the Northwest
 CHAPTER 5: PRIORITY HABITATS 159

included in the conservation network. Using those • About 1.8% of the Big Sagebrush-Bunchgrass type is
guidelines, the OBP determined that almost 10% of included in the conservation network.
Oregon’s total area, not including tribal or most private • About 3.1% of Oregon white oak woodlands are
lands, was included in the current conservation network. included in the conservation network.
Salmon Core Areas. Salmon Core Areas were identified • Only about 0.1% of bitterbrush steppe is included in
by the Oregon Coastal Salmon Restoration Initiative as the conservation network.
areas of critical importance to sustain salmon populations 3. Identification of statewide priorities for conservation
in individual basins. Maps identifying the historic based on widespread decline of habitat types and
distribution and current status of Oregon’s anadromous significance of the habitat in multiple ecoregions.
fish species50 also were used to identify areas important Results of the analyses indicated the following habitat
to salmon populations. types were priorities for conservation in Oregon:
Aquatic Diversity Areas. These areas were identified by • Oak savanna and woodlands,
the Oregon Chapter of the American Fisheries Society as • Wetlands,
being important to the conservation of aquatic diversity2
• Riparian,
because they represent Oregon’s best remaining aquatic
habitats and include connecting corridors and at-risk fish • Bottomland hardwood forests,
species. • Old-growth conifer, and
• Native grasslands and prairies.
Statewide Analysis
The OBP’s statewide analyses consisted of coarse-filter 4. Summary of the abundance and distribution of non-
analyses that provided an overview of current conditions native habitats. Non-native land cover types account for
in Oregon. To conduct the analyses, OBP used information greater than 16% of Oregon’s landscape.
about current and historic vegetation, aquatic ecosystems, • 11% of Oregon’s land is in farmland and developed
at-risk species, human population and land development pastures;
(e.g., population growth, road networks), and land • 4.5% of Oregon’s land is dominated by exotics such as
ownership and administration. General assessments cheatgrass;
included:
• 0.7% of Oregon’s land is in urban, industrial, and
1. Identification and description of Oregon’s network of residential classes or cover types;
conservation lands.
• Native habitats most affected by conversion to non-
• Greater than 1/3 of Oregon’s native vegetation types native types include grasslands, prairies, wetlands, and
have <5% of their distribution within the existing bottomland hardwood forests.
network of conservation lands.
• Largely due to wilderness designations and the Ecoregional Analyses
President’s Northwest Forest Plan, current management As a complement to the statewide analyses, OBP
for biodiversity is most extensive in alpine habitats and conducted analyses at the ecoregional level. The goal of
Westside forests. these analyses were to (1) identify the elements of
biodiversity that required additional protection and those
• Westside ecoregions (Coast Range, Klamath, and West
that were protected by the current conservation network,
Cascades) have large federal ownerships and
and (2) identify areas with the potential to address
approximately 25% of the land is included in the
statewide and ecoregional priorities. The results of the
network of conservation lands.
ecoregional analyses included a general description of
• Willamette Valley and Columbia Basin Ecoregions each ecoregion, its current and historic vegetation, and
have a large proportion of land in private ownership information about climate, land forms, habitats, industry,
and have less than 2% of land in the network of and human population levels. Analyses included the
conservation lands. following:
• Eastside ecoregions (Owyhee Uplands, East Cascades, 1. Vegetation Analysis: identification of vegetation types
Blue Mountains, Basin and Range, and High Lava that are conservation priorities based on current
Plains) have large federal ownership, but only 2-7% of management status and an assessment of changes in
the land is included in the network of conservation abundance and distribution over the past century;
lands.
2. At-risk Species: analysis of abundance, distribution,
2. Statewide and ecoregional analyses identifying risks/threats, and representation in the current
vegetation types represented in the existing conservation network;
conservation network and how well each type was
3. Ecosystem changes: assessment of changes in
represented. Examples of results of these analyses
ecosystem processes and vegetation structure not
include the following:
addressed by analyses of the coarse-scale vegetation
• Over 90% of subalpine and alpine meadows are map, and the results and impacts of management
included in the conservation network. practices such as fire suppression, timber harvest, and
grazing;
 160 Wildlife–Habitat Relationships in Oregon and Washington

4. Analyses of aquatic species; is unique in that it was designed to be used by many user-
5. Summary of conservation issues for the ecoregion, groups, accommodating geographic and species-specific
such as: queries and thus the differing priorities of those groups.
For example, the system can be used by the CDOW to
• threats, human population growth, economic
help identify the habitats that should be protected in a
development, pollution, conversion to non-native
specific resource area to maintain native fauna or to protect
habitats;
areas important to a single species. Additionally, the
• changes in management practices (e.g., fire system can be used by county planners interested in
suppression, grazing, timber harvest, conversion of identifying and protecting habitats for economically
wetlands); important species in their county. The High Priority
• changes in natural disturbance regimes (e.g., fire Habitat system is available through an interactive Internet
suppression, flooding); site, the Colorado Natural Diversity Information Source
• invasions of non-native plant and animal species; (http://www.ndis.nrel.colostate.edu/), a site maintained
through a cooperative effort of state and private agencies.
• habitat fragmentation; and small private ownerships.
The goal of the Natural Diversity Information Source is to
6. Identification of “Conservation Opportunity Areas,” provide easy access to biological, geopolitical, and
or areas with the potential to address statewide and demographic data needed to understand potential impacts
ecoregional conservation priorities. Generally, for changes in land use on wildlife and natural
characteristics of these areas included the following: communities.
• large blocks of native habitats,
Data Sets
• vegetation or habitats that have declined,
To develop the High Priority Habitat System, the CDOW
• vegetation types not well represented in the used existing data sets, including a statewide vegetation
conservation network, map, species distribution maps, and the Colorado
• at-risk species, and Vertebrate Ranking System, an information system that
• potential to complement or connect the existing incorporates information from several existing databases
conservation network. to identify species at risk.
The OBP used a two-step approach to identify Statewide Vegetation Map, Fifty-two vegetation types
Conservation Opportunity Areas. The first step used GIS were identified and mapped by the Colorado Gap Analysis
data layers as a quantitative approach that enabled the Project. The mmu for the vegetation map was 250 acres
project to (1) identify gaps in the existing conservation (100 ha), and was determined by the National Gap
network, (2) assess changes in the conservation network, Analysis Program. Basin-wide classifications and maps
(3) assess changes from historic vegetation patterns, and provided more detailed information.
(4) display areas identified as having significant
Species Distributions Maps. CDOW used two sources
biodiversity values. The second step consisted of a
of information to delineate vertebrate distribution. For rare
subjective assessment of the potential of different areas to
species or economically important species, CDOW used
enhance the existing conservation network. This
data sets that tracked the occurrence of each species to
assessment included evaluations of land ownership,
develop maps identifying the known distribution of each
current management, existing and potential programs for
species. For most other species, the project used maps that
conservation, pending public policy discussions, and
predicted species distributions based on the model created
potential threats to the elements of biodiversity.
by the Colorado Gap Analysis Project. The Gap model
Defenders of Wildlife published a book, Oregon’s Living
used the vegetation map, vegetation associations, and
Landscape: Strategies and Opportunities to Conserve
elevation constraints to predict the distribution of each
Biodiversity,7 outlining the process followed and the results
wildlife species.
achieved by the OBP. The book is written in a style that is
appealing to the general public, and describes the Colorado Vertebrate Ranking System. The CDOW
components of biodiversity as well as a process that can developed the Colorado Vertebrate Ranking System10
be used to identify conservation needs and opportunities (COVERS) to provide information necessary to evaluate
in Oregon. the conservation needs of Colorado’s native vertebrates.
The system was modeled after a process developed by
Colorado: the Florida Non-game Wildlife Program,24 and is linked
to and incorporates information from several statewide
The High Priority Habitat System databases including the Colorado Natural Heritage
Recently, the Colorado Division of Wildlife (CDOW)
Program’s ranking databases, the Colorado Wildlife
developed a system, the High Priority Habitat system, to
Species Database, and the Colorado Bird Observatory
rank habitats in terms of their priority for conservation.4
database.
This approach to identification and conservation of
The COVERS ranking process is designed to identify
priority habitats and areas is the most recently developed
species at risk, and uses a two-step process to accomplish
and complex approach for the western states. The system
that task. First, all species are ranked for variables that
 CHAPTER 5: PRIORITY HABITATS 161

indicate (1) the degree of biological imperilment of the imperilment of the species, (2) the state of knowledge,
species and (2) the state of knowledge about the species’ protection, and management for the species, and (3) the
distribution, abundance, and ecology. Variables ranked in importance of the species to the health of the global
this step include the following: population and to biodiversity in Colorado.
1. Biological imperilment Ranking Habitat Value
• global population size The High Priority Habitat system is designed so that the
• population concentration user can specify the geographic area and group(s) of
• largest global population size species that are of interest to them. Possible geographic
• global population trend areas include the entire state of Colorado, any county or
• state population trend hydrounit, Bureau of Land Management District or
• global distribution Resource Area, National Forest or Grassland, and CDOW
• global distribution trend Regions or Game Management Units. The system
• ecological specialization continually is being modified and improved, and
dietary specialization eventually will allow for queries of user-defined groups
reproductive specialization of species and for evaluations of single species. Presently,
other specialization the system is capable of accommodating queries for the
• sensitivity to exotic/invasive organisms following groups of species:
• sensitivity to human-induced factors
• Rare species
2. Current state of knowledge and management Federally Listed Threatened and Endangered
• knowledge of state distribution State Listed Threatened and Endangered
• knowledge of state populations trend State Species of Special Concern
• knowledge of state population limitations Colorado Natural Heritage Program Imperiled
• ongoing management in Colorado Species
• number of protected sites or populations in • Taxonomic groups
Colorado amphibians
Species identified as “imperiled” though the first step reptiles
of the process underwent an additional rating process to birds
determine the degree of imperilment and to evaluate mammals
factors that could influence conservation of the species. native fish
Variables ranked at this stage of the process included: mollusks
3. Additional biological variables • Other Groupings
• condition of largest population all species
• current distribution in Colorado economically important species
• reproductive potential for recovery The High Priority Habitat system ranks the vegetation
number of offspring per year polygons identified in the Colorado Gap Analysis
years to first reproduction statewide vegetation map. These polygons were identified
• magnitude of habitat threat from remote sensing techniques and each polygon was
• immediacy of habitat threat assumed to represent a single vegetation type. Therefore,
• direct threat to species the High Priority Habitat system serves as a tool to help
• portion of available habitat currently occupied identify the value and importance of each vegetation
4. Importance of Colorado populations polygon identified in the project’s vegetation maps.
• systematic significance at the state level Three methods were developed for ranking vegetation
• percent of total range in Colorado polygons. The first two methods were developed to rank
• percent of global population in/visiting Colorado vegetation polygons according to their importance to (1)
• relation of Colorado distribution to global range economically important species and (2) rare species. These
• importance of species to other organisms methods used the distribution maps developed by CDOW
for rare species and economically important species.
5. Social considerations Unlike the maps developed through the Colorado Gap
• economic consequences of population decline Analysis Program, which predict the distribution of
• economic benefits of population increase species, the maps developed by CDOW for rare species
• current listing status and economically important species identify the known
other considerations distribution of each species. The third method for ranking
The scores for each variable were recorded in the vegetation polygons can be used for all species, including
COVERS database, and can be accessed to address or rare species and economically important species, and uses
evaluate specific conservation needs of each species. The the maps of potential distribution of each species predicted
COVERS database also calculates summary scores for each by the Colorado Gap Analysis model.
species that can be used to indicate (1) the biological
 162 Wildlife–Habitat Relationships in Oregon and Washington

Ranking Habitat Values for Economically Important and planning designations, tax incentives, technical
Species. For each economically important species, assistance, and cost sharing. The OBP published a booklet
“activity areas,” or areas used by the species for a specific that discusses these incentives, their strengths and
purpose (e.g., elk calving areas) or during a specific season weaknesses, and provides suggestions for modifications
(e.g., elk winter range), were mapped and then rated or additional incentives that will contribute to biodiversity
subjectively in terms of their importance to the species. conservation on lands managed primarily for commodity
For example, winter range was rated as relatively less production and other human uses.47 Another source of
important to a species’ survival than severe winter range. information about conservation incentives has been
A composite map for all economically important species compiled by the Oregon Coordinated Resource
was developed by incorporating ratings for the Management (OCRM) Task Group,29 a group of federal
importance of activity areas from the individual species and state agencies supporting coordination of renewable
maps. For the composite map, values assigned to natural resource planning and management on state,
vegetation polygons were not additive, but reflected the federal, and private lands. The OCRM Task Group has
highest score assigned to the polygon for any of the identified tax incentives and public funding sources for
economically important species. The composite map can individual landowners as well as public and private
be used to identify the vegetation polygons that are organizations. Projects funded by incentive programs
relatively more important to economically important include but are not limited to the following:
species. 1. Watershed improvement projects
2. Fish and wildlife habitat improvement projects
Ranking Habitat Values for Rare Species. Similar to the
3. Wetland/Riparian Area improvement projects
methods followed for economically important species, for
4. Timber stand development and improvement
each rare species, activity areas were mapped and then
projects
rated according to their importance to the species. In
5. Soil projection projects
addition, activity areas were weighted according to the
6. Landowner stewardship plan development
rarity of the species (e.g., Federally Endangered, Federally
projects
Threatened or State Endangered, State Threatened, or
7. Education projects
Federal Category 1). A composite map for all rare species
was developed by incorporating ratings for species rarity
and the importance of activity areas. For the composite Environmental Report Cards
map, values assigned to vegetation polygons were not The fundamental goal of conservation programs is to
additive, but reflected the highest score assigned to the maintain ecosystems and the biodiversity contained
polygon for any of the rare species. The composite map within them. Incorporating the concept of “ecosystems”
can be used to identify the vegetation polygons that are into conservation efforts serves to broaden our view of
relatively more important to rare species. the environment by recognizing that management of our
natural resources must integrate ecological relationships
Ranking Habitat Values for All Species. For each species, with social and political values so that our natural systems
vegetation polygons within the species’ distribution were can be protected and maintained over time. To be
assigned the value from the COVERS database indicating successful in our attempt to maintain functional and viable
the degree of biological imperilment. This score provides ecosystems, we will need to measure and monitor our
an indication of the biological imperilment of the species progress. One way of monitoring the success of our
in Colorado and ranks the species in terms of its needs for achievements is to report our progress in terms of our goals
conservation efforts and habitat protection. Composite and objectives. An ideal way to do this is to adopt a “report
maps were developed for rare species, different taxonomic card” that routinely apprizes the general public, scientific
groups, and economically important species by assigning community, and decision-makers about our progress.
each vegetation polygon a score calculated as the sum of
the scores for biological imperilment for all species Criteria for Developing a Report Card
predicted to occur in that polygon. Therefore, the overall The report card should document our progress towards
value of a vegetation polygon incorporates information achieving our desired conditions and should be written
on species richness and species imperilment, and provides so that it is understandable by a wide audience. This is
an indication of the importance of each polygon to the because clear communication of the desired conditions
conservation of wildlife species. will allow the public to follow and aspire to the same
vision. In addition, the report card should define
Conservation and Stewardship terminology as well as identify our desired conditions,
Incentives measures used to evaluate performance, the scientific basis
for assigning grades, and the current conditions/processes
Many programs exist nationally as well as regionally and
of the abiotic and biotic resources. Other information that
locally to encourage landowners and managers to adopt
is important to incorporate is identified by Harwell et al.11
alternative land management practices and conserve
and includes but is not limited to: (1) habitat quality—
biodiversity. Examples of incentives include relief from
landscape mosaic; spatial extent; landscape and
natural resource regulations, land acquisitions and
community diversity; connectivity, fragmentation,
exchanges, easements, management agreements, zoning
 CHAPTER 5: PRIORITY HABITATS 163

structural diversity; (2) integrity of biotic community— Wildlife Habitat Maps for
biodiversity; trophic structure; exotic, invasive or noxious
species; threatened and endangered species; economic or
Oregon and Washington
aesthetically important species; (3) ecological processes— As a starting point for the development of quantitative
primary and secondary productivity; succession; key processes for identifying and prioritizing areas important
ecological functions nutrient cycling; species dispersal and to conservation values, we have summarized existing
migration; (4) water quality—biological characteristics, digital data for Oregon and Washington. The frontispiece
physical characteristics, chemical characteristics; (5) of this book depicts the wildlife habitats for each state and
hydrological system—hydroperiod; surface and Tables 1, 2, and 3 (which follow the Literature Cited) list the
groundwater flow; water storage; channel complexity or acreage of each wildlife habitat and the amounts that are
sinuosity; sediment transport; (6) disturbance regime— protected within each ecoregion of each state. We hope
frequency and intensity of fire, flooding, storms, and these will provide a simple overview of the abundance and
drought; disease or pest outbreaks; anthropogenic distribution of our wildlife habitat and land cover types,
disturbances; (7) soil quality—biological, physical, and generating ideas and contributing to the development of
chemical characteristics; erosion and accumulation of soil quantitative processes for identifying priority areas and
and sediment; and (8) air quality—biological, physical, and habitats and conserving the biodiversity of the region.
chemical characteristics.
Literature Cited
In addition to reporting on the progress being made
1. Anselin, A., P. M. Meire, and L. Anselin. 1989. Multi-criteria techniques in
towards the project’s environmental goals, it also would ecological evaluation: an example using the analytical hierarchy process.
be helpful to identify 5-10 questions that the project is Biological Conservation 49: 215-229.
striving to answer. This step would be a tremendous help 2. Bottom, D., S. Beckwitt, J. Christy, S. Clarke, J. Dambacher, C. Rissell, R.
with initiating research coordination and inter- and intra- Hughes, H. Li, D. McCullough, A. McGee, K. Moore, R. Nawa, and S.
agency cooperation. Finally, any grade assigned should Thiele. 1993. Oregon critical watersheds database. Watershed
be based on a scientific rationale that gives the reader a Classification Subcommittee of the Natural Production Committee,
Oregon Chapter American Fisheries Society, Corvallis, OR.
clear understanding of the current status of the project 3. Bolton, M. P. and R. L. Specht. 1983. A method for selecting
and what steps are needed to achieve its goals. conservation reserves. Australian National Parks and Wildlife Service
For most projects, involving a variety of interest groups Occasional Paper 8.
in the development of the desired conditions will facilitate 4. Colorado Division of Wildlife. 1998. High priority habitats. Natural
and promote shared expectations and ownership of the Resource Ecology Laboratory, Colorado State University, Fort Collins,
ideas and goals. For example, since Oregon and CO.
5. Csuti, B., P. J. Kennelly, S. M. Meyers, M. M. P Huso, and J. Sifneos. 1998.
Washington share several diverse ecoregions, a regional
Multiscale biodiversity conservation: a prototype process for Oregon.
assessment of both state’s ecoregions would make a large Final report: Task 2A; Develop and refine methodology for analyzing
contribution to conservation efforts by allowing the region biodiversity at the state scale. U.S. Environmental Protection Agency,
and multiple agencies to address multi-jurisdictional Corvallis, OR.
issues in a cohesive and unified manner. Agencies like the 6. ———, A. J. Kimerling, T. A. O’Neil, M. M. Shaughnessy, E. P. Gaines, and
Northwest Power Planning Council, Bonneville Power M. M. P. Huso. 1997. Atlas of Oregon wildlife: distribution, habitat, and
Administration, U.S. Forest Service, and U.S. Fish and natural history. Oregon State University Press, Corvallis, OR.
7. Defenders of Wildlife. 1998. Oregon’s living landscape: Strategies and
Wildlife Service would be able to set their program goals
opportunities to conserve biodiversity. Oregon State University Press,
in a manner that would meet a regional vision and would Corvallis, OR.
transcend political boundaries. 8. Federal Geographic Data Committee. 1996. FGDC vegetation
Two recently completed efforts that have used classification and information standards. U.S. Geological Survey, Reston,
ecoregion-level analysis to address conservation concerns VA.
were the OBP and World Wildlife Fund’s review of the 9. Gotmark, F., M. Ahlund, and M. O. G. Eriksson. 1986. Are indices reliable
for assessing conservation value of natural areas? An avian case study.
Terrestrial Ecoregions of North America. 34 Both
Biological Conservation 38: 55-73.
approaches took a landscape-level view of the ecoregion 10. Gross, J. E., and Melcher. 1997. Colorado vertebrate ranking system.
and discussed its biological uniqueness, current Natural Resource Ecology Laboratory, Colorado State University, Fort
conservation status, and priority actions. However, Collins, CO.
conservation efforts and assessments should not be limited 11. Harwell, M.,V. Myers, T.Young, A. Barruska, N. Gassman, J. Gentile, C.
to regional and ecoregional analyses. Assessments need to Harwell, S. Appelbaum, J. Barko, B. Causey, C. Johnson, A. McLean, R.
be conducted at several scales including local scales and at Smola, P. Templet, and S. Tosini. 1999. A framework for an ecosystem
integrity report card. Bioscience 49:7: 543-556.
the level of hydrologic units comprising basins and
12. International Council for Bird Preservation. 1992. Putting biodiversity
watersheds. A multi-scale approach will depict results of on the map: priority areas for global conservation. Cambridge, England.
a shared vision along with noting the implemental 13. Kagan, J. S., and S. Caicco. 1992. Manual of Oregon actual vegetation.
achievements at the region to local levels. Because specific Idaho Fish and Wildlife Research Cooperative Unit, University of Idaho,
goals and objectives may vary at these different levels, it is Moscow, ID.
important to address the time requirement to meet 14. Karr, J. R. 1990. Biological integrity and the goal of environmental
strategies across all levels. legislation: lessons for conservation biology. Conservation Biology 4:
244-250.
 164 Wildlife–Habitat Relationships in Oregon and Washington

15. Kershaw, M., G. M. Mace, and P. H. Williams. 1995. Threatened status, 37. Scott, J. M., F. Davis, B. Csuti, R. Noss, B. Butterfield, C. Groves, H.
rarity, and diversity as alternative selection measures for protected Anderson, S. Caicco, F. D’Erchia, T. C. Edwards, Jr., J. Ulliman, and G.
areas: a test using Afrotropical antelopes. Conservation Biology 9: 324- Wright. 1993. Gap analysis: a geographical approach to protection of
334. biological diversity. Wildlife Monographs 123.
16. Kiester, A. R., J. M. Scott, B. Csuti, R. F. Noss, B. Butterfield, K. Sahr, and 38. ———, and M. D. Jennings. 1998. Large-area mapping of biodiversity.
D. White. 1996. Conservation prioritization using GAP data. Annals of the Missouri Botanical Garden 85:34-47.
Conservation Biology 10: 1332-1342. 39. Short, H. L. and J. B. Hestbeck. 1995. National biotic resource
17. Kiilsgaard, C. 1999. Oregon Vegetation: Mapping and Classification of inventories and GAP analysis. BioScience 45: 535-539.
Landscape Level Cover Types. Final Report. U.S. Geological Survey- 40. Smith, P. G. R., and J. B. Theberge. 1987. Evaluating natural areas using
Biological Resources Division: Gap Analysis Program. Moscow, ID. multiple criteria: theory and practice. Environmental Management 11:
18. Kiilsgaard C., and C. Barrett. 1999. Map of: Oregon Vegetation 447-460.
Landscape-Level Cover Types. Published by Northwest Habitat 41. Specht, R. L. 1975. The report and its recommendation. Pages 11-21 in:
Institute, Corvallis, OR. F. Fenner, editor. A national system of ecological reserves in Australia.
19. Kirkpatrick, J. B. 1983. An iterative method for establishing priorities Report 19. Australian Academy of Science, Canberra, Australia.
for the selection of nature reserves: an example from Tasmania. 42. Stoms, D. M. 1994. Scale dependence of species richness maps.
Biological Conservation 25: 127-134. Professional Geographer 46: 346-358.
20. Lawton, J. H., J. R. Prendergast, and B. C. Eversham. 1994. The numbers 43. Thomas, J. W., technical editor. 1979. Wildlife habitats in managed
and spatial distributions of species: analyses of British data. Pages 177- forests: the Blue Mountains of Oregon and Washington. Agriculture
195 in: P. L. Forey, J. Humphries, and R. I.VaneWright, editors. Handbook 553. U.S. Forest Service, Washington D.C.
Systematics and conservation evaluation. Clarendon Press, Oxford, 44. Thomas, C. D., and H. C. Mallorie. 1985. Rarity, species richness and
England. conservation: butterflies of the Atlas Mountains in Morocco. Biological
21. Margules, C. R. 1989. Introduction to some Australian developments Conservation 33: 95-117.
in conservation evaluation. Biological Conservation 50: 1-11. 45. Turpie, J. K. 1995. Prioritizing South African estuaries for conservation:
22. ———, C. R., A. O. Nicholls, and R. L. Pressey. 1988. Selecting a practical example using waterbirds. Biological Conservation 74: 175-
networks of reserves to maximize biological diversity. Biological 185.
Conservation 43: 63-76. 46. Usher, M. B. 1986. Wildlife conservation evaluation: attributes, criteria
23. ———, and M. B. Usher. 1981. Criteria used in assessing wildlife and values. In: M. B. Usher, editor.Wildlife conservation evaluation.
conservation potential: a review. Biological Conservation 21: 79-109. Chapman and Hall, London, England.
24. Millsap, B. A., J. A. Gore, D. E. Runde, and S. I. Cerulean. 1990. Setting 47. Vickerman, S. 1998. Stewardship incentives: conservation strategies for
priorities for the conservation of fish and wildlife species in Florida. Oregon’s working landscape. Defenders of Wildlife, Lake Oswego, OR.
Wildlife Monographs 111. 48. Washington Department of Fish and Wildlife. 1996. Priority habitats
25. Nicholls, A. O., and C. R. Margules. 1993. An upgraded reserve section and species list. Habitat Program, Washington Department of Fish and
algorithm. Biological Conservation 64: 165-169. Wildlife, Olympia, WA.
26. Noss, R. F. 1987. From plant communities to landscapes in 49. White, D., A. J. Kimerling, and W. S. Overton. 1992. Cartographic and
conservation inventories: a look at The Nature Conservancy (U.S.A.). geometric components of a global sampling design for environmental
Biological Conservation 41: 11-37. monitoring. Cartography and Geographic Information Systems 19: 5-22.
27. ———, and A.Y. Cooperrider. 1994. Saving nature’s legacy: protecting 50. Wilderness Society, The. 1993. Pacific salmon and federal lands, a
and restoring biodiversity. Island Press, Washington, D.C. regional analysis. The Wilderness Society, Bolle Center for Forest
28. O’Neil, T. A., R. J. Steidl, W. D. Edge, and B. Csuti. 1995. Wildlife habitat Ecosystem Management, Washington D.C.
clusters: using wildlife communities to improve vegetation classification 51. Williams, G. 1980. An index for ranking of wildfowl habitats, as applied
for conserving biodiversity. Conservation Biology 9: 1482-1491. to eleven sites in west Surrey, England. Biological Conservation 18: 3-
29. Oregon Coordinated Resource Management Task Group. 1996. Public 99.
funding sources for landowner assistance. Oregon CRM Task Group, 52. Williams, P., D. Gibbons, C. Margules, A. Rebelo, C. Humphries, and R.
Portland, OR. Pressey. 1996. A comparison of richness hotspots, rarity hotspots and
30. Prendergast, J. R., R. M. Quinn, J. H. Lawton, B. C. Eversham, and D. W. complementary areas for conserving diversity of British birds.
Gibbon. 1993. Rare species, the coincidence of diversity hotspots and Conservation Biology 10: 155-174.
conservation strategies. Nature 365: 335-337. 53. Wright, D. F. 1977. A site evaluation scheme for use in the assessment
31. Pressey, R. L., C. J. Humphries, C. R. Margules, and R. I. Vane-Wright. of potential nature reserves. Biological Conservation 11: 293-305.
1993. Beyond opportunism: key principles for systematic reserve
selection. Trends in Ecology and Evolution 8: 124-128.
32. ———, and A. O. Nicholls. 1989. Application of a numerical algorithm
to the selection of reserve in semi-arid New South Wales. Biological
Conservation 50: 263-278.
33. Rapoport, E. H., G. Borioli, J. A. Monjeau, J. E. Puntieri, and R. D. Oviedo.
1986. The design of nature reserves: a simulation trial for assessing
specific conservation value. Biological Conservation 37: 269-290.
34. Ricketts, T., E. Dinerstein, D. Olson, C. Loucks, W. Eichbaum, D.
DellaSala, K. Kavanagh, P. Hedao, P. Hurley, K. Carney, R. Abell, and S.
Walters. 1999. Terrestrial ecoregions of North America, a conservation
assessment. Supported by World Wildlife Fund. Island Press,
Washington, D.C.
35. Rojas, M. 1992. The species problem and conservation: what are we
protecting? Conservation Biology 6: 170-178.
36. Saetersdal, M., J. M. Line, and H. J. B. Birks. 1993. How to maximize
biological diversity in nature reserve selection: vascular plants and
breeding birds in deciduous woodlands, western Norway. Biological
Conservation 66: 131-138.
 CHAPTER 5: PRIORITY HABITATS 165

Table 1. The total acreage of wildlife habitat types that occur and are under conservation-oriented protection
strategies in Oregon and Washington.

Wildlife Habitat Oregon Washington

Total Protected1 Total Protected1

Westside Lowlands Conifer-Hardwood Forest9,349,756 1,544,745 9,064,128 469,747

Westside Oak and Dry Douglas-fir Forest and 433,132 2,596 425,038 13,832
Woodlands
Southwest Oregon Mixed Conifer Forest 4,020,321 900,642 Does Not Occur Does Not Occur
Montane Mixed Conifer Forest 2,949,586 933,681 4,653,306 2,061,798
Eastside (Interior) Mixed Conifer Forest 4,126,957 279,223 4,662,101 400,429
Lodgepole Pine Forest and Woodlands 532,587 13,834 119,201 17,720
Ponderosa Pine Forest and Woodlands 6,226,351 98,349 1,927,176 98,907
Upland Aspen Forest 19,685 7,783 100,621 3,823
Subalpine Parkland 84,240 48,410 327,442 190,593
Alpine Grassland and Shrublands 291,494 163,170 1,591,115 1,300,415
Westside Grasslands 1332 Not Mapped 22,491 1,320
Ceanothus-Manzanita Shrublands 52,104 22,016 Does Not Occur Does Not Occur
Western Juniper and Mountain Mahogany 4,037,221 105,708 Does Not Occur Does Not Occur
Woodlands
Eastside (Interior) Canyon Shrublands 358,250 68,709 Not Mapped3 Not Mapped
Eastside (Interior) Grasslands 1,935,794 132,070 1,002,076 65,535
Shrub-steppe 17,420,753 490,879 7,144,697 473,622
Dwarf Shrub-steppe 514,066 26,777 Not Mapped4 Not Mapped
Desert Playa and Salt Scrub Shrublands 719,503 48,105 Not Mapped4 Not Mapped
Agriculture, Pasture and Mixed Environs 6,197,887 28,134 9,251,107 43,991
Urban and Mixed Environs 575,087 487 1,204,680 4,967
Open Water - Lakes, Rivers, Streams 780,901 96,437 761,360 46,896
Herbaceous Wetlands 1,031,343 135,121 210,451 12,966
Westside Riparian-Wetlands 168,872 10,339 347,653 38,287
Montane Coniferous Wetlands 56,099 2,373 241,450 25,551
Eastside (Interior) Riparian-Wetlands 31,121 2,604 100,763 9,285
Coastal Dunes and Beaches 52,451 1,821 Not Mapped4 Not Mapped
Coastal Headlands and Islets 9,137 987 7,776 1,831
Bays and Estuaries 172,748 31,332 226,336 3,6896
Inland Marine Deeper Water Does Not Occur Does Not Occur 1,855,780 2,9076
Marine Nearshore 223,371 1,350 750,329 1,0446
Marine Shelf 3,905,164 Not Determined 4,780,625 Not Determined5
Oceanic 33,987,189 Not Determined 19,845,660 Not Determined5
Totals 100,263,303 70, 532,093
1
Oregon’s values were determined by using the Defenders of Wildlife’s biodiversity management rating (Defenders of Wildlife 1998).
Specifically, only those lands rated with a high degree of protection were used, and these were areas that had a rating of 8, 9, or 10.
Washington’s values were determined using the Washington Gap Analysis protected areas data layer; specifically, only lands with a
high degree of protection were used, and these were areas that had a status of 1 or 2 (Cassidy 1997:13). Marine protected areas
were determined from the following publications: Murray, M.R. 1998. The status of marine protected areas in Puget Sound.Volumes I
and II Puget Sound/Georgia Basin Environmental Report Series 8.Puget Sound Water Quality Action Team, Olympia, Washington,
USA; and Robinson, M.K. 1999. The status of Washington’s coastal marine protected areas. Washington Department of Fish and
Wildlife, Olympia, USA.
2
Because of difficulty in classifying this type using remote sensing (i.e., discerning native grasslands from pasture lands) native
westside grasslands have inadvertently been classified within the agriculture habitat type. Nonetheless, there are few areas known to
be native westside grasslands.
3
This type was only recognized along the Oregon and Washington border, otherwise it was not part of the vegetation classification
when the Washington Gap Project mapped the state of Washington. Thus, no wildlife habitat area was determined.
4
This type was not part of the vegetation classification when the Washington Gap Project mapped the state of Washington. Thus, no
wildlife habitat area was determined.
5
While not included in this analysis, there are 102 Marine Protected Areas (MPAs) in Puget Sound (Murray 1998); an additional 33
MPAs occur along the outer Washington coastline (Robinson 1999). While most MPAs are quite small in size, the Olympic Coast
National Marine Sanctuary along the outer coast of northwest Washington offers protection to some 2,112,000 acres of marine
nearshore, marine shelf, and oceanic habitat.
 166 Wildlife–Habitat Relationships in Oregon and Washington

Table 2. Amount of protected1 wildlife habitat types in Oregon by ecoregion.

Wildlife-Habitat2 Ecoregion3

1 2 3 4 5 6 7 8 9 10 11 12 13

1 790,718 8,005 98,483 628,149 0 19,389 0 0 0 0 0 0 0

2 85 1,757 0 662 0 92 0 0 0 0 0 0 0
3 3,714 8,143 608,870 248,819 31,095 0 0 0 0 0 0 0 0
4 0 0 7,941 247,269 391,740 77,026 42,611 167,093 0 0 0 0 0
5 0 0 0 0 19,802 81,002 150,958 26,769 74 0 0 185 0
6 0 0 0 316 5,317 8,201 0 0 0 0 0 0 0
7 0 850 0 8,519 15,733 26,319 39,466 641 0 0 0 130 0
8 0 0 0 0 0 0 0 59 1,102 0 0 0 0
9 0 0 185 712 40,540 2,712 0 0 0 0 0 0 0
10 0 0 133 0 16,424 0 3,988 141,516 0 0 0 0 0
11 (this habitat not mapped, thus no acreage figures currently available)
12 0 20,549 1,459 0 0 0 0 0 0 0 0 0
13 0 0 0 0 0 808 15,624 0 9,750 0 50 9,715 0
14 0 0 0 0 0 0 0 0 0 0 2,351 0 0
15 0 0 0 0 0 2,206 125,246 4,798 0 0 0 0 0
16 0 0 0 0 0 679 3,045 44 226,119 0 3,045 7,569 0
17 0 0 0 0 0 231 0 0 25,027 0 0 0 0
18 0 0 0 0 0 0 0 0 10,323 0 0 0 0
19 575 6,655 259 2,6230 2,769 1,142 0 129 0 0 0 0
20 0 294 0 0 0 120 0 0 0 0 0 0 0
21 128 468 63 1,413 1,215 2,268 110 802 40,618 0 102 0 0
22 885 834 688 203 343 46,320 1,310 0 7,008 0 28 0 0
23 7,324 2,330 69 315 0 0 0 0 0 0 0 0 0
24 0 0 0 988 585 692 106 0 0 0 0 0 0
25 0 0 0 0 0 0 0 0 238 0 0 0 0
26 371 0 1,451 0 0 0 0 0 0 0 0 0 0
27 369 0 621 0 0 0 0 0 0 0 0 0 0
28 31,030 13 26 268 0 0 0 0 0 0 0 0 0
30 731 0 619 0 0 0 0 0 0 0 0 0 0
31 0 0 0 0 0 0 0 0 0 0 0 0 0
32 0 0 0 0 0 0 0 0 0 0 0 0 0
1
Oregon’s values were determined by using the Defenders of Wildlife’s biodiversity management rating. Specifically, only those rated
with a high degree of protection were used, and these were areas that had a rating of 8, 9, or 10.
2
Wildlife Habitats: (1) Westside Lowlands Conifer-Hardwood Forest, (2) Westside Oak and Dry Douglas-fir Forest and Woodlands,
(3) Southwest Oregon Mixed Conifer Forest, (4) Montane Mixed Conifer Forest, (5) Eastside (Interior) Mixed Conifer Forest, (6)
Lodgepole Pine Forest and Woodlands, (7) Ponderosa Pine Forest and Woodlands, (8) Upland Aspen Forest, (9) Subalpine Parkland,
(10) Alpine Grassland and Shrublands, (12) Ceanothus-Manzanita Shrublands, (13) Western Juniper and Mountain Mahogany
Woodlands, (14) Eastside (Interior) Canyon Shrublands, (15) Eastside (Interior) Grasslands, (16) Shrub-steppe, (17) Dwarf Shrub-
steppe, (18) Desert Playa and Salt Scrub Shrublands, (19) Agriculture, Pasture and Mixed Environs, (20) Urban and Mixed Environs,
(21) Open Water-Lakes, Rivers, Streams, (22) Herbaceous Wetlands, (23) Westside Riparian-Wetlands, (24) Montane Coniferous
Wetlands, (25) Eastside (Interior) Riparian-Wetlands, (26) Coastal Dunes and Beaches, (27) Coastal Headlands and Islets, (28) Bays
and Estuaries, (30) Marine Nearshore, (31) Marine Shelf, (32) Oceanic.
3
Ecoregion: (1) Oregon Coast Range, (2) Western Oregon Interior Valley, (3) Siskiyou Mountains, (4) Westside Oregon Cascades, (5)
High Oregon Cascades, (6) Eastside Oregon Cascades, (7) Oregon Blue and Wallowa Mountains, (8) High Oregon Blue and Wallowa
Mountains, (9) Basin and Range, (10) Columbia Basin, (11) Owyhee Uplands, (12) High Lava Plains, (13) Snake River Plains.
 CHAPTER 5: PRIORITY HABITATS 167

Table 3. Amount of protected1 wildlife habitat types in Washington by ecoregion.

Wildlife Habitat2 Ecoregion3

1 2 3 4 5 6 7 8 9 10 11 12 13

1 101,073 21,038 0 0 0 746 2,979 0 0 93,120 58,577 857 191,353

2 0 3,532 0 0 17 0 8,921 0 0 0 1,222 139 0
4 273,608 0 11,984 0 17 195,521 91,259 32,658 277,140 677,505 336,052 0 166,054
5 0 0 71,349 23 1,248 178,324 17,704 44,307 69,145 11,775 6,554 0 0
6 0 0 384 0 11 4,776 2,296 0 5,049 3,375 1,829 0 0
7 0 0 18,053 474 8,940 25,270 11,362 18,677 15,433 693 5 0 0
8 0 0 190 63 29 297 0 0 2,680 564 0 0 0
9 0 0 135 0 0 44,703 7,793 0 27,015 50,477 23,838 0 36,632
10 75,390 0 4,298 0 0 153,578 12,187 5,872 203,560 582,801 145,585 0 117,143
11 0 1,276 44 0 0 0 0 0 0 0 0 0 0
14 (this habitat not mapped, thus no acreage figures currently available)
15 0 0 2,826 2,240 3,784 19,492 6,140 13,723 14,737 2,593 0 0 0
16 0 0 9,120 2,042 420,826 11,292 4,329 17,077 8,927 10 0 0 0
19 287 1,658 3,770 557 34,256 441 196 137 1,378 37 25 1,123 126
20 273 3,074 405 897 0 0 0 0 0 0 0 112 206
21 5,743 2,072 3,011 665 8,640 4,252 441 57 2,175 11,162 6,204 0 2,473
22 1,890 1,456 111 0 6,212 0 0 0 0 595 0 1,148 1,554
23 21,230 2,723 0 0 0 0 0 0 0 5,971 2,501 1,129 4,733
24 225 0 3,790 60 0 4,140 4,655 20 6,038 2,751 3,870 0 0
25 0 0 0 0 3,726 2,418 60 860 2,220 0 0 0 0
26 (this habitat not mapped, thus no acreage figures currently available)
27 1,831 0 0 0 0 0 0 0 0 0 0 0 0
28 2,166 882 0 0 0 0 0 0 0 0 0 478 163
29 0 2,907 0 0 0 0 0 0 0 0 0 0 0
30 718 326 0 0 0 0 0 0 0 0 0 0 0
31 0 0 0 0 0 0 0 0 0 0 0 0 0
32 0 0 0 0 0 0 0 0 0 0 0 0 0
1
Washington’s values were determined using the Washington Gap Analysis protected areas data layer. Specifically, only those lands
with a higher degree of protection were used, and these were areas that had a status of 1 or 2.
2
Wildlife Habitats: (1) Westside Lowlands Conifer-Hardwood Forest, (2) Westside Oak and Dry Douglas-fir Forest and Woodlands,
(4) Montane Mixed Conifer Forest, (5) Eastside (Interior) Mixed Conifer Forest, (6) Lodgepole Pine Forest and Woodlands, (7)
Ponderosa Pine Forest and Woodlands, (8) Upland Aspen Forest, (9) Subalpine Parkland, (10) Alpine Grassland and Shrublands, (11)
Westside Native Grasslands, (14) Eastside (Interior) Canyon Shrublands, (15) Eastside (Interior) Grasslands, (16) Shrub-steppe, (19)
Agriculture, Pasture and Mixed Environs, (20) Urban and Mixed Environs, (21) Open Water-Lakes, Rivers, Streams, (22) Herbaceous
Wetlands, (23) Westside Riparian-Wetlands, (24) Montane Coniferous Wetlands, (25) Eastside (Interior) Riparian-Wetlands, (26)
Coastal Dunes and Beaches, (27) Coastal Headlands and Islets, (28) Bays and Estuaries, (29) Inland Marine Deeper Water, (30) Marine
Nearshore, (31) Marine Shelf, (32) Oceanic.
3
Ecoregion: (1) Outer Olympic Peninsula, (2) Puget Sound, (3) Northeastern Corner, (4) Okanogan Highlands, (5) Columbia Basin, (6)
East Central Cascades, (7) Southeast Cascades, (8) Blue Mountains, (9) Northeast Cascades, (10) Northwest Cascades, (11)
Southwest Cascades, (12) Willamette Valley, (13) Inner Olympic Peninsula.
 CHAPTER 5: PRIORITY HABITATS 165

Table 1. The total acreage of wildlife habitat types that occur and are under conservation-oriented protection
strategies in Oregon and Washington.

Wildlife Habitat Oregon Washington

Total Protected1 Total Protected1

Westside Lowlands Conifer-Hardwood Forest9,349,756 1,544,745 8,952,281 A 469,747

Westside Oak and Dry Douglas-fir Forest and 433,132 2,596 186,862A 13,832
Woodlands
Southwest Oregon Mixed Conifer Forest 4,020,321 900,642 Does Not Occur Does Not Occur
Montane Mixed Conifer Forest 2,949,586 933,681 4,649,516A 2,061,798
Eastside (Interior) Mixed Conifer Forest 4,126,957 279,223 4,763,842� 400,429
Lodgepole Pine Forest and Woodlands 532,587 13,834 119,047A 17,720
Ponderosa Pine Forest and Woodlands 6,226,351 98,349 2,197,650� 98,907
Upland Aspen Forest 19,685 7,783 100,617� 3,823
Subalpine Parkland 84,240 48,410 327,430� 190,593
Alpine Grassland and Shrublands 291,494 163,170 1,599,056A 1,300,415
Westside Grasslands 1332 Not Mapped 18,612A 1,320
Ceanothus-Manzanita Shrublands 52,104 22,016 Does Not Occur Does Not Occur
Western Juniper and Mountain Mahogany 4,037,221 105,708 Does Not Occur Does Not Occur
Woodlands
Eastside (Interior) Canyon Shrublands 358,250 68,709 Not Mapped3 Not Mapped
Eastside (Interior) Grasslands 1,935,794 132,070 996,858A 65,535
Shrub-steppe 17,420,753 490,879 7,130,399A 473,622
Dwarf Shrub-steppe 514,066 26,777 Not Mapped4 Not Mapped
Desert Playa and Salt Scrub Shrublands 719,503 48,105 Not Mapped4 Not Mapped
Agriculture, Pasture and Mixed Environs 6,197,887 28,134 9,254,527A 43,991
Urban and Mixed Environs 575,087 487 1,204,636A 4,967
Open Water - Lakes, Rivers, Streams 780,901 96,437 761,307A 46,896
Herbaceous Wetlands 1,031,343 135,121 210,437A 12,966
Westside Riparian-Wetlands 168,872 10,339 347,638 A 38,287
Montane Coniferous Wetlands 56,099 2,373 241,824A 25,551
Eastside (Interior) Riparian-Wetlands 31,121 2,604 100,735A 9,285
Coastal Dunes and Beaches 52,451 1,821 Not Mapped4 Not Mapped
Coastal Headlands and Islets 9,137 987 7,775A 1,831
Bays and Estuaries 172,748 31,332 226,327A 3,6895
Inland Marine Deeper Water Does Not Occur Does Not Occur 1,855,713A 2,9075
Marine Nearshore 223,371 1,350 750,302A 1,0445
Marine Shelf 3,905,164 Not Determined 4,780,450A Not Determined5
Oceanic 33,987,189 Not Determined 19,844,932A Not Determined5
Totals 100,263,303 70, 628,773A
1
Oregon’s values were determined by using the Defenders of Wildlife’s biodiversity management rating (Defenders of Wildlife 1998).
Specifically, only those lands rated with a high degree of protection were used, and these were areas that had a rating of 8, 9, or 10.
Washington’s values were determined using the Washington Gap Analysis protected areas data layer; specifically, only lands with a
high degree of protection were used, and these were areas that had a status of 1 or 2 (Cassidy 1997:13). Marine protected areas
were determined from the following publications: Murray, M.R. 1998.The status of marine protected areas in Puget Sound.Volumes I
and II Puget Sound/Georgia Basin Environmental Report Series 8.Puget Sound Water Quality Action Team, Olympia,Washington,
USA; and Robinson, M.K. 1999.The status of Washington’s coastal marine protected areas.Washington Department of Fish and
Wildlife, Olympia, USA.
2
Because of difficulty in classifying this type using remote sensing (i.e., discerning native grasslands from pasture lands) native
westside grasslands have inadvertently been classified within the agriculture habitat type. Nonetheless, there are few areas known to
be native westside grasslands.
3
This type was only recognized along the Oregon and Washington border, otherwise it was not part of the vegetation classification
when the Washington Gap Project mapped the state of Washington. Thus, no wildlife habitat area was determined.
4
This type was not part of the vegetation classification when the Washington Gap Project mapped the state of Washington. Thus, no
wildlife habitat area was determined.
5
While not included in this analysis, there are 102 Marine Protected Areas (MPAs) in Puget Sound (Murray 1998); an additional 33
MPAs occur along the outer Washington coastline (Robinson 1999). While most MPAs are quite small in size, the Olympic Coast
National Marine Sanctuary along the outer coast of northwest Washington offers protection to some 2,112,000 acres of marine
nearshore, marine shelf, and oceanic habitat.
 6
Key Ecological Functions of Wildlife Species
Bruce G. Marcot & MadeleineVander Heyden

Introduction
An ecosystem is the set of all its component organisms
and populations and their ecological interactions with each To ensure sustainable wildlife populations, conserving
other and with the abiotic world. Traditionally, wildlife- threatened or endangered species should not stop at
habitat relationships (WHR) programs, models, and addressing only their individual habitat needs, that is, the
databases have focused on how the presence of terrestrial W = f(H) relation. In fact, a primary purpose of the
vertebrates is influenced by environmental conditions, and Endangered Species Act is “. . . to provide a means
have mostly ignored ecological interactions. WHR whereby the ecosystems upon which endangered species
approaches have assumed that wildlife (W) basically is a and threatened species depend may be conserved . . . .”
function of habitat (H), or W = f(H). Further, most (Sec. 2[b]). As well, the U.S. Forest Service has moved to
evaluations of biodiversity patterns and of imperiled or an “ecosystem management” basis for planning and
scarce species have focused only on mapping species management on national forests and grasslands.34 To
counts or evaluating the role of habitat on species’ presence conserve and manage ecosystems means understanding
and abundance. their dynamics and processes, including the ecological
In most of these worthy pursuits, community functions of species.
interactions and ecological roles of species—how
organisms change their environments by what they do— Key Ecological Functions of
have been largely absent. That is, as much as organisms Wildlife Species
are a function of their habitat, environmental conditions, The term key ecological functions (KEFs) refers to the
including habitat and resources available for other species, principal set of ecological roles performed by each species
are influenced by the ecological roles of organisms. This in its ecosystem. KEFs refer to the main ways organisms
chapter provides a foundation for explicitly considering use, influence, and alter their biotic and abiotic
the ecological roles and interactions of vertebrate wildlife environments. “Key” refers to the main roles played by
and for integrating those roles with the more traditional each species. As we show in this chapter, categories of
habitat-based WHR focus. KEFs can be depicted for each species and used in
multiple-species planning, biodiversity conservation, and
The Functional Foundation of Sustainable other facets of ecosystem analysis and management.
Ecosystems
Much has been written on sustainable management of Background and Theory
natural resources and conservation of wildlife populations
and natural ecosystems.6 Sustainability may be defined Expanding the WHR Paradigm
as resource use habits that do not outstrip the capacity of The explicit use of functional categories for evaluating
an ecosystem to produce desirable conditions and effects of land use planning on ecological communities
commodities, and often has been seen as a de facto appeared in the terrestrial ecology science assessments of
conservation goal. But what influences sustainability? the Interior Columbia Basin Ecosystem Management
Sustainable use of resources is made possible by ensuring Project.23, 24, 28 In that work, the traditional WHR paradigm
that rates of production are not exceeded by rates of loss was expanded by depicting the environmental and
plus extraction. 43 Further, natural ecosystems are functional influences on a species in a “species influence
sustainable only when their native biodiversity (the diagram” (Figure 1). The species influence diagram
variety of life and its processes) and the functional basis of illustrates how the distribution and abundance of a species
productivity are maintained. Ultimately, it is the set of (or species group) are influenced by key environmental
ecological roles played by its component organisms, correlates; that a species performs specific KEFs; that KEFs
including humans, that influences ecosystem biodiversity, in turn influence the biodiversity, productivity, and
productivity, and sustainability. sustainability (BPS) of the ecosystem; that management

168
 CHAPTER 6: ECOLOGICAL FUNCTIONS 169

Key environmental correlates Key ecological functions • rodents can serve to disseminate beneficial
mycorrhizal fungi in western U.S. forests;18 and
KEC1 KEF1 • reptilian primary burrow excavators can provide
KEC2 KEF2 for avian secondary burrow users.46
Species Many other examples can be found in the literature,
f1 abundance, f2 especially on ecological functions of plants and
fitness, or invertebrates.
viability (S) What is the manager to do with all this information?
The types and ramifications of ecological functions and
interactions in an ecosystem can be overwhelming. This
is why we have developed a structured classification of
Management activities, Biodiversity/ KEFs and provide here a way of considering functions
directives, prescriptions productivity/ when formulating and evaluating effects of land use
sustainability (BPS) management actions. Through the species influence
Figure 1. A generalized species influence diagram, showing diagram and use of the matrixes on the CD-ROM,
(1) the relations of species (S) to their key environmental management activities can be linked to habitat elements
correlates (KECs or habitat elements; f1 functions) and key and structures, and thence to species, their KEFs, and their
ecological functions (KEFs); (2) that species’ KEFs influence influence on ecosystem BPS.
ecosystem biodiversity, productivity, and sustainability (BPS;
f2 functions); and (3) that management activities can be based Ecological Functions Central to the Trophic
on BPS and can influence KECs and habitats, in turn Structure of Wildlife Communities
influencing the viability of species. The matrices on the CD- Ecological functions of organisms support the trophic
ROM provide data on each of these diagram elements. Arrows structure of ecosystems, that is, energy flows, food webs,
represent rates of influence among the elements. and nutrient cycling. More biodiverse systems support
wider arrays of ecological functions,16, 37 and thus might
support a broader or deeper trophic structure. Ecosystems
goals for BPS can help establish management guidelines; differ in their rates and patterns of primary production
and that management activities influence species’ key (photosynthesis by plants), which largely supports
environmental correlates. The term “key environmental vertebrate primary consumers and much of the rest of the
correlate” refers to wildlife habitats, habitat elements, and food web and trophic pyramid. Trophic structures differ
other nonhabitat influences on the distribution and among ecosystems in that different sets of vertebrate
abundance of organisms. species participate in various ways in providing major
We distinguish KEFs of species, which are the biotic and standing crop biomass, types and rates of consumption
ecological roles of organisms, from abiotic ecosystem and decay or recycling functions, energy transfer and
processes, which include weather disturbance events and nutrient cycling, and interspecific interactions including
geochemical cycles. In this chapter we focus on the link competition and predation. We provide some examples
between species and their environments through KEFs below.
and demonstrate how other components in this overall
species influence diagram can be used in evaluations. Building a Database of Species’ Key
Ecological Functions of Vertebrates Ecological Functions
It has long been recognized that the ecological roles of
vertebrate species influence ecosystems. Only recently,
Purpose
however, has this been integrated into theory and The purpose of building a matrix of KEFs of vertebrates
practice.7, 42, 44 Examples of some ecological functions of is to provide a consistent framework from which to
vertebrate species include how: consider the ecological roles of wildlife in the management
• browsing or grazing by ungulates can change plant of populations, habitats, and ecosystems. A related
communities;1, 3, 14, 41 purpose is to provide a means of posing working
• animals can act as “environmental engineers” and hypotheses of the ecological roles of wildlife and effects
influence geomorphology5 and ecosystem of management actions on those roles.
processes;17
Methods
• frugivores can support viable fruit-bearing plants;25
We developed the KEF matrix by first refining the existing
• pollinators can support plant diversity;36
hierarchical classification system of KEFs developed by
• seed dispersers8 and frugivores13 can influence
Marcot et al.,23 and then populating the KEF matrix by
forest succession and regeneration;
denoting the pertinent KEF categories for each species.
• carrion feeding can support the trophic structure of
Marcot et al.’s KEF classification was developed for use
a community;12
with soil microorganism groups, lichens, bryophytes,
• carnivore predation can influence populations of
vascular plants, and invertebrates, as well as vertebrates.
ungulate prey species;48
To maintain some similarity, we retained their major KEF
 170 Wildlife–Habitat Relationships in Oregon and Washington

categories and numbering system, trimmed out distribution that is slightly skewed to the right (Figure 2).
subcategories not pertinent to vertebrates, and expanded That is, few species are coded with very few, or very many,
some subcategories. As with the original system, our KEF categories, and there are slightly more species with
revised KEF classification is strictly hierarchical so that many KEF categories than predicted by a normal
one can query the matrix at various levels of specificity. distribution. The species with few KEF categories are
In populating the KEF species matrix, we started with functional specialists; like the condor and vulture, they
that of the Interior Columbia Basin Ecosystem perform only a very few functions within their ecosystems.
Management Project.21 We added pertinent wildlife species The species with many KEF categories tend to be functional
not included in that database, including coastal and generalists; they perform many functions. We discuss the
marine birds and mammals, and deleted species not found ecological implications of such functional patterns below.
in Washington and Oregon. We also reviewed published
literature, used our own experience and professional Patterns of Key Ecological Functions of
judgment, and sought peer review of the species or KEF
categories we added. We lacked data by which to describe
Vertebrate Species in Washington and
KEFs for a species for each wildlife habitat, habitat Oregon
element, or habitat structure individually. Following are some examples of various functional
patterns of wildlife communities in Washington and
Database Results Oregon.
The KEF classification system includes some 85 categories
of ecological functions, including major headings and Trophic Structure of Wildlife Communities
subheadings (Table 1). The 8 major headings are trophic The KEF matrix can be used to depict general trophic
relations, nutrient cycling, organismal relations, disease structures of communities by summing number of species
vectors, soil relations, wood structure relations, water according to their trophic and diet relations. In Washington
relations, and vegetation relations. and Oregon as a whole, 50% of all wildlife species are
The KEF matrix on the CD-ROM includes 598 wildlife primary consumers (herbivores), 87% are secondary
species out of the 755 species identified for the project area. consumers (primary predators), and only 1% are tertiary
The 157 species not included are considered “accidental” consumers (secondary predators), with bird species
or are subspecies of KEF-coded species. The KEF matrix playing a proportionally greater role across this spectrum
has 7,319 records, each record being a specific KEF (Figure 3). Other minor trophic categories include carrion-
category for a species. Thus, species average 12 KEF feeders (6%, mostly birds and mammals), cannibalistic
categories each (including KEF categories and feeders (1%, amphibians, birds, and mammals), and
subcategories, so there is some duplication in this count) coprophagous feeders (2%, all mammals).
and range from 3 (California condor and turkey vulture, The main (>50 species) primary-consumer categories
both carrion-feeder specialists, the former of only historical are spermivores (seed-eaters), frugivores (fruit-eaters),
occurrence), to 30 (American black bear) KEF categories. grazers (grass/forb-eaters), foliovores (leaf-eaters), and
As might be expected, the frequency of species by aquatic herbivores (Figure 4). Amphibians play the main
number of KEF categories roughly follows a normal role of feeding on plant material in water on decomposing

1.1.6 Coprophagous
Key ecological function

1.1.5 Cannibalistic
Number of species

1.1.4 Carrion feeder

1.1.3 Tertiary
consumer
1.1.2 Secondary
consumer
1.1.1 Primary
consumer
0 100 200 300 400 500
Number of species

Number of KEF categories Amphibians Reptiles Birds

Mammals Marine mammals
Figure 2. Frequency histogram of number of vertebrate
wildlife species in Washington and Oregon by number of
categories of key ecological functions (KEFs) they perform, as Figure 3. Trophic level functions of wildlife in Oregon and
denoted in the KEF matrix. Washington.
 CHAPTER 6: ECOLOGICAL FUNCTIONS 171

Table 1. The classification of key ecological functions of wildlife in Washington and Oregon.

1 Trophic relationships *
1.1 heterotrophic consumer (an organism that is unable to manufacture its own food and must feed on other organisms)*
1.1.1 primary consumer (herbivore; an organism that feeds primarily on plant material) (also see below under Herbivory) *
1.1.1.1 foliovore (leaf eater) *
1.1.1.2 spermivore (seed eater) *
1.1.1.3 browser (leaf, stem eater)
1.1.1.4 grazer (grass, forb eater)
1.1.1.5 frugivore (fruit eater) *
1.1.1.6 sap feeder
1.1.1.7 root feeders *
1.1.1.8 nectivore (nectar feeder)
1.1.1.9 fungivore (fungus feeder) *
1.1.1.10 flower/bud/catkin feeder
1.1.1.11 aquatic herbivore
1.1.1.12 feeds in water on decomposing benthic substrate (benthic is the lowermost zone of a water body)
1.1.1.13 bark/cambium/bole feeder
1.1.2 secondary consumer (primary predator or primary carnivore; a carnivore that preys on other vertebrate or
invertebrate animals, primarily herbivores) *
1.1.2.1 invertebrate eater
1.1.2.1.1 terrestrial invertebrates
1.1.2.1.2 aquatic macroinvertebrates (e.g., not plankton)
1.1.2.1.3 freshwater or marine zooplankton
1.1.2.2 vertebrate eater (consumer or predator of herbivorous or carnivorous vertebrates) *
1.1.2.2.1 piscivorous (fish eater) *
1.1.2.3 ovivorous (egg eater)
1.1.3 tertiary consumer (secondary predator or secondary carnivore; a carnivore that preys on other carnivores)
1.1.4 carrion feeder (feeds on dead animals)
1.1.5 cannibalistic (eats members of its own species)
1.1.6 coprophagous (feeds on fecal material)
1.1.7 feeds on human garbage/refuse
1.1.7.1 aquatic (e.g., offal and bycatch of fishing boats)
1.1.7.2 terrestrial (e.g., garbage cans, landfills)
1.2 prey relationships
1.2.1 prey for secondary or tertiary consumer (primary or secondary predator)
2 Aids in physical transfer of substances for nutrient cycling (C,N,P, etc.) *
3 Organismal relationships *
3.1 controls or depresses insect population peaks *
3.2 controls terrestrial vertebrate populations (through predation or displacement) *
3.3 pollination vector
3.4 transportation of viable seeds, spores, plants, or animals (through ingestion, caching, caught in hair or mud on feet, etc.) *
3.4.1 disperses fungi
3.4.2 disperses lichens
3.4.3 disperses bryophytes, including mosses
3.4.4 disperses insects and other invertebrates (phoresis)
3.4.5 disperses seeds/fruits (through ingestion or caching)
3.4.6 disperses vascular plants *
3.5 creates feeding, roosting, denning, or nesting opportunities for other organisms *
3.5.1 creates feeding opportunities (other than direct prey relations) *
3.5.1.1 creates sapwells in trees
3.5.2 creates roosting, denning, or nesting opportunities *
3.6 primary creation of structures (possibly used by other organisms) *
3.6.1 aerial structures (typically large raptor or squirrel stick or leaf nests in trees or on platforms, or barn swallow/cliff
swallow nests)*
3.6.2 ground structures (above-ground, non-aquatic nests and ends and other substrates, such as woodrat middens, nesting
mounds of swans, for example)*
3.6.3 aquatic structures (muskrat lodges, beaver dams)*
3.7 user of structures created by other species
3.7.1 aerial structures (typically large raptor or squirrel stick or leaf nests in trees or on platforms, or barn swallow/cliff
swallow nests)
 172 Wildlife–Habitat Relationships in Oregon and Washington

3.7.2 ground structures (above-ground, non-aquatic nests and ends and other substrates, such as woodrat middens, nesting
mounds of swans, for example)
3.7.3 aquatic structures (muskrat lodges, beaver dams)
3.8 nest parasite
3.8.1 interspecies parasite (commonly lays eggs in nests of other species)
3.8.2 common interspecific host (parasitized by other species)
3.9 primary cavity excavator in snags or live trees (organisms able to excavate their own cavities)
3.10 secondary cavity user (organisms that do not excavate their own cavities and depend on primary cavity excavators or natural
cavities)
3.11 primary burrow excavator (fossorial or underground burrows)
3.11.1 creates large burrows (rabbit-sized or larger)
3.11.2 creates small burrows (less than rabbit-sized)
3.12 uses burrows dug by other species (secondary burrow user)
3.13 creates runways (possibly used by other species; runways typically are worn paths in dense vegetation)
3.14 uses runways created by other species
3.15 pirates food from other species
3.16 interspecific hybridization (species known to regularly interbreed)
4 Carrier, transmitter, or reservoir of vertebrate diseases
4.1 diseases that affect humans *
4.2 diseases that affect domestic animals
4.3 diseases that affect other wildlife species
5 Soil relationships *
5.1 physically affects (improves) soil structure, aeration (typically by digging) *
5.2 physically affects (degrades) soil structure, aeration (typically by trampling) *
6 Wood structure relationships (either living or dead wood) *
6.1 physically fragments down wood *
6.2 physically fragments standing wood *
7 Water relationships *
7.1 impounds water by creating diversions or dams *
7.2 creates ponds or wetlands through wallowing
8 Vegetation structure and composition relationships *
8.1 creates standing dead trees (snags) *
8.2 herbivory on trees or shrubs that may alter vegetation structure and composition (browsers)
8.3 herbivory on grasses or forbs that may alter vegetation structure and composition (grazers)

* Key ecological functions of Homo sapiens.

1.1.1.13 Bark/cambium/bole
1.1.1.12 Benthic substrate
1.1.2.3 Ovivorous
1.1.1.11 Aquatic herbivore
1.1.2.2.1 Piscivorous
1.1.1.10 Flower/bud
1.1.1.9 Fungivore
1.1.2.2 Vertebrate eater
1.1.1.8 Nectivore
1.1.1.7 Root feeder
1.1.2.1.3 Zooplankton
1.1.1.6 Sap feeder
1.1.1.5 Frugivore 1.1.2.1.2 Aquatic macroinvertebrates
1.1.1.4 Grazer
1.1.1.3 Browser 1.1.2.1.1 Terrestrial invertebrates
1.1.1.2 Spermivore
0 100 200 300 400
1.1.1.1 Foliovore
0 50 150 200 Number of species
Number of Species
Amphibians Reptiles Birds
Amphibians Reptiles Birds Mammals Mammals Marine mammals

Figure 4. Primary consumption (herbivory) functions of Figure 5. Secondary consumption (carnivory) functions of
wildlife in Washington and Oregon. wildlife in Washington and Oregon.
 CHAPTER 6: ECOLOGICAL FUNCTIONS 173

benthic substrates. Birds play main roles as frugivores, Organismal Relations Within
sap feeders, nectivores (nectar-feeders), and flower, bud, Wildlife Communities
and catkin feeders. Mammals play main roles as browsers, Some 26 categories of organismal relations within wildlife
grazers, root feeders, and fungivores (fungus-feeders). communities can be evaluated with the KEF matrix.
Both birds and mammals participate more equally as Patterns presented here are for all communities in
foliovores and spermivores. Washington and Oregon (Figures 6-8); patterns among
The main (>100 species) secondary consumer categories specific wildlife habitats may differ.
are feeders on terrestrial invertebrates, vertebrates, aquatic A number of examples of symbiosis can be
macroinvertebrates, and fish (piscivores) (Figure 5). demonstrated. Seven species of birds (hummingbirds and
Amphibians participate in all secondary consumer orioles) serve as pollination vectors for plants. Among
categories, but proportionally are the greatest consumers terrestrial vertebrates, mammals are the sole dispersers
of freshwater invertebrates. Birds proportionately of fungi and lichens; among terrestrial vertebrates,
dominate all categories except that of consuming primarily birds disperse invertebrates and vascular plant
freshwater invertebrates. Mammals participate in most parts; and both birds and mammals disperse seeds and
categories with their greatest proportionate representation fruits (Figure 6). Birds create sapwells for feeding by other
as terrestrial invertebrate-feeders, vertebrate feeders, and species. Both birds and mammals create roosting, denning,
ovivores (vertebrate egg-eaters). Tertiary consumers or nesting structures in aerial, ground, and aquatic
include a few birds and mammals only. situations that other amphibian, reptile, bird, and mammal
These patterns differ among native wildlife habitats. species also use (Figure 7).
For example, avian primary consumers tend to be most Three bird species act as nest (brood) parasites
numerous (total number of species and proportion of all (although only the brown-headed cowbird regularly
species) in both eastside (east of the Cascade Mountains parasitizes nests of other species within the Oregon-
crest) and westside wetlands and in dry forest types, Washington area) and 58 bird species serve as their hosts.
whereas avian secondary consumers are most numerous Eighteen birds and 1 mammal species act as primary cavity
in westside (wetland, forest, woodland, and riparian) excavators, serving 32 bird and 24 mammal secondary
types. Mammalian primary consumers are most numerous cavity-using species. Twenty-one mammal species dig
in higher elevation or eastside grassland, shrubland, or large (rabbit size) burrows and 9 bird and 53 mammal
forest types, whereas mammalian secondary consumers species dig small burrows, which also provide for 7
are most numerous in westside types (riparian, wetland, amphibian, 14 reptile, 1 bird, and 59 mammal secondary
grassland, woodland, and forest types alike). Much of this burrow-using species. Forty-six mammal species create
can be explained by the kinds of climates and vegetation runways in terrestrial vegetation that also can be used by
physiognomies present in various geographic locations, 2 amphibian and 41 other mammal species (Figure 8).
such as the dry eastside rainshadow effects where Managers might wish to determine which species of
grasslands, shrublands, and grazing and browsing such symbiotic, organismal relations occur in the wildlife
mammals (mammalian primary consumers) are more habitats and communities under their charge to ensure
common. that the habitat requirements of all members are duly
Understanding such trophic patterns and their addressed, and that full ramifications of modifying habitat
biogeographic relations may be important to managers at for one set of species on their symbiotic partners are
a broad scale, as land ownership patterns and acknowledged.
management activities might affect habitats differently,
favoring one trophic category over another in different Soil, Wood, Water, and Vegetation Relations
areas. For example, the two anthropogenic (human- Some examples of soil, wood, water, and vegetation
created) habitats—Agriculture, Pastures, and Mixed relations are shown in Figure 9. Seventy-five species of
Environments, and Urban and Mixed Environments— amphibians, reptiles, birds, and mammals may help
provide poorly for mammalian secondary consumers. improve or maintain soil structure and aeration through
Comparing historical to current habitat patterns may tunneling, digging, and turning over soil. This also has
reveal broad-scale and longer-term shifts in trophic the benefit of increasing aerobic bacterial decay in soils
structures of wildlife communities, which in turn can and incorporating coarse decaying wood as organic matter
disclose differential repercussions on other ecosystem into the soil. One bird and 20 mammal species physically
functions and biodiversity patterns. It may be of interest help fragment down wood, and 1 bird and 1 mammal
to compare current and potential future conditions to species fragment standing wood; these functions, too, help
extrapolated historical conditions, to see if and how hasten the wood decay process.
wildlife communities have changed in terms of their Seven mammal species, including 2 elk subspecies,
trophic patterns and to help set broad-scale planning impound water by creating diversions or dams, or create
objectives. Such objectives might include restoring or ponds or wetlands through wallowing. Such ponds and
maintaining all historical trophic categories by geographic wetlands can then provide for a wide array of other
area or habitat, or mitigating the adverse reduction of wildlife species; the habitat elements database lists 242
trophic categories in anthropogenically disturbed species of amphibians, reptiles, birds, and mammals as
environments. being associated with ponds and wetlands. Three
 174 Wildlife–Habitat Relationships in Oregon and Washington

3.4.6 Disperses vascular 8.3 Grazing alters vegetation structure

plants 8.2 Browsing alters vegetation structure
3.4.5 Disperses seeds/fruits 8.1 Creates snags
3.4.4 Disperses invertebrates
7.2 Creates ponds/wetlands
3.4.3 Disperses bryophytes
7.1 Impounds water
3.4.2 Disperses lichens

3.4.1 Disperses fungi 6.2 Fragments standing wood

3.3 Pollination vector 6.1 Fragments down wood
3.2 Control vertebrate
populations 5.2 Degrades soil structure & aeration
3.1 Control insect 5.1 Improves soil structure & aeration
populations
0 50 100 0 20 40 60 80
Number of species Number of species

Amphibians Reptiles Birds Mammals Amphibians Reptiles Birds Mammals

Figure 6. Organismal functional relations of wildlife in

Figure 9. Soil, wood, water, and vegetation functional
Washington and Oregon: population control, and pollination
relations of wildlife in Washington and Oregon.
and dispersal vectors.

3.7.3 Uses aquatic structures mammal species create snags through shredding, digging
3.7.2 Uses ground structures
into, or girdling of trees. The habitat elements matrix lists
131 species of amphibians, birds, and mammals as being
3.7.1 Uses aerial structures
associated with snags. Browsing by 10 mammal species
and grazing by 1 bird and 7 mammal species can alter
3.6.3 Creates aquatic structures vegetation structure and composition. This in turn can
3.6.2 Creates ground structures change habitat conditions for a wide array of other wildlife
3.6.1 Creates aerial structures associated with grass, forb, and shrub species and cover.

3.5.2 Creates roost/den/nest Taxonomy and Definitions of

3.5.1.1 Creates sapwells Functional Patterns
0 5 10 15 20 25 30 Beyond the simple species tallies discussed above, what
Number of species are the functional characteristics of ecosystems that can
be evaluated by using the KEF matrix? We suggest a
Amphibians Reptiles Birds Mammals
taxonomy of 15 types of patterns of ecological functions
(Table 2, follows Literature Cited at end of chapter) and
Figure 7. Organismal functional relations of wildlife in
provide some examples here of using the matrix for
Washington and Oregon: creation and use of feeding,
evaluating them. Some of these patterns are discussed in
breeding, and resting structures.
the literature but many we pose here for the first time as
working hypotheses of how wildlife communities operate
3.14 Uses runways
3.13 Creates runways
and can be evaluated. All of the types of patterns need
empirical study to determine actual rates of influence of
3.12 Secondary burrow user
the functional roles of animals.
3.11.2 Creates small burrows
3.11.1 Creates large burrows
Patterns to Watch
Community patterns. Community patterns of KEFs in
3.10 Secondary cavity user wildlife communities include functional redundancy,
3.9 Primary cavity user functional richness, total functional diversity, functional
webs, functional profiles, and functional homologies.
3.8.2 Interspecific host Functional redundancy is the number of wildlife species
3.8.1 Interspecific nest
performing a specific ecological function, and is
parasite
0 20 40 60 80 determined by tallying species in the KEF matrix for
Number of species specific KEF categories for specific habitats conditions.
Functional richness is the number of KEF categories in a
Amphibians Reptiles Birds Mammals community. Total functional diversity is functional richness
weighted by functional redundancy,4 analogous to species
Figure 8. Organismal functional relations of wildlife in diversity.
Washington and Oregon: nest parasitism and hosts, and A number of wildlife habitats in Washington and
cavity, burrow, and runway excavation and use. Oregon are more or less equally functionally rich except
 CHAPTER 6: ECOLOGICAL FUNCTIONS 175

Oceanic
Marine shelf
Marine nearshore
Inland marine deeper waters
Bays and estuaries
Coastal headlands & islets
Coastal dunes & beaches
Eastside riparian wetlands Figure 10. Total functional
Montane coniferous wetlands richness (number of key ecological
Historical occurrence—Westside riparian wetlands
Westside riparian wetlands
function categories) by wildlife
Herbaceous wetlands habitat in Washington and
Open water—lakes, rivers, & streams Oregon.
Urban
Agriculture & pastures
Desert playa & salt scrub shrublands
Dwarf shrub-steppe
Shrub-steppe
Eastside grasslands
Eastside canyon shrublands
Western juniper & mountain-mahogany woodlands
Ceanothus-manzanita shrublands
Westside grasslands
Alpine grasslands & shrublands
Subalpine parkland
Upland aspen forest
Ponderosa pineforests & woodlands
Lodgepole pine forest & woodlands
Eastside mixed conifer forest
Montane mixed conifer forest
Southwest Oregon mixed conifer-hardwood forest
Westside oak & dry Douglas-fir forest & woodlands
Westside lowlands conifer-hardwood forest

0 20 40 60

for the less rich marine and coastal habitats (Figure 10). conservation attention on the latter and ensure the
Agriculture, Pastures, and Mixed Environments, and continued integrity of the former.
Urban and Mixed Environments are the most functionally A functional web is the full array of all KEFs associated
rich, but the matrixes might overstate the viability of with a set of species that may be specified by some habitat
wildlife species within these anthropogenic environments. element or structure. An example given in Morrison et
Among native wildlife habitats, a number of forest, al.28 from the interior Columbia Basin illustrates an array
woodland, and wetland environments rank as the most of 22 KEF categories just of mammals associated with
functionally rich. down wood in eastside ponderosa pine forest
Total functional diversity differs more among wildlife communities. Many of these functions, such as dispersal
habitats in Washington and Oregon than does functional of fungi or primary burrow excavation, extend well
richness, although it follows much of the same ordinal beyond the confines of down wood substrates per se. This
pattern (Figure 11). An exception is with various alpine demonstrates the concept of how functions supported in
and shrubland wildlife habitats that have low functional part by specific habitat elements can influence parts of
diversity. This means that even though a relatively high the ecosystem well beyond those habitat elements.
number of categories of KEFs are represented in alpine Marcot et al.24 provided functional profiles, or graphs
and shrubland wildlife communities, there is lower showing functional redundancy for each KEF category,
functional redundancy (fewer wildlife species) performing and of total functional diversity of all KEF categories,
many of those functions than found in forest, woodland, across all vegetation communities of the interior Columbia
and wetland communities; there are far fewer still in Basin. The concept of functional homologies, or how
marine and coastal communities. This may imply a greater communities compare in their functional categories and
sensitivity of alpine, shrubland, coastal, and marine redundancies, is new but can be easily evaluated using
communities to change than with forest, woodland, and the KEF database (Table 2) and similarity analyses.
wetland communities. Managers may wish to evaluate functional webs, profiles,
This illustrates the premise that functionally redundant, and homologies to help describe the unique functional
rich, and diverse communities may be more resistant or roles of communities in their charge.
resilient to perturbations20, 29, 33 and can support greater
Geographic Patterns. Species’ range maps or distributions
levels of biodiversity45 than less functionally redundant,
by wildlife habitats can be used to depict geographically
rich, or diverse communities. Managers may wish to
many of the functional patterns discussed here. This opens
develop functional profiles (see below) not just of local
the door to a new way to display functional patterns
wildlife habitats but also of structural stages and
spatially, and may be termed “geographic functional
combinations of habitat elements. Then they can identify
ecology”. For example, stacking the range maps of species
those communities that are naturally rich, diverse, and
with a particular KEF category produces a map of
redundant in their ecological functions, and those that are
functional redundancy. Areas of lowest redundancy that
not, to help prioritize further evaluations and to focus
serve to link areas of high redundancy can be delineated
 176 Wildlife–Habitat Relationships in Oregon and Washington

Oceanic
Marine shelf
Marine nearshore
Inland marine deeper waters
Bays and estuaries
Coastal headlands & islets
Figure 11. Total functional Coastal dunes & beaches
diversity (functional richness Eastside riparian wetlands
Montane coniferous wetlands
times mean number of species per Historical occurrence—Westside riparian wetlands
function) by wildlife habitat in Westside riparian wetlands
Washington and Oregon. Herbaceous wetlands
Open water—lakes, rivers, & streams
Urban
Agriculture & pastures
Desert playa & salt scrub shrublands
Dwarf shrub-steppe
Shrub-steppe
Eastside grasslands
Eastside canyon shrublands
Western juniper & mountain-mahogany woodlands
Ceanothus-manzanita shrublands
Westside grasslands
Alpine grasslands & shrublands
Subalpine parkland
Upland aspen forest
Ponderosa pine forests & woodlands
Lodgepole pine forest & woodlands
Eastside mixed conifer forest
Montane mixed conifer forest
Southwest Oregon mixed conifer-hardwood forest
Westside oak & dry Douglas-fir forest & woodlands
Westside lowlands conifer-hardwood forest

0 10 20 30 40

as functional bottlenecks or cold spots, just as areas of highest feed on different kinds of carrion in different habitats and
redundancy can be identified as functional linkages or hot on different substrates than does the turkey vulture. Also,
spots. In this way, many other aspects of KEFs can be there may be fewer carrion-feeding wildlife species in any
mapped. It may be of interest and importance to managers given wildlife habitat; this is determined by linking the
to map such geographic functional patterns, to identify KEF database to the wildlife habitats database. For
areas and communities worthy of greater conservation example, there are only 17 carrion-feeding species found
attention. in Westside Lowland Conifer-Hardwood Forest. In fact,
the fewest number (8 species) occurs in Inland Marine
Species’ Functional Roles. Two major patterns of the
Deeper Waters and Marine Nearshore habitats, but the
functional roles of species are functional keystones and
turkey vulture does not occur in either of these habitats.
functional specialization. Functional keystones are species
Where it does occur, the carrion-feeding function is shared
whose removal would most alter and degrade the
by other species. Therefore, although the turkey vulture
structure or function of the community. If the species is
is a functional specialist, its function is usually performed
the only one that performs a particular ecological function
by at least some other species, so, although specialized,
within a community, it is a critical functional link species.
its carrion-feeding role, in general, is not entirely a
The function it performs is then a critical function.
keystone role. It may be a keystone role, however, for
Functional breadth is the number of ecological functions
specific kinds of carrion on specific substrates or in specific
performed by a species. When a species performs very
seasons; there are many dimensions to a species’ niche.
few functions, it is a functional specialist.
Some KEFs, however, have few participants, even if
Functional keystones, functional link species, and
the associated species are not functional specialists. An
functional specialists can be identified by querying the
example is the American beaver, which is the only species
KEF database. For example, the turkey vulture was
coded in the database with KEF code 7.1. “impounds water
identified above as being an extreme functional specialist
by creating diversions or dams”. So for this function, this
(it performs the fewest total functions). Its function is KEF
species is a critical functional link and a keystone.
category 1.1.4 (see Table 1), which means that it is a
American beaver is also coded with 24 other KEF
heterotrophic consumer and, more specifically, a carrion
categories (these include subcategories as well), so it is
feeder. It has no other trophic categories, which means
not much of a functional specialist . However, it is the only
that it is a specialist in this function. Loss of carrion means
species that performs KEF 7.1, which is a function that
loss of this species.
influences many other species. In fact, the habitat elements
But is the reverse also true, that is, is the turkey vulture
database lists 49 other wildlife species directly and
critical to the carrion-feeding function? This is determined
positively associated with habitat element 2.3, “Beaver/
by identifying the functional redundancy of that function.
muskrat activity, including dams, lodges, and ponds” in
Querying the KEF database shows that there are 39 other
Washington and Oregon.
wildlife species also coded as carrion feeders in
The lesson for the manager here is to pay attention to
Washington and Oregon. Granted, many of these species
both the array of functions by species, as well as the array
 CHAPTER 6: ECOLOGICAL FUNCTIONS 177

of species by function. The concept of a keystone species 5. Functional breadth is not necessarily related to
or keystone function has many dimensions and potential critical functional links. That is, species, which are
management implications. We caution against careless use the sole (or few) performer(s) of a given function
of the general term “keystone” without clarifying its in a community might also perform very few, or
context and significance.26 For example, species that are many other, functions.
functional specialists are probably highly vulnerable to 6. Critical functional links cannot be predicted from
changes in their environment if they depend on that functional breadth.
function for survival. A species that is a functional Of course, working hypotheses also can be stated as
generalist with broad functional breadth, that is, that null hypotheses for the purpose of critical testing of their
performs the widest array of functions-might greatly veracity. For example, the first working hypothesis can
influence an ecosystem upon its removal, but this depends be stated as the following null hypothesis: there is no
on what and how other species also perform such influence on functional resistance of a community from
functions. A function with one or few species may itself degree of functional switching of its component species.
be vulnerable and of management interest for ensuring Then, an appropriate research study can be devised that
its continuance in an ecosystem. compares the functional constitution of communities
before and after a disturbance, for communities that differ
Functional Responses of Communities. The categories
in their degree of functional switching of their component
discussed above can be used to characterize the functional
species.
responses of communities. Community functional
responses refer to how communities respond functionally
to disturbances. Responses can be categorized as Lost Functions of Extirpated Species
functional resilience, resistance, attenuation, and shifting, Thirteen species are recognized in this effort as extirpated
and imperiled functions. from either Oregon and/or Washington (see Chapter 18).
Functional resilience is the capacity of a community to Extirpated species performed 49 ecological functions
rebound to a starting functional state after a perturbation, (including subcategories) that were potentially reduced
such as a natural or human-caused disturbance event. or eliminated as a result of extirpation. Because ecosystem
Functional resistance is the degree of inertia of a community sustainability is dependent on the maintenance of
to retain its initial functional state and not change under biodiversity and native ecological functions, the loss of
stressors. Functional attenuation is the degree to which a these species may have affected the functional integrity
community loses functions after a disturbance, and of the communities in which they formerly occurred. The
functional shifting is the difference in functions before and degree of such influence, however, is largely unstudied.
after disturbance when the community has finally
stabilized in species composition. Imperiled functions are Key Ecological Functions of Extirpated
those functions that are represented by few species that Species in Oregon and Washington
are themselves at risk. Trophic Relations. The 13 extirpated species varied widely
in their degree of functional specialization, with most
Some Hypotheses on Ecological Functions species performing >1 trophic-related KEF. The grizzly
Ecological implications of functional patterns of species bear, for example, had the widest functional breadth of
and communities, as discussed above, can be taken as trophic KEFs (7) among the extirpated species, whereas
testable hypotheses about the roles of wildlife and how California condor was a functional specialist (1), feeding
ecosystems work (Table 1). Some additional hypotheses exclusively on carrion. The upland sandpiper and yellow-
include the following: billed cuckoo also were narrow in their trophic relations,
1. Functional switching imparts functional resistance: feeding solely on terrestrial invertebrates and serving as
a greater degree of functional switching by species prey for secondary or tertiary consumers. Overall, trophic
imparts a greater functional resistance to the relations of extirpated species included spermivores
community. (grizzly bear, Wyoming ground squirrel, trumpeter swan,
2. Functional redundancy imparts resilience: for a and sharp-tailed grouse), grazers (pronghorn, bison,
particular function, the higher the functional bighorn sheep, and trumpeter swan) frugivores (gray wolf,
redundancy, the greater is the functional resilience grizzly bear, and sharp-tailed grouse), root-feeders (grizzly
(or resistance) for that function. bear), and foliovores (trumpeter swan and sharp-tailed
2. Functional diversity imparts low attenuation: the grouse). Eight species were secondary consumers,
greater the functional richness and diversity of a including 4 species that consumed vertebrates; 1 species
community, the lower is the degree of functional was a tertiary consumer (gray wolf); 2 were carrion-
attenuation (loss of functions). feeders; 1 was cannibalistic, and 9 were prey for other
3. The more functionally diverse a community, the species.
greater is its productivity and its biodiversity.
Organismal Relations. The 13 extirpated species
4. The greater the functional redundancy, the more
performed 6 categories of KEFs related to organismal
sustainable is the set of resources that the function
relations. Three mammal species were primary burrow
provides.
excavators; 3 mammals were creators or users of runways;
 178 Wildlife–Habitat Relationships in Oregon and Washington

2 mammals potentially controlled vertebrate populations; the extirpation of wolves, coyotes increased dramatically.27
2 birds and 3 mammals dispersed insects and/or seeds; Studies currently are assessing the impacts and dynamics
and 2 mammals created feeding opportunities for other of wolf recolonization on coyotes in the Greater
species. Other functions of extirpated species included Yellowstone Ecosystem.
carriers of disease (4 mammals), and functions related to
Provision of Feeding Opportunities. The grizzly bear,
soil (1 mammal), wood (3 mammals), and water relations
now rare in Washington and extirpated from Oregon,
(1 mammal).
performed the function of providing carrion in 11 habitats.
To assess the potential impact of losing these functions,
The species function profile for the KEF “provides feeding
we calculated functional redundancy by tallying the
opportunities” revealed that 6 other species (minus the
number of species with selected KEFs by wildlife habitat.
gray wolf, which was also extirpated from Oregon) also
These species function profiles depict the degree of
perform this function. However, only 1, the mountain lion
functional redundancy among habitats, thus serving to
performs this KEF in essentially the same manner, by
identify imperiled or lost functions. From among the set
providing carrion. It does so in all 12 of the habitats that
of KEFs performed by each extirpated species, we selected
the grizzly bear once occupied. This is an example of a
for analysis those functions that might be expected to vary
potentially imperiled function. Although this function was
among communities, as we discuss next.
not completely lost, the elimination of grizzly bears and
Carrion-feeding. Feeding exclusively on carrion, the gray wolves from their historical range may have greatly
California condor was an extreme functional specialist. constricted the occurrence and rate of this carrion-
Historically, it inhabited the Westside Grasslands, providing function. Because providing carrion is a
Westside Riparian-Wetlands, and Coastal Dunes and symbiotic, organismal relation, other species that use
Beaches of Oregon and Washington. Querying the carrion might be affected by the scarcity of this function.
matrices shows that 24 individual species perform this
Creation of Ponds or Wetlands Through Wallowing.
function in these 3 habitats. Twelve species perform this
Wallowing bison likely once created small ponds and
function in Westside Grasslands, 20 in Westside Riparian-
wetlands in 8 habitats in Oregon and Washington. Four
Wetlands, and 16 in Coastal Dunes and Beaches. Similar
ungulate species currently perform this function in subsets
to the turkey vulture, the condor was not a keystone
of these habitats. Of the 4 species, the Rocky Mountain
species for the maintenance of the carrion-feeding
elk is the only ungulate that can create small ponds and
function. Nonetheless, functional specialists may be highly
wetlands in all 8 of the habitats formerly occupied by
sensitive to changes in their environment, thus increasing
bison, and it is the only ungulate that performs this
their vulnerability to extinction or extirpation.31 The
function in Lodgepole Pine Forests and Woodlands.
extreme degree of specialization (for carrion as well as
Although not imminent, extirpation of the Rocky
perhaps other needs) of the California condor may have
Mountain elk from this habitat would result in the loss of
contributed to its demise.
this potentially imperiled function.
Control of Vertebrate Populations. This KEF was The Rocky Mountain elk, therefore, is a critical
examined for redundancy within the range of habitats functional link species for the critical function of creating
historically inhabited by gray wolves in Oregon. Thirty small ponds and wetlands in Lodgepole Pine Forests and
other species potentially perform this function, including Woodlands. Twenty wildlife species that occur in this
1 amphibian, 1 reptile, 16 birds, and 11 mammals habitat are associated with ephemeral pools, including 8
(excluding the grizzly bear, also extirpated from Oregon). species of amphibians. Interestingly, 4 species of frogs that
Obviously, not all of these species perform this KEF in occur in Brazil breed only in peccary wallows and other
comparable ways. Among the assemblage of mammals small permanent ponds; conserving these amphibians
associated with this function, coyotes, mountain lions, and necessitates preserving the wallowing function of
black bears might perform this KEF similarly to gray peccaries.40 This is an example of how the loss of one
wolves, although they may selectively prey on other species (bison or peccary) can influence the distribution
species. of another.
Importantly, the loss of top predators from a We found no evidence that the extirpation of the
community may have reverberating effects on the rest of pronghorn, Wyoming ground squirrel, sea otter, or upland
the community. Predators lost from a complex food web sandpiper resulted in the loss or imperilment of any
“release” their herbivore prey, thus potentially leading to specific ecological function in their former habitats. The
significant changes in vegetation structure, cover, and trophic KEF categories of these species (e.g., grazer,
species from increases of the prey population.31 In the spermivore, invertebrate-eater) were and are shared by
Greater Yellowstone Ecosystem, for example, the other species; thus the habitats they occupied were highly
extirpation of wolves led to an unprecedented growth in redundant for these functions. However, we recognize that
the moose population, which had profound effects on the level of specificity in our database is insufficient to
willow communities, and consequently, on neotropical characterize the complete set of ecological functions for
migrants associated with those communities (Berger et al., every species; simply performing the same general KEF
unpubl. data). Furthermore, wolves are dominant over in the same habitat does not necessarily confer
coyotes and aggressively exclude or displace them. With substitutability. At some level, one species is never
 CHAPTER 6: ECOLOGICAL FUNCTIONS 179

completely interchangeable with another, regardless of feed on carrion (2 reptiles and Virginia opossum), 2
functional similarity. mammals are coprophagous, and 23 are prey for
Evaluating the KEFs of extirpated species may help secondary or tertiary consumers.
land managers understand the influences that historically Twenty-three exotic species participate in organismal
contributed to ecosystem function. From this relationships including the potential control of terrestrial
understanding, restoration goals can then be formulated vertebrate populations (bullfrog), and transportation of
to restore lost functions and to protect rare ones. Species seeds (9 birds and 4 mammals). Two exotic squirrel species
that perform functions with low functional redundancy create structures that are used by other species, and 3 exotic
in a particular habitat could receive priority to ensure species use structures created by other species. Exotic
perpetuation of that function in the community. Because secondary cavity users, the eastern fox squirrel, eastern
most species are involved in several to many trophic, gray squirrel, English or house sparrow, and European
organismal, and symbiotic relations, the conservation of starling, compete for cavities in their respective habitats
overall biodiversity can surely lend to community stability. with up to 40 native species. Declines of Lewis’
By minimizing species loss, the integrity of ecosystem woodpecker and purple martin have been attributed, in
function may be maintained.45 Understanding how species part, to competition with starlings for cavities. Three exotic
contribute to community function therefore may be mammals are primary burrow excavators and 7 are
imperative to crafting “ecosystem management” secondary burrow users, with 2 exotic rats (Norway rat
guidelines and defining ecosystem restoration objectives. and black rat) alone competing with up to 51 native
Also, KEF analyses can be used to formulate and test secondary burrow users. Five exotic mammal species
hypotheses related to the effects of function loss on create runways and 5 other exotic mammal species use
community stability and ecosystem integrity. The manager runways created by others.
can apply this approach at a variety of scales. That is, a Exotic species can affect gene pools (e.g., eastern
species need not go extinct in a two-state area before its cottontail hybridizes with native species) and spread
more local reduction or extirpation of its populations and disease (at least 5 exotic mammal species carry diseases
associated functions are addressed. that may affect humans). Soil-related KEFs include
potentially physically improving soil structure by digging
Competing Functions of Exotic Species or burrowing (4 exotic mammals and 1 exotic reptile) and
Thirty-one non-native (exotic) species occur in Oregon and physically degrading soil structure by compaction (wild
Washington, including 2 amphibians, 3 reptiles, 14 birds, burro, feral horse). Three exotic species fragment down
and 12 mammals (see Chapter 16). Collectively, they wood (wild burro, feral horse, and feral pig) and the same
perform 54 KEFs in 30 wildlife habitats, and thus exotic 3 species can create small ponds or wetlands through
species are able to exploit niches occupied by native wallowing. Lastly, the eastern cottontail, feral pig, and
animals. The most successful non-native species have wild burro can alter vegetation structure and composition
KEFs that confer a competitive advantage, often to the through grazing.
detriment of native species. Exotic species threaten Understanding how exotic species compete with native
biodiversity by displacing native species either directly wildlife may help land managers predict which native
through predation or displacement, or indirectly through species are most likely to be affected, and in which habitats.
the alteration of ecosystems, and have been described as Although the mechanisms by which systems resist
one of the greatest threats to native species and human- introductions are not well understood, they seem to be
disturbed ecosystems in the world.35 related, in part, to trophic complexity.31 Introductions of
The exotic species found in Oregon and Washington more trophically generalized mammals to species-poor,
compete with native species through trophic and trophically simple systems (e.g., islands) have had extreme
organismal relations; soil, wood, and water relations; and effects on the native species. Conversely, trophically
by altering vegetation structure, and composition. Trophic complex, species-rich systems may be more resilient.
relations involve competition for food or predation. Most Maintaining the functional web, or full set of KEFs, within
exotic primary consumers are foliovores (9 bird species a community may, therefore, increase resistance or
and 2 mammals), spermivores (12 bird species and 6 resilience to the effects of exotic species whether
mammals), and frugivores (8 bird species and 5 mammals). introduced intentionally or not.
Seven exotic species are grazers, including the wild burro
and feral horse, which compete with native ungulates for The Functional Roles of Marine Species
forage. Seven exotic species are aquatic herbivores, and a
few are root feeders, browsers, flower feeders, and Marine Birds
decomposing benthic substrate feeders (2 species each). There are 83 species of coastal and pelagic birds associated
None of the exotic species consumes sap, nectar, or bark. with the 4 oceanic wildlife habitats: Inland Marine Deeper
Of the 25 exotic species that are secondary consumers, Waters, Marine Nearshore, Marine Shelf, and Oceanic
most consume invertebrates (24 species) and 12 prey on habitats. Collectively, they comprise 53 KEF categories.
vertebrates, including 3 exotic species that feed on fish They are all secondary consumers; 13 are also primary
(bullfrog and 2 reptiles) and 2 species that are ovivorous consumers, mostly including aquatic herbivores, but a few
(Eastern fox squirrel and Virginia opossum). Three species (mostly waterfowl) are also foliovores, spermivores,
 180 Wildlife–Habitat Relationships in Oregon and Washington

browsers, and grazers. Many consume invertebrates (76 So, what is it about the ecological functions of humans
bird species) and/or vertebrates (63 bird species), that so greatly influence native ecosystems and wildlife
including fish (60 bird species) and eggs (12 bird species). communities? It is not that humans play any critical
Seventeen bird species (skua, eagle, gulls, and jaegers) functional roles, or that humans are functional specialists.
pirate food from other species, an adaptation found much Rather, it seems to be our amazing functional breadth—
less in terrestrial habitats; 13 species (waterfowl, fulmar, that is, the number of functions we perform and our
eagle, gulls, storm-petrels, and shearwaters) are carrion resounding influence on the rest of the functional web of
feeders; and 12 species (skua, fulmar, gulls, albatrosses, natural communities. Such functional influences should
shearwaters, and jaegers) feed on human garbage or refuse be considered when evaluating effects of human activities.
in the ocean, principally offal and bycatch of fishing boats. Managers, in fact, may wish explicitly to include the KEF
Some help disperse invertebrates (28 bird species) or plants categories of humans in their evaluations of functional
(25 bird species). patterns of wildlife species and communities.
Because most coastal and pelagic bird species nest on
stacks, sea cliffs, and other coastal substrates, and some Management Implications
are fossorial and nocturnal, there are 7 species (guillemot, Throughout the text above, we have offered some ideas
auklets, kingfisher, puffin, and storm-petrels) that are on how managers might use this new approach to describe
primary excavators of small burrows and 11 species the functional patterns of wildlife species and
(grebes, 1 eagle, 1 gull, 1 tern, 1 pelican, and osprey) that communities and influences of human activities. More
create structures used by other species. There are even 4 specifically, the manager can link the KEF matrix with the
secondary cavity users (waterfowl) and 3 species other matrixes, especially those depicting wildlife habitats,
(cormorants) that can kill trees and create snags. Again, habitat elements, habitat structures, and management
many of these are examples of how species’ ecological roles activities.
extend beyond just the realm of their primary habitat, in For example, one could begin by specifying a particular
this case, coastal and open ocean. management activity, such as using herbicides in
shrubland or grassland management (Management
Marine Mammals
Activities 5c; see Chapter 20) within a particular wildlife
The matrix includes 18 species of marine mammals.
habitat such as Eastside Grasslands (Wildlife Habitat 15;
Collectively, they comprise some 20 KEF categories. All
see Chapter 20). Linking the management activities,
18 species consume aquatic invertebrates and 3 (whales)
wildlife habitats, and habitat elements matrixes, one can
consume marine zooplankton. Seventeen species consume
determine which wildlife species are potentially
vertebrates (15 of these eat fish) and 7 serve as prey for
influenced by this activity in this habitat, that is, which
other predators. Four whale species create feeding
species are coded for using habitat elements that are
opportunities for other species, and a number of species
affected by this activity in this wildlife habitat. In this case,
aid in nutrient cycling and carry or transmit disease,
it is a set of 2 amphibians, 7 reptiles, 75 birds, and 44
although these functions are poorly studied in marine
mammals.
environments.
Next, one can link this species set to their KEFs and
determine which KEF categories might be most influenced
The Functional Roles of Humans by this management activity by comparing their functional
Although not explicitly included in the KEF matrix, Homo redundancies to those of all wildlife species in this habitat.
sapiens nonetheless is one of the most effective ecological For example, the ecological function of fungi dispersal is
change agents that can adversely affect BPS. 9, 19 performed overall by two wildlife species in Eastside
Anthropogenic change of habitats is a major disturbance Grasslands: the deer mouse and Rocky Mountain elk.
event in native ecosystems of Washington and Oregon (see These two species both are also potentially influenced by
chapter 20 on Management Activities). use of herbicides in this habitat, because herbicide use is
Humans perform some 35 categories of KEFs (Table 1) linked to at least one habitat element used by each of these
(this tally includes category headings and subheadings, species, namely the habitat elements of herbaceous layer
so there is some repetition in this value). This exceeds the (Habitat Element code 1.2.1), fruits/seeds (1.2.2), shrubs
influence of the two most functionally broad native (1.2.6), fungi (1.2.7), forbs (1.2.8), grasses (1.2.10), and
wildlife species, the American black bear (30 KEF plants (2.1.1). Herbicide use may have a less salient
categories) and the American beaver (24), and helps influence on some other KEFs depending on how many
explain how humans have so greatly altered ecosystem wildlife species perform the functions that are not
functions and processes. Humans certainly rate as a influenced by the activity. In this and other ways, the
functional keystone species, in that their removal from a manager can use the matrixes to evaluate the potential
system would likely result in major redistributions of other influence of specific management activities on ecological
species and their functions. However, none of the functions and determine the functions that may be most
ecological functions performed by humans are necessarily sensitive to the activities, and perhaps those worthy of
critical link functions, which are not performed in at least special mitigation or conservation attention. We suggest
some general way by other wildlife species. using the categories and analysis procedures listed in Table
2 to address this and other patterns of ecological function.
 CHAPTER 6: ECOLOGICAL FUNCTIONS 181

Ultimately, a goal of wildlife conservation may be to Need for Validation and

help ensure fully functional ecosystems. To this end, the Primary Field Studies
manager could use the approaches we offer in this chapter Ecological implications of the KEF pattern categories
to determine, for individual wildlife habitats: discussed in this chapter should be viewed as working
1. the full set of native ecological functions, hypotheses. Further, the relations depicted among the
2. the full functional redundancy of each KEF elements in the species-influence diagram may be
category and total functional diversity, quantified as specific rates (e.g., rates of soil digging and
3. habitat elements contributing to maintaining turnover, rates of pollination, etc.). To do this for the entire
functional diversity of wildlife communities, and set of species, KECs, KEFs, and their relations to BPS and
4.which management activities simplify, extirpate, or management activities would entail a major research
restore functions. program. Thus, the matrixes can be used to prioritize
The manager can use functional profiles to compare critical relations and identify functions most in need of
among communities, or a community over time under empirical, quantitative study.
different management scenarios, to determine: (1) which Scientific knowledge of some functions and species is
scenario would provide for the fullest array of native KEF more complete than for other functions or species. We did
categories; (2) the degree to which full functionality of an not denote this variation in the database per se, but wildlife
ecosystem can be maintained; (3) which functions might specialists or systems ecologists should be able to
be at higher risk; and (4) which species, habitat elements determine areas needing further, or primary, empirical
and structures, and management activities account for study.
reduction or loss of specific functions. This provides a basis
for identifying threats to KEFs as influenced by human Resolution of the KEF Categories
(or other) disturbance events. Linking to maps of species Some KEF categories are broadly generalized whereas
and functions as discussed above, the manager can then others are very specific. This is inevitable, given the
determine spatially where such threats might occur. variation in scientific knowledge and understanding
across species and functional categories. The user of the
Caveats KEF matrix should not expect it to accurately predict
We emphasize that the information in the KEF database ecological functions within small or local geographic areas
largely represents a collective expert judgment and is such as single forest stands; if linked to the wildlife habitats
mostly not based on scientific field studies. As well, the and habitat elements matrixes, the KEF matrix will likely
KEF information is coded largely as categorical data and err on the side of commission of functions rather than
not quantified as rates of influence. Thus, patterns omission. The KEF matrix might serve the manager better
generated by querying the KEF matrix should be at broader geographic scales.
considered as tentative working hypotheses requiring
empirical validation and refinement. We view the major Ecological Functions of
value of the KEF matrix as providing (1) a way of explicitly Other Taxonomic Groups
and repeatably evaluating the functional patterns of The user needs to integrate the information provided in
species and communities and the potential influences from the KEF matrix with some understanding of the functions
human activities, and (2) generating working hypotheses of other taxonomic groups, namely algae, plankton, plants,
on such influences for management consideration and plant allies (cryptogams), and invertebrates. The user
testing. should not expect to be able fully to describe (and
Following Marcot,22 we offer the following caveats prescribe!) ecosystem processes only in terms of vertebrate
when using the KEF matrix: wildlife species. This is especially true with ecological
functions that are poorly represented by vertebrates, such
Completeness of the KEF Database as those pertaining to soil productivity, wood decay,
Although all vertebrate species are included, there may carbon sequestration, filtering of xenobiotics, and many
be many ways to represent their ecological functions, and other functions.23
some functions likely are incompletely depicted. An When managing wildlife in an ecosystem context, it is
example is KEF category 2 on nutrient cycling relations. important not to ignore the functional influence of plants
This category was originally developed more for plants and invertebrates, which really perform most ecological
and invertebrates23 but could be more fully developed for functions in ecosystems. For example, plants and their
vertebrates. Some KEF categories require additional allies—including algae, fungi, lichens, bryophytes (mosses
attention. An example is KEF category 4, disease vector. and liverworts), and cryptogamic soil crusts, as well as
Species listed under this category need review and vascular plants—provide the foundation of net primary
refinement by wildlife disease experts. There are likely to production, and trophic and physical structures of
be other errors of omission and commission in the matrix ecosystems (except for rare autotrophic, chemosynthetic
and other needs to revise or expand the KEF classification and other extremophile-based systems found in a few
system (Table 1). locations). Invertebrates, including soil microorganisms
such as mites, springtails, nematodes, rotifers, protozoa,
and bacteria, also play central functional roles in
 182 Wildlife–Habitat Relationships in Oregon and Washington

ecosystems, especially in soil and decaying wood; E.O. Literature Cited

Wilson called invertebrates the “little things that run the 1. Augustine, D. J., and S. J. McNaughton. 1998. Ungulate effects on the
world.”47 functional species composition of plant communities: herbivore
Ecological functions of plants and invertebrates may selectivity and plant tolerance. Journal of Wildlife Management 62:
best be addressed in functional groups of species,10, 30, 32 1165-1183.
for example, as suggested for wetland plants,2 exotic plant 2. Boutin, C., and P. A. Keddy. 1993. A functional classification of wetland
plants. Journal of Vegetation Science 4: 591-600.
species,11, 15 plants central to ecosystem process and global
3. Brelsford, M. J., J. M. Peek, and G. A. Murray. 1998. Effects of grazing by
change,39 and invertebrates that play ecological roles in wapiti on winter wheat in northern Idaho. Wildlife Society Bulletin 26:
managed forests.38 203-208.
Vertebrates also take a back seat to other organisms in 4. Brown, J. H. 1995. Macroecology. The University of Chicago Press,
terms of species richness. In the interior Columbia River Chicago, IL.
Basin in the U.S., vertebrates constituted only 4% of all 5. Butler, D. R. 1995. Zoogeomorphology: animals as geomorphic agents.
Cambridge University Press, New York, NY.
known species and 1% of all expected (known and
6. Callicott, J. B., L. B. Crowder, and K. Mumford. 1999. Current normative
unknown) species of macroorganisms, and even far less concepts in conservation. Conservation Biology 13: 22-35.
if microorganisms are included.23 Nonetheless, the relation 7. Chapin, F. S., III, and others. 1996. The functional role of species in
of plants, plant allies, and invertebrates to vertebrate terrestrial ecosystems. In B. Walker and W. Steffen, editors. Global
species is pertinent and may be of interest to managers change and terrestrial ecosystems. Cambridge University Press, New
for maintaining overall ecological functions and ecosystem York, NY.
processes. Plants support vertebrate primary consumers 8. Duncan, R. S., and C. A. Chapman. 1999. Seed dispersal and potential
forest succession in abandoned agriculture in tropical Africa. Ecological
including browsers and grazers; invertebrates support
Applications 9: 998-1008.
insectivores and other secondary consumers. Plants and 9. Forester, D. J., and G. E. Machlis. 1996. Modeling human factors that
plant allies provide many kinds of physical habitat affect the loss of biodiversity. Conservation Biology 10: 1253-1263.
structures, help filter water and detoxify soils, support soil 10. Golluscio, R. A., and O. E. Sala. 1993. Plant functional types and
structures for burrowing, and many other functions of use ecological strategies in Patagonian forbs. Journal of Vegetation Science
to vertebrates. Invertebrates or their byproducts (e.g., 4: 839-846.
honey) serve as a food source; they also foster wood decay, 11. Gordon, D. R. 1998. Effects of invasive, non-indigenous plant species
on ecosystem processes: lessons from Florida. Ecological Applications 8:
create snags and down wood, serve as disease
975-989.
transmission vectors, and play many other ecological roles 12. Green, G. I., D. J. Mattson, and J. M. Peek. 1997. Spring feeding on
affecting vertebrates. ungulate carcasses by grizzly bears in Yellowstone National Park.
Journal of Wildlife Management 61: 1040-1055.
Conclusion 13. Green, R. J. 1995. Using frugivores for regeneration: a survey of
knowledge and problems in Australia. Pages 651-657 in J. A. Bissonette
Ecosystem managers can address the biotic component and P. R. Krausman, editors. Integrating people and wildlife for a
of ecosystems by explicitly including key ecological sustainable future. The Wildlife Society, Bethesda, MD.
functions of species (including humans) in assessments 14. Hobbs, N. T. 1996. Modification of ecosystems by ungulates. Journal of
and in formulating management direction. Many Wildlife Management 60: 695-713.
ecosystem management approaches to wildlife and land 15. Horvitz, C. C., J. B. Pascarella, S. McMann, A. Freedman, and R. H.
use planning are already considering ecosystem processes Hofstetter. 1998. Functional roles of invasive non-indigenous plants in
hurricane-affected subtropical hardwood forests. Ecological
of disturbance events (e.g., fire, wind, drought, and Applications 8: 947-974.
flooding). The picture needs to be made complete by also 16. Huston, M. A. 1997. Hidden treatments in ecological experiments: re-
considering the functional role of organisms, and the evaluating the ecosystem function of biodiversity. Oecologia 110: 449-
influence of those functions on overall ecosystem 460.
biodiversity, productivity, and sustainability. The 17. Jones, C. G., J. H. Lawton, and M. Shachak. 1996. Organisms as
framework we offer here provides a rigorous, repeatable ecosystem engineers. Pages 130-147 in F. B. Samson and F. L. Knopf,
editors. Ecosystem management: selected readings. Springer, New York,
means of evaluating at least the qualitative influence of
NY.
wildlife on their ecosystems, and should be viewed as 18. Li, C. Y., C. Maser, Z. Maser, and B. A. Caldwell. 1986. Role of three
starting hypotheses from which to focus conservation or rodents in forest nitrogen fixation in western Oregon: another aspect
research attention. We fully expect that many KEF of mammal-mycorrhizal fungus-tree mutualism. Great Basin Naturalist
categories and even their qualitative relations with wildlife 46: 411-414.
species may change given empirical study. 19. Mack, M. C., and C. M. D’Antonio. 1998. Impacts of biological
invasions on disturbance regimes. Trends in Ecology and Evolution 13:
195-198.
Acknowledgments 20. MacNally, R. C. 1995. Ecological versatility and community ecology.
We thank Tom O’Neil, Kelly Bettinger, and Sally Olsen- Cambridge University Press, New York, NY.
Edge for helpful comment on the manuscript. We 21. Marcot, B. G. 1997. Species-environment relations (SER) database. U.S.
gratefully acknowledge the contributions of Marla Forest Service, Pacific Northwest Research Station, Portland, OR.
Interior Columbia Basin Ecosystem Management Project database
Trevithick in developing the databases. We benefited from
available on-line at URL: http://www.icbemp.gov/spatial/metadata/
ecological discussions with Mikell O’Mealy and Susan databases/dbase.html
Tank.
 CHAPTER 6: ECOLOGICAL FUNCTIONS 183

22. Marcot, B. G. 1997. The species-environment relations (SER) database: 42. Stone, R. 1995. Taking a new look at life through a functional lens.
an overview and some cautions in its use. U.S. Forest Service, Pacific Science 269: 316.
Northwest Research Station, Portland, OR. Interior Columbia Basin 43. Urbanska, K., N. Webb, and P. Edwards, editors. 1998. Restoration
Ecosystem Management Project database documentation available on- ecology and sustainable development. Cambridge University Press,
line at URL: http://www.icbemp.gov/spatial/metadata/databases/ New York, NY.
792aux2.html 44. van Gardingen, P. R., G. M. Foody, and P. J. Curran, eds. 1997. Scaling-up:
23. Marcot, B. G., M. A. Castellano, J. A. Christy, L. K. Croft, J. F. Lehmkuhl, from cell to landscape. Cambridge University Press, New York, NY.
R. H. Naney, K. Nelson, C. G. Niwa, R. E. Rosentreter, R. E. Sandquist, B. 45. Walker, B. H. 1992. Biodiversity and ecological redundancy.
C. Wales, and E. Zieroth. 1997. Terrestrial ecology assessment. Pages Conservation Biology 6: 18-23.
1497-1713 in: T. M. Quigley and S. J. Arbelbide, editors. An assessment of 46. Williams, J. B., B. E. Tieleman, and M. Shobrak. 1999. Lizard burrows
ecosystem components in the interior Columbia Basin and portions of provide thermal refugia for larks in the Arabian desert. Condor 101:
the Klamath and Great Basins. Volume III. U.S. Forest Service General 714-717.
Technical Report PNW-GTR-405. 47. Wilson, E. O. 1987. The little things that run the world (the
24. Marcot, B. G., L. K. Croft, J. F. Lehmkuhl, R. H. Naney, C. G. Niwa, W. R. importance and conservation of invertebrates). Conservation Biology
Owen, and R. E. Sandquist. 1998. Macroecology, paleoecology, and 1: 344-346.
ecological integrity of terrestrial species and communities of the 48. Young, D. D., Jr, and T. R. McCabe. 1997. Grizzly bear predation rates
interior Columbia River Basin and portions of the Klamath and Great on caribou calves in northeastern Alaska. Journal of Wildlife
Basins. U.S. Forest Service, General Technical Report PNW-GTR-410. Management 61: 1056-1066.
25. Mawdsley, N. A., S. G. Compton, and R. J. Whittaker. 1998. Population
persistence, pollination mutualisms, and figs in fragmented tropical
landscapes. Conservation Biology 12: 1416-1420.
26. Mills, L. S., M. E. Soule, and D. F. Doak. 1993. The keystone-species
concept in ecology and conservation. BioScience 43: 219-224.
27. Moore, G. C., and G. R. Parker. 1992. Colonization by the eastern
coyote (Canis latrans). Pages 23-37 in A. H. Boer, editor. Ecology and
management of the eastern coyote. Wildlife Research Unit, University
of New Brunswick, Fredericton, New Brunswick, Canada.
28. Morrison, M. L., B. G. Marcot, and R. W. Mannan. 1998. Wildlife-habitat
relationships: concepts and applications. Second edition. University of
Wisconsin Press, Madison, WI.
29. Naeem, S. 1998. Species redundancy and ecosystem reliability.
Conservation Biology 12: 39-45.
30. Parueo, J. M., and W. K. Lauenroth. 1996. Relative abundance of plant
functional types in grasslands and shrublands of North America.
Ecological Applications 6: 1212-1224.
31. Pimm, S. L. 1994. Biodiversity and the balance of nature. Pages 347-
359 in E. D. Shulze and H. A. Mooney, editors. Biodiversity and
ecosystem function. Springer-Verlag, Berlin, Heidelberg, Germany.
32. Pugnaire, F. I., and F. Valladares, editors. 1999. Handbook of functional
plant ecology. Marcel Dekker, New York, NY.
33. Rastetter, E. B., L. Gough, A. E. Hartley, D. A. Herbert, K. J. Nadelhoffer,
and M. Williams. 1999. A revised assessment of species redundancy and
ecosystem reliability. Conservation Biology 13: 440-443.
34. Rauscher, H. M. 1999. Ecosystem management decision support for
federal forests in the United States: a review. Forest Ecology and
Management 114: 173-198.
35. Reid, W.V., and K. R. Miller. 1989. Keeping options alive-the scientific
basis for conserving biodiversity. World Resources Institute,
Washington, D.C.
36. Robertson, A. W., D. Kelly, J. J. Ladley, and A. D. Sparrow. 1999. Effects of
pollinator loss on endemic New Zealand mistletoes (Loranthaceae).
Conservation Biology 13: 499-508.
37. Schlapfer, F., and B. Schmid. 1999. Ecosystem effects of biodiversity: a
classification of hypotheses and exploration of empirical results.
Ecological Applications 9: 893-912.
38. Schowalter, T., E. Hansen, R. Molina, and Y. Zhang. 1997. Integrating the
ecological roles of phytophagous insects, plant pathogens, and
mycorrhizae in managed forests. Pages 171-189 in K. A. Kohm and J. F.
Franklin, editors. Creating a forestry for the 21st century: the science
of ecosystem management. Island Press, Washington, D.C.
39. Smith, T. M., H. H. Shugart, and F. I. Woodward, editors. 1997. Plant
functional types: their relevance to ecosystem properties and global
change. Cambridge University Press, New York, NY.
40. Soule, M. and K. A. Kohm. 1989. Research Priorities for Conservation
Biology. Island Press, Washington, D.C.
41. Stohlgren, T. J., L. D. Schell, and B. Vanden Heuvel. 1999. How grazing
and soil quality affect native and exotic plant diversity in Rocky
Mountain grasslands. Ecological Applications 9: 45-64.
 184 Wildlife–Habitat Relationships in Oregon and Washington

Table 2. A taxonomy of patterns of key ecological functions (KEFs) of wildlife species and communities, and
how to evaluate them using the KEF database.*

Functional pattern Definition Ecological implications How to evaluate

Community patterns
Functional redundancy The number of species High redundancy imparts greater Tally number of species by KEF
performing the same ecological resistance of the community to category for specific wildlife
function in a community. changes in its overall functional habitats, comparing changes over
integrity. Low redundancy suggests time or among habitats.
critical functions to watch.
Functional richness The total number of KEF Denotes degree of functional Tally number of KEF categories
categories in a community. complexity; greater functional among all species present in a
richness means more functionally wildlife habitat and, optionally,
diverse systems. Also denotes the habitat structure. Compare such
degree to which the full “functional tallies resulting from changes in
web” of a community would be habitats and structures.
provided or conserved.
Total functional diversity The total array of KEF Denotes total functional capacity Tally number of species by KEF
categories weighted by their of a community. High functional category for a given wildlife
redundancy, i.e., the number of diversity means many functions habitat; calculate mean number of
functions times the mean and even redundancy among species per KEF category. Can
functional redundancy across functions; low functional diversity assign weights to some KEF
all functions. means few functions and skewed categories if they pertain to
redundancy (some functions with specific management objectives.
few species).
Functional web The set of all KEFs within a Depicts how habitat elements Identify habitat elements of
community and their provide for species, and the array management interest; list all
connections among species of ecological functions performed species within a wildlife habitat
and thence to habitat elements. by those species. Functions typically that are associated with those
extend well beyond the specific habitat elements; list all KEF
habitat elements. categories associated with those
species. Compare changes in
habitat elements.
Functional profile The degree of functional Identifies communities with low Tally number of species by KEF
redundancy compared across (or high) redundancy of particular category and by wildlife habitat.
communities. KEF categories. This can help Identify wildlife habitats with
prioritize habitat management, lowest tallies for each KEF
e.g., to ensure continuance of category.
low-redundancy functions.
Functional homologies The functional similarity of Two communities are functionally Produce functional profiles across
communities even if species homologous if they have similar all KEF categories for several
composition differs. functional profiles and patterns communities or for a community
of functional redundancy, even if over time based on its expected
the species performing the changes in habitat elements,
functions differ. Functionally habitat structures, etc. Compare
homologous communities can be the profiles (e.g., via contingency
expected to operate in similar analysis) and identify statistically
ecological ways. similar (functionally homologous)
communities.

* Many of these categories are unstudied in wildlife communities of Washington and Oregon; thus, ecological implications should be
viewed as working hypotheses. (See Table 1 for KEF categories.)
 CHAPTER 6: ECOLOGICAL FUNCTIONS 185

Functional pattern Definition Ecological implications How to evaluate

Geographic patterns
Functional bottlenecks Geographic locations with Functional bottlenecks denote Map wildlife habitats and/or
or cold spots very low functional redundancy areas of higher risk of severing distributional ranges of wildlife
of an otherwise widely- functionally connected species. For a given KEF category,
distributed functional category. communities across the landscape. map the number of wildlife
Severing functions might set the species in each habitat or overlay
stage for degradation of their range maps. Identify
functional ecosystems. locations with lowest species
richness bordering higher richness
on each side; these are geographic
functional bottlenecks (“cold
spots”).
Functional linkages or Geographic locations with Functional hot spots denote areas Map species richness for a
hot spots very high functional where many species provide a particular KEF category as above.
redundancy. specific ecological function; such Identify locations with highest
communities may be more species richness; these are
resilient to changes in environment functional linkages (“hot spots”).
or habitat for that function. Determine which species occur in
a given hot spot and their wildlife
habitats, habitat elements, and
habitat structures, and how
changes might influence the
persistence of the species and
thus the redundancy of the
function.

Species’ functional roles

Functional keystone species, Functional keystone species Reduction or extirpation of For a given wildlife habitat, tally
critical functional link are species whose removal populations of functional keystone the number of species for each
species, and critical functions would most alter the structure species and critical functional links KEF category (functional
or function of the community. may have a ripple effect in their redundancy). For KEF categories
One form of this may be critical ecosystem, causing unexpected or with only 1 species, determine
functional links, which are undue changes in biodiversity, which species performs this
species that are the only ones biotic processes, and the function. This is a critical
that perform a specific functional web of a community. functional link species for this
ecological function in a particular function in this habitat.
community. A critical function
therefore is the associated
functional category represented
by only 1 (or very few) species
within a community.
Functional breadth and The number of ecological Species with the narrowest For a given wildlife habitat, tally
functional specialization of functions performed by a functional breadth (i.e., fewest the number of KEF categories for
species species. functions) are functional specialists each species. Identify the species
and may be more vulnerable to with the fewest number of
extirpation from changes in categories. These are functional
conditions supporting that specialists. Determine their
function. habitat elements and structures
and thus their potential
vulnerability to changes thereof.
Functional specialists that are also
functional keystones may be of
high priority for conservation
attention.
 186 Wildlife–Habitat Relationships in Oregon and Washington

Functional pattern Definition Ecological implications How to evaluate

Functional responses of communities

Functional resilience The capacity of a community to Functionally resilient Determine the total functional
return to a starting pattern of communities are better able to diversity, functional richness, and
total functional diversity, maintain their biotic processes in functional redundancy of a pre-
richness, and redundancy, the face of disturbances. disturbance community. Then
following a disturbance event. Conversely, it is important to know determine the types and rates of
how far a community can be recovery of its wildlife habitats,
changed by some anthropogenic habitat elements, and habitat
disturbance event and still be able structures following some
to return to its starting functional disturbance; the wildlife species
pattern. associated with such recovery
stages; and the species’ KEF
categories and functional diversity,
richness, and redundancy for each
recovery stage. Compare stages
for functional similarity and thus
resilience.
Functional resistance The ability of a community to Functionally resistant communities Analyze as above and determine
resist changing its functional can be counted on to continue to the degree to which functional
diversity, richness, and provide specific ecological functions diversity, richness, and redundancy
redundancy, following a in spite of and during disturbances. do not change for each post-
disturbance event. They may provide a bastion for a disturbance stage. This is a
specific desired function in a measure of functional resistance.
disturbed or managed landscape.
Functional attenuation The degree to which the set Functionally attentuated Analyze as above and determine
of ecological functions within communities provide fewer or which KEF categories likely drop
a community simplify following lower redundancies of ecological out and which remain over post-
a disturbance event. functions. It may be particularly disturbance recovery stages as
important to know the degree of compared with initial conditions.
functional attentuation to be Calculate functional diversity,
expected following anthropogenic richness, and redundancy for each
disturbances. stage to determine the rate of
functional attenuation. Compare
final stage to initial conditions to
determine overall functional
attenuation.
Functional shifting The degree to which a Communities with low functional Analyze as above and compare
community changes to a new, resilience or resistance may end KEF categories and functional
stable, functional constitution up with a new array of functions diversity, richness, and redundancy
following a disturbance event. and a new pattern of functional between predisturbance and final,
diversity, richness, and stable stages. Identify which
redundancy. It may be particularly functions are lost or gained, and
important to know how a which change in redundancy.
community might functionally shift
following anthropogenic
disturbances and thus which
functions might be weakened,
strengthened, lost, or gained.
Imperiled functions A function that is represented Loss of imperiled functions serve For a given wildlife habitat,
by very few species (critical to degrade overall ecosystem determine KEF categories with
functional link species) or by integrity. Even seldom-performed the lowest functional redundancy,
species that are themselves ecological functions might be and the risk level of the associated
scarce, declining, or moribund, critical to maintaining ecosystems, species. Imperiled functions are
where extirpation of the such as occasional dispersal of those with 1 or few species which
species would mean loss of the plant seeds in the face of shifting are themselves at risk.
function. Imperiled functions climates.
also can be identified
geographically.
 7
Wildlife of Westside and High Montane Forests
Deanna H. Olson, Joan C. Hagar,Andrew B. Carey, John H. Cissel, &
Frederick J. Swanson

Introduction
More than 300 vertebrate species are associated with site, microhabitat, and microclimate conditions, and recent
western forests of Oregon and Washington (Table 1). disturbance events contribute to explanations of species
Western and montane conifer-hardwood forests and oak distribution patterns. Habitat assessments of forest-
woodlands are some of the more species-rich areas within associated wildlife conducted for this volume are
the two states. Both the productivity and the mosaic of compiled to summarize species-habitat relations. Across
conditions within western forests contribute to the higher spatial scales, species life history, behavior, and intra- and
vertebrate diversity. These forests are exceeded in richness interspecific interactions are significant elements for our
only by habitats encompassed by riparian-wetlands, understanding of wildlife habitat associations. These
urban, and agriculture and pasture designations. abiotic and biotic components are outlined to more fully
Interestingly, these are habitats that either border or are conceptualize their roles in the organization of forest
found nested within western forests. A high overlap of faunal assemblages. Hotspots of wildlife diversity are
species occurs between these habitat types and western addressed at both broad landscape and finer forest stand
forests, especially along their interfaces. Forest species spatial scales.
include those taxa that are obligates to forested habitats Although all vertebrate classes have some coverage in
for all or part of their life history, more generalist species this chapter, amphibians are used for many lead examples.
that occur in the forest matrix but also in other nonforest Whereas amphibians do not comprise a large percentage
types, and transient species that are found incidentally in of the overall wildlife fauna in these habitats (about 13%),
forests because of their proximity to other habitats. western forests are key habitats for many amphibians. Of
This chapter provides an overview of the broad- and 31 native amphibian species occurring in Oregon and
fine-scale patterns of western forest wildlife assemblages, Washington, 29 (93%) occur in western forests (Table 1)
emphasizing the main faunal habitat associations with and 22 (71%) are restricted to this region.39 These restricted
forest conditions. Drivers of the geographic distributions species are either endemic to western Oregon and
of many taxa include climate conditions, the legacy of past Washington or have ranges within the larger ecological
natural disturbances, and vegetation types. This mix of extent of the western forest landscape.
physical and biological conditions has been elegantly There are many unique amphibian taxa reliant on these
consolidated into ecoregion designations for western forested habitats. Among these are (1) the tailed frog
forests. Ecoregions provide a context for broad-scale (Figure 1); (2) the largest terrestrially occurring
species richness pattern assessment. At finer spatial scales, salamander, the Pacific giant salamander (Figure 2); (3)

Table 1. Species occurrences (no. species) in western and montane forest habitat types of
Washington and Oregon.*

Taxon n Montane mixed SW Oregon Westside oak and Westside lowlands

conifer mixed conifer- dry Douglas-fir conifer-hardwood
hardwood & woodlands

Amphibians 29 27 19 18 26
Reptiles 20 5 19 18 14
Birds 164 107 131 119 120
Mammals 104 82 67 74 73
Total 317 221 236 229 233
* Data are compiled from expert panel assessments for this book, and no historic occurrences are included (e.g., bison, gray wolf,
merlin). n = total no. species evaluated.

187
 Figure 2. Pacific giant salamanders are stream breeders, but
after metamorphosis they are terrestrial, occurring in upslope
Figure 1. Tailed frogs (Ascaphus truei) are primitive anurans forests. J. S. Applegarth, photographer.
strongly associated with conditions of western Oregon and
Washington old-growth forests. J. S. Applegarth,
photographer.

the torrent salamanders, Rhyacotriton spp.; and (4) several for species. These have and will mold habitats, and
endemic plethodontid salamanders34 (Figure 3, page 196). incidentally or by design, wildlife distribution and
Western forests are recognized as a region of phylogenetic abundance patterns. We are learning how to be stewards
radiation for amphibians, yet our understanding of the for the long-term economic productivity and ecological
amphibian biodiversity is incomplete. As a result of sustainability of these forested landscapes. As our
ecological, morphometric, and genetic studies, formal knowledge gaps are filled and standards for biological
designations of distinct populations (e.g., evolutionarily conservation are adjusted, we can expect an adaptive
significant units [ESUs]30) or species across the western framework for forest management policies for the next
forest landscape are expected to change. Within this millennium.
vertebrate class, the tendency for philopatry and limited
dispersal capability has resulted in isolation of Broad Scale Patterns
subpopulations, that now seem to have diverged Western forests in Oregon and Washington harbor a
ecologically, morphologically, and potentially diverse and unique fauna. From a broad regional
taxonomically. Support for newly recognized species, or perspective, the habitats provided by western coniferous
proposals to recognize distinct populations or species, forests across the landscape can be easily distinguished.
have developed for Rhyacotriton, Dicamptodon, several Their high productivity and structural complexity are
Plethodon salamanders, 2 Rana frogs, and 1 Aneides captured by remote sensing images (Figure 4, page 197).
salamander. The potentially hidden biodiversity within This is not a homogeneous landscape, however, as might
amphibians is expected to be a topic of research for some seem apparent at this broad spatial scale. It is the habitat
years to come. As species are examined in this light, heterogeneity, both across the region and within
concern for the status of unique subpopulations or species landscapes, that is pivotal to understanding the wildlife
is heightened. This has ramifications for species diversity of the western forests.
conservation, and consequently western forest The large-scale mosaic of habitats across westside and
management policies. montane forests is exemplified by the multiple
Final sections of this chapter address forest physiographic provinces or ecoregions for this area.
management and its role in maintaining the persistence Physiographic province designations take primarily
of wildlife populations in the western forest region. physical factors into consideration, including soils,
Anthropogenic disturbances over the last 100 years have geomorphology, natural disturbance history, and climate.
reshaped forest habitats in Oregon and Washington. A When discussed in light of wildlife distributions,
current pivot in trajectories for public land management Nussbaum et al.37 recognized 3 physiographic provinces
is introducing new standards for landowner stewardship over the western forest landscape (Cascade Mountains,
of biological resources. Consequences of both the past Coast Ranges, and Klamath Mountains). These are
management policies that emphasized economic resources subdivided into 8 physiographic provinces in the Recovery
and the current new management directions balancing Plan for the northern spotted owl56 (Cascade Mountains
socioeconomics with ecosystem integrity are significant into 4 provinces: eastern and western Cascade Ranges in

188
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 189

both Oregon and Washington; Coast Ranges into 3 western forest ecoregions. Nine general species range
provinces: Olympic Peninsula, Washington lowlands, patterns were derived from avian range maps in Csuti et
Oregon Coast Range; and the Klamath Mountains). al.16 (Figure 7, >240 species evaluated, shorebirds restricted
Ecoregions are geographic descriptors of environmental to coastal shores were excluded). Proportion of species
conditions; they are based on spatial patterns of per pattern and example species are (1) 2%, northern
physiography, climate, disturbance, vegetation, and spotted owl and harlequin duck; (2) 8%, MacGillivray’s
wildlife. Omernik40 recognized 5 Level III ecoregions for warbler and varied thrush; (3) 3%, purple finch and
the area of interest in this chapter (Coast Range, Cascades, chestnut backed chickadee; (4) 22%, American dipper and
Eastern Cascades and foothills, North Cascades, Klamath dark-eyed junco; (5) 2%, plain titmouse and black phoebe;
Mountains); these are analogous to some of the (6) 7%, boreal owl and three-toed woodpecker; (7) 6%,
physiographic provinces above. Pater et al.42 provided a calliope hummingbird and flammulated owl; (8) 9%,
further subdivision of the Level II ecoregions into 42 Level northern flicker and common raven; and (9) 39%, western
IV regions (Figure 5, page 198). meadowlark and yellow-headed blackbird. Half of the
Whereas general wildlife patterns were criteria for bird species in Oregon and Washington have the majority
ecoregion determination, correspondence of wildlife of their ranges in westside forests (patterns 1-7, Figure 7.)
species distributions with ecoregions is not fully Most western forest birds have some occurrences east of
investigated. Evaluation of species’ associations with the the Cascade Range (e.g., patterns 2, 4, 5, and 7), and
newly developed Level IV ecoregions for western Oregon relatively few have ranges restricted to westside forests
and Washington have not been conducted. However, some (e.g., patterns 1, 3, and 6).
initial pattern assessments have developed using Oregon Such a species range assessment provides a slightly
Level III ecoregions. The Oregon Biodiversity Project41 shifted perspective of the spatial arrangements of species
presented species richness categories on a hexagonal grid in comparison to species richness maps (e.g., Figure 6).
for the State, with 150,000-acre (60,750-ha) hexagons, for For comparison among wildlife vertebrate taxa and
butterflies, amphibians, fish, reptiles, and mammals between Oregon and Washington, additional range
(Figure 6, page 200). patterns were assessed (Figure 8). To facilitate
Across Oregon, fish species richness was greatest in the comparisons, patterns 1 through 4 were combined to
western portion of the state coincident with western forests consolidate species with general west-side forest
(many hexagons with >20 species and >29 species), relationships, and patterns 6 and 7 were combined to more
compared to east-side areas (<8 species in much of the easily represent species restricted to the Cascade Range.
southeastern portion of the state, and patches from <8 to In Oregon, forests in the Cascade Range and westward
20-29 species in the northeastern portion). In the Cascades, (Figures 7 and 8, general range patterns 1 through 7) are
fish species richness decreased with increasing elevation. focal portions of the ranges of >50% of the species in each
Mammal and butterfly richness patterns were similar to of these vertebrate classes. However, for all vertebrate
each other, although mammals were less variable in classes assessed, east-side landscapes provide significant
richness numbers across the western forested landscape. habitats for some species (Figures 7 and 8, general range
These 2 taxa had greatest diversity in the southern Cascade pattern 9). This fact might be overlooked if only species
Range and up the ridge of the Cascade crest (including richness maps were examined. For example, amphibian
portions of the Klamath, and eastern and western Cascade species richness (Figure 6) is clearly tied to western forests,
ecoregions; >102 butterfly and >64 mammal species in yet east-side landscapes are of critical importance for the
much of this area). In contrast, among the western forested ranges of about 20% of Oregon and Washington
regions, butterfly and mammal richness were least in the amphibian species (Figure 8, general range pattern 9).
Coast Range. Amphibian patterns in this analysis were Interestingly, reptiles in Washington are much more reliant
extremely patchy. Like fish, diversity was greater in the on east-side habitats than those in Oregon (Figure 8). In
western forests, with the least richness in the eastern Oregon, the west-side Klamath ecoregion contributes a
Cascades. Among the other 3 western forested ecoregions, significant reptile species “hotspot.”
each had some hexagons with higher amphibian diversity Herpetological distribution patterns in the Klamath
(>15 species), most with intermediate (7-15 species) ecoregion are further addressed by Bury and Pearl.4 They
richness categories, and a number of patches with <5 reported that this region has the most species-rich
amphibians. Reptile richness showed the most remarkable herpetofauna of any similar-sized mountain range in the
concordance with ecoregion boundaries. Among western Pacific Northwest (38 native species of amphibians and
forested regions, reptile richness was highest (>15 species) reptiles; this number reflects total richness for the region,
in the Klamath region. Much of the Coast Range had the as opposed to richness per hexagon in the previous
lowest reptile richness category (<6 species), and the discussion). As already noted, this is largely attributable
western Cascades showed a decreasing richness with to the higher reptile richness in the Klamath region. Bury
elevation, with 12-15 species in the foothills and 6-8 species and Pearl4 provided explanation for this patterning based
in the higher landscape. on organism ecology, life history and behavior, and the
Although Oregon bird species richness patterns were common data elements that go into defining ecoregions
not compiled for the Oregon Biodiversity Project,41 bird (e.g., elevation, latitude, climate, legacy of past natural
species ranges correspond to several patterns related to disturbance events, vegetative structure and composition).
 190 Wildlife–Habitat Relationships in Oregon and Washington

Figure 7. General range patterns of

Oregon birds, derived from species
range maps in Csuti et al.16 Dark
shading indicates the main portion of
species ranges (areas in which most
localities or habitats are known),
light stippling indicates some
occurrences, white indicates no or
very few localities are known. Level
III ecoregion boundaries (Figure 5)
are used for general range
boundaries.

80

60
Figure 8. General range patterns
(1-9 from Figure 7) of Oregon
40
wildlife and extension of those
geographic patterns in
Washington. Patterns 1-4 from 20
Fig. 7 are combined to represent
taxa with predominantly west-side
ranges, and patterns 6 and 7 are 0
combined to represent species Oregon
occurring primarily in the Cascade
Range. Data are compiled from Birds
range maps in Csuti et al.,16
Johnson and Cassidy,22 Leonard et Mammals
al.,25 Smith et al.,46 and Storm and
Reptiles
Leonard.48
80
Amphibians

60

40

20

0
1-4 5 6-7 8 9
Washington
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 191

These are the puzzle pieces that must be evaluated to species in the southwestern U.S., which presumably
understand the broad-scale species richness trends for any moved north with the retreating glaciers and
wildlife taxa. This rationale for herpetofauna of the establishment of warmer, more xeric conditions.
Klamath ecoregion can be extended to demonstrate Likewise, amphibians responded to the same
probable drivers of regional patterns in the western forests combination of events, the broad-ranging temperate
of Oregon and Washington. forests subject to subsequent mountain-building,
Across western Oregon and Washington forests, many glaciation, and drought. Yet rather than these events
wildlife species diversity patterns are established with resulting in colonization from adjoining regions, as in
latitude, elevation, and climate, some of the basic data reptiles, many amphibian populations became isolated as
elements of ecoregions. These factors interact to produce the temperate rain forests contracted from their historical
different biotic and abiotic environments for wildlife, and extent, becoming fragmented, with inhospitable
species composition changes markedly with these factors. conditions developing between remnants. Both glaciation
Taxa reliant on temperature and moisture regimes, such and drought exacerbated isolation of such species
as reptiles and amphibians, show strong physiographic dependent on milder conditions. Welsh51 elaborated on
associations.37 For example, species richness decreases this scenario in his presentation of relictual amphibian
with increases in latitude and elevation for reptiles and species adapted to the ancient primeval coniferous forest
amphibians in western forests as temperature and ecosystem. Several species now associated with old-
moisture tolerance limits are encountered. Among growth forest conditions may be relics of this historical
amphibians occurring in the Cascade Range, forest landscape, now strongly tied to stands retaining a
approximately 20 species generally occur at elevations semblance of the microhabitat and microclimate
<4,000 ft (1,219 m), 13 species at 4,000-5,000 ft (1,219-1,524 conditions of yore. Welsh’s59 analyses from southwestern
m), 9 species at 5,000-6,000 ft (1,524-1,829 m), and 7 species Oregon and northwestern California identified the Del
at >6,000 ft (1829 m, range limits compiled from Leonard Norte salamander, the tailed frog, and the southern torrent
et al.25). Latitudinally, reptile species distributions become salamander as such relics that generally do not tolerate
much more restricted to the north: only 3 of about 18 conditions in highly managed forest stands. As presented
species retain broad distributions, 8 species are found by Welsh,59 relatively small sedentary organisms with
primarily within the inland valleys (e.g., Willamette restricted distributions and narrow habitat requirements
Valley), and 7 species’ ranges end with the Klamath and climatic tolerances would be sensitive to disturbance
Mountains ecoregion. In southern Oregon, amphibian or environmental change. Many of the 22 native
species gradients are found with distance from the coast. amphibian species restricted to the western forest
Presumably the cooler, moister climates of the coastal zone landscape occur in association with old-growth forest
may explain this; conditions become xeric rapidly inland. conditions, and are in essence phylogenetically symbolic
Of 9 salamanders occurring at the coast in the extreme of a past legacy of events and conditions in this region.
southwestern corner of the state, only 5 have likely ranges Isolation scenarios based on disturbance and changing
extending through Jackson County, about 100 miles (161 environments can be conceived for many of them (e.g.,
km) inland (Pacific giant salamanders, roughskin newts, the plethodontids in Figure 3). The current distribution of
Ensatina, clouded salamanders, and Del Norte the Larch Mountain salamander (Figure 3) likely reflects
salamanders are “replaced” by Siskiyou Mountain volcanic disturbance events from Mount Saint Helens,
salamanders; Figure 3). Mount Rainier, and Mount Hood, in addition to spatio-
Current broad distributions of many taxa also reflect a temporal fluctuations in microhabitat and microclimate
legacy effect of past natural disturbances or environmental conditions. Such dot map distributions also reflect biases
conditions. In particular, less vagile taxa with specific of survey locations (i.e., remote areas are less sampled).34
habitat requirements are not as resilient to disturbances, Synthesizing the data contributing to ecoregion
and their distributions may retain a signature of past designations can lead to a better understanding of
events for extended time periods. The development of the “hotspots” of wildlife diversity at the broad spatial scale
Pacific Northwest herpetofauna and current herpetofaunal of western and montane forests. As discussed, reptile
distributions were summarized by Nussbaum et al.37 in diversity is highest in the more xeric Klamath Mountain
light of the regional-to-continental extent of the northern region. The margins of several species ranges occur here,
temperate rain forests comprising the Arcto-Cretaceo- at their northern or western extent, whereas those that
Tertiary Geoflora, and subsequent orogenic, glaciation, are found to the north often show more restricted
and drought events. They surmised that some currently distributions to lowland areas. Birds mirror this pattern
depauperate areas can be accounted for by both (see Case History 1, below). DellaSala et al.17 referred to
insufficient time for reinvasion and the current relatively the Klamath-Siskiyou ecoregion as an area of
cold, harsh environments restricting colonization. They “extraordinary biodiversity”, ranking its biodiversity
proposed that reptiles with broader distributions “among the world’s most outstanding temperate
elsewhere and peripherally distributed in the Pacific coniferous forests”. Of 2,377 terrestrial animals they
Northwest may be relatively recently introduced to this analyzed (snails, butterflies, birds, mammals, reptiles and
region subsequent to the last glaciation (25,000-10,000 amphibians), 168 (7%) occurred nowhere else. They found
years ago). Many of the region’s reptiles are derived from high endemism among aquatic animals: 42% of fish (n=33),
 192 Wildlife–Habitat Relationships in Oregon and Washington

and 60% of mollusks (n=235). At the subspecies level, 281 Changes in riparian bird communities along an elevational
(8%) of >3,500 plants were cited as endemic. A dominant gradient contribute to the decline in species richness with
contributor to the Klamath-Siskiyou wildlife diversity increasing elevation. Several riparian obligate species,
hotspot is the convergence of several mountain ranges, in such as great blue heron, wood duck, osprey, and
an “H” pattern. As a result, there is juxtaposition of several kingfisher, are most abundant at low elevations, where
highly dissected stream drainage networks, extensive forests or trees adjacent to large streams, rivers, and
elevational and microclimatic gradients, and relative estuaries provide habitat for nesting, roosting, and
proximity of different life zones or ecoregions: the high foraging. At higher elevations, fewer species are strongly
desert of the Great Basin (east); California mountains (i.e., associated with steep, highly constrained streams and
Sierra Nevada), plains and valleys (south); Coast (west); riparian vegetation that typically is only subtly
and the inland wet-temperate river valley systems (in the distinguished from surrounding upslope forest. Dippers,
Klamath region and north to the Willamette Valley). In however, specialize in foraging and nesting along such
addition to the diverse array of physical conditions and clear, swift mountain streams throughout western Oregon
vegetation types from the intersection of these zones, the and Washington.
survival of relictual flora and fauna due to reduced glacial Bird community composition varies with seral stage,
impacts and, consequently, remnant habitats in this area with differences being most pronounced between very
adds a legacy effect to its biological diversity. For early open canopy (i.e., grass-forb-shrub) and closed
amphibians, at a provincial or ecoregion level, a broader canopy stages. Species richness of birds tends to be similar
ranging diversity hotspot occurs in the more productive in early and late stages of forest development, and lowest
wet temperate forests of the Coast Range.37 Broad-scale in the structurally simple mid-seral stages of managed
diversity hotspots for mammals are less easily identified. forests.2 Although relations between abundance and seral
Diversity hotspots may be distinguished at finer spatial stage vary for many species on a geographic scale,44 species
scales, and are discussed below relative to patterns that are typically associated with early seral conditions
resulting from more fine-grained habitat elements. west of the Cascade Range include willow flycatchers,
white-crowned sparrows, song sparrows, and spotted
towhees.2, 11, 57 Species that typically are more abundant in
old forests include Pacific-slope flycatcher, varied thrush,
Case History 1
and many members of the bark-foraging guild (e.g., brown
Patterns of Bird Communities in Coniferous creeper, chestnut-backed chickadee, red-breasted
Forests of Western Oregon and Washington nuthatch, hairy woodpecker44). The avian species that are
Joan C. Hagar most closely associated with old-growth forests are
marbled murrelets, northern spotted owls, and Vaux’s
The majority of bird species occurring in conifer-
swifts.44 However, most species that reach their greatest
dominated forests west of the Cascade crest in Oregon
abundance in older forests nonetheless also will use early
and Washington are widely distributed within this zone.
seral habitats as long as key structural features are present.
Chestnut-backed chickadees and varied thrushes are
For example, species that forage on bark and nest in
notable residents of moist, low elevation (<5,000 ft [1,524
cavities, such as chestnut-backed chickadees and red-
m]) coniferous forests west of the Cascade crest because
breasted nuthatches, occur in recent harvest units where
they are broadly endemic to the Pacific Northwest and
green trees and snags have been retained.11, 57
coastal southern California. Other year-round resident
Hardwood trees and shrubs may be one of the most
species that are characteristic of these forests include
important factors influencing bird community
northern spotted owl, pileated woodpecker, Steller’s jay,
composition in the conifer-dominated landscape of the
winter wren, and golden-crowned kinglet. During the
Pacific Northwest. The abundance and diversity of birds
breeding season, migrant species swell the ranks of the
has been correlated positively with the abundance and
avifauna in terms of both numbers of species and density
distribution of hardwoods.7, 19, 20, 21, 26, 32, 43 Deciduous
of individuals. Among the most broadly distributed and
hardwoods provide different resources for foraging and
abundant migrant species in these conifer forests are the
nesting than conifers, and thus provide unique habitat
Swainson’s thrush, Pacific-slope flycatcher, and hermit
with which several bird species are strongly associated.
warbler. The breeding distribution of hermit warblers is
Warbling vireos are predictably found in alder groves, and
largely restricted to Washington, Oregon, and California,
several species of neotropical migrant warblers (e.g.,
defining this species as an endemic.
MacGillivray’s, orange-crowned, Wilson’s) typically
Avian community composition is influenced by
forage and nest in thickets of deciduous shrubs.16, 31 Unique
elevation, seral stage, vegetation structure and
associations between individual avian species and either
composition, presence of water and other special features,
hardwoods or conifers result in high bird diversity where
and the interaction of all these factors. Total bird
hardwoods are mixed with or adjacent to conifers.
abundance in western Oregon and Washington is
Examples include the margins of large valleys where
generally negatively correlated with elevation. 21 A
Oregon white oak occurs and the Klamath region.
negative relationship between elevation and bird species
richness in old-growth forests also has been reported.24
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 193

Provincial Bird Patterns communities. In addition, merlins, northwestern crows,

Few bird species have a close affinity to only one black swifts, and pine grosbeaks distinguish the avifauna
physiographic province or ecoregion, but species richness of the north Cascades in Washington from those south of
and abundances of many species vary among provinces. the Columbia River. Although both hermit warblers and
Although direct comparisons of avian species richness Townsend’s warblers breed in both states, hermit warblers
across all provinces are not available, some general reach the northern extent of their contiguous range in
patterns are evident. Total bird abundance in western Washington, and Towsend’s warblers do not breed south
Oregon and Washington is generally correlated negatively of northern California. Thus, Townsend’s warblers are
with latitude and positively with longitude.21 Thus, mean more abundant in the Washington Cascades, while hermit
abundance and species richness of diurnal breeding birds warblers are prevalent in Oregon, and the entire region
is higher in the Oregon Coast Range than the Cascades of comprises a zone of hybridization for these two species.
Oregon and southern Washington,21 and species richness The Klamath region in southern Oregon stands out
is highest in the Klamath region. Characteristics of the from the rest of the Pacific Northwest for two reasons:
avifauna that distinguish each province are described high plant species diversity, and the convergence of several
below. ecological zones and their associated fauna. The highest
The avifauna of the tall, dense forests of the Coast Range avian species richness west of the Cascade crest in Oregon
and Puget lowlands shows some influence of the and Washington occurs in the Klamath Mountains. The
proximity to ocean, bays, and estuaries. The marbled high diversity of birds in this region has been attributed
murrelet is a unique example of this coastal influence, to the diversity of vegetation, and in particular the
being the only seabird that nests in forest habitats. Limited abundance of hardwoods.43 Several avifaunas converge
by the necessity to feed at sea, this species nests only within in the Siskiyou-Klamath mountains. Species such as
60 miles of the ocean, and predominantly in old-growth Allen’s hummingbird, black phoebe, oak titmouse, and
forests where branches of sufficient diameter provide nest blue-gray gnatcatcher reach the northwestern extent of
platforms. No bird species is particularly associated with their geographic ranges in this region. Species that are
the coastal band of Sitka spruce, but wrentits are notable typical of pine habitats to the south and east (e.g., Lewis’
for the area because they do not regularly occur west of woodpecker, white-headed woodpecker), and of Great
the coastal scrub along the southern and central Oregon Basin habitats (e.g., calliope hummingbird, green-tailed
coast. towhee, ash-throated flycatcher) occur in forested areas
As in the Coast Range, dense coniferous forests within this region. Other representatives of southern or
dominate habitats of the lower west slopes of the Cascade arid regions are more abundant here than further north
Range, so it is not surprising that the avifaunas of these (e.g., acorn woodpecker), but the full compliment of
two ecoregions are very similar. In Oregon, a slightly lower species that are characteristic of moist coastal forests also
average avian species richness on the lower west slopes occur (e.g., varied thrush, marbled murrelet, hermit
of the Cascades21 may be partially attributable to a lower warbler).
abundance of broad-leafed deciduous trees, such as big-
leaf maple and red alder, than in the Coast Ranges.35 On
the other hand, some habitats that are unique to the Patterns within Landscapes
Cascades contribute some distinctive members to the bird Narrowing our focus from the broad regional scale
community. For example, two duck species, bufflehead perspective of species distribution patterns to a finer
and Barrow’s goldeneye, breed on lakes in the high within-landscape approach (i.e., within the five Level III
Cascades, and are considered forest associates because western forest ecoregions) allows a more concise
they use cavities, usually in snags, for nesting. Harlequin discussion of wildlife-habitat relations. Whereas coarse-
ducks are associated with fast-moving mountain streams, grained elements such as legacy disturbance effects and
and find ideal breeding habitat along drainages of the west regional climate gradients may help our understanding
slope of the Cascades. Breeding populations of these three of regional taxonomic diversity trends, individuals of a
duck species are patchy and local throughout the Cascade species survive and reproduce at finer scales. In the
Range in Oregon and Washington. Other species that are western and montane forests of Oregon and Washington,
more likely to occur in the Cascades than in other micro- to macrohabitat conditions at the forest stand level
provinces include golden eagle and goshawk. are of critical importance to the individual. Aggregating
Changes in bird communities that can be attributed to up in biological organization, populations similarly
elevation are more obvious in the Cascades and Olympics function within these narrower bounds. In this section,
than in the Coast Ranges. For example, Clark’s nutcrackers the dominant finer scale habitat associations of western
do not occur below the high elevation spruce-fir forests forest wildlife within landscapes are presented, including
in the Cascades, and mountain chickadees replace forest plant species and stand structural conditions. Key
chestnut-backed chickadees above the Douglas-fir zone.16 habitat elements within western forests to which wildlife
The presence of species such as great gray owls, boreal have strong ties (e.g., logs, rock substrates, litter, snags,
owls, black-backed woodpeckers, and boreal chickadees and large trees) are distinguished. Old growth, young seral
in the high Cascades (>5,000 ft [1,524 m]), North Cascades, stages, riparian forests, and forest edges are highlighted
and high Olympics indicates the boreal influence on these because of their roles as wildlife habitat hotspots for
 194 Wildlife–Habitat Relationships in Oregon and Washington

various taxa within our current managed landscapes. At Forest Habitat Types and Structural
this finer spatial scale, heterogeneity among these habitat Conditions
types and microhabitat features remains a dominant driver Western forests are a complex mix of vegetative conditions.
of the western wildlife species diversity. Herb, shrub, and canopy tree structure and composition
Current knowledge of wildlife species’ use of habitats are key predictors of the occurrence of various wildlife
and general ecology in Oregon and Washington was species. Wildlife species were assessed relative to 4 western
compiled by panels of species-experts and is presented in forest habitat types: (1) Westside Lowlands Conifer-
several matrixes on the CD-ROM accompanying this book. deciduous Forest; (2) Westside Oak and Dry Douglas-fir
This expert knowledge includes data supported by Forest and Woodlands; (3) Southwest Mixed Conifer-
research, personal observations, or expert opinion. In the Hardwood Forest; and (4) Montane Mixed Conifer Forest.
tables below, these data are summarized to show wildlife Fairly similar numbers of total species occurred in these
relationships with forest habitat type, structural habitats (Table 1), with differences reflecting some of the
conditions, habitat elements, and trophic and organismal larger scale patterns already discussed (e.g., reptiles).
relations.

Table 2. Western forest wildlife habitat associations with vegetation height and successional stage.1

Taxon Grass/forb Shrub/seedling Sapling/pole Small trees Medium trees Large & giant trees

Amphibians2 19 19 20 21 28 28
Reptiles3 19 16 16 18 18 17
Birds3 61 92 98 130 140 140
Mammals3 66 65 64 72 79 78
Total 165 192 198 241 265 263
1
Data are compiled from expert panel assessments for this book, see accompanying CD-ROM. Numbers of species are indicated.
2
These species require specific habitat elements to occur within these structural conditions.
3
These species may require specific habitat elements to occur within these structural conditions.

Table 3. Numbers of western forest wildlife species associated with tree size (large includes giant trees) and
canopy complexity (single = single story canopies, multi = multiple story canopies).*

Taxon Small trees Medium trees Large trees

Canopy Cover single multi single multi single multi

Amphibians
Open 21 21 22 22 24 22
Moderate 21 21 22 28 24 28
Closed 20 21 22 28 28 28
Reptiles
Open 17 18 16 15 17 16
Moderate 10 9 10 9 10 9
Closed 6 6 6 6 6 8
Birds
Open 106 96 115 104 121 114
Moderate 89 85 97 101 102 104
Closed 68 67 74 76 76 96
Mammals
Open 61 62 68 58 71 73
Moderate 54 54 65 60 69 68
Closed 33 41 43 47 53 65
* Data are compiled from expert panel assessments for this book.
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 195

Table 4. Numbers of species using (use) and closely associated (close assoc.) with old-growth forests
in western Oregon and Washington, and their relative dispersal ability
(stand is within approximately 60 acres [24 ha]; range = across provinces).1

Taxon Use Close assoc. Close assoc. Close assoc. Dispersal of close assoc.
in Oregon inWashington

Amphibians 312 16 16 10 Stand

Reptiles 10 0 — — —
Birds 119 38 38 37 Stand to Range
Mammals 67 26 21 21 Stand to Range
Total 227 80 75 68
1
Data compiled from Thomas et al.,51 and includes species from northwestern California forests.
2
The former Olympic salamander is represented here as the 4 torrent salamander species.

Within these 4 habitats, forest/woodland structural related to vertical stratification: large tree size, multiple
conditions were identified. Three components were used story, and closed canopies. These create habitats or
to classify structure: vegetation height and successional conditions to which some old-growth associates are reliant
stage (i.e., grass/forb, shrub/seedling, sapling/pole, and (see Habitat Elements below).
small-to-giant tree categories); number of canopy layers Some old-growth associated species do not spend their
(single or multiple); and canopy cover (open, moderate, entire life in these forests. For example, neotropical
closed). Several patterns of wildlife associations with these migrant birds spend their winters in Mexico or Central
components are apparent (Tables 2 and 3). Total species America (7 species), and some waterfowl migrate in winter
richness increases with vegetation height, and this holds to lowland bays, lakes, and surf zones (6 species). Similarly,
true for each taxon except reptiles (Table 2). Bird species many aquatic-breeding amphibians reproduce in lentic
show a particularly dramatic response to vegetation and lotic waters within the forested landscape, and move
height, successional stage, and canopy layers. Mammals to the upslope forest matrix after metamorphosis from
and amphibians show a similar but dampened pattern. their aquatic stages.
All 4 wildlife groups show a pattern with canopy closure Early successional stands are used by many western
(Table 3). Greater numbers of birds, mammals and reptiles forest-dependent wildlife. From Table 2, herb, shrub and
are associated with open rather than closed canopies. The sapling conditions are used by a high percentage of species
reverse is apparent for amphibians. in every taxonomic group. Open conditions across tree
Associations of western forest fauna with forest age- size categories are especially important for birds and
size or successional stage categories have been analyzed mammals (Table 3). Similarly, 13 (25%) of 53 mammal
in numerous studies and assessments across the region. species were associated with early seral stages in the
Results from these studies and existing knowledge of Augusta Creek watershed, in the western Oregon Cascade
species’ forest associations were compiled for the Report Range.18 McGarigal and McComb27 reported about a third
of the Scientific Analysis Team51 for areas within the range of bird species (n=99) used early seral forests in western
of the northern spotted owl. The particular assignment Oregon.
for this assessment was to identify those species likely to Riparian forests are critical to the life history of
be closely associated with late-successional and old- numerous vertebrates in western Oregon and Washington.
growth forest conditions, from the longer list of those that The western forest landscape in the Pacific Northwest is
used forest habitats (Table 4). Close associates met one of highly dissected by stream channels, and the stream-
several criteria, such as having greater abundance in old- upslope interface is not far from any locale in the forest
growth forest than in mature or pole stands and requiring matrix. Riparian areas are distinguished by their cool,
habitat components that are contributed by old-growth moist environments, and by their multitude of conditions
forest (Table 5-1 in Thomas et al.51). From their data across a watershed. Small headwater streams are
compilation, about a third of the vertebrates using western ecologically distinct from their downstream mainstem
forests were identified as likely close associates with the counterparts. Along this entire aquatic network, riparian
older forest conditions (Table 4). No reptiles and about areas are prone to small-scale disturbances, such as
half the amphibians were close associates. These patterns flooding, bank and slope slippage, landslides, and
may be reflected in the current habitat assessments. About treefalls. These add heterogeneity to riparian forest
a third more forest wildlife species occurred in association conditions. Heterogeneity also is added as slope gradients
with large trees than the earliest successional stages. vary, as streams flow through constrained and
However, there was only about a 10% increase between unconstrained reaches, and with temporal variation in
small and large trees (Table 2). Amphibians were more foliage cover (i.e., deciduous trees) and rainfall patterns.
associated with closed canopies. Structural conditions of Unconstrained reaches and tributary junctions, in
old-growth forests include components in Table 3 that are particular, are proposed as higher species diversity areas
 196 Wildlife–Habitat Relationships in Oregon and Washington

Figure 3. Four terrestrial salamanders restricted to forested

habitats of the Pacific Northwest. These species are covered
under the federal Survey and Manage provision of the
Northwest Forest Plan, for which surveys are conducted prior
to ground disturbing activities and known sites are currently
managed for salamander persistence.34, 54, 55 W. P. Leonard,
photographer. R. S. Nauman, GIS technician.
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 197

Table 5. Primary habitat element associations of species with close ties to old-growth forest conditions.1

Taxon n Talus Logs Duff/litter Large snags Large trees Riparian

Amphibians 16 12 11 9 0 0 132
Birds 38 0 7 0 25 10 8
Mammals 26 4 14 7 14 10 11
Total 80 16 32 16 39 20 32
1
n = total number of species per taxon. Species may be represented in multiple habitat element columns. Compiled from Thomas et
al.51 (includes northwestern California species).
2
The former Olympic salamander is represented here as the 4 torrent salamander species.

for several aquatic-dependent taxa. As habitat complexity on riparian habitats for foraging or refuge. Assessments
may beget diversity, riparian forests are predictable species in Thomas et al.51 determined that 29 (36%) of 80 old-
hotspots in western forests because they may encompass growth forest associated vertebrates were strongly
numerous habitat and microhabitat conditions. associated with riparian areas (Table 5). These include 53%
As discussed for old-growth forests above, whereas of the amphibians (10 species, primarily stream- and pond-
some taxa use riparian areas for their entire lives, others breeders, and bank associates), 42% of the mammals (7
are users for critical life history functions. For example, bats, Pacific shrew, shrew-mole, deer mouse, marten), and
several myotis bats use riparian corridors for foraging 21% of the birds (marbled murrelet, bald eagle, and 6
habitat and travel corridors, but roost upslope. The waterfowl species) that were assessed. As can be seen in
terrestrial stages of aquatic-breeding amphibians may rely Table 5, riparian-associated species include users of several
forest habitat elements (talus, logs, snags, large trees).
Extending the diversity hotspot concept to within-
landscape, forest stand-scale areas, both old-growth and
riparian forests are areas of higher richness and diversity
for many taxa. However, overall reduced breeding bird
diversity was found along streams in comparison to
upslope forest habitats by McGarigal and McComb26 in
the Oregon Coast Range. They suggested differences in
vegetation structure and composition could explain their
results: fewer large conifers and snags were noted along
their study streams. More generally, riparian areas within
late-successional or old-growth conditions are particularly
high in wildlife numbers. An understanding of the habitat
elements and microclimate components of these areas is
important to fully grasp the habitat complexities of these
areas. Riparian-dependent wildlife are discussed further
in Chapter 13.

Habitat Elements
Several forest habitat elements are used for foraging,
refuge, or reproduction by western forest wildlife. These
include features such as downed logs, snags, duff or litter,
and rocky substrates. They also include biotic or live
vegetative components, including shrubs, live tree
branches, or live remnant trees. Many of the old-growth
forest associated species may occur in younger stands if
these critical habitat elements are found there. Table 6
summarizes habitat elements associations for forest
wildlife. Table 5 shows the primary habitat elements of
old-growth associated vertebrates, compiled from Thomas
et al.51 Each element in Tables 5 and 6 can be considered a
dominant contributor to western forest wildlife habitats
Figure 4. Satellite image of the Pacific Northwest. Western across taxa. Microhabitats provided by logs and snags,
and montane forests are shown as darker areas on the left, and rock, duff and litter, cavities, shrubs, and large trees are
are coincident with the range of the northern spotted owl and
the Northwest Forest Plan. M. Fiorella, GIS technician. Text continues on page 201
 198 Wildlife–Habitat Relationships in Oregon and Washington

Figure 5. Level III and IV

ecoregions for western
Oregon and Washington,
compiled at a scale of
1:250,000 by Pater et al.42
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 199

1 Coast Range 4 Cascades

1a Coastal Lowlands 4a Western Cascades Lowlands and Valleys
1b Coastal Uplands 4b Western Cascades Montane Highlands
1c Low Olympics 4c Cascade Crest Montane Forest
1d Volcanics 4d Cascades Subalpine/Alpine
1e Outwash 4e High Southern Cascades Montane Forest
1f Willapa Hills 4f Umpqua Cascades
1g Mid-Coastal Sedimentary 4g Southern Cascades
1h Southern Oregon Coastal Mountains
1i Redwood Zone
9 Eastern Cascades Slopes and Foothills
9a Yakima Plateau and Slopes
2 Puget Lowland 9b Grand Fir Mixed Forest
2a Fraser Lowland 9c Oak/Conifer Eastern Cascades Columbia Foothills
2b Eastern Puget Riverine Lowlands 9d Ponderosa Pine/Bitterbrush Woodland
2c San Juan Islands 9e Pumice Plateau Forest
2d Olympic Rainshadow 9f Cold Wet Pumice Plateau Basins
2e Eastern Puget Uplands 9g Klamath/Goose Lake Warm Wet Basins
2f Central Puget Lowland 9h Fremont Pine/Fir Forest
2g Southern Puget Prairies 9i Southern Cascades Slope
2h Cowlitz/Chehalis Foothills 9j Klamath Juniper/Ponderosa Pine Woodland
2I Cowlitz/Newaukum Prairie Floodplains

77 North Cascades
3 Willamette Valley 77a North Cascades Lowland Forests
3a Portland/Vancouver Basin 77b North Cascades Highland Forests
3b Willamette River and Tributaries Gallery Forest 77c North Cascades Subalpine/Alpine
3c Prairie Terraces 77d Pasayten/Sawtooth Highlands
3d Valley Foothills 77e Okanogan Pine/Fir Hills
77f Chelan Tephra Hills
77g Wenatchee/Chelan Highlands
Level III ecoregion 77h Chiwaukum Hills and Lowlands
Level IV ecoregion 77i High Olympics
County boundary
State boundary
International boundary 78 Klamath Mountains
78a Rogue/Illinois Valleys
78b Siskiyou Foothills
78c Umpqua Interior Foothills
78d Serpentine Siskiyous
78e Inland Siskiyous
78f Coastal Siskiyous
Universal Transverse Mercator Projection, Zone 10 78g Klamath River Ridges

Principal authors: David E. Pater (Dynamac Corporation), Sandra

A. Bryce, (Dynamac Corporation), Thor D. Thorson (NRCS),
Jimmy Kagan (Oregon Natural Heritage Program), Chris
Chappell (Washington DNR), James M. Omernik (USEPA), Sandra
H. Azevedo (OAO Corporation), and Alan J. Woods (Dynamac
Corporation).
Collaborators and contributors: Terry L. Aho (NRCS), Duane
Lammers (USFS), Thomas Atzet (USFS), Robert Meurisse (USFS),
Kenneth Radek (USFS), Carl Davis (USFS), Thomas Loveland
(USGS), M. Frances Faure (OAO Corporation), and Jeffrey A.
Comstock (OAO Corporation).
This project was partially supported by funds from the USEPA –
Office of Research and Development – Regional Applied
Research Effort (RARE) program.
 200 Wildlife–Habitat Relationships in Oregon and Washington

Amphibians Reptiles

<5 <6
5-6 6-8
7-13 9-11
14-15 12-15
16-18 16-19
Fish

<8
8-13
14-19
20-29
30-50

Mammals

Butterflies

<58 <65
58-59 65-76
60-61 77-93
62-64 94-102
65-75 103-117

Figure 6. Species richness maps for Oregon, derived from data

compiled for the Oregon Biodiversity Project41 CD-ROM.
Species richness (numbers of species) categories per
taxonomic group per hexagon (approximately 150,000 acres
[60,750 ha]) are shown. Bird data were not available.
(Courtesy of R.S. Nauman, GIS technician)
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 201

Figure 12. Satellite images of the

eastern Oregon Cascade Range,
showing the Three Sisters Wilderness
from an eastern perspective. Upper
image shows forests of the eastern
Cascade ecoregion (red) and areas of
downslope forest management. Lower
image shows the likely impact areas of
recreation (within 328 ft [100 m])of
roads and trails, focussed along
riparian areas). M. Richmond, GIS
technician.

key elements for forest-dwelling species. It is interesting particularly large down logs, are the habitat elements most
to note that there are several differences between the frequently used by amphibians and mammals. Large
results of these two assessments (Tables 5 and 6). Table 6 wood decays gradually, its rate dependent on the tree size,
shows that many more species use these habitat elements ambient conditions, and tree species. Wildlife species may
in western forests than the more limited pool of species have affinities for specific wood decay classes.
that are both closely tied to old-growth forest conditions For vertebrates, wood provides foraging, cover, and
and associated with these elements (Table 5). Criteria for sites for reproduction. For some species within each
use versus association are different. Among elements vertebrate class, large downed wood provides thermal
considered in both assessments, logs and snags were the refugia, buffering temperature and moisture extremes.
dominant habitat elements represented in Table 5, whereas Blessing et al.1 demonstrated the temperature buffering
surface rock and live trees also are dominant in Table 6. capacity of a log, 20 in x 13 ft (50 cm x 4 m), containing a
Dead and downed wood occurs in numerous forms Van Dyke’s salamander nest site. While ambient air
(Figure 9). In western forests, downed wood includes logs, temperatures ranged 43-76½F (6.3 to 24.7½C) in the shade
rootwads and stumps, wood piles and slash, roots, 6.6 ft (2 m) from the log, temperatures inside the log cavity
branches, loose bark, and bark piles. Different species tend at the salamander nest ranged 46-63½F (7.7-17½C). For
to be associated with the different downed wood types. several days in the summer, the maximum nest
Among amphibians, for example, both ensatina and temperature was cooler than the minimum outside air
clouded salamanders can be found within decaying logs, temperature. For amphibians, logs provide cool, moist,
but the clouded salamander is frequently found under the and stable microhabitats suitable for their physiological
bark of logs and the ensatina is often found in bark piles temperature and moisture requirements. Some
associated with decaying snags. Coarse woody debris, plethodontid salamander species have limited home
 202 Wildlife–Habitat Relationships in Oregon and Washington

Landscape Plan Interim Plan

Terrestrial reserves Terrestrial reserves

Management Aquatic reserves Riparian reserves
Areas Landscape area 1 Matrix
Landscape area 2 Non-National Forest
Landscape area 3
Non-National Forest

Future Conditions

Young, light overstory

Young, heavy overstory
Mature
Old
Non-forest

Figure 13. Management areas and projected future landscape

structures for the Blue River watershed managed under
Matrix and Riparian Reserve designations of the Northwest
Forest Plan54 (“Interim Plan”), and for the Blue River
landscape management strategy (“Landscape Plan”).14 T.
Turner, GIS technician.

ranges, remaining at log sites for indefinite periods. They important for cavity excavators, and for species using
find sufficient foraging opportunities at logs, in addition snags for foraging. Some snag-users have preferences for
to using them for cover and reproduction. For mammals, snag size. Protection from predation is considered a
downed wood habitats similarly provide resting, nesting selective force of snag use, and again, thermal buffering
and denning, and foraging habitat for numerous species is thought to be a critical component of these habitat
(Table 6). Of the 14 log-associated mammals listed in Table elements. Standing dead trees within intact stands will
5, 2 species are bats that potentially use logs for roosting, provide suitable thermal refugia for species sensitive to
and the remaining 12 species, 10 rodents and insectivores temperature extremes. However, some species tend to use
and 2 forest carnivores, use the downed wood for multiple snags in open conditions, or in a variety of closed and
life history functions. Although fewer birds rely on logs, open forest types. Many bats have roosting and
their use of downed wood includes perching and lookout hibernacula in larger snags, some preferring snags with
in addition to foraging sites, cover, and nesting. The loose bark. Fishers use cavities for denning and resting,
sharptail snake is one of the few western forest reptiles preferring large snags. Cavity-nesting birds include many
associated with downed wood, and often is found in logs. waterfowl, owl, and woodpecker species.
Snags are used extensively by birds and mammals Duff, litter, live trees, and surface rock are additional
(Tables 5 and 6). Cavities, cracks, crevices, and loose bark western forest components required by various wildlife
on or in snags are used by numerous species as cover and species (Table 5 and 6). Many amphibians in western
resting, roosting and nesting sites. Snag decay class is forests use duff, litter and surface rock, but only 1 species
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 203

Table 6. Habitat elements used by forest wildlife in western Washington and Oregon, as determined during
the panel assessments for this book.

Taxon Surface rock Logs Duff/litter Snags Live trees Moss Cavities Shrubs

Amphibians 3 12 10 4 3 1 0 1
Reptiles 10 5 4 0 0 0 0 0
Birds 27 18 5 57 72 5 38 21
Mammals 35 50 19 22 37 4 22 11
Total 75 85 38 83 112 10 60 33

(clouded salamander, in California) has been found in nesting species of birds and mammals is another useful
trees. Rock associates include some plethodontid assemblage in this context. Assemblages have increasing
salamanders, such as the Del Norte salamander. Surface validity as distinct ecological entities when they are
rock can be covered by litter and not readily identified as populated by alternative members across landscapes that
a likely site.38 Mammals use all of these habitat elements, span multiple species’ ranges.
yet relatively little is known about the ecology of some For amphibians, 3 main assemblages are generally
arboreal mammals. The northern red tree vole is thought partitioned for the Pacific Northwest (Figure 10). These
to live almost exclusively within the forest canopy. are separated by breeding habitat: terrestrial; “pond”
Numerous birds use live trees. Cavities and crevices are a (inclusive of all lentic habitats; e.g., lakes and wetlands)
main refuge and nesting place whereas the various canopy (e.g., Figure 11); and “stream” (inclusive of all lotic waters;
layers are used for foraging, cover, and nesting. Large e.g., streams and seeps). From a finer-grained habitat
conifers may be of particular importance to several species. assessment of species in western and montane forests
(lower portion of Figure 10), we have added complexity
Forest Wildlife Assemblages to this model. In western forests, distinct assemblages of
Sorting wildlife species by their main habitat associations stream-breeders are found in association with stream size.
helps us understand their roles in western forest In headwater streams and seeps, Rhyacotriton torrent
ecosystems. Such assemblages or communities can be salamanders dominate assemblages. Cope’s giant
derived from the classifications of taxa relative to the salamanders and tailed frogs may occur in some
habitat parameters presented here, including forest type, headwater channels and seeps as well. Larger downstream
structural components, and habitat elements. The scale at channels are dominated by Pacific giant salamanders, co-
which assemblages are identified might vary with the occurring with cottid and salmonid fishes in many
context that is being examined. The reptile assemblage of systems. The stream bank community is distinct, often
the Klamath ecoregion might be identified for 1 purpose, comprised of terrestrial-breeding Dunn’s salamanders,
whereas in another vein Klamath woodland snakes might western red-backed salamanders, and Van Dyke’s
be distinguished. Ground-dwelling small mammals and salamanders. We have split the terrestrial assemblage into
forest canopy mammals are examples of 2 assemblages two groups: the rock and downed wood associates. Both
around which hypotheses of ecological function may be groups are highly fossorial, spending much of their time
developed (Case History 2, below). Given the large subsurface. When temperature and moisture regimes at
numbers of species using snag or tree cavities, cavity- the forest surface are suitable, they can be found in

Figure 9. Downed wood provides habitat for

multiple vertebrate species. H. J. Andrews
Experimental Forest, western Oregon
Cascades. J. Means, photographer.
 204 Wildlife–Habitat Relationships in Oregon and Washington

Figure 11. Red legged frog (Rana aurora) is a pond breeder in

western forests. W. P. Leonard, photographer.

inhospitable warm, dry surface conditions away from

aquatic habitats. Pond-breeding amphibians such as
Cascades frogs often are found along montane creeks in
the summer. Foraging opportunities may be enhanced in
such locations, and they may function as dispersal
corridors. The relatively high species richness found along
banks also might result from this being an edge between
habitat types. As a boundary between habitats, you might
be more likely to find members of neighboring
assemblages along its interface. Or alternatively, as an
edge, this region may represent a barrier to further
movement.
Among terrestrial forms, rock and wood associates
have the potential for relatively high habitat overlap as
individuals opportunistically use cover as it is available
Figure 10. Fundamental (top) and finer-grained (bottom) and as suitable microclimate conditions warrant. This
habitat assemblages of western and montane forest occurs in two ways. First, within populations, there might
amphibians. be use of both wood or rock, depending on its availability
and suitability as habitat cover. Individuals in such
association with wood or rock cover. At some locations, populations might use the different cover types for
these animals may be found year-round if suitable surface different functions, such as dispersal cover, foraging areas,
refugia are available. Wood associates include the slender or reproduction. Dispersal cover, in particular, might be
salamanders, the black salamander, and those species used more opportunistically. Second, for some species,
associated with logs that were mentioned previously. Rock there seems to be segregation of habitat use geographically
associates include the plethodontids in Figure 3. As stream among populations. For example, the Larch Mountain
or pond breeders move into upslope forests, they often salamander is found in association with surface rock along
opportunistically use both wood and rock microhabitats, the Columbia River Gorge, but is found associated with
as well as subsurface retreats. downed wood and loamy soils elsewhere.15 Similarly, the
Overlap among assemblages is considerable for Del Norte salamander appears to be a rock associate at
amphibians because of their complex life history. Adults inland locations, and can be found with downed wood at
are generally not restricted to breeding habitats. Thus, coastal sites.38 The critical factor in this might still be habitat
stream and pond breeders venture from the aquatic and availability and suitability. Suitable microclimate and
riparian forest landscapes into upslope forests. Likewise, microsite conditions for these terrestrial salamanders
terrestrial forms occur in riparian zones and may be found might occur through combinations of either of these cover
streamside. The bank seems to be the primary area of types.15, 38
assemblage overlap, as members of all assemblages are We discuss western forest wildlife assemblages again
found in this near-water riparian zone. This may occur below. First, as trophic relations are identified,
because bank conditions in western and montane forests assemblages may link to ecological functions and
may be almost ideally suited for amphibians, having cool processes (see also Case History 2). Second, assemblages
moist microclimates. During summer, in particular, banks often are used during the development of protective
might offer surface refugia for amphibians from measures for wildlife when forest management activities
are proposed.
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 205

Species’ Life History, Behavior, Table 7. Distribution of species by trophic

and Biotic Interactions relationships in western Washington and Oregon.*
Distribution patterns of western forest wildlife are highly
dependent on several aspects of their behavior and Taxon Primary Secondary Tertiary Carrion
ecology. In particular, species’ life history, behavior, and consumer consumer consumer feeder
species interactions may need to be understood to fully
explain species-habitat relations. Amphibians 10 29 0 0
Dispersal limitations seem to contribute to the isolation Reptiles 1 20 0 2
of many amphibian populations. Amphibians generally Birds 96 161 6 8
are reliant on relatively narrow windows of temperature Mammals 70 62 1 12
Total 177 272 7 22
and moisture conditions for surface activity. Spring and
fall rainy seasons are primarily when dispersal occurs for * Data are compiled from expert panel assessments for this
most taxa. Compounded by their relatively low mobility, book.
amphibians’ survival during migrations across
heterogeneous forested landscapes can be affected. on only small slugs, and most small salamanders prey on
However, few studies on western forest amphibian various invertebrates.
dispersal have been conducted. Some taxa may move Amphibians and mammals are likely key vertebrate
hundreds of feet to several miles (Ambystoma, Taricha, connectors within the western forest system. Both groups
Dicamptodon, and Rana spp.). The terrestrial plethodontid are the chief prey for secondary predators (= tertiary
salamanders are considered less vagile. Studies of some consumers), and as primary consumers and primary
log-associates found movement of only a few feet over predators are conduits of energy from the lower trophic
seasons to years. levels, particularly the diverse arthropod and fungal
A tendency for site fidelity may interact with amphibian communities. In eastern U.S. forests, amphibians may
dispersal capabilities. There may be philopatric tendencies comprise a major component of the vertebrate biomass;3
among both the more and less mobile amphibians. Many however, such a biomass estimation has not been done
toads known to trek miles in montane forests have high for western Oregon and Washington forests. Rodents
breeding site fidelity. Traditionally used breeding sites are comprise about half the mammal species of the region,
common in pond breeding salamanders and frogs, and functioning as prey for numerous species. Bats are
may occur in the other forest amphibian assemblages. believed to consume enormous quantities of insect prey,
Mammals and birds seem to be less restricted in their mainly over streams, ponds, and riparian areas. Shrews
movements (Table 3), yet many have relatively small home and moles are carnivorous, eating predominantly
ranges (i.e., <10 acres [4 ha]27). Even those with large home arthropods. The fungi-feeding mammals link the
ranges may be restricted in their movements by perceived vertebrate trophic network with the diverse fungal
barriers on the landscape. Clearcuts and roads are community, of pivotal importance in the forest
suspected barriers for some species. ecosystem.52 Fungi also are food for microorganisms and
Interactions among species are the threads that weave invertebrates, and have critical forest functions as
the fabric of the integrated living forest ecosystem. decomposers, nutrient cyclers, and through their
Intraspecific interactions are often dominated by mutualistic or symbiotic relationships with other forest
competition for food, space, and mates, and interspecific species.28
interactions involve both competition and predation. Both Competition and trophic relationships are only a part
intra- and interspecific interactions may displace of the interspecific interactions among wildlife in forests
individuals or populations, affect survivorship, and result (Table 8). Wildlife mediate the reproduction of numerous
in altered distribution and abundance patterns across the forest species as dispersers or pollinators. Birds and
landscape. mammals are dominant players in organismal
Trophic relations are established for most western and relationships (Table 8). Their actions can alter habitats
montane forest wildlife species (Table 7). Typical of food significantly, making them either suitable or unsuitable
webs, most numerous among forest wildlife are the for other species. For example, cavity, burrow, and runway
primary consumers (herbivores) and secondary creators provide critical habitats for many other wildlife
consumers (primary predators). Primary consumers eat a species.
host of plant material (e.g., leaves, seeds, sap, roots, bark, The stability of vertebrate communities can be
fruit). Primary consumers are birds (58% of this taxon) dependent on the balance of intraspecific, trophic, and
and small-to-large bodied mammals (e.g., rodents, deer, other organismal relations with species’ habitat conditions.
elk; 67% of mammals), and larval pond-breeding Measures of biotic integrity attempt to determine the status
amphibians. Secondary consumers prey on invertebrates, of communities by assessing indicator conditions or
vertebrates, and eggs. In western forests, some mammals, species. These can be examined relative to disturbance
most birds and all reptile and amphibian species are impacts, such as forest management activities, to monitor
secondary consumers (Table 7). Many of these are gape- forest system functions. In western forests, arboreal rodent
limited predators that often change diet opportunistically and forest-floor small mammal assemblages are suggested
with their size or age. Sharptail snakes are noted as feeding to be ideal indicators of forest biotic integrity (Case History
2).
 206 Wildlife–Habitat Relationships in Oregon and Washington

Table 8. Number of species by organismal relationships in western Washington and Oregon.*

Organismal relationship Amphibians Reptiles Birds Mammals

Controls insect populations 8 0 29 13

Controls terrestrial vertebrate populations 1 1 14 12
Pollination vector 0 0 6 0
Transportation of seeds, spores, plants or animals 0 0 65 36
Nest parasite 0 0 40 0
Creates feeding, roosting, denning, or nesting sites for other 0 0 4 4
species
Creates structures 0 0 13 11
Uses structures created by other species 1 3 5 11
Cavity excavator 0 0 14 1
Cavity user 0 0 29 12
Burrow excavator 0 0 0 44
Burrow user 0 9 0 38
Runway creator 0 0 0 32
Runway user 0 0 0 33
Pirates food from others 0 0 4 0
Interspecific hybridization 0 3 12 6
* Data are compiled from expert panel assessments for this book.

are listed in Carey and Johnson.8 This community can be

Case History 2 described with various techniques: live trapping, pitfall
Biotic Integrity trapping, and snap trapping.
Andrew B. Carey Another interesting community is the arboreal rodent
community. In Washington, the community consists of
How can the effectiveness of ecosystem and landscape northern flying squirrel, Douglas’ squirrel, and
management be evaluated? One attractive method is to Townsend’s chipmunk. In Oregon, one has the option of
measure the integrity of select vertebrate communities.10 adding bushy-tailed woodrat, dusky-footed woodrat, and
Ideally, these communities would consist of a limited red tree vole. Individually and collectively, the biomass
number of year-round resident species, common enough of these species is indicative of carrying capacities for a
to be found in most, if not all, patches of suitable habitat, variety of vertebrate predators including mustelids,
yet sensitive enough to management that >1 species would hawks, and owls. In addition, in managed forests,
be absent or severely reduced in abundance in unsuitable community arrangement will diverge from the high
or low-quality environments. A limited number of species abundances of all members found in old growth, to
is desirable because operational practicality requires use communities dominated by 1 or 2 species when
of one technique of high reliability. Year-round resident management has failed to adequately address key
species are desirable because population levels of migrants ecosystem components.6 In particular, the arboreal rodent
incorporate variability due to wintering and migration community measures ecological productivity—the energy
factors that are independent of the area being managed. that the system of fungi, understory plants, and overstory
Relatively high abundance is required to ensure that all trees diverts to reproduction (truffles, mushrooms, fruits,
species can be measured effectively when present. Two seeds, and nuts). This community integrates production
communities fit these requirements: forest-floor small of fruit with the decadence process that produces cavities
mammals and arboreal rodents. and affects spatial arrangements of habitat elements. This
Forest-floor small mammals are interesting because results in niche diversification. Thus, the sum of the
complete communities with moderately high abundances population sizes of species in the arboreal rodent
of each species depends on Hutchinsonian preinteractive community measures, in large part, the energy the system
niche diversification—in other words, biocomplexity in is putting into reproduction.9 This community (with the
the structure and processes of the forest floor. Thus, the exception of red tree vole) can be effectively described with
community represents diverse forest-floor functions, live trapping.5
abundance of coarse woody debris, and understory Arboreal rodents and forest-floor small mammals can
development. On the Olympic Peninsula in Washington, be used both to monitor effectiveness6 but also to predict
for example, the ranked relative abundances of the species results of forest management strategies through
in the community are Trowbridge’s shrew (5), southern modeling.10 The use of biotic integrity, when supported
red-backed vole (4), montane shrew (4), deer mouse (4), by basic research, offers an alternative approach for
forest deer mouse (3), shrew-mole (3), creeping vole (3), populations of rare, cryptic, or other species that are
and vagrant shrew (2). Assemblages in other provinces difficult to study, survey, and manage.
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 207

Management Issues
Western and montane forest management policies in woody debris management, specified snag and green tree
Oregon and Washington are undergoing tremendous retention levels, and forest management for red tree vole
changes, largely to address the long-term persistence of and northern spotted owl dispersal corridors. For species
species and to ensure the maintenance of ecosystem that were not adequately protected by these series of
integrity. The main human uses of western and montane measures, a “Survey and Manage” provision was
forests include timber production, water resources, special created.53, 54, 55 For those of the most concern, sites proposed
forest products, recreation, mining, and the associated for ground disturbance would be surveyed for the species
support systems of roads and trails (Figure 12, page 201). of interest, and if found, managed to maintain the
These forests also maintain the treasuries of biophysical persistence of that species54, 55 (Figure 3). These protection
legacies, functions, and processes having aesthetic, ethical, measures are additive, together addressing the long-term
and cultural values within our society. However, across persistence of old-growth associated species on the federal
this landscape, timber management has been the dominant forest landscape in this region.
focus on both public and private lands for >100 yrs. By Since implementation of this Plan, alternative
the 1970s and 1980s, concern for high profile species such landscape management plans have developed. The
as the northern spotted owl grew as late-successional and measures developed for the Northwest Forest Plan might
old-growth forests were diminished and fragmented be considered experimental, since nothing of that scope
because of logging, and as the U.S. Endangered Species has ever before been attempted in this region. The Plan
Act (ESA) of 1973 prohibited harm to species listed under may not be the sole means to maintain ecosystem integrity
the Act, and to their habitats. Studies were initiated to and biological legacies in western forests, while also
understand main habitat associations of the northern providing timber products and other socioeconomic
spotted owl, as well as the requirements of numerous other values. For the federally administered Augusta Creek
suspected obligates to the ancient forests of this region. watershed, an alternative scenario of forest management
By 1994, >1000 taxa were identified as likely associates of was developed using the natural disturbance processes
old-growth forest habitats. (i.e., fire) of the landscape as criteria for forest
management.13 Forest rotations and green tree retention
Federal Forest Management levels were matched to fire frequency intervals (100, 200,
The range of the northern spotted owl has close and 300 years) and intensities (15-50% retention). The
resemblance to the western and montane forests Plan’s Riparian Reserve system was reduced to provide
considered in this chapter (Figure 4), but also extends buffers along only the mainstem fish-bearing streams;
latitudinally beyond Oregon and Washington. In the U.S., however, tree retention would be weighted along other
42% of this landscape is federally administered (24.3 stream channels in harvested units. Aquatic reserves also
million acres [9.8 million ha]53). This landscape across 3 were placed in small basins for species or areas of concern.
states was used to develop the first ecosystem The resulting landscape was modeled and evaluated after
management plan for the nation which spanned land several hundred years and found to have advantages over
ownerships: the federal Northwest Forest Plan.54 The the Northwest Forest Plan landscape at the watershed
Plan’s goals included providing for the highest scale. Importantly, the fragmented and edgy “spaghetti”
contribution to the socioeconomic needs of the region landscape of the Plan (i.e., spatial pattern resulting from
while ensuring the long-term viability of the old-growth the highly dendritic stream network and its accompanying
forest ecosystem and associated species. Although the Plan Riparian Reserves, between which harvested units are
has been in place for 6 years, the balance of socioeconomic located) is consolidated into larger contiguous forest
productivity and protection of biological integrity is a blocks. Benefits for wildlife included improved habitat
challenge that is still being developed as we enter the 21st connectivity and maintenance of interior old-growth
century. The Plan is based on adaptive management, and conditions in much of the landscape because of reduced
thus the adaptive phase is expected to continue as new edge effects on microclimate.12 This scenario, with some
knowledge and management tools develop for western adjustments, is being tested in a neighboring watershed
forests. of the Willamette National Forest in the western Oregon
For species protective measures, the foundation of this Cascade Range (Case History 3).
Plan relies on several land allocations, habitat provisions,
and species-specific mitigations.54 A backbone of reserved
lands was created across the region for the maintenance Case History 3
of well-distributed populations of most of the broad The Blue River Landscape Study
ranging taxa considered. For habitats of fish and riparian- John H. Cissel and Frederick J. Swanson
dependent species of concern in the region, an Aquatic
A team of scientists and managers working on the H. J.
Conservation Strategy45, 54 was formulated, including
Andrews Experimental Forest and the Blue River Ranger
protections of key watersheds, development of watershed
District of the Willamette National Forest have been
analysis, watershed restoration, and identification of
cooperating for most of this decade to develop and test a
Riparian Reserves, primarily along streams and around
landscape management approach based on natural
unstable areas. Forest habitat provisions included coarse
disturbance regimes.13, 14 The team has been motivated to
 208 Wildlife–Habitat Relationships in Oregon and Washington

a significant degree by concern over the loss and implementation of the Northwest Forest Plan as if it were
fragmentation of older forests, and the lack of a coherent applied to the Blue River watershed (“Interim Plan”; e.g.,
long-term strategy for conservation of older forest systems Riparian Reserves occur on all streams, and 80-year
in managed landscapes. The underlying assumption of rotation regeneration harvests with 15% retention occur
this approach is that by approximating key aspects of in the “Matrix”54).
important disturbance regimes in management regimes, A watershed restoration strategy is an integral
risks posed to native species and ecological processes are component of the Blue River landscape management
reduced as compared to other historical and contemporary approach. Restoration activities are intended to reestablish
landscape management approaches.23, 29, 49 a resilient, interconnected aquatic network capable of
The Blue River Landscape Study is intended to evaluate maintaining aquatic habitats and processes while
the potential effects of implementing a landscape plan management activities are occurring in the watershed.
based on historical landscape dynamics. The landscape Road restoration activities are planned to occur first in
management approach used in the study is intended to areas where risks to aquatic ecosystems are high.
meet the same general objectives underlying the Future timber harvest and forest successional patterns
Northwest Forest Plan:54 provide habitat to sustain species were projected across the watershed for the next 200 years
associated with late-successional forests, maintain and for both plans (Figure 13, page 202) and analyzed. Results
restore aquatic ecosystems, and provide a sustainable show that the landscape plan will produce more late-
supply of timber. A combination of effectiveness successional habitat (71% of the watershed versus 59%)
monitoring, long-term plots, retrospective studies, and in a less fragmented landscape as compared to the interim
modeling assessments are being used to evaluate and plan.14 Larger patches in the landscape plan create more
adjust this landscape management approach. The Blue interior habitat, thought to benefit some wildlife species
River watershed provides an ideal setting for the study such as the northern spotted owl. Less edge between old
due to its size (approximately 56,790 acres [23,000 and young forests in the landscape plan reduces edge
hectares]) and the presence of the H. J. Andrews effects such as altered microclimates and increased plant
Experimental Forest and numerous long-term studies mortality, and may reduce habitat for certain species that
within the watershed. In addition, the Blue River favor edges, such as elk. More complex stand structures
watershed is a part of the Central Cascades Adaptive are present in the landscape plan due to generally higher
Management Area, a federal land allocation in the overstory canopy retention levels. Retention of live and
Northwest Forest Plan that encourages development and dead trees in young stands has been found to favor
evaluation of new approaches. cyanolichens, certain fungi and invertebrates associated
The dynamics of historical landscapes in this area were with older forests, amphibians with life histories requiring
heavily influenced by fire of varying frequency, severity both stream/riparian and upland habitats, provide more
and spatial extent. General patterns of past fire behavior options for protection of rare species, and to moderate
were interpreted into three fire regimes based on a 500- understory environments. The landscape plan also
year dendrochronological record.33, 50, 58 For example, wet, maintains a substantial component of mature forest (80-
cool sites burned infrequently while warmer, drier sites 200 years old). In contrast, the interim plan nearly
burned more frequently. Characteristics of these three eliminates the mature forest component because almost
regimes were used to establish timber and fire all lands are either in a reserve, where all stands grow old
management regimes in actively-managed portions of the and large-scale disturbance is eliminated, or in matrix
landscape. Timber harvest frequency and rotation age lands where a relatively short rotation (approximately 80
(100-260 years) were based upon historical fire frequency, years) prevents regrowth of mature forest. We feel that
timber harvest intensity (15-50% overstory canopy the absence of mature forest in the interim plan poses
retention) was based upon historical fire severity, and the substantial risk when mortality due to disturbance, climate
spatial patterns of timber harvest were based upon the change or senescence eliminates older Douglas-firs in the
spatial patterns of historical fires. Implementation reserves.
guidelines are intended to reflect natural disturbance Landscape structures resulting from both the landscape
patterns to the extent feasible while protecting ecological management plan in this study14 and from the interim plan
values. are historically unprecedented. For that reason we feel it
An aquatic reserve system also was established to help is critical that an adaptive management approach be
meet the aquatic ecosystem objectives in the Northwest followed for both plans. We are pressing ahead with
Forest Plan.45, 54 These reserves are of two types: small- implementation, monitoring, modeling, and research to
watershed reserves and corridor reserves. Small- better define and evaluate a historically-rooted approach
watershed reserves are strategically located throughout in the Blue River watershed based on the landscape
the watershed to encompass areas of particular importance dynamics inherent to the area. We hope these concepts
to aquatic ecosystems and spotted owls. In addition, can be tested in other provinces in the region, and that the
corridor reserves are established on all fish-bearing Matrix and Riparian Reserve approach of the interim
streams. Figure 15 depicts the landscape management plan plan54 can be similarly tested.
(“Landscape Plan”), and, for comparison, a literal
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 209

State and Private Forest Management protection is needed in different portions of the western
In the last decade, several forest management plans at forest region, by ownership and location, the science of
watershed to landscape scales have been designed by state forest management is rapidly changing.
agencies and industrial land owners. These alternative Sustainable forestry techniques currently being tested
approaches to forest management reflect the diverse across western Oregon and Washington forests are
alignment of roles as wildlife stewards of these land numerous and innovative. Forest density management
owners. On federal lands, the more conservative standard and alternative silviculture is being examined for both
has been set for species maintenance or restoration as a restoration and regeneration harvests by interagency
priority equal to or greater than providing economic collaborative partners. Selective harvests are being more
returns. On state lands, timber revenue is an identified broadly implemented. Mosaics of thinning levels, clearcut
priority, and consequently a more intensive timber islands, and green tree leave islands may achieve multiple
management program is implemented. Although species forest objectives, retaining localized patches of rare species
persistence is addressed by states and many rare species or species hotspots while opening other patches for
protective measures are implemented, a relatively greater regeneration of shade intolerant tree species and
risk to native habitats and species is perceived with their production of greater wood volumes. Such practices are
more intensive timber harvest practices, reduced reserved more costly, involving greater site reconnaissance and site
lands, and narrower riparian buffers. Whether or how preparation, and more complex logging directions, but
states might alter their role as ecosystem and biodiversity they also may attain goals for compatible wood production
stewards is currently a debated issue regionally and and biological resource protection at the site level. The
nationally. Private and industrial forest land owners seek role of leave islands for vertebrate species persistence
to maximize timber returns, and while they actively design within a managed forest landscape needs further study.
provisions to maintain biodiversity as legally required, Several forest management approaches and provisions
their provisions may minimally protect species habitats, are being tested, or are in need of being tested. Riparian
rarely identify all taxa associated with the forest landscape, management approaches are being examined at the site
and do not necessarily restore habitats to allow rare species level. Alternative stream buffer widths are being examined
to [re]colonize their lands. Industrial land owners with (e.g., Figure 14). Concurrently, forest biological resources
more extensive land holdings have been more proactive are being investigated. Are there habitats or taxa that
for species protections, yet on the broader spectrum, they require special consideration in various portions of the
seem to be held less accountable for species persistence stream network? If so, what are their responses to
than state and federally administered lands. Although alternative forest management designs? Stream buffers
there is acknowledgment of the different roles of land may not be the only mechanism for aquatic and riparian
owners for ecosystem, habitat or species stewardship, a resource maintenance. Those in current use do not provide
good model for a managed alignment of these diverse roles interior old-growth microclimate conditions along
has not been developed for a landscape with multiple streams, but rather mitigate for slope slippage, water
ownerships. The “Coastal Landscape Analysis and temperatures, wood inputs to streams, or perhaps
Management System” project is hoped to advance such a development of late-successional structural components
model for the Oregon Coast Range province.47 (e.g., large tree size). Patch reserves along streams have
Standards for species and particularly wildlife been proposed but not tested. Patches and buffers might
conservation are changing and we are still mid-pivot. be used together, like beads on a string, to provide
“Precautionary principles” are more often cited as intermittent riparian habitats with interior conditions and
rationale for conservative forest management decisions, narrower intervening sections designed to retain stream
and there has been a shift in the burden of proof for species temperatures, limited downed wood recruitment, and
and habitat protections: we’ve gone from needing to prove near-stream habitat.
a value needs protection before providing it, to proving it Management of downed wood recruitment is another
has adequate protection before lifting it.36 As mentioned topic in need of additional attention. As presented in this
above, there also has been a switch from addressing a few chapter, many wildlife habitat associations rely on dead
threatened and endangered wildlife species to a broader and downed wood. Yet are we maintaining and managing
spectrum of species (e.g, fungi, lichens, bryophytes, for the recruitment of sufficient large logs and snags?
mollusks) and assemblages (e.g, arthropod functional Examination of log decay classes in managed forests
groups).53, 54, 55 Increased public concern and review add reveals a paucity of hard logs, and in some locations,
complexities to processes that now seem to require full mostly just legacy large wood from high-grading harvests
consensus, whereas they were more authoritatively of half a century or more ago. Loss of coarse woody debris
controlled previously. Litigation or the threat of litigation has implications for mammals and amphibians, key taxa
has been an effective driver of these changing land in food webs, linking producers to consumers.
management ethics. As adaptive management approaches Spatial scale of protection is an issue that needs to be
are being advocated, long-term contracts for state and addressed for multiple wildlife species and habitats across
private Habitat Conservation Plans are becoming more Oregon and Washington. What resources should be
and more difficult to achieve. And finally, while the policy maintained at the site, at the watershed or land-ownership
arena is embroiled in controversy over how much wildlife block, or at the landscape and region? Is the intent to
 210 Wildlife–Habitat Relationships in Oregon and Washington

Figure 14. Alternative riparian buffer widths being examined streamside trees for bank stability, those within about 20 feet
in headwaters, with upslope density management. Tree (6 m) of the stream (D. Olson, Riparian Buffer Study,
heights correspond to federal interim Riparian Reserves for interagency research conducted by the Bureau of Land
the Northwest Forest Plan.54 The variable buffer has a 50-feet Management and the U.S. Forest Service in western Oregon
(15 m) minimum and varies with topographic or vegetation forests). K. Ronnenberg, graphics artist.
conditions, and the streamside retention buffer retains only

maintain all rare species and key habitats at all localities? 3. Integrated habitat-based and species-specific
At what level of rarity can sites be prioritized for management approaches are being designed by
maintenance at larger scales, such as a watershed or land federal, state, and private landowners.
ownership, so that losses at individual sites are acceptable 4. Collaborative efforts are being initiated to
if larger scale persistence is assured? What levels of risk investigate alternative management approaches to
are acceptable for these different species-rarities, habitats, achieve multiple resource production and
and spatial scales? Can protective mitigations be nested protection across landscapes.
among sites, watersheds, provinces, landscapes, and
ownerships? Mechanisms and processes for multiscale Literature Cited
and inter-landowner management approaches need 1. Blessing, B. J., E. P. Phoenix, L. L. C. Jones, and M. G. Raphael. 1999. Nests
advancement. These need to be tied to effective monitoring of Van Dyke’s salamander (Plethodon vandykei) from the Olympic
strategies and adaptively managed. Peninsula, Washington. Northwestern Naturalist 80: 77-81.
2. Brown, E. R., technical editor. 1985. Management of wildlife and fish
habitats in forests of western Oregon and Washington. Part 2,
Conclusion Appendices. U.S. Forest Service R6-F&WL-192-1985, Portland, OR.
Several common themes are presented for new directions 3. Burton, T. A., and G. E. Likens. 1975. Salamander populations and
in western and montane forest management. These are biomass in the Hubbard Brook Experimental Forest, New Hampshire.
sure to develop further in the next few years and decades. Copeia 1975: 541-546.
1. The burden of proof is shifting to demonstrable 4. Bury, R. B., and C. A. Pearl. 1999. Klamath-Siskiyou herpetofauna:
biogeographic patterns and conservation strategies. Natural Areas
stewardship of species and their habitats prior to
Journal 19: 341-350.
implementation of west-side forest management 5. Carey, A. B. 1995. Sciurids in Pacific Northwest managed and old-
activities, with increased use of the “precautionary growth forests. Ecological Applications 5: 648-661.
principle” and conservative approaches to hedge 6. Carey, A. B. 2000. Effects of new forest management strategies on
uncertainties and risk to species persistence. squirrel populations. Ecological Applications 10:248-257.
Increased public involvement and oversight forest 7. Carey, A. B., M. M. Hardt, S. P. Horton, and B. L. Biswell. 1991. Spring bird
land management activities is partly responsible communities in the Oregon Coast Range. Pages 123-142 in: L. F.
Ruggiero, K. B. Aubry, A. B. Carey, M. H. Huff, technical coordinators.
for this trend, resulting in greater land-owner
Wildlife and vegetation of unmanaged Douglas-fir forests. U.S. Forest
accountability for the maintenance of natural Service General Technical Report PNW-GTR-285, Portland, OR
resources. 8. Carey, A. B., and M. L. Johnson. 1995. Small mammals in managed,
2. Well-defined goals are needed for wildlife naturally young, and old-growth forests. Ecological Applications 5: 336-
management in western forests among federal and 352.
state agencies, private and industrial land 9. Carey, A. B., J. Kershner, B. Biswell, and L. D. de Toledo. 1999. Ecological
ownerships. scale and forest development: squirrels, dietary fungi, and vascular
plants in managed and unmanaged forests. Wildlife Monographs 142: 1-
71.
 CHAPTER 7: WESTSIDE/MONTANE FORESTS 211

10. Carey, A. B., B. R. Lippke, and J. Sessions. 1999. Intentional Ecosystem 25. Leonard, W. P., H. A. Brown, L. L. C. Jones, K. R. McAllister, and R. M.
Management: managing forests for biodiversity. Journal of Sustainable Storm. 1996. Amphibians of Washington and Oregon. The Trailside
Forestry 9: 83-125. Series, Seattle Audubon Society, Seattle, WA.
11. Chambers, C. L., W. C. McComb, and J. C. Tappeiner II. 1999. Breeding 26.McGarigal, K., and W. C. McComb. 1992. Streamside versus upslope
bird responses to three silvicultural treatments in the Oregon Coast breeding bird communities in the central Oregon Coast Range. Journal
Range. Ecological Applications 9(1): 171-185. of Wildlife Management 56: 10-23.
12. Chen, J., J. F. Franklin, and T. A. Spies. 1995. Growing-season 27.McGarigal, K., and W. C. McComb. 1993. Research problem analysis on
microclimatic gradients from clearcut edges into old-growth Douglas- biodiversity conservation in western Oregon forests. Special Research
fir forests. Ecological Applications 5:74-86. Report, Bureau of Land Management, The Pacific Forest and Basin
13. Cissel, J. H., F. J. Swanson, G. E. Grant, D. H. Olson, S. V. Gregory, S. L. Rangeland Systems Cooperative Research and Technology Unit,
Garman, L. R. Ashkenas, M. G. Hunter, J. N. Kertis, J. H. Mayo, M. D. Corvallis, OR.
McSwain, S. G. Swetland, K. A. Swindle, and D. O. Wallin. 1998. A 28. Molina, R., T. O’Dell, S. Dunham, and D. Pilz. 1999. Biological diversity
landscape plan based on historical fire regimes for a managed forest and ecosystem functions of forest soil fungi: management implications.
ecosystem: the Augusta Creek study. U.S. Forest Service General Pages 45-58 in: R. T. Meurisse, W. G.Ypsilantis, and C. Seybold, technical
Technical Report PNW-GTR-422, Portland, OR. editors. Proceedings: Pacific Northwest forest & rangeland soil
14. Cissel, J. H., F. J. Swanson, and P. J. Weisberg. 1999. Landscape organism symposium. U.S. Forest Service General Technical Report
management using historical fire regimes: Blue River, OR. Ecological PNW-GTR-461, Portland, OR.
Applications 9(4): 1217-1231. 29. Morgan, P., G. H. Aplet, J. B. Haufler, H. C. Humphries, M. M. Moore, and
15. Crisafulli, C. M. 2000. Survey protocol for the Larch Mountain W. D. Wilson. 1994. Historical range of variability: a useful tool for
salamander (Plethodon larselli). Chapter VII pages 253-310 in: D. H. evaluating ecosystem change. Journal of Sustainable Forestry 2: 87-111.
Olson, editor. Survey protocols for amphibians under the Survey and 30. Moritz, C. 1994. Defining “evolutionarily significant units” for
Manage provision of the Northwest Forest Plan. Interagency conservation. Trends in Ecology and Evolution 9(10): 373-375.
publication of the Regional Ecosystem Office, Portland, OR. U.S. GPO 31. Morrison, M. L. 1981. The structure of western warbler assemblages:
[2000-589-124/04022 Region No. 10] available at http:// analysis of foraging behavior and habitat selection in Oregon. Auk 98:
www.or.blm.gov/surveyandmanage/ 578-588.
16. Csuti, B., A. J. Kimerling, T. A. O’Neil, M. M. Shaughnessy, E. P. Gaines, 32. Morrison, M. L., and E. C. Meslow. 1983. Avifauna associated with early
and M. M. P. Huso. 1997. Atlas of Oregon wildlife: distribution, habitat, growth vegetation on clearcuts in the Oregon Coast Ranges. U.S.
and natural history. Oregon State University Press, Corvallis, OR.. Forest Service General Technical Report PNW-GTR-305, Portland, OR.
17. DellaSalla, D. A., S. B. Reid, T. J. Frest, J. R. Strittholt, and D. M. Olson. 33. Morrison, P., and F. J. Swanson. 1990. Fire history and pattern in a
1999. A global perspective on the biodiversity of the Klamath-Siskiyou Cascade Range landscape. U.S. Forest Service General Technical Report
ecoregion. Natural Areas Journal 19: 300-319. PNW-GTR-254, Portland, OR.
18. Garman, L. R., and M. G. Hunter. 1998. Phase 4-Evaluation: Birds and 34. Nauman, R. S., and D. H. Olson. 2000. Survey and Manage salamander
mammals. Pp. 59-61 in: J. H. Cissel, F. J. Swanson, G. E. Grant, D. H. Olson, known sites. Chapter II in: D. H. Olson, editor. Survey protocols for
S. V. Gregory, S. L. Garman, L. R. Ashkenas, M. G. Hunter, J. N. Kertis, J. H. amphibians under the Survey and Manage provision of the Northwest
Mayo, M. D. McSwain, S. G. Swetland, K. A. Swindle, and D. O. Wallin. A Forest Plan. Interagency publication of the Regional Ecosystem Office,
landscape plan based on historical fire regimes for a managed forest Portland, OR. U.S. GPO [2000-589-124/04022 Region No. 10] available
ecosystem: the Augusta Creek study. U.S. Forest Service General at http://www.or.blm.gov/surveyandmanage/
Technical Report PNW-GTR-422, Portland, OR. 35. Niemiec, S. S., G. R. Ahrens, S. Willits, and D. E. Hibbs. 1995.
19. Gilbert, F. F., and R. Allwine. 1991. Spring bird communities in the Hardwoods of the Pacific Northwest. Forest Research Laboratory,
Oregon Cascade Range. Pages 145-158 in: L. F. Ruggiero, K. B. Aubry, A. Oregon State University, Research Contribution 8. Corvallis, OR.
B. Carey, and M. H. Huff, technical coordinators. Wildlife and vegetation 36. Noss, R. F., M. A. O’Connell, and D. D. Murphy. 1997. The science of
of unmanaged Douglas-fir forests. U.S..Forest Service General Technical conservation planning: habitat conservation under the Endangered
Report PNW-GTR-285, Portland, OR. Species Act. Island Press, Washington, D.C.
20. Hagar, J. C., W. C. McComb, and W. H. Emmingham. 1996. Bird 37. Nussbaum, R. A., E. D. Brodie, Jr., and R. M. Storm. 1983. Amphibians
communities in commercially thinned and unthinned Douglas-fir stands and reptiles of the Pacific Northwest. University of Idaho Press,
of western Oregon. Wildlife Society Bulletin 24(2): 353-366. Moscow, ID.
21. Huff, M. H., and C. M. Raley. 1991. Regional patterns of diurnal 38. Ollivier, L. M., and H. H. Welsh, Jr. 2000. Survey protocol for the Del
breeding bird communities in Oregon and Washington. Pages 177-205 Norte salamander (Plethodon elongatus). Chapter V pages 163-200 in: D.
in: L. F. Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff, technical H. Olson, editor. Survey protocols for amphibians under the Survey and
coordinators. Wildlife and vegetation of unmanaged Douglas-fir forests. Manage provision of the Northwest Forest Plan. Interagency
U.S. Forest Service General Technical Report PNW-GTR-285, Portland, publication of the Regional Ecosystem Office, Portland, OR. U.S. GPO
OR. [2000-589-124/04022 Region No. 10] available at http://
22. Johnson, R. E., and K. M. Cassidy. 1997. Terrestrial mammals of www.or.blm.gov/surveyandmanage/
Washington state: location data and predicted distributions. Volume 3 39. Olson, D. H. and W. P. Leonard. 1997. Amphibian inventory and
in: K. M. Cassidy, C. E. Grue, M. R. Smith, and K. M. Dvornich, editors. monitoring: a standardized approach for the Pacific Northwest.
Washington State Gap Analysis Project-final report, Washington Chapter 1 in:D. H. Olson, W. P. Leonard, and R. B. Bury, editors. Sampling
cooperative fish and wildlife research unit, University of Washington, amphibians in lentic habitats: methods and approaches for the Pacific
Seattle, WA. Northwest. Northwest Fauna 4:1-22.
23. Landres, P. B., P. Morgan, and F. J. Swanson. 1999. Overview of the use 40. Omernik, J. M. 1987. Ecoregions of the conterminous United States.
of natural variability concepts in managing ecological systems. Map Supplement (Scale 1:7,500,000). Annals of the Association of
Ecological Applications 9(4): 1179-1188. American Geographers 77: 118-125.
24. Lehmkuhl, J. F., L. F. Ruggiero, and P. A. Hall. 1991. Landscape-scale 41. Oregon Biodiversity Project. 1998. Oregon’s living landscape:
patterns of forest fragmentation and wildlife richness and abundance in strategies and opportunities to conserve biodiversity. A Defenders of
the southern Washington Cascade Range. Pages. 425-442 in: L. F. Wildlife Publication, Oregon State University Press, Corvallis, OR.
Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff, technical coordinators. 42. Pater, D. E., S. A. Bryce, T. D. Thorson, J. Kagan, C. Chappell, J. M.
Wildlife and vegetation of unmanaged Douglas-fir forests. U.S. Forest Omernik, S. H. Azevedo, and A. J. Woods. 1998. Ecoregions of western
Service General Technical Report PNW-GTR-285, Portland, OR. Oregon and Washington. Interagency poster produced by U.S.
Geological Survey, Reston, VA.
 212 Wildlife–Habitat Relationships in Oregon and Washington

43. Ralph, C. J., P. W. C. Paton, and C. A. Taylor. 1991. Habitat association 56. U.S. Department of Interior. 1992. Recovery plan for the northern
patterns of breeding birds and small mammals in Douglas-fir/hardwood spotted owl—final draft. U.S. Department of the Interior, 2 volumes.
stands in northwestern California and southwestern Oregon. Pages Portland, OR.
379-393 in: L. F. Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff, 57. Vega, R. M. S. 1993. Bird communities in managed conifer stands in the
technical coordinators. Wildlife and vegetation of unmanaged Douglas- Oregon Cascades: habitat associations and nest predation. M.S. thesis.
fir forests. U.S. Forest Service General Technical Report PNW-GTR- Oregon State University, Corvallis, OR.
285, Portland, OR. 58. Weisberg, P. J. 1996. Blue River fire regime analysis and description.
44. Ruggiero L. F., L. L. C. Jones, K. B. Aubry. 1991. Plant and animal habitat Unpublished report, available at the Blue River Ranger District Office,
associations in Douglas-fir forests of the Pacific Northwest: an U.S. Forest Service, Blue River, OR.
overview. Pages 447-462 in: L. F. Ruggiero, K. B. Aubry, A. B. Carey, and M. 59. Welsh, H. H., Jr. 1990. Relictual amphibians and old-growth forests.
H. Huff, technical coordinators. Wildlife and vegetation of unmanaged Conservation Biology 4: 309-319.
Douglas-fir forests. U.S. Forest Service General Technical Report PNW-
GTR-285, Portland, OR.
45. Sedell, J. R., G. H. Reeves, and K. M. Burnett. 1994. Development and
evaluation of Aquatic Conservation Strategies. Journal of Forestry
92(4): 28-31.
46. Smith, M. R., P. W. Mattocks, Jr., and K. M. Cassidy. 1997. Breeding birds
of Washington state.Volume 4 in: K. M. Cassidy, C. E. Grue, M. R. Smith,
and K. M. Dvornich, editors. Washington State Gap Analysis Project-final
report, Seattle Audubon Society Publications in Zoology No. 1, Seattle,
WA.
47. Spies, T. A., G. H. Reeves, K. M. Burnett, W. C. McComb, K. N. Johnson,
G. Grant, J. L. Ohmann, S. L. Garman, and P. Bettinger. In press. Assessing
the ecological consequences of forest policies in a multi-ownership
province in Oregon. In: J. Liu and W. W. Taylor, editors. Integrating
landscape ecology into natural resources management. Cambridge
University Press, NY.
48. Storm, R. M., and W. P. Leonard, coordinating editors. 1995. Reptiles of
Washington and Oregon. The Trailside Series, Seattle Audubon Society,
Seattle,WA.
49. Swanson, F. J., J. A. Jones, D. O. Wallin, and J. H. Cissel. 1994. Natural
variability - implications for ecosystem management. Pp. 89-106 in: M. E.
Jensen, and P. S. Bourgeron, technical editors. Eastside forest ecosystem
health assessment - Volume II: ecosystem management: principles and
applications. U. S. Forest Service General Technical Report PNW-GTR-
318, Portland, OR.
50. Teensma, P. D. A. 1987. Fire history and fire regimes of the central
western Cascades of Oregon. Ph.D. Dissertation, University of Oregon,
Eugene, OR.
51. Thomas, J. W., M. G. Raphael, R. G. Anthony, E. D. Forsman, A. G.
Gunderson, R. S. Holthausen, B. G. Marcot, G. H. Reeves, J. R. Sedell, and
D. M. Solis. 1993. Viability assessments and management considerations
for species associated with late-successional and old-growth forests of
the Pacific Northwest. Report of the Scientific Analysis Team. U. S.
Forest Service, Portland, OR.
52. Trappe, J. M., and D. L. Luoma. 1992. The ties that bind: fungi in
ecosystems. Chapter 2 in: G. C. Carroll, and D. T. Wicklow, editors. The
fungal community: its organization and role in the ecosystem, Second
edition. Marcel Dekker, Inc., NY.
53. U.S. Department of Agriculture, Forest Service, U. S. Department of
Commerce (NOAA), U.S. Department of Interior (Bureau of Land
Management, National Park Service and U.S. Fish and Wildlife Service)
and the Environmental Protection Agency. 1993. Forest ecosystem
management: an ecological, economic, and social assessment. Report of
the Forest Ecosystem Management Assessment Team [FEMAT]. U. S.
GPO 1993-793-071. Available at: Regional Ecosystem Office, Portland,
OR.
54. U.S. Department of Agriculture, Forest Service, and U.S. Department
of Interior, Bureau of Land Management. 1994. Record of decision for
amendments for Forest Service and Bureau of Land Management
planning documents within the range of the northern spotted owl
(Northwest Forest Plan). U.S. D.A. Forest Service and U.S. D.I. Bureau
of Land Management, Portland, OR.
55. U.S. Department of Agriculture, Forest Service, and U.S. Department
of Interior, Bureau of Land Management. 1999. Draft-Supplemental
environmental impact statement: for amendment to the survey and
manage, protection buffer, and other mitigating measures standards and
guidelines. [Draft SEIS]. U.S. Forest Service and U.S. Bureau of Land
Management, Portland, OR.
 8
Wildlife of Eastside (Interior)
Forests and Woodlands
Rex Sallabanks, Bruce G. Marcot, Robert A. Riggs,
Carolyn A. Mehl, & Edward B.Arnett

Introduction
Eastside (interior) forests and woodlands are those forest Habitat Characterizations
and woodland environments that occur in Oregon and Eastside (interior) forests and woodlands are characterized
Washington east of the crest of the Cascade Mountain by highly variable climate, landscape topography,
Range. Eastside (interior) forests and woodlands are elevational gradients, and varying degrees of natural and
broadly classified into 4 forest habitats based on conifer anthropogenic disturbances. Mountain ranges, rolling
composition: (1) Eastside (Interior) Mixed Conifer Forest hills, and deep canyons characterize much of eastern
(EMCF), (2) Lodgepole Pine (Pinus contorta) Forest and Oregon and Washington where Eastside (interior) forests
Woodlands (LPFW), (3) Ponderosa Pine (Pinus ponderosa) and woodlands dominate the landscape. Elevation ranges
Forest and Woodlands (PPFW), and (4) Upland Aspen from 100 feet (30 m) above sea level (asl) (e.g., PPFW in
(Populus tremuloides) Forest (UAF). We specifically do not the Columbia River Gorge) to 9,500 feet (2,896 m) asl (e.g.,
address forested riparian habitats (see Kauffman et al.87). UAF).38 As a result of this broad elevational gradient, and
In this chapter, we first highlight the principal the extensive geographic distribution of Eastside (interior)
characteristics of Eastside (interior) forests and woodlands, forests and woodlands, physical site conditions are highly
describe key habitat features that make them suitable for variable. In general, however, mature vegetation
wildlife, provide an overview of the wildlife assemblages conditions of Eastside (interior) forests and woodlands
that inhabit them, and discuss aspects of biological are controlled over broad regions by climate, with soils
diversity and wildlife residency. With an emphasis on and topography playing secondary roles; that is, most
ponderosa pine, we then focus on management issues mature vegetation is zonal in nature. Variation in
relevant to Eastside (interior) forests and woodlands, disturbance regimes also has influenced species
consider historical disturbance regimes, threats, and composition and density in forests of the inland West.42
trends, and then relate these issues to wildlife. Finally, we Historically, the presence and structure of forests
provide several case histories that illustrate specific dominated by Douglas-fir (Pseudotsuga menziesii) and
management issues and/or wildlife-habitat relationships grand fir (Abies grandis) throughout much of the inland
and provide some suggestions for future research and West were influenced by frequent, low-intensity ground
monitoring. fires that reduced densities of trees and surface vegetation.
A broad discussion of all Eastside (interior) forests and For low elevation cover types (primarily PPFW), this
woodlands and their associated wildlife throughout disturbance regime produced open, park-like stands of
Oregon and Washington is beyond the scope of both this all ages but predominantly large ponderosa pine, with a
chapter and our collective working knowledge. grass-dominated understory. These forests were regularly
Throughout this chapter, therefore, our comments are interrupted by nonforest openings (e.g., sclerophyllous
often based on our own research experiences and reflect shrubs, shrubsteppe, grassy parks, or balds, and
certain biases. Most notably, these biases are nongame herbaceous- or broadleafed-dominated riparian areas)57,
landbirds and the Blue Mountains of northeastern Oregon. 153
(Figure 1). As a result, low elevation forests of the inland
Where appropriate, however, we focus on other West, including Eastside (interior) forests and woodlands,
ecologically significant habitats (e.g., ponderosa pine) or were historically fragmented, at least with respect to forest
wildlife taxa (e.g., large herbivores). For more specific cover.138
information on UAF, see DeByle and Winokur48a and Eastside (Interior) Mixed Conifer Forest currently is
DeByle et al.48b In summary, our intent is to describe key dominated by grand fir, Douglas-fir, western larch (Larix
aspects of Eastside (interior) forests and woodlands, occidentalis), and ponderosa pine. These forests occur on
highlight significant management issues, and by way of the periphery of the maritime climatic region that supports
specific examples, illustrate wildlife-habitat relationships cedar-hemlock forests.41, 146 Mixed conifer forest is more
and potential interactions with land-use practices. diverse along eastern slopes of the Cascade Mountains
than elsewhere in the interior Columbia Basin. For

213
 214 Wildlife–Habitat Relationships in Oregon and Washington

example, in the very southern Cascades and northern elevation lodgepole pine forests for feeding and nesting.
Sierras and Klamath mountain convergence areas within This woodpecker selects for smaller-diameter snags over
Oregon, white fir (A. concolor), incense cedar (Calocedrus larger-diameter ones,130 but most other Eastside cavity
decurrens), Jeffrey pine (Pinus jeffreyi), and sugar pine (P. nesters select for the largest available snag (e.g., pygmy
lambertiana) also can be found in association with Douglas- nuthatch and white-headed woodpecker). The pinyon jay
fir and ponderosa pine.42, 118 However, these species do not is well known to cache viable seeds of pinyon and other
typically occur in most EMCF. Extensive dry forests of pine species, thereby acting as a pine dispersal agent.
PPFW occur at elevations below the mixed conifer forest
zone. Lodgepole Pine Forest and Woodlands occur Distributional Aspects of Eastside
throughout the mixed conifer forest zone, primarily in Environments
cold, relatively dry environments and usually where Within Oregon and Washington, EMCF occurs primarily
winter snowpack is persistent. Pumice soil lodgepole pine throughout the east Cascades, Blue Mountains (northeast
habitat (see case history 4) is typically intermixed with Oregon), and Okanogan Highlands (northeast
PPFW and lies in between EMCF and either western Washington). Lodgepole Pine Forest and Woodlands have
juniper (Juniperus occidentalis) woodland or shrub-steppe a similar geographic distribution to EMCF;38 in particular,
habitat.38 Finally, UAF, found from 2,000 to 9,500 feet (610 this forest type is common in the pumice zone of
to 2,896 m) can occur as seral stands in both EMCF and southcentral Oregon from near Mt. Jefferson south to the
PPFW or as climax forest on well-drained mountain slopes Crater Lake area (also see case history 4).
or canyon walls that have some moisture. Ponderosa Pine Forest and Woodlands are also
Some wildlife species are more or less closely associated widespread throughout foothills of the Blue Mountains,
with each of these forest types in Washington and Oregon. east Cascades, Okanogan Highlands, and central Oregon
Examples of species largely confined to each of these types pumice zone, as well as in the Columbia Basin in
(i.e., that also occur within 3 or fewer other habitat types, northeastern Washington. Upland Aspen Forest is
according to the Wildlife-habitat Relationship [WHR] primarily found in the Steens Mountains (southeast
matrixes [see CD-ROM]) include (in increasing order of Oregon) and in the northeastern Cascades of Washington.
number of other habitat types also used): Cordilleran Mixed conifer, lodgepole pine, and ponderosa pine
flycatcher, pygmy shrew, red-tailed chipmunk, boreal owl, habitats are widespread and relatively common, often
and gray catbird in EMCF; pygmy nuthatch, pygmy shrew, occurring in large contiguous tracts throughout the Blue
pinyon jay, and gray flycatcher in PPFW; boreal owl and Mountains, eastern slopes of the Cascades, and Okanogan
gray flycatcher in LPFW; and yellow warbler, American Highlands. In contrast, UAF is relatively uncommon and
redstart, and boreal owl in UAF. None of these species is highly patchy in its distribution across the Eastside
confined to only one habitat type. (interior) forest landscape. At least currently, where it
Other wildlife species found within the Eastside occurs as a seral stage within other forest types, aspen
(interior) forest and woodland array have evolved stands are small and isolated from one another (Figure 2).
associations with specific substrates or environmental Whereas many wildlife species that inhabit Eastside
conditions. Examples include the northern three-toed (interior) forests and woodlands might be considered
woodpecker, which invades recently burned-over, high- “associates” of specific forest communities (see above),

Figure 1. Eastside Mixed

Conifer Forest in the Blue
Mountains. Note the
landscape is naturally
fragmented with forested
north-facing slopes (left) and
nonforested south-facing
slopes (right).
 CHAPTER 8: EASTSIDE FORESTS 215

Snags. Dead standing trees or snags are widely recognized

to be an essential forest component for numerous wildlife
species.130 In the 4 Eastside (interior) forest and woodland
types, 77 species of vertebrates associate with snag
substrates during some part of the annual cycle, including
2 amphibians, 51 birds, and 24 mammals, which use
specific snag characteristics for diverse purposes. Primary
cavity excavators include 15 species (2 chickadees, 3
nuthatches, 9 woodpeckers, and black bear). A few species
use bark piles at the base of snags, including the
northwestern salamander, which also uses down logs, and
the Larch Mountain salamander, which uses bark piles
particularly during dry seasons and on sites with moist
talus and low soil content. 105 Partially decomposed
standing wood provides nest, feeding, and roost sites for
many other species; decomposing down wood is also used
by various other species.
Most species of bats occurring in Eastside (interior)
forests and woodlands use snags and large trees with
structural defects (e.g., broken tops) for roosting.17, 35, 104
Bats generally select tall, large-diameter snags for
roosting.18 However, the influence of forest structure and
landscape pattern surrounding roost sites is not well
understood for most species in this region. Snags and trees
selected for roosting by bats occur in more open habitats
(e.g., less canopy closure and understory vegetation) or
near edges, and are close to water. 17, 18, 35 Anecdotal
information on local declines of snag-associated bats may
be related to reduction in the number of large snags.105
Species experiencing habitat losses over the historical
period were closely associated with ponderosa pine
habitats (especially single-story old-forest stands).
Figure 2. Patches of Upland Aspen Forest, often intermixed
with other Eastside forest and woodland habitats, are Down Wood. Sixty-eight species of wildlife found in
currently small and isolated. Eastside (interior) forests and woodlands have correlations
with down wood, including 6 amphibians, 5 reptiles, 13
birds, and 44 mammals. A few amphibian species, such
they are not often considered “obligates.” Thus, we do as the Larch Mountain salamander, may lay eggs in moist
not generally consider the distributional aspects of any of habitats but are associated with down wood or talus as
the Eastside (interior) forest and woodland habitats to be adults, rather than aquatic environments. The distribution
limiting for wildlife. Possible exceptions to this might exist and abundance of many amphibian species are more
for species that inhabit upland aspen and old-growth closely associated with specific substrates (such as down
ponderosa pine, because of the patchy distribution of these wood) and microhabitat conditions (such as deep, moist
habitats across the landscape. Species associated with talus), than with general vegetation cover types and
aspen habitat include the red-naped sapsucker46 and structural stages.105 Collectively, these amphibians are most
several species of bats, including the big brown bat.44, 86 sensitive to changes in down wood, litter, and duff depths
Species known to be associated with old-growth and characteristics, still and flowing water quality and
ponderosa pine include the white-headed woodpecker,60 quantity, and precipitation quality and weather patterns.
white-breasted nuthatch,80a and flammulated owl.74 Bull et al.32 provide details of use of down wood by
numerous other wildlife species in the interior Columbia
Associations Between River Basin. Some species, such as the pileated
Habitat Elements and Wildlife woodpecker in the Blue Mountains of northeastern
Here we emphasize wildlife associations with 4 habitat Oregon, frequently forage on down wood for carpenter
features (elements) that seem to be especially significant ants30 (Camponotus spp.) (Figure 3).
in Eastside (interior) forests and woodlands. The Habitat
Closed-canopy Forest. Some 233 wildlife species found
Element matrixes (see CD-ROM) can be used to list habitat
in Eastside (interior) forests and woodlands are generally
elements, including substrates and microhabitat and
associated with closed-canopy forest structures, including
conditions, that are used by species associated with each
single- and multi-story structures of shrub/seedling,
of the 4 Eastside (interior) forests and woodlands.
sapling/pole, small-tree, medium-tree, and large-tree
 216 Wildlife–Habitat Relationships in Oregon and Washington

Figure 3. Down wood is an important

habitat element for many wildlife species in
Eastside forests and woodlands. The cavities
seen in this photograph were likely excavated
by the pileated woodpecker in search of
carpenter ants.

forest stages. No wildlife species within this set has an advantages of these associations are largely unstudied.
obligate association with closed-canopy conditions of Most Eastside (interior) forests did not commonly occur
shrub/seedling, sapling/pole, small-tree, medium-tree or historically as closed-canopy structures (with high
large-tree forest structures. Such associates use 10 other volumes of dead wood), which developed only in recent
structural conditions (see CD-ROM matrixes). Examples decades with the advent of fire suppression management
include the least flycatcher, willow flycatcher, dusky- (see below), whereas early successional grass/forb and
footed woodrat, fox sparrow, green-tailed towhee, and shrub/seedling stages did occur naturally following
yellow-pine chipmunk. Most of these species occur more intermittent natural fires.
frequently in sapling/pole and small-tree stages. Species
frequenting medium- and large-tree stages include only Wildlife Assemblages
the least flycatcher and dusky-footed woodrat. The In this section we provide an overview of all vertebrate
western red-backed vole and western wood-pewee use wildlife taxa that inhabit Eastside (interior) forests and
closed-canopy conditions of medium- or large-tree woodlands and discuss species residency (i.e., migratory
structures and also a dozen other structural conditions. status). This section is intended to be general, although in
In EMCF of northeastern Oregon, Townsend’s warbler is some cases little information is available on distribution,
highly associated with closed-canopy conditions, but not abundance, and habitat relationships for wildlife species,
exclusively so (see case history 1). All of the species listed like reptiles. Some species, such as birds, are relatively
here constitute the closest set of closed-canopy associates well studied.
in medium- and large-tree stages of Eastside (interior)
forests and woodlands. Overview of Wildlife
Grass/forb and Shrub/seedling. In contrast to closed- According to the WHR data matrices, there are 287
canopy forest, many wildlife species are closely associated vertebrate wildlife species that currently inhabit Eastside
with the closed-canopy conditions of grass/forb and (interior) forests and woodlands (8 salamanders, 10 frogs/
shrub/seedling stages of Eastside (interior) forests and toads, 2 turtles, 6 lizards, 13 snakes, 98 mammals, and 150
woodlands. Species nearly tied (that is, that use only 1 birds). This species richness is roughly equivalent to
other structure) to such closed grass/forb structures Eastside shrubland and grassland habitats (285 species),
include the killdeer, meadow vole, American pika, coastal and marine environments (278 species), and urban
California ground squirrel, northern bog lemming, and environments (265 species), subtantially greater than
Brazilian free-tailed bat. Species closely tied (that is, that alpine and subalpine habitats (173 species), and less than
use only 3 other structures; none uses less) to closed shrub/ freshwater, riparian, and wetland habitats (429 species),
sapling structures include the rough-legged hawk, barn agriculture and pastures (344), and westside and high
swallow, and Brewer’s sparrow. A lack of shrub cover in montane forest habitats (320). Three species occurring
EMCF of Oregon may actually influence the abundance historically in Eastside (interior) forests and woodlands
of some shrub-nesting songbirds, many (83%) of which are now locally extirpated: northern leopard frog, bison,
are Neotropical migrants34 (R. Sallabanks, 1995 Annual and sharp-tailed grouse (recently reintroduced). Much of
Report, Sustainable Ecosystems Institute, Meridian, ID). the following information has been summarized from
One can speculate on why more wildlife species seem previous publications.105, 159
to be more closely associated with closed structures of Amphibians. Amphibians of Eastside (interior) forests and
grass/forb and shrub/seedling structures than with older woodlands are largely unstudied. For many species, even
forest structures; evolutionary conditions and adaptive those listed as sensitive, such as the tailed frog in Oregon,28
 CHAPTER 8: EASTSIDE FORESTS 217

data on populations, distribution, and habitat populations. Whereas some species seem to be negatively
requirements are scarce to nonexistent. In general, the 4 impacted by excessive grazing (e.g., Cascade frog and
Eastside (interior) forest and woodland habitats were Columbia spotted frog), others seem tolerant (e.g., Pacific
considered to play a supportive (but not essential) role in chorus frog). Other land-use practices also may affect
species’ maintenance and viability (i.e., most amphibian amphibians. Irrigation canals or field flooding can provide
species were classified as being generally “associated” adequate habitat for egg laying and larval development,
with each habitat) (Figure 4a). but if water is shut off prior to hatching or metamorphosis,
Within Eastside (interior) forests and woodlands, species will fail to reproduce.
standing or flowing water is required for egg laying and
Reptiles. Distribution and abundance of reptile species
larval development for most amphibian species. Frogs are
have been poorly studied. Habitat selection for snakes and
predominantly associated with forested habitats; riparian
lizards is driven more by the need for warm climates,
areas provide essential habitat for most adult forms. At
rocks, talus, and soils than by the presence of general
lower elevations, in dryer forest types characterized by
vegetation types. Most reptile species occur in the drier
ponderosa pine and Douglas-fir, several amphibian
PPFW (21 species) at lower elevations (Table 1) than in
species occur, most of which are locally endemic and more
the other 3 Eastside (interior) forest and woodland
common west of the Cascade Range. Ponderosa Pine
habitats. Moister forest habitats at intermediate elevations
Forest and Woodlands and EMCF provide habitat for 13
also have many reptile species, but to a lesser extent
and 12 amphibian species, respectively (Table 1). Within
compared to PPFW. At higher elevations, in cold forest
EMCF, the moister cedar-hemlock and grand fir habitat
habitats, reptiles are relatively rare.
types support the richest amphibian communities because
Like amphibians, most reptile species are considered
of the damp climate and greater abundance of aquatic
generally associated with Eastside (interior) forest and
habitats.
woodland habitats; none are considered “closely
At higher elevations, in colder forest habitat types such
associated” (dependent on a habitat or structural condition
as Engelmann spruce (Picea engelmannii), lodgepole pine,
for part or all of its life history requirements) and few are
and subalpine fir (A. lasiocarpa), temperatures are generally
considered “present” (known to occasionally use a habitat
too cold and the breeding season too short to support
or structural condition) (Figure 4b).
diverse amphibian communities. Lodgepole Pine Forest
and Woodlands and UAF support only 9 and 4 amphibian Birds. Eastside (interior) forests and woodlands provide
species, respectively (Table 1). habitat for many bird species, most of which are known
In some cases, exotic fish species introduced into high to occur in PPFW and EMCF (Table 1). Riparian vegetation
elevation lakes have impacted indigenous amphibian within Eastside (interior) forests and woodlands is of
species, such as the Cascades frog. Elsewhere, exotic particular importance to many species, especially
amphibian species (e.g., the bullfrog) may be directly Neotropical migratory songbirds.136 The distribution and
responsible for the decline or elimination of native abundance of many birds seems to be more heavily
amphibian species (e.g., the northern leopard frog99). Other influenced by the presence of key structural attributes
species, such as the southern race of the Columbia spotted (habitat elements) within forested stands rather than
frog and the western toad have declined or become locally habitat types or seral stages per se (see case histories 1
extirpated for unknown reasons.105 Little is documented and 4). In general, the 4 Eastside (interior) forest and
about the effects of livestock grazing on amphibian woodland habitats were considered to play a supportive

Table 1. Numbers of vertebrate wildlife species known to occur

in Eastside forest and woodland habitats.*

Taxonomic class Mixed conifer Lodgepole pine Ponderosa pine Upland aspen

Amphibians 12 9 13 4
Reptiles 11 12 21 5
Birds 116 83 131 77
Mammals
Small mammals 43 26 31 24
Bats 11 9 15 5
Carnivores 18 13 14 10
Ungulates 9 8 7 5
All mammals 81 56 67 44
All species 220 160 232 130
* Information derived from a query of Wildlife-Habitat Relationships Project data matrixes.
Species with historical or unknown occurrences are excluded.
 218 Wildlife–Habitat Relationships in Oregon and Washington

A—Amphibians D—Mammals

Forest habitat
Forest habitat

closely associated
closely associated
generally associated
generally associated
present
present

0 2 4 6 8 10 12 14 0 10 20 30 40 50 60 70 80 90
Number of amphibian species Number of mammal species

Figure 4. Degree of association of amphibian (A), reptile (B),

B—Reptiles bird (C), and mammal (D) species among the 4 Eastside forest
and woodland habitats (top: Mixed Conifer; next: Lodgepole
Pine; next: Ponderosa Pine; bottom: Upland Aspen). Closely
associated—a species is widely known to depend on a habitat
or structural condition for part or all of its life history
requirements; generally associated—a species exhibits a high
Forest habitat

degree of adaptability and may be supported by a number of

habitats or structural conditions; present—a species
demonstrates occasional use of a habitat or structural
condition.

(but not essential) role in species’ maintenance and

closely associated
viability (Figure 4c).
generally associated
Most bird species are wide-ranging and use several, if
present
not all, of the 4 Eastside (interior) forest and woodland
habitats. Relatively few species were classified as being
0 4 8 12 16 20 24
closely associated with any of the forest habitats,
Number of reptile species
suggesting that few species depend on any 1 specific forest
habitat for maintenance and viability (Figure 4c).
C—Birds Examples of such “closely associated” species include
cordilleran flycatcher and EMCF, boreal owl and LPFW,
northern saw-whet owl and PPFW, and red-naped
sapsucker and UAF.
Most birds using Eastside (interior) forest and
woodlands are “foliage-gleaners” (Figure 5a). Such species
Forest habitat

(e.g., warblers and chickadees) forage primarily by

gleaning insects or fruit from vegetation (foliage and
occasionally branches), not from the ground. Many
foliage-gleaning bird species occur in PPFW and EMCF.
Other foraging guilds (swooping divers, ground-gleaners,
aerial insectivores, and bark-gleaners) are relatively evenly
closely associated distributed among Eastside (interior) forest and woodland
generally associated habitats in terms of species numbers (Figure 5a). Nesting
present guilds are dominated primarily by open-cup (e.g.,
thrushes and warblers) and cavity (e.g., woodpeckers and
0 20 40 60 80 100 120 140 owls) nesters; platform (e.g., accipiters) and ground (e.g.,
Number of bird species grouse and quail) nesters are less common (Figure 5b).
Many open-cup nesting bird species occur in PPFW and
EMCF. Cavity nesters seem to be distributed similarly
 CHAPTER 8: EASTSIDE FORESTS 219

among the 4 Eastside (interior) forest and woodland

habitats (Figure 5b).
Small Mammals. Few data exist relative to distribution
and abundance of small mammal species in Eastside
(interior) forests and woodlands. Several are exotic to the
region, including Virginia opossum, which was recorded
in Umatilla County, northeast Oregon, as recently as
1994.161 There are more small mammal species in EMCF
than the other 3 habitats (Table 1). Most use a wide variety
of forest types, however, and like many species in this
assessment, use key habitat elements such as burrows,
litter, down wood, rock outcrops, and forest openings in
all seral stages. Many squirrels, mice, woodrats, and other
species depend on seeds from trees, especially large
ponderosa pine seeds. Some species, especially ground
squirrels, have benefited from agricultural intensification
of dryer, low-elevation forest communities. Others, such
as the northern pocket gopher and common porcupine
have benefited from forestry practices.105
More mammal species exhibit close associations with
Eastside (interior) forest and woodland habitats (especially
EMCF) than any other taxonomic group considered in this
assessment (Figure 4d). Examples include bushy-tailed
woodrat and EMCF, snowshoe hare and LPFW, western
pocket gopher and PPFW, and least chipmunk and UAF.
Many species are considered generally associated with
Eastside (interior) forests and woodlands; relatively few
are considered present (Figure 4d).
Bats. Of all vertebrate wildlife groups, we perhaps know
least about bats. Bats occupy a unique and important niche
in Eastside (interior) forests and woodlands, as they are
the only mammal with powered flight capability and the
only nocturnal predator of many forest insects. Bats spend
more than half their lives subjected to the selective
pressure of roosting, and availability and quality of roost
sites are thought to be critical factors influencing
population size and distribution of some species. 95
Proximity to drinking and foraging sites also influences
bat use of habitat and may limit populations.35, 39, 45, 95, 165 In Figure 5. Distribution of bird species, summarized by
general, forest-dwelling bats use large-diameter snags and foraging guild (A) and nesting guild (B), among the 4
trees as roosts, and frequently switch roosts, which may Eastside forest and woodland habitats.
be related to the type and permanency of roost occupied,
and inversely related to the availability of roosts.100 High
levels of bat activity in mature and old-growth forests123, to occur (Table 1). Important management issues relative
151, 152
are thought to be partially a consequence of to bats include timber harvest, particularly removal of
availability of older trees for roosting in these stands.35, 39, snags, and disturbance at mine and cave sites. Campbell
95, 152
However, western long-eared myotis are flexible in et al.35 suggested that retention and recruitment of snags
selection of roost sites, and their patterns of use of roost in managed forests are important for the conservation of
structures may differ with landscape characteristics165 (e.g., bats in this region. Selective harvest techniques that retain
habitat pattern, snag availability). Bats also use various trees and snags distributed across the landscape could
human-made structures such as bridges, buildings, and provide habitat to maintain populations of bats.35, 163
mines,95 thus human development may favor some species Restricting human entry to caves and mines where bats
of bats by providing roost sites, hibernacula, and are known to occur may reduce potentially adverse effects
nurseries.52, 104 from recreation.
Information on basic ecology, abundance, distribution,
Carnivores. Eastside (Interior) Mixed Conifer Forest
and habitat selection of bats in Eastside (interior) forests
provides habitat for 18 species of carnivores; among the 3
and woodlands is generally rare.17, 35, 122 Bats reach their
other Eastside (interior) forest and woodland habitats,
richest communities in PPFW where 15 species are known
carnivores are relatively evenly distributed (Table 1). Many
 220 Wildlife–Habitat Relationships in Oregon and Washington

of the carnivores that inhabit Eastside (interior) forests and Like elk and deer, bighorn sheep are popular for
woodlands are habitat generalists, dependent more on hunting and viewing, and at higher elevations within
availability of prey and lack of disturbance from humans EMCF, frequently use cliff and rock walls. Whereas some
(e.g., black bear). Others, such as the American marten populations of bighorns seem to be maintaining current
and Pacific fisher, are closely associated with late- numbers, others are generally declining, perhaps due to
successional conifer forests and riparian habitats or with widespread habitat changes164 such as replacement of
interior forest environments;61 these 2 mustelids also use grass, forbs, and low shrubs with tall shrubs and trees,
down logs for den sites.105 However, marten also use which bighorns generally avoid.128 Fire suppression has
intensively managed lodgepole pine forests in the central contributed significantly to these changes. Bighorn sheep
Oregon Pumice Zone126 (see case history 4). benefit from management activities that maintain early-
The distribution and abundance of some carnivores seral vegetative conditions, such as prescribed fire9, 78, 121a,
have changed dramatically from historical conditions.117 127a
and timber harvest.10
Species such as the grizzly bear, gray wolf, and mountain
lion were (and still are) considered a threat to human safety Aspects of Wildlife Residency
and livestock production. As a result, they were pursued Among wildlife taxa considered in this assessment, only
by trappers from the mid-1800s through the mid-1900s. bird species migrate entirely out of the region seasonally.
Grizzly bear and gray wolf populations were extirpated In fact, there are almost as many Neotropical migratory
from Eastside (interior) forests and woodlands within birds in Eastside (interior) forests and woodlands as there
Oregon and significantly reduced in Washington. are year-round resident birds (Figure 6). Most Neotropical
Although likely reduced during the trapping era, migrants spend the winter in Central and South America,
mountain lion populations have increased toward the end returning to Oregon and Washington in April and May to
of the century subsequent to changes in harvest breed. Fall migration to wintering grounds typically occurs
regulations. Other carnivorous species such as coyotes, in August and September. Numbers of migratory bird
foxes, and skunks have apparently increased because of species vary among habitat communities in relation to
reduced competition and removal of their major overall numbers of bird species; no particular habitat
predators105 as well as the encroachment of agricultural seems to have disproportionately more or less (Figure 6).
lands at lower elevations. The status, trends, and general habitat relationships of
Neotropical migratory birds in the interior Columbia River
Ungulates. Ungulates are significant components of Basin, which includes Eastside (interior) forests and
Eastside (interior) forests because of their profound woodlands, have been reviewed elsewhere.136
influence on vegetation (see case history 3). Ungulate In general, however, wildlife species that inhabit
species are similarly distributed among EMCF, LPFW, and Eastside (interior) forests and woodlands are primarily
PPFW. Upland Aspen Forest has the fewest species of year-round residents. Of the 287 species that currently
ungulates (Table 1). Vegetation communities at mid- inhabit these environments, 227 (79%) remain in Oregon
elevations, especially open plant communities in
ponderosa pine and Douglas-fir zones, and mixed conifer
communities at higher elevations, support the largest
numbers of herbivores.82 These forest zones dominate the
Blue Mountains landscape, particularly at mid-elevations
where animals must gravitate to meet nutritional
demands. Hence it is the strategic position of these types
and their broad distribution that dictate their importance
to ungulates. Highly productive habitats (e.g., UAF) can
be extraordinarily important to large herbivores, despite
their limited distribution because of their high
productivity.
Deer and elk have expanded their ranges in recent
times, and where densities have exceeded the capacity of
remaining wildlands, they have become depredators of
crops in some agricultural areas, particularly on private
lands near the interface of valley and montane zones. In
some forest settings, elk and deer use dense stands of
shade-tolerant understory trees for cover, which would
not have been as available under natural fire regimes.
Within UAF, aspen and its associated understory plants
are heavily used forage sources, sometimes throughout
the year. Many aspen communities have become degraded Figure 6. Distribution of bird species, summarized by
as a result of persistent heavy foraging by seasonal migratory status, among the 4 Eastside forest and woodland
concentrations of high ungulate numbers (both wild and habitats.
domestic).
 CHAPTER 8: EASTSIDE FORESTS 221

and Washington year-round (although not necessarily in that operates in drier, low- to mid-elevation Eastside
Eastside [interior] forests and woodlands). There is some (interior) forests has been a primary impact of modern
seasonal movement within eastern Oregon and land-management activities in the region. This low to mid-
Washington by several ungulate species, but there is no elevation range corresponds primarily with the historical
migration out of the region. Some bat species are thought occurrence of ponderosa pine forest2 and woodlands in
to migrate southward during the winter months (e.g., eastern Washington.97 Fire exclusion policies, coupled with
hoary bat and spotted bat), but this has yet to be confirmed. and accelerated by selective harvest of the seral tree
Many bird species that are considered permanent (i.e., component and livestock grazing, have allowed many
nonmigratory) residents exhibit “migrations” between PPFWs to progress farther along successional pathways
breeding and nonbreeding locations within Eastside than would have occurred under historical fire regimes.3,
43, 97
(interior) forests and woodlands. Such species can be At the same time, a return to a more natural
summarized using the classification system derived by disturbance regime would result in the loss of some old-
Marcot.103 Elevational migrants include (1) species that forest characteristics (e.g., closed-canopy, dead wood) that
breed locally, but move to lower elevations in winter (e.g., have developed from fire exclusion.
Cooper’s hawk); (2) species that breed at higher elevations, Whereas management issues exist in many or all
but move into local areas in winter (e.g., ruby-crowned Eastside (interior) forests and woodlands (i.e., clearcutting,
kinglet and Lincoln’s sparrow); and (3) species that breed roads), the primary anthropogenic impacts of the last 100-
at low elevations/bottomlands, but move to local areas 150 years have resulted in extensive changes in
post-breeding (e.g., American goldfinch, Brewer ’s distribution, structure, and species composition of
blackbird, and brown-headed cowbird). Latitudinal ponderosa pine forests,2 and to a lesser extent (because of
migrants include “displacement” species that move limited historical range in eastern Washington)
latitudinally within the region (e.g., sharp-shinned hawk, woodlands.97 The corresponding impacts to associated
band-tailed pigeon, and winter wren). Finally, there are wildlife are poorly understood and opportunities for
several nomadic permanent residents that exhibit less future research are limited because of the extensive loss
predictable movements within Eastside (interior) forests of these habitats in eastern Washington.
and woodlands throughout the year (e.g., cedar waxwing Efforts to restore and maintain biodiversity within
and red crossbill). targeted landscapes often concentrate on the most
significantly altered environment relative to historical
Ecological Functions of Wildlife Species in distribution and conditions.114 For this reason, we confine
Eastside (Interior) Forests and Woodlands most of our discussion of management impacts within
The CD-ROM matrixes were queried to determine the key Eastside (interior) forests and woodlands to PPFW.
ecological functions (KEFs) performed by wildlife
associated with Eastside (interior) forests and woodlands. Natural Disturbance Regimes
All 4 Eastside (interior) forest and woodland habitats Historically, landscape vegetation patterns in the Pacific
contained the same categories of KEFs.107 A high degree of Northwest and in many Eastside (interior) forest and
functional similarity was found in patterns of species woodland habitat types were primarily influenced by fire
redundancy (richness) by KEF category, with a few at return intervals ranging from 7-20 years (ponderosa pine
interesting exceptions. There were fewer foliovores and forests), 50-100 years (Douglas-fir and grand fir forests),
browsers in PPFW, and EMCF (more in UAF and LPFW) and >100 years (subalpine fir forests).2, 3 In dry, low- to
than expected by overall patterns of species richness mid-elevation PPFW, the frequency of fire kept fuel
among the 4 forest types. There were fewer grazers, loading of the forest floor and understory to a minimum
fungivores, and bark feeders in PPFW than expected. which, in turn, prevented fire from reaching the overstory
Ponderosa Pine Forest and Woodlands were also and killing trees.2, 97, 172 The result was an overall reduction
depauperate in species that disperse fungi and lichens. in fire intensity, dependent on the fire return interval. One
This was also true for species that fragment down wood response of vegetation was to favor more fire-tolerant tree
and impound water. This might be related to the greater species (e.g., ponderosa pine and/or white oak [Quercus
aridity of PPFW than the other 3 forest habitats. However, garryana] in the overstory and bunchgrasses in the
overall trophic structures (percent of all species in primary, understory) that were well adapted to a frequent low-
secondary, and tertiary consumer categories) were nearly severity fire regime.5, 13, 172
identical among all 4 Eastside (interior) forest and Whereas many of these fires were ignited naturally,
woodland habitats. This seems to be an ecosystem Native Americans used Eastside (interior) forests and
constant. woodlands, especially PPFW, for food and building
materials for thousands of years prior to Euro-American
Management Issues in Eastside settlement14 and their most significant influence on these
forests was their propensity to intentionally start fires.
(Interior) Forests and Woodlands Possible reasons for setting fires were to improve desired
Wildfire is a driving force in shaping the structure and wildlife habitat, increase desired plant species, drive game
function of all Eastside (interior) forests and woodlands.2, animals into traps, and open transportation routes.67
3
In the last century, alteration of the natural fire regime Historically then, these fires generally burned extensively
 222 Wildlife–Habitat Relationships in Oregon and Washington

Figure 7. Fire exclusion policies throughout

Eastside forests and woodlands during the past
100 years have created densely stocked stands of
trees. Note the high fuel load on the forest floor,
which in combination with the crowded
overstory, would increase the likelihood of a
stand-replacing wildfire in this forest.

throughout low- to mid-elevation Eastside (interior) intervals (say, >50 years) and high-severity fire regimes
forests and woodlands, perhaps only being extinguished occurring within the dry, low-elevation ponderosa pine
by fall rains or lack of fuel due to previous fires. Under zone.12, 54, 145, 170, 172 On similar ponderosa pine sites in central
such a natural disturbance regime, the potential for high- Idaho, Steele144 found these high-severity fires to have
severity wildfires,2 insect outbreaks, and disease events dysfunctional effects on the ecosystem through reduced
were low.168 Volume of dead wood was probably low as loss of site production, seed sources, nutrients, and even
well.127b, pages 308-9 soil.
Fire Suppression. Along with logging and grazing, fire Timber Harvest. Effects of timber harvest on Eastside
suppression has been responsible for significant changes (interior) forests and woodlands have changed over the
in forest structure and species composition of Eastside years.3, 96 Early timber harvests usually targeted the largest
(interior) forests and woodlands; this is especially true for trees, which primarily included ponderosa pine and
PPFW.2, 13 Such management action has lengthened the fire western larch. More recently, species such as Douglas-fir,
return interval and has significantly altered the species grand fir, lodgepole pine, and subalpine fir have also been
composition and structure, compared to forests existing used.3 Historical harvest regimes, coupled with fire
under a low-severity fire regime.3 Today, many PPFWs suppression, allowed smaller, more shade tolerant, late
no longer could support a low-severity fire regime; many successional species to capture the growing space of
of these forests have not burned since the 1800s and are ponderosa pine forests.140 The result has been a rapid shift
characterized by considerable fuel loading in the on many sites from forests dominated by seral species to
understory.3, 172 those dominated by late successional species, and from
Fire exclusion policies during the past 100 years have relatively open stands of old-growth trees to relatively
interrupted the developmental pattern of clumped groups closed stands of young trees which have high fuel and
of trees by allowing regeneration to survive across an large woody debris loads.3, 172
entire site and not just in openings.5 As a result, extensive More recently, timber management programs have
areas of dry, low- to mid-elevation Eastside (interior) forest used more intensive harvest practices. Clearcut areas in
landscapes, including EMCF and LPFW, have become low-elevation forests generally recover slowly from
increasingly homogeneous, with “dog-hair thickets” of logging disturbance, and efforts to reforest clearcuts have
trees in many areas (Figure 7).3 Most, if not all, low- to been generally unsuccessful.146 Today, selective harvest
mid-elevation potential vegetation types are believed to that favors retention of early seral species, while also
be experiencing tree densities outside the historical range thinning the stand and reducing fuels, is considered the
of conditions for these sites.3, 144, 172 Dense overstory and most ecologically responsible harvest method in
understory conditions provide vertical continuity of fuels ponderosa pine systems, especially for those sites where
that effectively creates a “ladder” for fire to spread from under-burning is considered a viable management
the understory to tree crowns. As a result, fires now kill practice over the long term.144
more trees through loss of overstory in addition to Eastside white oak woodlands have relatively little
understory, as well as being more severe in terms of economic value as a timber species and are more
temperature and site impacts. Consequently, Eastside commonly harvested for firewood purposes.97
(interior) forest landscapes are now characterized by
Grazing. Greater livestock densities at lower elevations
infrequent fire return intervals and large expanses of high-
have led to greater grazing impacts in PPFW relative to
severity, stand-replacing fires.3 Research on historical fire
the other Eastside (interior) forest and woodland habitats.3
return intervals throughout the Intermountain West has
Grazing began in the late 1800s when enormous herds of
uncovered little to no evidence of infrequent fire return
 CHAPTER 8: EASTSIDE FORESTS 223

Figure 8. Livestock grazing, in addition to fire

suppression and timber harvest, is a
management activity that has played a major
role in changing historical fire regimes of
Eastside forests and woodlands.

domestic sheep and cattle were allowed to graze freely moisture that results in reduced ponderosa pine
throughout PPFW. By 1860, in addition to feral horses and regeneration and increased mortality over the long-term.144
sheep, there were 200,000 cattle in Oregon.59 Steele et al.146 Little forest remains today that represents historical old-
observed and documented substantial damage to soils and growth (>200 years68) ponderosa pine forest.74, 114 Where
vegetation on ponderosa pine sites in Idaho, especially stands still contain old-growth ponderosa pine, tree
where herds were concentrated. Perennial bunchgrasses, densities and fuel accumulations present a significant risk
in particular, are still recovering from severe overgrazing to long-term survival and future restoration of this forest
of the early days.3, 96, 146 type.144 Lightening and accidental fire starts are expected
Grazing pressure at this level can also contribute to fire to cause future burns of unprecedented and uncontrollable
suppression by reducing continuity of understory intensity and magnitude.4
vegetation and preventing low intensity fires from Allowing these forests to burn under a high-severity
spreading in their normal pattern across the landscape.43 fire regime to “reset the balance” is considered by some
Today, livestock grazing continues throughout all Eastside to be a nonviable alternative for restoration of these
(interior) forests and woodlands, but at much lower forests.144 Such fires could potentially eliminate what little
densities (Figure 8). Localized damage to vegetation and old-growth remains, as well as have detrimental effects
soils may still occur where animals concentrate, on wildlife.96 Replacing these old forests would take >200
particularly in riparian areas and forest openings.64 On years, thereby foregoing any immediate restoration
the other hand, grazing by wild herbivores increased options and flexibility for spatial distribution. To
dramatically during the twentieth century.82 complicate matters, the occurrence of high-severity fires
Grazing continues to affect forest ecology in terms of on these sites often damages the soil or promotes species
succession, species composition, and species diversity.127c better adapted to a “hot” fire regime, making these
Specific influences on forest structure have been increased landscapes more prone to future high-severity fires.3 The
tree numbers, decreased native grasses, increased result can be delayed recolonization by all species where
accumulation of downed woody material, increased soil is damaged, or recolonization by shrubs that are able
spread of exotic and noxious weeds, and increased forest to shade out seral tree species and delay their
floor duff.69, 174 However, changes in the densities of wild establishment for many years to come (R. Steele, forestry
and domestic herbivores, relative to one another, suggest consultant, pers. comm.). The end result is the same,
that the focus of future research on grazing should extend however: the extended loss of the old-growth ponderosa
beyond merely the effects of livestock grazing.127c pine ecosystem from the landscape.
Kay88 concludes that, historically, Native Americans
Status, Threats, and Trends of commonly determined the structure of entire plant and
Habitats and Communities animal communities by hunting and by setting fires (also
Collectively, fire suppression, grazing, and logging have see papers cited in Knowles and Knowles 93), and that a
contributed to the loss of most fire-maintained old-growth current “natural regulation” approach to management
Eastside (interior) forests and woodlands. Ponderosa Pine does not recognize, and thus probably would not replicate,
Forest and Woodlands have been more heavily impacted such historical conditions.106 Perhaps with careful thinning,
by these factors relative to the other three Eastside along with prudent reintroduction of fire into these
(interior) forest and woodland habitats. Today, the two ecosystems, old-growth ponderosa pine forests can be
greatest immediate threats to future viability of ponderosa extensively restored. Given the dynamic nature of
pine forests are high-severity fire occurrences and ecological communities in Eastside (interior) forests and
increased site-specific competition for nutrients and woodlands, particularly regarding potential effects of fire,
 224 Wildlife–Habitat Relationships in Oregon and Washington

perhaps the very concept of defining “desired future mountain and western bluebirds, are associated with sites
conditions” for planning could be replaced with a concept where fires have killed the forest overstory (also see case
of describing “desired future dynamics.” That is, long- history 2). Black-backed and three-toed woodpeckers also
term evolutionary potentials can be met only by associate with mid- to high-elevation forests following
accounting for potential future changes in conditions. No irruptions of insect populations such as bark beetles.22
condition can be static for long. Impending changes in Hutto80b documented differential responses by bird
regional climates, too, have the capacity for causing great species to stand-replacement fires in the northern Rocky
shifts in composition of ecological communities. Mountains, and noted that one species, the black-backed
woodpecker, was nearly restricted to sites with standing
Relationships Among Management Issues, fire-killed trees. He also listed other bird species that may
Wildlife, and Disturbance Regimes benefit over the long term from stand-replacement fires.
Fire suppression has resulted in conifer encroachment on (For more details on relationships between severity of
certain shrub habitats and has contributed to loss of habitat wildfire and bird communities, see case history 2).
for many reptiles.105 Little information is available on the Other western studies, although not conducted in
effects of fire and fire suppression, season of burn, and Oregon or Washington, provide data on the response of
changes in vegetation pattern resulting from burns and bird communities to snag harvest following fire (i.e.,
fire suppression on birds in Eastside (interior) forests and salvage logging). Raphael125 explored the response of birds
woodlands. 156a Although general trends have been to reduced densities of snags, simulating various snag
reported, species-specific effects on bird population harvest levels immediately following fire using simulation
density and trends from livestock grazing, changes in models. He predicted that optimum snag densities under
distribution and density of nest parasites (principally the the constraints tested should be between 17-37 snags/acre
brown-headed cowbird), changes in insect populations, (7-15 snags/ha). Also using models, Raphael and White127
and availability of free water are poorly understood. predicted drastic declines in cavity-nesting bird density
Livestock grazing is well known to indirectly influence following removal of snags from burned areas in the Sierra
some nongame bird populations via its effects on Nevada.
vegetation.148, 149 In ponderosa pine forests of southwestern Idaho, Saab
Resource management, including fire suppression and and Dudley135 found Lewis’ woodpecker to be the most
timber management, and resulting changes in the abundant and successful cavity nester in 2- to 4-year-old
structure and distribution of vegetation communities, has burns. At least during the first 4 years post-fire, nesting
also influenced the distribution and abundance of many densities of cavity-nesting birds continued to increase.
avian species. For example, the amount of old-growth Lewis’ woodpeckers seemed to be positively affected by
ponderosa pine forest that has been maintained by salvage logging, experiencing highest nesting success in
frequent, low-severity fires has declined by approximately salvage-logged units. All bird species selected nest sites
85% from historical conditions to present across the entire with higher tree densities than those measured at random
region; in some areas, such as the Klamath Plateau and sites, and cavity nesters as a group selected clumps of
eastern slopes of the Cascade Mountains, <5% remains snags as opposed to snags retained in uniform, evenly
today.74 Species associated with this community, such as spaced distributions. Retaining snags in clumps during
the white-headed woodpecker and flammulated owl, salvage logging operations is therefore a management
likely have declined in abundance and are considered recommendation. Another is to maintain snags (especially
sensitive by federal and state agencies. Lewis’, black- broken-topped snags) in forests that are susceptible to
backed, and three-toed woodpeckers, along with

Figure 9. Salvage logging of fire-killed trees is

a common management practice in Eastside
forests and woodlands today. More studies are
needed on the effects of this type of forestry on
wildlife species.
 CHAPTER 8: EASTSIDE FORESTS 225

stand-replacement wildfires to provide nest trees during

post-fire years when most trees cannot yet be excavated. Case History 1
Hutto80b stated that post-fire salvage cutting (Figure 9) Bird-habitat Relationships in Grand Fir
might be conducted too frequently to be justified on the Forests of the Blue Mountains, Oregon
basis of sound ecosystem management because some bird Habitat selection by birds has been well studied, both
species require burned forests for maintenance of viable empirically40, 84, 169 and theoretically.58, 131, 160 Since Thomas,153
populations. He pointed out that some bird species differ however, few additional data have been added to our
in the microhabitats that they occupy within a burn; knowledge of forestry-avifauna relationships in the
salvage prescriptions that tend to “homogenize” forest interior Pacific Northwest. Prior to the study reported here,
structure (e.g., selective removal of all trees of a certain Mannan and Meslow102 and Bull et al.33 provided the only
size) are therefore unlikely to maintain the necessary data on songbirds in the Blue Mountains. This case history
variety of microhabitats within a burned forest. Hutto80b summarizes a three-year study of bird populations in
suggested that rather than implementing a selective tree grand fir forests of the Blue Mountains. The objectives of
removal salvage operation, it may be better to take trees the study were to (1) assess how forest stand attributes,
from one part of a burn and leave another part completely such as canopy cover, snag density, and understory
untouched, dependent on management objectives. structure influence habitat selection by birds; (2)
Unfortunately, the paucity of empirical data on effects of investigate bird use of forest structural classes in Eastside
snag harvest on wildlife populations following fire makes mixed conifer (grand fir) forests; and (3) relate habitat use
it difficult to suggest viable management alternatives. by birds to forest management options and the potential
for timber harvest practices to provide suitable breeding
Research Activities in Eastside habitat for birds. We present data on avian community
(Interior) Forests and Woodlands composition, relationships with specific structural
attributes of forest stands, and on how birds were
To provide practical examples of relationships between
distributed among forest structural classes (i.e.,
wildlife species and management issues in Eastside
developmental stages). Other potential drivers of habitat
(interior) forests and woodlands, we present the following
use by birds, such as landscape features and abiotic factors,
four case histories. These case histories offer details on
were controlled for wherever possible. To enhance the
specific wildlife species, habitat relationships, and effects
relevance of this study to current management, we
of management activities. With one exception (case history
describe avian abundance with respect to a forest
3), we emphasize birds and/or the Blue Mountains to
classification scheme formulated for the Interior Columbia
illustrate wildlife use of Eastside (interior) forests and
Basin Ecosystem Management Project (ICBEMP).124
woodlands. Based on our review of literature and current
This study was conducted in the Marine Physiographic
state of knowledge of wildlife communities in Eastside
Zone 36 of the northern Blue Mountains Ecological
(interior) forests and woodlands, we have identified
Province57 in northeastern Oregon. Study sites were
several information needs. We conclude with suggestions
located in forests managed by Boise Cascade Corporation
for future research.
(n = 59), and the U.S. Forest Service, Umatilla and
Wallowa-Whitman National Forests (NF) (n = 24). All
sample stands were in the Abies grandis series of forest
habitat types.85 Sample stands ranged from 40-124 acres
(16-50 ha). By limiting the study to grand fir forests,

Figure 10. Stem exclusion, open canopy forest is

a common structural class in Eastside Mixed
Conifer Forests of the Blue Mountains. Our
research has found this structural class, which is
often management-induced (note stumps in the
foreground), to have the richest avifauna of all
grand fir forest types in the region.
 226 Wildlife–Habitat Relationships in Oregon and Washington

variation among study plots with respect to abiotic factors Several of the relationships observed are consistent with
such as climate, soil conditions, and potential previous studies. The relative abundance of Townsend’s
evapotranspiration was minimized. Grand fir forests are warbler, for example, was consistently correlated with
part of EMCF. canopy cover across years, and canopy cover explained
Data were collected in 83 forest stands between 1994 almost 50% of the variation in Townsend’s warbler
and 1996. Data were collected annually in each stand and numbers among structural classes when data were pooled
vegetation data were collected in the first year that a among years. Mannan and Meslow102 also reported that
particular stand was sampled (either 1994 or 1995). The Townsend’s warblers nested in sites that had high canopy
same 45 stands were sampled in all three years and the volumes of grand fir and Douglas-fir, and that canopy
same 70 stands were sampled in both 1995 and 1996. All volume of grand fir was the primary attribute separating
stands were classified into 1 of 7 structural classes nest sites from random sites.
according to structure115: (1) stand initiation (SI); (2) stem In general, however, research of this nature has not
exclusion, open canopy (SEOC) (Figure 10); (3) stem occurred in Eastside (interior) forest and woodlands and
exclusion, closed canopy (SECC); (4) understory most of the habitat relationships we report have not been
reinitiation (UR); (5) young forest, multistory (YFMS); (6) previously described. Only 4 species showed sufficiently
old forest, single-story (OFSS); and (7) old forest, consistent relationships with structural attributes to
multistory (OFMS). This scheme has been adopted by warrant development of habitat functions to predict
ICBEMP, and it essentially represents an expansion of the relative abundance. Such functions may be used to predict
4 “stand development stages” first described by Oliver.116 the abundance of a species if we know the values of the
Bird communities were censused using standard point different parameters in that species’ habitat model. Thus,
count methodology.81 Vegetation was sampled using the potential for a particular forest stand to provide
protocols modified from James and Shugart.83 For more suitable habitat for a species can be evaluated for different
details see R. Sallabanks (1996 Annual Report, Sustainable management scenarios. Overstory removal, for example,
Ecosystems Institute, Meridian, ID). can reduce numbers of Townsend’s warblers at the stand
In total, 36,602 detections of 90 bird species were level (note, however, that any stand-level effects should
recorded. Rank order of abundance changed only slightly not be considered without respect to landscape-level
from year to year. Overall, the red-breasted nuthatch, modification and resulting dynamics). Similarly, the
Townsend’s warbler, and dark-eyed junco dominated the orange-crowned warbler is a ground-nesting songbird that
pooled sample. Among the 30 species for which data were was more abundant in stands with well-developed
analyzed, 29 exhibited significant correlations between understories (>1,236 shrubs, seedlings, and saplings/acre
their abundance and one or more structural attributes, in [>500/ha]) and substantial ground cover (>60% ground
at least 1 year of study. Only abundance of the pine siskin cover). Nest predators presumably have more difficulty
was apparently unrelated to the structural attributes locating nests in such stands, where nests built on the
measured. Adjusted R2 ranged from 0.07 (hermit thrush ground are well hidden beneath shrubs, grasses, forbs,
in 1995) to 0.63 (chipping sparrow in 1994) with an overall and conifer seedlings. Thus, where bird densities are
mean of 0.24. Four species (American robin, chipping strongly linked to shrub density and/or ground cover, it
sparrow, orange-crowned warbler, and Townsend’s is possible that prescribed burning and/or grazing by
warbler) showed significant correlation with the same large ungulates (i.e., wild or domestic) may be expected
structural attribute(s) in all 3 years; habitat functions were to influence nesting or foraging habitat, and thereby
subsequently developed for these species by pooling influence bird density or reproductive success. Overstory
census data among years. manipulation is relatively unimportant in this context. On
Abundance of 23 bird species (64%) varied significantly the other hand, understory suppression may improve
among forest structural classes. All species except ruffed habitat for the American robin, a species found to be more
grouse, brown creeper, Swainson’s thrush, Townsend’s common in stands with little understory (<988 shrub
warbler, house wren, and mountain bluebird were present stems, seedlings, and saplings/acre [<400/ha]). This
in all structural classes. The house wren and mountain ground-foraging thrush requires flat, open forest floors
bluebird were found almost exclusively in SI, and in which to search for invertebrates.139 Hence, what is
Hammond’s flycatcher was far more abundant in SECC potentially deleterious for one species may be beneficial
than in other structures. The following species were most for another (see also Sallabanks et al.137). The chipping
abundant in, but not necessarily restricted to, particular sparrow would apparently benefit from silviculture that
single structural classes: dusky flycatcher, orange-crowned produces open forest canopy (10-30%) and removal of
warbler, chipping sparrow, northern flicker, house wren, larger trees (>89 feet [>27 m] in height and 12.6 inches
dark-eyed junco, and mountain bluebird (SI); western [>32 cm] diameter at breast height [dbh]). This species was
tanager (SEOC); Hammond’s flycatcher, yellow-rumped consistently found to be more common in stands with
warbler, pine siskin, and golden-crowned kinglet (SECC); short trees (89 feet [27 m]) and open canopies (<40%). In
hermit thrush (UR); ruffed grouse, Swainson’s thrush, and fact, for the chipping sparrow, we found that canopy
brown-headed cowbird (YFMS); and brown creeper height explained the greatest amount of variation (65%)
(OFMS). in bird abundance.
 CHAPTER 8: EASTSIDE FORESTS 227

On average, SEOC had the richest avifauna; YFMS was species. Instead, to provide suitable habitat for most
also relatively species rich. Those forest types at the species, a rather broad range of the structural continuum
extremes of the structural class spectrum (SI and OFMS) may be required, as long as the key habitat elements (e.g.,
tended to have the most depauperate bird assemblages. snags) and structural attributes (canopy cover, understory)
These results are the first to document relative bird are provided at different levels throughout that range.
abundance among these structural classes; they can also
be used to assess the degree to which different bird species
are “associates” of any one structural class (i.e., found more Case History 2
or less exclusively in only one class). The implication for Response of Breeding Bird Communities to
management of such structural class associates is that Severity of Wildfire in Subalpine Fir Forests of
landscape planners may use decision rules for projecting the Blue Mountains, Oregon
bird abundance and thereby evaluate the potential effects Fire is the dominant disturbance agent in the interior
of proposed forestry practices on particular bird species. Pacific Northwest2 and especially in the Blue Mountains
Despite the fact that fewer bird species were found in SI of northeastern Oregon.3 Hence, a better understanding
than elsewhere, this class had more structural class of how fire influences ecosystem structure and function
associates than any other class studied. Two ground is necessary if we are to improve management and
nesters (dark-eyed junco and orange-crowned warbler), conservation of natural systems in this region. The Blue
2 low-mid canopy nesters (chipping sparrow and dusky Mountains province is a fire-dominated ecosystem3 where
flycatcher), and 3 cavity nesters (house wren, northern forest health issues such as effects of stand-replacement
flicker, and mountain bluebird) were all significantly more wildfire on wildlife populations are much needed areas
common in this structural class than elsewhere. Two of study. In particular, management of forested ecosystems
species (house wren, mountain bluebird) were almost throughout the interior Pacific Northwest would greatly
exclusively observed in SI. Clearly, the loss of this benefit from knowledge of how changes in forest habitat
particular structural class from the Blue Mountains’ following wildfire influence the wildlife species inhabiting
landscape, at least in the Abies grandis zone, would leave them, especially for those species perceived to be at risk
species like the house wren and mountain bluebird with or in decline. For example, the decline of some Neotropical
few alternatives. The only other clear structural class migratory bird species, such as the olive-sided flycatcher,
specialist was the Hammond’s flycatcher, in SECC. is perhaps linked to a century of fire suppression in
Only one species (brown creeper) was found to be western forests.6
significantly more common in OFMS, but not to the extent Early studies of the effects of fire on bird populations
to be considered an old forest associate. The brown creeper focused primarily on game species such as quail and
has been found to be associated with structures grouse.16, 50, 132 More recently, research has focused on snag
characteristic of old forest by others.1, 108 Other species in use by cavity-nesting birds in forested systems48, 135 and
the Blue Mountains have also been referred to as “old general effects of fire on songbird habitat.79, 101 In their
forest specialists,” primarily because of their apparent examination of post-fire succession of avifauna in the
dependency on large snags (e.g., pileated woodpecker30, Olympic Mountains of western Washington, Huff et al.79
31, 102
and Vaux’s swift27, 29, 102). This was not so in our study, found bird communities to be similar in forest 1 year post-
where the presence of pileated woodpecker did not differ fire compared with nearby old-growth (preburn) forest.
significantly among structural classes. Avian breeding density and diversity did decline,
The fact that so few species exhibited significant however, in burned forest in years 2 and 3 post-fire.
“selection” for structural class suggests that the structural Studying avian community structure during early post-
attributes and habitat elements required by most species fire succession in the Sierra Nevada, Bock et al.25 found
are present in several structural classes rather than being species richness, diversity, and evenness all to be highest
restricted to just one. Stand Initiation possibly contained on a burned plot 8 years post-fire, but to be lowest on the
the most structural class associates because it differed the same plot 15 years post-fire. Blake21 reported reduced
most from the other structural classes with respect to key species richness of nonbreeding bird communities in
habitat elements and structural attributes that influence burned compared with unburned ponderosa pine habitat
the distribution and abundance of birds, such as canopy in Arizona. Other studies, in other habitat types, also found
cover. Unless a species requires an entirely open canopy bird populations declining following fire.98, 154
(e.g., house wren) or large decadent trees with flaking bark In general, however, most studies report few, if any,
for nesting cover (e.g., brown creeper), any number of the differences between the avifaunal composition of recently
intermediate structural classes from SEOC to YFMS burned and unburned forests.16, 51, 101, 147, 150 A few studies
provide suitable habitat. Those structural classes that report increases in avifaunal richness following fire.23, 24
represent the extremes of the successional spectrum (SI In a survey of breeding bird communities from 34 burned
and OFMS) are likely to be the most different structurally, forest sites in western Montana and northern Wyoming,
have the most specialized avifauna, and the fewest Hutto 80b discovered that within 2 years after stand-
generalists. When attempting to maintain avian replacement fire, the relative abundance of woodpeckers,
biodiversity, therefore, structural classes per se may not flycatchers, and seed-eaters (a total of 15 species) was
be relevant and are not likely to be limiting for most significantly higher, presumably due to increased foraging
 228 Wildlife–Habitat Relationships in Oregon and Washington

and nesting opportunities. Given the influence of fire and

current forest health issues, the Blue Mountains provide
us with an ideal setting in which to study this topic.
In 1994, the Twin Lakes Fire burned approximately
22,000 acres (8,900 ha) of high elevation EMCF in the Blue
Mountains. This case history reports briefly on the
influence of the Twin Lakes Fire on structure and
composition of breeding bird communities during the first
3 years (1995-97) following the fire (for more details on
study design and sampling protocols, see R. Sallabanks,
1995 Annual Report, Sustainable Ecosystems Institute,
Meridian, ID).
To address the effects of wildfire on bird communities,
4 treatments were selected: (1) unburned forest (i.e.,
control); (2) low severity burn forest (defined as 0-33%
tree mortality with few differences in ground and/or
understory vegetation compared with unburned forest);
(3) medium severity burn forest (defined as 34-66% tree
mortality with approximately 50% of live ground and/or
understory vegetation absent); and (4) high severity burn
forest (defined as 67-100% tree mortality with little or no
live ground and/or understory vegetation present) (Figure
11). Five replicate plots of each of the no burn, low severity
burn, and medium severity burn forest, and 10 replicate
plots of the high severity burn forest were located in Spring
1995 and 25 permanent plots were established. “Plots” can
best be described as forest patches, which may incorporate
multiple stands that were homogeneous with respect to
severity of burn. All plots were at least 40 acres (16 ha).
Most plots (n = 20) were located in the Pine Ranger District,
southeast of the Eagle Cap Wilderness Area, of the
Figure 11. High intensity wildfire can cause 100% mortality
Wallowa-Whitman NF. The remaining plots (n = 5) were
in overstory and understory vegetation, such as seen in this
in the Hells Canyon National Recreation Area, and 2 of
photograph taken 1 year post-fire. Many Eastside forests and
these 5 were within the boundary of the Wilderness Area. woodlands are now characterized by large, infrequent, stand-
Each year from 1995 through 1997, breeding bird replacing fire regimes.
communities were sampled using standard fixed-radius
point count censusing techniques. In 1995 and 1997, habitat bird species. We concur with Hutto and emphasize the
characteristics were measured using a relatively intensive need for species-specific relationships to be assessed in
vegetation sampling protocol. These data helped facilitate all studies of this nature.
an evaluation of the effects of wildfire on forest habitat Species-specific analyses were conducted on all birds
structure and a better understanding of the causal that were detected 10 times (within 164 feet [50 m] of count
mechanisms underlying changes in the bird community. stations) in 2 of the 3 years of this study (n = 23 species).
Due to space limitations, however, this case history focuses For analysis we used repeated measures ANOVA, with
on the response of avifauna only; changes in habitat “treatment” (degree of burn) as the independent variable,
features and their relationships with the distribution and “year” (1995, 1996, 1997) and “visit” (1, 2, 3) number as
abundance of bird species will not be discussed. the repeated measures, and “relative abundance” (mean
During the 3-year study, 8,399 bird detections no. birds detected per point count per visit per species) as
representing 74 bird species were recorded. The pine the dependent variable. Fourteen of the 23 species
siskin, dark-eyed junco, and chipping sparrow dominated analyzed were found to vary significantly with severity
the avian community in the recently burned Blue of burn. Most species, including those with the most
Mountains forest landscape. Overall, wildfire had a significant effects, such as the golden-crowned kinglet and
somewhat negative effect, reducing bird species richness mountain chickadee, were negatively affected. “Tree
wherever burn intensity was the highest. Hutto80b argues foliage-searching” species such as these also were found
that many previous studies of wildfire effects may not have to be more common in unburned compared with burned
found a response of bird communities to fire because plots by Bock and Lynch24 in conifer forests in the Sierra
analyses were based on composite statistics such as total Nevada. Lyon and Marzluff101 found similar results for
bird density, species richness, and within-guild this guild, reporting that foliage-insect feeders, in
abundances; these measures may “hide more than they particular, declined to only one-third of the numbers
reveal” in terms of the biological effects of fire on specific prefire. In mixed conifer forests of the Grand Teton and
 CHAPTER 8: EASTSIDE FORESTS 229

Yellowstone national parks, foliage-feeding bird species

also were more abundant in unburned and moderately Case History 3
burned compared with heavily burned sites.147 Modification of Forest Vegetation by Wild and
Although significant year effects were found for 12 Domestic Ruminant Herbivores in Mixed
species, significant interaction effects between treatment Conifer Forests of the Blue Mountains,
and year were only found for 3 species, suggesting that
Oregon
relationships between severity of burn and species’
Ruminant herbivores usually manipulate the composition
abundance were generally consistent among years. Cavity
of vegetation to some degree, especially when their density
nesting birds in general, but especially the hairy
is high relative to food supply.8, 37, 94, 129, 155 Ruminants also
woodpecker, mountain bluebird, and black-backed
may regulate community processes by modifying cycling
woodpecker, responded positively to the fire. Woodpecker
and availability of nutrients.55, 110, 120 Their dung and urine
species such as the black-backed and three-toed in
contribute directly to the pool of nitrogen that is
particular rapidly colonize recently burned forests.20, 71, 80b,
101, 147, 173
immediately available for plant growth and increase
The increased availability of snags and open
nitrogen to carbon ratio. 56, 111, 120, 134 This accelerates
foraging habitat are identified as potential causal
decomposition of litter by microbes and mineralization
mechanisms for these species.
of nitrogen,120, 133, 142 thereby accelerating nutrient cycling
Two species of management concern,53 the black-backed
and increasing production of some plants. Ruminants also
woodpecker and olive-sided flycatcher, warrant closer
regulate community processes by foraging selectively.
attention. Both species are associated with recently burned
Selective feeding directly alters the composition of plant
forest.80b The relative abundance of the black-backed
communities, influencing the amount and quality of plant
woodpecker increased significantly in the most severely
litter that becomes available to decomposers over time.120
burned forest of the Blue Mountains in 1996 compared
Through nutrient processing and selective suppression
with 1995, but numbers were back down again in 1997 (R.
of plants, ruminants influence forest attributes such as
Sallabanks, unpubl. data). The olive-sided flycatcher, on
wood production,62, 63, 120, 166 herbivore carrying capacity,82,
the other hand, continued to increase in abundance 109
and biodiversity.26, 49, 112 Their propensity to stimulate
throughout the 3-year study, reaching its highest numbers
plant growth has been documented in grasslands110 and
3 years post-fire.
in subarctic tree-line communities.113 Their ability to alter
The Twin Lakes Fire had mixed effects on the bird
the succession and stability of aspen ecosystems also is
community in the Oregon Blue Mountains. In general,
well-documented.15, 89, 90, 91 Where herbivores suppress
fewer species used the most severely burned forest
deciduous plants in forests, they advance succession,
compared with unburned and less severely burned forest.
increase nutrient flow to the conifer overstory, and thereby
However, several species seemed to be opportunistically
may reduce forest productivity over time.119, 120, 121
fire-dependent and clearly benefited from the 1994 burn.
Influences of herbivores can be most perplexing when
Changes in nesting and foraging opportunities
they interact with episodic disturbance. In Abies and
simultaneously triggered positive responses from some
Pseudotsuga forests, for example, successional potential is
species (e.g., cavity nesters and aerial feeders) and negative
predisposed by site characteristics, past disturbance, and
responses from others (e.g., overstory nesters and foliage
by the reproductive strategies and abundance of
gleaners). One relatively rare species (black-backed
individual plants (i.e., seed banks, rootstocks). Secondary
woodpecker) and 1 rapidly declining species (olive-sided
succession is initiated by episodic agents (e.g., silviculture
flycatcher) were both found to be positively affected by
and fire), and its starting point and trajectory are
the fire. These species, it seems, are precariously
influenced by the attributes that define each disturbance
dependent on stand-replacement wildfires occurring on
agent’s regime (e.g., intensity and size of disturbance).72,
the landscape, even though their use of a recently burned 73, 92, 141, 156
Subsequent episodic disturbances and herbivory
forest may be short term (1-3 years). Additional research
may then further modify succession. Under these
to better understand the role of wildfire as a natural
circumstances, the influence of any disturbance agent can
disturbance event on bird communities in forested
be contingent, or dependent, on the influences of other
landscapes is needed.
disturbance agents. Such contingent influences have been
described for grasslands77 and in aspen forests, 15 and
obviously have implications for any land management
strategy that is based on disturbance management.
Nonetheless, multi-agent contin-gencies have not been
deeply explored in forests.
Herbivore exclosures have been used to study the
influences of herbivory on vegetation development in
various settings, and where several exclosures are used
to document a range of episodic disturbance, they can
provide insight on the role of disturbance-agent
interactions, and on the relative importance of individual
agents to development of communities over time. The
 230 Wildlife–Habitat Relationships in Oregon and Washington

study on which this case history is based127c explored herbivores feed selectively when given choices implies that
influences of ruminants on community composition, some taxa are always predisposed to suppression, even
biomass accumulation, and nutrient pools under varied when grazing pressure is low on average across the plant
episodic-disturbance regimes that were defined by community.143
variation in logging history, fire regime, and grass seeding. Results of this study also implicated herbivores in
Seven exclosures, built in the mid-1960s, were studied controlling understory biomass accumulation and nutrient
in the following 3 localities of the Blue Mountains dynamics. At the end of the experiment, total understory
Ecological Province:57 Mottet (n = 1), Hoodoo (n = 3), Hall and forest-floor biomasses respectively averaged 2.1 and
Ranch (n = 3). All but 1 of the sites were dominated by 1.5 times greater inside exclosures than outside. Amounts
vegetation types within the Abies grandis forest series and of total and exchangeable Ca at the forest floor were greater
had long histories of use by wild and domestic herbivores. inside exclosures than outside (18% and 11%, respectively).
Although undocumented, the sites may have been grazed Amounts of total and exchangeable Mg at the forest floor
by the horses of Native Americans in the eighteenth and were also greater inside than outside (20% and 16%,
early nineteenth century. Elk and mule deer were respectively). Concentration of total N at the forest floor
indigenous, but were nearly extirpated between 1870 and was 26% higher inside exclosures than outside, but
1900. Domestic ruminants increased dramatically during concentration of mineral N did not differ significantly. No
the same period.82 All 7 sites probably were grazed to some differences were detected for nutrients in soil.
extent by domestic sheep in the first half of the twentieth Shrubs were major contributors to the litterfall inside
century. Sheep grazing extended to present day in the the exclosures, particularly in clear-cuts (where potential
vicinity of Mottet, but no evidence of sheep was observed for shrub expression was greatest). Also, shrub foliage (and
at the exclosure itself during the 27 years of the experiment. that of forbs) contained relatively high concentrations of
During experiments, cattle were the dominant livestock N, phosphorus (P), sulfur (S), Ca, and Mg compared to
at all sites except Mottet. All 7 study sites were also subject graminoids. Greater amounts of these nutrients inside
to use by elk and both white-tailed and mule deer. Whereas exclosures probably reflected the contribution of shrub
livestock were more prevalent than wild herbivores early foliage to forest-floor biomass as well. Greater
in the century, wild ruminants were variously more accumulation of forest-floor Ca inside exclosures probably
common during the exclosure experiments summarized reflected translocation of Ca from the lower soil profile
here. by some shrubs.19
Plant communities were sampled periodically between In conclusion, herbivores influence forests by
1965 and 1995 inside and outside of all 7 exclosures. selectively suppressing plant taxa and by accelerating the
Similarly, total biomass and nutrients of nonconiferous cycling of nutrients. Based on results in some shrub-
understory and forest floor were estimated in 1995. dominated riparian communities,66, 148 one can logically
Composition of herbivore diets was assessed using hand- hypothesize that both of these influences have implications
reared elk in 1992.127c for ecosystem management. First, the herbivory regime
Throughout the experiments, several key influences of can influence the trajectory of succession and change its
herbivory were documented, including (1) suppressed terminus, thereby confounding the seral and climax
development of shrub canopies; (2) suppressed vegetations that managers use in forest planning.66, 127c
accumulation of standing biomass and litter; and (3) Second, herbivores can suppress the forest’s carrying
reduced nutrient accumulations at forest floors (total N, capacity for fauna linked to shrubs in forest understories.49,
148
total and exchangeable calcium [Ca] and magnesium This may have its greatest relevance in predicting
[Mg]). Suppression of deciduous shrubs is a common avifaunal composition because many birds typically nest
result of ruminant herbivory in forest ecosystems.7, 75, 89, 120, on the ground and in shrub canopies, rather than in forest
155, 171
Regression and cluster analyses suggested a overstories. Finally, by suppressing some plants via
continuum of vegetation response to elk dietary selective feeding, herbivores can reduce the rate at which
preferences on the study sites. Deciduous shrubs certain nutrients are replenished on sites following
predominated at one extreme, whereas unpalatable herbs episodic disturbance. This observation probably has its
dominated at the other. Forage preferences of elk greatest relevance in the context of frequent burning.
correlated with development of plant taxa over time Burning volatizes nutrients, especially N and S, thereby
(P<0.002), and several taxa were predisposed to contributing to their loss.60a, 154a The stability of the burned
suppression by their high palatability and low abundance. site’s nutrient pool is thus influenced by the rate of loss
These included several indicators of climax community via volitization (as functions of the frequency and intensity
(e.g., Acer glabrum, Taxus brevifolia, Linnaea borealis), shrubs of burning) and by the rate of nutrient replenishment (as
that contribute vertical structure and leaf litter during a function of passive input and N fixing, the latter being
forest succession (Acer glabrum, Amelanchier alnifolia, Salix potentially controlled by herbivores through their selective
spp., Sorbus sitchensis), and plants that actively replenish herbivory). In this study, the most profound influence of
N pools after episodic loss (Astragalus canadensis, Ceanothus wild and domestic ruminant herbivores may have been
sanguineus, C. velutinus). The consequences of “over- their modification of site production potential, via selective
grazing” by herbivores at or near maximum carrying suppression of N-fixing plants.
capacity are universally recognized. However, the fact that
 CHAPTER 8: EASTSIDE FORESTS 231

American Marten. In summer 1993, the U.S. Forest Service

Case History 4 Pacific Northwest Research Station began a five-year study
A Review of Wildlife-habitat Relationships in on American marten in the central Oregon Pumice Zone
Managed Lodgepole Pine Forests of the (U.S. Forest Service, 1999 Annual Report, U.S. Forest
Central Oregon Pumice Zone Service, Olympia, WA). Objectives of this study were to
Perhaps one of the least understood habitats in Eastside (1) obtain basic natural history and habitat selection
(interior) forests and woodlands is the lodgepole pine information on martens in lodgepole pine-bitterbrush
forest of the central Oregon Pumice Zone, particularly with communities; (2) evaluate movement and behavior of
respect to wildlife-habitat relationships. This area lies east martens during winter; (3) evaluate marten response to
of the Cascade crest and encompasses >2.7 million acres different density, size, and configuration of slash piles at
(>1.1 million ha) within the Deschutes, Fremont, and the stand- and landscape-scale; and (4) determine the
Winema National Forests.162 The topography of this region influence of salvage logging on marten ecology. The study
strongly reflects its volcanic origin, with numerous cinder area is on the Chemult Ranger District of the Winema NF,
cones and volcanic domes arising from broad basins approximately 12.4 miles (20 km) northeast of Chemult,
formed by basalt flows.162 The entire region is blanketed OR. Martens were livetrapped and fitted with radiocollars
by recent pumice eruptions from Mount Mazama (Crater to evaluate home range size, use of resting and denning
Lake), Newberry Caldera, and other sources; pumice soil sites, and habitat selection patterns. One hundred and
sustains a climax forest of lodgepole pine with an antelope fifteen martens (71 M, 44 F) were captured during the
bitterbrush (Purshia tridentata) shrub component.57, 162 study and 6,251 radio relocations were obtained from
Wildfires and mountain pine beetle (Dendroctonus marked individuals. Martens were found resting or
ponderosae) epidemics were historical disturbances denning on 1,851 occasions (30% of all telemetry locations)
associated with lodgepole pine forests in central Oregon.76 in 821 resting sites, 70 denning sites, and 27 resting and
A history of fire suppression and recent mountain pine denning sites. Slash piles were most commonly used for
beetle epidemics, coupled with drought conditions, have resting and denning (44 and 40%, respectively). Down
resulted in extensive mortality in lodgepole and ponderosa logs, live trees, and snags also were used extensively by
pine forests in this region.76 Subsequently, salvage logging marten for resting and denning. Raphael and Jones126
of these forests has been implemented in recent years on reported seasonal differences in structures used for rest
several national forests east of the Cascade Mountains. sites by martens; 60% of resting sites were in slash piles
Salvage logging of forests subjected to fire, insects, and during snow-free periods, whereas only 17% were in slash
forest pathogens has increased over the past two decades. during times of snow. Additionally, martens in this area
This forestry practice typically influences vegetative generally used down logs more for denning (39%) than
structure by removing snags and down wood, but resting (29%). Raphael and Jones126 also reported that live
generally retains varying densities of green trees and trees, snags, and logs with diameters >19.7 inches (>50
snags. Salvage harvest prescriptions vary and can mimic cm) dbh were selected significantly more than predicted
clearcut or selective harvest techniques. Government from availability.
agencies often use a “pay-as-cut” timber sale contract, These data support the importance of managing
where purchasers pay only for wood volume removed. structural woody features in forests to provide habitat for
This differs from purchasing a fixed volume estimate, with marten and their prey. Retaining slash piles following
economic incentives to remove as much of the available timber harvest seems to provide important habitat for
wood volume as possible, unless logistics and/or costs martens126 (U.S. Forest Service, 1999 Annual Report, U.S.
are prohibitive. With a pay-as-cut sale, contractors Forest Service, Olympia, WA), and other species of wildlife
typically seek areas with concentrations of wood volume (U.S. Forest Service, Chemult Ranger District, unpubl.
(dead trees in this case) to reduce operational expenses, data). Retaining large (>19.7 inches [>50 cm] dbh) snags
and do not generally spend time and money to search for and down logs also is important for providing resting and
individuals and small groups of trees. Consequently, snags denning sites for marten. Raphael and Jones126 estimated
and down wood can be retained throughout harvest units that 0.5 live trees >19.7 inches dbh, 0.7 snags >19.7 inches
in varying densities and distributions. There are few dbh, 1.5 logs, and 3.2 slash piles per acre (0.2 live trees
published studies on wildlife-habitat relationships from >50 cm dbh, 0.3 snags >50 cm dbh, 0.6 logs, and 1.3 slash
the central Oregon Pumice Zone,126 and available data piles per ha) retained across the landscape would provide
generally are limited to work in progress (e.g., E. B. Arnett, habitat for resting and denning, and suggested that these
B. Altman, W. P. Erickson, and K. A. Bettinger, 2000 Final estimates of structure distribution could be relatively easy
Report, Weyerhaeuser Company, Springfield, OR), agency to accommodate in managed forests.
reports,65 and other unpublished “gray literature.” In this Forest Songbirds. Habitat relationships of forest songbirds
case history, we summarize available information on occupying lodgepole pine forests of the central Oregon
wildlife-habitat relationships in lodgepole pine forests of Pumice Zone are poorly understood. Herein, we
the central Oregon Pumice Zone, focusing on studies of summarize results of bird-habitat relationships as studied
American marten126 and forest songbirds (E. B. Arnett, B. in this region from 1995 to 1999 (E. B. Arnett, B. Altman,
Altman, W. P. Erickson, and K. A. Bettinger, 2000 Final W. P. Erickson, and K. A. Bettinger, 2000 Final Report,
Report, Weyerhaeuser Company, Springfield, OR). Weyerhaeuser Company, Springfield, OR). Objectives of
 232 Wildlife–Habitat Relationships in Oregon and Washington

this study were to (1) estimate bird species composition mammals and gray jays, abandonment, and possibly
and relative abundance in stands of lodgepole pine; (2) weather-related mortality. Cowbird parasitism was not
model relationships between structural habitat features observed during nest monitoring, though cowbirds were
and bird species relative abundance and probabilities of detected during point counts.
use in lodgepole pine stands; (3) monitor nests of key To illustrate wildlife-habitat relationships of birds in
species to evaluate reproductive success and nesting lodgepole pine forests in this region, we present summary
habitat relationships; and (4) evaluate relationships results of nest and relative abundance-habitat selection
between bird species abundance and nesting success and models for 3 species: chipping sparrow, dark-eyed junco,
salvage logging of lodgepole pine forests. The study area and hermit thrush (E. B. Arnett, B. Altman, W. P. Erickson,
lies in the northeast corner of the Chemult Ranger District and K. A. Bettinger, 2000 Final Report, Weyerhaeuser
of the Winema NF,157 and in the northwest portion of the Company, Springfield, OR). The probability of presence
Silver Lake Ranger District of the Fremont NF.158 of male chipping sparrows and dark-eyed juncos was
Forty-two bird species were recorded within 164 feet greater in areas containing fewer total live trees and
(50 m) during point count surveys on both the Fremont saplings, less canopy closure, and greater shrub cover.
and Winema national forests. Among species detected,17, Conversely, presence of male hermit thrushes was
15, and 10 species were permanent residents, Neotropical positively related to the number of live trees and saplings,
migrants, and short-distance migrants, respectively. The percent canopy closure, and percent seedling cover. When
mountain chickadee, yellow-rumped warbler, dusky comparing nest-site habitat characteristics with those at
flycatcher, dark-eyed junco, and chipping sparrow were random points (37.1 feet [11.3 m] fixed-radius plot), all
the most common species detected on the Fremont NF, three species selected nest sites with greater percent
whereas the mountain chickadee, yellow-rumped warbler, canopy closure. Chipping sparrow nest sites also were
chipping sparrow, dark-eyed junco, and American robin positively associated with greater percent seedling cover
were most common on the Winema NF. Results indicated and number of saplings. Dark-eyed juncos selected nest
that salvage logged stands consistently had a higher total sites in areas having greater percent cover of grasses and
number of species and mean number of species per stand forbs, greater percent seedling cover, and less bare ground
compared to reference stands. In general, relative cover.
abundance for the most common species, all species Results of this study generally support the
combined, and mean number of species per stand did not characterization of avian community guilds and species
differ (P >0.10) between reference and treatment stands composition of lodgepole pine forests. Because these
on both forests. For species having >20 detections, the forests are often single-story, dense, mono-typical forests
hermit thrush and red-breasted nuthatch were with limited structural complexity and a sparsely
significantly more abundant in reference stands compared vegetated understory, species richness in lodgepole pine
to treatment stands on the Fremont NF, whereas dark-eyed is typically low and species dominance is often high.
juncos, American robins, and Cassin’s finches were Despite geographic and climatic differences, avian species
significantly more abundant in treatment stands on this composition in lodgepole pine forest seems to be relatively
forest. The dusky flycatcher, hairy woodpecker, and red- consistent throughout the range of this forest type.
breasted nuthatch all were significantly more abundant Although no species is known to be dependent exclusively
in treatment stands on the Winema NF, while the gray on lodgepole pine forests, the abundance and consistency
flycatcher was more abundant in reference stands on this in occurrence suggest the importance of this forest type
forest. for several bird species, such as black-backed
Between 1997 and 1999, 420 nests of 21 different species woodpeckers.
were monitored within 12 100-acre (40-ha) nest plots (6 in Preliminary results of habitat selection models
salvage-logged stands and 6 in reference stands). Eighty- generally corroborate nesting and foraging strategies of
one per cent of these nests (n = 336) were from 6 species: avian species highlighted in this case history. Chipping
mountain chickadee (n = 83), yellow-rumped warbler (n sparrows and dark-eyed juncos usually are ground
= 63), dark-eyed junco (n = 58), chipping sparrow (n = foragers that are often associated with more open forest
57), American robin (n = 44), and dusky flycatcher (n = conditions, while hermit thrushes appear more associated
31). Average Mayfield estimates of nest success for the 6 with increasing density of trees and saplings, percent
most common species ranged from 30% (dark-eyed junco) canopy closure, and shrub cover. Birds in these lodgepole
to 70% (yellow-rumped warbler). Among nesting guilds, pine forests selected nest sites with specific structural
mean nest success was lowest for ground nesters (28.8%, habitat features related to cover for protection from
n = 2 species, n = 72 nests), highest for cavity nesters (66.1%, predators and weather, often features that were not
n = 8 species, n = 131 nests), and moderate for open-cup selected for by singing males at larger spatial scales. Thus,
foliage nesters (55.3%, n = 11 species, n = 217 nests). Nest identifying structural habitat components important to
success was similar between reference and treatment birds at multiple spatial scales will be valuable to
stands for 5 of the 6 common species: American robin nest managers as they incorporate the needs of birds in
success was greater in treatment stands compared to management planning.
reference stands. Though not specifically quantified, It may be possible to maintain forest bird relative
causes of nest failure included predation by small abundance and reproductive success in lodgepole pine
 CHAPTER 8: EASTSIDE FORESTS 233

stands that have been salvage logged under certain 1. The implications of prescribed burning on plant
prescriptions (E. B. Arnett, B. Altman, W. P. Erickson, and and subsequent invertebrate diversity are
K. A. Bettinger, 2000 Final Report, Weyerhaeuser unknown. Factors such as fire interval, intensity,
Company, Springfield, OR). The same authors duration, season, patchiness, and spatial extent
hypothesized that more diverse and structurally complex need to be examined to determine their effects on
harvest stands have a much greater probability of plant community composition and diversity and
maintaining bird species relative abundance, diversity, and abundance of invertebrate herbivores and
reproductive success. Thus,we believe that structure predators;
retained after this particular harvest treatment (e.g., live 2. Assessments of the viability of species, both in
tree and snag density, down logs, understory vegetation) space and time, that inhabit landscapes managed
could explain findings presented in this case history. for the natural range of variability;
3. Studies on effects of fire occurrence, fire
Future Research Needs suppression, season of burn (see A. L. Turner and
Inevitably, our synthesis of wildlife communities of R. Sallabanks, 1999 Annual Report, Sustainable
Eastside (interior) forests and woodlands generated more Ecosystems Institute, Meridian, ID), and changes
questions than it answered. Research in Eastside (interior) of vegetation pattern resulting from burns and fire
forests and woodlands, as elsewhere, has been heavily suppression on ground-nesting birds and their
biased toward charismatic megafauna, birds, and habitats (see Figure 12);
ungulates (especially game species). Nongame species, 4. Studies to determine the effects of herbivory on
especially smaller vertebrates such as bats, reptiles, and forest understory composition and associated bird
amphibians, have been relatively unstudied. In many cases communities;
we have yet to describe accurately the distribution and 5. Effects of salvage logging of fire- and insect-killed
abundance of a species, let alone study its relationship trees on predator-prey dynamics, behavior, habitat
with habitat structure or management activities. In suitability, and productivity and survivorship of
general, more information is needed on relationships wildlife species;
between wildlife species and the 3 predominant 6. Studies that address the effects of unevenaged
management issues in Eastside (interior) forests and management (e.g., thinning from below) in PPFW
woodlands: fire, timber harvest, and grazing. In addition, and EMCF on habitat selection and population
forest fragmentation, a process found to severely impact dynamics of wildlife species;
wildlife populations elsewhere in the U.S., has not yet been 7. Restoration efforts, such as management action, to
studied in Eastside (interior) forests and woodlands.138 restore aspen or old-growth ponderosa pine to
Relative to EMCF and PPFW, UAF and LPFW have been Eastside (interior) forests and woodlands. Such
poorly studied. Many research needs identified by forest efforts should incorporate a research component
managers in the Pacific Northwest (Oregon, Washington, to examine the response of wildlife habitat
and Idaho) are relevant to Eastside (interior) forests and restoration over the long-term; and
woodlands and have been discussed elsewhere.11 8. Studies that address how manipulation of natural
Below we offer a few specific examples of research processes, such as fire and ecological succession,
needs: can influence genetics and viability of populations
clearly dependent on those processes.

Figure 12. Ponderosa Pine Forest and

Woodlands have been heavily impacted
by fire suppression, grazing, and
logging. Restoration efforts today
include controlled burning to reduce
fuels. In this stand on the Umatilla
National Forest in northeast Oregon,
smoke still lingers from a recent fire
prescription.
 234 Wildlife–Habitat Relationships in Oregon and Washington

Acknowledgments 17. Betts, B. J. 1996. Roosting behaviour of silver-haired bats (Lasionycteris

noctivagans) and big brown bats (Eptesicus fuscus) in northeast Oregon.
The research discussed here has been funded by the U.S. Pages 55-61 in R. M. R. Barclay and R. M. Brigham, editors. Bats and
Forest Service, Boise Cascade Corporation, National forests symposium, Victoria, British Columbia, Canada. BC Ministry of
Council for Air and Stream Improvement, Blue Mountains Forests, Victoria, BC.
Natural Resources Institute, Weyerhaeuser Company, 18. ———. 1998. Roosts used by maternity colonies of silver-haired bats
Oregon Department of Fish and Wildlife, U.S. Fish and in northeastern Oregon. Journal of Mammalogy 79:643-650.
Wildlife Service, Oregon Agricultural Experiment Station, 19. Binkley, D., and D. L. Husted. 1983. Nitrogen accretion, soil fertility,
and Douglas-fir nutrition in association with redstem ceanothus.
and Sustainable Ecosystems Institute. We are especially
Canadian Journal of Forest Research 13:122-125.
grateful to D. H. Johnson and K. A. Bettinger for providing 20. Blackford, J. L. 1955. Woodpecker concentration in burned forest.
technical and logistical support throughout this project. Condor 57:28-30.
A. L. Turner provided Figure 1, and S. Edge made 21. Blake, J. G. 1983. Influence of fire and logging on nonbreeding bird
numerous editorial comments on an earlier version of this communities of ponderosa pine forests. Journal of Wildlife Management
manuscript. All other photographs courtesy of R. 46:404-415.
22. Bock, C. E., and J. H. Bock. 1974. On the geographical ecology and
Sallabanks.
evolution of the three-toed woodpeckers, Picoides tridactylis and P.
arcticus. American Midland Naturalist 92:397-405.
Literature Cited 23. ———, and ———. 1983. Responses of birds and deer mice to
1. Adams, E. M., and M. L. Morrison. 1993. Effects of forest stand structure prescribed burning in ponderosa pine. Journal of Wildlife Management
and composition on red-breasted nuthatches and brown creepers. 47:836-840.
Journal of Wildlife Management 57:616-629. 24. ———, and J. F. Lynch. 1970. Breeding bird populations of burned and
2. Agee, J. K. 1993. Fire ecology of pacific northwest forests. Island Press, unburned conifer forest in the Sierra Nevada. Condor 72:182-189.
Covelo, CA. 25. ———, M. Raphael, and J. H. Bock. 1978. Changing avian community
3. ———. 1996. Fire in the Blue Mountains: a history, ecology, and structure during early post-fire succession in the Sierra Nevada. Wilson
research agenda. Pages 119-145 in R. G. Jaindl and T. M. Quigley, editors. Bulletin 90:119-123.
Search for a solution: sustaining the land, people, and economy of the 26. Bowers, M. A. 1997. Influence of deer and other factors on an old-
Blue Mountains. American Forests, in cooperation with the Blue field plant community. Pages 310-326 in W. L. McShea, H. B. Underwood,
Mountains Natural Resources Institute, Washington, D.C. and J. H. Rappole, editors. The science of overabundance: deer ecology
4. ———. 1997. The severe weather wildfire: too hot to handle? and population management. Smithsonian Institution Press, Washington,
Northwest Science 71:153-56. D.C.
5. ———. 1998. The landscape ecology of western forest fire regimes. 27. Bull, E. L. 1991. Summer roosts and roosting behavior of Vaux’s swift
Northwest Science 72:24-34. in old-growth forests. Northwestern Naturalist 72:78-82.
6.Altman, B., and R. Sallabanks. Olive-sided flycatcher (Contopus cooperi). 28. ———, and B. E. Carter. 1996. Winter observations of tailed frogs in
In A. Poole, and F. Gill, editors. The birds of North America, No. 502. northeastern Oregon. Northwestern Naturalist 77:45-47.
2000. The Birds of North America, Inc., Philadelphia, PA. 29. ———, and J. E. Hohmann. 1992. The association between Vaux’s swift
7. Ammer, C. 1996. Impact of ungulates on structure and dynamics of and old-growth forests in northeastern Oregon. Western Birds 24:38-
natural regeneration of mixed mountain forests in the Bavarian Alps. 42.
Forest Ecology and Management 88:43-53. 30. ———, and R. S. Holthausen. 1993. Habitat use and management of
8. Anderson, R. C., and A. J. Katz. 1993. Recovery of browse-sensitive tree pileated woodpeckers in northeastern Oregon. Journal of Wildlife
species following release from white-tailed deer (Odocoileus virginianus Management 57:335-345.
Zimmerman) browsing pressure. Biological Conservation 63:203-208. 31. ———, and E. C. Meslow. 1977. Habitat requirements of the pileated
9. Arnett, E. B. 1990. Bighorn sheep habitat selection patterns and woodpecker in northeastern Oregon. Journal of Forestry 75:335-337.
response to fire and timber harvest in south-central Wyoming. Thesis, 32. ———, C. G. Parks, and T. R. Torgersen. 1997. Trees and logs
University of Wyoming, Laramie, WY. important to wildlife in the Interior Columbia River Basin. U.S. Forest
10. ———, L. L. Irwin, F. G. Lindzey, and T. J. Hershey. 1990. Use of Service, General Technical Report PNW-391.
clearcuts by Rocky Mountain bighorn sheep in south-central Wyoming. 33. ———, T. R. Torgersen, A. K. Blumton, C. M. McKenzie, and D. S.
Proceedings of the Northern Wild Sheep and Goat Council 7:194-205. Wyland. 1995. Treatment of an old-structure stand and the effect on
11. ———, and R. Sallabanks. 1998. Land manager perceptions of avian birds, ants, and large woody debris: a case study. U.S. Forest Service,
research and information needs: a case study. Pages 399-413 in J. M. General Technical Report PNW-353.
Marzluff and R. Sallabanks, editors. Avian conservation: research and 34. Bunnell, F. L., L. L. Kremaster, and R. W. Wells. 1997. Likely
management. Island Press, Covelo, CA. consequences of forest management on terrestrial, forest-dwelling
12. Arno, S. F., H. Y. Smith, and M. A. Krebs. 1997. Old-growth ponderosa vertebrates in Oregon. Report M-7 of the Centre for Applied
pine and western larch stand structures: influences of pre-1900 fires Conservation Biology, University of British Columbia,Vancouver, BC.
and fire exclusion. U.S. Forest Service, Research Paper INT-RP-495. 35. Campbell, L. A., J. G. Hallett, and M. A. O’Connell. 1996. Conservation
13. Barnhardt, S. J., J. R. McBride, C. Cicero, P. da Silva, and P. Warner. 1987. of bats in managed forests: use of roosts by Lasionycteris noctivagans.
Vegetation dynamics of the northern oak woodland. Pages 53-58 in T. R. Journal of Mammalogy 77:976-984.
Plumb and N. H. Pillsbury, technical coordinators. Proceedings of the 36. Caraher, D. L., J. Henshaw, F. Hall (and others), panel members. 1992.
Symposium on multiple-use of California’s hardwood resources. U.S. Restoring ecosystems in the Blue Mountains: a report to the regional
Forest Service, General Technical Report PSW-100. forester and the forest supervisors of the Blue Mountain forests. U.S.
14. Barret, S., and S. Arno. 1982. Indian fires as an ecological influence in Forest Service, Pacific Northwest Region.
the Northern Rockies. Journal of Forestry 80:647-651. 37. Chadde, S. W., and C. E. Kay. 1991. Tall-willow communities on
15. Bartos, D. L., J. K. Brown, and G. D. Booth. 1994. Twelve years of Yellowstone’s northern range: a test of the “natural-regulation”
biomass response in aspen communities following fire. Journal of Range paradigm. Pages 231-262 in R. B. Keiter and M. S. Boyce, editors. The
Management 47:79-83. Greater Yellowstone ecosystem: redefining America’s wilderness
16. Bendell, J. F. 1974. Effects of fire on birds and mammals. Pages 73-138 heritage. Yale University Press, New Haven, CT.
in T. T. Kozlowski and C. E. Ahlgren, editors. Fire and ecosystems. 38. Chappell, C.B., R.C. Crawford, C. Barrett, J. Kagan, D.H. Johnson, M.
Academic Press, New York, NY. O’Mealy, G.A. Green, H.L. Ferguson, W.D. Edge, E.L. Greda, and T.A.
 CHAPTER 8: EASTSIDE FORESTS 235

O’Neil. 2001. Wildlife habitats: descriptions, status, trends, and system 59. Galbraith, W. A., and E. W. Anderson. 1991. Grazing history of the
dynamics. Pages 22-114 in D. H. Johnson and T. A. O’Neil, managing Northwest. Rangelands 13:213-218.
directors. Wildlife-habitat relationships in Oregon and Washington. 60. Garrett, K. L., M. G. Raphael, and R. D. Dixon. 1996. White-headed
Oregon State University Press, Corvallis, OR. woodpecker (Picoides albolarvatus). In A. Poole and F. Gill, editors.The
39. Christy, R. E., and S. D. West. 1993. Biology of bats in Douglas-fir birds of North America, No. 252. The Birds of North America, Inc.,
forests. U.S. Forest Service, General Technical Report PNW-GTR-308. Philadelphia, PA.
40. Cody, M. L., editor. 1985. Habitat selection in birds. Academic Press, 60a. Geist, J.M. 1974. Chemical characteristics of some forest and
New York, NY. grassland soils of northeastern Oregon. U.S. Forest Service, Research
41. Cooper, S. V., K. E. Neiman, and D. W. Roberts. 1991. Forest habitat Note PNW-217.
types of northern Idaho: a second approximation. U.S. Forest Service, 61. Gibilisco, C. J. 1994. Distributional dynamics of modern Martes in
General Technical Report INT-236. North America. Pages 59-71 in S. W. Buskirk, A. S. Harestad, M. G.
42. Covington, W. W., R. L. Everett, R. Steele, L. L. Irwin, T. A. Daer, and A. N. Raphael, and R. A. Powell, editors. Martens, sables, and fishers biology
D. Auclair. 1994. Historical and anticipated changes in forest ecosystems and conservation. Cornell University Press, Ithaca, NY.
of the inland west of the United States. Pages 13-63 in R. N. Sampson 62. Gill, R. M. A. 1992a. A review of damage by mammals in north
and D. L. Adams, editors. Assessing forest ecosystem health in the temperate forests: 1. Deer. Forestry 65:145-169.
Inland West. The Haworth Press, New York, NY. 63. ———. 1992b. A review of damage by mammals in north temperate
43. ———, and M. M. Moore. 1994. Postsettlement changes in natural fire forests: 3. Impact on trees and forests. Forestry 65:363-388.
regimes and forest structure: ecological restoration of old growth 64. Gillen, R. L., W. C. Krueger, and R. F. Miller. 1985. Cattle use of riparian
ponderosa pine forests. Pages 153-181 in R. N. Sampson and D. L. meadows in the Blue Mountains of northeastern Oregon. Journal of
Adams, editors. Assessing forest ecosystem health in the Inland West. Range Management 38:205-209.
The Haworth Press, New York, NY. 65. Goggans, R., R. D. Dixon, and L. C. Seminara. 1987. Habitat use by
44. Crampton, L. H., and R. M. R. Barclay. 1998. Selection of roosting and three-toed and black-backed woodpeckers. Oregon Department of
foraging habitat by bats in different-aged aspen mixed-wood stands. Fish and Wildlife, Nongame Report 87-3-02, Bend, OR.
Conservation Biology 12:1347-1458. 66. Green, D. M., and J. B. Kauffman. 1985. Succession and livestock
45. Cross, S. P. 1986. Bats. Pages 497-517 in A. Y. Cooperrider, R. J. Boyd, grazing in a northeastern Oregon riparian system. Journal of Range
and H. R. Stuart, editors. Inventory and monitoring of wildlife habitat. Management 48:307-314.
U.S. Bureau of Land Management, BLM Service Center, Denver, CO. 67. Gruell, G. E. 1985. Indian fires in the Interior West: a widespread
46. Csuti, B., A. J. Kimerling, T. A. O’Neil, M. M. Shaughnessy, E. P. Gaines, influence. Pages 68-74 in Proceedings: Symposium and workshop on
and M. M. P. Huso. 1997. Atlas of Oregon wildlife: distribution, habitat, wilderness fire. U.S. Forest Service, General Technical Report INT-182.
and natural history. Oregon State University Press, Corvallis, OR. 68.Hamilton, R. C. 1993. Characteristics of old growth forest in the
48. Davis, J. W., G. A. Goodwin, and R. A. Ockenfels, technical coordinators. Intermountain Region. U.S. Forest Service, Intermountain Resource
1983. Snag habitat management: proceedings of the Symposium. U.S. Station, Ogden, UT.
Forest Service, General Technical Report RM-99. 69. Harrod, R. J., R. J. Taylor, W. L. Gaines, T. Lillybridge, and R. Everett. 1996.
48a. DeByle, N.V., and R.P. Winokur, editors. 1985. Aspen: Ecology and Pages 107-117 in R. G. Jaindl and T. M. Quigley, editors. Search for a
management in the western United States. U.S. Forest Service, General solution: sustaining the land, people, and economy of the Blue
Technical Report RM-119. Mountains. American Forests, in cooperation with the Blue Mountains
48b. ———, C.D. Bevins, and W.C. Fischer. 1987. Wildlife occurrence in Natural Resources Institute, Washington, D.C.
aspen in the interior western United States. Western Journal of Applied 70. Hein, D. 1980. Management of lodgepole pine for birds. Pages 238-246
Forestry 2:73-76. in R. M. DeGraaf, technical coordinator. Management of western forests
49. deCalesta, D. S. 1994. Effects of white-tailed deer on songbirds within and grasslands for nongame birds. U.S. Forest Service, General Technical
managed forests in Pennsylvania. Journal of Wildlife Management Report INT-86.
58:711-718. 71. Heinselman, M. L. 1973. Fire in the virgin forests of the Boundary
50. Doerr, P. D., L. B. Keith, and D. H. Rusch. 1971. Effects of fire on a Waters Canoe Area, Minnesota. Quarterly Research (NY) 3:329-382.
ruffed grouse population. Tall Timbers Fire Ecology Conference 72. ———. 1978. Fire in wilderness ecosystems. Pages 248-278 in J. C.
Proceedings 10:25-46. Hendee, G. H. Stankey, and R. C. Lucas, editors. Wilderness
51. Emlen, J. T. 1970. Habitat selection by birds following a forest fire. management. U.S. Forest Service, Miscellaneous Publication No. 1365.
Ecology 51:343-345. 73. ———. 1981. Fire intensity and frequency as factors in the
52. Fenton, M. B. 1997. Science and the conservation of bats. Journal of distribution and structure of northern ecosystems. Pages 7-57 in H. A.
Mammalogy 78:1-14. Mooney, T. M. Bonnicksen, N. L. Christensen, J. E. Lotan, and W. A.
53. Finch, D. M. 1992. Threatened, endangered, and vulnerable species of Reiners, technical coordinators. Fire regimes and ecosystem properties.
terrestrial vertebrates in the Rocky Mountain Region. U.S. Forest U.S. Forest Service, General Technical Report WO-26.
Service, General Technical Report RM-215. 74. Henjum, M. G., J. R. Karr, D. L. Bottom, D. A. Perry, J. C. Bednarz, S. G.
54. Finch, R. B. 1984. Fire history of selected sites on the Okanogan Wright, S. A. Beckwitt, and E. Beckwitt. 1994. Interim protection for
National Forest. U.S. Forest Service, Pacific Northwest Region, late-successional forests, fisheries, and watersheds: national forests east
Okanogan National Forest, Okanogan, WA. of the Cascade crest, Oregon and Washington. The Wildlife Society,
55. Frank, D. A., and R. D. Evans. 1997. Effects of native grazers on Bethesda, MD.
grassland N cycling in Yellowstone National Park. Ecology 78:2238- 75. Hernandez, M. P. G., and F. J. Silva-Pando. 1996. Grazing effects of
2248. ungulates in a Galician oak forest (northwest Spain). Forest Ecology
56. ———, R. S. Inouye, N. Huntly, G. W. Minshall, and J. E. Anderson. 1994. and Management 88:65-70.
The biogeochemistry of a north-temperate grassland with native 76. Hessburg, P. F., R. G. Mitchell, and G. M. Filip. 1994. Historical and
ungulates: nitrogen dynamics in Yellowstone National Park. current roles of insects and pathogens in eastern Oregon and
Biogeochemistry 10:163-180. Washington forested landscapes. U.S. Forest Service, General Technical
57. Franklin, J. F., and C. T. Dyrness. 1988. Natural vegetation of Oregon Report PNW-327.
and Washington. Oregon State University Press, Corvallis, OR. 77. Hobbs, N. T., D. S. Schimel, C. E. Owensby, and D. S. Ojima. 1991. Fire
58. Fretwell, S. D., and H. L. Lucas. 1970. On territorial behavior and other and grazing in the tallgrass prairie: contingent effects on nitrogen
factors influencing habitat distribution in birds. Acta Biotheoretica budgets. Ecology 72:1374-1382.
19:16-36.
 236 Wildlife–Habitat Relationships in Oregon and Washington

78. ———, and R. A. Spowart. 1984. Effects of prescribed fire on 97. Larsen, E. M., and J. T. Morgan. 1998. Management recommendations
nutrition of mountain sheep and mule deer during winter and spring. for Washington’s priority habitats: Oregon white oak woodlands.
Journal of Wildlife Management 48:551-560. Washington Department of Fish and Wildlife, Olympia, WA.
79. Huff, M. H., J. K. Agee, and D. A. Manuwal. 1985. Postfire succession of 98. Lawrence, G. E. 1966. Ecology of vertebrate animals in relation to
avifauna in the Olympic Mountains, Washington. Pages 8-15 in Fire’s chaparral fire in the Sierra Nevada foothills. Ecology 47:278-291.
effects on wildlife habitat-Symposium proceedings. U.S. Forest Service, 99. Leonard, W. P., H. A. Brown, L. L. C. Jones, K. R. McAllister, and R. M.
General Technical Report INT-186. Storm. 1993. Amphibians of Washington and Oregon. Seattle Audubon
80a. Hutto, R. L. 1995. 1995 Annual Report. University of Montana, Society, Seattle, WA.
Missoula, MT. 100. Lewis, S. E. 1995. Roost fidelity of bats: a review. Journal of
80b. ———. 1995. Composition of bird communities following stand- Mammalogy 6:481-496.
replacement fires in northern Rocky Mountain (U.S.A.) conifer forests. 101. Lyon, L. J., and J. M. Marzluff. 1985. Fire’s effects on a small bird
Conservation Biology 9:1041-1058. population. Pages 16-22 in Fire’s effects on wildlife habitat—
81. ———, S. M. Pletschet, and P. Hendricks. 1986. A fixed-radius point Symposium proceedings. U.S. Forest Service, General Technical Report
count method for nonbreeding and breeding season use. The Auk INT- 186
103:593-602. 102. Mannan, R. W., and E. C. Meslow. 1984. Bird populations and
82. Irwin, L. L., J. G. Cook, R. A. Riggs, and J. M. Skovlin. 1994. Effects of vegetation characteristics in managed and old-growth forests,
long-term grazing by big game and livestock in the Blue Mountains northeastern Oregon. Journal of Wildlife Management 48:1219-1238.
forest ecosystems. U.S. Forest Service, General Technical Report PNW- 103. Marcot, B. G. 1985. Habitat relationships of birds and young-growth
325. Douglas-fir in northwestern California. Dissertation, Oregon State
83. James, F. C., and H. H. Shugart, Jr. 1970. A quantitative method of University, Corvallis, OR.
habitat description. Audubon Field Notes 24:727-736. 104. ———. 1996. An ecosystem context for bat management: a case
84. ———, and N. O. Wamer. 1982. Relationships between temperate study of the interior Columbia River Basin, U.S.A. Pages 19-36 in R. M.
forest bird communities and vegetation structure. Ecology 63:159-171. R. Barclay and R. M. Brigham, editors. Bats and forests symposium,
85. Johnson, C. G., Jr., and R. R. Clausnitzer. 1992. Plant associations of the Victoria, British Columbia, Canada. BC Ministry of Forests, Victoria, BC,
Blue and Ochoco mountains. U.S. Forest Service, Pacific Northwest Canada.
Region, R6-ERW-TP-036-92. 105. ———, M. A. Castellano, J. A. Christy, L. K. Croft, J. F. Lehmkuhl, R. H.
86. Kalcounis, M. C., and R. M. Brigham. 1998. Secondary use of aspen Naney, K. Nelson, C. G. Niwa, R. E. Rosentreter, R. E. Sandquist, B. C.
cavities by tree-roosting big brown bats. Journal of Wildlife Wales, and E. Zieroth. 1997. Terrestrial ecology assessment. Pages
Management 62:603-611. 1497-1713 in T. M. Quigley and S. J. Arbelbide, editors. An assessment of
87. Kauffman, J. B., M. Mahrt, L. Mahrt, and W.D. Edge. 2001. Wildlife of ecosystem components in the interior Columbia Basin and portions of
riparian habitats. Pages 361-388 in D.H. Johnson and T.A. O’Neil, the Klamath and Great Basins. Volume III. U.S. Forest Service, General
managing directors. Wildlife-habitat relationships in Oregon and Technical Report PNW-405.
Washington. Oregon State University Press, Corvallis, OR. 106. ———, L. K. Croft, J. F. Lehmkuhl, R. H. Naney, C. G. Niwa, W. R.
88. Kay, C. E. 1995a. Aboriginal overkill and native burning: implications Owen, and R. E. Sandquist. 1998. Macroecology, paleoecology, and
for modern ecosystem management. Proceedings of the 8th George ecological integrity of terrestrial species and communities of the
Wright Society Conference on research and resource management on interior Columbia River Basin and portions of the Klamath and Great
public lands. Basins. U.S. Forest Service, General Technical Report PNW-410.
89. ———. 1995b. Browsing by native ungulates: effects on shrub and 107. ——— and M. Vander Heyden. 2001. Key ecological functions of
seed production in the Greater Yellowstone Ecosystem. Pages 310-320 wildlife species. Pages 168-186 in D.H. Johnson and T.A. O’Neil,
in B. A. Roundy, E. D. McArthur, J. S. Haley, and D. K. Mann, compilers. managing directors. Wildlife-habitat relationships in Oregon and
Proceedings: wildland shrub and arid land restoration symposium. U.S. Washington. Oregon State University Press, Corvallis, OR.
Forest Service, General Technical Report INT-315. 108. Mariani, P., and D. A. Manuwal. 1990. Factors influencing brown
90. ———. 1997.The condition and trend of aspen, Populus tremuloides, in creeper (Certhia americana) abundance patterns in the southern
Kootenay and Yoho national parks: implications for ecological integrity. Washington Cascade Range. Studies in Avian Biology 13:53-57.
Canadian Field-Naturalist 111:607-616. 109. McCullough, D. R. 1979. The George Reserve deer herd. The
91. ———, and S. Chadde. 1992. Reduction of willow seed production by University of Michigan Press, Ann Arbor, MI.
ungulate browsing in Yellowstone National Park. Pages 92-99 in W. P. 110. McNaughton, S. J. 1976. Serengeti migratory wildebeest: facilitation
Clary, E. D. McArthur, D. Bedunah, and C. L. Wambolt, compilers. of energy flow by grazing. Science 191:92-94.
Proceedings of Symposium on ecology and management of riparian 111. ———. 1992. The propagation of disturbance in savannas through
shrub communities. U.S. Forest Service, General Technical Report INT- food webs. Journal of Vegetation Science 3:301-314.
289. 112. McShea, W. J., and J. H. Rappole. 1997. Herbivores and the ecology of
92. Kilgore, B. M. 1981. Fire in ecosystem distribution and structure: forest understory birds. Pages 298-309 in W. J. McShea, H. B.
western forests and scrublands. Pages 58-59 in H. A. Mooney, T. M. Underwood, and J. H. Rappole, editors. The science of overabundance:
Bonnicksen, N. L. Christensen, J. E. Lotan, and W. A. Reiners, technical deer ecology and population management. Smithsonian Institution
coordinators. Fire regimes and ecosystem properties. U.S. Forest Press, Washington, D.C.
Service, General Technical Report WO-26. 113. Molvar, E. M., R. T. Bowyer, and V.Van Ballenberghe. 1993. Moose
93. Knowles, C. J., and P. R. Knowles. 1993. A bibliography of literature and herbivory, browse quality, and nutrient cycling in an Alaskan treeline
papers pertaining to presettlement wildlife and habitat of Montana and community. Oecologia 94:472-470.
adjacent areas. FaunaWest Wildlife Consultants, Boulder, MT. 114. Noss, R. F., E. T. LaRoe, III, and J. M. Scott. 1995. Endangered
94. Kroll, J. C., W. D. Goodrum, and P. J. Behrman. 1986. Twenty-seven ecosystems of the United States: a preliminary assessment of loss and
years of over-browsing: implications to white-tailed deer management degradation. Biological Report 28, U.S. Department of the Interior,
on wilderness areas. Pages 294-303 in D. L. Kulhavy and R. N. Conner, National Biological Service, Washington, D.C.
editors. Wilderness and natural areas in the eastern United States: a 115. O’Hara, K. L., P. A. Latham, P. F. Hessburg, and B. G. Smith. 1996. A
management challenge. Center for Applied Studies, School of Forestry, structural classification for inland northwest forest vegetation. Journal
Stephen F. Austin University, Nacagdoches, TX. of Applied Forestry 11:97-102.
95. Kunz, T. H. 1982. Roosting ecology of bats. Pages 30-55 in T. H. Kunz, 116. Oliver, C. D. 1981. Forest development in North America following
editor. Ecology of bats. Plenum Press, New York, NY. major disturbances. Forest Ecology and Management 3:153-168.
96. Langston, N. 1995. Forest dreams, forest nightmares. University of
Washington Press, Seattle, WA.
 CHAPTER 8: EASTSIDE FORESTS 237

117. Paquet, P., and A. Hackman. 1995.Large carnivore conservation in the 133. Ruess, R. W. 1987. The role of large herbivores in nutrient cycling of
Rocky Mountains: a long-term strategy for maintaining free-ranging and tropical savannas. Pages 93-141 in B. H. Walker, editor. Determinants of
self-sustaining populations of carnivores. World Wildlife Fund, tropical savannas. International Union of Biological Scientists, Paris,
Washington, D.C. France.
118. Parsons, D. J., and S. H. DeBenedetti. 1979. Impact of fire suppression 134. ———, and S. J. McNaughton. 1987. Grazing and the dynamics of
on a mixed-conifer forest. Forest Ecology and Management 2:21-33. nutrient and energy regulated microbial processes in the Serengeti
119. Pastor, J., and Y. Cohen. 1997. Herbivores, the functional diversity of grasslands. Oikos 49:101-110.
plant species, and the cycling of nutrients in ecosystems. Theoretical 135. Saab,V. A., and J. G. Dudley. 1998. Responses of cavity-nesting birds to
Population Biology 51:165-179. stand-replacement fire and salvage logging in ponderosa pine/Douglas-
120. ———, and R. J. Naiman. 1992. Selective foraging and ecosystem fir forests of southwestern Idaho. U.S. Forest Service, Research Paper
processes in boreal forests. American Naturalist 139:691-705. RM-11.
121. ———, ———, B. Dewey, P. McInnes, and Y. Cohen. 1993. Moose 136. ———, and T. D. Rich. 1997. Large-scale conservation assessment
browsing and soil fertility in the boreal forests of Isle Royale National for Neotropical migratory land birds in the Interior Columbia River
Park. Ecology 74:467-480. Basin. U.S. Forest Service, General Technical Report PNW-399.
121a. Peek, J.M., D.A. Demarchi, and D.E. Stucker. 1985. Bighorn sheep and 137. Sallabanks, R., E. B. Arnett, T. B. Wigley, and L. L. Irwin. In press.
fire: seven case histories. Pages 36-43 in Fire’s effects on wildlife Accommodating birds in managed forests of North America: A review
habitat—Symposium proceedings. U.S. Forest Service, General of bird-forestry relationships. National Council for Air and Stream
Technical Report INT-186. Improvement Technical Bulletin: In press.
122. Perkins, J. M., J. M. Barss, and J. Peterson. 1990. Winter records of 138. ———, P. J. Heglund, J. B. Haufler, B. A. Gilbert, and W. Wall. 1999.
bats in Oregon and Washington. Northwestern Naturalist 71:59-62. Forest fragmentation of the Inland West: issues, definitions, and
123. ———, and S. P. Cross. 1988. Differential use of some coniferous potential study approaches for forest birds. Pages 187-199 in J. A.
forest habitats by hoary and silver-haired bats in Oregon. The Murrelet Rochelle, L. A. Lehmann, and J. Wisniewski, editors. Forest
69:21-24. fragmentation: wildlife and management implications. Brill Academic
124. Quigley, T. M., R. W. Haynes, and R. T. Russell, technical editors. 1996. Publishers, Leiden, The Netherlands.
Integrated scientific assessment for ecosystem management in the 139. ———, and F. C. James. 1999.American robin (Turdus migratorius). In
interior Columbia Basin and portions of the Klamath and Great Basins. A. Poole and F. Gill, editors. The birds of North America, No. 462. The
U.S. Forest Service, General Technical Report PNW-382. Birds of North America, Inc., Philadelphia, PA.
125. Raphael, M. G. 1983. Cavity-nesting bird response to declining snags 140. Sampson, R. N., D. L. Adams, S. S. Hamilton, S. P. Mealey, R. Steele, and
on a burned forest: a simulation model. Pages 211-215 in J. W. Davis, G. D.Van De Graaff. 1994. Assessing forest ecosystem health in the Inland
A. Goodwin, and R. A. Ockenfels, technical coordinators. Snag habitat West. Pages 3–10 in R. N. Sampson and D. L. Adams, editors. Assessing
management: Proceedings of the Symposium. U.S. Forest Service, forest ecosystem health in the Inland West. The Haworth Press, New
General Technical Report RM-99. York, NY.
126. ———, and L. L. C. Jones. 1997. Characteristics of resting and 141. Schimmel, J., and A. Granstrom. 1996. Fire severity and vegetation
denning sites of American martens in central Oregon and western response in the boreal Swedish forest. Ecology 77:1436-1450.
Washington. Pages 146-165 in G. Proulx, H. N. Bryant, and P. M. 142. Seagle, S. W., S. J. McNaughton, and R. W. Ruess. 1992. Simulated
Woodard, editors. Martes: taxonomy, ecology, techniques and effects of grazing on soil nitrogen and mineralization in contrasting
management. The Provincial Museum of Alberta, Edmonton, Alberta, Serengeti grasslands. Ecology 73:1105-1123.
Canada. 143. Smith, A. D. 1965. Determining common use grazing capacities by
127.———, and M. White. 1984. Use of snags by cavity-nesting birds in application of the key species concept. Journal of Range Management
the Sierra Nevada. Wildlife Monographs 86:1-66. 18:196-201.
127a. Riggs, R.A., and J.M. Peek. 1980. Mountain sheep habitat use 144. Steele, R. 1994. The role of succession in forest health. Pages 183–
patterns related to post-fire succession. Journal of Wildlife Management 190 in R. N. Sampson and D. L. Adams, editors. Assessing forest
44:933-938. ecosystem health in the Inland West. The Haworth Press, New York,
127b. ———, S. Bunting, and S.E. Daniels. 1996. Prescribed fire. Pages NY.
295-319 in P.R. Krausemann, editor. Rangeland wildlife. Society for 145. ———, S. F. Arno, and K. Geier-Hayes. 1986. Wildfire patterns
Rangeland Management, Denver, CO. change in central Idaho’s ponderosa pine-Douglas-fir forest. Western
127c. ———, A.R. Tiedemann, J.G. Cook, T.M. Ballard, P.J. Edgerton, M. Journal of Applied Forestry 1:16-18.
Vavra, W.C. Krueger, F.C. Hall, L.D. Bryant, L.L. Irwin, and T. DelCurto. In 146. ———, R. D. Pfister, R. A. Ryker, and J. A. Kittams. 1981. Forest
press. Modification of mixed-conifer forests by ruminant herbivores in habitat types of central Idaho. U.S. Forest Service, General Technical
the Blue Mountains ecological province. U.S. Forest Service, Research Report INT-114.
Paper PNW-RP-527. 147. Taylor, D. L., and W. J. Barmore, Jr. 1980. Post-fire succession of
128. Risenhoover, K. L., and J. A. Bailey. 1985. Foraging ecology of avifauna in coniferous forests of Yellowstone and Grand Teton national
mountain sheep: implications for habitat management. Journal of parks, Wyoming. Pages 130-145 in Workshop Proceedings: management
Wildlife Management 49:797-804. of western forests and grasslands for nongame birds. U.S. Forest
129. Rose, A. B., and K. H. Platt. 1987. Recovery of northern fiordland Service, General Technical Report INT-86.
alpine grasslands after reduction in the deer population. New Zealand 148. Taylor, D. M. 1986. Effects of cattle grazing on passerine birds nesting
Journal of Ecology 10:23-33. in riparian habitat. Journal of Range Management 39:254-258.
130. Rose, C., J. Ohmann, K. Waddell, B. Schreiber, D. Lindley, K. Mellen, 149. ———, and C. D. Littlefield. 1986. Willow flycatcher and yellow
and B. G. Marcot. 2001. Decaying wood in Pacific Northwest forests: warbler response to cattle grazing. American Birds 40:1169-1173.
Concepts and tools for habitat management. Pages 580-623 in D.H. 150. Theberge, J. B. 1976. Bird populations in the Kluane Mountains,
Johnson and T.A. O’Neil, managing directors. Wildlife-habitat southwest Yukon, with special reference to vegetation and fire.
relationships in Oregon and Washington. Oregon State University Canadian Journal of Zoology 54:1346-1356.
Press, Corvallis, OR. 151. Thomas, D. W. 1988. The distribution of bats in different ages of
131. Rosenzweig, M. L. 1985. Some theoretical aspects of habitat Douglas-fir forests. Journal of Wildlife management 52:619-626.
selection. Pages 517-540 in M. L. Cody, editor. Habitat selection in birds. 152. ———, and S. D. West. 1991. Forest age associations of bats in the
Academic Press, New York, NY. southern Washington Cascade and Oregon Coast Ranges. Pages 295-
132. Rowe, J. S., and G. W. Scotter. 973. Fire in the boreal forest. Quarterly 303 in L. F. Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff, technical
Research (NY) 3:444-464. coordinators. Wildlife and vegetation of unmanaged Douglas-fir forests.
U.S. Forest Service, General Technical Report PNW-285.
 238 Wildlife–Habitat Relationships in Oregon and Washington

153. Thomas, J. W., technical editor. 1979. Wildlife habitats in managed 173.Yunick, R. P. 1985. A review of recent irruptions of the black-backed
forests: The Blue Mountains of Oregon and Washington. U.S. Forest woodpecker and three-toed woodpecker in eastern North America.
Service, PNW Region, Agricultural Handbook No. 553. Journal of Field Ornithology 56:138-152.
154. Tiagwad, T. E., C. M. Olson, and R. E. Martin. 1982. Single-year 174. Zimmerman, G. T., and L. F. Neuenschwander. 1984. Livestock grazing
response of breeding bird populations to fire in a curlleaf mountain influences on community structure, fire intensity, and fire frequency
mahogany-big sagebrush community. Pages 101-110 in E. E. Starkey, J. F. within the Douglas-fir/Ninebark habitat type. Journal of Range
Franklin, and J. W. Matthews, editors. Ecological research in national Management 37:104-110.
parks of the Pacific Northwest. Forest Research Laboratory, Oregon
State University, Corvallis, OR.
154a. Tiedemann, A.R., R.R. Mason, and B.E. Wickman. 1998. Forest floor
and soil nutrients five years after urea fertilization in a grand fir forest.
Northwest Science 72:88-95.
155. Tilghman, N. G. 1989. Impacts of white-tailed deer on forest
regeneration in northwestern Pennsylvania. Journal of Wildlife
Management 53:524-533.
156. Turner, M. G., W. H. Romme, R. H. Gardner, and W. W. Hargrove. 1997.
Effects of fire size and pattern on early succession in Yellowstone
National Park. Ecological Monographs 67:411-433.
156a. Turner, A.L., and R. Sallabanks. 1999 Annual Report, Sustainable
Ecosystems Institute, Meridian, ID.
157. U.S. Forest Service. 1992a. Environmental assessment for the First
Timber Sale. U.S. Forest Service, Winema National Forest, Klamath
Falls, OR.
158. ———. 1992b. Antelope salvage planning area: environmental
assessment. U.S. Forest Service, Fremont National Forest, Lakeview,
OR.
159. ———, and U.S. Bureau of Land Management. 1997. Eastside Draft
Environmental Impact Statement, Volume 1. Interior Columbia Basin
Ecosystem Management Project (ICBEMP), Walla Walla, WA.
160.Verner, J., M. L. Morrison, and C. J. Ralph, editors. 1986. Wildlife 2000:
Modeling habitat relationships of terrestrial vertebrates. University of
Wisconsin Press, Madison, WI.
161.Verts, B. J., and L. N. Carraway. 1998. Land mammals of Oregon.
University of California Press, Berkeley, CA.
162. Volland, L. 1985. Plant associations of the central Oregon Pumice
Zone. U.S. Forest Service, Pacific Northwest Region, R6-ECOL-104-
1985, Portland, OR.
163.Vonhof, M. J. 1996. Roost-site preferences of big brown bats
(Eptesicus fuscus) and silver-haired bats (Lasionycteris noctivagans) in the
Pend d’Oreille Valley in southern British Columbia. Pages 62-80 in R. M.
R. Barclay and R. M. Brigham, editors. Bats and forests symposium,
Victoria, British Columbia, Canada. BC Ministry of Forests, Victoria, BC,
Canada.
164. Wakelyn, L. A. 1987. Changing habitat conditions on bighorn sheep
ranges in Colorado. Journal of Wildlife Management 51:904-912.
165. Waldien, D. L. , J.P. Hayes, and E.B. Arnett. 2000. Day-roosts of female
long-eared myotis in western Oregon. Journal of Wildlife Management
64:785-796.
166. Weigand, J. F., R. Haynes, A. R. Tiedemann, R. A. Riggs, and T. M. Quigley.
1993. Ecological and economic impacts of ungulate herbivory on
commercial stand development in forests of eastern Oregon and
Washington. Forest Ecology and Management 61:137-155.
168. Wickman, B. E. 1992. Forest health in the Blue Mountains: the
influence of insects and diseases. U.S. Forest Service, General Technical
Report PNW-295.
169. Wiens, J. A., and J. T. Rotenberry. 1981. Habitat associations and
community structure of birds in shrubsteppe environments. Ecological
Monographs 51:21-41.
170. Wischnofske, M. G., and D. W. Anderson. 1983. The natural role of
fire in the Wenatchee Valley. U.S. Forest Service, Pacific Northwest
Region, Wenatchee National Forest, Wenatchee, WA.
171. Woodward, A., E. G. Schreiner, D. B. Houston, and B. B. Moorhead.
1994. Ungulate-forest relationships in Olympic National Park:
retrospective exclosure studies. Northwest Science 68:97-110.
172. Wright, C. S. 1996. Fire history of the Teanaway River drainage,
Washington. Thesis, University of Washington, Seattle, WA.
 9
Wildlife in Alpine and Subalpine Habitats
Kathy M. Martin

Introduction
The landscape of western North America is defined by elevation ‘islands’ year-round, and leave only to travel to
vast areas of striking mountainous terrain. Since other alpine patches. However, the majority of species
settlement, humans have been fascinated and frustrated move to lower elevation habitats at some life history stage.
with these imposing habitats that offer more vertical than Thus, connectivity is a key ecological process to maintain
horizontal relief. In the old world, humans have used for alpine wildlife. Anthropogenically-induced changes
alpine areas for hunting and agriculture since the at both high and low elevations constitute potential threats
beginning of recorded history. Europeans and Asians have for alpine animals which are well adapted for extreme
practiced alpine agriculture for over 10 centuries, conditions, but not so well for increased warming,
cultivating crops and moving livestock up to alpine competition or predation. Alpine sites have the potential
pastures in summer and down to lower elevations in fall.9 to serve as natural experiments that allow ‘space for time’
In North America, our relationship with alpine areas is perspectives in predicting animal responses to global
much more recent. Mountains posed serious barriers to climate change.
exploration and the development of agriculture in the
previous century. Today, however, alpine areas are valued 1. The Nature, Distribution, and Diversity of
for their intrinsic beauty and wildness, for their Alpine Habitats—Global to Local
recreational potential, and as a refuge from dense urban A mountain is a landmass arising above the general
areas. landscape that induces a change in climate that affects
The alpine zone consists of rugged, partially vegetated vegetation and animal life111 The word alpine comes from
terrain with snowfields and rocky ridges, above the the Alps, and refers to the zone above the natural treeline,
natural treeline. Alpine ecosystems are structurally simple with persistent or permanent snowfields, rocky ridges,
with few plant species compared to most lower elevation occasional wind-shaped trees and continuous to scattered
habitats. High elevation habitats are characterized by high tundra vegetation.92 The treeline, the lower boundary of
winds, prolonged snow cover, steep terrain, extremes of the alpine life zone, is often fragmented over several
heat and cold, and intense ultra-violet radiation. With hundred meters of altitude.81 Several factors define the
increasing elevation, time for breeding decreases and alpine zone, including elevation, aspect and high relief,
environmental stochasticity increases; at the highest but climate is probably the best determinant of where
elevations, hypoxic conditions add additional energetic alpine zones begin.81, 111 Alpine climates are characterized
living costs. These factors result in short, intense breeding by high winds, low temperatures, low effective moisture,
seasons for wildlife, and the need for seasonal movements and short growing seasons.11, 21 Alpine zones increase in
to and from patchy breeding habitats and wintering areas. elevation from north to south and from coastal areas to
Despite being highly valued for their intrinsic beauty interior. Northward across the mid-latitudes of North
and wildness, alpine vertebrates and their high elevation America, treeline decreases about 152 m (500 ft) in
habitats are a neglected area for research and management. elevation per 160 km (100 miles).43 In the Cascades, treeline
Alpine ecosystems recently have experienced large increases in elevation from about 2000 m (6560 ft) on the
increases in amount and kinds of human use, and some west side to 2500 m (8200 ft) on the eastern continental
areas now show significant deterioration. In this chapter, side.5 In this chapter, the alpine zone includes two habitat
I describe the wildlife communities inhabiting the alpine types, alpine treeless and partially vegetated areas at the
zone above natural treeline. I describe what is special about top, and the sub-alpine, which is the zone between closed
high elevation communities, and what is changing. About upper montane forest and the upper limits of small trees
one third of the vertebrate fauna in the Pacific Northwest in open parklands.
is connected to alpine and sub-alpine habitats across one The global alpine landmass comprises about 4 million
or more seasons. Alpine habitats are essentially vertical km2, about 3% of the global landmass,81 an area about half
islands. Some animals, such as white-tailed ptarmigan, the size of the continental United States (~7 million km2,
hoary marmots and mountain goats, remain in their high 2.98 million sq. mi.). About 30% of this alpine landmass is

239
 240 Wildlife–Habitat Relationships in Oregon and Washington

vegetated.81 In Washington, the total area of alpine and several impressive natural disturbance events. The Mt. St.
sub-alpine habitat is about 4.4% of the landbase and in Helens volcano in Washington erupted in 1980, scorching
Oregon, about 0.6%. 76 Further north in the Pacific or leveling 500 km2 (200 mi2) of surrounding forest up to a
Northwest, 17% of British Columbia is classified as alpine distance of 25 km (15 mi).59 Recovery of the mountain flora
(BC Ministry of Forests Database). By global and and fauna after this natural disturbance is of special
continental standards, the mountains in Washington and ecological interest to biologists. Shortly after Mt. St. Helens
Oregon are modest in elevation. Mt. Everest 29,029 ft (8,848 cooled, wildlife species began to recolonize and, recovery
m), on the Tibet-Nepal border is the highest elevation on continues. Species with the most complete recovery tend
earth. Mt. Denali in Alaska 20,320 ft (6194 m) is the highest to be associated with standing dead trees, stream
peak in North America. In the Pacific Northwest, Mt. ecosystems, or living beneath the soil surface.41, 94
Rainier is the highest mountain in Washington at 4,409 ft The Cascades Mountains, the major mountain range
(4392 m) (Figure 1), while Mt. Hood is the highest in in the region, extend from British Columbia south through
Oregon at 11,239 ft (3426 m). Washington and Oregon to California. The Selkirk
The North Cascades Range provides a rugged set of Mountains run from southern British Columbia to
young mountains, geologically complex, and heavily northeastern Washington. The Olympic Mountains lie
glaciated. Mt. Baker, Glacier Peak and Mt. Rainier, high west, and the Okanogan Highlands lie east of the Cascades
ice-covered volcanoes, are the most conspicuous Range. The major mountain ranges in Oregon are the
mountains scattered along the crest of the Range. Below Wallowa Mountains, Blue Mountains, Steens Mountain,
the ice, these “island” mountains support well-developed and the Siskiyous.
alpine plant communities in moist and cool
environments.11, 46 The Cascades Range is distinguished 2. High Elevation Wildlife Habitats in
by being one of the snowiest places on the planet. Mt. Washington and Oregon
Rainier and Mt. Baker have the right combination of winter In Washington and Oregon, the two main high elevation
precipitation and oceanic air currents, as well as steep wildlife habitats classified were Sub-alpine Parklands (#9)
temperature and elevational gradients to generate and Alpine Grasslands and Shrublands (#10).36, 76 These
impressive snowfall. During the winter of 1998-99, Mt. habitats comprise about 2.2% of the Washington and
Baker recorded a world record snowfall of 95.0 ft (28.9 Oregon land base (Table 1), and include a broad diversity
m)(U.S. Dept. of Commerce and the National Oceanic and of alpine habitats and conditions. Coastal or maritime
Atmospheric Administration, Press release, August 3, alpine located on the western side of the mountains differs
1999). dramatically from interior mountain ranges on the eastern
Alpine and sub-alpine habitats are among the most side.11, 55 On the western side, there is more coastal alpine,
undisturbed habitats remaining in the Pacific Northwest. connected to Upper Aspen habitat, montane conifers such
The amount or distribution of high elevation habitat in as Engelmann spruce (Picea engelmannii), and sub-alpine
Washington and Oregon has changed little since pre- fir (Abies lasiocarpa), and coniferous wetland habitat types.
settlement times. Alpine ecosystems, however, are On the eastern side, it is drier, alpine habitats are at higher
sensitive to sustained and heavy use. Olympic and Mt. elevations, and these are connected to lodgepole pine
Rainier National Parks show significant deterioration of (Pinus contorta) and high shrub Steppe habitat types.35
alpine fellfields and sub-alpine meadows caused by Locally, high elevation habitats also differ in climate,
recreational activities, grazing, and air-borne vegetation, and biological processes such as treeline limit
contaminants.15, 69, 78, 114 This region has also experienced and resilience to impacts depending upon location, slope,
elevation, and aspect.

Figure 1. Sub-alpine flowers in the

foreground, open montane forest with hints of
tree islands, and a spectacular view of snow-
capped Mt Rainier in the background. Slide
by Steve Ogle (#17), mid to late August
1999.
 ALPINE/SUBALPINE 241

Sub-alpine Parkland habitats occur below alpine or extreme variation in snow pack between years in high
krummholz (the zone where growth of trees and shrubs elevation habitats can kill and desiccate plants or shorten
is stunted and deformed) and above continuous montane growing seasons. Because seedling establishment is
forest. The parklands are mosaics of patches of herbaceous generally poor, most alpine plants also propagate by clones
or dwarf-shrub vegetation and tree islands or scattered or stolons.81 Trees often invade from wind-dispersed seeds,
trees with 10 to 30% canopy cover. Sub-alpine Parklands but seed dispersal by mountain birds such as Clark’s
occur throughout the high mountain ranges of Washington nutcrackers may determine upper treeline, especially for
and Oregon, extending north into Canada, and south to whitebark pine.87 Tree invasion rates into Sub-alpine
the Sierra Mountains. Mountain hemlock (Tsuga Grasslands are slow compared to sub-alpine tree or shrub
mertensiana) sub-alpine parklands occur along the communities.56, 83 Herbivory and associated trampling
Cascades Crest and in the Olympic Mountains. In the west disturbance by elk and mountain goats creates patches of
Cascades and Olympic Mountains, Sub-alpine Parklands open ground.69 Avalanches and snow-slumping convert
are mosaics of tree patches and heath shrublands or coniferous forest to open meadows or deciduous forests
wetlands. Sub-alpine Parklands in the east Cascades and providing nutritious forage for breeding or migrating
Wallowa Mountains, occurring at slightly higher birds, rodents, bears, and ungulates. 82 Sub-alpine
elevations (up to 2438 m, 8000 ft; Table 1), contain grasslands burn on occasion, but since 80-90% of sub-
Whitebark pine (Pinus albicaulis)-subalpine fir with ground alpine plant biomass is underground, fire does not affect
cover typically dominated by sedges and grasses, and with the structure of sub-alpine grasslands greatly. In Sub-
less heath than Alpine Grasslands and Shrublands. alpine Parklands, fire suppression has contributed to
changes in habitat structure and function. During wet
Alpine Grasslands and Shrublands include all vegetated
climatic cycles, reduced fire frequency can lead to tree
areas above the upper treeline in the highest mountains,
islands coalescing and, parklands becoming a more closed
as well as significant expanses of grassland just below the
forest. 1 Area of alpine grassland, however, may be
upper treeline within the sub-alpine zone. Upper treeline
increased by fires in Sub-alpine Parklands.83 Periodic shifts
is defined as the elevation above which trees are unable
in climatic factors such as drought, and depth or duration
to grow in an upright form. Alpine vegetation is
of snow pack may either lower treeline or allow tree
dominated by sedge species, grasses, hardy forbs and/or
invasions into meadows and shrublands creating more
dwarf shrubs such as heathers. This habitat type includes
parkland habitat.
the krummholz. Alpine Grasslands and Shrublands occur
in the high mountains throughout the Cascades, Olympic Status and trends. There has been little change in
Mountains, Okanogan Highlands, Wallowa Mountains, abundance of alpine habitat over the past 150 years. Most
Blue Mountains, Steens Mountain in southeast Oregon, areas are dominated by native species and are still in good
and occasionally in the Siskiyous. Alpine heath condition.76 Current trends for most alpine grasslands are
communities are found primarily along the Cascade crest considered stable, but there are increasing threats from
and west, especially from Mt. Rainier north. Sub-alpine recreational pressures and livestock grazing, and possibly
and alpine wetland habitat occurs throughout the range, some slow loss of sub-alpine grassland to recent tree
and is more common in the high mountain ranges of invasion. Conditions are changing, however, as over the
Washington.35 past half century, mountain recreational activities have
Most natural disturbances in Alpine Grasslands and increased markedly. Some alpine areas on the eastside are
Sub-alpine Parklands involve weather events or animal degraded physically. Recreational impacts are noticeable
activities, and usually have small-scale impacts. Frost in some national parks and wilderness areas. Mt. Rainier
heaving can have small-scale but important effects on National Park gets two million visitors per year, most of
alpine vegetative communities. 49 Wind blasting and whom converge on Paradise and Sunrise sub-alpine

Table 1. Areas of high elevation habitats, and elevational ranges for Washington and Oregona

Washington Oregon
Acres Sq. mi Km2 % Protectedb Acres Sq. mi Km2 % Protectedb

#9. Sub-alpine Parklandsc 327,442 512 1325 58.2 84,240 132 341 57.5
#10. Alpine Grasslands 1,599,115 2,499 6,471 81.3 291,494 455 1,180 56.0
and Shrublandsd
Total high elevation area 1,926,557 3,010 7,796 77.4 395,734 618 1,601 56.3
Total state land base 43,164,632 67,443 174,678 61,974,831 96,834 250,800
a
Kiilsgaard (1999).
b
Percent of habitat type protected in Category 1 and 2 (Shaughnessy and O’Neil 2000).
c
Elevation west of Cascade Crest (1372-1829 m; 4500-6000 ft), East of Cascade Crest (1524-2438 m; 5000-8000 ft).
d
Elevation: 1524 m (5000 ft) to > 3050 m (10,000 ft).
 242 Wildlife–Habitat Relationships in Oregon and Washington

Table 2. Living and breeding in high elevation environments: constraints,

consequences and wildlife adaptations

A. Environmental Constraints C. Biological adaptations

Cold and extreme temperatures Physiological
High winds Increased visual acuity
Open habitats Large thermal neutral zone for heat and cold
Strong temporal resource gradient (melting snow fields) Biochemical adaptations to hypoxia
Strong spatial resource gradient (food phenology) Molting patterns for cryptic plumage
Aridity Night-time torpor
UVB light Flexible reproductive timing
Hypoxia High tolerance for environmental toxins
Airborne toxins Cold-tolerant embryos, antifreeze solutions
Fragmented habitats Structural
B. Ecological consequences Larger body size
High energetic costs for living and breeding Thicker, warmer pelage/plumage
Cooling and warming adaptations required Appendages modified for wind stabilization
Patchy distribution and strong seasonality of resources Behavioral
Low parasitism levels, good health Cryptic behaviour
Delayed breeding schedules Non-directional vocalizations
Increased reproductive synchrony Social and or sub-nivean roosting
Increased reproductive stochasticity Energy-minimizing behaviors such as gliding and walking uphill
Fewer broods/litters per season D. Life history adaptations
Longer development times Cryptic plumage/pelage
High predation risk on young and adults Extensive molting patterns
Small, low density populations Increased parental care
Need to disperse across unsuitable habitats Monogamous mating system
Strong seasonality in habitat use
Hibernation or dispersal in winter
Increased longevity with elevation
Strong age dependence and senescence
Excellent dispersal/migration abilities
External recruitment for rescue

3. Alpine Environments and Wildlife

Adaptations for High Altitudes
meadows. Sedge turfs are the most resilient to trampling Alpine environments are characterized by low mean
and heaths the least resilient. When alpine heath is opened temperatures, high winds, prolonged snow cover, and
up to bare ground it typically does not revegetate for intense ultra-violet radiation12 (Table 2A). Topography
decades, unless active restoration programs are initiated. varies from steep to gentle, and winter snow pack ranges
Sunrise Meadow in Mt. Rainier National Park has had from several meters in gullies to exposed windswept
success in restoration of the sub-alpine meadows.78, 114 The ridges. Although the vegetation structure is simple,
major human impacts to high alpine grasslands are variation in slope and aspect results in rapid changes in
trampling and associated impacts caused by tent sites on habitat types with only modest changes in elevation, thus
Mt. Rainier National Park.78 Exotic ungulates, such as the generating significant structural complexity to alpine
mountain goats introduced to the Olympic Mountains, can habitats. Steep local gradients in snow cover and soil
have profound impacts on high elevation grasslands.69 moisture often govern the productivity and distribution
Grazing by domestic animals also has negative impacts,19 of alpine plants.11 Precipitation is more important than
but these effects are not well studied in the Pacific temperature for alpine plant growth, as most plant species
Northwest. Sub-alpine Parkland habitat trends are are cold tolerant and can carry on photosynthesis down
believed to be generally stable. Whitebark pine may be to -6oC.12
declining due to effects of blister rust or fire suppression.83 Wildlife living at high elevations must be able to cope
For Alpine Grasslands and Shrublands, only one of 40 with high winds, cold temperatures, and desiccation, since
plant associations listed in the National Vegetation often little precipitation originates from rainfall and it
Classification is considered threatened, and less than 10% drains quickly. Although alpine soils are normally cold,
of the Pacific Northwest Sub-alpine Parkland community daily temperatures on the ground vary over a range of
types are identified as threatened.65 47oC144 (Box 1). Thus, during mid day, overheating can be
a problem for alpine wildlife species. On elevational
gradients, resources can be patchily distributed in narrow
 ALPINE/SUBALPINE 243

Animals respond to living in extreme environments in

1. Getting by on high: several ways. They can migrate to warmer environments,
Avian adaptations to high altitude and hibernate, or stay active all year. Alpine animals that stay
life history adjustments active all year have developed biological and life history
Alpine and arctic birds face similar problems of adaptations to survive in these extreme habitats (Table 2).
living in habitats characterized by low Vertebrate species living at high elevations have developed
temperatures, high winds, short growing seasons impressive physiological, behavioral and morphological
and delayed breeding schedules. At high altitudes, adaptations to conserve energy (Table 2C). The type of
hypoxia and aridity further elevate energetic costs adaptation may increase with elevation, or with increased
for survival and reproduction. Physiological time spent at high elevation. For example, species that use
adaptations and life history traits of white-tailed low elevation alpine, or high alpine for only short periods,
ptarmigan in the Colorado Rocky Mountains may require only behavioral adjustments like moving to
(3600-4400 meters) are compared with arctic more benign or sheltered sites to conserve energy.38, 86
willow ptarmigan (0.5-800 m). Ptarmigan embryos Hummingbirds exploiting the rich resources in sub-alpine
in the alpine must develop at cellular oxygen meadows during migration go into nighttime torpor if
tension levels that are normally lethal for tissue their energy reserves at nightfall are below a critical set
maintenance.32, 33 To deal with hypoxic conditions, point.34 With less vegetative cover for concealment, many
white-tailed ptarmigan embryos have elevated alpine mammals and birds have developed cryptic
levels of blood hematocrit and citrate synthase appearances and behaviors to enhance blending into the
enzyme in muscle tissue at a much earlier stage of landscape. Although ptarmigan are renowned for their
development than willow ptarmigan that show cryptic plumage, larger mammals such as bighorn sheep
similar levels to other avian embryos at low and coyotes blend exceptionally well in these open
elevations. Hematocrit and red blood cell organic landscapes. Often species living in the alpine such as
phosphate levels decline when white-tailed marmots, pika and ptarmigan have territorial and social
ptarmigan clutches are incubated at lower contact calls that are hard to localize, which may reduce
altitudes, indicating that high elevation levels are risk of detection by predators when vocalizing.22 Like arctic
a physiological acclimatization to hypoxia, rather wildlife, animals living at high elevations develop fat
than a genetic adaptation.33, 47 Daily temperatures deposits, extra feathers or thicker fur to increase insulation.
in the alpine range from -2oC to >45oC and, Red foxes in the alpine develop a thick pelage and a faded
females adjust incubation schedules based on nest cryptic gray coat color.
site type and ambient temperature.145 Cooling as True alpine species may develop structural adaptations.
well as generating heat to maintain homeothermy Species living at the highest elevations for extended
is likely a problem for most alpine vertebrates. periods make biochemical adjustments such as increasing
Embryo viability does not differ between the two their blood haematocrit concentrations26, 32, 33 (Box 1). In the
species, but living at high elevation likely imposes Himalayas, alpine passerines tend to have long pointed
significant life history costs. Alpine ptarmigan wings for efficient flight to scattered resources and
have smaller clutches, slower egg laying rates, improved flight performance in strong and variable winds.
longer incubation periods and higher predation Finches and chats at the highest elevations have square-
than arctic ptarmigan.97 Alpine birds show ended or only shallow forked tails for flight stability in
stronger age-dependent effects on survival and wind, and strong hind limbs with small feet for ground
reproduction compared to arctic ptarmigan.145 foraging.84, 85, 86 Winds may pose problems for smaller
animals, but larger animals such as raptors use wind to
remain aloft and move efficiently.
The reduced time for breeding within a season may
bands of diverse habitat types that vary sharply in time alter life history traits. Generally, vertebrate species living
within a season (e.g., plant phenology or insect emergence; at high elevations exhibit lifestyles where it takes longer
Table 2B). Spatial and temporal variation in resources can to achieve independence, there is lower expected annual
extend the availability of food and cover to wildlife, but fecundity and, at least for some species, accompanied by
also requires good mobility and longer migration distances longer life spans and more developed social behaviors.
from patchy breeding habitats to winter areas. Alpine White-tailed ptarmigan, a high alpine grouse, showed
environments also show significant stochasticity in slower laying rates, smaller clutches, longer incubation
environmental conditions annually. Some years have low periods with consequent higher reproductive failure
snow cover leading to an absence of cover for sub-nivean compared to arctic breeding willow ptarmigan97 (L.
species and, in other years, like 1999 in the Pacific lagopus; Box 1). A study of golden-mantled ground
Northwest, much breeding habitat remained snow- squirrels across five elevations in the Sierra Nevada in
covered for the entire season (K. Martin unpubl. data). At California showed reduced time above ground for adults
the highest elevations, hypoxic conditions add an and juveniles, later age of maturity, lower litter size and
additional energy cost.26, 97 These factors result in short, greater survivorship of females with increasing elevation.24
intense breeding seasons for wildlife. Parental care patterns vary with elevation in marmots.
 244 Wildlife–Habitat Relationships in Oregon and Washington

2. Sleeping through winter with family members:

Marmot hibernation and sociality
Marmots are large diurnal, ground dwelling smallest marmot, lives at medium to high
rodents that occupy middle and upper elevation altitudes57 and is moderately social. The basic
alpine and sub-alpine mountains across North social units are polygynous groups usually
America and Eurasia. At the high elevations in developed through recruitment of daughters.
Washington and Oregon, the three marmot species Dispersal of young typically occurs after the first
living in alpine and sub-alpine habitats have non- hibernation.4 European alpine marmots (M.
overlapping ranges. Yellow-bellied marmots occur marmota), living at the highest elevations, are the
in habitats #9 and #10 only in Oregon, and in most social species with up to 20 individuals in a
Washington are replaced in the high elevation group defended by a dominant male and female.8
habitats by hoary marmots, which live only in the Juveniles typically do not disperse for two or more
Cascades. The Olympic marmot is restricted to the years, and their over-winter survival is enhanced
Olympic Peninsula. Marmots are the largest true by increased group size in joint hibernacula.6, 7, 8
mammalian hibernators, with species spending Like birds, the principal life history costs incurred
from 4.5 to 8.5 months per year underground.4 by marmots living in alpine and sub-alpine
Marmots thus have a short active season above environments appear to be time delays. Young
ground, on average 4.8 months, when they must require more (longer) parental care at higher
grow, reproduce, and prepare for hibernation. elevations and thus remain with their parents or
Marmots have complex social systems, delayed extended family for one or more years after they
maturity of young, and flexible patterns of reach maturity.4, 8 Delayed dispersal results in
dispersal and recruitment as a consequence of groups of high relatedness, which in some species
their large body size and living in extreme and leads to reproductive suppression of subordinate
patchy environments.4, 6, 7 The degree of sociality members in the group.4 Reproductive skipping
across the 14 marmot species correlates with between years occurs in hoary and Olympic
environmental harshness and increases with marmots due to extreme conditions, but not in
altitude and latitude.8 The low elevation yellow-bellied marmots.4 Higher longevity is
woodchuck (Marmota monax) is solitary, young of observed in high elevation marmots, and this may
the year disperse from natal burrows, and be due to delayed dispersal, reproductive
individuals usually reproduce after their first skipping, or higher sociality that may reduce
hibernation.51 The yellow-bellied marmot, the predation risk.4

Alpine marmots have prolonged parental care, live and dispersal from and to alpine habitats is more challenging
hibernate in larger groups and take longer to reach for alpine mammals than birds. The frequent speciation
maturity compared to marmot species living at lower that occurred in the high elevation marmots (Olympic,
elevations4, 6, 7 (Box 2). Hoary, Yellow-bellied, and Vancouver Island) and ground
Among mountain ungulates, there is considerable squirrels (Beldings, Columbian, golden-mantled and
variation in patterns of parental investment and time to Cascade golden-mantled) may have been in response to
offspring independence in high and low elevation isolation imposed by dispersal barriers.
populations, but it is not clear what influence alpine
habitats have in explaining these patterns. In order to 3. Advantages to Living and Breeding
maximize maternal survival, mountain goats may reduce in Alpine Habitats
their investment in reproduction with increasing altitude, Why would any animal live at high elevation given the
but species spending more time at high elevations may extreme environmental conditions and the high energetic
prolong the period of maternal care to young.52, 60 Delayed costs to living there? Is it worth it? The answer is that there
primiparity may be the life history trait most likely to vary are definite advantages to living in high elevation
with elevation for ungulates (M. Festa-Bianchet, pers. environments. In winter, despite extensive snow pack,
comm.). One expects other life history tactics such as most alpine areas have wind-swept ridges with exposed
increased probability of bet hedging against the regular herbaceous stems and seeds for foraging, and as winter
but stochastic reproductive failure, 24 but such progresses, increasing snow levels allow herbivores access
demographic parameters are usually not available for to new layers of vegetation. In spring, vast numbers of
species living at high elevation. cold-numbed insects are swept up from lower elevations
Despite the tendency for small population sizes, birds to land on high elevation snowfields. These insects provide
adapted to live in naturally fragmented alpine habitats a nutritious and abundant food supply for alpine-breeding
appear to have well developed dispersal abilities (Box 3). birds and mammals.104,122 Snowfields melting through the
Pikas also are excellent dispersers, 118 but generally season create a gradient in plant phenology that provides
 ALPINE/SUBALPINE 245

3. Patterns and mechanisms of rescue and dispersal in alpine ptarmigan

The periodic “rescue” of populations that are Ptarmigan have adapted to breeding in highly
declining with recruitment from productive fragmented and stochastic alpine habitats with a
populations may be an important feature of system of extensive external recruitment that
population biology for many species. Dispersal functions at a landscape scale. Local populations
and recruitment are crucial life history parameters were able to avoid extinction due to external
for persistence of populations or metapopulations, recruitment, since about 95% of females and 75%
but data on patterns and mechanisms are lacking of males recruiting to sites did not originate from
for most vertebrates. We studied four populations any of the studied populations. Excellent dispersal
of white-tailed ptarmigan breeding in highly abilities allowed recruitment from populations
fragmented alpine habitats, with corresponding during a productive episode to “rescue”
small populations, in the Rocky Mountains of populations at risk of collapse were they
Colorado, from 1987 to 1996. Multiple populations dependent solely on internal recruitment. Studies
allowed us to distinguish between local and of radio-marked juveniles in Colorado revealed a
regional events. Populations showed dramatic spatial scale of about 35 km (22 mi), and
variation in offspring production and local movement patterns of juveniles were not related
survival (return) of adults across years and sites, to locations or movements of their mothers.95 The
yet populations remained relatively stable. enabling factors for this extensive external
Variation in reproductive success and survival recruitment appear to be low costs to dispersal
were unlinked across sites and appeared to be and low benefits to philopatry. Observations and
driven mainly by internal ecological processes experiments on mate and territory replacement
such as depredation of eggs and young.22 Sites showed no apparent reproductive costs to females
varied temporally in predation risk,22 perhaps that switch mates or territories66 (K. Martin
because the generalist predators that prey on unpubl. data), suggesting that dispersal to
ptarmigan live for several years and their hunting unfamiliar sites is not costly. Populations will be
skills likely improve with age. Computer maintained as long as there is a balance of
simulation models of individual populations populations producing recruits in reasonable
predicted that all populations should go extinct in proximity to populations requiring rescue. Rescue
2 to 10 years, assuming no linkage between them. by external recruitment appears to apply to other
Treating the four populations as different grouse such as red grouse (Lagopus lagopus
fragments of a single large closed population scoticus) , sage grouse and capercaillie (Tetrao
resulted in predicted persistence times of about urogallus),109, 125 and perhaps generally to other
nine years.98 Yet all populations have persisted for taxonomic groups. For small populations in
at least 30 years and likely substantially longer. In stochastic environmentals, maintaining
one site, the population remained stable despite connectivity between populations is crucial
zero production the previous year and no return because populations producing recruits must be
of breeding females. within reasonable proximity to those requiring
rescue.95

an extended supply of high quality food for herbivores undetected, although they are clearly successful in doing
that can migrate along the green-up line. In late summer, so. In northern British Columbia, caribou give birth to their
leaf budding, flowering, and fruiting of huckleberry calves at alpine sites because they are considered to be
(Vaccinium deliciosum) or bearberry (Arctostaphylos uva-ursi) refugia from predators, and maybe also from biting
may co-occur in close proximity grading away from the insects.117
edge of a snowfield. Avalanche chutes at high elevation Alpine habitats appear to be healthy environments with
provide lush vegetation adjacent to forest cover for bears, low levels of parasitism or disease, at least for alpine
marmots and songbirds. Availability of spring forage may grouse, which have few blood infections or intestinal
be crucial for breeding in a number of species such as parasites.22, 123 Blood parasite infections in white-tailed
bighorn sheep and mountain goats. 52, 110 Generally, ptarmigan varied with degree of fragmentation in the low
compared to other habitats, there are few intraspecific or elevation coastal alpine on Vancouver Island as 84% of 25
interspecific competitors relative to food supplies. birds sampled on isolated mountains in the north and
Predation risk in the alpine may or may not differ from south had blood parasites compared to only 57% of 53
lower elevation sites, but it appears easier for prey to detect individuals in the more continuous alpine in Strathcona
aerial and mammalian predators in open alpine habitats. Provincial Park (K. Martin and N. J. K. Braun, unpubl.
It is a challenge for medium and large predators such as data). The higher rates on smaller, more isolated
foxes, mountain lion and wolves to approach prey mountains were likely related to ptarmigan mixing more
 246 Wildlife–Habitat Relationships in Oregon and Washington

with blue grouse at these sites. In contrast to birds, 4. Resident Wildlife and Migrant Visitors to
parasites and infectious viruses can be a problem for alpine Alpine and Sub-alpine Habitats
marmots 8 and other mammals. Alpine ungulates, Year-round and summer residents. Only a few North
particularly bighorn sheep, are susceptible to a range of American bird species, white-tailed ptarmigan, American
infectious diseases including pneumonia and blood and pipit, black-crowned rosy finches, gray-crowned rosy
organ parasites, especially when animals are in contact finches, the alpine sub-species of horned lark (Eremophila
with domestic livestock.54, 107, 113 alpestris alpinus), and golden-crowned sparrow breed
Relative to the arctic, where long distance migrations exclusively in alpine and sub-alpine habitats. Of these,
to winter sites are required for most seasonal breeding only ptarmigan are year round residents (Figures 2, 3).
species, winter habitat for alpine species often occurs in Mammals, such as hoary marmots, Olympic marmots,
close proximity to breeding sites. In alpine habitats in the Cascade golden-mantled ground squirrels, mountain
Pacific Northwest, animals may simply descend one or a goats, pikas and water voles, associate primarily with high
few kilometers to habitats that offer greater shelter or more elevation habitats year-round. Pikas can live at low
benign weather. In the low elevation coastal alpine in elevations if appropriate habitat features occur, such as
central Vancouver Island, white-tailed ptarmigan moved talus, but high elevation populations are non-migratory.
around the mountain to a southern or western aspect to Overall, more species of mammals than birds are closely
winter in openings in the montane forest or parkland only associated with high alpine habitats (Table 3). There are
100 to 400 m (300-1000 ft) lower in elevation than their no true alpine amphibians or reptiles in the Pacific
breeding habitat. In the Cascades Range, ptarmigan likely Northwest, and only the Cascade frog is closely associated
make similar movements in winter. with sub-alpine parkland habitats (Table 3).
Despite their reputation for unfavorable thermal Alpine songbirds face special challenges to survive and
conditions, alpine habitats offer thermal advantages for breed in alpine environments. With their small body size
wildlife species in both winter and summer. Winter is they must cope with cold temperatures and high winds
reliably cold, usually remaining below freezing for periods as well as incubate small eggs that cool rapidly in near
of seven to eight months. Thus arousal for hibernating freezing ambient temperatures. All alpine and sub-alpine
animals is likely to occur at appropriate times. There is passerines provide bi-parental care to their altricial young.
generally a reliable supply of snow to provide well- True alpine passerines (birds breeding only at high
insulated and safe sub-nivean habitats. Summers are
reliably cool and breezy and, a number of birds and
mammals escape the late summer heat at lower elevations
by moving up to alpine meadows to forage and to escape
insects. Temperature inversions are common in mountain
landscapes and provide an exception to the general rule
of decreased temperature with increased elevation. A
thermal belt results in warmer temperatures at mid-slope
than either the valley bottom or the upper slopes.111 On
the north slope of Mt. Hood in Oregon, a sharply delimited
thermal belt between the Hood River valley bottom and
the upper slopes allows suitable growing conditions for
such crops as cherries.
In winter, thermal inversions occur in high elevation
areas, and temperatures may remain high for several
weeks. In the Northeastern Asian highlands, temperatures
at timberline from December to February can be 25-30½C
warmer than the low forest conditions of -50½C. Moose
migrate to high elevation habitats during thermal
inversions, possibly to capitalize on the favorable
temperatures, although they may also be attracted to sub-
alpine shrublands (A. Andreev, pers. comm.).
In summary, there may be high ecological costs to living
in open habitats at high elevations as the need to move
for food or cover may result in increased risk of detection
by predators. However, alpine animals likely experience
lower levels of inter-specific competition and less habitat
degradation than wildlife occupying lower elevation
habitat types. Figure 2. White-tailed ptarmigan in winter plumage and in
winter habitat at Guanella Pass, Colorado, with Mount
Evans (breeding habitat) in the background. Photo by K.
Martin, January 1998.
 ALPINE/SUBALPINE 247

dead wind-blown arthropods deposited on snowfields.122

Verbeek 135 emphasized the importance of having
snowfields to provide an insect food supply within
territories of pipits in Wyoming. However, horned larks
and pipits foraged more frequently in snow-free areas than
on snow on Mt. Baker in Snoqualmie National Forest,
although both species did use snowfields in late afternoon
after updrafts from low elevations deposited insects on
snow.104 The proportion of energy budget provided by
chilled insects on snowfields was not determined in any
of these studies.
With such limited vertical structure in the alpine,
passerines need to choose nest sites that are protected as
much as possible from predators and extremes of wind,
Figure 3. Male (left) and female (upper right) white-tailed precipitation, or temperature. Rosy finches and mountain
ptarmigan in cryptic spring breeding plumage feeding on bluebirds nest in inaccessible cliffs or rock crevices, while
alpine territory, Loveland Pass, Colorado. Photo by Robert E. pipits and horned larks nest on open grassy tundra slopes
Bennetts. that are potentially more exposed to predators and climatic
extremes. Johnson74 reported 81% of 139 rosy finch nests
elevations) are hardy and appear to survive well despite were in cliff crevices, and suggested that rodents might
storms and inclement weather. They take shelter from limit rosy finch nesting to cliffs. Nest sites in cliff crevices
wind beside or under rocks, and sit on dark rocks to warm and under rocks are colder thermal environments than
up in cold sunny weather (K. Martin, unpubl. obs.). A open cup nest sites in the alpine and arctic tundra.93, 144
study of alpine finches showed ecological and Water pipits (Anthus spinoletta) in Austria spent three to
morphological differentiation across an elevational four weeks choosing potential nest sites within their
gradient in the Himalayas with the heaviest species territories.14 Nest sites were on steep slopes and oriented
occupying the highest elevations.84, 85, 86 Both sexes of the NW-NE, providing shelter from drifting snow or rain. All
alpine sub-species of horned lark, which lives only in the nests were placed below ground level with a solid nest
Washington Cascades and Olympic Mountains, have roof resembling a nest cavity more than a nest cup and
longer wings and tarsi than low elevation sub-species in providing considerable insulation against the cold alpine
the same region.10 temperatures.14, 136 American pipit nests are sunk into the
Most alpine passerines are both granivorous and ground on steep grassy slopes.99, 135
insectivorous.136 Within alpine sites, species tend to use The breeding schedules of alpine bird species are three
different microhabitats and foraging modes. Rosy finches, to six weeks later than those for related species at low
associated with steep rocky slopes, are slow searchers for elevations. Initiation of egg-laying in white-tailed
insects and seeds, while horned larks and pipits are active ptarmigan populations above 3200 m (10,500 ft) in
searchers in more grassy areas.38 Wheatears (Oenanthe Colorado may begin on 25 May and extend to 11 July
oenanthe) frequent open slopes and plains and pounce from (mean of 8 June; 5 years),22 about six weeks later than
boulder perches to capture ground and low-flying insects. ruffed grouse, a low elevation ground nesting tetraonid,
Food supplies in alpine sites for songbirds are plentiful that commences egg laying by 10 April and extends to 19
generally, but of short duration.38 Receding snowfields in July in British Columbia.28 Some alpine species extend
summer continually uncover new food for rosy finches breeding later into the season to compensate for their late
and pipits.133 Many alpine passerines forage on chilled and start in spring. There was no overlap in laying dates for

Table 3. Vertebrate species use of sub-alpine and alpine habitats in Washington and Oregon1

Mammals Birds Herptiles

A2 B C D A B C D A B C D

Sub-alpine parklands (#9) 11 39 15 65 7 48 47 102 1 7 3 11

Alpine grasslands (#10) 13 22 14 49 5 16 52 73 0 4 1 5
Number of species 65 112 11
1. Associations derived from Expert Panels of the Species Habitat Project and supplemented by atlases from Washington (Smith et al
1997), Oregon (Csuti et al 1997), British Columbia (Campbell et al 1990, 1996), and Colorado (C. E. Braun 1969), as well as
unpublished field data from K. Martin (University of British Columbia—Website: http://www.forestry.ubc.ca/alpine/index.htm), and
field counts from L. Steiner, Washington.
2. A = Closely Associated; B = Generally Associated; C = Present; D = Total.
 248 Wildlife–Habitat Relationships in Oregon and Washington

rosy finches in the Montana Rocky Mountains (egg laying to compensate for reduced number of breeding attempts
16 June to 6 July), over seven weeks later than on Amchitka per season. However, clutch size did not increase with
Island, Alaska (28 April to 26 May).75 The alpine sub- elevation in garden warblers.141 Despite less favorable
species of horned lark at 3200 m in Wyoming laid eggs thermal conditions, both incubation and nestling periods
between 25 June and 10 July, up to three months later than were one day shorter in high elevation garden warblers
low elevation populations.10, 134 compared to lowland populations; this was attributed to
Some mountain goat and bighorn sheep herds remain higher levels of male parental care and thicker nests.141
at high elevation year-round, subsisting in winter on the Nest structures in the high elevation population weighed
windswept alpine grasslands and meadows. Herds consist almost twice as much as those at 200 m and presumably
of small groups of bachelors or female with young. conferred greater insulative warmth. Success of individual
Mountain goat populations may be limited by quality and nesting attempts was good, but the frequency of total
quantity of winter food as suggested by Houston and failure for a season was higher at increased elevation.141
Stevens,69 but recent work indicates that spring weather For most ‘alpine-tolerant’ species, fewer broods per
or timing of access to new plant growth in spring is more season are produced at higher elevations. For example,
important than winter conditions110 (M. Festa-Bianchet, American robins breeding at low elevation in British
pers. comm.). Columbia can produce up to five clutches and three
broods28 (J. N. M. Smith, pers. comm.), but in the sub-alpine
Species breeding across wide elevational gradients.
they may only have time to produce one brood, or possibly
Some birds and mammals range naturally across wide
two clutches. Dippers and horned larks also produce
elevational gradients with their breeding habitats
multiple broods per season at low elevations and single
sometimes extending from coastline into the alpine zone
broods at high elevations.77, 134
(Table 3). Allee and Schmidt2 referred to this group as
Population densities of species that breed across a range
‘alpine-tolerant’. Examples include Canada geese, blue
of elevations are expected to decrease with increasing
grouse, rufous hummingbird, killdeer, dipper, mountain
elevation, but this was not found for garden warblers.142
bluebird, rock wren, and Townsend’s solitaire, and a
Densities of territorial male black redstarts (Phoenicuros
variety of mammals such as long-legged Myotis bat, white-
ochruros) in Austria increased with elevation.79 Densities
tailed jackrabbit, yellow-bellied marmot, Beldings and
of alpine chats also remained high with increasing
Columbian ground squirrels, northern pocket gopher,
elevation in the Himalayas.84 However, these studies had
western jumping mouse, heather vole, long-tailed vole,
small study plots and reported densities might be inflated
fox, coyote, black bear, grizzly bear, wolverine, mountain
as a result of plots being located in optimal habitats. Across
lion, and big horn sheep. At high elevations, Columbian
elevational gradients, individuals inhabiting higher
ground squirrels have lower litter size, lower female body
elevations were not younger or less competitive birds
weight, lower proportion of young females breeding, and
unable to obtain good territories at lower elevations. High
higher adult survival relative to low elevation
elevation populations of garden warblers in Switzerland
populations.45 Some within-species differences in ground
and redstarts in Austria did not contain a higher than
squirrel life history with elevation, such as litter size and
average proportion of first time breeders, and natal
proportion of young females with litters, disappeared
philopatry and breeding site fidelity of birds in the upper
when high elevation populations were food-
elevations was high.79, 142
supplemented.44 They suggested that ground squirrel
Poikilothermic (cold-blooded) animals such as frogs,
populations showed a phenotypically plastic life history
salamanders, and snakes have special problems moving
response to variation in food availability with elevation.
and developing eggs in cold alpine climates as low
Pika range from high alpine to sea level where there are
temperatures result in prolonged development times for
grasses and forbs close to talus. Pika show similar life
amphibians and reptiles. In alpine habitats, frogs
history patterns to Columbian ground squirrels as high
commonly pass the first winter as tadpoles and require
alpine populations have smaller litters and greater
an extra year to achieve full development,2 in contrast to
longevity of adults than those in low elevation talus.119, 120
lower elevation individuals that usually achieve full
There are few directed studies for ‘alpine tolerant’ birds
maturity in their first year. In the Alps, the three species
on whether and how life history or behavior shifts with
of reptiles, green lizard (Lacerta vivipara:), common viper
increasing elevation. However, we can make some
(Vipera berus) and the blindworm (Angius fragilis) that reach
inferences. Breeding schedules such as arrival dates and
the alpine zone are all viviparous (bear live young).2
initiation of breeding are significantly later with increases
Reptile eggs cannot develop and hatch in such cold
in elevation. Garden warblers (Sylvia borin) breeding in
climates, but when viviparous female snakes or lizards at
Switzerland at 4,921 ft (1500 m) arrived on territories about
high elevations retain eggs in their body, they can speed
three weeks later than birds breeding at 656 ft (200 m).142
embryonic development by basking in the sun during the
The high elevation population of garden warblers did not
day, and moving to sheltered locations at night. 111
extend their egg laying period beyond lower elevation
Interestingly, the green lizard is oviparous (egg laying)
populations and thus, had one or two fewer broods in a
throughout most of its lower elevational range in Europe.
season than low elevation pairs.141 If restricted gene flow
Snakes in the European Alps show delayed sexual
results in local adaptation, clutch size or offspring survival
maturity and reproduce only every two to four years in
is predicted to be higher in high elevation environments
 ALPINE/SUBALPINE 249

the alpine, rather than annually as in low elevation Table 4. Characteristics of alpine and sub-alpine
populations.31, 116 Cold-blooded animals living in mountain habitats in late summer for migrants and residents
environments, including amphibians, reptiles and
invertebrates are almost universally dark-colored. 1. Rich food resource gradients—plants, insects and small prey
Melanism apparently contributes to heat absorption and 2. Open habitats to detect predators
helps protect against ultraviolet radiation. 2, 111 For 3. Energetically more favorable for ascent (updrafts)
hibernation, cold-blooded animals move under rocks or 4. Habitat features similar to high latitude environments
into animal burrows. They tend to aggregate in multi-
species groups (insects, snakes, and lizards), and one can
migrants of local origin. Some forest birds, such as
find predators and prey congregated under one rock for
nuthatches and pine siskins, certainly moved up the
winter hibernation.111
mountain to track emerging food resources (insects/
In Washington and Oregon, five species of amphibians
flower/ fruit) from snow melt and green-up. For North
and reptiles occur in alpine grasslands and 11 species of
America, over 200 species of birds were recorded using
salamanders, frogs, toads, and garter snakes live in sub-
alpine and sub-alpine habitats in late summer (literature
alpine parklands if appropriate habitat features such as
and field surveys, web site: http://www.forestry.ubc.ca/
standing water are available. Western toads and Pacific
alpine/index.htm).
Treefrogs breed in high elevation ponds.39 Western toads
There are a number of reasons why alpine habitats are
occur at elevations up to 7,400 ft (2255 m)39 in Washington,
suitable for fall migrants (Table 4). In late summer, food
and in Colorado to 12,000 ft (3658 m, C. E. Braun, pers.
resources have declined at lower elevations whereas in
comm.). Garter snakes occur in the alpine and sub-alpine
the alpine, late July to early September is the peak period
on southern and central Vancouver Island at elevations
of flowering and invertebrate emergence.81 Migrants from
ranging from 5,056 to 5,904 ft (1,540 to 1,800 m, K. Martin,
the north use alpine and sub-alpine habitats as refueling
unpubl. data). Little is known about the reproductive
points. The proximity of alpine habitats to forest,
success and specific adaptations to life in alpine and sub-
grasslands, shrub steppe or coastal habitats at lower
alpine habitats of amphibians and reptiles in the Pacific
elevations means that animals can move between these
Northwest.
habitats in a short time. In late summer, few robins are
In sum, the ecology, behavior, and life history of the
observed on low elevation coastal breeding areas, a time
upper elevational ranges for ‘alpine-tolerant’ vertebrates
when large numbers are observed in nearby alpine and
is poorly understood. Quite possibly there is more
sub-alpine habitats, as well as in lowland berry-rich
restricted gene flow and greater local adaptation than we
wetlands (J. N. M. Smith, pers. comm.). Rufous
realize. Other species, in addition to horned lark, marmots
hummingbirds move up to sub-alpine meadows and
and ground squirrels, may have well differentiated high
parklands in late summer and defend territories around
elevation sub-species or eco-types yet to be discovered.
flower patches.128 Territorial boundaries of migrant
We cannot calculate relative reproductive success and
hummingbirds shift daily in relation to temporal patterns
survival for wildlife species living at high and low
of flowering plants such as paintbrush (Castilleja spp.) or
elevations, mainly because longitudinal studies of
columbine (Aquilegia spp.).61, 62 From mid August to early
individuals at high elevation sites have not been done.
September, alpine grasslands support large numbers of
Seasonal migration of wildlife to high elevation habitats. grasshoppers that are eaten by kestrels, mountain
The seasonal migration of vertebrates to alpine and sub- bluebirds and Townsend’s solitaire (S. Ogle, pers. obs.).
alpine habitats has been noted, 108, 111 and naturalists Yellow-rumped warblers, common passerine migrants in
acknowledge the use of alpine habitats by migrating the alpine, fly-catch from cliff edges in late September. The
birds. 30 Mammalian wildlife also move up to high increasing abundance of prey attracts predators. Migrating
elevations seasonally. However, the importance of the raptors use the open mountain terrain to hunt for land
phenomenon is not widely acknowledged and, the birds and small mammals, often using updrafts along cliff
ecological processes involved have not been determined. faces and ridges. The duration that individual altitudinal
At Chinook Pass and Lake Valhalla, north of Stevens Pass migrants spend in the alpine is unknown, but temporal
in the Cascades Mountains, Washington, 32 bird species patterns are most likely related to resource availability and
were observed during 8 surveys in July, September and may vary with species. Some altitudinal migrants could
October (Les Steiner, unpubl. data). During field studies move to alpine and sub-alpine areas on a daily basis and
in central and southern British Columbia, over 113 bird spend their nights at lower elevations. Band-tailed pigeon
species were recorded foraging or resting in high elevation could be one such species where availability of late
habitats over a three-month period (Box 4). About 80% of summer fruits in the sub-alpine, such as huckleberry,
bird species observed in late summer do not breed at high crowberry (Empetrum nigrum), and bearberry, may help
elevation. Some species were clearly migrants from higher compensate for the loss of traditional fruit and nutbearing
latitudes using alpine sites as migration corridors, shrubs due to habitat loss at lower elevations.
including arctic shorebirds like Baird’s sandpipers and In summary, about one-third of the vertebrate fauna in
greater yellowlegs, and northern raptors such as sharp- Washington and Oregon use high elevation habitats at
shinned hawks and northern goshawks. We suspect a high some period in their life history. The chief period of
proportion of birds arriving in the alpine were elevational occupation for many species is between late July and late
 250 Wildlife–Habitat Relationships in Oregon and Washington

4. Elevational migration: use of high elevation habitats in late summer

Studies in British Columbia suggest that alpine alpine habitat, 69 km (43 mi) in sub-alpine, and 52
areas support a high diversity and abundance of km (32.3 mi) in montane parkland (approximately
birds and mammals during late summer and fall. 270 survey hours on 62 days, years pooled).
Since 1996, information was collected on temporal Species richness was similar in all three habitats in
and spatial patterns of use of high elevation 1998, a warm and relatively dry summer (Figure
habitats by wildlife, especially migrating birds, on 4). Species richness was much reduced in 1999, a
Vancouver Island and mainland British Columbia cold, wet summer with greatly delayed snowmelt
(K. Martin, University of British Columbia and and early onset of winter conditions. Except
Environment Canada). In 1996 and 1997, we during peak migration (5-19 September),
collected data opportunistically on species use proportionately fewer species used alpine habitats
while censusing ptarmigan on 25 mountains than either the sub-alpine or montane in 1999
across Vancouver Island, including relatively (Figure 4). In 1999, one interior site that was snow-
contiguous alpine in the central Island and more free by late June had the greatest number of
fragmented alpine habitats to the north and south. species (9 more species than in 1998). One coastal
During this time, we observed 53 avian species site, where the snow-line extended down to 1100
with peak migration in these coastal alpine and m (3610 ft.) in late August, had only 26
sub-alpine habitats starting in late August and detections/km surveyed in 1999, compared to 93
remaining high through September. Only 11 detections/km in 1998. Overall, during a 10-week
(21%) of these species regularly breed in high period in late summer, 113 species from a diverse
elevation habitats.28, 130 array of avian families used coastal and interior
In 1998 and 1999, we surveyed alpine, sub- high elevation habitats on Vancouver Island and
alpine and upper montane parkland habitats in the central and southern mainland British
mainland British Columbia ranging from 58o N Columbia. Data summaries are available on
latitude to the Washington border. We recorded a Centre for Alpine Studies web site: http://
total of 107 species and 8,969 detections, using line www.forestry.ubc.ca/alpine/index.htm.
transect sampling, over a total of 82 km (51 mi) in
Number of species

100
90
Figure 4. Avian use of high elevation habitats during intervals. Since many species were observed in more
migration in south and central British Columbia. Ten than one of alpine, sub-alpine, or montane parkland
mountains were surveyed in 1998 (left), and five sites habitats, lines represent the total species counted
were re-censused in 1999 (right). Bars represent total during each period.
numbers of species observed by habitat type for 10-day
 ALPINE/SUBALPINE 251

October (Figure 4). Thus, the biodiversity of high elevation wildlife are not well monitored in Washington and
ecosystems is often greatly underestimated, and, clearly, Oregon.
we need to consider life history periods outside the
breeding season. Local Impacts
Wildlife conflicts in winter—downhill skiing and other
Winter residents. Few wildlife species winter in alpine alpine recreation activities. Winter may be a period of
areas, and fewer still remain active in winter. Some birds, great sensitivity, with human disturbances possibly
including white-tailed ptarmigan, raven, and rosy finch, causing the most adverse impacts on wildlife species. At
remain at the highest elevations during winter, and a few this time, grouse and ungulates occur in social groups or
arctic or northern species like snowy owls and snow flocks and live at about the same elevational range as
buntings migrate south to use alpine areas in the Pacific downhill ski facilities. On Vancouver Island, British
Northwest. Rosy finches forage for exposed seeds on Columbia, for example, white-tailed ptarmigan move from
windswept alpine ridges.58 Gray jays, raven, Clark’s an average summer elevation of 5,500 ft (1,676 m) down
nutcracker, and hardy songbirds like golden-crowned to treeline at 4,500 ft (1,372 m) in winter (K. Martin, unpubl.
kinglets and dark-eyed juncos use tree islands in sub- data), while blue grouse move up to treeline.121 Winter
alpine parklands in winter. A number of mammals make ptarmigan flocks can include individuals from a distance
use of alpine and sub-alpine habitats in winter, including of 14 mi (23 km).67 Thus activities that impact flocks
pika, white-tailed jackrabbits, mountain caribou, and adversely may be observed over considerable distances.
mountain goats, and their predators, foxes, coyotes, pine Ski resort developments impose both on- and off-site
marten, wolverine, lynx, bobcat, and mountain lion, hunt impacts. Tourist activities in winter cause serious
along or above treeline. Mammals develop thick insulated disturbance to black grouse (Tetrao tetrix), now found
fur coats and are adept at finding shelter from alpine winds mainly in mountain forests due to extensive habitat loss
and winter temperatures. Three Myotis bats hibernate in at low elevations in central Europe.126 The construction of
sub-alpine winter caves or mines in Washington and ski stations in the Alps between 1950 and 1980 removed
Oregon. Most species use snow roosts or snow burrows or reduced the quality of many black grouse winter
for sleeping or during storms. White-tailed jackrabbits habitats in France and Germany. Ski stations also cause
winter as individuals at the interface of wind swept ridges major disturbances in winter feeding behavior, winter
and the krummholz.18 For marmots and ground squirrels range and disturbance on leks, resulting in dramatic
that hibernate overwinter at high elevations, choice of population declines.100, 148 Downhill and back country
burrow and amount of snow cover can be crucial to skiers searching for deep powder snow cause additional
overwinter survival.4, 7, 8 Pika, weasels, and voles remain disturbance as these sites are also sought by grouse for
active in the sub-nivean layer where they have temporary snow burrows to escape the cold and predators (A. Zeitler,
reprieve from life-threatening predators and winter pers. comm.). The use of explosives to precipitate
storms. controlled avalanches causes ptarmigan to flush from
snow burrows on ski areas in the Alps and Pyrenees.100
5. Threats/Status for Wildlife Species Mountain goats are particularly sensitive to disturbance,
Most local and regional scale threats to wildlife species in as they do not habituate to alpine recreational activities
alpine and sub-alpine habitats relate to human activities such as helicopter hiking or skiing.40
(Table 5). These include on and off-site impacts due to ski Adverse effects from ski hill facilities can occur year-
resorts, hiking, mountain biking, climbing, fishing, group round. After a road and chairlifts were constructed in 1960-
camping, off-road use of all-terrain vehicles, and 1961 at Cairngorm Mountain in the Scottish highlands,
helicopter-supported skiing, hiking, fishing and sight- allowing easier human access, numbers of crows and gulls
seeing activities. In Mt. Rainier National Park, 12% of on the mountain increased, largely sustained by food
plants (107 of 894 species) and 14% of birds (21 of 147) are scraps,137, 139 but they also preyed on ptarmigan (L. mutus)
exotic species.53 Other potential threats to ecological eggs and dotterel (Charadrius morinellus).102, 131 Density and
conditions in alpine areas include livestock grazing, breeding success of red grouse (L. l. scoticus) and
mining, montane forestry, ground source contaminants, ptarmigan on Cairngorm Mountain did not change in
and barriers imposed by roads and trails.19, 88 Except in a relation to skier activities, but ski lift cables caused
few parks, anthropogenically induced impacts on alpine significant mortality for both species when birds flew into
them, and harassment by free-running dogs was
Table 5. Land uses and possible threats in alpine and reported.138 In France, 91 dangerous ski cables were
sub-alpine habitats identified, and measures proposed to improve visibility
of these cables.106 Mortality of grouse due to ski area
1. Ski area developments, on-site and off-site impacts activities is almost certainly additive since it occurs in late
2. Hiking, fishing, and hunting, especially with helicopter access winter and early spring, after the period when highest
3. Mountain biking, rock climbing, para-gliding overwinter mortality in grouse populations was recorded
4. Livestock grazing, especially sheep grazing in Norway (G.E. Frilund and H. Chr. Pedersen, pers.
5. Mining, montane forestry comm.) and in Iceland (O.K. Nielsen, pers. comm.).
6. Airborne toxins, ground source contaminants
 252 Wildlife–Habitat Relationships in Oregon and Washington

Wildlife conflicts in summer. Alpine species may be Regional Impacts

heavily impacted directly by recreational pressures during Air and water quality. Alpine areas are considered pristine
the breeding season. Recreational activities like rock habitats where the air is pure and water is clean and both
climbing or hang gliding can disturb cliff-nesting raptors are continually renewed by wind and snow melt.
or ungulates with young. No differences were observed However, emerging information suggests that alpine sites
in reproductive success of black grouse inside and outside are not as pristine as we think. Mt. Rainier National Park
areas visited by tourists in the French Alps.101 However, in western Washington consistently had the highest
black grouse use of habitat close to a heavily frequented average weekly concentrations of ozone.15 Although a
path in a French National Park was reduced by 40 to 50%, natural component of the earth’s trophosphere, in high
perhaps due to the presence of wandering dogs.68 Alpine concentrations ozone can injure vascular plants and be
marmots in the Swiss and French Alps adjust well to potentially hazardous to human health. 90 Ozone
human presence once they become habituated to large concentrations are higher at high elevations, which is
numbers of hikers.91, 103 However, chamois (Rupicapra partly a natural atmospheric phenomenon, but also partly
rupicapra) and mouflon (Ovis orientalis) that are hunted in due to upwind transport of pollutants from lower
the Alps and Pyrenees retreat to more inaccessible alpine elevation urban areas. Thus, alpine plants and animals in
areas when they encounter humans, even when there are Washington are exposed to high ozone concentrations on
high numbers of hikers. Mouflon came down to forage warm and sunny days.15 Blais et al.13 found high levels of
on alpine grasslands when there were few hikers or when organochlorines in mountain parks in western Canada.
fog reduced visibility, but they quickly retreated upslope Airborne toxins may influence snow melt patterns in
when disturbed (J.-L. Martin and K. Martin, pers. obs.). alpine sites and thus change environmental conditions for
Thus, alpine ungulates are less likely to habituate to wildlife species as well as quality of the water supply for
disturbance from tourists where they are hunted than nearby urban sites.
where they are protected, and the impact of disturbance Alpine habitats are similar to arctic ecosystems because
will be greatest when high alpine refuges offer only low both have shallow soils that are impervious much of the
quality forage. year and have low levels of biological activity (e.g., soil
Visitors feeding wildlife in parks and recreation areas bacteria);37, 81 thus neither are effective at filtering toxic
can cause local negative impacts. Wildlife feeding can materials. They differ in that the alpine atmosphere is less
change habitat use patterns and inappropriate food can resistant to airborne transport of particulate matter. Air
be injurious to health of the animals. Wildlife feeding may and water quality in many mountain wilderness sites in
result in naivete, particularly of young animals. Mountain western North America are also relatively more
goats and bighorn sheep have been killed by cars on compromised than arctic systems, given their proximity
mountain roads because they were unwary and associated to large urban centers such as Seattle, Vancouver, and
cars with food (K. Martin, pers. obs.), or because they were Denver.15
attracted to road salt (M. Festa-Bianchet, pers. comm.). Ground source contaminants are also a potential risk
Densities of such generalist predators as chipmunks, to wildlife at high elevations (Table 5). In the metal-rish
corvids or canids, or herbivore species such as marmots ore belt of the Colorado Rocky Mountains, high
or golden-mantled ground squirrels may increase due to concentrations of cadmium, an extremely toxic but
accidental or intentional feeding of wildlife. This might naturally occurring heavy metal, were found in white-
change predation risk regimes for alpine species due to a tailed ptarmigan tissue and also in their foods.88 High
direct increase in predator abundance or the amount of cadmium concentrations were associated with kidney
time predators spend in the alpine.138 damage, particularly for overwintering female ptarmigan.
Grazing by domestic species in alpine sites, particularly This study also reported a possible reduction in ptarmigan
by sheep, reduces habitat cover for predator avoidance survival and lower population densities in the ore belt
and may reduce forage quality for native herbivores.21 If regions. Since willow was shown to biomagnify cadmium,
areas are heavily grazed and trampled, species other herbivores such as elk and deer living in these alpine
composition can change dramatically, with many regions could also be at risk of cadmium toxicity.88 Effects
herbaceous plants being extirpated locally. Large areas of of changes in the quality of air, water, and soils on alpine
the Okanogan Highlands in the Paysaten Wilderness have wildlife merit further study.
been heavily grazed by thousands of domestic sheep since
The increasing value of high elevation habitats. Alpine
the late 1800s. Recently, the U.S. Forest Service reduced
and sub-alpine areas have increased in value as wildlife
the number of sheep on this area to a few hundred. Studies
habitat, given changes in habitats or processes at low
on long-term effects of sheep grazing in alpine and sub-
elevations. Wildlife species that formerly used habitats
alpine grasslands are needed for this region. However,
over a broad range of elevations are becoming restricted
other studies have suggested that domestic livestock have
to the upper elevations of their former range. The problem
a greater impact on native ungulates through transmission
of upward shifting is well advanced in central Europe.
of exotic diseases rather than by grazing competition.54,
107, 113
Here most forest grouse species that originally occupied
a range of habitats from low elevation bogs, heaths, and
moorlands up to the sub-alpine treeline have been
 ALPINE/SUBALPINE 253

extirpated from the low elevations in the past century, due Ultimate factors—climate change and global
to intensive agricultural and forestry activities. Black warming
grouse in central Europe are now ‘de facto’ sub-alpine Climate change has strong impacts on wildlife species at
parkland species due to the current unsuitability of high elevations.27, 70, 73 Increased concentrations of CO2
traditional low elevation habitats. 126 The relative available for photosynthesis and changes in temperature
productivity for wildlife species in high and low elevation and moisture availability increase growth rates,
habitats was not established previous to extirpation at low abundance and elevational range of many alpine plants
elevation. However, high elevation habitats regularly at higher elevations.81 In Austria, surveys of alpine plants
experience total reproductive failure and many species are growing 15-20 m from the peak in 1992 and 1993 found 26
endangered.126 Formerly, high elevation populations may (87%) of 30 mountains had higher species richness than
have been rescued in years of low production by low surveys done in identical sites 40 to 100 years earlier.64, 112
elevation populations, but this option is less and less Upslope movement rates for alpine plants for these
possible. Predation risk might be increasing at high mountains were calculated as 3.2 to 13 ft (1 to 4 m) over a
elevations if predators have increased food availability as 10-year period.64 In the context of global warming, the
a result of human presence. In Europe, wolves and brown small-scale patterns of environmental conditions in the
bears are largely restricted to high latitudes and mountain alpine zone, allow ‘space for time’ studies, because
habitats. 129 For large carnivores, the retreat to high vegetation has had no time to adjust to these temperature
elevation habitat patches results in species distributions gradients, and thus is likely to produce a more realistic
being more fragmented than formerly.23 Eurasian lynx picture of longterm responses to changing climates.81
(Lynx lynx) in Switzerland may travel over 100 km to an Research on wildlife responses to climate change is
adjacent mountain group to circumvent barriers imposed beginning to emerge.70 With climate change, upper limits
by highways and densely settled areas.23 of plant growth will increase, and food availability per
On first consideration, it might appear that we need unit area for alpine herbivores may increase. However,
not worry about mean elevational increases for species in with the increasing altitude of treeline, another
the Pacific Northwest. However, in western Washington consequence of climate change, alpine habitats will
and Oregon, low elevation coniferous forests have been become more fragmented, with smaller and more isolated
converted to intensive agricultural, industrial, or high patches. Animals living in these patches will have smaller
density urban areas, and many low elevation wetlands populations and be required to disperse longer distances
have disappeared. 48, 50 Hence, wildlife species that to other alpine patches, or pay the consequences of not
traditionally occupied a broad elevational range of habitats dispersing.95, 115
here have also experienced significant range reduction, Alpine animals are well adapted for living in cold
accompanied with an ‘upward shift’, including such climates, but most have not developed sophisticated
species as white-tailed jackrabbit, western toad, other mechanisms for coping with warm temperatures. 119
amphibians, blue grouse, ruffed grouse, wolf, and other Daytime ground temperatures in the Colorado alpine
large predators. For example, blue grouse numbers have regularly exceeded 450C, and thus cooling can be a concern
declined in many low elevation areas of Washington and for alpine birds in summer. 144 Alpine animals cool
Oregon, while high elevation populations remain themselves by going underground or by moving to
unchanged (M.A. Schroeder, Washington Department of snowfields. In the low elevation alpine of Washington and
Fish and Wildlife, pers. comm.). In the case of grizzly bear, Oregon, an increase of 1 to 20C in mean temperature could
the ‘upward shift’ in population distribution is also have a large impact on the extent of snow pack and
facilitated by greater conflicts with humans at low
elevations, in addition to low elevation habitat loss.
For wildlife species living at higher elevations, we need
to determine which species have experienced the largest
elevational shifts and whether any appear in difficulty.
One problem relates to potential increased energetic costs
of living at higher elevations as discussed in Section 4, as
well as the impact on population viability of more frequent
and greater stochasticity in annual production.141 We
should establish whether wildlife populations at low
elevation rescue high elevation populations following
years of reproductive failure at high elevations. Risks to
wildlife population viability in spatially separated high
elevation ‘islands’ that result from removal of low
elevation populations will increase due to habitat
fragmentation and reduced effective population size. Figure 5. Cattle Creek in Stein Park shows range of habitat
types from riparian valley to mountaintop with treeline going
up much higher on north side (left) than on the drier south
side. Photo by Steve Ogle, August 1998.
 254 Wildlife–Habitat Relationships in Oregon and Washington

snowmelt patterns. Faster melting snowfields might not importance of maintaining connectivity across time and
persist through summer (Figure 5). Fewer or smaller space for species with seasonal altitudinal shifts. For
snowfields leave animals with restricted habitat ungulates and large carnivores, we need to ensure that
availability and longer dispersal distances to other alpine species have access to valley lowlands and other
habitats. With less area to search, predators may be more important wintering habitats. In national parks and
successful in capturing prey. wilderness sites, more research and application is required
In response to increasing temperatures in spring, pied on restoration of impacted alpine grasslands and sub-
flycatchers (Ficedula hypoleuca) increased their egg size over alpine meadows, particularly in the North Cascades
a 19-year period in the mountains of northern Finland; Range.114 Finally, to detect and predict the ecological effects
larger eggs had higher hatching success, but there was no of global climate change, it will be crucial to establish long-
improvement in numbers of young raised.71, 72 Increasing term alpine habitat and wildlife monitoring programs
spring temperatures over a 28-year period were also similar to programs in the Austrian Alps,64 or in northern
correlated with earlier initiation of egg laying and larger Finland,71, 72 or in the Rocky Mountains.70
clutches.71 However, duration of abundant food resources
Migration corridors in the alpine: the need to consider
in late summer will decrease if plant phenology gradients
habitat connectivity. Alpine habitats function as seasonal
are reduced by faster snowmelt. Global warming is thus
migration and dispersal corridors for wildlife. High
expected to have large consequences for high elevation
latitude birds use alpine and sub-alpine habitats as
wildlife species, including reduced habitat availability and
migration highways to move from northern breeding
suitability, reduced duration of food abundance, but
territories to southern winter areas (Box 4). Nocturnal and
perhaps greater per unit area of food availability and
diurnal migrants may use mountain ridges for
possibly increased predation risk. Unfortunately, empirical
navigation. 25 The continued loss of low elevation
data for alpine plants and climate change have been
migration habitat such as riparian and coastal areas
ambiguous with responses by plant species being quite
strengthens the need to understand and manage for such
specific.81 Overall responses will be difficult to predict until
ecological functions as alpine migration corridors.
the key ecological or physical factors are determined. For
Increasing use of alpine areas is expected by large area-
example, temperature may be less important than
sensitive animals, particularly predators, where
moisture in determining the distribution of mountain flora
landscapes have been fragmented.
and fauna.147 Long-term monitoring programs for alpine
The issue of connectivity between high and low
and sub-alpine wildlife will have high efficacy, as even
elevation habitats is an important biological and
small increments in warming have significant impacts on
management question. Parks in the Rocky Mountains have
habitat quantity and quality for breeding and migration.
a disproportionate amount of their habitat at high
6. Information Needs and Conservation elevation, and most parks do not contain year-round
habitat requirements for a number of wildlife species.140
Recommendations
For example, some elk are year-round residents in Banff
Need to develop concerns for alpine conservation and
National Park, but others are migratory; some migratory
management. Probably the largest concern about alpine
elk use up to six different management jurisdictions in a
habitats is the lack of concern about high elevation
single year, including both hunted and non-hunting zones
conservation. The idea prevails that much alpine area is
in two provinces.146 For alpine residents, we need to ensure
already protected, maybe too much, and that most alpine
that connectivity is maintained from alpine habitats to
areas are inaccessible and thus not at great risk. Alpine
adjacent lower elevation forests and valley bottoms (Figure
areas are given lower priority than low elevation habitats
5). In mountain habitats, landscape context can be critical
because they have modest diversity of breeding species.
as access to large areas may contain bottlenecks.105 Urban
Such attitudes account for a lack of concern and
areas and roads in valley bottoms can sub-divide
knowledge regarding status and trends of alpine wildlife
populations or restrict access to seasonally used habitats.
populations, in particular for non-game species. Most
Mountain passes are also likely important for seasonal
research for alpine fauna has been done in summer and,
access or population connectivity, and these tend to be
thus our knowledge of seasonal use of alpine areas is
where roads, trails and recreational huts are situated.
limited. We are not in a good position to detect early
The most severe wildlife-human conflicts occur in
population declines, especially if ecological processes such
mountain landscapes with narrow valleys that are densely
as predation are shifting at large cross-habitat scales.
settled or developed. Unfortunately, mammalian wildlife
Wildlife research is needed to determine the important
in western North America often require access to valley
ecological processes and habitat functions required by both
bottoms to move between mountain chains, since alpine
‘alpine obligate’ and ‘alpine tolerant’ vertebrate species.
areas tend to be closed off by cliffs. Thus the presence of
Research is also needed to improve predictions on how
urban or highway development is almost guaranteed to
wildlife species will cope with changing climatic and
interrupt wildlife movements, act as a dispersal barrier
environmental conditions. Life history studies are needed
and fragment populations of large ungulates and
for high elevation populations of ‘alpine tolerant’ species
predators. All U.S. Interstate highways in mountains
that are experiencing extensive habitat loss at low
impede movements of large mammals (C.E. Braun, pers.
elevations. Better understanding is required about the
comm.). Also, this is a particularly important problem in
 ALPINE/SUBALPINE 255

large mountain parks such as Banff National Park where garbage and educate people to avoid feeding wildlife in
urban developments and major highways in the narrow and adjacent to alpine areas to ensure natural population
Bow Valley have greatly impeded movements of large densities of generalist predators.
mammals and fragmented their populations.105 Species
Coexistence of alpine wildlife and winter recreation
vary in their willingness to cross such dispersal barriers
activities. Ski areas and adjacent resorts have negative
and their wariness when doing so.140 Determining the
ecological impacts on alpine habitats such as soil erosion
ecological value of the alpine as migration habitat and
from sub-alpine forest cutting, which can in turn impact
minimizing interruptions of wildlife movements in
water quality. Wildlife may be displaced or their
mountains are two research priorities.
movements restricted due to cleared areas for ski runs or
Changes in ecological parameters at high elevations. building facilities. Snow making for fall skiing occurs in
Increases in predator density or the amount of time many North American ski operations, and can draw down
predators spend in the alpine could significantly increase water tables in the drier interior mountain conditions.89
the predation risk for alpine and sub-alpine species, as Little is known about impacts on alpine plants and animals
well as for ‘upward shifted’ alpine tolerant species. of additives (ice-nucleating bacteria, fertilizer) to water to
Medium and large-sized predators such as grizzly, wolf improve snowmaking efficiency.89 Before approval of
or lynx that do not persist easily in high-density urban skiing and other recreational developments in alpine and
areas have probably moved higher. For them, ungulates montane habitats, careful surveys should be done in
provide a reliable food resource through the winter period. winter to determine which areas to avoid to minimize
Forestry activities at high elevation may also result in wildlife conflicts. Patterns of winter habitat use by high
changes to wildlife. Research is needed on the influence elevation wildlife species (e.g., blue grouse, ptarmigan,
of montane forest cutting in the Pacific Northwest on elk, bighorn sheep) should be determined before facilities
distribution and abundance of generalist predators, and are approved. For example, after it was determined that a
consequent impacts on survival and reproduction of proposed expansion of ski runs and lifts disrupted an elk
wildlife in adjacent alpine and sub-alpine habitats. migration route in Colorado, the Aspen Skiing Company
Increases in recreational use of high elevation areas revised their plans in consultation with the Pitkin County
increases food availability for generalist predators. High Commission, Aspen town council, Colorado Division of
elevation habitats are generally unsuitable for crows or Wildlife, and The Sierra Club local chapter63 (web site:
Steller ’s jays, but populations can exist if there are http://www.skiaspen.com/environment). Construction
dwellings or dumps nearby.121 Along the Cascade Crest, of ski runs and buildings was done in winter. Trees were
crows occur in ski areas and towns. Raven densities may cut, but no bulldozer grading was done, to avoid
increase along roads as a result of road kills. A study disturbing sub-alpine vegetation. Little is known, or being
during the breeding season in northern Finland found investigated, regarding potential impacts to alpine or sub-
more magpies and crows around three ski hill centers than alpine species from other disturbances associated with ski
in pristine forest of the same type and age in the region. facilities, such as large volumes of sewage and garbage,
Also, predation of artificial nests was much higher for the compacted sub-alpine vegetation, light pollution from
ski hill centers than the undisturbed forest (J. Jokimaki, night skiing, and the consequences of later spring green-
unpubl. data, [email protected]). up caused by delayed snow melt from compacted snow
Human activities at lower elevations might also and snow making.89
influence ecological processes in the alpine. Mid-sized In the past two decades, there has been a dramatic
predators like ravens and coyotes have substantial home increase in alpine cross-country ski touring, and in the
ranges and, could easily travel 5 to 10 kilometers from use of helicopters to access remote areas for skiing, fishing,
lower elevations to the alpine, but it is unknown whether and hiking. Such activities do not result in many visits
they do so. Over-winter survival of generalist predators per mountain; however, the overall size of area impacted
is improved by road-kills, dumps in adjacent urban areas, has increased dramatically. The impacts of such
or from feeding in backyards at holiday homes or disturbances on wildlife are difficult to monitor, control
ranchettes in montane forest. Generalist predators no or to predict. Ski touring groups in Bavaria cause
longer depend year-round on traditional prey to survive considerable disturbance to wintering populations of
when anthropogenic food supplies are available, but if grouse and ungulates. 148 Proactive multi-partner
they kill the natural prey they encounter even in just one consultations have had good success in designing ski
season, they could profoundly impact alpine prey touring routes with leave areas for wintering wildlife in
population densities. Given that many high elevation Bavaria.127, 148
wildlife species are long-lived, a reduction in adult The increased use of alpine facilities in fall, particularly
survival has a relatively higher demographic impact than with late summer hiking and mountain biking, may result
for r-selected species. We need to conduct field research in conflicts with seasonal movements of wildlife from late
on possible disturbance factors that increase generalist summer habitats to winter sites. Careful evaluation of
predator densities in or near high elevation areas, whether habitat use should be made, and seasonal wildlife travel
these predators spend more time at high elevation, and corridors determined in advance of designing and
whether these increases significantly impact alpine approving new ski developments or expanding existing
wildlife populations. At the same time, we need to manage operations. The effects of ski developments cover a much
 256 Wildlife–Habitat Relationships in Oregon and Washington

larger area than proposed development sites, particularly habitats has declined. Thus, we need to include life history
since they are often positioned at the headwaters of periods outside the breeding season when evaluating high
watersheds.89 A secondary effect of ski area developments elevation habitats for wildlife. Connectivity is a key
is the urbanization of mountain valleys. Increasingly, ski ecological process to maintain high elevation wildlife
resort areas are becoming amenities to support luxury populations. Connectivity needs to be maintained (1)
second home communities. Reviews of proposed new among patchy alpine habitats, (2) along mountain
developments or expansions of ski resort areas should corridors for north-south migrants, and perhaps most
consider off-site impacts from this secondary urbanization, importantly (3) between alpine and adjacent lower
including all concerns relevant to urban areas near alpine elevation habitats and valley bottoms.
habitats. Potential impacts should be evaluated at a Despite being highly valued for their intrinsic beauty
sufficiently large scale to include connectivity issues and wildness, alpine vertebrates and high elevation
discussed earlier for maintaining viable mountain wildlife habitats are a neglected area for management and research.
populations. Proposed alpine recreational developments Even though currently most alpine species appear secure
should demonstrate that they have allowed for in mostly stable habitat conditions, change within and
maintenance of normal ecological processes. adjacent to alpine and sub-alpine habitats is happening
rapidly and over extensive areas. Alpine ecosystems have
7. Summary experienced large increases in amount and kinds of human
The alpine zone consists of rugged, partially-vegetated use, with some areas showing significant deterioration
terrain with snowfields and rocky ridges, above the caused by recreational activities, livestock grazing, mining,
natural treeline. Alpine ecosystems are structurally simple and air-borne contaminants. High elevation habitats are
with few plant species compared to most lower elevation vulnerable to erosion, especially near melting snowfields
habitats. In Washington and Oregon, the main wildlife or when soil crusts are broken above volcanic ash, and
habitats are (1) alpine grasslands and shrublands, and (2) vegetation recovers slowly after such disturbances. To
sub-alpine parklands. These habitats are characterized by prevent elevated predation risk for alpine wildlife, we
high winds, prolonged snow cover, steep terrain, extremes should avoid inadvertent increases in food supplies for
of heat and cold, and intense ultra-violet radiation. With generalist predators in and near high elevation habitats.
increasing elevation, time for breeding decreases and On a landscape scale, developed valley bottoms constitute
environmental stochasticity increases; at the highest dispersal barriers for many wildlife species with seasonal
elevations, hypoxic conditions add additional energetic vertical movements. Since we lack comprehensive
living costs. Other ecological costs may also be high, such understanding of what is required to maintain ecological
as high predation risk in the open alpine habitats, and the processes in alpine habitats and have limited experience
need for seasonal movements from patchy breeding in restoring these sensitive ecosystems, the precautionary
habitats to wintering areas. These factors result in short, principle is strongly advised for managing alpine and sub-
intense breeding seasons for wildlife. alpine communities. Finally, we need to establish long-
About one third of the vertebrate fauna (188 species) term monitoring programs to detect and predict the
in Washington and Oregon use alpine or sub-alpine ecological effects of climate change on alpine habitats and
habitats at some period in their life history. These include wildlife populations.
65 species of mammals, 112 birds and 11 reptiles and
amphibians. Only a few species such as white-tailed Acknowledgments
ptarmigan, hoary marmots and mountain goats live I thank Environment Canada for allowing me time to
exclusively in the alpine, while the majority breed in both develop the ideas for this chapter. I thank I. Storch, The
alpine and lower elevation habitats or use high elevation Munich Wildlife Society, Germany, and J. L. Martin, cefe-
habitats during migration. For true alpine breeders, some CNRS, Montpellier, France for providing office logistics
life history traits may vary with altitude as reproductive to produce this paper. Numerous colleagues from Europe
rates are often lower, development times longer, with and North America provided insights and discussed ideas
accompanying increases in longevity. For wildlife species concerning alpine ecology and conservation: W. Arnold,
breeding across elevational gradients, reproductive A. Bernard-Laurent, C. Böhn, C. E. Braun, U. Breitenmoser,
success at high elevation appears similar to those for Ch. Breitenmoser-Würsten, R. W. Campbell, E. R. Dunn,
populations at lower elevations. However, with L. E. Ellison, M. Festa-Bianchet, C. L. Gass, H. Gossow, M.
compressed reproductive seasons, wildlife at high Guentert, A. Landmann, J.-L. Martin, M. A. Schroeder, I.
elevation produce fewer offspring in a season. Storch, D. B. A. Thompson, H. Zeiler, and A. Zeitler. D. H.
An unexamined aspect of biodiversity for high Johnson and K. Bettinger provided vital local material and
elevation ecosystems is their use by migrating wildlife guidance during the chapter production. K.E.H. Aitken,
during late summer and fall. From late July through C. L. Hitchcock, and M.D. Mossop provided logistical
October., alpine areas support a rich diversity and assistance. I thank C. E. Braun, M. Festa-Bianchet, C. M. I.
abundance of birds and mammals that move up from Meslow, B. K. Sandercock, M. A. Schroeder, J. N. M. Smith,
lower elevations, as well as high latitude avian migrants. and N.A.M. Verbeek for reviewing various drafts.
This is a time when alpine habitats offer rich food
resources, while productivity in many low elevation
 ALPINE/SUBALPINE 257

Literature Cited 26. Bullard, R. W. 1972.Vertebrates at altitudes. Pages 209-226 in M. K.

Yousef, S. M. Horvath, and R. W. Bullard editors. Physiological
1. Agee, J.K. and L. Smith. 1984. Subalpine tree establishment after fire in
adaptations, desert and mountain. Academic Press, New York, NY.
the Olympic Mountains, Washington. Ecology 65: 810-819.
27. Burton, J. F. 1995. Birds and climate change. Helm, London, UK.
2. Allee, W. C. and K. P. Schmidt. 1951. Ecological Animal Geography. John
28. Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W.
Wiley and Sons, Inc. New York, NY.
Kaiser, M. C. E. McNall, G. E. J. Smith. 1990 and 1996. The Birds of British
3. Armitage, K. B. 1986. Individuality, social behavior, and reproductive
Columbia.Volumes 1-3. University of British Columbia Press,Vancouver,
success in yellow-bellied marmots. Ecology 67: 1186-1193.
BC, Canada.
4. ———. 1999. Evolution of sociality in marmots. Journal of Mammalogy
29. Cannings, R. A., R. J. Cannings, and S. G. Cannings. 1987. The birds of
80: 1-10.
the Okanagan Valley. Royal BC Museum,Victoria, BC, Canada.
5. Arno, S. F. and R.P. Hammerly. 1984. Timberline: Mountains and arctic
30. Cannings, R. J., and S. G. Cannings. 1996. British Columbia, A Natural
forest frontiers. The Mountaineers, Seattle, WA.
History. Douglas and McIntyre, Vancouver, BC, Canada.
6. Arnold, W. 1990a. The evolution of marmot sociality: I. Why disperse
31. Capula, M, L. Luiselli, and C. Anibaldi. 1992. Complementary study on
late? Behavioural Ecology and Sociobiology 27: 229-237.
the reproductive biology in female adder, Vipera berus (L.) from eastern
7. ———. 1990b. The evolution of marmot sociality: II. Costs and benefits
Italian Alps.Vie Milieu 42: 327-336.
of joint hibernation. Behavioural Ecology and Sociobiology 27: 239-246.
32. Carey, C. 1980. Adaptation of the avian egg to high altitude. American
8. ———. 1993. Social evolution in marmots and the adaptive value of
Zoologist, 20: 449-459.
joint hibernation. Verh. Deutsch Zool. Ges. 86: 79-93.
33. ———, and K. Martin. 1997. Physiological ecology of incubation of
9. Bätzing, W. 1991. Die alpen: entstehung und Gefährdung einer
ptarmigan eggs at high and low altitudes. Wildlife Biology 3: 211-218.
europäischen kulturlandschaft. Beck, Munich, Germany.
34. Carpenter, F. L., and M. A. Hixon. 1988. A new function for torpor: fat
10. Beason, R.C. 1995. Horned lark (Eremophila alpestris). Pages 1-24 in A.
conservation in a wild migrant hummingbird. The Condor 90: 373-378.
Poole and F. Gill, editors. The Birds of North America, No. 195. The
35. Cassidy, K. M., C. E. Grue, M. R. Smith, and K. M. Dvornich, editors.
Academy of Natural Sciences; Philadelphia.
1997. Washington State Gap Analysis - Final Report.Volumes 1-5.
11. Billings, W. D. 1989. Alpine vegetation. Pages 392-420 in W. D. Billings
Washington Cooperative Fish and Wildlife Research Unit, University of
and R. K. Peet, editors. Terrestrial vegetation of North America. Paragon
Washington, Seattle, WA.
Books.
36. Chappell, C.B., R.C. Crawford, J. Kagan, D.H. Johnson, M. O’Mealy, G.A.
12. ———, and H. A. Mooney. 1968. The ecology of arctic and alpine
Green, H.L. Ferguson, and W.D. Edge. 2001. Wildlife habitats:
plants. Biological Review 43: 481-529.
descriptions, status, trends, and system dynamics. Pages 22-114 in D.H.
13. Blais, J. M., D, W. Schindler, D. C. G. Muir, L. E. Kimpes, D. B. Donald, B.
Johnson and T.A. O’Neil, managing directors. Wildlife-habitat
Rosenburg. 1998. Accumulation of persistent organochlorine
relationships in Oregon and Washington. Oregon State University
compounds in mountains of western Canada. Nature 395: 585-588.
Press, Corvallis, OR.
14. Böhn, C., and A. Landmann. 1995. Nest-site selection and nest
37. Chapin, F. S. III, and C. Körner, editors. 1994. Arctic and alpine
construction in the water pipit (Anthus spinoletta). Journal für
biodiversity: patterns, causes and ecosystem consequences. Springer-
Ornithologie 136: 1-16.
Verlag, New York, NY.
15. Brace, S. and D. L. Peterson. 1998. Spatial patterns of tropospheric
38. Cody, M. L. 1985. Habitat selection in grassland and open country
ozone in the Mount Rainier region of the Cascade Mountains, U.S.A.
birds. Pages 191-226 in Habitat selection in birds. Academic Press. New
Atmospheric Environment, 32: 3629-3637.
York, NY.
16. Braun, C. 1969. Population dynamics, habitat, and movements of
39. Corkran, C.C. and C. R. Thoms. 1996. Amphibians of Oregon,
White-tailed Ptarmigan in Colorado. Ph.D. thesis, Colorado State
Washington, and British Columbia. Lone Pine,Vancouver, BC, Canada.
University, Fort Collins, CO.
40. Côte, S. D. 1996. Mountain goat responses to helicopter disturbance.
17. Braun, C. E. 1971. Habitat requirements of Colorado white-tailed
Wildlife Society Bulletin 24: 681-685.
ptarmigan. Proceedings of Western Association of State Game and Fish
41. Crisafulli, C. M., and C. P. Hawkins. 1998. Ecosystem recovery
Commissioners. 51: 284-292.
following a catastrophic disturbance: Lessons learned from Mount St.
18. Braun, C. E., and R. G. Streeter. 1968. Observations on the occurrence
Helens. in M. J. Mac, P. A. Opler, C. E. Puckett Haecker, and P. D. Doran,
of white-tailed jackrabbits in the alpine zone. Journal of Mammalogy 49:
editors. Status and trends of the nations biological resources.Volumes
160-161.
1-2. U.S. Department of Interior, U.S. Geological Survey, Reston, VA.
19. Braun, C. E., R. W. Hoffman, and G. E. Rogers. 1976. Wintering areas
42. Csuti, B., A.J. Kimerling, T.A. O’Neil, M.M Shaughnessy, E.P. Gaines, and
and winter ecology of white-tailed ptarmigan in Colorado. Special
M. M. P. Huso. 1997. Atlas of Oregon wildlife: Distribution, habitat and
Report No. 38, Colorado Division of Wildlife, Denver, CO.
natural history. Oregon State University Pres, Corvallis, OR.
20. Braun, C. E., R. K. Schmidt, and G. E. Rogers. 1973. Census of
43. Daubenmire, R. 1954. Alpine timberlines in the Americas and their
Colorado white-tailed ptarmigan with tape-recorded calls. Journal of
interpretation. Butler University Botanical Studies 11: 119-136.
Wildlife Management 37:90-93
44. Dobson, F. S. and J.O. Murie. 1987. Interpretation of intraspecific life
21. ———. 1980. Alpine bird communities of western North America:
history patterns: evidence from Columbian ground squirrels. American
implications for management and research. Pages 280-291 in R. M.
Naturalist 129: 382-397.
DeGraff and N. G. Tilghman, compilers. Workshop Proceedings:
45. ———, R. M. Zammuto, and J.O. Murie. 1986. A comparison of
Management of western forests and grasslands for nongame birds. U.S.
methods for studying life history in Columbian ground squirrels.
Department Agriculture Forest Service General technical report INT-
Journal of Mammalogy 667: 154-158.
86.
46. Douglas, G.W. and L.C. Bliss. 1977. Alpine and high subalpine plant
22. ———, K. Martin, and L. A. Robb. 1993. White-tailed ptarmigan
communities of the north Cascades Range, Washington and British
(Lagopus leucurus). Pages 1-24 in A. Poole, and F. Gill, editors.The birds
Columbia. Ecological Monographs 47: 113-150.
of North America, No. 68. Philadelphia: The Academy of Natural
47. Dragon, S., C. Carey, K. Martin, and R. Bauman. 1999. Effect of high
Sciences; Washington, D.C.
altitude and in vivo adenosine/β-adrenergic receptor blockade on ATP
23. Breitenmoser, U. 1998. Large predators in the Alps: the fall and rise of
and 2,3BPG concentrations in red blood cells of avian embryos.
man’s competitors. Biological Conservation 83: 279-289.
Journal of Experimental Biology 202:2787-2795
24. Bronson, M. T. 1979. Altitudinal variation in the life history of the
48. Edge, W.D. 2001. Wildlife of agriculture, pastures, and mixed environs.
golden-mantled ground squirrel (Spermophilus lateralis). Ecology 60: 272-
Pages 342-360 in D.H. Johnson and T. A. O’Neil, managing directors.
279.
Wildlife-habitat relationships in Oregon and Washington. Oregon State
25. Bruderer, B. 1982. Do migrating birds fly along straight lines? Pages 3-
University Press. Corvallis, OR.
14 in F. Papi and H. G. Wallraff, editors. Avian Migration. Springer,
Heidelberg, Germany.
 258 Wildlife–Habitat Relationships in Oregon and Washington

49. Edwards, O. M. 1980. The alpine vegetation of Mount Rainier National 72. ———. 1995. Effects of climate change on mountain bird populations.
Park: structure, development and constraints. Ph.D. dissertation, Univ. Pages 73-74 in A. Guisan et al., editors. Potential Ecological impacts of
of Wash., Seattle,WA. climate change in the Alps and Fennoscandian Mountains.
50. Ferguson, H.L., K. Robinette, K. Stenberg. 2001. Wildlife of urban Conservatoire et Jardin botaniques de la Ville de Geneve: Geneva,
habitats. Pages 317-341 in D.H. Johnson and T.A. O’Neil, managing Switzerland.
directors. Wildlife-habitat relationships in Oregon and Washington. 73. Jenik, J. 1997. The diversity of mountain life. Pages 199-231 in B.
Oregon State University Press. Corvallis, OR. Messerli, and J. D. Ives, editors. Mountains of the world: a global priority.
51. Ferron, J., and J. P. Ouellet. 1989. Temporal and intersexual variations Parthenon Publishing Group, London, UK.
in the use of space with regard to social organization in the 74. Johnson, R. E. 1965. Reproductive activities of rosy finches, with
woodchuck Marmota monax. Canadian Journal of Zoology 67: 1642- special reference to Montana. Auk, 82: 190-205.
1649. 75. ———. 1983. Nesting biology of the rosy finches on the Aleutian
52. Festa-Bianchet, M., M. Urquhart, and K.G. Smith. 1994. Mountain goat Islands, Alaska. Condor 85: 447-452.
recruitment: kid production and survival to breeding age. Canadian 76. Kiilsgaard, C. W. 1999. Oregon vegetation: mapping and classification
Journal of Zoology 72: 22-27. of landscale level cover types. Final report submitted to the U.S.
53. Filley, B. 1996. The big fact book about Mount Rainier. Dunamis House Geological Survey, Biological Resources Division: GAP Analysis
Publishing, Issaquah,WA. Program, Moscow, ID.
54. Foreyt, W. J., K. P. Snipes, and R. W. Kasten. 1994. Fatal pneumonia 77. Kingery, H. E. 1996.American Dipper (Cinclus mexicanus). Pages 1-28 in
following innoculation of healthy bighorn sheep with Pasturella A. Poole, and F. Gill, editors. The birds of North America, No. 229. The
haemolytica from healthy domestic sheep. Journal of Wildlife Disease Academy of Natural Sciences; Philadelphia. Washington, D.C.
30: 137-145. 78. Kirk, R. 1999. Sunrise to paradise: the story of Mount Rainier
55. Franklin, J. F., and C. T. Dyrness. 1973. Natural Vegetation of Oregon National Park. University of Washington Press, Seattle, WA.
and Washington. Oregon State University Press, Corvallis, OR. 79. Kollinsky, C., and A. Landmann. 1996. Altitudinal distribution of male
56. Franklin, J. F., W.H. Moir, M. A. Hemstrom, S. E. Greene, and B. G. Smith. black redstarts: are there age-dependent patterns? Bird Study 43: 103-
1988. The forest communities of Mount Rainier National Park. U.S.D.I. 107.
National Park Service Scientific Monograph Series 19, Washington, D.C. 80. Körner, C. 1994. Alpine plant diversity, a global survey and functional
57. Frase, B. A. and R. S. Hoffman. 1980. Marmota flaviventris. Mamalian interpretation. Pages 45-64 in F. S. Chapin III, and C. Körner, editors.
Species 135: 1-8. Arctic and alpine biodiversity: patterns, causes and ecosystem
58. French, N. R. 1959. Life history of the black rosy finch. Auk 76: 158- consequences. Springer-Verlag, New York, NY.
180. 81. ———. 1999. Alpine Plant Life: functional plant ecology of high
59. Frenzen, P. M. and C. M. Crisafulli. 1990. Mount St. Helens ten years mountain ecosystems. Springer-Verlag, Heidelburg, Germany.
later: past lessons and future promise. Northwest Science 64: 263-267. 82. Krajick, K. 1998. Animals thrive in an avalanche’s wake. Science 279:
60. Gaillard, J. -M., M. Festa-Bianchet, and K. G. Smith. 1998. Population 1853.
dynamics of large herbivores: variable recruitment with constant adult 83. Kuramoto, R. T. and L. C. Bliss. 1970. Ecology of subalpine meadows in
survival. Trends in Research in Evolution and Ecology 13: 58-63. the Olympic Mountains, Washington. Ecological Monographs 40:317-
61. Gass, C. L., and K. P. Lertzman. 1980. Capricious mountain weather: a 347.
driving variable in hummingbird territorial dynamics. Canadian Journal 84. Landmann, A. and N. Winding. 1993. Niche segregation in high-altitude
of Zoology 58: 1964-1968. Himalayan chats (Aves, Turdidae): does morphology match ecology?
62. ———, and G. Sutherland. 1985. Specialization by territorial Oecologia 95: 506-519.
hummingbirds on experimentally enriched patches of flowers: energetic 85. ———, and ———. 1995a. Adaptive radiation and resource
profitability and learning. Canadian Journal of Zoology 63: 2125-2133. partitioning in Himalayan high-altitude finches. Zoology Analysis of
63. Giezentanner, K. 1994. Wildlife and biodiversity analysis for the Complex Systems 99: 8-20.
Snowmass Ski Development expansion. U.S. Forest Service report, CO. 86. ———, and ———. 1995b. Guild organization and morphology of
64. Grabherr, G. and M.G. H. Pauli. 1994. Climate effects on mountain high-altitude granivorous and insectivorous birds: convergent evolution
plants. Nature 369: 448. in an extreme environment. Oikos 73: 237-250.
65. Grossman, D.H., D. Faber-Langendoen, A.W. Weakley, M. Anderson, P. 87. Lanner, R. M. 1988. Dependence of Great Basin Bristlecone pine on
Bourgeron, R. Crawford, K. Goodin, S. Landaal, K. Metzler, K.D. Clark’s nutcracker for regeneration at high elevations. Arctic and Alpine
Patterson, M. Pyne, M. Reid and L. Sneddon. 1998. International Research, 20: 358-362.
classification of ecological communities: terrestrial vegetation of the 88. Larison, J.R., G.E. Likens, J.W. Fitzpatrick, and J.G. Crock. 2000.
United States.Vol. 1: The National Vegetation Classification Standard. Cadmium toxicity among wildlife in the Colorado Rocky Mountains.
The Nature Conservancy. Nature 406: 181-183.
66. Hannon, S.J. and K. Martin. 1996. Mate fidelity and divorce in 89. Legault, S. 1997. Down a slippery slope: ski resorts and their effect on
ptarmigan: polygyny on the tundra. Pages 192-210 in J.M. Black, editor. ecological, social and economic health. Unpublished Discussion paper,
Partnerships in birds, the study of monogamy. Oxford University Press, UTSB Research Inc., Banff, Alberta, Canada.
Oxford, England. 90. Lippman, M. 1989. Health effects of ozone: a critical review. Journal of
67. Hoffman, R. W. and C. E. Braun. 1975. Migration of a wintering the air and waste management association 39: 672-695.
population of white-tailed ptarmigan in Colorado. Journal of Wildlife 91. Louis, S., and M. Le Berre. 1997. Visitor impact on alpine marmot
Management 39: 485-490. foraging behavior. Journal of Wildlife Research 2: 133-136.
68. Houard, T., and M. Mure. 1997. Les tetras-lyre des vallons de Salese et 92. Love, D. 1970. Subarctic and subalpine: where and what? Arctic and
Mollieres. Parc National du Mercantour. Domaine vital et influence du alpine research 2: 63-73.
tourisme. Revue d’Ecologie (La Terre et la Vie), Supplemente 4: 165-197. 93. Lyon, B. E. and R. D. Montgomerie. 1987. Ecological correlates of
69. Houston, D. B., and V. Stevens. 1988. Resource limitation in mountain incubation feeding: a comparative study of high arctic finches. Ecology
goats: a test by experimental cropping. Canadian Journal of Zoology 66: 68: 713-722.
228-238. 94. Manuwal, D. A., M. H. Huff, M. R. Bauer, C. B. Chappell, and K. Hegstad.
70. Inouye, D.W., B. Barr, K.B. Armitage and B.D. Inouye. 2000. Climate 1987. Summer birds of the upper subalpine zone of Mount Adams,
change is affecting altitudinal migrants and hibernating species. Mount Rainier, and Mount St. Helens, Washington. Northwest Science,
Proceedings of the National Academy of Science 97: 1630-1633. 61: 82-92.
71. Järvinen, A. 1994. Global warming and egg size of birds. Ecography 17: 95. Martin, K., C. E. Braun, and P. B. Stacey. 2000. Recruitment, dispersal,
108-110. and demographic rescue in spatially structured white-tailed ptarmigan
populations. Condor 102:503-506.
 ALPINE/SUBALPINE 259

96. ———, C. Doyle, F. Mueller and S. J. Hannon. 2001. Forest grouse and 119. Smith, A. T. 1974. The distribution and dispersal of pikas: influences of
ptarmigan. Chapter 11In C.J. Krebs, S. Boutin, and R. Boonstra, editors. behavior and climate. Ecology 55: 1368-1376.
Ecosystem dynamics of the Boreal Forest: The Kluane Project. Oxford 120 Smith, A. T. 1978. Comparative demography of pikas (Ochotona):
University Press, in press. effect of spatial and temporal age-specific mortality. Ecology 59: 133-
97. ———, R. F. Holt and D. W. Thomas. 1993. Getting by on high: 139.
ecological energetics of arctic and alpine grouse. Pages 33-41 in C. 121. Smith, M. R., P. W. Mattocks, Jr., and K. M. Cassidy. 1997. Breeding
Carey, G. L. Florant, B. A. Wunder, and B. Horwitz, editors. Life in the birds of Washington State.Volume 4 in K. M. Cassidy, C. E. Grue, M. R.
Cold: Ecological, physiological, and molecular mechanisms. Westview Smith, and K. M. Dvornich, editors. Washington State Gap Analysis—
Press, Boulder, CO. Final report. Seattle Audubon Society Publications in Zoology No.1,
98. ———, P. B. Stacey, and C. E. Braun. 1997. Demographic rescue and Seattle, WA.
maintenance of population stability in grouse - beyond 122. Spalding, J. B. 1979. The aeolian ecology of White Mountain Peak,
metapopulations. Wildlife Biology 3: 295-296. California: wind-blown insect fauna. Arctic and Alpine Research 11: 83-
99. Medin, D. E. 1987. Breeding birds of an alpine habitat in the southern 94.
Snake Range, Nevada.Western Birds 18: 163-168. 123. Stabler, R. M., N. J. Kitzmiller, and C. E. Braun. 1974. Hematozoa from
100. Menoni, E., and Y. Magnani. 1998. Human disturbance of grouse in Colorado birds, IV. Galliformes. Journal of Parasitology 60: 536-537.
France. Grouse News, No. 15: 4-8. 124. Storch, I. 1993. Patterns and strategies of winter habitat selection in
101. Miquet, A. 1986. Contribution a la etude des relations entre Tetras- alpine capercaillie. Ecography 16: 351-359.
lyre Tetrao tetrix L.Tetraonidae et tourisme hivernal en Haute- 125. ———. 1997. The role of the metapopulation concept in
Tarentaise. Acta Oecologia, Oecologia Applications 7: 325-355. conservation of European woodland grouse. Wildlife Biology, 3: 272.
102. Nethersole-Thompson, D., and A. Watson. 1981. The Cairngorms. 126. ———, compiler. 2000. Grouse Status Survey and Conservation
Melven Press, Perth, U.K. Action Plan 2000-2004. WPA/Birdlife/SSC Grouse Specialist Group.
103. Neuhaus, P. and B. Mainini. 1998. Reactions and adjustment of adult IUCN, Gland. Switzerland and Cambridge, UK and the World Pheasant
and young alpine marmots Marmota marmota to intense hiking Association, Reading, UK
activities. Wildlife Biology, 4: 119-123. 127. Suchant, R., and R. Roth. 1998. Tourism in the Black Forest - danger
104. Norvell, J. R., and P. D. Creighton. 1990. Foraging of horned larks and for the capercaillie. Grouse News, No. 15: 13-17.
water pipits in alpine communities. Journal of Field Ornithology 61: 128. Sutherland, G. D., C. L. Gass, P. A. Thompson, and K. P. Lertzman. 1982.
434-440. Feeding territoriality in migrant rufous hummingbirds: defense of
105. Noss, R. F., H.B. Quigley, M.G. Hornocker, T. Merrill, and P.C. Paquet. yellow-bellied sapsucker (Sphyrapicus varius) feeding sites. Canadian
1996. Conservation biology and carnivore conservation in the Rocky Journal of Zoology 60: 2046-2050.
Mountains. Conservation Biology, 10: 949-963. 129. Sutherland, W. J. and J. D. Reynolds. 1998. Sustainable and
106. Observatoire des Galliformes de Montagne. 1997. Rapport annuel unsustainable exploitation. Pages 90-115 in W. J. Sutherland, editor.
de l’Observatoire des Galliformes de Montagne. Conservation science and management. Blackwell, Oxford, UK.
107. Onderka, D. K., S. A. Rawluk, and W. D. Wishart. 1988. Susceptibility 130. Taylor, K. A. 1994. Pages 165-185 in A Birder’s Guide to Vancouver
of Rocky Mountain bighorn sheep and domestic sheep to pneumonia Island. Keith Taylor Birdfinding Guides, Victoria, BC, Canada.
induced by bighorn and domestic livestock strains of Pasturella 131. Thompson, D. B. A. and D. P. Whitfield. 1993. Research on mountain
hemolytica. Canadian Journal of Veterinary Research 52: 439-444. birds and their habitats. Scottish Birds 17: 1-8.
108. Pattie, D. L. and N.A.M. Verbeek. 1966. Alpine birds of the Beartooth 132. ———, A. Watson, S. Rae, and G. Boobyer. 1996. Recent changes in
Mountains. Condor 68: 167-176. breeding bird populations in the Cairngorms. Botanical Journal of
109. Piertney, S. B., A. D. C. MacColl, P. J. Bacon, and J. F. Dallas. 1998. Local Scotland. 48: 99-110.
genetic structure in red grouse (Lagopus lagopus scoticus): evidence from 133. Twining, H. 1940. Foraging behavior and survival in the Sierra Nevada
microsatellite DNA markers. Molecular Ecology, 7: 1645-1654. rosy finch. Condor 42: 64-72.
110. Portier, C., M. Festa-Bianchet, J. M. Gaillard, J. T. Jorgenson and N. G. 134.Verbeek, N. A. M. 1967. Breeding biology and ecology of the horned
Yoccoz. 1998. Effects of density and weather on survival of bighorn lark in alpine tundra. Wilson Bulletin 79: 208-217.
sheep lambs (Ovis canadensis). Journal of Zoology, London 245: 271-278. 135. ———. 1970. Breeding ecology of the water pipit. Auk 87: 425-451.
111. Price, L. W. 1981. Mountains and man: a study of process and 136. ———. 1981. Nesting success and orientation of water pipit Anthus
environment. University of California, Berkeley, CA. spinoletta nests. Ornis Scandinavica 12: 37-39.
112. Price, M. F. and R. G. Barry. 1997. Climate change. Pages 409-445 in B. 137. Watson, A. 1979. Bird and mammal numbers in relation to human
Messerli, and J. D. Ives, editors. Mountains of the world: a global priority. impact at ski lifts on Scottish hills. Journal of Applied Ecology, 16: 753-
Parthenon Publishing Group, London. 764.
113. Pybus, M. J., S. Groom, and W. M. Samuel. 1996. Meningeal worm in 138. ———. 1982. Effect of human impact on ptarmigan and red grouse
experimentally infected bighorn and domestic sheep. Journal Wildlife near ski lifts in Scotland. Page 51 in Annual Report of ITE for 1981,
Disease 32: 614-618. Cambridge, UK.
114. Rochefort, R. M. and S.T. Gibbons. 1992. Mending the meadow: high 139. ———. 1991. Increase of people on Cairngorm plateau following
altitude meadow restoration in Mount Rainier National Park. easier access. Scottish Geographical Magazine 107: 99-105.
Restoration and Management Notes 10: 120-126. 140. Weaver, J.L., P.C. Paquet and L.F. Ruggiero. 1996. Resilience and
115. Roland, J., N. Keyghobadi, and S. Fownes. 2000. Alpine Parnassius conservation of large carnivores in the Rocky Mountains. Conservation
butterfly dispersal: effects of landscape and population size. Ecology 81: Biology 10: 964-976.
1642-1653. 141.Widmer, M. 1993. Breeding biology of the garden warbler Sylvia borin
116 Saint Girons, H. 1992. Strategies reproductrices des viperidae dans in a subalpine habitat in the central Swiss Alps. Der Ornithologische
les zones temperees fraiches et froides. Bulletin de la Société Beobachter 90: 83-115. German with English abstract.
zoologique de France 117: 267-278. 142. ———. 1996. Phänologie, Siedlungsdichte und Populationsökologie
117. Shackleton, D. 1999. Hoofed mammals of British Columbia. der Gartengrasmucke Sylvia borin in einem subalpinen Habitat der
University of British Columbia Press,Vancouver, BC, Canada. Zentralalpen. Journal für Ornithologie, 137: 479-501.
118. Shaughnessy, M. M, and T.A. O’Neil. 2001. Conservation of 143. ———. 1999. Altitudinal variation of migratory traits in the garden
biodiversity: Considerations and methods for identifying and warbler Sylvia borin. Ph. D.Thesis, University of Zurich, Zurich,
prioritizing areas and habitats. Pages 154-167 in D.H. Johnson and T.A. Switzerland.
O’Neil, managing directors. Wildlife-habitat relationships in Oregon and 144. Wiebe, K. L., and K. Martin. 1997. Effects of predation, body
Washington. Oregon State University Press. Corvallis, OR. condition and temperature on incubation rhythms of ptarmigan.
Wildlife Biology 3: 219-227.
 260 Wildlife–Habitat Relationships in Oregon and Washington

145. ———, and ———. 1998. Age-specific patterns of reproduction in

white-tailed and willow ptarmigan (Lagopus leucurus and L. lagopus). Ibis
140:14-24.
146. Woods, J.G. 1991. Ecology of a partially migratory elk population.
Ph.D. Thesis. University of British Columbia, Vancouver, BC, Canada.
147. Woodward, F. I. (editor). 1992. Advances in Ecological Research.
Volume 22. The ecological consequence of global climate change.
Academic Press, London.
148. Zeitler, A. and U. Glanzer. 1998. Skiing and grouse in the Bavarian
Alps. Grouse News, 15: 8-12.
Web sites:
Centre for Alpine Studies (UBC): http://www.www.forestry.ubc.ca/alpine/
index.htm
Centre of Alpine ecology (Italy): http://www.itc.it/cea/index_e.html
.
 10
Wildlife of Westside Grassland and
Chaparral Habitats
Bob Altman, Marc Hayes, Stewart Janes,
& Richard Forbes

Introduction
Westside Grasslands and Chaparral (Ceanothus- Westside valley grasslands include glacial outwash
Manzanita Shrublands) are among the most distinctive prairies of the Puget Lowlands of Washington, particularly
and endangered wildlife habitats in Oregon and the South Puget Sound area; and early seral fire-
Washington. They are characterized by a limited maintained grasslands of the Willamette, Rogue, and
distribution either naturally (i.e., chaparral) or from Umpqua valleys in Oregon.43 Most discussion in this
extensive losses associated primarily with post-European chapter focuses on these grasslands, which comprise the
settlement (i.e., grassland). Although not as species rich largest area of westside grasslands. Limited reference is
or biologically diverse as many habitats, they contain an made to coastal headland and dune grasslands and
array of vertebrates that occur rarely elsewhere in Oregon shrublands, montane grass balds, and balds in the San
and Washington. These habitats also occur where most of Juan Islands, which are minor components of westside
the human population lives, thus they have been grasslands and for which few data exist on wildlife
extensively altered since European settlement. relationships. Coastal headland grasslands and
Loss of westside native grassland habitat has been shrublands include flat coastal plain grasslands (e.g.,
especially dramatic; >90% in the Puget Lowlands27 and Quillayute Prairie, Washington) and the rocky, herbaceous
>99% in the Willamette Valley.23 Remaining parcels are and low shrub-dominated exposed headlands along the
invariably small, moderately to heavily disturbed, and ocean (e.g., Heceta Head, Oregon; Cascade Head, Oregon).
geographically disjunct. Consequences for grassland- These grasslands are generally minor components within
associated wildlife have been local and regional the Sitka spruce (Picea sitchensis) vegetation zone. Montane
extirpation of some species, and severe declines among balds are grass-dominated, south-facing hillsides of Coast
other species with a concomitant increase in exotic species. Range and Olympic Peninsula summits. They occur as
scattered but conspicuous openings in an otherwise forest-
Habitat Types dominated landscape.43 Balds in the San Juan Islands are
low elevation (<2,000 feet [610 m]) dry sites on south to
Grasslands west aspects, moderate to steep slopes, with shallow to
Throughout this chapter we use the term grassland to bedrock soils22 (C. Chappell, Washington Natural Heritage
include both prairie and savanna habitats (see Chapter 2 Program, pers. comm.).
for descriptions). We emphasize to the extent possible We categorized grasslands into three types: wet prairie,
native grasslands, which occur as scattered historical dry prairie, and savanna. In the Willamette Valley, wet
remnants. We also acknowledge agricultural grasslands prairies occur primarily on poorly drained (high clay
(e.g., hayfields, pasture, annual and perennial grass fields, fraction) soils134 (see Chapter 2 this volume—herbaceous
fallow agricultural fields) because these dominate the wetlands). Dry prairies occur on better drained lowland
current landscape, and some wildlife associated with them and valley soils, especially along valley margins. 134
were integral components of native grasslands. Savanna refers to dry prairie with singular widely
Westside grasslands are open landscapes characterized scattered trees or small open-canopy tree groves with
by herbaceous plants, particularly graminoids. Periodic <30% canopy cover.1 The tree species is usually savanna-
fire, the degree of seasonal flooding, and local variation form Oregon white oak (Quercus garryanna breweri), but
in soils likely created a mosaic of vegetation and in some areas it may be ponderosa pine (Pinus ponderosa)
successional conditions within native grasslands. or Douglas-fir (Pseudotsuga menziesii).43, 103, 111
Perennial bunchgrasses dominated native sites,1 but exotic
annual grasses are prevalent in many remnant Chaparral
grasslands.43, 123 Some sites, particularly wet prairies, are Chaparral occurs in southwestern Oregon in the interior
rich in forbs. Vegetation height, density, cover, and species valley and foothills below 6,800 feet (2,100 m) in the Rogue
composition are dependent upon several factors including River basin and patchily northward into the Umpqua
soil type, aspect, climate, and land use and management. River basin (see Chapter 2—ceanothus-manzanita

261
 262 Wildlife–Habitat Relationships in Oregon and Washington

shrublands). Valley chaparral, a diluted extension of the by agriculture. Most grasslands with some resemblance
California chaparral types,33 occurs mostly below 3,600 feet to the historical condition are small pockets on tablelands
(1,100 m). A higher elevation montane chaparral with a north of Medford.
different species composition occurs between 3,600-6,800 Key factors that changed historical conditions in
feet (1,100-2,100 m) in the Siskiyou Mountains and westside grasslands were:
southern Cascades. Some chaparral communities may be 1. alteration of natural disturbance processes (fire
climax (e.g., wedgeleaf ceanothus [Ceanothus cuneatus] on and flooding), which shaped and maintained the
the Rogue Valley floor), and others are believed to be fire- pre-European structure of plant communities;
maintained. Chaparral consists of a shrubland with a 2. agricultural development (livestock grazing and
highly variable canopy and herbaceous understory, both cropland), which accelerated the conversion of
of which may depend on fire history. Dominant plants native grasslands; and
are multi-stemmed shrubs usually 1.6-13 feet (0.5-4 m) tall. 3. urbanization, which resulted in the irretrievable
Chaparral can either be a climax or a successional loss of native grasslands.
community. Regardless of the seral condition, a tree Fire suppression became significant in the mid-1800s1
component may be present. Although gradients exist when the expanding population became dominated more
between shrub- and tree-dominated communities, by Europeans than Native Americans. Before that, the
chaparral comprises the shrub-dominated end of this extent and persistence of native grasslands were closely
spectrum. tied to the regular disturbance of fire, either wildfire or
fires set by Native Americans.1, 14, 124, 130 Europeans deemed
Habitat Distribution and Impacts that fires were a threat to property. Fire suppression
allowed succession in fire-maintained grasslands,
Grasslands resulting in widespread shrub (including exotics) and tree
Before European settlement, the landscape of the (particularly Douglas-fir) encroachment into native
Willamette Valley, particularly south of Salem, was largely grasslands and savannas.1, 73 For example, in the South
an open expanse of prairie and savanna.43 An estimated Puget Lowlands, 32% of prairie loss has been attributed
1,030,604 acres (417,249 ha) of prairie and 539,938 acres to forest invasion and conversion.55
(218,598 ha) of savanna existed within the Willamette Agricultural development resulted in the loss and
Valley at the time of European settlement (E. Alverson, degradation of native grasslands from conversion to
The Nature Conservancy, pers. comm.). Among prairie cropland and livestock grazing, respectively. The effect of
habitats, approximately one-third, 301,472 acres (122,053 conversion to cropland was the immediate and permanent
ha), was wet prairie. Native prairie and savanna habitats loss of native prairie on a local scale. Until the early 1900s,
in the Willamette Valley have been reduced from being largely animal-dependent farming limited the size of areas
the most abundant vegetative communities, covering that could be cultivated. Thus, the landscape remained a
approximately 45% of the landscape (30% prairie and 15% mosaic of cultivated areas and native prairie habitats.
savanna), to a few small, scattered parcels of semi-natural Mechanical agriculture and fossil-fueled machinery
remnants amid farmland and urban and rural residential resulted in progressive conversion of native prairie and
development totaling approximately 2,000-3,000 acres savanna to agricultural land. By the mid-1970s, only tracts
(800-1,200 ha) or <1% of the historical extent (E. Alverson, too small, tracts otherwise poorly suited for cultivation,
The Nature Conservancy, pers. comm.). or tracts under some protection, were untouched. These
In the South Puget Lowlands, prairies were a less are among the few scattered areas that remain today as
significant part of the pre-European landscape than in the refugia for native prairie plants. Many savanna-form oaks
Willamette Valley. Assessment of pre-European vegetative also were lost in the conversion to cropland, and those
conditions based on soil type indicates 149,360 acres that remain are relicts in the midst of a sea of cultivation.66
(60,470 ha) of prairie habitat, or about 10% of the Livestock grazing has altered grassland habitat
landscape.27 Current estimates of prairie habitat are 12,582 primarily through degradation. However, moderate to
acres (5,094 ha), and most of this (91%, 11,500 acres [4,656 high levels of grazing can also result in the permanent
ha]) occurs on Fort Lewis Military Installation.27 Only 3% loss of the more palatable native grasses and forbs (E.
(2,993 acres [1,212 ha]) of the pre-European total is Alverson, The Nature Conservancy, pers. comm.). Nearly
considered “intact prairie” and again most of this (71%, all remnant grassland parcels not converted to cropland
2,130 acres [862 ha]) occurs on Fort Lewis. Thus, pre- have been grazed. Most grazing was open range until the
settlement prairie vegetation in the South Puget Lowlands late 1800s, when a shift to fenced pastureland occurred.66
has sustained losses >90%. Losses also have been extreme This practice, in conjunction with repeated disturbance
for grasslands and balds in the northern Puget Lowlands of soil and vegetative communities from cultivation,
(C. Chappell, Washington Natural Heritage Program, pers. importation of selected forage grasses and forbs, and the
comm.). seasonal movement of livestock between parcels, set the
In the Rogue and Umpqua valleys, no estimates of the stage for the invasion of exotic plants. Many remnant
extent of pre-European grassland habitat are available. prairie parcels are exotic species-dominated. Heavily
Most of these grasslands, and essentially all of them at grazed sites tend to have the highest proportion of exotic
the lowest elevations, are thought to have been modified species.123 Colonial bentgrass (Agrostus capillaris), sweet
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 263

vernalgrass (Anthoxanthum odoratum), Scot’s broom subdivisions and clearing associated with low density
(Cytisus scoparius), medusa-head wildrye (Elymus caput- rural development present a different threat to chaparral
medusae), and reed canarygrass (Phalaris arundinacea) are habitat. As development of private lands abuts public
prominent among exotic species present. Consequences lands, land managers are increasingly required to manage
of exotic plant invasion were the displacement or local lands to reduce fire hazard to adjacent residences.
extirpation of native plant species, which also may have Typically this involves manual removal of chaparral plants
disrupted food webs for wildlife. to minimize the likelihood of fire.
Attempts at quantifying prairie loss to agriculture are The chaparral community is adapted to recurring fire.
rare, but these losses are significant. In the South Puget Fire suppression inevitably will lead to profound changes
Lowlands, 30% of prairie losses have been attributed to in the distribution and abundance of plant species.10 For
agricultural uses.55 In addition, factors that reduced native example, the recruitment of new generations of shrubs
ungulate populations (e.g., habitat fragmentation [e.g., likely will be affected. The germination of many chaparral
fencing], hunting) altered successional relationships plants is triggered by the heat of fire or the chemicals found
between native browsers and grazers and grassland in smoke. Without wildfire, new recruitment is limited,
species. In Washington, declining recruitment of savanna especially among chaparral herbaceous plants, and as
oaks after 1870 is thought to be linked to heavy grazing1, 95 chaparral shrubs become decadent, their productivity is
Urbanization resulted in local and permanent losses of reduced. 11, 57 Given our limited knowledge of the
grassland,1 particularly in the South Puget Lowlands and functioning of this ecosystem along the northern margin
northern Willamette Valley. Beyond the near-complete loss of its range, many changes that might arise as a
of native grassland habitat, urbanization continues to consequence of fire suppression cannot be anticipated.
reduce the availability of agricultural grassland habitats.
This pattern is most common in the central and southern Wildlife Communities
Willamette Valley and central Rogue Valley, where urban Wildlife communities in grassland, and to a lesser extent
growth boundaries have extended into active and passive chaparral habitats, tend to be ecologically simple, with
agricultural lands, resulting in loss of habitat for several relatively low species diversity and richness. Within each
grassland-associated species that have been able to persist habitat type, the more uniform the vegetative structure,
in those areas.3 Attempts to quantify losses resulting from the lower the species diversity. As different structural
urban development are also rare, but the levels of loss components are added (e.g., shrubs in grassland, trees in
identified are significant. In the South Puget Lowlands, chaparral), the wildlife community becomes more diverse.
one-third of prairie losses were attributed to urban However, extensive encroachment of these structural
development.55 elements may make the habitat less suitable to its highly
associated species.
Chaparral
Numerous site-specific factors result in differences in
Little is known about the pre-European distribution of
vegetative composition within grassland and chaparral
chaparral habitat in the Rogue and Umpqua valleys.43 The
habitats. However, from a wildlife standpoint, coarse-level
only available current estimate has identified 38,412 acres
vegetative structural similarities exist that are linked to
(15,551 ha) of chaparral occurring in the Rogue and
wildlife use. The principal feature in grasslands is the
Umpqua river basins96 (T. O’Neil, Northwest Habitat
dominance of herbaceous vegetation, with low, but
Institute, pers. comm.). However, this figure is an
varying numbers of shrubs and/or scattered trees. The
underestimate because it excludes the montane subtype
principal feature in chaparral is the dominance of shrubs,
and successional stages of other communities that have a
interspersed with varying amounts of herbaceous
chaparral structure.
vegetation and hardwood or, less frequently, coniferous
Chaparral habitat has declined in extent primarily
trees. Differences in wildlife use within grassland and
because of conversion to agriculture and urban/residential
chaparral habitats are often related to differences in
development, and the quality of chaparral habitat has been
vegetation structure within physiognomic layers other
degraded because of fire suppression. Subsequent to
than the dominant layer. Additionally, microhabitat
European settlement, much chaparral was converted to
features, such as rocks or water, often account for the
pasture and cropland, particularly on the lower slopes and
presence/absence of many amphibians, reptiles, and small
valley bottoms. In the Rogue Valley, little chaparral
mammals.
remains on the valley floor. Steeper slopes have been less
Extensive loss of grasslands has resulted in a
affected. Where chaparral remains, fire suppression
fragmented habitat pattern that presents significant
probably has increased the predominance of dense, tall,
obstacles to the persistence of wildlife populations. Two
shrub stands instead of the more open, patchy structure.
important issues are (1) genetic isolation and dispersal,
Where grazing has been excluded, savannas or open
particularly for some amphibians, reptiles, and small
woodlands have developed a chaparral-like structure.1
mammals with limited mobility; and (2) the inability of
The latter would account for minor local increases in
small patches of habitat to meet the areal requirements of
chaparral habitat.
some birds, reptiles, and medium-sized to large mammals.
More recently, population growth in the Rogue Valley
Consequences of genetic isolation may be local losses of
has further encroached on chaparral habitat. New
genetic diversity and greatly increased susceptibility to
 264 Wildlife–Habitat Relationships in Oregon and Washington

local extirpation. Pronounced fragmentation may result agricultural grassland habitats. This is largely a
in habitat patches with varying amounts of a consequence of native grasslands having nearly
depauperated fauna, whose populations may have lost disappeared, and the fact that some wildlife species have
their genetic flexibility. been able to persist in agricultural grasslands.

Historical Knowledge and Trends Mammals. Most information on the mammal fauna is for
Information on wildlife communities in native habitats at medium-to-large mammals, and a significant portion of
the time of European settlement are sparse, and evidence it is based on hunting and trapping data. Suspected
of population status and trends is mostly anecdotal. population changes for most mammal species are based
Additionally, historical information on the vertebrate on an evaluation of potential conflicts between species’
fauna is disproportionate; greatest for large mammals and habitat requirements and types of habitat change, and our
birds, and almost nonexistent for small mammals and relatively limited knowledge about a species’ ability to
herpetofauna. Additionally, population trend information tolerate change.68, 90, 128
is available only for birds, and only for the last thirty years Mammals associated with native grasslands were
from the Breeding Bird Survey (BBS). Only recently have probably reduced with post-settlement habitat losses and
systematic studies been conducted on wildlife use of alteration, but suggestions of this pattern are based entirely
westside grassland and chaparral habitats (for example, on large mammals. For example, at the time of settlement,
see Case Histories). elk foraged in valley grasslands, but hunting and
Where data are unavailable on historical wildlife cultivation drove them into more secluded forested
populations, our knowledge of wildlife habitat areas.58, 119, 128 Declines in elk and other large mammals in
relationships is based on current conditions within these lowland grassland habitats undoubtedly reflected loss and
habitats. For example, in grasslands, an understanding of alteration of their requisite habitat. However, the pattern
wildlife species relationships with native habitats is partly is difficult to disentangle from concomitant spatial and
based on current knowledge of species relationships with temporal changes in burning, climate variation, forestry

Table 1. Bird species regularly associated with westside grassland and chaparral habitat with significantly
declining population trend estimates based on Breeding Bird Survey data (Sauer et al. 1997).

Species Habitat associationa BBS population trendb Comments

Grassland Chaparral 1966-98 1980-98

American kestrel x -7.3** -9.6**

Ring-necked pheasant x x -5.8*** -0.7 introduced species
Mourning dove x x -3.3*** -4.3***
Common nighthawk x x -6.8** 8.6
Vaux’s swift x -3.9* -1.8
Northern flicker x x -1.2* -0.6
Acorn woodpecker x x -1.8 -2.2*** Oregon only (i.e., westside)
Western wood-pewee x x -2.9*** -0.8
Ash-throated flycatcher x x -0.8 -3.5*** Rogue and Umpqua valleys only
Bushtit x -12.1*** -9.9***
Oak titmouse x -3.8* -2.2 Rogue valley only
Western bluebird x x -3.6** -7.7
House wren x -3.4 -6.8*
Wrentit x -0.9 -1.7*** Oregon only
Lazuli bunting x x -4.0** -0.6
Chipping sparrow x x -8.3*** -4.5**
Lark sparrow x -5.2** -4.2 Rogue and Umpqua valleys only
Song sparrow x -1.1*** -1.0**
Dark-eyed junco x -2.7*** -2.2**
Western meadowlark x x -7.0*** -4.9*
Brown-headed cowbird x x -2.8*** -1.8**
Brewer’s blackbird x x -3.3 -2.7**
Bullock’s oriole x -3.0** -2.8
American goldfinch x x -2.3* -1.6
a
Grassland includes dry and wet prairie and savanna; chaparral encompasses valley and montane subtypes; x = species association
with this habitat type.
b
Trend expressed as percent annual change. Only includes species that have occurred on •14 routes. Bold indicates significantly
declining trends. Significance values are * P <0.10, ** P <0.05, and *** P <0.01.
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 265

practices, and hunting. Although Columbian black-tailed

deer have been influenced by the same factors, they seem
significantly more tolerant to land conversion and human-
caused disturbances than elk.128
Birds. Populations of bird species associated with native
grasslands were undoubtably reduced to some extent with
post-settlement losses of native grassland habitats. Some
species such as the streaked horned lark, western
meadowlark, and Oregon vesper sparrow were apparently
able to persist in newly created agricultural grasslands
such as hayfields and pasture. These species were referred
to as “abundant or common” in the Willamette Valley and
Puget Lowlands by all authors from the time of European
settlement122 through the late 1800s and early 1900s5, 30, 65,
100
up until the mid-1900s.44, 54, 72 Since the late 1960s,
however, many westside grassland bird species have
experienced substantial population declines (Table 1). Of
the 44 species with either long-term (1966-98) or short- Western meadowlark. (Photograph by Richard B. Forbes)
term (1980-98) significantly declining trends in the BBS
Region encompassing westside grasslands and chaparral Valley BBS routes from a mean of approximately 11 birds/
(Southern Pacific Rainforest), 24 are regularly (closely and route in the early 1970s to approximately 2 birds/route in
generally) associated with grasslands and/or chaparral. the mid 1990s.
A few of these species also use forest habitats (e.g., western In contrast to declining species, only 16 species have
bluebird, Bewick’s wren), thus declines may be affected significantly increasing trends.106 Only 2 of those are
by populations in forest habitats. Several grassland species grassland associates (common yellowthroat and red-
that lack sufficient BBS data for analysis (e.g., short-eared winged blackbird in wet prairie), and 2 are chaparral
owl, northern harrier, streaked horned lark, Oregon vesper associates (California towhee and American robin). The
sparrow) are considered to be declining in western Oregon remaining 12 species are forest, riparian, or urban
and Washington based on anecdotal information. associates.
Statistically descriptive indications of population Concurrent trends in land use during the BBS time
declines can also be made for some grassland species based frame (1966-96) include extensive suburban and
on BBS relative abundance data. In the Willamette Valley, residential development, and conversion to row crop
western meadowlark abundance declined from a mean agriculture and large, intensively cultivated grass fields,
of approximately 13 birds/route (n = 11) in the early 1970s particularly in the Willamette Valley. Row crop agriculture
to <1 bird/route in the mid 1990s.4 The chipping sparrow, provides almost no habitat for grassland birds, and
a savanna-associate, declined in abundance on Willamette although cultivated grass fields (e.g., hayfields, annual and
perennial grass fields) can provide habitat for a few native
Table 2. The 12 most abundant species during point- species (e.g., savannah sparrow, western meadowlark),
count censusing in Willamette Valley agricultural they can also function as ecological traps when harvesting
grasslands, May-June, 1996.a aborts nesting attempts and reduces reproductive success.
Two broad patterns of change that have occurred in
contemporary grassland avifauna worldwide are declines
Species Birds/point count
in native and endemic species and increases in exotic
species.75 These patterns also are evident in the Willamette
Valley based on censusing of agricultural grassland
Savannah sparrow 1.62
European starling 1.11 habitats in 1996.3 Among the 12 most abundant species,
Brewer’s blackbird 0.80 only the savannah sparrow, the most abundant species, is
Red-winged blackbird 0.54 considered highly associated with native grassland habitat
Barn swallow 0.53 (Table 2). The second most abundant species, the European
American goldfinch 0.51 starling, is an exotic, and the third, fourth, and twelfth
Song sparrow 0.47 most abundant species (Brewer’s blackbird, red-winged
American robin 0.46 blackbird, and killdeer, respectively) are characteristic of
Common yellowthroat 0.39 agricultural lands. Of the remaining species, 3 swallows
Cliff swallow 0.37 (barn, cliff, and violet-green) are open-country aerial
Violet-green swallow 0.20 foragers, and 4 other species (American robin, American
Killdeer 0.18 goldfinch, song sparrow, and common yellowthroat) are
a
Based on detections within 328 feet (100 m) of 544 point-count relatively common edge or generalist species that use
stations visited once in May and once in June.3 grasslands opportunistically. Species highly associated
with native grasslands such as the western meadowlark,
 266 Wildlife–Habitat Relationships in Oregon and Washington

Table 3. Historical records of herpetofauna of the Southern Puget Trough outwash

prairies (modified from Leonard and Hallock83).

Species Selected Reproductive Occurrenceb

sources modea Historical Current

Northwestern salamander 79
Sb C C
Long-toed salamander 79
Sb C C
Rough-skinned newt 79
Sb C C
Western toad 107, 109
Sb C U
Pacific treefrog 79, 109
Sb C C
Northern red-legged frog 79, 109
Sb C C
Oregon spotted frog 109
Sb U R
Bullfrog 79, 109
Sb C C
Western pond turtle 109
El C X
Northern alligator lizard 109
Lb C C
Western fence lizard 109
El C X
Rubber boa 109
Lb U U
Racer 2
El R X
Gopher snake 122
El R X
Western terrestrial garter snake 79, 109
Lb C C
Northwestern garter snake 79, 109
Lb C C
Common garter snake 79, 109
Lb C C
a
Sb = stillwater breeding, El = egg laying, Lb = live bearing
b
C = common, U = uncommon, R = rare, X = functionally extirpated

Oregon vesper sparrow, and streaked horned lark were Current Populations and Habitat
uncommon to rare. Relationships
Herpetofauna. Leonard and Hallock 83 provided an A few mammal studies have been recently conducted in
overview of the historical information on herpetofauna westside grassland and chaparral habitats. Research on
in the South Puget Lowlands prairie landscape. Most the Western pocket gopher in the Puget Lowlands has
records are from museum collections and personal field emphasized biology and habitat use, 135 and species
notes of amateurs and professional biologists. These distribution relative to the degree of soil rockiness in
records date back approximately 150 years and provide prairies.115 Several studies have been conducted on the
at least qualitative documentation of the relative ecology of gray-tailed voles in the Willamette Valley,136, 137,
138
abundance of several species. It is noteworthy that several including effects of habitat fragmentation,139 mowing,34
species documented as occurring historically are now and pesticides.35, 94 An examination of the relationship of
extremely rare or extirpated (Table 3). Additionally, a small mammals and fire in wet prairie was conducted at
number of species seem to be much less abundant than Finley National Wildlife Refuge in the Willamette Valley
described in these historical accounts. (see Case Histories).
Historical data addressing grassland-associated Several recent bird community studies, which include
Willamette Valley herpetofauna were erratically collected, work done in the Willamette Valley,3 Umpqua Valley,28 and
both geographically and temporally. Early surveys Puget Lowlands 101, 104, 105 have examined habitat
provided distributional data from selected areas, e.g., relationships of grassland and savanna species.
Portland,70 Benton and Linn Counties,50 or anecdotal Additionally, species-specific studies include the blue-gray
information as part of larger regional compendiums,49, 52, gnatcatcher in the Rogue Valley 112 and the western
108, 125
but data on habitat relationships were generally bluebird in the Willamette Valley.36 Among the grassland
sparse (although see Graf 50). Because few historical studies, Altman3 focused on agricultural grasslands
grasslands remain, reduction or disappearance of at least because native grasslands are nearly extirpated in the
some of the herpetofauna characteristic of those systems Willamette Valley, and Rogers104 focused on a few of the
is strongly suspected. Historical data on grassland- remaining parcels of native prairie in the South Puget
associated herpetofauna for the Umpqua Valley are Lowlands. The three Puget Lowlands studies focused on
lacking. Some data on herpetofauna exist for the Rogue remnant grasslands on military installations at Fort Lewis
Valley,41 but only a few of these data apply to grassland- and McChord.
associated herpetofauna. Several recent surveys, some of which occurred on
grasslands, have attempted to assess the current status
and habitat relationships of herpetofauna in the South
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 267

Puget Lowlands.18, 56, 82 Most have focused on state-listed Umpqua valleys, whereas the Camas pocket gopher and
or federal candidate species such as the western pond gray-tailed vole are restricted to Willamette Valley
turtle42 and the peripherally grassland-associated Oregon grasslands. In a similar pattern, the broad-footed mole
spotted frog.47, 92 Several of these studies were conducted rather than Townsend’s mole is the more common mole
on remnant prairies found on military installations at Fort in grasslands in the Rogue Valley.
Lewis and McChord. Several additional insectivores and rodents are
Studies focused on grassland-associated herpetofauna common in westside grasslands, but are widespread in
in Oregon are limited (R. Goggans, Oregon Department distribution and occur in several habitat types. These
of Fish and Wildlife, pers. comm.). St. John116, 117, 118 provided include the creeping vole, Townsend’s vole, and Western
some habitat data on grassland herpetofauna in the Rogue, pocket gopher in western Oregon. The only large mammal
Umpqua, and Willamette valleys, respectively. that could be viewed as a grassland specialist, the
Additionally, there is an ongoing effort to characterize the pronghorn, was extirpated historically across the western
snake fauna of several habitats in the Willamette limit of its distribution in the Rogue Valley;7 evidence is
hydrographic basin, although only some of this effort lacking that it was once more widespread in westside
addresses grasslands or grassland-containing grasslands.
assemblage91 (R. Mason, Oregon State University, pers. Several groups of mammals, although frequent and
comm.). widespread in westside grasslands, lack representative
species specialized for westside grassland existence per
Grasslands: Mammals. The mammal fauna of westside
se. These include bats, carnivores, rabbits, shrews, and
grasslands includes a diverse array of approximately 55
selected ungulates (elk, deer). All species in these groups
species that occupy a variety of ecological niches (Table
either use grassland habitat only for foraging or require
4). This fauna includes a large suite of grazing, browsing,
some structural element not present in exclusively
or seed-eating species, represented by exposed
herbaceous grassland (e.g., shrub or tree structure, an
surface-active (deer, elk), vegetation tunneling (selected
aquatic habitat, or both). Shrews and most bats require
rabbits, voles), or burrowing forms (pocket gophers,
proximity to a mesic association or an aquatic habitat,
selected squirrels). The mammal fauna also includes a
whereas rabbits, many carnivores, and ungulates need
smaller group of aerial (bats) and terrestrial (shrews,
some woody structure for concealment. For example,
moles) insectivores, and a species rich and diverse group
black-tailed jackrabbits require the relatively open matrix
of carnivores. This fauna includes forms that range from
found in many grasslands to effectively use running as
being closely tied to grassland habitats (e.g., gray-tailed
an anti-predator strategy. However, they also require
vole, Camas pocket gopher), to generalized species that
scattered shrub clumps to conceal themselves following a
occur across a range of habitat types (e.g., bobcat, coyote,
predator-escape run. Additional species are common in
weasels, skunks).
habitats outside grasslands (oaks, woodland, or forest
Mammals most characteristic of westside grasslands
margins), but also may be abundant on grassland margins
are all small species dependent on an herbaceous structure
(e.g., California ground squirrel).
or the invertebrate fauna supported by that structure.
Some of the aforementioned patterns contribute to a
These include Camas pocket gopher, gray-tailed vole,
generally increasing species richness gradient among
Townsend’s mole, and local forms of the Western pocket
mammals with decreasing latitude (Table 4), a frequent
gopher in the Puget Lowlands. Regionally, some of these
pattern observed in nearly all faunal groups on a broader
species are replaced by ecologically equivalent but more
(global) scale. For example, of the 3 grassland-specialized
generalized species. For example, Botta’s pocket gopher
mammals previously discussed, only 1, Townsend’s mole,
and California vole occupy grasslands in the Rogue and
is widespread in Washington; another, Camas pocket
gopher, does not occur in the state; and the third,
gray-tailed vole, is only known from Clark County.
Grasslands: Birds. The breeding avifauna in westside
grasslands is characterized by relatively low species
diversity, and a unique assemblage of cryptically colored
ground nesters. Species diversity is greater in savanna
habitat, where several cavity nesters are associated with
the tree component.
Grassland birds have evolved several adaptations to
accommodate to life in an herbaceous, open landscape.
Most nest on the ground, and most nests are well
concealed in vegetation for protection from predators.
Some species, such as the western meadowlark, enhance
nest concealment by building a domed nest and a short
runway to the nest. Others, such as the burrowing owl,
Gray-tailed vole. (Photograph from W. Daniel Edge) nest underground in mammal burrows. Many passerines
developed behavioral adaptations to avoid nest predation,
 268 Wildlife–Habitat Relationships in Oregon and Washington

Table 4. Mammal distribution in westside grassland and chaparral habitats.

Grasslanda,b Chaparrala,b Grasslanda,b Chaparrala,b

P W U R U R P W U R U R

Striped skunk x x x x x x
Insectivores
Mountain lion x x x x x x
Fog shrewc pxd px
Bobcat x x x x x x
Vagrant shrew x x x x
Broad-footed mole x px Ungulates
Coast mole x x x x Elk x x x x x x
Townsend’s mole x x x x Black-tailed deer x x x x x x
Columbian white-tailed x x x
Bats
deer
California myotis x x x x px px
Pronghorn h
Long-eared myotis x x px x x x
Totals 38 46 43 42 31 33
Little brown myotisc x x px x
Long-legged myotis x x px px px x a
Grasslands include dry and wet prairie and savanna; chaparral
Yuma myotisc x x x x encompasses valley and montane subtypes (see text for details).
Hoary bat m m m x px px b
P = Puget Lowlands, W = Willamette Valley, U = Umpqua Valley,
Silver-haired bat x px px x x? x? R = Rogue Valley.
Big brown bat x x px x x x c
Species typically needs some kind of aquatic habitat beyond that
Townsend’s big-eared bat x x x x px x
found in seasonally wet prairie (e.g., pond, stream, marsh) and
Pallid bat m x x px x
thus, is only weakly linked to grasslands as defined here.
Brazilian free-tailed bat x x px? x?
d
x = known occurrence in the geographic areas indicated for
Rabbits each habitat type (p preceding the designation indicates probable
Brush rabbit x x x x x occurrence); m = present, but of marginal occurrence with a
Black-tailed jackrabbit x x x greater distribution outside the region; and h = historically
Rodents present, but functionally extirpated. Data are based on a
Siskiyou chipmunk x combination of Ingles68, Maser et al.90,Verts and Carraway128,
California ground squirrel x x x x x other sources, and personal observations.
Golden-mantled m x e
Data on natural range expansions and introduced populations
ground squirrel in Oregon are confusing (Verts and Carraway128).
Botta’s pocket gopher m x x px x f
Portions of what is known as Western pocket gopher is treated
Camas pocket gopher x as Mazama pocket gopher by some investigators.115
Western pocket gopherf m m x x x x
California kangaroo rat x including landing away from the nest and running
Deer mouse x x x x x x concealed along the ground to get to the nest, injury-
Pinon mouse x feigning distraction displays (killdeer), and acoustical
Dusky-footed woodrat x x x x x mimicry of rattlesnakes (juvenile burrowing owls).
California vole m x x x Grassland birds often have shorter incubation and nestling
Gray-tailed vole m x periods than other passerines, and young leave the nest
Long-tailed vole x m px x prior to being able to fly, but can run effectively along the
Creeping vole x x x x x x ground until capable of flight. Other ecological
Townsend’s vole x x x x adaptations unique to grassland birds with limited or no
Pacific jumping mousec x x x m perches available include flight songs (horned lark),
Common porcupine m m px x m x
singing from the ground (many species), and hovering in
Carnivores the air as an aerial perch for hunting (American kestrel).
Coyote x x x x x x Approximately 42 bird species are regularly associated
Gray wolf h h h h h h breeding species in westside grassland habitats (Table 5).
Gray fox x x x Species richness increases from wet prairie (16 species),
Red foxe x x x x to dry prairie (23), to savanna (27). Some species occur
Black bear x h x x x px only within 1 type of habitat along the grassland
Grizzly bear h h h h h h continuum (e.g., common snipe in wet prairie, acorn
Ringtail x x woodpecker in savanna). Others are equally associated
Raccoon x x x x x x with multiple grassland types (e.g., western meadowlark,
Ermine x x px px px x streaked horned lark). Wet prairie is characterized by
Long-tailed weasel x x px x x x species associated with mesic sites such as common snipe,
American badger h h
Western spotted skunk x x x x x x
Text continues after Table 5, on page 271
 Table 5. Breeding bird species regularly associated with grassland and chaparral habitats in western Oregon and Washington.a

Speciesb Statusc Guilds Grasslandd Chaparrald Grasslandse Chaparrale

Foragef Nestg Wet Prairie Dry Prairie Savanna Chaparral Chaparral Oak P W U R U R
Grassland Shrub Chaparral

White-tailed kite T G, C F x x x m m m
Northern harrier T G, C G X x x x m m
American kestrel T G, C C x x x x x x x
Red-tailed hawk T G, C F x x x x x x x x x x x
Wild turkey (I) R G, G G x x x m m x x x x
Ring-necked pheasant (I) R G, O G x x x x x x x x
Northern bobwhite (I) R G, G G x m m
Mountain quail R G, H G x x x x x
California quail R G, G G x x x x x x x x x
Mourning dove T G, G F x x x x x x x x x x
Sandhill crane T G,O G x h
Killdeer R G, I G x x x x x x x x x x
Western screech owl R G. C C x x x
Burrowing owl T G, C G X h h h
Short-eared owl T G, C G X x m m m m
Common snipe T G, O G X x x x x
Common nighthawk N A, I G x x m m m m m m
Common poorwill N A, I G x x m x
Vaux’s swift N A, I C x x x
Anna’s hummingbird T F, O F x x x x x
Lewis’ woodpecker T A, I C X x h h m m m m
Acorn woodpecker R B, G C x x m x x x x
Northern flicker R G, I C x x x x x x x x
Western wood-pewee N A, I F x x x x x x x x
Ash-throated flycatcher N A, I C x x x x x x
CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 269

Dusky flycatcher N A, I F X x m x
Western kingbird N A, I F x x x m m x x x x
Say’s phoebe N A, I C x h h h
Streaked horned lark R G, O G x X x x
Tree swallow N A, I C x x x x x
Violet-green swallow N A, I C x x x x x x x x
Western scrub jay R G, O F x x x x m x x x x x
Oak titmouse R F, O C X x x
Bushtit R F, I F x x x x
White-breasted nuthatch R B, I C x x h x x x x x
Bewick’s wren R G, I C x x x x x
House wren N F, I C x x x
Blue-gray gnatcatcher N F, I F X x
 270 Wildlife–Habitat Relationships in Oregon and Washington

a
The list includes only species that
regularly nest in grassland and
chaparral habitats, currently or
historically. Species that only forage

47
Chaparrale

x
x
x
x

x
x
x

x
x
x
x

x
x
x
x
x
x
x
in these habitats during the
U R

breeding season, such as rufous

43
m
m
x
x
x
x

x
x
x

x
x

x
x
x
x
x
x
hummingbird (wet prairie) and
rock dove (dry prairie, savanna),
some raptors and other species

39 41
m

m
that require specific structural
x
x

x
x

x
x

x
x
x

x x
x x
P W U R

elements for nesting, such as

Grasslandse

m
x
x

x
x

x
x
x
common raven and peregrine

35 41
falcon (cliffs) or barn swallows and
x
x

h
x
x

x
x

x
x

x
x

x
x
cliff swallows (outbuildings) are not
x
x

x
x

x
x

x
included.
b
I = Introduced species.
Chaparral

c
Status refers to the predominant

38
x
x
x

x
x
x

x
x

x
x

x
x
x
x
x

pattern for the majority of

Chaparral Oak

individuals of that species that

breed in western Oregon and
Chaparrald

Washington. It is based on

22
X

X
X
x
x

x
x

x
information presented in Puchy and
Shrub

Marshall99 with professional

opinion from others. Populations of
some species considered
Chaparral
Grassland

temperate migrants may be

27
X
x
x

x
x

x
x

x
x
x
x

x
x
resident in the Rogue Valley (e.g.,
mourning dove, American robin,
western bluebird, Anna’s
hummingbird, and American
goldfinch), but the majority of the
Wet Prairie Dry Prairie Savanna

27
X

population in westside grassland

x

x
x
x

x
x

and chaparral habitats is migratory.

R = resident; N = nearctic (long-
Grasslandd

distance) migrant; T = temperate

(short-distance) migrant
23
X

X
X
x
x

d
X = species has a high degree of
association with this habitat during
the breeding season (obligate or
16

semi-obligate) such that species

x

x
x

x
x
x
x

abundance is significantly higher in

this habitat than any other in
western Oregon and Washington; x
= species has a relatively strong
Foragef Nestg

—
G

G
G
G

G
G

G
G
G
C
C

F
F
F

F
F

F
F
F
F

association with this habitat during

Guilds

the breeding season; i.e., it is as

abundant in this habitat as others,
G, O

G, O
G, O

G, O

G, O
G, O
G, O
G, O
G, O
G, O

G, O

G, O
G, O
G, O
G, O

G, G
G, G
G, F
G,V

F, O
G, I

G, I

but it is not obligate to this habitat

F, I

F, I
F, I

F, I

for breeding.
x = regular breeding species in the
Statusc

geographic areas indicated for each

N

N
N

N
N

N
N
R

T
T
T
R

R
T
T
R
R
T

R
T
R

habitat type; m = marginal

Oregon vesper sparrow

presence or in low abundance as a

Brown-headed cowbird
Black-headed grosbeak
Common yellowthroat

Grasshopper sparrow

Western meadowlark
Red-winged blackbird

breeding species in the geographic

Green-tailed towhee
European starling (I)

American goldfinch
Brewer’s blackbird
Savannah sparrow

areas indicated for each habitat

California towhee

Chipping sparrow
Western bluebird

Dark-eyed junco
Spotted towhee

Lesser goldfinch
American robin

Bullock’s oriole

type (i.e., much greater distribution

Song sparrow
Lazuli bunting

Lark sparrow

Total Species
Fox sparrow

House finch

outside the geographic area); h =

historically present as a breeding
Wrentit
Speciesb

species, but believed to be

extirpated in the geographic areas
indicated for each habitat type.
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 271

f
First letter is predominant foraging zone during the breeding species rich than the 3 Oregon valleys (Table 5). Several
season: G = ground, A = aerial; F = foliage; B = bark. Second westside grassland or savanna species (i. e., white-tailed
letter is predominant food type during the breeding season: C = kite, acorn woodpecker, ash-throated flycatcher, lark
carnivore (vertebrates); F = frugivore (fruits); G = granivore sparrow, grasshopper sparrow, lesser goldfinch) reach the
(nuts, seeds); H = herbivore (plants); I = insectivore (insects); O northern limit of their geographic ranges in Oregon. There
= omnivore (plant and animal matter); V = vermivore (worms). are no northern grassland species that reach the southern
g
Nesting zones: G = ground (includes low shrub <1 m off limit of their breeding range in the Puget Lowlands,
ground); F = foliage (open-cup in shrub or tree >1 m off ground); although historically sandhill cranes maintained a small
C = cavity or mostly enclosed structure (e.g., ledge). breeding population in Puget Lowland prairies.72
The nonbreeding-season bird community in grasslands
differs considerably from that of the breeding season.
northern harrier, and short-eared owl. Dry prairie is
Species diversity can be highly variable, dependent on
characterized by species such as Oregon vesper sparrow,
several factors such as weather, field type and
burrowing owl, and grasshopper sparrow. Where a
management, fluctuating invertebrate populations, degree
scattered shrub component is able to persist in dry prairie,
of seasonal surface water, and timing of spring green-up.
species diversity may increase to include the lazuli bunting
Use is characterized by flocking species and aerial hunting
and American goldfinch, and species abundance may
behavior. Breeding species present in winter flocks include
increase for some obligate birds such as the Oregon vesper
western meadowlark, streaked horned lark, and killdeer,
sparrow and western meadowlark. The presence of
but the dominant avian presence is nonbreeding flocking
singular or small groves of scattered oaks, Douglas-fir, or
species such as Canada geese, sandhill cranes, American
ponderosa pine (savanna) results in an entirely new group
pipits, and several duck, gull, and shorebird species (e.g.,
of highly associated species such as American kestrel,
dunlin and dowitchers). There also is an influx of raptors
western bluebird, chipping sparrow, western kingbird,
that hunt for small mammals and birds in native and
and acorn woodpecker. Many of these species forage on
agricultural grasslands.
the ground or in the air above the grassland, but are
A few species of note occur in the coastal and dune
responding to availability of cavities for nesting (e.g.,
grassland and low shrubland types. Streaked horned lark
American kestrel, western bluebird), presence of acorns
breeds in coastal dune grasslands,110 which are also
as a food source (e.g., acorn woodpecker), elevated perches
important habitat for several wintering raptors and for
for singing (e.g., chipping sparrow), and/or the
small populations of migrant and wintering longspurs and
availability of branch structure for nesting substrate (e.g.,
snow buntings. The fox sparrow subspecies fuliginosa
western kingbird).
breeds in coastal headland shrublands north of Grays
As expected in an herbaceous vegetative community,
Harbor in Washington. 110 Yellow-rumped warblers
grassland bird assemblages are dominated by ground
concentrate in coastal dune shrublands in winter,
foragers (71%, n = 30) (Table 5). Nesting guilds are less
especially in California wax myrtle (Myrica californica) (C.
dominated by ground nesters (41%, n = 17), and more
Chappell, Washington Natural Heritage Program, pers.
equally represented by cavity nesters (29%, n = 12) and
comm.).
foliage nesters (27%, n = 11). However, if those species
exclusively associated with the tree component in savanna Grasslands: Herpetofauna. No amphibians and few
(n = 13) are not considered, ground foraging and ground reptiles can be considered grassland dependent; most use
nesting guilds become more dominant (89% and 57%, these habitats opportunistically. All are facultatively
respectively). It is in the savanna community that bird associated with grasslands through the presence of wet
species of other foraging and nesting guilds become more prairies and aquatic habitats within the grassland matrix.
representative. This includes foraging guilds such as aerial Facultative use of grasslands may occur for various aspects
insectivores (western kingbird) and granivores (acorn of a species life history, such as foraging, breeding,
woodpecker), and nesting guilds such as cavity nesters thermoregulation, and refuge/overwintering sites (Table
(American kestrel, western bluebird) and open-cup 6). In general, grassland habitat use by herpetofauna
foliage-nesters (chipping sparrow, western kingbird). requires the presence of at least one key feature on which
The composition of breeding bird species in westside these species are dependent, either within or next to
grasslands is predominantly migrants (64%, n = 27) (Table grasslands. For many amphibians, the key feature is
5). When distinguishing type of migrant (temperate vs. usually some type of surface water, but other species
nearctic) there is relatively equal representation among require rocks, sizable woody debris, or some kind of tree
resident (36%, n = 15), temperate migrant (26%, n = 15), canopy. Westside grasslands possess temporary pools
and nearctic migrant (29%, n = 13) species. Further similar to the more expansive grasslands in California, but
delineation excluding savanna-only species results in the amphibian fauna characteristic of the latter (e.g.,
similar increases in species richness from nearctic migrants spadefoot toads and tiger salamanders) is absent from the
(27%, n = 8) to residents (20%, n = 9) to temperate migrants grassland pools of the Pacific Northwest.
(43%, n = 13).
The most distinctive geographic pattern is that the Text continues after Table 6 on page 274
Puget Lowlands grassland breeding avifauna is less
 Table 6. Relationships of herpetofauna with grassland and chaparral habitats in western Oregon and Washington.

Species Reproductiona Foodb Refuge/Overwinteringc Grasslandsd Chaparrale Comments

WP DP S V M

Amphibians
Northwestern salamander E-P Stlw B: BR, ST L: Aq Mc, Inv, Am L L: AV, SS, WDs xf x x Ephemeral sites must have a 4.5
PM: Inv, Sl, Ea PM: MB, RC month or more hydroperiod
Long-toed salamander E-P Stlw B: VA L: Aq Mc, Inv L: SS, AV, WDs x x x x Ephemeral sites can have a 2.5-3
272

PM: Inv, Sl, Ea PM: MB, RC month hydroperiod

Rough-skinned newt P Stlw B: AV L: Aq Mc, Inv L: AV, SS, WDs x x x x x Oviposition typically on dense
PM: Inv, Sl, Ea PM: MB, RC submerged aquatic vegetation
Ensatina Terr MB, WD Co, Inv UB, WDt, MB, RC x x x Not in WA grasslands; prefers
woody debris
Black salamander Terr UN Co, Inv RK, MB?, RC? x Extreme SW OR only; open
shrubland near Ashland
Western toad P Stlw NB L: Al, Po, Oa, Car L: N, AV x x x x x Large woody debris may be
PM: Inv PM: MB, WDt preferred for overwintering
Pacific treefrog E-P Stlw B: VA L: Al, Oa, Car L: AV, SS x x x x x Ephemeral sites can have a 2.5-3
PM: Inv PM: WD, UB, LI month hydroperiod
Northern red-legged frog E-P Stlw B: EV, BR L: Al, Oa, Car L: AV, SS x x x Ephemeral sites must have a 4.5
PM: Inv, Sl, Ea, PM: TV, MB, PD month or more hydroperiod
Sm Am
Foothill yellow-legged frog P Flw B: RK, WDs L: Al, Oa, Car L: RS, WDs x x x x x Most frequently in medium-sized
PM: Inv, Sm Am PM: CM, O (3rd to 6th order) stream
Oregon spotted frog E-P Stlw NB L: Al, Oa, Car L: AV, SS PM: SP, O x x x Adult habitat P Stlw;
PM: Inv, Sm Am PM: SP, O reproductive site may be
ephemeral
Reptiles
Western pond turtle Egg SO Inv, Car, Aq Vg R: AV, SS, WD, RK xR xR x Active-season habitat in
O: LI, AV permanent water, but highly
variable in type
Long-nosed leopard lizard Egg SO Inv, Sm Lz R: SH x x SW OR only; western fringe of
O: MB? range; few occurrences known
Wildlife–Habitat Relationships in Oregon and Washington

Sagebrush lizard Egg SO Inv R: TV, SH, RK, MB x SW OR only; western fringe of
O: MB? range; fragmented occurrence
Western fence lizard Egg MB, SO Inv R: TV, SH, RK, MB x x x x Coastal Puget Sound non-
grassland in WA; in chaparral
O: MB, UB, O abundant; needs exposed
vertical structure for basking
Western skink Egg UR, MB Inv R: MB, RK, LI, WD x x x Prefers rocky habitat; Columbia
O: MB? Gorge non-grassland in WA
 Species Reproductiona Foodb Refuge/Overwinteringc Grasslandsd Chaparrale Comments
WP DP S V M

Northern alligator lizard Live Inv R: TV, SH, WD, RK x x x Present in low-elevation
O: UN grassland only in WA
Southern alligator lizard Egg MB, WD, SO Inv, NMa R: TV, SH, WD X X X Not in western WA grasslands;
O: MB, WD dense vegetation matrix key
Rubber boa Live NMa, Nb R: UB, WD x x x Needs sizable woody debris
O: WD, O
Ringneck snake Egg UR, WD Sm Am, Sl, Ea R: RK, WD, MB, LI x x Preferred refuges beneath well-
O: UN insolated rocks
Sharptail snake Egg WD Sl, Sm Sa R: WD, LI x x Associated with woody debris;
O: UN historic only in WA
Racer Egg MB Sm Lz, Be, Nb, R: TV, MB x x X Prefers a shrub-level matrix;
Sm Ma, Inv O: MB? historic only in WA
Striped whipsnake Egg UN Sm Lz, Be, Nb, R: TV, MB x SW OR only; western fringe of
Sm Ma O: MB? range; few occurrences known
Gopher snake Egg MB Sm Ma, Sm Lz, R: MB, TV, RK x X X x x Presumed extirpated in western
Be, Nb O: MB, O WA
Common kingsnake Egg MB Sm Lz, Sm Ma, R: MB, WD x X x Often near riparian corridors;
Be, Nb, Sm, Sn O: MB, O SW OR only
California mountain kingsnake Egg UN Sm Lz, Be, Nb, R: RK, WD, MB x In mixed decid-conif forest
Sm Ma, Sm, Sn O: UN or riparian; SW OR only
Pacific coast aquatic garter snake Live F, Sm Am, AmL R: RK, TV x x x SW OR only; fishing snake;
O: UN needs coarse substrate stream
Western terrestrial garter snake Live Sm Ma, Sm Am, R: TV, MB, WD x x x x x Most common along low, but
F, Sl, Nb O: ¡ººWD, O well-vegetated aquatic margins
Northwestern garter snake Live Sl, Ea, Sm Sa R: TV, LI, WD x x x Common where habitat favors
O: MB, O native slugs—primary prey
Common garter snake Live Sm Am, Am L, R: AV, TV, MB, WD x x x x x Most common along stillwater
F, Sl O: RK, O aquatic margins
CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 273

Western rattlesnake Live Sm Ma, Sm Lz R: RK, TV, MB, WD x x x x Most common in rocky talus or
O: RK, MB fractured rock areas
a

Aq Mc = aquatic
larva (L) and post-
debris (terrestrial).

categories are listed.

microorganisms (not

lizards, Sm Sa = small
birds, NMa = nestling
= egg laying, Live = Live

= branches (live woody

mammals, Sm Lz = small

snakes. Only major food

vegetation), AV = aquatic
bearing; Location (where

Sl = slugs, Sm Am = small
woody vegetation), UN =

metamorphic (PM) diet is

terrestrial amphibians are

of reptiles are considered

vegetation, MB = mammal
of decreasing importance.

microcrustaceans, Aq Vg =
Where multiple codes are

vegetation, EV = emergent

amphibians, Sm Ma = small
= Ephemeral, Flw = flowing

salamanders, Sm Sn = small
collectively: Food types: Al =
(submerged), WDt = woody

aquatic vegetation (non-algal

considered collectively. Post-
with a free-living larval stage,
debris, WDs = woody debris

forms), Be = bird eggs, Car =

water, P = Permanent, Stlw =
given, they are listed in order

excavation (not in burrow or

burrow, RK = rock, SO = soil

Food codes: For amphibians

partitioned; non-egg stages of

carrion, Co = springtails, Ea =

= invertebrates, Nb = nestling
earthworms, F = small fish, Inv

mammals, Oa = Other aquatic

under rock), ST = sticks (dead
known): B = oviposition brace,
Reproduction codes: Mode: E

NB = no oviposition brace, BR

= highly variable, WD = woody

stillwater, Terr = terrestrial, Egg

unknown, UR = under rock,VA

algae, Am L = amphibian larvae,

microcrustaceans), Po = pollen,

Notes continue on next page

hatching or post-partum stages
 274 Wildlife–Habitat Relationships in Oregon and Washington

c
Refuge/Overwintering site codes: For amphibians with a free-
living larval stage, larval and postmetamorphic sites are
partitioned; too little is known to partition refuge (active-season)
and overwintering sites categories; Notations for refuge (R)
versus overwintering (O) sites are partitioned; AV = aquatic
vegetation, CM = channel margin (instream), LI = litter (leaves),
MB = mammal burrow, N = none, O = other (this is noted
wherever other site type(s) besides the one(s) listed is(are)
suspected), PD = pond, RC = root channel (terrestrial), RK =
under rocks (terrestrial), RS = rocky substrate (aquatic only), SH
= shrub matrix, SP = spring, SS = soft substrate (aquatic only), TV
= terrestrial vegetation, UB = under bark, UN = unknown, WDT Southern alligator lizard. (Photograph by Richard B. Forbes)
= woody debris (terrestrial; not under bark of debris), WD =
woody debris, WDs = woody debris (submerged), ? = indicated fragmented distribution along their northern range
site type is suspected. margins, which may make them vulnerable to local
d
WP = wet prairie, DP = dry prairie, S = savanna,
extirpation. Puchy and Marshall99 viewed populations of
western rattlesnakes in the southern Willamette Valley as
e
V = valley, m = Montane relict because of their confinement to a few south-facing
f
Occurrence codes: x = present, X = strong association, xR = foothills and buttes. However, whether the western
used as reproductive habitat because active season habitat does rattlesnake was ever common on the valley floor is
not provide requisite conditions. unknown because of the few historical records.
Because of their terrestrial life histories, reptiles exhibit
stronger links to grassland habitats than amphibians.
However, only 2 reptiles, the gopher snake and southern
An evaluation of historical and current data for the alligator lizard, can be described as having an obligatory
Puget Lowlands listed 1 amphibian (Oregon spotted frog) grassland link; the others are best regarded as generally
and 3 reptiles (northwestern pond turtle, yellow-bellied associated with grasslands. The gopher snake has high
racer, and gopher snake) as obligate in prairie habitat.83 insolation requirements, which make it among the most
An obligatory association for the 3 reptiles is easily surface active of Pacific Northwest snakes;98 insolation
justified—all three lay eggs in terrestrial nests and need needs likely restrict the gopher snake to grasslands at
thermally adequate oviposition sites.15, 66 Moreover, the 2 higher latitudes. However, lack of burrowing mammals
snakes forage preferentially in upland habitat with a low can also exclude this species from grasslands. Burrowing
vegetative structure, and the turtle uses terrestrial mammals are not only prey, but also create most refuge
overwintering locations with Oregon white oaks.66 and oviposition sites for gopher snakes in a grassland
In contrast, justifying the Oregon spotted frog as a landscape. The other potential obligate, the southern
grassland obligate is difficult. This frog is a warmwater alligator lizard, favors a high-density herbaceous matrix,
marsh specialist,59, 60, 62 and its occurrence within grassland through which it literally “swims,” pushing off closely
habitat seems dependent on the presence of warmwater spaced stems with its small limbs.40 Two additional snakes,
marsh and appropriate overwintering sites (e.g., springs the racer and western terrestrial garter snake, seem to be
or selected flowing water channels). Moreover, available most common in grassland associations in the northern
data indicate that Oregon spotted frog use of upland parts of their ranges in the Pacific Northwest, also likely
habitats is not only limited, but its focal habitat, because of their insolation requirements. However, despite
warmwater marsh, may occur within matrices of different the fact that both have relatively diverse diets,113 their
upland habitat types without grassland habitat being highest densities are attained in grasslands with some
present. At best, the Oregon spotted frog should be viewed shrub structure and an aquatic margin, respectively. These
as only generally associated with grassland habitats. patterns may be related to favored prey achieving their
The Willamette Valley encompasses the northern limit highest densities in these grassland subtypes. The racer
(Pacific slope portion) of the main body of the geographic favors a scattered shrub matrix for foraging on eggs,
range for 10 species of reptiles (western pond turtle, nestling birds, and lizards; whereas the western terrestrial
western fence lizard, western skink, southern alligator garter snake robs the nests of voles.
lizard, ringneck snake, sharptail snake, racer, gopher Latitudinal patterns linked to insolation requirements
snake, western rattlesnake, and western terrestrial garter result in southwestern Oregon (Rogue and Umpqua
snake).97, 114 Most (8 of 10) are egg-layers and all are known valleys) having a greater species richness of reptiles than
or presumed to deposit eggs in a terrestrial nest. The only the Puget Lowlands (Table 7). Some egg-laying snakes
4 of these egg-laying reptiles recorded from the greater (e.g., kingsnakes) have their northern limits of distribution
Portland area (northern Willamete Valley) are known from in southwestern Oregon, whereas others (e.g., ringneck
fewer than a dozen records,84 and all were from well snake, sharptail snake) are geographically highly restricted
insolated, south-facing slopes that at least historically outside this region in the Pacific Northwest.41, 97, 113, 114
consisted of a grassland or Oregon white oak savanna Nonetheless, except for the gopher snake and southern
association. These reptiles seem to have a patchy or alligator lizard, all other reptiles occurring in Rogue and
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 275

Table 7. Herpetofaunal species distribution in westside grassland and chaparral habitats.

Grasslanda Chaparrala
Pb
W U R U R

Frogs and toadsc

Western toad md x x
Pacific treefrog x x x x x x
Northern red-legged frog x x x m m h
Foothill yellow-legged frog m x x x x
Bullfrog (E) x x x x x x
Oregon spotted frog x h
Salamanders
Northwestern salamanderc x x x
Long-toed salmanderc x x x x
Rough-skinned newtc x x x x x x
Ensatina x x x x x x
Black salamander x
Lizards
Long-nosed leopard lizard m
Sagebrush lizard m
Western fence lizard h x x x x x
Western skink m x x x x x
Northern alligator lizard x m m
Southern alligator lizard m x x x x
Snakes
Rubber boa x x x m x x
Ringneck snake x x x x x
Sharptail snake h x x x x x
Racer h x x x x x
Striped whipsnake m m
Gopher snake h x x x x x
Common kingsnake x x m m
California mountain kingsnake m x x
Pacific coast aquatic garter snake x x x x
Western terrestrial garter snake x x x x x x
Northwestern garter snake x x x m x x
Common garter snake x x x x x x
Western rattlesnake m x x x x
Turtles
Western pond turtle h x x x x x
Totals 20 23 24 25 24 26
a
Grasslands include dry and wet prairie and savanna; chaparral encompasses valley and montane subtypes (see text for details).
b
P = Puget Lowlands, W = Willamette Valley, U = Umpqua Valley, R = Rogue Valley.
c
Species typically needs some kind of aquatic habitat beyond that found in seasonally wet prairie (e.g., pond, stream, marsh) and thus,
is only weakly linked to grasslands as defined here.
d
x = known occurrence in the geographic areas indicated for each habitat type; m = present, but of marginal occurrence with a
greater distribution outside the region; and h = historically present, but functionally extirpated without or with few recent records.
Data are based on a combination of Corkran and Thoms26, Leonard and Hallock83, numerous other sources, and personal
observations.
 276 Wildlife–Habitat Relationships in Oregon and Washington

Umpqua grasslands require that some structural or habitat wrentit and lesser goldfinch, have a restricted distribution
element be present for these species to occur within a in the Pacific Northwest, but attain their highest densities
grassland landscape. For example, the western skink and in chaparral.
ring-necked snake are typically not found in grasslands Approximately 48 bird species regularly breed within
lacking rocks (J. Applegarth, Bureau of Land Management, chaparral habitats of southwestern Oregon (Table 5).
pers. comm.), because these seem to lack appropriate Species composition changes markedly as structure of the
thermal requirements for nest sites. Sharptail snakes, chaparral changes. Some species are limited to chaparral
specialized slug consumers, are typically not found in of a specific configuration, whereas others occur across
grasslands lacking a large woody matrix generator, the full range of habitat types. In general, bird species
typically oaks,25 probably because the slugs they consume, richness is lowest in chaparral shrub (Table 5), which lacks
mostly members of the tail-dropping slug genus trees and extensive herbaceous openings. Only 22 species
Prophysaon, are tied to this habitat element. regularly breed among wedgeleaf ceanothus and whiteleaf
manzanita (Arctostaphylos vicida) in chaparral shrub. Some
Chaparral: Mammals. Approximately 39 species of
of the most common species include the western scrub-
mammals occur in chaparral habitats of southwestern
jay, spotted towhee, California towhee, and lesser
Oregon (Table 4). Species richness is slightly less than in
goldfinch. Diversity is slightly higher if chaparral contains
grasslands. Few mammals are tied exclusively to the
substantial amounts of grassland. This feature adds
westside chaparral of the Rogue and Umpqua basins. The
species such as western meadowlark and lark sparrow.
best indicators of chaparral in Oregon are mammals that
Species richness climbs sharply on chaparral sites that
have a fundamental tie to the shrubby structure that
include hardwood trees. On the most mesic sites, which
characterizes this habitat. The California kangaroo rat,
include madrone (Arbutus menziesii) and oak, common
fairly common on brushy hillsides in the upper Rogue
species include mountain quail, oak titmice, white-
River Valley, is likely the best candidate, although its
breasted nuthatch, black-headed grosbeak, chipping
distribution covers only part of what may be characterized
sparrow, and dark-eyed junco. A number of factors
as chaparral in Oregon. Other candidates include the
contribute to this increase in diversity. Trees provide a
dusky-footed woodrat, which often builds stick nests in a
broader range of foraging opportunities because of
shrub matrix, and brush rabbit, usually no more than a
increased structural complexity and nesting opportunities
few feet away from a shrubby refuge.128 However, dusky-
for cavity users. A different group of birds uses high-
footed woodrats attain their highest densities where tree
elevation chaparral sites. Some of the more closely
cover is >90% and has not been fire-disturbed,19 and brush
associated species are the green-tailed towhee, fox
rabbits attain highest densities in more mesic coastal
sparrow, and dusky flycatcher.
sites,141 so both species seem to respond to habitat features
Migrant species represent a greater portion of the
that chaparral does not provide. Many other species occur
breeding bird species in chaparral (62%, n = 29) than
in varying abundance in chaparral, but are common only
residents (38%, n = 18) (Table 5). Among migrants, there
in the presence of selected trees (conifers: long-eared
are more nearctic migrant species (55%, n = 16) than
myotis, golden-mantled ground squirrel); riparian areas
temperate migrant species (45%, n = 13). Nearctic migrants
(several bats and shrews); or rocks or rocky outcrops
particularly contribute to the increased species richness
(California ground squirrel). One of the striking absences
in chaparral with hardwood trees. Nearly absent from
from Oregon chaparral is the brush or chaparral mouse,
treeless shrublands, nearctic migrants represent a
an indicator species for California chaparral,68 recorded
significant proportion of the bird fauna in chaparral with
within 2 miles (3.2 km) of the Oregon border.53, 128
hardwood trees, comprising 42% (n = 14) of the breeding
Chaparral: Birds. The chaparral habitat of southwestern species (Table 5).
Oregon harbors a unique assemblage of bird species, some Ground foragers dominate (58%, n = 27) chaparral bird
of which are found in few places elsewhere in the Pacific assemblages (Table 5). The percentage of ground foragers
Northwest. Shrublands, including the chaparral of declines with increasing site moisture due, in part, to
southwestern Oregon, are intermediate in structural additional bird species exploiting alternative foraging
complexity between grasslands and forest, and elsewhere opportunities. Bark foragers, foliage and twig gleaners,
have been demonstrated to support intermediate numbers and flycatchers all increase in species richness along the
of bird species.9, 37, 88 However, Pacific region chaparral moisture gradient. Among nesting guilds, foliage nesters
contains more bird species than many other shrublands, (41%, n = 19), cavity nesters (30%, n = 14), and ground
including the shrub-steppe community of eastern Oregon nesters (28%, n = 13) are roughly similar in number (Table
and Washington.133 5).
The bird assemblage of Oregon chaparral is similar to The winter bird community displays many similarities
that observed in chaparral to the south. 6 The most with the breeding bird community. Chaparral shrub
noticeable difference is the absence of the California contains the fewest species in both seasons. Likewise, more
thrasher in Oregon. Also, Bewick’s wren seems to be species occur along the interface with grasslands, and the
numerically more important in California chaparral, assemblage is richest in the presence of hardwoods. In
where it is one of the most frequently detected species in winter, several species leave the drier sites, but other
both chamise and mixed-chaparral. Two species, the migrants largely compensate for the loss, including the
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 277

golden-crowned and white-crowned sparrows from the snake and western skink); and Oregon white oak and
north as well as altitudinal migrants such as dark-eyed madrone (rubber boa, western skink, and common
juncos. Nearctic migrants leaving the mesic habitats are kingsnake). Three garter snakes (common, Pacific coast
replaced by fewer migrants in the winter (kinglets and aquatic, and western terrestrial garter snake) occur within
chickadees), resulting in a net loss of about 3-8 species. In chaparral largely because the requisite aquatic habitat
winter, bird densities are greatest near the valley floor and (stillwater or stream) occurs within the chaparral habitat
decrease with increasing elevation. matrix. How these species use upland habitat such as
Somewhat surprisingly, foraging guild composition chaparral is not well known, but the most significant use
changes relatively little seasonally. Ground foragers is probably for overwintering or short-term retreat sites.66
dominate in both seasons and show the same relative Both kingsnakes seem to be more common in chaparral
decline with increasing moisture. In winter, the decline is where a riparian zone exists, which likely reflects more
both absolute as well as relative. The contribution of suitable levels of prey; this probably also applies to the
foliage and twig gleaners also changes little on a seasonal western skink and northern alligator lizard.
basis. Changes that do occur include the disappearance In the Pacific Northwest, no amphibian species exhibits
of flycatchers in winter, and increases in bark foragers and strong ties to chaparral. The ensatina salamander is the
drillers. amphibian most likely to be found in chaparral, especially
Besides their importance to breeding and wintering where oaks are present. However, this species is
birds, chaparral habitats are important to migrants, significantly more abundant in more mesic westside
especially in spring. Species such as the warbling vireo, associations. Black salamanders have been recorded
and yellow, Wilson’s, MacGillivray’s, and yellow-rumped patchily from chaparral-like habitats on the north slopes
warblers stop to feed before continuing north to breed in of the Siskiyou Mountains, but this species also has been
other habitats. Northbound migrants typically travel most frequently recorded along permanent riparian
through the lower elevations, taking advantage of the corridors in the region (D. Clayton, Rogue River National
transient abundance of canopy-dwelling arthropods in Forest, pers. comm.). Remaining amphibians that occur
chaparral shrubs and hardwoods. In late April, wedgeleaf within the chaparral landscape are there because the
ceanothus experiences a brief flush of arthropods that aquatic habitat they require is present; their use of the
drops quickly to very low levels for the rest of the year. upland with a chaparral structure is likely restricted to
Oregon white oak experiences a similar pulse at about the seasonally important refuge sites. This applies mostly to
same time, but arthropod volume does not fall as the Pacific chorus (tree) frog, foothill yellow-legged frog,
precipitously and persists at a moderate level into the fall. and rough-skinned newt.
The ephemeral abundance of arthropods also helps
explain the relatively low numbers of foliage gleaners Endemics, Extirpations, and
breeding in chaparral, particularly in chaparral lacking Peripheral Species
hardwoods. Wildlife species population patterns in westside grassland
and chaparral habitats reveal several features:
Chaparral: Herpetofauna. The herpetofauna of chaparral
1. losses are prominent, but gains have been rare,
habitats includes a diverse array of 24 species, including
2. losses have been greater in grasslands than in
4 frogs, 4 salamanders, 3 lizards, and 13 snakes (Table 6).
chaparral, which reflects the relative magnitude of
No reptile or amphibian species can be considered
change in each habitat type,
exclusively tied to chaparral. The western whiptail,
3. no species inhabiting grasslands or chaparral has
perhaps the reptile species known to be most strongly
become extinct, but a number of species have been
linked to chaparral in California,114 does not occur in
extirpated over large areas, a pattern especially
chaparral in southwestern Oregon, although it occurs in
pronounced in grasslands,
a large area of high desert shrub-steppe in eastern
4. regional endemics and species that are
Oregon.97 However, among the habitat associations found
distributionally peripheral or naturally rare seem
in southwestern Oregon, the western fence lizard and
to have sustained the greatest losses, and
southern alligator lizard reach their highest densities in
5. a few species have expanded their ranges in
chaparral, especially wedgeleaf ceanothus- dominated
westside grasslands.
areas. Chaparral shrubs, especially in combination with
rocks, provide the complex, vertical structure for foraging, Grasslands: Mammals. The Camas pocket gopher and
and elevated vantage sites for predator-detection gray-tailed vole are endemic to the Willamette Valley sensu
characteristic of western fence lizard habitat.87 Wedgeleaf lato.128 The gray-tailed vole formerly was associated with
ceanothus also provides the dense stem matrix within prairie habitats, and now occurs in agricultural grasslands,
which the southern alligator lizard can forage and particularly pastures, hayfields, grass-seed fields, and
maneuver to escape predators.40 The sharptail snake and grain fields. The Camas pocket gopher occurs primarily
western rattlesnake may also attain their highest densities in cultivated cropland such as grain fields. A well-drained
in wedgeleaf ceanothus chaparral, but data on variation field and the presence of plants with reserves stored in
in snake assemblages in southwestern Oregon are lacking. bulbs or tap roots characterizes suitable habitat.128
Several reptiles occur or are more common in chaparral Local forms of the Western pocket gopher are declining
if selected structural elements are present: rocks (ringneck in the Puget Lowlands,115 and this species is patchily
 278 Wildlife–Habitat Relationships in Oregon and Washington

distributed in westside grasslands. In the Puget Lowlands, At the time of European settlement, the streaked horned
Western pocket gopher occurrence is negatively associated lark was a “very abundant summer resident on gravelly
with increasing levels of rocky soils.115 prairies of Puget Sound”.122 In the late 1800s, it was “a
At the time of European settlement, gray wolves rather common summer resident” in Washington County,
occurred in low densities throughout westside OR;5 in the early 1900s, “common resident in western
grasslands,17, 122, 128 grizzly bears were present in grasslands Oregon and southwestern Washington;”65 in the 1930s, “a
in western Oregon,128 and the pronghorn, a species with a common breeding bird of the open fields in western
strong grassland tie, was present in the upper Rogue Oregon;” 44 and in the 1940s, “common permanent
Valley.7 Disappearance of grizzly bears, gray wolves, and resident” in the southern Willamette Valley.54 Currently, it
pronghorn from westside grasslands occurred shortly after is rare to locally uncommon, with scattered small
settlement, probably a function of hunting and/or populations in the Willamette Valley, the largest breeding
incompatability with humans, rather than habitat population occurring on and north of Baskett Slough
alteration, because most habitat alteration was subsequent National Wildlife Refuge.3 Breeding habitat is prairie and
to their disappearance.90, 128 most types of agricultural grasslands where the vegetation
Black-tailed jackrabbits have declined substantially in is short, <1 ft (<0.3 m), with patches of bare or sparsely
the Willamette Valley. 89 Bailey 7 reported them as vegetated ground.3 In Washington, it has been extirpated
“commonly reaching as far north as the country about from the San Juan Islands and much of the Puget
Salem, and more rarely to the Columbia River.” On one Lowlands.85 A few scattered breeding populations exist
day in November in 1930, he observed 9 road-killed black- in the South Puget Lowlands, the largest on prairies at
tailed jackrabbits on a trip between Salem and Eugene. In Fort Lewis Military Installation.104 A few pairs also occur
addition to loss of grassland habitat, sport hunting and in coastal dune habitats on the southern Washington
disease may have adversely impacted this species.119 A few coast 110 (C. Chappell, Washington Natural Heritage
were seen in the 1970s,128 and jackrabbits were seen at Program, pers. comm.).
approximately 15 locations from 1996 to 1999 (BA; MH). Several bird species have been extirpated as breeding
Most sightings were in Christmas tree farms, and occurred species from all or some westside grasslands since
throughout the valley south of Portland. European settlement. The burrowing owl was considered
Columbian white-tailed deer occurred throughout “a familiar sight”44 and a “common breeding species”16 in
grasslands of the Willamette and Umpqua valleys in the 1920s and 1930s in the Rogue Valley, and may have
Oregon at the time of European settlement.128 They now nested in the Willamette Valley50 and Puget Lowlands.110
are restricted to remnant populations in Umpqua Valley The last suspected nesting in westside grasslands was in
near Roseburg, and a few sites in bottomland forests and the early 1980s (O. Swisher, retired, pers. comm.). An
agricultural grasslands along the Lower Columbia River.46 experimental reintroduction program in the Rogue Valley
This form is federally endangered, although the Douglas in the mid 1980s was unsuccessful.89 Burrowing owls
County populations are proposed for delisting (D. occasionally winter in grassland habitats in the Willamette,
Peterson, U.S. Fish and Wildlife Service, pers. comm.). Loss Rogue, and Umpqua valleys.
of habitat, increased human density, and hunting have Lewis’ woodpeckers formerly nested in oak savannas
reduced use of grassland habitats by Columbian white- of the Puget Lowlands72 and Willamette Valley.44 Prior to
tailed deer. 1965, they were fairly common residents in Columbia
River bottomlands,45 and nested in a few oak savanna
Grasslands: Birds. Two subspecies of birds, the Oregon
locations in the Willamette Valley (e.g., Finley National
vesper sparrow and streaked horned lark, are endemic to
Wildlife Refuge) into the early 1970s (A. Contreras,
westside grasslands. At the time of European settlement,
University of Oregon, pers. comm.). A drastic reduction
the Oregon vesper sparrow was “rather abundant on the
in the population occurred in the late 1960s and early
Nisqually plains, Puget Sound”.122 In the late 1800s, it was
1970s, and the last documented breeding record in the
“abundant summer resident, found everywhere in open
Willamette Valley was in 1977 near Scapoose along the
country” in Washington County, Oregon;5 in the early
Columbia River.48 They have been extirpated primarily
1900s, “fairly common on the prairies and grassy fields of
because of (1) competition with the European starling for
western Oregon and Washington;” 65 in the 1930s,
nest cavities since the arrival of that species in Oregon in
“abundant in Willamette Valley native and agricultural
the 1950s and its proliferation in the 1960s, and (2) loss of
grasslands;”44 in the 1940s, “common summer resident”
large savanna-form oaks to development and agriculture.
in the southern Willamette Valley;54 and in the early 1950s,
Lewis’ woodpeckers may nest infrequently in Rogue and
“numerous about pastures and prairies of the Puget Sound
Umpqua valley savanna habitats, but have declined in
Region.”72 Currently, it is rare to locally uncommon in
those areas also.89 It occurs as a regular wintering species
widely scattered areas of the Willamette Valley.3 In the
in the Rogue Valley, and as an occasional migrant and
Puget Lowlands, it regularly occurs only in the San Juan
wintering bird in the other valleys.
Islands85 and on prairies in the South Puget Lowlands,
The common nighthawk was “very abundant on the
especially at Fort Lewis Military Installation.104 In the
prairies near Puget Sound,”122 and nested in grasslands in
Willamette Valley, it occurs almost exclusively in light- to
the Willamette Valley,44 but may be extirpated there.3 The
moderately-grazed pastures and weedy Christmas tree
loss of nesting nighthawks may have resulted from
farms in the valley foothills.3
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 279

reductions in the insect prey base, and increases in However, 2 frogs with a peripheral link to grasslands have
populations of urban and rural predators, such as the experienced significant regional extirpation. The Oregon
American crow. Nests have been recently reported in spotted frog is thought to be extirpated on the Willamette
Christmas tree farms and clearcuts in the foothills of the Valley floor and has been nearly extirpated in the Puget
Willamette Valley.3 Lowlands, where only 2 populations about 10 miles (16
Two species, Say’s phoebe and the sandhill crane, km) apart are known to remain in the Black River
occurred in small local breeding populations in westside watershed in Thurston County 83, 92 (K. McAllister,
grasslands, but disappeared as breeding species many Washington Department of Fish and Wildlife, pers.
years ago. Say’s phoebe was considered a “regular resident comm.). Surveys for the Oregon spotted frog conducted
of Rogue, Umpqua, and Willamette valleys.” 44 It in 1993-94 failed to detect the species at any of the 10
maintained a small breeding population near Corvallis at verifiable historical sites on the Willamette Valley floor,60,
least through the 1940s,39 but has not been reported as a 62
and surveys of historical sites in Washington have
nesting species since. Sandhill cranes formerly were a similarly failed to detect the species.92 Similarly, the foothill
“summer resident in the prairies of western yellow-legged frog is thought to be nearly extirpated in
Washington.”72 At the time of European settlement they the Willamette Valley, as the species was found at only 1
were considered a “common summer resident.”122 of 10 verifiable historical sites.13 Both species are presumed
The lark sparrow has been extirpated as a breeding to have declined because of changes in their aquatic
species in the Willamette Valley, but still maintains habitats, rather than changes in the associated grassland
breeding populations in the Rogue and Umpqua valleys. matrix.
Shortly after European settlement, lark sparrows were Three of the 4 egg-laying reptiles (western pond turtle,
reported as “common breeders in the Willamette Valley,”74 racer, and gopher snake) that occurred historically in the
and into the early 1900s were considered an “uncommon South Puget Lowland prairies are presumed extirpated
summer resident at Corvallis and Dayton.”140 It has not (Table 3). Confusion exists regarding the fourth species,
been reported as a breeding species in the Willamette the western fence lizard. Leonard and Hallock83 listed it
Valley for approximately 50 years. as historically common, but indicated that the association
One obligate grassland species, the grasshopper of this species with prairie appears to be in error. Moreover,
sparrow, is considered naturally rare in westside current reports may involve misidentifications.
grasslands. It maintains a small, tenuous breeding Uncertainty about the historical distribution arising from
presence in the Willamette, Umpqua, and Rogue valleys. comments by Suckley and Cooper122 make it best to
This inconspicuous bird was first reported in the Rogue conservatively interpret western fence lizard as
Valley in 1963.102 It has been reported periodically since functionally extirpated (K. McAllister, Washington
then, but only a few birds at most. It was not recorded in Department of Fish and Wildlife, pers. comm.). Regardless
the Willamette Valley until the early 1970s.93 Generally, 1- of their historical abundance, all egg-laying grassland
3 birds have been reported most years since, until an reptiles in the South Puget Lowlands may be extirpated.
extensive study in 1996-97 reported 20-25 singing males, Based on knowledge of the magnitude of habitat
mostly on private pasture lands.3 Habitat in the Willamette change on the Willamette Valley floor, selected reptiles
Valley includes relatively short grass (<18 in [<0.5 m]) may have sustained major reductions in geographic range
pasture lands with little to no shrub cover.3 in this region. The western terrestrial garter snake that
The white-tailed kite and acorn woodpecker are species occurs in the Willamette Valley is a morphologically
that have expanded their breeding range in westside distinct variant that currently lacks taxonomic recognition
savanna habitats. The acorn woodpecker has always been (A. St. John, Independent consultant, pers. comm.). This
a common to uncommon species in oak savanna of the snake occurred across the southern half of the Willamette
Rogue and Umpqua valleys, but has expanded its range Valley with a narrow extension north along the toe of the
in the Willamette Valley in the last 50 years.48 Prior to 1940, Coast Range to the vicinity of Forest Grove.97, 114 It is now
there were only a few records of this species near Eugene,39, uncommon in the southern Willamette and has not been
44
but now it occurs north to Portland. Its distribution in encountered at the northern end of the western extension
the Willamette Valley is spotty, and it is less abundant despite extensive searches (J. Applegarth, Bureau of Land
further north. It does not occur in the Puget Lowlands. Management, pers. comm.; A. St. John, Independent
Only a few incidental nonbreeding season records of consultant, pers. comm.).
white-tailed kite existed in westside grasslands prior to
Chaparral: Mammals. Only 1 mammal, the California
the 1960s and 1970s.48 Sightings have increased since then,
kangaroo rat, may be viewed as endemic to chaparral in
and kites are now a regularly occurring wintering species
Oregon, and no mammals known to occur in Oregon
throughout westside valley grassland habitats. They are
chaparral are extinct. However, small mammal surveys
also a rare breeding species in grasslands of all of the
focused in Oregon chaparral to detect endemics like the
interior valleys of Oregon.48
brush mouse have not been conducted.
Grasslands: Herpetofauna. No amphibian or reptile
Chaparral: Birds. Three bird species are endemic or near-
species that occurs in westside grasslands in Oregon and
endemic to chaparral habitats of southwestern Oregon.
Washington is endemic to this habitat, and no species is
All have a broader geographic distribution in California.
known to be completely extirpated from this habitat type.
 280 Wildlife–Habitat Relationships in Oregon and Washington

The blue-gray gnatcatcher occurs in the zone where widespread in the upper Rogue; thus, they can be
Oregon white oak and chaparral shrubs coexist. It rarely considered distributionally peripheral and naturally rare.
occurs in chaparral without oaks, and is not encountered The black salamander is distributionally the most
in oak savanna lacking wedgeleaf ceanothus. Since the peripheral chaparral-associated species, occurring only
first reported breeding in 1963,102 blue-gray gnatcatchers across a small area of the Siskiyou Mountains of
have been found at scattered chaparral sites throughout southwestern Oregon. However, recent surveys show it
Jackson and Josephine counties. Densities are relatively to be somewhat more widespread than previously thought
low as might be expected for a peripheral population. (D. Clayton, Rogue National Forest, pers. comm.).
Between 1.4 and 2.7 individuals/25 ac (10 ha) have been
recorded on Lower Table Rock (SJ). Of 8 nests monitored Exotic Species
by Speer and Felker,112 one fledged a brown-headed Several introduced species of wildlife maintain
cowbird and 3 successfully fledged young gnatcatchers. populations in westside grassland and chaparral habitats.
Nests are constructed on a horizontal limb usually in an At least 8 introduced vertebrates occur in these habitats
oak, and birds forage in both oak and ceanothus. (Virginia opossum, eastern cottontail, house mouse,
Oak titmice have similar habitat requirements and a Norway rat, wild turkey, ring-necked pheasant, northern
similar range to the blue-gray gnatcatcher. Being cavity bobwhite, and European starling), and at least 4 other
nesters, they are further restricted to habitat containing introduced vertebrates occur in aquatic habitats to which
mature Oregon white oaks with dead limbs and enough grasslands or chaparral are peripherally linked (nutria,
rot to permit excavation of a nest site. Densities of between bullfrog, common snapping turtle, and red-eared slider).
4.1-7.3 individuals/25 ac (10 ha) were recorded on Lower Additionally, ambiguous data suggest that the Willamette
Table Rock (SJ). Titmice display a broad range of foraging Valley—Puget Lowland populations of red fox were
activities. Not only do they forage among both oaks and introduced around the turn of the century.128 Some of these
ceanothus, but they also regularly feed on the ground. exotic species are well established in grassland and
Another species endemic to chaparral is the California chaparral habitats, but others, (i.e., house mouse and
towhee. Unlike the blue-gray gnatcatcher and oak Norway rat) are only rarely dissociated from human
titmouse, it occurs over a broad range of chaparral habitats dwellings and highly modified human habitats into
and as far north as the Umpqua Valley. The California adjacent grasslands or chaparral. Data regarding whether
towhee is a terrestrial forager that forages in openings exotic species impact native species vary greatly. Evidence
between shrubs, usually ceanothus or manzanita. Despite of negative impacts exist, and positive effects have not
feeding in more open areas than the spotted towhee, it is been found. Most data suggesting some kind of exotic
seldom found far from dense cover. species effects are confounded with alternative factors that
have potential to influence native species.20, 64
Chaparral: Herpetofauna. No amphibians or reptiles are The Virginia opossum, first introduced to Oregon
endemic to chaparral habitats of southwest Oregon, and between 1910 and 1921,71 is now established throughout
no amphibians or reptiles recorded from Oregon chaparral western Oregon and Washington. 128 Opossums are
habitats are extinct. Several infrequently observed species characterized as highly opportunistic carnivores,67 but
of snakes (California mountain kingsnake, common seem to be too slow to be efficient predators on even
kingsnake, ringneck snake, and sharptail snake) that occur moderately mobile prey. However, immobile or poorly
in chaparral are thought to have a restricted distribution mobile prey, including bird’s eggs and nestlings and
in Oregon. Although these snakes are generally immature and nursing small mammals are vulnerable.
infrequently encountered and may be considered rare (e.g., Because opossum diet in western Oregon and Washington
sharptail snake), 89 the extent of their distribution is has not been evaluated, the impact of this exotic on
unknown because few systematic surveys have been populations of ground nesting birds and mammals is
conducted when these species are near-surface active. unknown.
Moreover, recent surveys in the Umpqua Basin suggest The eastern cottontail was introduced into Benton and
that the sharptail snake has a much broader altitudinal Linn counties in 1937 and 1941 from Ohio and Illinois,
range than previously thought61, 63 (R. Hoyer, retired, pers. respectively.51 From these introductions, it has spread at
comm.). Since similar biases probably exist with the least through the mid-Willamette Valley.127, 129 The source
remainder of these rarely observed snakes, their status will of eastern cottontails in the northern Willamette Valley
remain ambiguous until enough surveys are completed. and Puget Lowlands is unknown, but they may be
Regardless of how abundant they may be, these snakes descendants from Missouri stock originally introduced in
are not restricted to chaparral habitats. 1933 near Battle Ground, Washington.29 Chapman and
Three species, long-nosed leopard lizard, sagebrush Verts21 provided evidence that eastern cottontails were
lizard, and striped whipsnake, have been recorded from behaviorally dominant over native brush rabbits. Verts and
few localities in southwestern Oregon. 77, 97 These are Carraway126 offered evidence that eastern cottontails and
members of the Great Basin herpetofauna that reach the brush rabbits hybridize under natural conditions at least
western fringe of their geographic ranges in the upper occasionally. However, Chapman and Trethewey 20
Rogue Valley. Distributional information on these species indicated that the ecological and economic impacts of the
is lacking, but it is unlikely that any of these species are introduction of eastern cottontails into Oregon were
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 281

unknown. Despite available evidence, the expansion of in local declines in ring-necked pheasants. However, the
eastern cottontails in the Willamette Valley, and ring-necked pheasant remains a visible part of the fauna,
contraction of brush rabbit distribution in the same region particularly in the agricultural grasslands of western
has not been demonstrated to have a direct connection. Oregon. No data exist regarding interactions between ring-
The date of introduction of the house mouse to Oregon necked pheasant and the native fauna in westside
and Washington is not known, but it was probably early grasslands.
considering its near ubiquitous association with humans. The northern bobwhite was introduced to agricultural
The infrequent capture of house mice in the field, and their areas of Oregon during 1900-1960, but has been largely
almost uniform occurrence near human habitation extirpated from most areas where it once seemed to have
suggests that populations in open fields are typically been established, such as the Willamette Valley. 48
temporary, near human structures, and limited to warmer Nonetheless, a few local sightings continue, some of which
and drier seasons.128 The apparent failure of house mice are attributable to escaped birds used in dog training,
to become established in the field in the Pacific Northwest private introductions, and in some cases, populations that
may be because native voles interfere with their have persisted for decades without reintroduction.
reproduction, 12 recruitment, 32 or a combination of Attempts to introduce the European starling to the
population processes.86 Pacific Northwest were first made at Portland, Oregon in
The Norway rat, like the house mouse a native of China 1889 and 1892.81 Both of these early efforts resulted in brief
and Siberia, is almost ubiquitous in its association with establishment, but eventually failed in 1901 or 1902.48 The
humans.128 The first record of the species in the Pacific eventual permanent introduction can be traced to Eugene
Northwest was an individual collected at Astoria by Scheifflin, a New York drug manufacturer whose bird and
Lieutenant Trowbridge in 1855.8 The Norway rat nearly Shakespeare hobbies engrossed him. Scheifflin introduced
always resides close to human dwellings, but in California, starlings in New York in 1906, where they began nesting
noncommensal populations of Norway rats occurred in the same year. The species spread like a wave westward
riparian woodlands and adjacent fallow agricultural fields and by 1928, was usually spoken of as anywhere east of
several miles from human habitation.121 As Norway rats the Mississippi.81 The first starlings recorded in the
have been trapped in agricultural and fallow lands in the westside Pacific Northwest were at Eugene and Meadow
Willamette Valley, noncommensal populations may also View (both Lane County, Oregon) on 26 December 1947.48
occur there. Nonetheless, data are lacking regarding Today, the starling is common to abundant year-round in
interactions between Norway rats and native small urban areas, towns, and agricultural lands through the
mammals in westside grasslands. westside Pacific Northwest. Starlings are aggressive cavity
At least 3 grassland and/or chaparral bird species were nesters and compete directly with many native birds for a
introduced as game species: the wild turkey, ring-necked limited cavity supply.48 Three westside grassland species
pheasant, and northern bobwhite. Wild turkey, introduced are thought to have declined due at least in part to the
to the Pacific Northwest just after 1900, are now most European starling: the western bluebird, American kestrel,
visible in the rolling oak savannas and, to a lesser extent, and Lewis’ woodpecker.
adjacent chaparral of the interior lowland Umpqua and Nutria (also known as coypu),native to the Patagonian
Rogue valleys. Several different forms of the wild turkey subregion of South America, were imported to North
met with poor to variable success until Rio Grande turkeys America for a fur-farm at Elizabeth Lake (California) in
from Texas were introduced during the early 1980s (T. 1899.38 Unknown numbers of nutria were liberated from
Farrell, Oregon Department of Fish and Wildlife, pers. a fur farm in Tillamook County during a flood (S. Jewett
comm.). This form has proven extraordinarily successful, in Larrison80). From this release, a colony was established
largely because adults attempt to renest after nesting in the vicinity of Garrison Lake. Subsequently, feral nutrias
failures even very late in the season. were observed along the Nestucca River and appeared to
The oldest of the 3 game bird introductions is the be doing well (F. Wire in Larrison80). By 1946, nutrias were
ring-necked pheasant. Ring-necked pheasants were first established across the westside interior valleys, including
brought from China in 1881 by Judge Owen Denny. This the lowland Willamette.76 Nutria use of grassland habitat
introduction failed, but subsequent Denny-encouraged adjacent to wetlands is limited to movements between
introductions resulted in its establishment by 1884.81 By aquatic habitats and marginal foraging. Based on this
1891, it was well enough established in Willamette Valley pattern of habitat use, effects of nutria are likely to be
agricultural grasslands that Oregon declared the first open greater in wetland habitats than in the adjacent upland
hunting season on pheasants, during which an estimated (e.g., grasslands).
50,000 individuals were taken. Even before that, pheasants The eastern bullfrog was originally introduced to
were being live-trapped for transport to other areas, and westside habitats in the Pacific Northwest by Matt
western Washington was among the first to receive Ryckman, who brought bullfrogs to the McKenzie Fish
translocated pheasants. Soon after 1900, the ring-necked Hatchery in 1921.78 Several subsequent introductions led
pheasant was established over most of the Pacific to the bullfrog-farming craze of the 1930s, which saw a
Northwest agricultural grasslands. Increasingly large number of entrepreneurial bullfrog ventures appear
manicured agricultural practices (e.g., reduction of through the West.69 All of these ventures failed and the
hedgerows) that began during the mid-1970s have resulted purported frog farmers either left their stock in situ or
 282 Wildlife–Habitat Relationships in Oregon and Washington

actually seeded them in aquatic sites nearby. By 1950, the Protection and restoration may be approached through
bullfrog was well established in westside habitats across several modes. These include acquisition, conservation
the Pacific Northwest (R. Storm, Oregon State University, agreements, landowner outreach and incentive programs,
pers. comm.). Like nutria, bullfrogs do not use upland legislative enactments, and management agency policies.
habitats frequently; such use is restricted to some foraging Where opportunities occur, acquisition by land
adjacent to and movements between aquatic sites, management agencies or organizations may facilitate long-
typically under wet or high humidity conditions. As a term protection and management. Mechanisms to
consequence, much of the potential impact of bullfrogs accomplish this include purchase, land-swaps, and deed-
has focused on native aquatic species such as the Oregon trusts. Frequently, extensive acquisitions are economically
spotted frog and foothill yellow-legged frog,13, 59, 60, 62 and unfeasible, so alternative approaches must be considered.
the effects bullfrogs have on these species is arguable.64 Most land ownership of westside grassland and
What impact bullfrogs have, if any, on those species chaparral habitat is private, so actions to protect or
associated with grassland or chaparral habitat proper, enhance habitat for wildlife must be in partnership with
which they only occasionally encounter, is unknown. private landowners, many of whom make their living on
The common snapping turtle is a large aquatic turtle these same lands. This approach is especially important
native to most of the eastern U.S. 24 The original where protected (public) and private land interdigitate,
introduction is not known and likely represents a series and the management of private land has opportunity to
of releases by pet owners. Similar to the red-eared slider, significantly influence system dynamics on public land.
its use of upland habitat is restricted to nesting-related For these reasons, land management agencies and
activities by females and to a lesser extent, movement organizations (e.g., U.S. Fish and Wildlife Service, The
between aquatic sites and to overwintering sites. Like the Nature Conservancy), and private landowner government
bullfrog, a diverse diet and imposing feeding mode have outreach agencies (e.g., Natural Resources Conservation
been the basis of suggestions that it might negatively Service, Division of State Lands) must play a major role
influence native species.66 However, the potential negative in wildlife conservation. A significant portion of this role
impacts of this species have not been investigated, and must be cooperative development of guidelines for
the impacts it may have on species in grassland or conservation with private landowners. For grassland
chaparral adjacent to aquatic sites, if any, are unknown. wildlife in particular, managing existing and future
The date of introduction of the red-eared slider, an agricultural lands to provide suitable habitat is crucial
aquatic turtle native to and widespread in the eastern U.S,24 because of the near disappearance of native grasslands,
is not known. The species appeared in the pet trade for and the low likelihood that extensive areas adequate for
many years, and similar to the common snapping turtle, supporting viable populations can be restored to native
it is likely that scattered point introductions from pet turtle conditions.
owners are responsible for its establishment in the On public lands, efforts must be directed towards
westside Pacific Northwest. Red-eared sliders now exist including all wildlife in habitat management plans, and
at scattered points in lowland drainages in western Oregon minimizing human-wildlife conflicts when allowing
and Washington. Evidence of nesting has been found at multiple-use activities. This must include
several sites, and indirect evidence of reproduction (e.g., institutionalizing policy that directs management plans
appearance of tiny hatchlings) also exists. As with the to encompass conservation of the entire spectrum of
previous aquatic species, use of upland habitats is limited, wildlife species. Some public lands also may offer the best
being largely restricted to nesting and nest site assessment potential for large areas of quality habitat where
activities by females as well as movements to alternative population viability for area-sensitive wildlife species (e.g.,
aquatic sites or to overwintering sites. Further, any western meadowlark, several large mammals) is more
evidence of influence on native species is restricted to likely. Public lands such as State Parks and Federal Wildlife
aquatic forms,66 so if red-eared sliders affect species that Refuges should be a conservation priority, since these
use grasslands or chaparral proper, those interactions have lands potentially represent stable, long-term
not been addressed. commitments.
Land use on westside agricultural lands often changes
Management and Conservation yearly, and this instability likely adversely impacts
Conservation of wildlife in westside grassland and grassland wildlife populations. Thus, the establishment
chaparral habitats will require integration of a suite of of designated conservation areas on public and private
activities on both private and public lands. These activities, lands may be important to maintain stable populations at
which may or may not be interdependent, include: multiple locations. Examples of these efforts include the
1. inventory of habitat types and their degree of Oregon Biodiversity Project31 and a Partners in Flight
modification, particularly where data are lacking Landbird Conservation Plan for Westside Lowlands and
or limited (i.e., Rogue and Umpqua valleys); Valleys.4
2. protection of high-quality sites and restoration of Beyond protection and restoration through cooperative
degraded habitats; and efforts, site-specific management requires attention to
3. management to maintain habitat quality and system dynamics. Fire, the primary natural disturbance
wildlife populations. process that shaped westside grassland and chaparral
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 283

habitats, occurs rarely relative to historical conditions. various Carex and Juncus species; forbs include Hypericum
Thus appropriately, current restoration and management perforatum, Myosotis laxa, Veronica scutellata, Plagiobothrys
of grasslands has focused on restoring fire back into the figuratus, and Camassia quamash; shrubs include Rosa spp.
system as well as removal and control of exotic plants, and Spirea douglasii; and trees include Fraxinus latifolia and
and seeding of native species.135 Significant attempts also Crataegus douglasii.120 Vegetation sampling during trap
have been made to identify and acquire remaining parcels periods showed that the height of woody shrubs was
of prairie habitat (E. Alverson, The Nature Conservancy, substantially reduced in burned sections, but percent cover
pers. comm.). was not greatly affected. Diversity and percent cover of
The expanding human population of western Oregon grasses were greater in the burned sections than the
and Washington has resulted in continual loss and unburned. Litter cover and depth increased with time
degradation of grassland and chaparral wildlife habitat. since fire and was consistently greater in the unburned
Current conservation emphasis has focused on rare and section.
declining species on public lands. A significant need exists
Small Mammals. Trapping surveys for small mammals
for more community-based conservation and wildlife
were conducted for 5 consecutive days in late summer
partnerships with private landowners. Without these
prior to burning in 3 years (1996-98), with an additional
components, the future of wildlife in westside grassland
trap period 2 weeks after burning in 1998. Small mammals
and chaparral habitats will probably be relegated to
were live-trapped along 9 permanent transects, 3 in each
protection and management of highly disjunct
prairie section.
populations and sites.
In decreasing order of abundance, small mammal
species captured on the prairie were the deer mouse,
vagrant shrew, Townsend’s vole, gray-tailed vole, short-
Case Histories
tailed weasel, dusky-footed woodrat, and Pacific jumping
1. Small Mammals and Fire in a Wet Prairie of mouse. Numbers of individuals captured over 3 trap
periods differed between prairie sections for most species
Western Oregon
(Table 8). The deer mouse, a habitat generalist, was fairly
James Faulkner abundant in all prairie sections, but was not evenly
Periodic fire and flooding, and variations in distributed among sections (X2 = 35.3, df = 2, P <0.01),
microtopography, soil types, and hydrology most likely having greatest abundance in Section C. The vagrant
created a mosaic of vegetation composition and shrew was not evenly distributed among sections (X2 =
successional stages in the historical prairies of the 17.1, df = 2, P <0.01), having greatest abundance in Section
Willamette Valley. Current prairie restoration and B and least in Section A. The distribution of Townsend’s
management efforts use prescribed fire to set back voles did not differ among prairie sections (X2 = 4.7, df =
succession and attempt to recreate historical prairie 2, P >0.08), nor did the distribution of gray-tailed voles
conditions. This case history investigates the influence of (X2 = 0.2, df = 2, P >0.25). Dusky-footed woodrats were
prescribed fire on the small mammal community of a captured only in Section C, and no nests were observed
native wet prairie in western Oregon. outside of that section.
Small mammal species had different direct responses
Study Site. The study site was a 339 acre (137 ha) section to fire, as indicated by the pre- and post-fire trap periods
of native wet prairie in the Willamette Floodplain Research in 1998. The chance of recapturing an individual deer
Natural Area (RNA). The RNA is located at W. L. Finley mouse from the pre-fire trap period during the post-fire
National Wildlife Refuge, 10 miles (16 km) south of trap period did not differ between the combined burned
Corvallis, Benton County, Oregon. Elevation ranges from sections (A and B) and the unburned section (P = 0.49).
269-289 feet (82-88 m). Soils are silty clay loams of alluvial Likewise, the chance of capturing a vole (species
origin. The impermeable clay holds winter precipitation, combined) in a burned area before fire did not differ from
causing shallow flooding of low areas until late spring. after fire (P = 0.45). This suggests little direct fire mortality
The prairie is surrounded by fields cultivated for
commercial grass seed and the production of goose forage.
A program of prescribed burning was initiated in 1990 on Table 8. Numbers of individuals of common small
the RNA prairie to control invasive alien plant species and mammal species captured during 3 trap periods
(1996-98) within the 3 (A, B, and C) differentially
reduce shrub and tree cover.120 The small mammal study
burned sections of prairie.
site consisted of a 69-acre (28 ha) section of prairie burned
annually since 1990 (Section A), a 91-acre (37 ha) section
burned triennially since 1991 (Section B), and a 178-acre Species A B C Total
(72 ha) section unburned since 1979 (Section C). Burning
during this study took place in September 1996 and 1998 Deer mouse 73 62 136 271
in Section A, and September 1998 in Section B. Vagrant shrew 11 37 17 65
Townsend’s vole 9 16 21 46
The RNA wet prairie has a high diversity of plant
Gray-tailed vole 10 11 9 30
species. Of the most common species, graminoids include
Dusky-footed woodrat 0 0 6 6
Deschampsia cespitosa, Agrostis tenuis, Holcus lanatus, and
 284 Wildlife–Habitat Relationships in Oregon and Washington

or fire-induced emigration from burned areas for deer

mice or voles. Conversely, 7 of the 11 vagrant shrews
captured in the pre-fire trap period were in the sections to
be burned, whereas 0 of 11 were trapped in those burned
sections following fire. This suggests that fire-induced
mortality and/or emigration may be significant for
vagrant shrews.
Discussion. The wet prairies of the Willamette Valley
historically were shaped by fire and water. Grassland is
not the climax state in this system, as fire suppression
results in shrubland and ultimately ash woodland in some
areas. The combination of prairie sections with different
burn frequencies on the RNA prairie creates different
stages of succession, which may allow for a greater
diversity of small mammal species. In this study, each
species had different direct responses to fire, as well as
different associations with the prairie sections (i.e., stages
of succession). Despite this variability in species
associations, none of the species were ever most abundant
in the annually burned section. A period of 2-3 years may
be needed to overcome the effects of fire. Therefore, a
prairie management scheme that takes the diversity and
population characteristics of small mammal species into
account may benefit by providing a variety of successional
stages, while allowing adequate time between fire events
for populations of fire-sensitive species to recover.

2. Managing Agricultural Grasslands for

Landbirds: The Role of Scale and Context
Bob Altman & Rebecca Goggans
In 1996, the Oregon Department of Fish and Wildlife
initiated a 2-year project to develop management and
conservation strategies for several state sensitive
grassland-associated bird species in the Willamette Valley,
particularly the western meadowlark, Oregon vesper
sparrow, and streaked horned lark. All 3 species were
formerly common breeding species in native grasslands,
but have suffered extensive population declines.
Methods. Data collection integrated several methods at
extensive and intensive scales. Broad-scale (extensive)
censusing was conducted at 544 roadside point-count Figure 1. Occurrence of three grassland-associated bird
stations throughout the Willamette Valley in 1996 to species based on point-count censusing in the Willamette
characterize the grassland breeding bird community, and Valley, May - June 1996.
quantify species abundance and habitat associations. MG – mowed grass
Territory-mapping (intensive scale) was conducted CG – cultivated grass seed
primarily in 1997 at selected sites to quantify species area- HRC – herbaceous row crop
requirements and habitat relationships. CTF – Christmas tree farm
Habitat characterization was conducted within a 328 BG – bare ground
feet (100 m) radius of each point-count station. Among PF – plowed field
habitat variables described were estimates of percent cover PS – pasture
of habitat type and percent cover of vegetation height WP – wet prairie
classes in 6-inch (15.2 cm) increments (e.g., 0-6 in [0-15.2 FF – fallow field
cm], 6-12 in [15.2-30.5 cm], etc.). Habitat data were MF – maintained field
collected within each territory along three 164-foot (50 m) RD – gravel/dirt road
transects randomly located from the center of the territory. RRW – roadside right-of-way
Habitat variables described included estimates of percent
cover by growth form, and percent cover of vegetation
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 285

height classes as described above. Territories were

delineated using the “repeat-flush” technique.131 Territory
size was generated from boundary point data collected
with a Global Positioning System (GPS) and downloaded
into Arc View software for calculations.
Results: Censusing. Patterns of species occurrence by
habitat type indicated a relatively high degree of
specialization for vesper sparrows and western
meadowlarks (each predominantly in 2 habitat types), and
a relatively wide range of habitat types used by horned
larks (Figure 1). Pasture was a predominant habitat for
the western meadowlark and vesper sparrow, accounting
for nearly 50% of the detections of both species, despite
<10% of the sampling effort in pasture. Christmas tree
farms were the other high-use habitat for vesper sparrow Figure 2. Horned lark and western meadowlark occurrence in
(approximately 45% of the detections). Grass-seed fields, annual and perennial grass fields based on point-count
which comprised 53% of the sampling effort, were not censusing in Willamette Valley, May - June 1996.
used by vesper sparrows, but were the second primary
habitat for western meadowlarks (approximately 35% of
the detections). Percent detections (10.6) of western
meadowlarks in fallow fields were also greater than the
percent of this habitat type sampled (6.3). The 4 principal
habitats used by horned larks were gravel and dirt roads,
roadside rights-of-way, plowed fields, and grass-seed
fields. Horned lark detections in bare ground, Christmas
tree farms, and herbaceous row crops were also greater
than the percent sampling effort in those habitat types.
Western meadowlark and horned lark occurrence in
grass-seed fields was further defined by the type of field
(i.e., annual or perennial). Western meadowlarks (n = 80)
were detected nearly equally in annual and perennial grass
fields, but horned larks (n = 29) were substantially more
abundant in annual grass fields (Figure 2). This difference
is likely because of the presence of more bare ground in
annual grass fields in the spring due to planting, whereas Figure 3. Vesper sparrow and western meadlowlark
perennial grass fields are vegetated to a greater extent occurrence in pasture types based on point-count censusing
during this time period. in Willamette Valley, May - June 1996.
Western meadowlark and vesper sparrow occurrence
in pastures was further defined by the amount of vertical
seven percent (n = 14) of the territories were 10-19 acres
structure provided by shrubs and trees. Both species
(4-7.7 ha). Only 19% (n = 4) were <10 acres (4.7 ha), and
occurred primarily in entirely herbaceous pastures or
14% (n = 3) >20 acres (8.1 ha). Only 24% (n = 5) of western
pastures with <10% shrub-tree cover, although western
meadowlark territories included grass-seed fields, and
meadowlarks were most often associated with
none were >50% grass-seed fields, despite efforts to locate
predominantly herbaceous pastures, and vesper sparrows
territories in areas that were exclusively or near-
with a higher percentage of shrub/tree cover (Figure 3).
exclusively grass-seed fields. Territory sizes for vesper
Pastures with >10% shrub-tree cover were moderately
sparrows and horned larks were considerably smaller than
used by vesper sparrows, but rarely used by western
those of western meadowlarks. Mean territory size for
meadowlarks. Shrubs and trees are important to both
vesper sparrow (n = 38) was 3.1 acres (1.3 ha). However,
species for singing perches.132
all but 1 of the territories were >6 acres (2.4 ha). Mean
Patterns of species occurrence relative to grass height
territory size for the horned lark was 1.9 acres (0.8 ha),
were different for all 3 species. Most horned lark detections
although the sample size was only 3 territories.
were in short grass (0-6 inches [0-15.2 cm]), vesper
Mean percent occurrence of each grass height class in
sparrows occurred in relatively short grass (1-12 inches
western meadowlark territories was relatively similar.
[2.5-30 cm]), and western meadowlarks were most tolerant
However, individual territories were highly variable, such
of different grass heights (up to 36 inches [91.4 cm])
that each height class was dominant or codominant in at
depending on the type of field.
least 1 territory. This suggests habitat selection by western
Results: Territory Mapping. Mean territory size for meadowlarks may not have been dependent on grass
western meadowlark (n = 21) was 14.3 acres (5.8 ha). Sixty- height. Vesper sparrow territories were distinctive for
 286 Wildlife–Habitat Relationships in Oregon and Washington

particular grass height classes, which suggests some

3. Squaw Flat:
degree of habitat selection by grass height. Over 80% of
mean percent occurrence of each height class was in the 3 A Fire Link between Grasslands and Reptiles
classes <18 inches (45.7 cm). Vesper sparrow territories Marc Hayes
were dominated (46%) by grass heights of 6-12 inches In southwestern Oregon, grasslands intermix with a
(15.2-30.5 cm). complex of conifer and deciduous tree-dominated
Conservation and Management. Habitat issues of greatest assemblages. At low elevations, relatively continuous
importance to the western meadowlark were size and grasslands dominate in an areal sense. Forested patches
context. Western meadowlark was relatively tolerant of cover progressively larger areas as elevation increases. At
various local habitat conditions (e.g., grass height) and mid-elevations, grasslands of a lowland character are
field types (e.g., annual or perennial grass fields, pasture, reduced to scattered patches. At higher elevations yet, this
fallow fields), but more area-sensitive than the other type of grassland vanishes from the forest matrix. Regions
species, and more sensitive to the context and juxtaposition of grassland-forest contact are not static. These areas
of habitat types. Thus, the management focus needs to be change in a manner that can markedly influence the fauna
on coarse-level conditions at landscape scales. With a associated with their adjoining vegetation assemblages.
relatively large territory size for a passerine, western The example provided here uses the reptile fauna to
meadowlarks require relatively large areas of suitable illustrate the grassland maintenance element of such a
contiguous habitat to maintain populations. However, dynamic for a mid-elevation site in the South Umpqua
since cultivated grass fields apparently do not meet their Basin.
habitat requirements exclusively, large patches of habitat Study site. The study site is the proposed Squaw Flat
must include at least half their preferred habitat types Research Natural Area (hereafter simply Squaw Flat).
(pasture and fallow fields). Squaw Flat comprises 558 acres (226 ha) of mixed conifer-
Habitat issues of greatest importance to vesper deciduous forest at the intersection of Jackson and Squaw
sparrows were type and condition. They regularly creeks in the South Umpqua Basin 40 miles (64 km)
occurred in small patches (e.g., <10 acres [4 ha]) of southeast of Roseburg, Oregon. The site ranges from 1,740
grassland habitat, but only within 1 grassland type feet (533 m) to 2,450 feet (747 m) in elevation, slopes mostly
(pastures) and with relatively specific local conditions for west and northwest, and has a complex volcanic substrate
grass height and shrub-tree cover. Thus, management does altered through substantial landflows. Except for 2 tiny
not need to be conducted at landscape scales, but must be ponds (<1.7 acre [0.7 ha]), scattered seeps, a few ephemeral
within the context of a specific agricultural grassland type, drainages, and the riparian margins of Jackson and Squaw
and be designed to create and maintain relatively specific creeks, Squaw Flat is uplands. Uplands can be
structural conditions within that habitat type. fundamentally divided into forest and prairies. Forest, the
Horned larks used a wide variety of habitat types, but dominant upland category, has a continuous but highly
were relatively specific at the microhabitat scale for grass variable conifer-dominated matrix that covers about 95%
height and the presence of bare ground. Thus, habitat of the upland area. The forest ranges from mesic (Douglas-
management can be accomplished at small scales by fir and western hemlock as dominants) to xeric (ponderosa
creating or maintaining bare ground or sparsely vegetated pine dominated) in a diffuse west-to-east gradient. The
areas within or adjacent to a wide variety of suitable other 5% (ca. 20 acres [8 ha]) of upland is 7 small prairies
habitat types. It also seems that suitable habitat can (0.6-10.3 acres [0.2-4.2 ha]) imbedded in the forest matrix
support relatively dense populations because of their small (Figure 4; A-G). Native grasses, graminoids, and forbs are
territory size and their propensity to forage communally dominant on these prairies. Prairies are fundamentally
during the breeding season at good foraging sites. open with low-stature vegetation, but Oregon white oaks
The near extirpation of native grasslands in the ring most of their peripheries.
Willamette Valley leaves few opportunities to maintain Squaw Flat is in as close to an undisturbed condition
populations of grassland bird species on these lands. Thus, as a site in this elevation range can be expected to be.
conservation of declining and state sensitive grassland Timber harvest, which has occurred at points along its
birds must occur primarily within the context of these periphery, has not occurred on the site, recently or
species’ use of non-native agricultural grasslands. Habitat historically (Figure 4). As a consequence, much of the forest
requirements of the western meadowlark, Oregon vesper is old growth that exceeds a few hundred years in age.
sparrow, and streaked horned lark share some similarities, The only significant disturbance to Squaw Flat over the
but also have unique differences that can be managed for last 30 years has been the irregular trespass of livestock
in agricultural grasslands. However, conservation of these from adjacent Umpqua National Forest land; appropriate
species in Willamette Valley agricultural grasslands will exclusion barriers are lacking.
require integrated management actions at several different
scales. Decision making will be dependent on Reptile Fauna. Squaw Flat has a rich reptile fauna with
opportunities available, and the context of type of land 14 species recorded (northern alligator lizard, southern
use occurring. alligator lizard, western skink, western fence lizard, rubber
boa, racer, western rattlesnake, ringneck snake, sharptail
snake, Pacific coast aquatic garter snake, western terrestrial
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 287

Figure 4. Squaw Flat aerial

photograph. Based on a 19 July 1992
overflight. Letters indicate prairies;
ponds are P1 and P2 (see text). True
north is 12 degrees left of the vertical
axis of the page.

Figure 5. Number of reptiles encountered at Squaw Flats Figure 6. Number of reptiles encountered at Squaw Flats
during surveys with effort-adjusted for the area of major during surveys with equal effort in forest and prairie habitats.
habitat categories.
asymmetry was even more skewed (Chi-square test: X2 =
32.3, P <0.0001; Figure 6). Egg-laying and live-bearing
garter snake, northwestern garter snake, common garter
reptiles both contributed to the pattern, but the pattern
snake, western pond turtle) but as many as 3 other species
was more pronounced among egg layers. Twenty-two
potentially present. Preliminary surveys of this fauna were
reptile nests were found in prairie or prairie-edge
completed during 1 May-15 June 1996. Surveys were
locations; 5 different species were represented (western
habitat proportional; effort was adjusted to the area of each
pond turtle, n = 8; western fence lizard, n = 2; western
habitat type. Despite the enormous asymmetry in effort
skink, n = 10; gopher snake, n = 1; racer, n = 1). A female
that favored forested habitat because of its much greater
western pond turtle was observed in the process of nesting
area, reptiles were found on prairies in much greater
on the northernmost prairie (A in Fig. 4). No nests of egg-
numbers than those observed in forested habitats (Chi-
laying reptiles were found in the forested habitats despite
square test: X2 = 16.7, P <0.005; Figure 5).
over 4 times the search effort allocated to these areas.
Subsequent equal effort surveys between forest and
prairie habitats completed during 15 April—15 September Vegetation Changes. Aspects of Squaw Flat vegetation
in 1997 and 1998 showed that this distributional reveal that changes have occurred during the last 60 years.
 288 Wildlife–Habitat Relationships in Oregon and Washington

First, young true firs, primarily of the grand/white fir Literature Cited
complex (Abies grandis/A. concolor), and young western 1. Agee, J. K. 1993. Fire ecology of Pacific Northwest forests. Island Press,
hemlocks dominate the forest understory. Over most of Washington, D.C.
the area, saplings and seedlings of true firs and western 2. Alcorn, G. D. 1935. The western yellow-bellied racer in western
hemlock outnumber the younger trees of other canopy Washington. Copeia 1935(2):135.
dominants by 10 to 1. This contrasts sharply with an 3. Altman, B. 1999a. Status and conservation of grassland birds in the
Willamette Valley. Final report submitted to Oregon Dept. of Fish and
overstory dominated by Douglas-fir, ponderosa pine, and
Wildlife, Corvallis, OR.
in a few areas western hemlock. Second, the forested side 4. ———. 1999b. Conservation strategy for landbirds in the lowlands
of the white oak fringe of most prairies consisted of a and valleys of western Oregon and Washington. Version 1.0. American
melange of healthy conifers and deciduous trees, including Bird Conservancy and Oregon-Washington Partners in Flight.
frequently senescent white oaks. The latter have partially 5. Anthony, A.W. 1886. Field notes on the birds of Washington County,
or completely dead crowns. This pattern is consistent with Oregon. Auk 3:161-172.
6. Avery, M. L., and C. van Riper. 1990. Evaluation of wildlife-habitat
change inferable from historical photographs. Sixty-year-
relationships data base for predicting bird community composition in
old aerial photographs indicate that, except for 1 site, the central California chaparral and blue oak woodlands. California Fish
area of prairies has been reduced 10-20%. The sole and Game 76(2):103-117.
exception was prairie F (Figure 4), which was landflow- 7. Bailey,V. 1936. The mammals and life zones of Oregon. U.S. Department
created since 1960. Observed vegetation changes are of Agriculture, Bureau of Biological Survey, North American Fauna
consistent with the fire suppression management policy (55):1-416.
that the Umpqua National Forest has engaged in during 8. Baird, S. F. 1857[1858]. Mammals. Pages 1-757 in Reports of
explorations and surveys, to ascertain the most practicable and
this period. Fire suppression across this elevation range
economical route for a railroad from the Mississippi River to the
in this region of the Pacific Northwest results in the Pacific Ocean. Beverly Tucker, Printer, Washington, D.C., 8 (Part 1).
characteristically successful recruitment of fire-intolerant, 9. Balda, R. P. 1975. Vegetation structure and breeding bird density. Pages
shade-tolerant conifers that progressively encroach on 59-80 in Proceedings of the symposium on management of forest and
other vegetation. range habitats for nongame birds. U.S. Forest Service General Technical
Report WO-1.
Future Outlook. Egg-laying reptiles depend on prairies 10. Barbour, M. G., and J. Major. 1977. Terrestrial vegetation of California.
for nesting sites and development. This is probably John Wiley and Sons. New York, NY.
because prairies may be the only significant areas available 11. Biswell, H. H. 1974. The effects of fire on chaparral. Pages 321-364 in T.
on Squaw Flat with the appropriate temperature T. Kozlowski and C. E. Ahlgren, editors. Fire and ecosystems. Academic
characteristics for incubating eggs, basking, or warming Press, New York, NY.
12. Blaustein, A. R. 1980. Behavioral aspects of competition in a
surface objects needed to gain heat by convection. three-species rodent guild of coastal southern California. Behavioral
Estimation of the rate of encroachment on the Squaw Flat Ecology and Sociobiology 6:247-255.
prairies indicates that >50% of prairie area will be lost over 13. Borisenko, A. N., and M. P. Hayes. 1999. Status of the foothill
the next 120 years. The level of closure that may alter yellow-legged frog (Rana boylii) in Oregon. Final report prepared for
relative abundance or species composition of the egg- The Nature Conservancy under contract with the U.S. Fish and
laying reptiles present is not known. However, if current Wildlife Service with assistance from the Oregon Department of Fish
and Wildlife, the U.S. Army Corps of Engineers, and the U.S. Geological
trends continue, the abundance and species richness of
Survey.
egg-laying reptiles is likely to be reduced. Maintaining 14. Boyd, R. 1986. Strategies of Indian burning in the Willamette Valley.
the high diversity of amphibians and reptiles at Squaw Canadian Journal of Anthropology 5:65-86.
Flat is a basic RNA objective. The meeting of that objective 15. Brodie, E. D., Jr., R. A. Nussbaum, and R. M. Storm. 1969. An egg-laying
cannot be accomplished with passive management, it will aggregation of five species of Oregon reptiles. Herpetologica
require adopting a management policy that can reverse 25(3):223-227.
the existing prairie-reduction trend. The most direct way 16. Browning, M. R. 1975. The distribution and occurrence of the birds of
Jackson County, Oregon, and surrounding areas. North American Fauna
to meet that objective is to engage in a trend-reversing
70.
fire management policy: a prescribed burn schedule that 17. Burroughs, R. D., editor. 1995. The natural history of the Lewis and
will limit encroachment of fire-intolerant conifers. Clark Expedition. Michigan State University Press, East Lansing, MI.
Grassland encroachment at forest interfaces is not 18. Bury, R. B. 1992. Fort Lewis biodiversity study: western pond turtle
unique to Squaw Flat. Mid-elevations in southwestern and herpetofauna study. National Ecology Resource, U.S. Fish and
Oregon and prominent latitudinal vegetation changes in Wildlife Service, U.S. Forest Service, PNW Research Laboratory,
Olympia, WA.
central western Oregon abound with grassland-forest
19. Carraway, L. R., and B. J.Verts. 1991. Neotoma fuscipes. Mammalian
transitions. Soil and hydrology can be responsible for Species (386):1-10.
grassland occurrence in such interface areas, but Squaw 20. Chapman, J. A., and D. E. C. Trethewey. 1972. Movements within a
Flat illustrates that fire may be fundamental in grassland population of introduced eastern cottontail rabbits. Journal of Wildlife
patch maintenance. That a suite of reptiles, including most Management 36:1221-1226.
egg-layers have their northern and altitudinal limits of 21. ———, and B. J.Verts. 1969. Interspecific aggressive behavior in
distribution in this grassland-forest interface-rich region rabbits. The Murrelet 50:17-18.
22. Chappell, C. B., and R. C. Crawford. 1997. Native vegetation of the
is no coincidence. As an important factor in the
south Puget Sound prairie landscape. Pages 107-122 in P. Dunn and K.
maintenance of grassland patches, fire is probably equally Ewing, editors. Ecology and conservation of the South Puget Sound
fundamental to the maintenance of reptile diversity where prairie landscape. The Nature Conservancy, Seattle, WA.
these patches exist.
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 289

23. Christy, J., E. Alverson, M. Dougherty, S. Kolar, L. Ashkenas, and P. 48. Gilligan, J., M. Smith, D. Rogers, and A. Contreras. 1994. Birds of
Minnear. 1999. Pre-settlement vegetation map for the Willamette Valley, Oregon: status and distribution. Cinclus Publishing, McMinnville, OR.
Oregon, compiled from records of the General Land Office Surveyors 49. Gordon, K. L. 1939. The Amphibia and Reptilia of Oregon. Oregon
(c. 1850). Oregon Natural Heritage Program, Portland, OR. . State College Monographs 1:1-81.
24. Conant, R., and J. T. Collins. 1991. A field guide to reptiles and 50. Graf, W. 1939. The distribution and habitats of amphibians and reptiles
amphibians: Eastern and Central North America. Houghton Mifflin, in Lincoln, Benton, and Linn counties. Thesis, Oregon State College,
Boston. Corvallis, OR.
25. Cook, S. F., Jr. 1960. On the occurrence and life history of Contia 51. Graf, W. 1955. Cottontail rabbit introductions and distribution in
tenuis. Herpetologica 16(2):163-173. western Oregon. Journal of Wildlife Management 19:184-188.
26. Corkran, C. C. and C. Thoms. 1996. Amphibians of Oregon, 52. Graf, W., S. G. Jewett, Jr., and K. L. Gordon. 1939. Records of
Washington, and British Columbia: a field identification guide. Lone Pine amphibians and reptiles from Oregon. Copeia 1939(2):101-102.
Press, Edmonton, Alberta, Canada. 53. Grinnell, J. 1933. Review of the recent mammal fauna of California.
27. Crawford, R. C., and H. Hall. 1997. Changes in the South Puget prairie University of California Publications in Zoology 40:71-234.
landscape. Pages 11-16 in P.V. Dunn and K. Ewing, editors, Ecology and 54. Gullion, G. W. 1951. Birds of the southern Willamette Valley, Oregon.
conservation of the South Puget Sound prairie landscape. The Nature The Condor 53:129-149.
Conservancy. Seattle, WA. 55. Hall, H. L., R. C. Crawford, and B. Stephens. 1995. Regional inventory
28. Cross, S. P., and J. K. Simmons. 1983. Bird populations of the mixed- of prairies in the southern Puget Trough: Phase I. File report.
hardwood forests near Roseburg, Oregon. Oregon Department of Fish Washington Department Natural Resources, Natural Heritage
and Wildlife Technical Report 82-2-05. Program, Olympia,WA.
29. Dalquest, W. W. 1941. Distribution of cottontail rabbits in Washington 56. Hallock, L. A., and W. P. Leonard. 1997. Herpetofauna of the Ft. Lewis
state. Journal of Wildlife Management 5:408-411. Military Reservation, Pierce and Thurston Counties, Washington.
30. Dawson, L. W., and J. H. Bowles. 1909. The birds of Washington. Washington Natural Heritage Program.
Occidental Press, Seattle, WA. 57. Hanes, T. L. 1971. Succession after fire in the chaparral of southern
31. Defenders of Wildlife. 1998. Oregon’s living landscape: strategies and California. Ecological Monographs 41:27-52.
opportunities to conserve biodiversity. Defenders of Wildlife, Lake 58. Harper, J. A. and numerous authors. 1987. Ecology and management of
Oswego, OR and The Nature Conservancy, Portland, OR. Roosevelt elk in Oregon. Revised edition. Oregon Department of Fish
32. DeLong, K. T. 1966. Population ecology of feral house mice: and Wildlife, Portland, OR.
interference by Microtus. Ecology 47:481-487. 59. Hayes, M. P. 1994a.The spotted frog (Rana pretiosa) in western
33. Detling, L. E. 1961. The chaparral formation of southwestern Oregon, Oregon. Oregon Department of Fish and Wildlife, Wildlife Diversity
with considerations of its postglacial history. Ecology 42:348-357. Program, Technical Report 94-0-01:1-19 + appendices.
34. Edge, W. D., J. O. Wolff, and R. L. Carey. 1995. Density-dependent 60. ———. 1994b. Current status of the spotted frog in western Oregon.
responses of gray-tailed voles to mowing. Journal Wildlife Management Oregon Department of Fish and Wildlife, Wildlife Diversity Program,
59:245-251. Technical Report 94-0-01:1-11.
35. ———, R. L. Carey, J. O. Wolff, L. M. Ganio, and T,. Manning. 1996. 61. ———. 1996. Assessment of the sensitive amphibian and reptile fauna
Effects of Guthion 2S on Microtus canicaudus: a risk assessment on selected culvert, pond, road, slide, and stream sites within the Tiller
validation. Journal Applied Ecology 32:269-278. Ranger District (Umpqua National Forest). Final report prepared for
36. Elzroth, E. K. 1987. Population study and breeding biology of the the Umpqua National Forest under subcontract to Resources
western bluebird (Sialia mexicana). Oregon Department of Fish and Northwest, Inc.
Wildlife, Nongame Wildlife Program, Technical Report 86-1-01. 62. ———. 1997. Status of the Oregon spotted frog (Rana pretiosa) in
37. Emlen, J. T. 1972. Size and structure of a wintering avian community in the Deschutes Basin and selected other systems in Oregon and
southern Texas. Ecology 53:317-329. northeastern California with a rangewide synopsis of the species
38. Evans, J. 1970. About nutria and their control. U.S. Department of the status. Final report prepared for the U.S. Fish and Wildlife Service with
Interior, Bureau of Sport Fisheries and Wildlife, Resource Publication assistance from the Oregon Department of Fish and Wildlife.
86:1-65. 63. ———. 1998. Assessment of the sensitive amphibian and reptile fauna
39. Evenden, F. G., Jr. 1949. Habitat relations of typical austral and boreal on Apple Jack and Johnnie Springs salvage units, Wildcat Timber Sale
avifauna in the Willamette Valley, Oregon. Dissertation, Oregon State Units, and ERFO sites within the Tiller Ranger District (Umpqua
College, Corvallis, OR. National Forest). Final report prepared for the Umpqua National
40. Fitch, H. S. 1935. Natural history of the alligator lizards. Transactions Forest under subcontract to EA Engineering, Science, and Technology,
of the Academy of Science of Saint Louis 39(1):3-38. Inc.
41. Fitch, H. S. 1936. Amphibians and reptiles of the Rogue River Basin, 64. ——— and M. R. Jennings. 1986. Decline of ranid frog species in
Oregon. American Midland Naturalist 17:634-652. western North America: are bullfrogs (Rana catesbeiana) responsible?
42. Forrester, B. and R. Storre. 1992. Western pond turtle survey: Ft. Journal of Herpetology 20(4):490-509.
Lewis, Washington. Unpublished report, Department of Army. David 65. Hoffman, R. 1927. Birds of the Pacific States: containing brief
Evans Associates., Bellevue, WA. biographies and descriptions of about four hundred species with
43. Franklin, J. F., and C. T. Dyrness. 1973. Natural vegetation of Oregon special reference to their appearance in the field. Boston, MA.
and Washington. U.S. Forest Service General Technical Report PNW-8. 66. Holland, D. C. 1994. The western pond turtle: habitat and history. Final
44. Gabrielson, I. N., and S. G. Jewett. 1940. Birds of Oregon. Oregon report to the U.S. Department of Energy, Bonneville Power
State Monograph No. 2. Oregon State College, Corvallis, OR. Administration, Division of Fish and Wildlife, and Wildlife Diversity
45. Galen, C. 1989. A preliminary assessment of the status of the Lewis’ Program, Oregon Department of Fish and Wildlife.
woodpecker in Wasco County, Oregon. Oregon Department Fish and 67. Hopkins, D. D., and R. B. Forbes. 1980. Size and reproductive patterns
Wildlife, Nongame Technical Report 88-3-01. of the Virginia opossum in northwestern Oregon. The Murrelet
46. Gavin, T. A. 1984. Whitetail populations and habitats: Pacific 60:95-98.
Northwest. Pages 487-496 in L. K. Halls, editor. White-tailed deer: 68. Ingles, L. G. 1965. Mammals of the Pacific States: California, Oregon,
ecology and management. Stackpole Books, Harrisburg, Pennsylvania. and Washington. Stanford University Press, Stanford, CA.
47. Gilbert, B., T. Williams, and J. Bottorff. 1991. 1991 spotted frog survey 69. Jennings, M. R., and M. P. Hayes. 1985. Pre-1900 overharvest of
Ft. Lewis Military Reservation, Pierce County, Washington. U.S. Fish and California red-legged frogs (Rana aurora draytonii): the inducement for
Wildlife Service, Olympia, WA. bullfrog (Rana catesbeiana) introduction. Herpetologica 41(1):94-103.
 290 Wildlife–Habitat Relationships in Oregon and Washington

70. Jewett, S. G., Jr. 1936. Notes on the amphibians of the Portland, 96. Northwest Habitat Institute. 1999. Oregon: Landscape-scale
Oregon, area. Copeia 1936(1):71-72. vegetation map. Northwest Habitat Institute, Corvallis, OR.
71. ———, and H.W. Dobyns. 1929. The Virginia opossum in Oregon. 97. Nussbaum, R. A., E. D. Brodie, Jr., and R. M. Storm. 1983. Amphibians
Journal of Mammal. 10:351. and reptiles of the Pacific Northwest. University of Idaho Press,
72. ———, W. P. Taylor, W. T. Shaw, and J. W. Aldrich. 1953. Birds of Moscow, ID.
Washington State. University of Washington Press, Seattle, WA. 98. Parker, W. S., and W. S. Brown. 1980. Comparative ecology of two
73. Johannessen, C. L., W. A. Davenport, A. Millet, and S. McWillaims. 1971. colubrid snakes in northern Utah. Milwaukee Public Museum,
The vegetation of the Willamette Valley. Annals of the Association of Publications in Biology and Geology (7):1-104.
American Geography 61:286-302. 99. Puchy, C. A., and D. B. Marshall. 1993. Oregon Wildlife Diversity Plan.
74. Johnson, O.B. 1880. List of the birds of the Willamette Valley, Oregon. Oregon Department of Fish and Wildlife, Portland, OR.
American Naturalist 14:485-491, 635-641. 100. Rathburn, S. F. 1902. A list of the land birds of Seattle, Washington,
75. Knopf, F. L. 1994. Avian assemblages on altered grasslands. Studies in and vicinity. Auk 19:131-141.
Avian Biology 15:247-257. 101. Resources Northwest, Inc. and Pentec Environmental Inc. 1995.
76. Kuhn, L. W., and E. P. Peloquin. 1974. Oregon’s nutria problem. Neotropical migratory bird survey, Fort Lewis Military Reservation.
Proceedings Vertebrate Pest Conference 6:101-105. Unpublished report submitted to U.S. Army Corps of Engineers and
77. Lais, M.C. 1969. Distribution of amphibians and reptiles in Jackson HQ, I Corps and Fort Lewis Directorate of Engineering and Housing.
County, Oregon. Thesis, Southern Oregon State College, Ashland, OR. 102. Richardson, C., and F. W. Sturges. 1964. Bird records from southern
78. Lampman, B. H. 1946. The coming of pond fishes. Binfords and Mort, Oregon. Condor 66:514-515.
Portland, OR. 103. Riegel, G. M., B. G. Smith, and J. F. Franklin. 1992. Foothill oak
79. Lardie, R. L. 1963. Checklist of the amphibians and reptiles of Pierce woodlands of the interior valleys of southwestern Oregon. Northwest
County, Washington, with special reference to those found on Science 66:66-76.
McChord Air Force Base. Tricor 3(1):10-17. 104. Rogers, R. 1999. Microhabitat selection and status of streaked
80. Larrison, E. J. 1943. Feral coypus in the Pacific Northwest. The horned lark, western bluebird, Oregon vesper sparrow, and western
Murrelet 24:3-9. meadowlark in western Washington. Thesis, Evergreen State College,
81. Laycock, G. 1966. The alien animals: the story of imported wildlife. The Olympia,WA.
Natural History Press, Garden City, New York, NY. 105. Rolph, D.N. 1998. Assessment of Neotropical migrant landbirds on
82. Leonard, W. P. 1995. Survey of amphibians and reptiles at McChord Air McChord Air Force Base, Washington. Unpublished report, The Nature
Force Base, Pierce County, Washington. Washington Natural Heritage Conservancy of Washington, Seattle,WA.
Program. 106. Sauer, J. R., J. E. Hines, I. Thomas, J. Fallow, and G. Gough,. 1999. The
83. Leonard, W. P., and L. A. Hallock. 1997. Herpetofauna of the South North American Breeding Bird Survey: results and analysis 1966-1998.
Puget Sound prairie landscape. Pages 65-74 in P.V. Dunn and K. Ewing, Version 98.1. U.S.G.S. Patuxent Wildlife Research Center, Laurel, MD.
editors, Ecology and conservation of the South Puget Sound prairie 107. Slater, J. R. 1955. Distribution of Washington amphibians. Occasional
landscape. The Nature Conservancy, Seattle, WA. Papers of the College of Puget Sound 24:212-242.
84. Lev, E., J. Fugate, M. P. Hayes, D. Smith, L. Wilson, and R. Wisseman. 108. Slevin, J. R. 1928. The amphibians of western North America: An
1994. The biota of Smith and Bybee Lakes Management Area. Final account of the species known to inhabit California, Alaska, British
report prepared for Metro Regional Parks and Greenspaces, Portland, Columbia, Washington, Oregon, Idaho, Utah, Nevada, Arizona, Sonora,
OR. and Lower California. Occasional Papers of the California Academy of
85. Lewis, M. G., and F. A. Sharpe. 1987. Birding in the San Juan Islands. The Sciences 16:1-151.
Mountaineers, Seattle, WA. 109. Slipp, J. W. 1940. The mammals, reptiles, and freshwater fishes of the
86. Lidicker, W. Z., Jr. 1966. Ecological observations on a feral house Tacoma area. Thesis, College of Puget Sound, Tacoma, WA.
mouse population declining to extinction. Ecological Monographs 110. Smith, M. R., P. W. Mattocks, Jr., and K. M. Cassidy. 1997. Breeding
36:27-50. birds of Washington State.Volume 4 in Washington State Gap Analysis
87. Lillywhite, H. B., G. Friedman, and N. Ford. 1977. Color matching and —Final Report in K. M. Cassidy, C. E. Grue, M. R. Smith, and K. M.
perch selection by lizards in recently burned chaparral. Copeia Dvornich, editors. Seattle, Audubon Society Publication In Zoology No
1977(1):115-121. 1, Seattle, WA.
88. MacArthur, R. H., and J. W. MacArthur. 1961. On bird species diversity. 111. Smith, W. P. 1985. Plant associations within the interior valleys of the
Ecology 42:594-598. Umpqua River basin, Oregon. Journal of Range Management
89. Marshall, D. B., M. Chilcote, and H. Weeks. 1996. Species at risk: 38:526-530.
sensitive, threatened, and endangered vertebrates of Oregon. Second 112. Speer, F. and J. Felker. 1991. Blue-gray gnatcatcher (Polioptila caerulea)
edition. Oregon Department of Fish and Wildlife, Portland, OR. study: Lower Table Rock Preserve. Unpublished report submitted to
90. Maser, C, B. R. Mate, J. F. Franklin, and C. T. Dyrness. 1981. Natural The Nature Conservancy, Medford, OR.
history of Oregon Coast mammals. U.S. Forest Service; and U.S. 113. Stebbins, R. C. 1954. Amphibians and reptiles of western North
Department of the Interior, Bureau of Land Management, General America. University of California Press, Berkeley, CA.
Technical Report PNW-133:1-496. 114. Stebbins, R. C. 1985. A field guide to the amphibians and reptiles of
91. Mason, R. T., D. T. Lerner, and I. T. Moore. 1997. Snake communities in western North America. Houghton Mifflin, Boston, MA.
the central Willamette Valley: effects of changing habitat. Presented at 115. Steinberg, E. and D. Heller. 1997. Using DNA and rocks to interpret
the Annual Meeting of the Society of Northwestern Vertebrate the taxonomy and patchy distribution of pocket gophers in western
Zoologist, Olympia,WA. Washington prairies. Pages 43-51 in P.V. Dunn and K. Ewing, editors,
92. McAllister, K. M., W. P. Leonard, and R. M. Storm. 1993. Spotted frog Ecology and conservation of the South Puget Sound prairie landscape.
surveys in the Puget Trough of Washington, 1989-1991. Northwestern The Nature Conservancy, Seattle, WA.
Naturalist 74:10-15. 116. St. John, A. D. 1984. The herpetology of Jackson and Josephine
93. McQueen, L. 1979. The grasshopper sparrows of the Willamette Valley. Counties, Oregon. Oregon Department of Fish and Wildlife, Nongame
Oregon Birds 5:32-35. Wildlife Program, Technical Report 84-2-05:1-78.
94. Meyers, S. M., and J. O. Wolff. 1994. Comparative toxicity of azinphos- 117. St. John, A. D. 1985. The herpetology of the Interior Umpqua River
methyl to house mice, laboratory mice, deer mice, and gray-tailed voles. Drainage, Douglas County, Oregon. Oregon Department of Fish and
Archives of Contamination and Toxicology 26:478-482. Wildlife, Nongame Wildlife Program, Technical Report 85-2-02:1-69.
95. Murray, K. 1968. The pig war. Washington Historical Society, Tacoma, 118. St. John, A. D. 1987. The herpetology of the Willamette Valley,
WA.
 CHAPTER 10: WESTSIDE GRASSLANDS/CHAPARRAL 291

Oregon. Oregon Department of Fish and Wildlife, Nongame Wildlife

Program, Technical Report 86-1-02:1-79.
119. Storm, R. M. 1941. Effect of white man’s settlement on wild animals
in the Mary’s River valley. Thesis. Oregon State University, Corvallis,
OR.
120. Streatfeild, R., and R. E. Frenkel. 1997. Ecological survey and
interpretation of the Willamette Floodplain Research Natural Area, W.
L. Finley National Wildlife Refuge, Oregon, U.S.A. Natural Areas Journal
17:346-354.
121. Stroud, D. C. 1982. Population dynamics of Rattus rattus and
norvegicus in a riparian habitat. Journal of Mammalogy 63:151-154.
122. Suckley, G., and J. G. Cooper. 1860. The natural history of
Washington Territory and Oregon. Final Reports. On the survey of the
Northern Pacific Railroad route. Bailliere Brothers, New York, NY.
123. Sugihara, N. G., L. J. Reed, and J. M. Lenihan. 1987. Vegetation of the
Bald Hills oak woodlands, Redwood National Park, California. Madrono
34:193-208.
124.Taylor, R. J., and T. R. Boss. 1975. Biosystematics of Quercus garryana in
relation to its distribution in the state of Washington. Northwest
Science 49:49-57.
125. Van Denburgh, J. 1922. Reptiles of North America: an account of the
species known to inhabit California, Alaska, British Columbia,
Washington, Oregon, Idaho, Utah, Nevada, Arizona, Sonora, and Lower
California. Volumes 1 and 2. California Academy of Sciences, San
Francisco, CA.
126. Verts, B. J., and L. R. Carraway. 1980. Natural hybridization of
Sylvilagus bachmani and introduced S. floridanus in Oregon.The Murrelet
61:95-98.
127. ———, and ——— 1981. Dispersal and dispersion of an introduced
population of Sylvilagus floridanus.The Great Basin Naturalist
41:167-175.
128. ———. and ———. 1998. Land mammals of Oregon. University of
California Press, Berkeley, CA.
129. ———., S. A. Ludwig, and W. Graf. 1972. Distribution of eastern
cottontail rabbits in Linn County, Oregon, 1953-1970. The Murrelet
53:25-26.
130. White, R. 1980. Land use, environment, and social change: The
shaping of Island County, Washington. University of Washington Press,
Seattle, WA.
131. Wiens, J. A. 1969. An approach to the study of ecological
relationships among grassland birds. Ornithology Monograph 8:1-93.
132. ———. 1973. Interterritorial habitat variation in grasshopper and
savannah sparrows. Ecology 54:877-884.
133. ———, and M. I. Dyer. 1975. Rangeland avifaunas: Their composition,
energetics and role in the ecosystem. Pages 146-181 in Proceedings of
the symposium on management of forest and range habitats for
nongame birds. U.S. Forest Service General Technical Report WO-1.
134. Wilson, M.V., E. R. Alverson, D. L. Clark, R. H. Hayes, C. A. Ingersoll,
and M. B. Naughton. 1995. The Willamette Valley Natural Areas
Network. Restoration and Management Notes 13(1):26-28.
135. Witmer, G. W., R. D. Sayler, and M. J. Pipas. 1996. Biology and habitat
use of the Mazama pocket gopher (Thomomys mazama) in the Puget
Sound Area Washington. Northwest Science 70(2): 93-98.
136. Wolff, J. O., and E. M. Schauber. 1996. Space use and juvenile
recruitment in gray-tailed voles in response to intruder pressure and
food abundance. Acta Theriologica 41:35-43.
137. Wolff, J. O., W. D. Edge, and R. Bentley. 1994. Behavioral and
reproductive biology of the gray-tailed vole. Journal Mammal. 75:873-
879.
138. Wolff, J. O., T. Manning, S. M. Meyers, and R. Bentley. 1996. Population
ecology of the gray-tailed vole (Microtus canicaudus). Northwest
Science 70:334-340.
139. Wolff, J. O., E. M. Schauber, and W. D. Edge. 1997. Effects of habitat
loss and fragmentation on the demography of gray-tailed voles.
Conservation Biology 11:945-956.
140. Woodcock, A.R. 1902. An annotated list of the birds of Oregon.
Oregon Agricultural Experiment Station Bulletin 68.
141. Zoloth, S. R. 1969. Observations of the population of brush rabbits
on Ano Nuevo Island, California. The Wasmann Journal of Biology
27:149-161.
 11
Wildlife of Eastside Shrubland
and Grassland Habitats
W. MatthewVander Haegen, Scott M. McCorquodale, Charles R. Peterson,
Gregory A. Green, & EricYensen

Introduction
The rain shadow of the Cascade Mountains gives rise to a Although pristine climax communities do exist for
suite of arid and semi-arid habitats that differ substantially eastside grassland and shrubland habitats, the majority
from those of the surrounding forest. O’Neil and of sites have been shaped by a legacy of past land uses
Johnson100 have classified these habitats into 6 different that includes continuous grazing by livestock and range
types: shrub-steppe, dwarf shrub-steppe, desert playa and improvements to increase livestock forage and that in turn
salt scrub, western juniper and mountain mahogany has facilitated invasion by exotic vegetation. This legacy
woodlands, eastside canyon shrublands, and eastside has modified the vegetation community in many areas,
grasslands. Most of these communities are dominated by with some changes occurring so long ago that they are
shrubs and herbaceous vegetation (grasses and forbs) and not apparent to the present day observer.22 Changes in the
typically have a microbiotic crust of lichens and mosses herb community brought about by excessive grazing and
binding the upper surface of the soil. With the exception exotic invaders are particularly damaging in these arid
of the western juniper and mountain mahogany habitats where the herb layer often contains the most
woodlands habitat type, trees in these communities are vegetation biomass. Moreover, the successional trajectory
limited mainly to riparian zones and ecotones with of vegetation communities in arid habitats can be modified
forested habitats and are entirely absent from extensive by influences such as grazing and fire, resulting in present
areas. Much of the historical vegetation in these habitat day “zootic” climax communities that differ greatly from
types, particularly in Washington and north central those which occurred historically.22, 128 Sites in southcentral
Oregon, has been converted to agricultural crops.27, 113 In Washington that were dominated by exotic annuals in the
some areas, the only remaining native communities are 1950s still have not been colonized by native plants some
on rocky soils or steep slopes unfit for agriculture. The four decades later.115
dominant land use in these shrubland and grassland The low vertical structural diversity inherent in these
habitats is livestock grazing, and few examples of habitats provides fewer habitat layers for wildlife,
undisturbed stands exist, limited primarily to sites where resulting in lower diversity in some taxa. There are, for
topography or remoteness from water has made access example, no arboreal mammals or canopy nesting birds.
for livestock grazing impractical.22

Photograph 1. Shrub-steppe habitats

dominate the landscape in much of
eastern Oregon and Washington. Moses
Coulee, Douglas County, Washington.
(M. Vander Haegen,WDFW).

292
 CHAPTER 11: SHRUB-STEPPE 293

Habitats with a shrub component tend to have more characteristics that allow them to tolerate arid conditions.
diverse wildlife communities than grass dominated Birds excrete nitrogenous waste as uric acid, a
habitats, a function of increased nesting and foraging characteristic that they share with reptiles and that uses
strata. For example, the shrub-steppe habitat has 49 closely <10% of the water required by mammals excreting urea.78
associated species, whereas eastside grassland has only Birds also operate at a higher body temperature than do
34. Sites dominated by native plants have more closely most other animals, allowing them to cool themselves by
associated species than sites dominated by exotics (34 convection at most ambient air temperatures. Some arid-
species closely associated with eastside grasslands vs. 2 land birds (e.g., black-throated sparrow) can drink and
with modified grasslands). process water with a high saline content.78 Reptiles are
Available water is a defining factor in these arid and particularly well adapted to arid environments. Their dry,
semi-arid habitats, and this strongly shapes the scaley integument and lack of exocrine glands reduces
composition of plant communities22 and influences the water loss to the environment17 and, unlike amphibians,
ecology and behavior of associated wildlife. In they can reproduce independent of standing water.
Washington, precipitation occurs primarily during late Many arid-land species modulate their activities to
autumn and winter with annual totals ranging from 5.9 avoid temperature extremes, seeking shade during the
inches (150 mm) in the lowest parts of the Columbia Basin heat of the day or, in the case of the rock wren, seeking the
to 21.7 inches (550 mm) in the higher elevations near the cooler environment offered by rock crevices.78 Fossorial
forest ecotone. Annual snowfall can be substantial at species find cooler temperatures and reduced water loss
higher elevations and snow can remain in colder areas underground. Seeking milder conditions in underground
into spring. Growth of vegetation in spring is affected by
available soil moisture, a result of “bioyear ”
Table 1. Living and breeding in arid and semi-arid
precipitation—water that falls as rain or snow from
environments: constraints, consequences, and animal
October-April and is stored in the soil. 22 Bioyear
adaptations.
precipitation varies widely from year to year, affecting
plant growth69 and influencing both forage for herbivorous
wildlife and populations of herbivorous insects that form A. Environmental Constraints
the food base for many reptiles and breeding birds. The High and low temperature extremes
marked seasonality in precipitation creates a flush of High winds
available food in spring and early summer that is Open habitats
capitalized on by breeding birds. In the lower rainfall Aridity
zones most vegetative growth is completed by early Strong temporal resource gradients
summer and many plants senesce in preparation for the Fragmented habitats
dry, hot months ahead. High annual variability in rainfall
B. Ecological Consequences
Adaptations to Arid, Seasonally Hot
Cooling and warming adaptations required
and Cold Environments Increased variability in annual reproductive output
Hot and arid conditions that prevail in many of these Need to disperse across unsuitable habitats
habitat types in summer play a part in determining the Water conservation required
animal life that can persist there. Daytime temperatures
can exceed 113oF (45oC) and free water can be scarce. Some C. Adaptations
species are physiologically adapted to survive in such an • Physiological
Facultative breeding
environment, whereas others modify their behavior—
Flexible reproductive timing
some survive by a combination of both strategies (Table
Ability to process hypersaline water
1). Species that require daily access to free water (e.g., bats,
Ability to survive without free water
elk) must restrict their daily use areas to include open
Large ears for cooling (jackrabbits)
water. Mourning doves must drink daily and frequently Speed for eluding prey (pronghorn)
fly great distances to reach free water.78 Other species drink • Behavioral
infrequently and conserve body water by minimizing Cryptic behavior
water lost through evaporation, respiration, and excretion. Shade-seeking
These species meet their daily water requirement through Burrow into soil to avoid heat extremes
moisture contained in food and from metabolic water Daily trips to available water
produced from oxidation as part of digestion.17 The
grasshopper mouse and the sage sparrow are examples D. Life History Adaptations
Cryptic plumage/pelage
of species that spend considerable time far from free water.
Strong seasonality in habitat use
Black-throated sparrows are particularly well adapted to
Colonial living
life in arid environments; experiments have shown that
Accelerated larval development
they can survive on a diet of dry seeds without access to
Hibernation or migration during winter
free water and without restricting their activity.78 Estivation during summer
Birds and reptiles have some physiological
 294 Wildlife–Habitat Relationships in Oregon and Washington

burrows can be a temporary tactic used during the heat

of the day, or it can be a way to escape harsh conditions
for much longer periods. Some fossorial mammals,
including ground squirrels and pocket mice, may estivate
during the driest months and emerge from their burrows
when the winter rains renew vegetation growth. Great
Basin spadefoot toads are one of few amphibians to live
in arid habitats, surviving the arid summer months by
estivating underground. Members of this genus minimize
water loss by burrowing underground and by allowing
urea levels in the blood to rise as the soil around them
dries. Sufficiently high internal osmotic pressures create
water potentials that should allow spadefoot toads to
absorb water from even very dry soil.111 Spadefoot toads
are also capable of tolerating considerable loss of body
water (up to nearly 50%).138
Spadefoot toads may spend many months burrowed
deep in the soil, emerging to feed or to breed on rainy
nights. Their facultative breeding system is well adapted
to arid habitats, allowing them to track local conditions
and breed after rain events or, in more recent times, after
irrigation has wetted the soil.98 They are adapted to
temporary breeding ponds by needing only 2-3 days for
egg hatching and only a few weeks for development to
metamorphosis. Some tadpoles may feed on carrion as
well as vegetation, which also may result in more rapid
development. Tadpoles of some other species of spadefoot
toads are known to be cannibalistic.98
Cold conditions may limit the activity times of
ectotherms daily and seasonally; however, their low
energetic requirements (typically <10% of those of similar
sized endotherms) allow amphibians and reptiles to Photograph 2. Cliffs provide valuable roost sites for bats as
survive for extended periods of time without eating. For well as nest sites for numerous species of shrub-steppe
example, an adult rubber boa, maintained in captivity wildlife. Grand Coulee, Douglas County, Washington. (M.
under simulated field thermal gradients, voluntarily Vander Haegen, WDFW).
fasted for 23 months before eating, with no apparent long-
term ill effects (Michael E. Dorcas, Davidson College, pers.
comm.). Such fasting abilities enable these animals to within territories (e.g., Mojave black-collared lizards),
survive conditions such as droughts when food nesting sites (e.g., ringneck snakes), and gestation sites
availability may be low. The low metabolic rates of (e.g., western rattlesnakes). Rocky features are often
amphibians and reptiles also result in low rates of needed to provide the temperature gradients required for
respiratory water loss.111 effective behavioral thermoregulation.56 Rocky features
such as talus slopes and crevices within lava flows are
Habitat Elements— especially important as overwintering sites for snakes, and
Special Habitat Features their availability may limit the distribution of some
Some wildlife species of eastside grassland and shrubland species.43 Often, overwintering sites are used by multiple
habitats are strongly associated with special features species (e.g., striped whipsnakes, gopher snakes, western
(habitat elements) on the landscape and are unlikely to be terrestrial garter snakes, and western rattlesnakes). Talus
found in their absence. Many species are associated with slopes and talus-like structures (rock piles, lava stringers)
geological formations such as rock outcrops, cliffs, and are associated with 22 species and provide refuge for small
talus slopes. Peregrine falcons, cliff swallows, and golden mammals like the least chipmunk, and hibernacula for a
eagles nest on cliffs and rock faces and are among the 35 variety of snakes including the western rattlesnake. Some
species associated with these structures. All of the snake amphibians (e.g., long-toed salamanders and Pacific
species and about half of the lizard species in shrub-steppe treefrogs) may be associated with talus. Rocky outcrops
habitats are associated with rocky features (individual provide nest sites for ferruginous hawks and habitat for
rocks, talus slopes, outcrops, cliffs, ridges, caves, crevices, rock wrens, yellow-bellied marmots, and 42 other species.
etc.) that serve a variety of functions including providing Caves are used by 18 species, including bobcats and
foraging locations (e.g., side-blotched lizards), retreat sites common ravens, and are critical habitats for bats for
from predators (e.g., western fence lizards), vantage points roosting and hibernation.
 CHAPTER 11: SHRUB-STEPPE 295

Burrowing owls and fossorial mammals like the Anthropogenic structures are frequently used for
Columbian ground squirrel and badger require deep soils nesting sites or as shelter by a variety of wildlife. Common
for constructing nests and dens. Washington ground ravens, American crows, and several buteonid hawks nest
squirrels used sites where soils were deeper, weaker, and on power transmission towers. Construction of such
contained less clay than adjacent, unoccupied sites.4 Soil towers was responsible for an increase in the number of
type also can affect the persistence of underground breeding ferruginous hawks on the Hanford Site in
burrows. In an Oregon study, nest burrows of burrowing southcentral Washington in the 1980s.32 Cement road
owls were more likely to be reused in subsequent years culverts and bridges are used as shelter or as nest sites by
when constructed in silty-loam soil; burrows in loamy- cliff swallows and 13 other species; they provide
sand soils were often silted in and thus unusable by the appropriate nesting structure previously missing in many
next breeding season.38 Loose soils are important to areas. Garter snakes often use the foundations of bridges
burrowing species like Great Basin spadefoot toads, and buildings for nocturnal retreat sites and overwintering
sagebrush lizards, and horned lizards. Several species of sites. Western fence lizards and racers use stone walls for
reptiles (e.g., long-nosed leopard lizards) bury their eggs retreat sites and foraging areas.
in loose soils. Abandoned homesteads and farm buildings provide
The burrows of other animals are often used by unique vertical structure in many areas of shrubland and
burrowing owls, many species of lizards and snakes,134 grassland habitat. Old buildings are used as nest sites by
and some species of amphibians (e.g., tiger salamanders kestrels, barn owls, deer mice, and numerous other
and western toads). Burrows may serve a variety of species. Abandoned buildings and trash piles may attract
functions, including providing retreats from predators small mammals and thus reptiles that prey on them (e.g.,
(e.g., long-nosed leopard lizards), foraging sites (e.g., western rattlesnakes). Gopher snakes may use wells for
rubber boas), egg deposition sites (e.g., collared lizards), estivation sites (C.R. Peterson, Idaho State University, pers.
and thermal gradients for regulating body temperature. obs.). Trees planted as windbreaks around homesteads and
The use of burrows by reptiles and amphibians in dry orchards provide nesting structure for red-tailed hawks,
shrub-steppe habitats is particularly important for black-billed magpies, orioles, and numerous other birds.
minimizing evaporative water loss. Abandoned farm equipment serves as nesting sites for
All amphibians in eastside shrubland and grassland European starlings, domestic pigeons, western kingbirds,
communities are associated with standing or slow-moving and other birds as well as various small mammals.
water sometime during their life cycle, because it is
required for breeding and larval development. Several Bird Communities in Shrub-steppe
species of reptiles also are associated with water (i.e., The following wildlife community profiles focus on
marshes, pools, ponds, lakes, streams, or rivers). Western shrub-steppe habitat, with references to other eastside
skinks and rubber boas are often found near moist areas. shrubland and grassland habitats where appropriate. We
Garter snakes often forage in the water or along shorelines. selected shrub-steppe as a focal habitat based on its
Painted turtles overwinter, breed, and forage in water. dominance in eastside landscapes and because of the
Human-made stock ponds have probably increased the severe management issues that this habitat currently faces.
amount of suitable habitat for some of these water Shrub-steppe bird communities are characterized by a
associated species. relatively small number of breeding species. The

Photograph 3. Talus slopes and talus-

like structures provide hibernacula for
western rattlesnakes. Grant County,
Washington. (M. Vander Haegen,
WDFW).
 296 Wildlife–Habitat Relationships in Oregon and Washington

assemblage of passerines, for example, typically totals 4- northern sites but spend all or part of the winter in the
8 birds and often is dominated by a single species.156 shrub-steppe, including rough-legged hawks and
Extensive surveys including many sites may generate a northern shrikes.
much larger species list;27 however, many species on these Two native galliforms, greater sage-grouse and sharp-
longer lists occur infrequently or breed in other habitats tailed grouse, live in shrub-steppe communities of Oregon
and are tallied on surveys as they forage or migrate and Washington. Both are ground nesters and require large
through the shrub-steppe. For example, three years of areas for their annual home ranges, including open areas
surveys on one site in the shrub-steppe of eastern for leks in the spring. Both grouse are listed as threatened
Washington produced a list totaling 28 species; however, in Washington, and the sharp-tailed grouse was only
only 5 species were documented to nest on the site recently reintroduced to Oregon after having been
(Washington Department of Fish and Wildlife [WDFW] extirpated in the 1960s.99 Loss of habitat and fragmentation
unpubl. data). Fragmentation of formally extensive shrub- by agriculture are believed to be primary causes for the
steppe and new habitats associated with agriculture and decline of these species. 16, 124 Exotic galliforms that
irrigation contribute species that are not typical of the frequently can be found nesting in shrub-steppe include
shrub-steppe community type. the chukar, gray partridge, and ring-necked pheasant.
What the shrub-steppe bird community may lack in Numerous raptors use shrub-steppe for nesting and
variety, it makes up for in specificity. Several species foraging, preying on a variety of small mammals, reptiles,
associated with shrub-steppe are so dependent on sage birds, and insects. Burrowing owls and short-eared owls
cover that they are termed sagebrush obligates. Sage and nest on the ground in shrub-steppe and can be found
Brewer’s sparrows, sage thrashers, and sage grouse are nesting in stands of big sage, antelope bitterbrush (Purshia
considered obligates, whereas vesper sparrows and green- tridentata), or open, low grasslands. Northern harriers
tailed towhees are classified as near-obligates.8 These often forage in shrub-steppe, especially along edges with
species often characterize the big sagebrush (Artemisia agriculture, but they require wetlands or similar areas of
tridentata) shrub-steppe community, although other shrub- thick reeds or grasses for their ground nests. Most raptors
steppe or grassland-associated species may dominate at require elevated nesting sites and historically nested on
specific sites. In big sagebrush communities in buttes, cliffs, and in riparian-associated trees. In
northcentral Oregon and southcentral Washington, the northcentral Oregon, juniper trees were important nesting
breeding bird community was dominated by sage platforms for ferruginous and Swainson’s hawks, and
sparrows and western meadowlarks54, 146 (WDFW unpubl. long-eared owls.39, 54 Juniper trees also were important as
data). In northcentral Washington, the breeding bird nest sites for ferruginous hawks in southcentral
community was dominated by Brewer’s sparrows and Washington. 31 Artificial planting of trees and the
vesper sparrows (WDFW unpubl. data). Disturbed sites proliferation of power transmission lines during the last
with few shrubs frequently are dominated by western century have increased the availability of suitable nest sites
meadowlarks, horned larks, grasshopper sparrows, and and likely have boosted raptor populations in some
long-billed curlews.54, 146 areas.119
Most passerines that breed in shrub-steppe eat insects A recent analysis of population trends using the
and other arthropods at some period during the year. Some Breeding Bird Survey (BBS)122 identified 8 shrub-steppe-
birds are primarily insectivorous, consuming a variety of associated species that are declining in the interior
invertebrates that they capture on the wing, or glean from Columbia River Basin. Four of these species, the Brewer’s
shrubs or herbaceous vegetation. Even granivorous birds sparrow, lark sparrow, loggerhead shrike, and western
that feed mostly on seeds and plant material as adults feed meadowlark, are closely associated with shrub-steppe.
invertebrates to their young to supply critical nutrients. Numerous other birds, including the sage sparrow,
Total precipitation during the bioyear affects primary and grasshopper sparrow, and burrowing owl, likely are not
secondary production and varies widely among years in monitored adequately by the BBS and will require
this ecosystem.22 This yearly variation in rainfall can affect specialized monitoring to detect changes in their
reproductive success of shrub-steppe nesting birds, populations.122
probably through its influence on availability of arthropod
prey.121 Organizing Principles: Birds
The community of breeding birds in shrub-steppe is Shrub-steppe communities extend from the northern
largely comprised of migrants, many (n = 16) wintering border of Washington to the southern border of Oregon.
south of the United States and therefore termed Neotropical Although some shrub-steppe-associated birds are
migrants. Spring migrants that pass through the shrub- common to much of this area, there are differences in avian
steppe on their way to more northern breeding grounds species assemblages across this latitudinal gradient. For
and that spend time in the sagebrush communities include example, black-throated sparrows and green-tailed
the white-crowned sparrow and ruby-crowned kinglet. towhees reach the northern extent of their range near
Resident species are largely gallinaceous birds (e.g., greater Oregon’s northern border and occur only sporadically in
sage-grouse and sharp-tail grouse) and corvids (e.g., Washington. Sharp-tailed grouse occur in several locations
common raven and black-billed magpie). The winter bird in northcentral and eastern Washington, but were
community is supplemented by species that breed in more extirpated from Oregon and have only recently been
 CHAPTER 11: SHRUB-STEPPE 297

Photograph 4. Sage sparrows are

sagebrush obligates and nest in
extensive tracts of shrub-steppe. Moses
Coulee, Douglas County, Washington.
(M. Vander Haegen, WDFW).

reintroduced.99 Bird communities also may change along billed magpies and a host of raptors, including red-tailed
an elevational gradient in shrub-steppe, as rainfall and Swainson’s hawks, and long-eared owls. Without the
increases with elevation and changes the vegetation vegetation structure to support nests and foraging
community. Vesper and Brewer’s sparrows occur rarely activities, these species are unlikely to establish territories
in sage communities on the low elevation Columbia River and breed successfully.
Plain, whereas they are regularly found on ridges only a Floristics also play a part in determining the
few kilometers to the west in sage communities above composition of the avian community in shrub-steppe. In
5,900 feet (1,800 m).146 As one proceeds north in the their extensive study of shrub-steppe birds that included
Columbia Basin, elevation increases and these species sites in Washington, Oregon, and Nevada, Wiens and
become common at all elevations. In fact, these two Rotenberry158 found that whereas habitat structure played
sparrows, along with the sage thrasher, typify sage a part in determining species occurrence at a
communities in northcentral Washington (WDFW unpubl. biogeographic scale, presence of particular plant species
data). was important to some birds at a regional scale. Sage
Presence and abundance of individual bird species vary sparrows, true to their name, prefer a shrub community
with a range of local and landscape variables. On a gross dominated by sagebrush over other species of shrub.158
scale, vegetation structure determines what species can Even within the sagebrush family, sage sparrows show a
breed in a community through presence of suitable nesting preference for stands of big sagebrush. Recent work in
and foraging strata. A suite of grassland-associated birds Washington has shown that both sage sparrows and
that breed in shrub-steppe includes the grasshopper Brewer ’s sparrows occur at greater abundance in
sparrow, horned lark, western meadowlark, and long- communities of big sagebrush than in stiff sagebrush (A.
billed curlew. These birds nest on the ground and depend rigida) communities typical of rocky soils.147 Wiens and
on grasses and forbs to conceal their nests. Native Rotenberry158 found abundance of Brewer’s sparrows to
perennial grasses generally dominate the ground layer in be negatively correlated with cover of spinescent shrubs
undisturbed shrub-steppe communities, and presence of such as hopsage and budsage. These authors suggested
several ground-nesting species, including the grasshopper that some shrub-steppe birds key on particular shrubs to
sparrow and horned lark, increased with percent cover of take advantage of arthropod foods particular to those
these grasses. 147 In contrast, long-billed curlews in shrub species.
northcentral Oregon104 and southeast Washington2 seemed Although we have only recently begun to examine
to prefer short-statured annual grasslands for nesting and spatial components of community structure, there is
foraging, an apparent adaptation to this recent, evidence that landscape characteristics influence the
disturbance-related vegetation community. occurrence of some shrub-steppe species. In Idaho, the
Shrub cover is a requirement for the suite of shrub- probability of finding sage sparrows and Brewer ’s
nesting birds that nest in Washington and Oregon’s shrub- sparrows in patches of sagebrush increased with size of
steppe. Sage, Brewer’s, and lark sparrows, along with sage the patch.63 Recent work in Washington has shown patch
thrashers, and loggerhead shrikes, nest in or immediately size to be very important to sage sparrows, with males
beneath shrubs. They also use shrubs for singing and establishing territories and nesting only on sites many
foraging perches, and Brewer ’s sparrows forage times larger than an average territory (WDFW unpubl.
extensively within the foliage of shrubs.162 Presence of data). The landscape context in which these spatial effects
trees, either along riparian areas or planted as windbreaks are examined also may have a bearing on how species
or near homesteads, provides nesting platforms for black- react. In extensive shrub-steppe in Idaho, where sagebrush
 298 Wildlife–Habitat Relationships in Oregon and Washington

Photograph 5. Young burrowing owls near their

nest burrow. Burrowing owls in the Columbia
Basin depend on fossorial mammals both for
nest sites and for much of their diet. Grant
County, Washington. (D. Hoyt, WDFW).

communities were fragmented by fire and subsequent Great Basin pocket mice, comprised the majority of the
cheatgrass (Bromus tectorum) invasion, sage thrashers biomass in pellets collected at nest sites.40
occurred more frequently in spatially similar sites with Brown-headed cowbirds may use different habitats for
low fragmentation of sage.63 However, in Washington, feeding and for breeding and selection of both may be
where sagebrush stands are fragmented by agriculture, driven by the presence of other species. Cowbirds likely
thrashers were regularly found in small fragments of sage evolved with large, grazing animals (i.e., bison) of the
and were more likely to occur in fragmented than short-grass prairies of the Midwest, following the animals
extensive sites. 147 For some species, these landscape as they grazed and eating seeds and insects exposed by
features likely interact with local vegetation characteristics trampling hooves.74 In the shrub-steppe of Oregon and
to determine suitability of a site. Landscape characteristics Washington, cowbirds are typically associated with
such as patch size and spatial similarity seem to have little livestock and can be found foraging among cattle and
effect on occurrence of some of the more generalist birds horses in pastures and feedlots. Cowbirds are “nest
such as western meadowlarks and horned larks, at least parasites” who lay their eggs in the nests of other birds,
at the scales studied.63, 147 and therefore require suitable populations of “host”
Another physical variable that influences the bird species to reproduce successfully. In big sagebrush
community in shrub-steppe is soil type. Texture and depth communities in eastern Washington, cowbirds typically
of the soil can affect its suitability for foraging and nesting, parasitized sage, Brewer’s, and vesper sparrows, although
and composition of the vegetation community is rates of parasitism were low compared to other
influenced by soil characteristics. Burrowing owls nest in communities.148 Cowbirds also parasitize birds that nest
underground burrows and require deep, friable soils. Sage in riparian communities within shrub-steppe,33, 102 and
and Brewer ’s sparrows in Washington were more their distribution likely is tied to that of suitable hosts in
abundant in deep soil communities characterized by big these communities as well.
sagebrush than in shallow soil communities dominated
by low-growing stiff sagebrush.147 Size of the sage shrubs Mammal Communities
affects their suitability as nest sites and perhaps also
determines availability of insect prey. In this same analysis,
in Shrub-steppe
loggerhead shrikes occurred in greatest numbers, and Species richness is typically related to the structural
western meadowlarks in lowest numbers, in sandy soil complexity of dominant vegetation.18, 116 Not surprisingly,
communities typical of the Columbia River Plain. the diversity of mammals in the shrub-steppe of western
Mourning doves were the only species that occurred at North America is lower than that typical of structurally
greater abundance in shallow soil communities. more complex habitats of the region. For example, 40 small
Occurrence of some shrub-steppe birds may be related mammal species are closely associated with forested
to the presence of other animal species. Burrowing owls habitats of Oregon and Washington, whereas only 20 are
in the West are closely tied to populations of fossorial closely associated with shrub-steppe habitat. Ten carnivore
mammals and the vegetation communities and soil types species are closely associated with forested habitats
that support them.38, 49 Burrowing owls may depend on compared to 2 in shrub-steppe. Because of low
badgers for nest sites in the Columbia Basin.38 Although precipitation, high incident solar radiation and wide
burrowing owls eat a variety of prey, small mammals are fluctuations in seasonal temperatures, the shrub-steppe
a key component and western populations often are is a challenging environment for homeotherms. Not
associated with colonies of ground squirrels and other surprisingly, many shrub-steppe mammals are relatively
small, burrowing mammals. In the Columbia Basin of specialized for life in this arid region.
Washington and Oregon, small mammals, particularly In addition to the relatively few mammals that are
 CHAPTER 11: SHRUB-STEPPE 299

Photograph 6. Pygmy rabbits are sagebrush

obligates, building their burrows in deep soils
beneath sagebrush plants and feeding on
sagebrush foliage. Douglas County, Washington.
(WDFW).

clearly associated with true shrub-steppe vegetation, dominated areas and black-tailed jackrabbits predominate
numerous species may be associated with very specialized in areas with a strong shrub component.19, 143 Areas
habitats that occur as minor components of shrub-steppe occupied by jackrabbits are obvious due to their practice
systems. For example, black bears, beavers, and muskrats of clipping vegetation to make runways, which they travel
are not considered shrub-steppe species, but all may occur repeatedly.97 Pygmy rabbits are uncommon and typically
in larger riparian corridors that extend from forested occur only in areas dominated by tall, dense stands of
communities into shrub-steppe. Great Basin or big sagebrush, which provide preferred
forage.19, 41, 153 Pygmy rabbits also excavate burrows in
Small and Meso-sized Mammals which they den, and therefore also require areas with
Small mammals of the shrub-steppe include ubiquitous friable soil. Mountain cottontails often are associated with
species such as the white-footed deer mouse and several distinct microhabitats such as riparian areas or rocky
species that are relatively restricted to shrub-steppe and ravines and also occur in areas developed by humans.19
other arid biotic associations. Hedlund and Rogers50 Yellow-bellied marmots are not associated with shrub-
trapped Great Basin pocket mice, northern grasshopper steppe associations except where rock piles occur as
mice, sagebrush voles, western harvest mice, and deer special, localized habitat features. Bushy-tailed wood rats,
mice in shrub-steppe habitat of the Hanford Site of Columbian ground squirrels, Belding’s ground squirrels,
southcentral Washington. Marr et al.,79 trapping another and a complex of smaller ground squirrels (discussed
area of the Hanford Site, caught deer mice, western harvest below) also occur in shrub-steppe habitats of eastern
mice, grasshopper mice, montane voles, Great Basin Oregon and Washington.
pocket mice, and northern pocket gophers. Rogers and
Rickard120 also listed vagrant shrews as occurring in shrub- Carnivores of the Shrub-Steppe
steppe habitat of southcentral Washington. Although small Because diversity and biomass tend to decline with
mammal communities of the shrub-steppe commonly increasingly higher order consumers,116 the diversity and
contain several of these species, a few species, notably abundance of shrub-steppe carnivores is a fraction of that
Great Basin pocket mice and deer mice, numerically of small and meso-sized mammals. Among the common
dominate most assemblages in eastern Washington.36, 50, 79 carnivores are the ubiquitous coyote, the more habitat
Although uncommon in Washington, kangaroo rats, such restricted badger, and the widespread long-tailed weasel.
as Ord’s kangaroo rat and the less common chisel-toothed The kit fox reaches its northern range limit in arid plant
kangaroo rat, inhabit shrub-steppe in eastern Oregon.19 associations of southeastern Oregon.19
The least chipmunk and dark kangaroo mouse also occur Coyotes, badgers, and weasels consume a diverse array
in some areas of shrub-steppe.19 of small mammals, insects, birds, and reptiles. Carnivore
Meso-sized mammals of the shrub-steppe include densities are related not only to densities of their prey but
several lagomorphs, such as white-tailed and black-tailed also to complex social factors.92, 126, 130 Coyote, weasel, and
jackrabbits, mountain cottontails, and pygmy rabbits. The badger occupancy of shrub-steppe habitats is probably
2 jackrabbit species are sympatric; however, white-tailed linked to vegetation indirectly via effects of vegetation on
jackrabbits are generally less common and tend to be prey diversity and abundance. The bobcat, another
distributed in smaller, isolated populations, at least in widespread species, occurs in shrub-steppe where rock
recent years.19, 80 Both species are prone to cyclic population outcrops, ravines, or lava formations are available for
growth, 44, 64, 97 and variation between high and low suitable den sites. 62 Mountain lions are the largest
population levels in black-tailed jackrabbits can be carnivores that use shrub-steppe habitat and typically
dramatic.3 When the 2 jackrabbit species occur together, occur only where mountainous terrain and shrub-steppe
white-tailed jackrabbits tend to occupy more open, grass- vegetation are contiguous.19, 20 Bobcats and mountain lions
 300 Wildlife–Habitat Relationships in Oregon and Washington

prefer prey at least as large as a hare, and are found only habitat features seem to be strong determinants of habitat
where suitable prey biomass is relatively high. Not suitability for some shrub-steppe carnivores.
surprisingly, bobcat and mountain lion reproductive Occurrence and density of primary consumers would
success and spatial use of the landscape varies with cyclic normally be expected to be directly linked to community
changes in densities of primary prey, whether lagomorphs floristics. However, generalists such as the larger
or ungulates.62 Mountain lions tend to be found only where herbivores seem to have broad tolerances for plant
large mammal prey also are available. associations as long as acceptable forage biomass is above
critical limits that relate to foraging efficiency thresholds.
Large Herbivores Primary consumers with specialized diets, such as pygmy
Native species of large herbivores in the shrub-steppe rabbits, may be very selective of plant associations.41 Black-
include generalists such as the mule deer and elk, as well tailed jackrabbits are seasonally selective of forage type,
as more specialized ungulates such as the pronghorn but will consume a variety of plant species143 and therefore
antelope. Bighorn sheep are not a true shrub-steppe display more generalized affinities for plant associations
species and have been historically rare in arid mountain than pygmy rabbits. Black-tailed jackrabbits will generally
ranges of eastern Oregon and Washington, but bighorns not consume cheatgrass, and therefore largely avoids pure
did occur prehistorically in some mountainous areas cheatgrass swards. 143 Where cheatgrass invasion is
adjacent to shrub-steppe habitats and have been occurring, jackrabbits foraging on native bunchgrasses can
reintroduced to several locations in Oregon 101 and convey a competitive advantage to cheatgrass and
Washington.151 All of these large herbivores require free facilitate further conversion towards an annual
water and therefore are not found in extensive tracts of grassland.132
shrub-steppe that lack surface water such as springs or Mammal species vary considerably in their affinity to
ponds. Large mammal ecology in the shrub-steppe is associations with a strong sagebrush component. Pygmy
discussed in greater detail in a later section. rabbits are strongly associated with sagebrush, rarely
occurring where sagebrush is a minor component or
Organizing Principles: Mammals lacking from the plant community. Sagebrush voles have
The determinants of habitat suitability for mammals in a strong affinity for sagebrush but occur in stands lacking
the shrub-steppe undoubtedly varies across species. For a sagebrush overstory if grass understories are thick
example, floristics may be important for some species enough to provide cover.19 Pronghorns are the only large
because of diet specialization. For some species, however, herbivore in the shrub-steppe that frequently forage on
the vegetative structure of the association is probably more sagebrush. Pronghorns often show an affinity for
important than the specific assemblage of plant species sagebrush81, 135 and are most successful where sagebrush
that comprise the association. species are available for winter forage, although they can
Secondary and tertiary consumers are usually linked occupy areas without sagebrush if other acceptable forage
to vegetation associations indirectly via habitat affinities is available.125
of their prey species. But because most predators in the
shrub-steppe will consume a variety of prey organisms,62 The Role of Ground Squirrels in
their generalized affinities for plant communities reflect Shrub-Steppe Communities
the diversity of habitats used by their smaller mammalian In pre-European times, shrub-steppe habitats in eastern
prey. However, as noted, several shrub-steppe carnivores Oregon and Washington and adjacent states were
directly select structural and geological habitat features
because of their importance to other life history needs (e.g.,
den availability). Accordingly, these important structural

Photograph 7. Badgers feed extensively on ground

squirrels and are ecologically linked to another
ground squirrel predator, the burrowing owl, that
uses badger dens for nest sites. Morrow County,
Oregon. (G. Green, Foster Wheeler Environmental
Corporation).
 CHAPTER 11: SHRUB-STEPPE 301

inhabited by the smaller ground squirrels of the genus bunchgrasses by about 20%. 67, 68 In shrub-steppe
Spermophilus. Like other ground squirrels they were environments, ground squirrels are important to the very
sedentary, and rivers and other barriers were significant plants they eat.
obstacles to gene flow.24, 37 Consequently, they diverged Badgers feed extensively on ground squirrels, and areas
into a complex of 7 species, each with a separate range with high ground squirrel densities usually have high
but all with similar ecological roles. Larger species of this densities of badger digs.167 These badger digs also increase
genus, such as Columbian and Belding’s ground squirrels, water infiltration and aeration of the soil53 and provide
also inhabited shrub-steppe habitats, but were generally nest sites and shelter for a variety of wildlife. Thus, the
in deeper soils, and frequently in more mesic habitats, and presence of ground squirrels has positive effects on other
are not considered here. Each species occupied a separate animal species, as well as on soil and vegetation.
geographic area. In Washington, Townsend’s ground Invasion of exotic annuals has changed both the fire
squirrel lived between the Cascade Mountains and the regimes and successional patterns in shrub-steppe
Yakima River,47, 52 whereas a similar, but chromosomally habitats. Range fires burn much hotter in exotic annual
distinct form occurred between the Yakima and Columbia infested rangelands, killing shrubs and allowing exotic
Rivers. 96 The Washington ground squirrel formerly annuals to out-compete native species. In many areas,
occupied the Columbia Basin between the Columbia River native communities of shrubs, bunch grasses, and forbs
and the range of the Columbian ground squirrel to the have been replaced by annual grasses (especially
east, and extended south into the Columbia Basin of cheatgrass) and dicots (particularly mustards). Ground
Oregon.4 Merriam’s ground squirrel (S. canus canus) squirrels will eat exotic plants,166 but the productivity of
occupied the shrub-steppe communities in central Oregon, these invasive plants varies annually with precipitation,
except in Malheur County where it was replaced by the providing an unstable food base for ground squirrels. In
similar S. c. vigilis.47, 55, 149 These small ground squirrels southwestern Idaho, ground squirrel populations are
remain poorly studied, but available information indicates unstable in areas dominated by exotic annuals and are
that they prefer deep, well-drained sandy to loamy soils4, prone to extinction.167 This instability is compounded by
55
but otherwise occur in a variety of native vegetation the large amounts of indigestible silica in cheatgrass and
types.129, 149, 167 especially in medusahead (Taeniatherium asperum) seeds,
There is little information available on the abundance making them a poor food source regardless of the
of ground squirrels in pre-European times. It is likely that quantity.123
their distribution was patchy, with subpopulations acting Fire also affects ground squirrels by reducing shrub
as sources and sinks. Their distribution may have been cover. Although ground squirrels can survive in areas
driven by foraging needs, as small herbivores have a low without shrubs as long as the herbaceous layer remains
digestive capacity and therefore depend on increasingly unchanged,106 in the long term their density decreases
higher quality forage as body size decreases.25 Ground when shrub cover drops below 11%, probably due to loss
squirrels depend upon high quality forage and an of protective cover, moisture changes, or other factors.
abundant supply of seeds in order to store the fat necessary Ground squirrel density also drops when shrub cover
to survive 8 months of hibernation. High quality forage exceeds 20%, apparently due to reduced productivity in
in turn depends on wildfire and disturbance providing the herbaceous layer.59
young, succulent, productive growth. In presettlement Agricultural conversion has had an even more drastic
times, wildfires were patchy and post-burn succession effect than exotic annuals on ground squirrel populations.
generally maintained a mosaic of patches in various In the Columbia Basin, agricultural conversion has
successional stages165 that would have allowed ground reduced the Washington ground squirrel to a handful of
squirrels to move about the landscape monopolizing new isolated populations with low probability of long-term
growth as it appeared. survival 5 (E. Yensen, Albertson College, pers. obs.).
Ground squirrels are important as a prey base for many Agricultural conversion likely has had similar effects on
of the predators in their ecosystems. In Idaho, Paiute the Townsend’s ground squirrel.45 No systematic survey
ground squirrels are a keystone species, providing a critical has been done on Merriam’s ground squirrel, but it seems
food source for prairie falcons and an important prey to be extirpated over much of its range (E. Yensen,
source for red-tailed and ferruginous hawks, badgers, Albertson College, unpubl. data). In this case, vegetation
western rattlesnakes, and gopher snakes. They also are changes favoring the larger Belding’s ground squirrel may
prey for long-tailed weasels, ravens, and others.167 be responsible.
As burrowers, ground squirrels are important in mixing Despite their importance in shrub-steppe ecosystems,
soils.1, 53, 137 For example, arctic ground squirrels were ground squirrels generally have not been appreciated by
documented to move 19 tons of soil/acre/yr.112 They are humans. Ground squirrels do invade agricultural fields,
important in soil aeration,57 and fertilize the soil with their eat alfalfa, grain and other produce, and dig holes in
feces and urine,42, 112, 136, 137, 141 resulting in significantly irrigation ditches. In return, ground squirrels have been
greener vegetation in the vicinity of their burrows.112 The poisoned, trapped, used for target practice, and reviled
burrows significantly increase water infiltration into the for over a century with little understanding of, or interest
soil, which increases plant productivity. In an Idaho study, in, their ecological roles. Ironically, their demise has
ground squirrel burrows increased productivity of unfortunate ecological consequences for shrub-steppe
 302 Wildlife–Habitat Relationships in Oregon and Washington

ecosystems that may portend economic losses as well. The Contemporary shrub-steppe habitats are exploited by
decline of ground squirrels in some areas implies reduced 3 large ruminant herbivores: pronghorn antelope, mule
productivity of the rangeland and, thus, lower value to deer, and elk. Bison occurred historically in the shrub-
grazing cattle. steppe of southeast Oregon,145 and there is archaeological
evidence that they once occurred in the shrub-steppe of
Ecology of Large Herbivores in the Columbia Plateau.103 Of the 3 contemporary shrub-
Shrub-Steppe steppe ruminants, only the pronghorn is a steppe
Living in warm, arid climates presents unique challenges specialist. Pronghorns are widespread in steppe-like
to large mammals in terms of water conservation and habitats of western and midwestern North America,135 but
maintenance of homeothermy.94 As previously noted, the the shrub-steppe of eastern Washington and Oregon
availability of drinking water is an important component represents the fringe of their distribution. The pronghorn
of suitable habitat for large herbivores in the shrub- is well suited for occupation of open steppe-like country.12
steppe.30, 86 Large herbivores also have relatively high As previously noted, pronghorns have a pronounced
forage intake demands; this intake rate is strongly affinity for plant associations with a strong sagebrush
correlated with the biomass of acceptable forage.15, 155 component,135 and their distribution conforms relatively
Accordingly, minimum forage biomass would also be well to the distribution of areas occupied by Artemisia
expected to limit the types of arid environments that large species. Pronghorns are the only ruminant herbivore in
herbivores can effectively exploit.84 Among ruminant North America that forages heavily on sagebrush, a
herbivores, smaller animals tend to have lower intake relatively unpalatable and poorly digested shrub.
demands, but because of morphological and digestive Pronghorns also seasonally consume a diverse array of
constraints, require higher quality forage relative to larger forbs, although they generally consume little grass.135, 168,
169
ruminants.48, 58 Smaller ruminants can afford the costs of Pronghorns are relatively common in eastern Oregon
selective herbivory typically required to obtain high but have been extirpated in eastern Washington.135 The
quality diets. Larger ruminants have high intake demands factors relating to their disappearance in Washington are
and must maintain high intake rates in order to meet these not well understood, but competition with livestock for
demands within the amount of time they invest in palatable forb species and the generally large-scale
foraging. Therefore, it has generally been theorized that conversion of sagebrush-dominated rangelands to other
smaller ruminants are typically forage-quality limited, in uses are implicated.
contrast with larger ruminants that are expected to be Mule deer are generalists and occur in a relatively broad
constrained more by forage quantity.58 array of habitats of western North America from densely
Because of low primary productivity in the shrub- forested coastal areas to arid deserts of the southwest.150
steppe relative to more mesic habitats of the region, it Mule deer are relatively common in the shrub-steppe of
would seem that the shrub-steppe would be a more eastern Oregon and Washington, although their
suitable environment for small ruminant herbivores. distribution is far from uniform.70 On the Hanford Site of
However, many shrub-steppe associations are dominated southcentral Washington, mule deer densities are highest
by bunchgrasses and nonpalatable shrubs such as big near riparian areas such as the Columbia River shoreline,
sagebrush. Because of high silica content and relatively waste ponds, and perennial springs. 29, 144 Individual
thick cell walls, grasses are not easily digested by small Hanford Site mule deer also make disproportionate use
herbivores. of areas near water.30 Carson and Peek13 found that mule

Photograph 8. The Rattlesnake Hills elk

herd resides entirely in shrub-steppe
habitat in eastern Washington.
Elsewhere, shrub-steppe serves as
winter habitat for some migratory
populations of elk. Arid lands ecology
reserve, Benton County, Washington.
(S. McCorquodale, Yakama Nation).
 CHAPTER 11: SHRUB-STEPPE 303

1: The Rattlesnake Hills Elk Herd

Until recently, shrub-steppe communities were not Washington shrub-steppe seem to be well suited
thought to be year-round habitat for elk.10, 95 to exploitation by elk; forage biomass typical of
Although elk were known to use shrub-steppe many shrub-steppe communities is above that
habitat contiguous with forested habitats during considered minimum for elk.84, 155 In grass-
severe winters, elk were generally thought to be dominated communities, elk seem to be
precluded from year-round occupation of shrub- competitively superior to mule deer.
steppe due to low forage productivity and the Elk in the shrub-steppe use a broad array of
inability to tolerate high thermal loads. In 1972, plant associations. Telemetry data indicate that
however, a small group of elk colonized the shrub- stands with a sagebrush overstory are used
steppe habitat of the Hanford Site in southcentral heavily by elk for bedding, particularly during the
Washington.114 This group, commonly known as heat of summer.89 Foraging elk, however, have
the Rattlesnake Hills elk herd, has been studied shown an affinity for grass swards where
extensively. The original colonizing group of 4-7 historical fires have eliminated sagebrush.83 Elk
elk grew rapidly because of high reproductive diets are diverse in the shrub-steppe, with grasses
output and extremely high first-year survival.88 dominating spring and winter diets, and forbs
Despite the fact that shrub-steppe had been dominating summer and fall diets.82, 85
considered unsuitable year-round habitat for elk, The Rattlesnake Hills elk population has shown
individual fitness, indexed by reproductive little dependence on shrub forage, even during
success in females and age-specific antler growth winter, but the predominant shrubs in the herd
in males, has been exceptional in the Rattlesnake range, sagebrush, greasewood (Sarcobatus
Hills elk population.86, 87, 88 Currently this vermiculatus), and spiny hopsage (Atriplex spinosa)
population numbers >700 individuals (B. Tiller, are not considered good elk forage.
Battelle Memorial Institute unpubl. data). Other factors have contributed to the success of
Although primary production is low in shrub- the Rattlesnake Hills elk population. The portion of
steppe relative to forested environments, energetic the Hanford Site occupied by elk is topographically
modeling has indicated that shrub-steppe diverse,82 and access restrictions associated with
communities may rival or exceed more productive Hanford Site security are believed to limit
environments in terms of forage energy harassment to levels tolerable to elk.89 The presence
availability because all production occurs in the of perennial springs in the southern portion of the
herbaceous and shrub layers.84 Grass-dominated Hanford Site are probably essential to the
associations typical of eastern Oregon and thermoregulatory strategy of elk in high thermal
loading environments.89, 105

deer in northcentral Washington preferred riparian cover Reptile and Amphibian Communities
types and areas with some topographic diversity as
opposed to flat expanses of shrub-steppe vegetation. Mule
in Shrub-Steppe
deer in the shrub-steppe consume diverse diets typical of Relative to other classes of terrestrial vertebrates, diversity
a generalist herbivore, but forbs and the early growth of of amphibians in shrub-steppe habitat of Oregon and
shrubs are particularly important,144 consistent with Washington is low (10 of the 32 species occurring in these
energetic predictions. Large areas dominated by two states). Only 3 of 21 species of salamanders occur in
bunchgrasses and lacking palatable shrubs do not seem shrub-steppe habitat: long-toed salamander, tiger
to be vegetation types exploited effectively by mule deer.70 salamander, and roughskin newt. Seven of 11 native
Although mule deer and elk may be residents of the species of anurans occur in shrub-steppe. Great Basin
shrub-steppe region (see Box 1), some migratory spadefoot toad, western toad, and Woodhouse’s toad are
populations also use shrub-steppe exclusively as winter the anuran species most likely to be found in shrub-steppe
habitat. Where shrub-steppe occurs adjacent to low- away from standing water. In southeastern Idaho, Great
elevation xeric forest, suitable arrays of forage species and Basin spadefoot toads may be found in shrub-steppe as
biomass and favorable climatological features make shrub- far as 5 km from standing water (S. Cooper Doering and
steppe rangeland highly valuable winter range, especially C. Peterson, Idaho State University unpubl. data). Ranid
during severe winters. In eastern Oregon and Washington, frogs are most closely associated with wetlands. Columbia
large numbers of migratory deer and elk display this spotted frogs in southwestern Idaho, for example, spend
seasonally intense use of shrub-steppe winter range, most of their time within several meters of water (T.
although during the warmer months they use higher Carrigan, Bureau of Land Management unpubl. data).
elevation forests. The occurrence and distribution of some anuran species
has changed within the last 30 years. Northern leopard
 304 Wildlife–Habitat Relationships in Oregon and Washington

frogs have disappeared from many sites in Oregon and years in some species), suitable breeding habitat does not
Washington where they occurred previously.131 Bullfrogs have to be present every year. Temperature variation
have been introduced into Oregon and Washington and among shrub-steppe communities does not seem to
pose a threat to native species such as northern leopard explain differences in the occurrence of amphibian species
frogs, spotted frogs, and western pond turtles.72, 131 within Oregon and Washington.
In contrast to amphibians, reptile diversity in shrub- Because reptiles are ectothermic, thermal conditions
steppe habitat is relatively high (21 of 28 native species in play a key role in determining the occurrence, distribution,
Oregon and Washington). Lizards are the group of reptiles and numbers of reptiles that will be found in shrub-steppe
most associated with shrub-steppe (9 of 11 species in habitats. In general, reptile species richness decreases with
Oregon and Washington). No lizard species occurs increasing latitude and altitude.60 The number of reptile
exclusively in shrub-steppe, but 3 species (Mojave black- species decreases by over 40% between southern Oregon
collared lizard, long-nose leopard lizard, and desert and northern Washington. For example, Mojave black-
horned lizard) occur only in shrub-steppe, dwarf shrub- collared lizards and ground snakes do not occur in
steppe, and desert playa/salt scrub shrublands. Washington, and side-blotched lizards and striped
Ten of the 15 (67%) snake species in Oregon and whipsnakes do not occur north of central Washington.
Washington occur in shrub-steppe habitat. The ground Within a landscape, the distribution and abundance of
snake occurs only in shrub-steppe, dwarf shrub-steppe, reptiles will vary with topography. For example, snake
and desert playa/salt scrub shrublands. Striped dens are usually located on south facing slopes43, 61 (S.
whipsnakes occupy these three habitats plus juniper and Cooper Doering and C. R. Peterson, Idaho State University
mountain mahogany woodlands, whereas night snakes unpubl. data).
are found in the same habitats as striped whipsnakes plus For many species of reptiles, it is likely that the time
eastside canyon shrublands. Three species (racer, gopher available for embryo development at appropriate
snake, and western rattlesnake) occur in a wide variety of temperatures is the condition that limits distribution. For
habitats, including shrub-steppe. Three more species example, biophysical analyses indicate that adult and
(rubber boa, western terrestrial garter snake, and common juvenile desert iguanas could survive in shrub-steppe
garter snake) also occur in a wide variety of habitats, habitat in Washington, but that soil temperatures are too
including shrub-steppe, especially near water. low to allow successful incubation of eggs.109 Gravid
Although both species of freshwater turtles (painted rubber boas and western rattlesnakes have been observed
turtle and western pond turtle) occur more frequently in at den sites in the late fall of cool years in southeastern
other habitats, they use shrub-steppe if near permanent Idaho, indicating that summer temperatures were not
water (marshes, slow rivers and streams, ponds, or lakes sufficiently high to allow the embryos to develop (M. E.
with abundant aquatic vegetation). These turtles venture Dorcas, Davidson College, pers. comm., J. Lee, Idaho State
onto land to bask or to disperse, and they may lay their University pers. comm.). These field observations,
eggs in nests up to 150 and 800 m, respectively, away from combined with laboratory studies of the thermal
water.98, 134 dependence of embryo development14 and observations
Although species richness of amphibians and reptiles of the inability of embryos to survive simulated
is lower than that of birds and mammals in Oregon and hibernation (M. Dorcas, Davidson College pers. comm.),
Washington shrub-steppe, amphibians and reptiles can be suggest that the amount of time at suitable temperatures
very important ecologically. Because their long-term available for embryo development may play an important
conversion efficiencies are many times higher than those role in setting the distributional limits of snakes. Indeed,
of birds and mammals, they can contribute the evolution of viviparity in reptiles is generally viewed
disproportionately to biomass production and make large as an adaptation to cool conditions.111
amounts of energy available to other trophic levels.110 For Vegetation structure and floristics seem to explain less
example, Turner at al.140 found that the annual biomass of the variation in occurrence and distribution of reptiles
produced by side-blotched lizards in the Nevada desert than in birds and mammals. In developing GIS models
was equal to or greater than that of birds and mammals for predicting reptile distributions for the Idaho Gap
in desert and grassland habitats. Analysis project, Butterfield et al.11 found that factors other
than vegetation appear to limit the distribution of many
General Organizing Principles: reptiles. These factors may include temperature, moisture,
Reptiles and Amphibians and the special habitat features previously described.
The presence of water is an essential habitat feature for Nevertheless, vegetation does influence reptiles in several
the amphibians occurring in Oregon and Washington ways. Vegetation structure may influence reptiles directly;
shrub-steppe. All of these amphibians breed in lentic for example, sagebrush lizards in northcentral Oregon use
environments (e.g., marshes, pools, ponds, side-channels, habitats with tall shrubs (thermal cover and protection
or oxbows). Consequently, the presence of standing or from avian predators) and sparse ground cover (ease in
slow-moving water for at least the time required for eggs detecting and pursuing prey) (G. Green, Foster Wheeler
to hatch and larvae to complete metamorphosis is required Environmental Corporation, unpubl. data). Because most
for successful reproduction. However, because many of reptiles inhabiting shrub-steppe are carnivorous,
these species can live for considerable lengths of time (>10 differences in vegetation probably influence reptile
occurrence indirectly via the habitat affinities of their prey.
 CHAPTER 11: SHRUB-STEPPE 305

Management Issues in nesting waterfowl and marshland birds, as well as

amphibians and aquatic mammals. These areas also serve
Shrub-Steppe as migration stop-over sites for waterfowl. Wetlands with
In this section we consider some of the management issues a woodland component provide stop-over sites for
that have had (and likely will continue to have) passerines that historically used naturally occurring
considerable impact on shrub-steppe communities at a wooded riparian habitats, a resource that has been greatly
regional scale. There are certainly other threats to shrub- depleted.
steppe communities, particularly at the local scale (e.g., Farm fields enrolled in the Conservation Reserve
real estate development and inundation by water projects), Program (CRP) can have considerable value to shrub-
but we limit our discussion here to more widespread steppe birds. These fields are taken out of production for
issues. >10 years and planted to tame grasses such as crested
wheatgrass, providing nesting habitat for greater sage-
Conversion to Agriculture
grouse (M. Schroeder, WDFW pers. comm.) and
There is little doubt that the conversion of native plant
grasshopper sparrows107 on lands that offered few such
communities to agricultural uses has had, and continues
values when under cultivation. As sagebrush colonizes
to have, profound effects on shrub-steppe habitats in the
CRP fields via seeds from adjacent shrub-steppe, habitat
Columbia Basin. Beginning with the westward migration
value increases because of added structure and food for
of Euro-Americans in the mid-1800s that brought farmers
sage-dependent species. The value of CRP fields to shrub-
into the deserts of Oregon and Washington, and
steppe wildlife could be increased by planting them with
accelerated by the damming of the Columbia River that
native vegetation and extending the period of enrollment.
made large-scale irrigation possible, about 14.8 million
acres (6 million ha) of shrub-steppe have been converted Habitat Fragmentation
to wheat fields, row crops, and orchards.113 Agricultural The pattern of agricultural conversion within the shrub-
development has been most pronounced in Washington steppe of eastern Washington and northcentral Oregon
where >50% of the land originally in shrub-steppe has has resulted in a highly fragmented landscape (Figure 1).
been converted27 (Figure 1). This large-scale displacement Where once native grasslands and shrublands stretched
of one habitat type for another has substantially reduced unbroken for thousands of square miles, there exists now
the area available to native shrub-steppe wildlife. only fragments of native habitats in a matrix of agricultural
Moreover, the addition of new, human-related habitats fields. This breakup of formerly contiguous habitats can
(agricultural and rural development) has elevated the food have detrimental effects on species occurrence and
base for some predators (e.g., magpies and gulls) and population dynamics. Much of the research documenting
likely their populations as well, with unknown impacts fragmentation effects has examined avian communities
on shrub-steppe wildlife. The addition of cattle feedlots, in forested ecosystems, although some recent work has
pastures, and lawns to the landscape has enhanced the focused on grasslands and shrublands.6, 63, 147
suitability of the area for brown-headed cowbirds, a nest Some forest bird species are area sensitive and will not
parasite that lays its eggs in the nests of other birds and inhabit habitat patches below a minimum size. 34, 117
thereby depresses the host bird’s reproductive success. Extensive surveys in Washington suggest that sage
Agricultural conversion has not occurred randomly sparrows are most likely to occur in blocks of shrub-steppe
across the landscape, but instead has focused on the most >2,470 acres (1,000 ha) and that male sage sparrows found
arable, deep soil communities. This has resulted in a singing in small fragments are unlikely to maintain a
disproportionate loss of these communities and an territory or attract a mate (WDFW unpubl. data).
increase in the proportion of shallow soil shrub-steppe Numerous studies have documented greater rates of nest
habitats on the landscape (Figure 2). Some species of predation28, 152, 163 and nest parasitism9, 118 in fragmented
shrub-steppe wildlife, such as badgers, ground squirrels, landscapes. Elevated rates of nest predation and
and burrowing owls, depend on deep soil communities. parasitism may result from an increase in the number of
The pygmy rabbit, listed as endangered in Washington, is predators and brown-headed cowbirds in fragmented
found only in deep, loamy soil sagebrush stands. landscapes and an increase in habitat edge. In Washington,
Furthermore, some shrub-steppe passerines occur in 3 shrub-steppe birds (Brewer’s sparrow, lark sparrow, and
greater abundance in loamy soil communities than in other sage thrasher) showed evidence of lower nesting success
soil types. 147 Conversion of deep soil shrub-steppe in fragmented shrub-steppe compared to continuous,
communities to irrigated agriculture will likely continue unbroken tracts (WDFW unpubl. data). Cameras
in the foreseeable future, making this one of our most monitoring artificial nests baited with quail eggs revealed
endangered arid land communities. that black-billed magpies and common ravens likely were
Although loss of native plant communities should be responsible, at least partly, for increased predation on nests
avoided, some habitats associated with agricultural in fragmented landscapes (WDFW unpubl. data). As
development can have values for wildlife. Wetlands remnant habitat becomes smaller and more fragmented,
associated with agricultural development provide it is under greater influence of the surrounding
breeding and feeding areas for species not typically landscape157 and more susceptible to external influences,
associated with shrub-steppe. Wetlands created as part of be they predators, nest parasites, potential competitors,
numerous irrigation projects provide habitat for various or the wind-blown seeds of exotic weeds.
 306 Wildlife–Habitat Relationships in Oregon and Washington

Landcover
Water/wetlands
Shrub-steppe
Agriculture
Forest

County boundary
miles kilometers

Figure 1. Current landcover of four counties in eastern using multi-temporal analysis (scene dates: May 1993 and
Washington, illustrating the degree of fragmentation in this August 1994). Counties illustrated are (clockwise from the
formerly shrub-steppe-dominated landscape. Landcover bottom right) Adams, Grant, Douglas, and Lincoln.
classes were derived from Landsat Thematic Mapper data
CHAPTER 11: SHRUB-STEPPE 307

Figure 2. (a) Historical

distribution of deep soil and
shallow soil shrub-steppe
communities in Grant and
Lincoln Counties, Washington.
(b) Current distribution of deep
soil and shallow soil shrub-steppe
communities in Grant and
Lincoln Counties, illustrating
the extensive conversion of deep-
soil shrub-steppe (primarily to
agricultural fields) and minimal
conversion of shallow-soil
shrubsteppe.

kilometers miles

Soil/landcover classes
Deep soil, shrub-steppe
Shallow soil, shrub-steppe
Deep soil, converted
Shallow soil, converted
Woodland
Water
 308 Wildlife–Habitat Relationships in Oregon and Washington

Photograph 9. Shrub-steppe
communities of eastern Washington
and northcentral Oregon have been
fragmented by agricultural conversion.
Douglas County, Washington. (M.
Vander Haegen, WDFW).

Fragmentation of the shrub-steppe landscape very herbivores, especially bison.77 For whatever reason, herds
likely has disrupted the dynamics of dispersal and of native ungulates never reached the numbers in the
immigration that allows populations to persist over large shrub-steppe regions of Oregon and Washington that they
areas. Stochastic events may cause the extirpation of a did east of the Rocky Mountains. Consequently, the
species from one habitat patch, necessitating recruitment Agropyron, Poa, and Festuca dominated grasslands of
from nearby patches to reestablish a population. Highly Oregon and Washington may have been poorly adapted
fragmented landscapes have lower connectivity, meaning to withstand the grazing pressures of European livestock
that dispersing individuals must cross unfavorable lands introduced over the last 200-300 years.
(in this case agricultural fields or developed lands) to move The legacy of livestock grazing in the shrub-steppe
from one habitat patch to another. In theory, the smaller regions of Oregon and Washington began about 1700 when
the patch and the more distant other population sources, the Shoshone brought horses into southeastern Oregon
the lower the probability that recolonization will occur.75, from the Spanish missionaries at Santa Fe.46 By 1730, horses
164
Species with small home ranges and limited dispersal had reached the Columbia Basin, where the Nez Perce and
capabilities, such as many small mammals and reptiles, Cayuse built herds into the thousands by 1800.35 The
are most likely to be affected (see Box 2). For species that impact of these horses on the local grassland ecology is
normally occur at relatively low densities, such as the unrecorded. Cattle grazing as an industry did not begin
northern grasshopper mouse and pygmy rabbit, small east of the Cascades until the 1860s, but quickly expanded,
breeding assemblages could become genetically isolated reaching its zenith in the late 1870s. At about the same
and vulnerable to extirpation. 65, 66 Likewise, for time cattle began competing with hundreds of thousands
populations that are prone to dramatic cyclic fluctuations of sheep and the thousands of horses needed for cattle
in population size, such as lagomorphs, fragmentation raising, plus thousands more unattended Indian ponies
may increase the probabilities of local extinctions called “cayuses.” By 1885 the range was showing signs of
associated with decline phases in isolated populations. deterioration. Farming in the latter part of the century not
Although research has addressed the consequences of only fueled greater competition on less and less range,
genetic isolation and the probability of persistence for but it brought with it exotic seed contaminants such as
small populations in fragmented forest communities,71, 93 cheatgrass, Jim Hill mustard (Sysimbrium altissimum), and
little work has been done in shrub-steppe. Russian thistle (Salsola kali), that facilitated further
deterioration of the range. As a result of public outcry
Livestock Grazing about poor range conditions, the Federal government
With the rise of the Cascade Range in the early Pliocene, finally gained control of all unclaimed rangelands with
maritime influences east of the range diminished and the the Taylor Grazing Act in 1934.
Intermountain West became drier, with rainfall patterns Today, grazing management is dictated by the science
for the most part centered in the fall and winter. Grasses of range management defined by Stoddart et al.133 as
capable of estivating during the dry summer months, such optimizing the returns from rangelands in those
as Agropyron, Poa, and Festuca, dominated the landscape.23, combinations most desired by and suitable to society
77
In turn, the lack of moisture in the plants during the through the manipulation of range ecosystems.
summer months, coupled with poor distribution of Manipulation in the Intermountain West mostly has
drinking water, may have imposed severe constraints on involved reseeding of deteriorated rangelands with non-
the establishment of significant numbers of large native grasses (largely because seed sources for many
 CHAPTER 11: SHRUB-STEPPE 309

2: Spatial Scale

The scale at which different species use a access to large tracts of habitat to be successful.29, 84,
86, 169
landscape varies widely, as does our knowledge of Large mammals such as elk have remarkable
this component of wildlife ecology. Sage and dispersal abilities, however, and thus are less
sharp-tailed grouse, both resident species, may prone than smaller mammals to becoming
cover a considerable area over the course of a year genetically isolated by fragmentation unless the
as they fill their various needs for breeding, food, distance between patches becomes very large.
and cover. A female sharp-tailed grouse must visit Meso-sized and small mammals have smaller
a male on his lek in the spring, then locate suitable home ranges, and therefore patch scale has
nesting cover to lay and incubate a clutch, rear her different implications relative to large mammals.
young in suitable brood cover, perhaps move to Small mammals can, in theory, continue to exploit
another area as she tracks changing food resources small patches of shrub-steppe.64 However, because
in the fall, and then complete the year in riparian of small home range sizes and limited dispersal
cover where hardwood trees provide buds as a abilities, fragmentation of shrub-steppe habitat
source of winter food. Her annual home range may effectively isolate many populations of meso-
may cover 1,000 acres (405 ha) and include a sized and small mammals, with undetermined
multitude of vegetation communities.16 A female consequences.
greater sage-grouse may include >10 miles2 (26 Although reptiles generally respond to their
km2) in her breeding season home range alone.124 environments on a finer spatial scale than birds or
In contrast, a Brewer’s sparrow arrives in the most mammals, a wide range of variation exists in
shrub-steppe after having spent the winter in the size of areas used by different species.
South America, establishes a <2.5 acre (1 ha) Territorial, sit-and-wait predators, such as
territory in a big sagebrush stand160 and may side-blotched lizards, move on a scale of feet and
spend the entire breeding season nesting and have home ranges of <5,000 feet2 (500 m2).139 In
rearing young on this small territory. contrast, active, widely foraging species, such as
Fragmentation and other changes in the landscape the western whiptail lizard, have home ranges
likely will affect these two species quite several times larger.90 Some species of snakes that
differently. However, our knowledge of the habitat use communal overwintering sites may seasonally
needs of most shrub-steppe passerines is meager migrate several miles in one direction and have
and is focused primarily on the breeding territory. home ranges of hundreds of acres (e.g., western
There may be other needs, such as critical rattlesnakes).14 Species with smaller spatial
premigratory habitats or post-fledging habitats, requirements may be better able to persist for
that we are not aware of and that would expand short periods of time in fragmented habitats;
the landscape use of species like Brewer’s however, it is unclear whether their long-term
sparrows. persistence will be lesser or greater than those
Large mammals have relatively large home species that require large home ranges but are
ranges, and in shrub-steppe they may require better able to recolonize areas where populations
have gone locally extinct.

native grasses were simply not available), and establishing trampling of the soil. The friable soils of the shrub-steppe
seasonal grazing regimes designed to prevent further zone, especially in the drier areas, are held together by
deterioration of rangeland.51, 133 However, these present- layers of cryptogamic mosses and lichens. This
day systems are still designed to maximize livestock cryptogamic crust can prevent establishment of annual
production, often to the detriment of shrub-steppe weeds and provide a moisture cap that reduces soil
wildlife. evaporation. However, the trampling action of livestock,
Do present day grazing practices benefit or adversely especially horses and cattle, can degrade these layers and
affect shrub-steppe wildlife? The answer is mixed. Some provide seed beds for cheatgrass and weedy forbs. Dense
wildlife species, such as long-billed curlews 104 and stands of cheatgrass not only outcompete native
burrowing owls38 may actually benefit from reduced bunchgrasses (especially for moisture in the early stages
vegetational structure (grazed perennials or low-statured of growth), but are also susceptible to hot wildfires that
annuals), and Great Basin pocket mice will attain high can virtually eliminate sagebrush. 22 Consequently,
population numbers in pure cheatgrass stands. 127 whereas the proximate effect of livestock grazing on
However, ground-nesting birds and small mammals that wildlife may be the removal of grass and forb biomass
require protective cover from vegetation may not benefit, important as forage and cover to many wildlife species,
especially if they become more susceptible to predation. the ultimate effect may be perpetuation of weedy annuals
The greatest impact from grazing, however, is probably that out-compete native plants that local wildlife have
the perpetuation of the weed legacy from livestock adapted to use.
 310 Wildlife–Habitat Relationships in Oregon and Washington

Photograph 10. Grazing by livestock

has profoundly influenced the
vegetation in many shrub-steppe
communities. Grand Coulee, Douglas
County, Washington. (M. Vander
Haegen, WDFW).

The Fire/Cheatgrass Cycle soil or cryptograms between plants. This discontinuous

Another human-caused agent of change that threatens to fuel layer does not carry a fire well, and wildfires in these
degrade shrub-steppe habitats, related in part to communities typically burn patches, creating a mosaic of
agricultural practices and to livestock grazing in particular, burned and unburned areas.154 Cheatgrass, in contrast,
is the conversion of extensive areas to simplified annual forms continuous stands that desiccate early in the season
grasslands. Cheatgrass is an exotic annual grass that was and carry fires well154 (Photograph 11). As a result, burns
introduced to the Intermountain West sometime in the late in cheatgrass are larger and more frequent. As cheatgrass
1800s, probably as an agricultural pest.76 Overgrazing of biomass increases, the ability of the community to carry
native bunchgrass communities by livestock led to fire also increases, resulting in a positive-feedback loop
deterioration of the range and opened the door for the where fire promotes cheatgrass dominance, leading to
widespread invasion of this exotic annual grass.76 Each of more frequent fires.21 On Idaho’s Snake River plain,
the arid and semi-arid grassland and shrubland habitat sagebrush communities evolved with a fire-return interval
types in eastern Oregon and Washington is susceptible to of 35-100 years; following invasion by cheatgrass, fire-
cheatgrass invasion.113 Characteristics of cheatgrass’s life return intervals in some shrub-steppe communities are
history and physical structure, coupled with a native flora now as low as 3-5 years.154
ill-equipped to compete with this new invader, have Increased fire frequency in steppe and shrub-steppe
allowed it to change the composition of shrub-steppe vegetation leads to a vegetation community with lower
communities and even alter ecosystem processes.21 species richness.154 On frequently burned areas of Idaho’s
Cheatgrass is a winter annual that goes to seed early in Snake River plain, almost all of the vegetation was
the year and generally desiccates in spring or early introduced annuals—primarily cheatgrass.154 Vegetation
summer. A native of the Mediterranean region, it is life forms react differently and predictably to the fire/
adapted to the climate of eastern Oregon and Washington cheatgrass cycle; essentially, annuals increase dramatically
and, unlike the native bunch grasses, can thrive under and all other plants decrease. 154 Wildfire kills big
heavy livestock grazing. Cheatgrass rapidly colonizes bare sagebrush, and because most species of sagebrush do not
soil and moves readily into disturbed sites. Intact resprout from root crowns after fires, regeneration
communities with native bunchgrasses and a healthy depends on the existing seed bank. Big sagebrush seeds
cryptogamic crust can keep the invader in check; however, are short-lived, and if fire returns before the new seedlings
heavy grazing of native grasses and mechanical reach reproductive age (4-6 years) the species can be
breakdown of the cryptogamic crust through trampling eliminated from the community.154 Even rabbitbrush, a
can provide a window for cheatgrass to spread and exert common successional shrub in sage communities that
dominance. Wildfire also can provide a window of readily sprouts after fire, can be lost from the community
opportunity by killing shrubs and making valuable soil with fire-return intervals of 2-4 years.154 As cheatgrass-
moisture available to cheatgrass for germination and fueled fires lead to more extensive burns, seed sources
growth. Cheatgrass recovers quickly following wildfires from adjacent, unburned areas become more distant and
and can out-compete native grasses.91 the patch dynamics that likely promoted revegetation in
The most extreme changes that cheatgrass has caused these communities historically no longer functions. This
in the West have resulted from repeated wildfires and the change in the fire frequency leads to a change in the
response of cheatgrass to post-fire conditions.21 Native trajectory of plant succession and represents an alteration
bunchgrasses generally grow sparsely, with forbs and bare of ecosystem processes.21
 CHAPTER 11: SHRUB-STEPPE 311

Photograph 11. Cheatgrass can form

continuous stands that burn frequently,
preventing reestablishment of the former
vegetation community. Hanford Site,
Benton County, Washington. Photo: M
Vander Haegen, WDFW.

Research Needs do we restore native vegetation on former agricultural sites

A host of wildlife species have received little attention in and maximize the chance of regaining the full range of
ecological studies, yet have great potential to be affected ecological function? How do we restore native vegetation
by changes in shrub-steppe landscapes. This list would and ecological function to highly degraded rangelands?
include many of the reptiles, amphibians, and small Agricultural lands enrolled in CRP are a significant part
mammals. We need studies examining demography and of the landscape in eastern Washington. Research on the
habitat affinities of the Great Basin spadefoot toad, long- value of CRP to wildlife will help us assess its place in the
nosed leopard lizard, night snake, sagebrush vole, and shrub-steppe ecosystem and guide future enrollments to
the full complex of ground squirrels, among others. We benefit native wildlife.
need a better understanding of the relationship between
fossorial mammals and the species that depend on them
Applying the Data Matrixes
for burrows and as prey. Fourteen species of bats are to a Management Example:
known to use shrub-steppe, yet we know little about their
population trends, population dynamics, and habitat Changes in the Wildlife Community
needs, or how disturbance by humans affects breeding or Following Wildfire and Conversion to
roosting activities. We need studies examining the effects
of habitat fragmentation on the distribution and Annual Grasslands
demography of shrub-steppe wildlife. We need to develop
and implement surveys for sage sparrows, grasshopper
Loss of Shrubs
sparrows, burrowing owls, and other birds that are not Little published work has examined the effects of repeated
covered sufficiently by the Breeding Bird Survey, as well wildfires and conversion to annual grasslands on shrub-
as for reptiles and mammals that are not easily observed. steppe wildlife communities. Loss of the shrub layer
We know little about how the condition of extant shrub- through repeated fires eliminates habitat for shrub nesting
steppe varies across the landscape. Mapping shrub-steppe birds, including some key shrub-steppe obligates (sage
communities across a gradient of conditions, from pristine sparrow, Brewer’s sparrow, and sage thrasher). Although
to highly degraded, would provide a more realistic fidelity to their breeding site will sometimes bring adults
assessment of the current status of the resource and would back in the year following a fire108 or other catastrophic
be invaluable for future modeling of wildlife distribution. loss of the shrub layer,159, 161 without suitable nesting shrubs
Livestock grazing has altered the vegetaion in many shrub- such returns are likely to be a short-term phenomenon.
steppe communities, yet we lack information on how these Loss of shrubs would be detrimental to some lizard
changes affect most wildlife species. We need research on species, reducing the availability of shaded sites needed
the role of microbiotic rusts in maintaining ecosystem for thermoregulation and cover used to avoid predators.
function and as an indicator of ecosystem integrity. Are When fire removes shrubs from a community, ungulates
microbiotic crusts an important component for some that browse on sagebrush, bitterbrush, and other shrubs
species? lose valuable winter habitat. Updike et al.142 documented
Expanding habitat linkages in rfagmented landscapes a decline in big game winter range following fire,
and increasing the amount of deep soil shrub-steppe for cheatgrass invasion, and suppression of shrub
species such as pygmy rabbits and ground squirrels will regeneration by cheatgrass. Species that depend on
require restoration of converted agricultural lands. How sagebrush for forage during all or part of the year, such as
 312 Wildlife–Habitat Relationships in Oregon and Washington

Table 2. Numbers of species associated with shrub-steppe habitat type, modified grassland structural
condition, and shrubs as a key habitat element, in Oregon and Washington.

Species Shrub-steppe Modified Grasslands Shrubs as a key

group Generally associated Closely associated Generally associated Closely associated element

Birds 44 22 31 2 22
Mammals 26 27 34 0 12
Reptiles 20 0 7 0 6
Amphibians 9 0 10 0 1
Totals 103 49 82 2 41

greater sage-grouse and pygmy rabbits, likely will be grasslands can be derived from the matrixes. Sites
excluded as sagebrush communities degrade to annual dominated by cheatgrass are classified as modified
grasslands. grasslands in the Structural Condition Matrix. A query of
The Habitat Element Matrix (contained on the CD- this matrix revealed 82 species “generally associated” with
ROM) can be used to determine which species depend on annual grasslands and only 2 species classified as “closely
shrubs as an important component of shrub-steppe habitat associated.” A query of the Wildlife-Habitat Matrix for
and to predict which species would be lost from the species in shrub-steppe found 103 species “generally
community as fire removes the shrub layer. A query of associated” and 49 species “closely associated” with this
the matrix revealed 22 birds, 12 mammals, 6 reptiles, and habitat type (Table 2). From the results of these queries
1 amphibian that are associated with shrub-steppe habitat we can conclude that the conversion of shrub-steppe
and require shrubs for some life function (Table 2). communities to annual grasslands through the fire/
cheatgrass cycle can cause dramatic changes in the wildlife
Conversion of the Herb Layer to Cheatgrass community. The change may be particularly severe in the
Changes in structure and composition of the herb layer breeding bird community, where 20 species closely
that follow cheatgrass invasion also affect the wildlife associated with shrub-steppe (several of them obligates)
community. Conversion of native forbs and bunchgrasses are predicted to be excluded. There are limitations to such
to exotic annuals results in a less stable food base for small an analysis, and one must give careful thought to other
herbivores like Townsend’s ground squirrel, increasing the factors that may influence species occurrence at a
amplitude of their population fluctuations and the particular site.
potential for localized extinctions.167 Long-billed curlews
seem to prefer cheatgrass dominated sites for nesting in Acknowledgments
southcentral Washington2 and northcentral Oregon,104 We thank the following individuals for reviewing previous
whereas some other ground-nesting birds have been drafts of the manuscript: S. Burton, J. Cossell, Jr., J. Lee,
found to occur at abnormally low densities in cheatgrass.7 W. Leonard, M. Schroeder, and P. Sherman. We also thank
Birds that prefer open ground for foraging (e.g., sage B. J. Verts for providing information on Merriam’s ground
sparrow and loggerhead shrike) avoid sites with dense squirrel and J. Jacobson for preparing Figures 1 and 2.
cheatgrass.27, 73 Conversion to cheatgrass probably would
decrease availability of prey for lizards and make it more Literature Cited
difficult for lizards to move about. Some snakes also would 1. Abaturov, B. D. 1972. The role of burrowing animals in the transport of
be negatively affected by such a conversion; night snakes mineral substances in the soil. Pedobiologia 12:261-266.
2. Allen, J. N. 1980. The ecology and behavior of the long-billed curlew in
and striped whipsnakes prey largely on lizards and thus
southeastern Washington. Wildlife Monographs 73.
would likely be negatively affected by the loss of lizards 3. Bailey,V. 1936. The mammals and life zones of Oregon. U. S.
due to habitat conversion. Preliminary results from a Department of Agriculture, Bureau of Biological survey, North
comparative trapping study conducted in the Snake River American Fauna 55:1-416.
Birds of Prey Area in 1978-197926 and 1997-1998 (J. Cossel, 4. Betts, B. J. 1990. Geographic distribution and habitat preferences of
Jr. and C. Peterson, Idaho State University unpubl. data) Washington ground Squirrels (Spermophilus washingtoni). Northwestern
are generally consistent with these predictions. Located Naturalist 71:27-37.
5. ———.1999. Current status of Washington ground squirrels in
in southwestern Idaho, much of this area has been Oregon and Washington. Northwestern Naturalist 80:35-38.
converted from natural shrubland to non-native grassland. 6. Bolger, D. T., T. A. Scott, and J. T. Rotenberry. 1997. Breeding bird
Although there is considerable year-to-year variation in abundance in an urbanizing landscape in coastal southern California.
the occurrence and trapping rates of reptiles, side-blotched Conservation Biology 11:406-421.
lizards and night snakes seem to have declined whereas 7. Brandt, C. A., and W. H. Rickard. 1994. Alien taxa in the North
more generalist species like racers have increased. American shrub-steppe four decades after cessation of livestock
grazing and cultivation agriculture. Biological Conservation 68:95-105.
Changes in the wildlife community that might be
8. Braun, C. E., et al. 1976. Conservation committee report on effects of
expected following conversion of shrub-steppe to annual alteration of sagebrush communities on the associated avifauna. Wilson
Bulletin 88:165-171.
 CHAPTER 11: SHRUB-STEPPE 313

9. Brittingham, M. C., and S. A. Temple. 1983. Have cowbirds caused forest technology in the management of impacted wildlife. Thorne Ecological
songbirds to decline? Bioscience 33:31-35. Institute, Boulder, CO.
10. Bryant, L. D., and C. Maser. 1982. Classification and distribution. Pages 33. Freeman, S., D. F. Gori, and S. Rohwer. 1990. Red-winged blackbirds
1-60 in J. W. Thomas and D. E. Toweill, editors. Elk of North America: and brown-headed cowbirds—some aspects of a host-parasite
ecology and management. Stackpole Books, Harrisburg, PA. relationship. Condor 92:336-340.
11. Butterfield, B. R., B. Csuti, and J. M. Scott. 1994. Modeling vertebrate 34. Freemark, K. E., and H. G. Merriam. 1986. Importance of area and
distributions for Gap Analysis. Pages 53-68 in R. I Miller, editor. Mapping habitat heterogeneity to bird assemblages in temperate forest
the diversity of nature. Chapman and Hall, London, England. fragments. Biological Conservation 36:115-141.
12. Byers, J. A. 1997. American pronghorn: social adaptations and the 35. Galbraith, W. A., and E. W. Anderson. 1971. Grazing history of the
ghosts of predators past. University of Chicago Press, Chicago, IL. Northwest. Journal of Range Management 24:6-12.
13. Carson, R. G., and J. M. Peek. 1987. Mule deer habitat selection 36. Gano, K. A. 1979. Analysis of small mammal populations inhabiting the
patterns in northcentral Washington. Journal of Wildlife Management environs of a low-level radioactive waste pond. Report PNL-2479,
51:46-51. Battelle, Pacific Northwest Laboratory, Richland, WA.
14. Cobb,V. A. 1994. The thermal ecology of pregnancy in free-ranging 37. Gavin, T. A., P. W. Sherman, E.Yensen, and B. May. 1999. Population
Great Basin rattlesnakes (Crotalus viridis lutosus). Dissertation, Idaho genetic structure of the northern Idaho ground squirrel. Journal of
State University, Pocatello, ID. Mammalogy 80:156-168.
15. Collins, W.B., and P. J. Urness. 1983. Feeding behavior and habitat 38. Green, G. A., and R. G. Anthony. 1989. Nesting success and habitat
selection of mule deer and elk on northern Utah summer range. relationships of burrowing owls in the Columbia Basin, Oregon.
Journal of Wildlife Management 47:646-663. Condor 91:347-354.
16. Connelly, J. W., M. W. Gratson, and K.P. Reese. 1998. Sharp-tailed 39. ———, and M. L. Morrison. 1983. Nest-site characteristics of
grouse (Tympanuchus phasianellus). In A. Poole and F. Gill, editors,The sympatric ferruginous and Swainson’s hawks. Murrelet 64:20-22.
Birds of North America, No. 354. The Birds of North America, Inc., 40. ———, R. E. Fitzner, R. G. Anthony, and L. E. Rogers. 1993.
Philadelphia, PA. Comparative diets of burrowing owls in Oregon and Washington.
17. Costa, G. 1995. Behavioural adaptations of desert animals. Springer- Northwest Science 67:88-93.
Verlag, New York, NY. 41. Green, J. S., and J.T. Flinders. 1980. Brachylagus idahoensis. Mammalian
18. Cox, C. B., and P. D. Moore. 1993. Biogeography: an ecological and Species 125:1-4, American Society of Mammalogists.
evolutionary approach. Blackwell Science, Oxford, England. 42. Greene, R. A., and C. Reynard. 1932. The influence of two burrowing
19. Csuti, B., A. J. Kimerling, T. A. O’Neil, M. M. O’Shaughnessy, E. P. Gaines, rodents, Dipodomysspectabilis spectabilis (kangaroo rat) and Neotoma
and M. M. P. Huso. 1997. Atlas of Oregon wildlife. Oregon State albigula albigula (pack rat) on desert soils in Arizona. Ecology 13:73-80.
University Press, Corvallis, OR. 43. Gregory, P. T. 1982. Reptilian hibernation. Pages 53-154 in C. Gans and
20. Currier, M. J. P. 1983. Felis concolor. Mammalian Species 200:1-7, F. H. Pough, editors. Biology of the Reptilia,Volume 13, Physiology D,
American Society of Mammalogists. Physiological Ecology. Academic Press Inc., New York, NY.
21. D’Antonio, C. M., and P. M. Vitousek. 1992. Biological invasions by 44. Gross, J. E., L. C. Stoddart, and F. H. Wagner. 1974. Demographic
exotic grasses, the grass/fire cycle, and global change. Annual Review of analysis of a northern Utah jackrabbit population. Wildlife Monographs
Ecology and Systematics 23:63-87. 40.
22. Daubenmire, R. 1970. Steppe vegetation of Washington. Washington 45. Hafner, D. J., E. Yensen, and G. L. Kirkland, Jr. 1998. North American
Agricultural Experiment Station Technical Bulletin 62. Washington State rodents. Status survey and conservation action plans. IUCN, Gland,
University , Pullman, WA. Switzerland.
23.———. 1975. Floristic plant geography of eastern Washington and 46. Haines, F. 1938. The northward spread of horses among the Plains
northern Idaho. Journal of Biogeography 2:1-18. Indians. American Anthropology 40:429-436.
24. Davis, W. B. 1939. The recent mammals of Idaho. Caxton Printers, Ltd., 47. Hall, E. R. 1981. Mammals of North America, 2nd edition. John Wiley &
Caldwell, ID. Sons, New York, NY.
25. Demment, M. W., and P. J.Van Soest. 1985. A nutritional explanation for 48. Hanley, T. A. 1980. Nutritional constraints of food and habitat
body-size patterns of ruminant and nonruminant herbivores. American selection by sympatric ungulates. Dissertation, University of
Naturalist 125:641-672. Washington, Seattle,WA.
26. Diller, L. V., and D. R. Johnson. 1982. Ecology of reptiles in the Snake 49. Haug, E. A., B. A. Millsap, and M. S. Martell. 1993. Burrowing owl
River Birds of Prey Area. U. S. Bureau of Land Management Snake River (Speotyto cunicularia). In A. Poole and F. Gill, editors,The Birds of North
Birds of Prey Research Project, Boise, ID. America, No. 61. The Birds of North America, Inc., Philadelphia, PA.
27. Dobler, F. C., J. Eby, C. Perry, S. Richardson, and M. Vander Haegen. 50. Hedlund, J. D., and L. E. Rogers. 1976. Characterization of small
1996. Status of Washington’s shrub-steppe ecosystem: extent, mammal populations inhabiting the B-C Crib environs. Report BNWL-
ownership, and wildlife/vegetation relationships. Research Report. 2181, Battelle Pacific Northwest Laboratory, Richland, WA.
Washington Department of Fish and Wildlife, Olympia, WA. 51. Heitschmidt, R. K., and J. W. Stuth. 1991. Grazing management, an
28. Donovan, T. M., F. R. Thompson III, J. Faaborg, and J. R. Probst. 1995. ecological approach. Timber Press, Portland, OR.
Reproductive success of migratory birds in habitat sources and sinks. 52. Hoffmann, R. S., C. G. Anderson, R. W. Thorington, Jr., and L. R. Heany.
Conservation Biology 9:1380-1395. 1993. Family Sciuridae. Pages 419-465 in D.E. Wilson and D.M. Reeder,
29. Eberhardt, L. E., E. E. Hanson, and L. L. Cadwell. 1982. Analysis of editors. Mammal species of the world: a taxonomic and geographic
radionuclide concentrations and movement patterns of Hanford Site reference. Smithsonian Institution Press, Washington, D.C.
mule deer. Report BNWL-4420, Battelle, Pacific Northwest Laboratory, 53. Hole, F. D. 1981. Effects of animals on soil. Geoderma 25:75-112.
Richland, WA. 54. Holmes, A. L., and G. R. Geupel. 1998. Avian population studies at
30. ———, S. M. McCorquodale, and G. A. Sargeant. 1989. Elk and deer naval weapons systems training facility Boardman, Oregon. Final report.
studies related to the Basalt Waste Isolation Project. Report PNL-6798, Point Reyes Bird Observatory, Stinson Beach, CA..
Battelle, Pacific Northwest Laboratory, Richland, WA. 55. Howell, A. H. 1938. Revision of North American ground squirrels, with
31. Fitzner, R. E., D. Berry, L. L. Boyd, and C. A. Rieck. 1977. Nesting of a classification of Sciuridae. North American Fauna 56:1-256.
ferruginous hawks (Buteo regalis) in Washington, 1974-75. Condor 56. Huey, R. B., C. R. Peterson, S. J. Arnold, and W. P. Porter. 1989. Hot
79:245-249. rocks and not-so-hot rocks: retreat-site selection by garter snakes and
32.———, and R. L. Newell. 1989. Ferruginous hawk nesting on the U.S. its thermal consequences. Ecology 70:931-944.
DOE Hanford Site: case history of a recent invasion caused by 57. Inouye, R. S., N. J. Huntly, D. Tilman, and J. R. Tester. 1987. Pocket
transmission lines. Pages 125-132 in Proceedings IV: issues and gophers (Geomys bursarius), vegetation, and soil nitrogen along a
successional sere in east central Minnesota. Oecologia 72:178-184.
 314 Wildlife–Habitat Relationships in Oregon and Washington

58. Jarman, P. J. 1974. The social organization of antelope in relation to 84. ———. 1991. Energetic considerations and habitat quality for elk in
their ecology. Behaviour 48:215-267. arid grasslands and coniferous forests. Journal of Wildlife Management
59. Johnson, K. H., R. A. Olson, and T. D. Whitson. 1996. Composition and 55:237-242.
diversity of plant and small mammal communities in tebuthiuron- 85. ———. 1993. Winter foraging behavior of elk in the shrub-steppe of
treated big sagebrush (Artemisia tridentata).Weed Technology 10:404- Washington. Journal of Wildlife Management 57:881-890.
416. 86. ———, and L. E. Eberhardt. 1993. The ecology of elk in an arid
60. Kiester, A. R. 1971. Species density of North American amphibians and environment: an overview of the Hanford elk project. Pages 56-63 in R.
reptiles. Systematic Zoology 20:127-137. L. Callas, D. B. Koch, and E. R. Loft, editors. Proceedings of the 1990
61. Klauber, L. M. 1956. Rattlesnakes. Volume 1. University of California Western States and Provinces Elk Workshop. California Department of
Press, Berkeley, CA. Fish and Game, Sacramento, CA.
62. Knick, S. T. 1990. Ecology of bobcats relative to exploitation and a 87. ———, ———, and G. A. Sargeant. 1989. Antler characteristics in a
prey decline in southeastern Idaho. Wildlife Monographs 108. colonizing elk population. Journal of Wildlife Management 53:618-621.
63. ———, and J. T. Rotenberry. 1995. Landscape characteristics of 88. ———, L. L. Eberhardt, and L. E. Eberhardt. 1988. Dynamics of a
fragmented shrub-steppe habitats and breeding passerine birds. colonizing elk population. Journal of Wildlife Management 52:309-312.
Conservation Biology 9:1059-1071. 89. ———, K. J. Raedeke, and R. D. Taber. 1986. Elk habitat use patterns in
64. ———, and D. L. Dyer. 1997. Distribution of black-tailed jackrabbit the shrub-steppe of Washington. Journal of Wildlife Management
habitat determined by GIS in southwestern Idaho. Journal of Wildlife 50:664-669.
Management 61:75-85. 90. McCoy, C. J. 1965. Life history and ecology of the Cnemidophorus tigris
65. Lacy, R. C. 1997. Importance of genetic variation to the viability of septentrionalis. Dissertation, University of Colorado, Boulder, CO.
mammalian populations. Journal of Mammalogy 78:323-335. 91. Melgoza, G., R. S. Nowak, and R. J. Tausch. 1990. Soil water exploitation
66. Lande, R. 1988. Genetics and demography in biological conservation. after fire: competition between Bromus tectorum (cheatgrass) and two
Science 241:1455-1460. native species. Oecologia 83:7-13.
67. Laundré, J. W. 1993. Effects of small mammal burrows on water 92. Messick, J. P., and M. G. Hornocker. 1981. Ecology of the badger in
infiltration in a cool desert environment. Oecologia 94:43-48. southwestern Idaho. Wildlife Monographs 76.
68. ———. 1998. Effect of ground squirrel burrows on plant productivity 93. Mills, L. S. 1996. Fragmentation of a natural area: dynamics of isolation
in a cool desert environment. Journal of Range Management 51:638- for small mammals on forest remnants. Pages 199-219 in R. G. Wright,
643. editor. National parks and protected areas: their role in environmental
69. Le Houerou, H. N., R. L. Bingham, and W. Skerbek. 1988. Relationship protection. Blackwell Science, Cambridge, MA.
between the variability of primary production and the variability of 94. Moen, A. N. 1973. Wildlife ecology: an analytical approach. W. H.
annual precipitation in world arid lands. Journal of Arid Environments Freeman, San Francisco, CA.
15:1-18. 95. Murie, O. J. 1951. The elk of North America. Stackpole Books,
70. Leckenby, D. A., D. P. Sheehy, C. H. Nellis, R. J. Scherzinger, I. D. Luman, Harrisburg, PA.
W. Elmore, J. C. Lemos, L. Doughty, and C. E. Trainer. 1982. Wildlife 96. Nadler, C. H. 1968.The chromosomes of Spermophilus townsendi
habitats in managed rangelands of the Great Basin of southeastern (Rodentia: Sciuridae) and report of a new subspecies. Cytogenetics
Oregon: mule deer. General Technical Report PNW-139, Pacific 7:144-157.
Northwest Forest and Range Experiment Station, Portland, OR. 97. Nowak, R. M. 1991. Walker’s mammals of the world. Fifth edition.
71. Lehmkuhl, J. F., S. D. West, C. L. Chambers, W. C. McComb, D. A. Johns Hopkins University Press, Baltimore, MD.
Manuwal, K. B. Aubry, J. L. Erickson, R. A. Gitznen, and M. Leu. 1999. An 98. Nussbaum, R. A., E. D. Brodie, and R. M. Storm. 1983. Amphibians and
experiment for assessing vertebrate response to varying levels and reptiles of the Pacific Northwest. University of Idaho Press, Moscow,
patterns of green-tree retention. Northwest Science 73:45- 63. ID.
72. Leonard W. P., H.A. Brown, L. L. C. Jones, K. R. McAllister, and R. M. 99. Olsen, B. 1976. Status report: Columbian sharp-tailed grouse. Oregon
Storm. 1993. Amphibians of Washington and Oregon. Seattle Audubon Wildlife 34:10.
Society, Seattle,WA. 100. O’Neil, T. A. and D. H. Johnson. 2001. Oregon and Washington wildlife
73. Leu, M. 1995. The feeding ecology and the selection of nest shrubs species and their habitats. Pages 1-21 In: D.H. Johnson and T.A. O’Neil,
and fledgling roost sites by loggerhead shrikes (Lanius ludovicianus) in managing directors. Wildlife-habitat relationships in Oregon and
the shrub-steppe habitat. Thesis, University of Washington, Seattle, WA. Washington. Oregon State University Press, Corvallis, OR.
74. Lowther, P. E. 1993. Brown-headed cowbird (Molothrus ater). In A. 101. Oregon Department of Fish and Wildlife. 1994. Oregon Species
Poole and F. Gill, editors, The Birds of North America, No. 47. The Birds Information System. Wildlife Research, Corvallis, OR.
of North America, Inc., Philadelphia, PA. 102. Orians, G. H., E. Roskaft, and L. D. Beletsky. 1989. Do brown-headed
75. MacArthur, R. H., and E. O. Wilson. 1967. The theory of island cowbirds lay their eggs at random in the nests of red-winged
biogeography. Princeton University Press. Princeton, NJ. blackbirds? Wilson Bulletin 101:599-605.
76. Mack, R. N. 1981. Invasion of Bromus tectorum L. into western North 103. Osborne, D. 1953. Archaeological occurrence of pronghorn
America: an ecological chronicle. Agro-Ecosystems 7:145-165. antelope, bison, and horse in the Columbia Plateau. Scientific Monthly
77. ———, and J. N. Thompson. 1982. Evolution in steppe with few large, 77:914-917.
hoofed mammals. American Naturalist 119:757-773. 104. Pampush, G. J., and R. G. Anthony. 1993. Nest success, habitat
78. Maclean, G. L. 1996. Ecophysiology of desert birds. Springer-Verlag. utilization and nest-site selection of long-billed curlews in the
New York, NY. Columbia Basin, Oregon. Condor 95:957-967.
79. Marr, N. V., C. A. Brandt, R. E. Fitzner, and L. D. Poole. 1988. Habitat 105. Parker, K. L., and C. T. Robbins. 1984. Thermoregulation in mule deer
associations of vertebrate prey within the Controlled Area study zone. and elk. Canadian Journal of Zoology 62:1409-1422.
Report PNL-6495, Battelle Pacific Northwest Laboratory, Richland, WA. 106. Parmenter, R. R., and J. A. MacMahon. 1983. Factors determining the
80. Marshall, D. B. 1992. Sensitive vertebrates of Oregon. Oregon abundance and distribution of rodents of a shrub-steppe ecosystem:
Department of Fish and Wildlife, Portland, OR. the role of shrubs. Oecologia 59:145-156.
81. Mason, E. 1952. Food habits and measurements of Hart Mountain 107. Patterson, M. P., and L. B. Best. 1996. Bird abundance and nest
antelope. Journal of Wildlife Management 16:387-389. success in Iowa CRP fields: the importance of vegetation structure and
82. McCorquodale, S. M. 1985.The ecology of a shrub-steppe elk (Cervus composition. American Midland Naturalist 135:153-167.
elaphus) population. Thesis, University of Washington, Seattle,WA. 108. Petersen, K. L., and L. B. Best. 1987. Effects of prescribed burning on
83. ———. 1987. Fall-winter habitat selection by elk in the shrub-steppe nongame birds in a sagebrush community. Wildlife Society Bulletin
of Washington. Northwest Science 61:171-173. 15:317-329.
109. Porter, W. P,. and C. R. Tracy. 1983. Biophysical analyses of energetics,
 CHAPTER 11: SHRUB-STEPPE 315

time-space utilization, and distributional limits. Pages 55-83 in R. B. ecological intruder in southern Idaho. Ecology 30:58-74.
Huey, E. R. Pianaka, and T. W. Schoener, editors. Lizard ecology. Harvard 133. Stoddart, L.A., A.D. Smith, and T.W. Box. 1975. Range Management.
University Press, Cambridge, MA. McGraw-Hill Book Company, New York, NY.
110. Pough, F. A. 1980.The advantages of ectothermy for tetrapods. 134. Storm, R. M., W. P. Leonard, H. A. Brown, R. B. Bury, D. M. Darda, L.V.
American Naturalist 115: 92-112. Diller, and C.R. Peterson. 1995. Reptiles of Washington and Oregon.
111. Pough, F. H., J. B. Heiser, and W. N. McFarland. 1996.Vertebrate life. Seattle Audubon Society, Seattle, WA.
Fourth edition. Prentice Hall, Upper Saddle River, NJ. 135. Sundstrom, C., W. G. Hepworth, and K. L. Diem. 1973. Abundance,
112. Price, L. W. 1971. Geomorphic effect of the arctic ground squirrel in distribution, and food habits of the pronghorn. Bulletin 10, Wyoming
an alpine environment. Geografiska Annaler 53A:100-106. Game and Fish Commission, Cheyenne, WY.
113. Quigley, T. M., and S. J. Arbelbide. 1997. An assessment of ecosystem 136. Taylor, W.P. 1935. Some animal relations to soil. Ecology 16:127-136.
components in the interior Columbia Basin and portions of the 137. Thorp, J. 1949. Effects of certain animals that live in soils. Scientific
Klamath and Great Basins. General Technical Report PNW-GTR-405. Monthly 68:180-191.
U.S. Forest Service, Pacific Northwest Research Station, Portland, OR. 138. Thorson, T., and A. Svihla. 1943. Correlation of the habits of
114. Rickard, W. H., J. D. Hedlund, and R. E. Fitzner. 1977. Elk in the shrub- amphibians with their ability to survive the loss of body water. Ecology
steppe of Washington: an authentic record. Science 196:1009-1010. 24:374-381.
115. ———, and B. E.Vaughan. 1988. Characteristics of contrasting 139. Turner, F. B., R. I. Jennrich, and J. D. Weintraub. 1969. Home ranges and
shrub-steppe plant communities. Pages 109-179 in W. H. Rickard, L. E. body size of lizards. Ecology 50:1076-1081.
Rogers, B. E. Vaughan, and S. F. Liebetrau, editors. Shrub-steppe: balance 140. ———, P. A. Medica, and B. W. Kowalewsky. 1976. Energy utilization
and change in a semi-arid terrestrial ecosystem. Elsevier, Amsterdam, by a lizard (Uta stansburiana). U.S. International Biological Programme
The Netherlands. Monograph No. 1. Utah State University Press, Logan, UT.
116. Ricklefs, R. E. 1983. Ecology (second edition). Chiron Press, New 141. Turner, G. T., R. M. Hansen,V. H. Reid, H. P. Tietjen, and A. L. Ward.
York, NY. 1973. Pocket gophers and Colorado mountain rangeland. Colorado
117. Robbins, C. S., D. K. Dawson, and B. A. Dowell. 1989. Habitat area State University Experiment Station Bulletin, 554S. Fort Collins, CO.
requirements of breeding forest birds of the Middle Atlantic states. 142. Updike, D. R., E. R. Loft, and F. A. Hall. 1990. Wildfires on big
Wildlife Monographs 103. sagebrush/antelope bitterbrush range in northeastern California:
118. Robinson, S. K., F. R. Thompson III, T. M. Donovan, D. R. Whitehead, implications for deer populations. Pages 41-46 in E. S. McArthur, R. M.
and J. Faaborg. 1995. Regional forest fragmentation and the nesting Romney, S. D. Smith, and P. T. Tueller, editors. Proceedings—symposium
success of migratory birds. Science 267:1987-1990. on cheatgrass invasion, shrub die-off, and other aspects of shrub
119. Rogers, L. E., R. E. Fitzner, L. L. Cadwell, and B. E.Vaughan. 1988. biology and management. General Technical Report INT-GTR-276. U. S.
Terrestrial animal habitats and population responses. Pages 182-250 in Forest Service, Ogden, UT.
W. H. Rickard, L. E. Rogers, B. E.Vaughan, and S. F. Liebetrau, editors. 143. Uresk, D. W., J. F. Cline, and W. H. Rickard. 1975. Diets of black-tailed
Shrub-steppe: balance and change in a semi-arid terrestrial ecosystem. hares on the Hanford Reservation. Report BNWL-1931, Battelle,
Elsevier, Amsterdam, The Netherlands. Pacific Northwest Laboratory, Richland, WA.
120. ———, and W. H. Rickard. 1977. Ecology of the 200 Area plateau 144. ———, and V. A. Uresk. 1980. Diets and habitat analyses of mule
waste management environs: a status report. Report PNL-2253, deer on the 200 Areas of the Hanford Site in southcentral Washington.
Battelle, Pacific Northwest Laboratory, Richland, WA. Report PNL-2461, Battelle, Pacific Northwest Laboratory, Richland,
121. Rotenberry, J. T., and J. A. Wiens. 1991. Weather and reproductive WA.
variation in shrub-steppe sparrows: a hierarchical analysis. Ecology 145. Van Vuren, D., and M. P. Bray. 1985. The recent geographic distribution
72:1325-1335. of Bison bison in Oregon. Murrelet 66:56-58.
122. Saab,V. A., and T. D. Rich. 1997. Large-scale conservation assessment 146.Vander Haegen, W. M. 1996. Survey of breeding bird communities on
for Neotropical migratory land birds in the interior Columbia River BRMaP sites, Hanford Site, 1996. Project completion report.
Basin. General technical report PNW-GTR-399. U.S. Forest Service, Washington Department of Fish and Wildlife, Olympia, WA.
Pacific Northwest Research Station, Portland, OR. 147. ———, F. C. Dobler, and D. J. Pierce. 2000. Shrubsteppe bird
123. Savage, D. E., J. A. Young, and R. A. Evans. 1969. Utilization of response to habitat and landscape variables in eastern Washington,
medusahead and downy brome caryopses by chukar partridges. Journal USA. Conservation Biology 14:1145-1160.
of Wildlife Management, 33:975-978. 148. ———, and B. Walker. 1999. Parasitism by brown-headed cowbirds
124. Schroeder, M. A., J. R.Young, and C. E. Braun. 1999. Sage Grouse in the shrub-steppe of eastern Washington. Studies in Avian Biology
(Centrocercus urophasianus). In A. Poole and F. Gill, editors,The Birds of 18:34-40.
North America, No. 425. The Birds of North America, Inc., Philadelphia, 149.Verts, B. J., and L. N. Carraway. 1998. Land mammals of Oregon.
PA. University of California Press, Berkeley, CA.
125. Schwartz, C. C., and J. G. Nagy. 1976. Pronghorn diets relative to 150. Wallmo, O. C. 1981. Mule and black-tailed deer distribution and
forage availability in northeastern Colorado. Journal of Wildlife habitats. Pages 1-26 in O. C. Wallmo, editor. Mule and black-tailed deer
Management 40:469-478. of North America. University of Nebraska Press, Lincoln, Nebraska.
126. Seidensticker, J. C., IV, M. G. Hornocker, W.V. Wiles, and J. P. Messick. 151. Washington Department of Fish and Wildlife. 1995. Washington State
1973. Mountain lion social organization in the Idaho Primitive Area. management plan for the bighorn sheep. Game Division, Washington
Wildlife Monographs 35. Department of Fish and Wildlife, Olympia, WA.
127. Small, R.J., and B.J. Verts. 1983. Responses of a population of 152. Weinberg, H. J., and R. R. Roth. 1998. Forest area and habitat quality
Perognathus parvus to removal trapping. Journal of Mammalogy 64:139- for nesting wood thrushes. Auk 115:879-889.
143. 153. Weiss, N. T., and B. J.Verts. 1984. Habitat and distribution of pygmy
128. Smith, E. L., et al. 1995. New concepts for assessment of rangeland rabbits (Sylvilagus idahoensis) in Oregon. Great Basin Naturalist 44:563-
condition. Journal of Range Management 48:271-282. 569.
129. Smith, G. W., and D. R. Johnson. 1985. Demography of a Townsend 154. Whisenant, S. G. 1990. Changing fire frequencies on Idaho’s Snake
ground squirrel population in southwestern Idaho. Ecology 66:171-178. River Plains: ecological and management implications. Pages 4-10 in E. S.
130. Springer, J. T. 1982. Movement patterns of coyotes in south central McArthur, R. M. Romney, S. D. Smith, and P. T. Tueller, editors.
Washington. Journal of Wildlife Management 46:191-200. Proceedings—symposium on cheatgrass invasion, shrub die-off, and
131. Stebbins, R. C., and N. W. Cohen. 1995. A natural history of other aspects of shrub biology and management. U. S. Forest Service,
amphibians. Princeton University Press, Princeton, NJ. Ogden, UT.
132. Stewart, G., and A. C. Hull. 1969. Cheatgrass, Bromus tectorum L.—an
 316 Wildlife–Habitat Relationships in Oregon and Washington

155. Wickstrom, M. L., C. T. Robbins, T. A. Hanley, D. E. Spalinger, and S. M.

Parish. 1984. Food intake and foraging energetics of elk and mule deer.
Journal of Wildlife Management 48:1285-1301.
156. Wiens, J. A. 1985. Habitat selection in variable environments: shrub-
steppe birds. Pages 227-251 in M. Cody, editor. Habitat Selection in
birds. Academic Press, New York, NY.
157. ———, C. S. Crawford, and J. R. Gosz. 1985a. Boundary dynamics: a
conceptual framework for studying landscape ecosystems. Oikos
45:421-427.
158. ———, and J. T. Rotenberry. 1981. Habitat associations and
community structure of birds in shrub-steppe environments. Ecological
Monographs 51:21-41.
159. ———, and ———. 1985. Response of breeding passerine birds to
rangeland alteration in a North American shrubsteppe locality. Journal
of Applied Ecology 22:655-668.
160. ———, ———, and B. Van Horne. 1985b. Territory size variations in
shrub-steppe birds. Auk 102:500-505.
161. ———, ———, and ———. 1986. A lesson in the limitations of
field experiments: shrub-steppe birds and habitat alteration. Ecology
67:365-376.
162. ———, B. Van Horne, and J. T. Rotenberry. 1987. Temporal and spatial
variations in the behavior of shrub-steppe birds. Oecologia 73:60-70.
163. Wilcove, D. S. 1985. Nest predation in forest tracts and the decline
of migratory songbirds. Ecology 66:1211-1214.
164. ———, C. H. McLellan, and A. P. Dobson. 1986. Habitat
fragmentation in the temperate zone. Pages 237-256 in M. E. Soule,
editor. Conservation biology: the science of scarcity and diversity.
Sinauer Associates, Sunderland, MA.
165.Yensen, D. L. 1980. A grazing history of southwestern Idaho with
emphasis on the Snake River Birds of Prey Area. U. S. Bureau of Land
Management, Boise District, Boise, ID.
166.Yensen, E. and D. L. Quinney. 1992. Can Townsend’s ground squirrels
survive on a diet of exotic annuals? Great Basin Naturalist 52:269-277.
167.———, ———, K. Johnson, K. Timmerman, and K. Steenhof. 1992.
Fire, vegetation changes, and population fluctuations of Townsend’s
ground squirrels. American Midland Naturalist 128:299-312.
168.Yoakum, J. 1958. Seasonal food habits of the Oregon pronghorn.
Interstate Antelope Conference Transactions 10:58-72.
169. ———. 1980. Habitat management guides for the American
pronghorn antelope. Technical Note 347, Bureau of Land Management,
Denver, CO.
 12
Wildlife of Urban Habitats
Howard L.Ferguson, Kevin Robinette,
& Kate Stenberg

Description, History, and Status

People often believe that when land is urbanized, it loses The low-density zone generally has the most native
its value for wildlife. The development of cities does not wildlife habitat available, the largest blocks of
eliminate all wildlife habitats—some are lost, some are unfragmented land, and the most diverse assemblage of
altered, and some are maintained. In the process of native plants and animals. This same trend is true for
urbanization, some species of wildlife are favored while habitat structure. The high-density zone typically has little
others are stressed or eliminated. Over the past two understory, and few, if any, snags, downed woody
decades researchers have studied wildlife in urban areas; material, and brush or rock piles.2, 102, 170 These key habitat
most of this urban research has been conducted in eastern structures increase as you move out from the inner city.198
North America and Europe, with little having been And, as expected, both the abundance and the diversity
conducted in the Pacific Northwest. More needs to be of wildlife present for a particular area depend on its
learned in this region because the issue of urbanization location along this continuum. The terms “urban” and
and the protection of our natural resources, including “suburban” refer to concentrated human development in
wildlife and their habitats, is being raised daily in Oregon and around towns and cities. Our use of the word “urban”
and Washington. encompasses all 3 urban zones as described above and in
Attempting to characterize urban habitats and urban Chapter 2. Rather than being circular, the 3 urban zones
wildlife is challenging. Urban ecosystems are in perpetual typically appear as zones of urbanization along highways,
change.43 They are a mix of natural and human-made rivers, or valleys (often looking like isotherms). Further
activities and disturbances managed and unmanaged complicating the continuum is the ontogeny of urban
lands, native and non-native plants, and natural and “sprawl,” the irregular or unplanned spread of
human-made structures all merged together.198 Instead of urbanization. It is caused by people, and in some cases,
one or two management goals, as is common in a forest businesses, in the inner suburbs moving farther and
stand, each urban landowner has a different goal and a farther out as the city continues to grow. State growth
different strategy for achieving that goal. management laws attempt to address and control urban
An urban area may be viewed as a continuum ranging sprawl (see section on “Growth Management”).
from the highly developed inner city out to the relatively The difference between urban conversions and many
undeveloped rural areas.18, 30, 43, 50, 67, 69, 137, 208 The position on other types of land conversions is the permanence of the
the urban-rural gradient has a significant impact on land- changes made to the landscape—changes are maintained
cover regimes and can actually induce substantially by humans and are likely to persist for hundreds of years.
different landscape structures.208 This concept implies that Typically, urbanization removes, alters, or replaces natural
environmental variability is spatially structured and that or existing vegetation with impervious surfaces.18, 179 Plant
this pattern influences ecological processes and annual species composition is changed by the culturing of some
population dynamics.18, 137 We have chosen to follow the native species at the expense of others and by the
concept of an urban gradient in this chapter. This introduction of exotic species. Selected, non-native plants
continuum is based on the level of urban development as flourish in urban settings because of the increased
determined by the percent of the land surface covered by availability of water, fertilizers, and cultivation. Exotic
impervious materials. At the center of this continuum is shrubs and trees outnumber native shrubs and trees 6:1
the highly developed area of the inner city, what we call in the understory layers.17
the high-density zone, having >60% impervious surface With the influx of exotics and the overall intensive
(area covered by roads, buildings, parking lots, sidewalks, maintenance of urban areas, succession is altered and the
and driveways). As you move out from the inner city, you community equilibrium changes.18, 134 The protection of
encounter the middle zone or the medium-density zone, human structures and property has led to intense fire
the area having 30-59% impervious surface. Farthest away suppression in and around urban areas, disrupting the
from the inner city is the low-density zone, with 10-29% natural fire disturbance regime. 133, 134, 157 Another
impervious surface. conspicuous consequence of urbanization is the

317
 318 Wildlife–Habitat Relationships in Oregon and Washington

fragmentation of the natural environment into a mosaic urbanization that has caused significant habitat alteration
of patches of different size, shape, composition, and age. has occurred during the last 50-100 years. Blondel19 stated:
Patches become surrounded by areas influenced by “Urban settlements (in the Old World) have existed for at
humans—cleared areas, roads, buildings, and other least 3,500 years, especially in the eastern Mediterranean”,
human-made structures—and, in general, become smaller and he described the progressive adjustments and
and more isolated toward the urban center46 (see section habituation of rural birds to urban life in Old World cities.
on “Habitat Loss and Fragmentation”). Succession is This can be contrasted with birds found in North American
further altered by human landscaping and maintenance cities, most of which have been occupied for <200 years.48,
104
activities. One example of a bird that seems to have adjusted to
Patches of native presettlement vegetation or special urban life in Europe but not in North America is the
or unique geological habitats, such as riparian areas, northern goshawk. It is recognized as an urban species in
canyons, rock outcrops, cliffs, and lakes, are often left Germany63, 64 and is increasing in human-influenced
undeveloped within urban zones. Wildlife found in and habitats in Poland,22 whereas in the U.S. it is considered
around these remnant habitats or inclusions are usually a an interior forest species that requires mature/old-growth
subset of the wildlife normally expected for each habitat. trees.
The species assemblage will be determined by the size of Urbanization is a widespread and ever-increasing
the remnant patch as well as the degree and amount of process. In 1989, 74% of the U.S. population (203 million
urbanization surrounding the remnant. Often, these people) lived in urban areas, and that number is expected
features are the reason for initial human settlement, and to increase to >80% by 2025.81, 99 This increase throughout
that is why many of our Northwest cities have such the country has resulted in the conversion of cropland,
habitats: Portland and Spokane are close to rivers; Tacoma pastures, and forests into urban and suburban
and Seattle are near Puget Sound waterways (Figure 1). environments.72 Between 1960 and 1970, urban land in the
These features can greatly enhance the wildlife abundance U.S. increased by 9 million acres (3,643,725 ha), and
and diversity normally expected to be found in an urban between 1970 and 1980 it increased by 13 million acres
area. These inclusions are the reason coyotes, bald eagles, (5,263,158 ha).83
ospreys, kingfishers, and even moose and cougars may In the Puget Sound area of Washington, between 1979
sometimes be found in the urban sections of our Pacific and 1989, suburban zone area increased by 7.3%, and
Northwest cities. Wildlife habitat, and the urban urban zone area increased by 15.7%. For example, nearly
environment, can be enhanced through the preservation 47% of the private land in Snohomish and Kitsap counties
of remnant forests, riparian areas, wetlands, cliffs, is now suburban.130 Between 1945 and 1970, 257,000 acres
meadows, fields, and orchards as cities expand.48 (104,049 ha) of Washington timberland were converted to
One factor that may influence the use of urban habitats urban use,24 and these westside forests are among the most
by wildlife in Oregon and Washington is the relatively productive in the U.S.24 From 1970 to 1992, nearly 10% of
young age of our urban areas. Urbanization has been all forests in the Puget Sound Region were converted to
extremely rapid in North America in comparison to other uses: roads, suburbs, cities, and farms. 130 An
Europe, Asia, and Africa. With few exceptions, events that estimated 30,000-80,000 acres (12,146-32,389 ha) of wildlife
took place slowly over many centuries in the Old World habitat is lost each year in Washington state; >90% of its
have been accomplished in just a few decades in the New wetlands in urban areas and 90-98% of its urban coastal
World; animals have had less time to adjust to urbanization wetlands are already gone. 206 The greatest losses of
in North America. In the western U.S., most of the Washington freshwater wetlands have probably resulted
from urban development.37 From 1992 to 1998, >16,000
acres (6,478 ha) of natural areas were lost to development
and other changes in the Portland metropolitan area (J.
Budhabhatti, Senior Planner, Portland Metro, pers.
comm.). The need for immediate conservation and
regional and statewide planning is essential, considering
that by 2045 our population is expected to nearly double.206

Common Ecological Features

of Urban Habitats
Buildings and Human Structures
Characteristic of urban areas are buildings, streets, roads,
parking lots, and other artifacts of human construction.18,
134
These structures occupy most of the ground surface and
form a largely impermeable and sterile covering of the
Figure 1. Spokane River in Spokane, Washington. soil, which once supported native vegetation. The water
(Photograph by Howard L. Ferguson) runoff from these impervious surfaces is high and fast,
with little infiltration into underlying strata, and results
 CHAPTER 12: URBAN ENVIRONS 319

in a reduced rate of recharge to natural groundwater Northwest, but it may benefit wildlife in the more arid
reservoirs, aquifers, and a lowering of the water table.126 areas of Oregon and Washington.
Emlen73 noted that the special features and artifacts of
human origin contributed importantly to the Light
physiognomy and diversity of the urban habitat. Rooftops In general, there is greater illumination in urban areas
and their superstructures, such as air-conditioner units because of increased reflective surfaces and widespread
and television antennae, provide song and resting perches artificial lighting. Artificial lighting prolongs the day
for birds. Supplemental nesting sites provided by human length in cities, which may affect diurnal cycles and trigger
structures include vent holes between bricks, vent holes earlier reproductive activity in some species.74, 200 It may
for dryers, openings around eaves and eave troughs, edges also influence migratory behavior and interfere with
of tile roofs, ledges under roofs, window shutters, and reproductive behavior of nocturnal species. Artificial
decorative boxes.23, 54 Overhead wires and other high song lighting may be a major factor contributing to the problem
perches may enhance the value of the urban area for of bird collisions with buildings.116, 117
territorial species, whereas traffic hazards, cat predation,
and disturbance or harassment by dogs and humans are Habitat Loss and Fragmentation
selectively detrimental to ground nesting and low-shrub The most obvious consequence of urbanization is the loss
foraging birds.23, 54 of rural or agricultural habitat. Much of the land converted
The major direct contribution of the human-built for urbanization may be considered loss of habitat because
environment to bird communities may be an increased of the permanence of the impervious surfaces covering
supply of holes, crevices, and ledges for nesting.73 The these lands. With urban development and loss of habitat
degree to which these human structures positively comes fragmentation (the division of habitat into smaller,
impacted birds was questioned by Lancaster and Rees123 more isolated patches of natural vegetation). As the
when they surmised that the major contributing elements landscape matrix shifts from natural vegetative cover
of buildings and other human-made structures in urban types to urban land uses, the natural habitats become
environments are, essentially, closed volumes and barren remnants or patches—smaller areas that are isolated and
surfaces with limited potential for diversification of niches. disconnected from each other. The ability of these patches
They noted that even the song and resting perches to support specific wildlife species is a function of their
associated with buildings, poles, and wires are size and isolation, as well as the neighboring vegetative
insignificant when compared with those made available type.4, 167
by comparable cover by trees and bushes.123 However, The impact of fragmentation depends on the
when natural features and structures are not present, such surrounding habitat;167 this may explain the different
as in downtown urban areas, these human structures do conclusions being reported for eastern U.S. and Pacific
provide some potential nesting and roosting sites for Northwest forests. Forest fragmentation in the eastern
animals able to adjust, or for those few commensal animals U.S., at both the landscape and site level, results in serious
already adapted to urban areas. impacts on wildlife.60, 211, 213 It can result in genetic isolation
of less-mobile species (e.g., reptiles and amphibians).175
Climate Because of species’ area needs, population isolation, and
The climate in large urban centers can differ substantially spatial heterogeneity, fragmentation further reduces the
from the surrounding exurban areas. Cities are warmer, amount of habitat available for certain species.34, 49, 202 In
with higher average daily temperatures and less variable southern California, Bolger 21 found that in areas
daily and annual temperatures, than the surrounding surrounded by urban development, fragmentation
areas.68, 74, 191, 201 In San Francisco, differences of up to 20oF seemed to cause more isolation than that reported for the
(11oC) have been recorded between the densely built-up forested landscapes of eastern and mid-western U.S. He
business districts of the city and the undeveloped urban reported that local extinctions in these isolated habitat
parkland of Golden Gate Park.69 In central California, this fragments were common, and that recolonization of these
temperature differential increased with increasing city areas was rare. Studies reporting significant negative
size.69 Warmer temperatures may be a factor that has impacts from fragmentation had either agriculture or
allowed, at least indirectly, some species to overwinter urban habitats as the surrounding or neighboring land
farther north than they did historically (e.g., blue jay182 use, whereas the patches of the Northwest are typically
and merlin110). Studies elsewhere have revealed some surrounded by different-aged forests—not agriculture or
evidence that breeding may begin earlier and end later in urban. These differences may explain why the few studies
the year partially due to this temperature differential in from the Northwest have reported fewer impacts from
cities.74, 88, 143 fragmentation on wildlife than the eastern U.S. Just what
More precipitation occurs over highly urbanized areas impact does fragmentation have on urbanized areas of
compared to similar rural areas of the same region.107 the Pacific Northwest? More research is needed to
Additional heat generated by cities causes air above cities determine the impact that fragmentation has on wildlife
to rise more rapidly than from the surrounding in urban areas of the Northwest.
countryside, resulting in more rapid cooling and rainfall. Another effect of fragmentation is the creation of more
Consequences of this are largely unknown for the Pacific edge, the interface where different plant communities
 320 Wildlife–Habitat Relationships in Oregon and Washington

meet. Moderate levels of urban development fragment decreased feeding, habitat avoidance and abandonment,
formerly continuous habitat, increasing the amount of and reproductive losses. Long-term exposure to noise
edge. If development occurs in non-continuous small causes excessive stimulation to the nervous system,
blocks, the amount of edge increases even more. As time resulting in chronic stress that may harm the overall health
goes on, and urbanization and development increase, of wildlife and their reproductive fitness.79
more and more native habitat is eliminated, and edge Predation rates are higher in urban areas16, 68, 94, 194, 196 in
habitat will eventually decrease, to the point where all comparison to similar exurban areas; with an increase in
native habitats are eliminated and no edge habitat exists.18 edge comes an increase in nest predation and brood
Whereas edges are beneficial to some wildlife species, they parasitism.181, 212 Avian nest predators and brood parasites
also can negatively impact others.4, 165, 213, 216 Higher nest are associated with strip corridors and edges between
predation and parasitism rates have been reported for fields and woodlands, resulting in decreased nesting
small, fragmented eastern U.S., where edge was bounded success for some bird species. One explanation for this is
by either urban or agriculture land use.131, 156, 211, 212 that predators can penetrate deeper211, 212 or are able to
search smaller areas more efficiently.131 However, Martin132
Roads believes nest predation cannot fully explain the losses of
Roads are a major component of urban areas and have bird species from fragments, and believes other ecological
major impacts on wildlife, wildlife habitat quality, and requirements, such as nest site preference, in combination
connectivity. In urban areas, roads or road corridors with increased predation may explain loss of species.
function ecologically as (1) habitat for animals, their Results from a study in the eastern U.S.31 attributed the
predators, and competitors; (2) sources and collection lack of nesting success of ovenbirds in fragmented forests
points of contaminants; (3) sources of human disturbance; to reduced food supplies.
(4) partial or complete barriers to animal movement; (5) Several authors have attributed increased predation in
conduits for animal movement or range expansion; (6) urban areas to human pets—cats and dogs.44, 114 Domestic
barriers to surface water percolation (lowering cats can be efficient small predators and may occur in high
groundwater tables); (7) sources of fragmentation of numbers in urban areas.40 Prey of free-ranging domestic
wildlife habitats; (8) facilitators of the spread of non-native cats in one study was approximately 70% small mammals,
species; and (9) mortality agents.80, 206 Roads divide once 20% birds, and 10% various other animals including
large populations into smaller groups, which are at greater reptiles and invertebrates.78 Cats often outnumber other
risk of extinction and genetic stagnation.80, 124, 125 Roads mid-sized native predators and may compete with them
often cut directly across long-established animal trails to for many of the same food sources.44 Buildings and barren
water and feeding grounds,169 and with increased road ground reduce and simplify vegetation within patches,
density, human access increases, which can result in more and provide hunting areas for domestic cats and dogs that
disturbances.80 Additional information on roads and the may effectively reduce the local abundance of vertebrate
impacts to wildlife can be found at http:// prey.60 For more information on feral cats, see http://
www.fhwa.dot.gov/environment/wildlife crossings/. www.dfg.ca.gov/hcpb/feralcat.html.

Human Disturbance, Noise, and Predation Food

Urban wildlife species are constantly exposed to human- The type and quantity of foods available in the urban
related disturbances, including noise and disturbance environment differ considerably from those found in
related to human activity around homes and buildings, undisturbed or more natural areas. Many authors have
road traffic, construction activities, and aircraft activity.207 commented on the abundance of food available in the
Some wildlife species become accustomed to regular and urban environment.32, 55, 56, 74, 94 This abundance is limited
predictable disturbances. Watson and Pierce207 found that to a few food types; quantities of seeds, fruits, and refuse
irregular noise and unpatterned disturbances—especially are unintentionally provided through spillage, waste,
human pedestrian disturbances—had more negative disposal, and collection and accumulation of food at refuse
impact on bald eagles than disturbances caused by autos, sites and landfills. Even the ubiquitous urban lawn can
boats, and aircraft. In Puget Sound, human activity altered be highly productive for certain types of food, e.g., insects
the normal feeding guild relationships of crows, gulls, and and worms,76, 82, 94 if chemical use is limited.
bald eagles.180 Eagles were least tolerant of human activity Considerable quantities of food are also provided
and gulls were most tolerant, even feeding more in the throughout the year by people. This food is primarily seeds
presence of humans. This behavior may negatively impact and nuts, the exception being nectar (hummingbird)
the less tolerant eagles in urban areas during years with feeders. In 1994, Americans spent >$2 billion on birdseed
limited food resources.180 alone; this figure does not include the cost of feeders and
For birds, noise may interfere with intraspecific other equipment.188 The urban environment provides these
communications required for successful reproduction, and artificial seed sources in much greater quantities and
noise may have contributed to the elimination of some concentrations than would ever occur in natural or
key interior species in the eastern U.S.115, 118, 161 In his book undisturbed habitats. Another center of human-supplied
on noise and wildlife, Busnel33 concluded that noise, at food is parks, where quantities of bread and other human
least indirectly, might cause bodily injury, energy loss, foods are provided to wildlife (Figure 2).
 CHAPTER 12: URBAN ENVIRONS 321

observed in ants,201 click beetles,151 birds,16, 73, 93, 94, 136 reptiles
and amphibians,140, 155 and mammals.198
A significant impact of urbanization is the removal,
alteration, and replacement of the natural vegetation.17 The
resultant urban vegetation differs from presettlement
vegetation in several ways: species composition is altered,
sterile varieties are common, and most importantly, woody
plants are sparsely distributed. Habitats are lost for those
species adapted to the naturally occurring vegetation types
and structures of an area. In urban areas, bird and small
mammal diversity is correlated with both the abundance
and diversity of vegetation, and also with habitat
heterogeneity;123 however, highest species richness is not
always in the most natural habitats, but often occurs in
moderately disturbed ones.157 Urban landscapes support
Figure 2. Riverfront Park, Spokane, Washington. a variety of wildlife species that are adapted to, or have
(Photograph by Howard L. Ferguson) adjusted to, urban areas. 73, 199 Refer to Table 1 for a
summary of the environmental conditions and constraints
Replanting with non-native plants similarly alters the confronting urban wildlife and the ecological
availability of foods.17 Native plant food sources, including consequences of these conditions. See Table 2 for a
native berries, fruits, and seeds, are typically unavailable summary of adjustments made by wildlife in response to
or present in limited amounts in urban areas. Exotic plants the urban environment.
support fewer insects than do native plants,133 and the
urban practice of controlling insects with broad-spectrum Birds
pesticides further limits the availability of insects in the A common result of increased urbanization is a decline in
urban environment. Removal of all down and dead wood, number of species and a simultaneous increase in total
diseased trees, and snags from the urban landscape bird density; i.e., an avifauna comprised of a few exotic
curtails the food base for those species that depend on species that become very abundant. 14, 56, 73, 123 The
insects or fungi from these sources.187 Consequently, the proportion of native bird species, especially shrub
urban food available to wildlife contributes to reduced dwelling, ground nesting, or insectivorous canopy feeders,
species diversity while it sustains high densities of favored, progressively declines as urban development intensifies.53
often exotic, species. 20, 123 The visible increase of the Flocks of gregarious, weakly territorial, omnivorous, and
abundance of a few species of ground feeding, cavity- granivorous species replace territorial73 and insectivorous
nesting omnivores (e.g., house sparrows and starlings) is migrant species.51, 53 The tempering of the urban climate
consistent with the nature and availability of urban food, may reduce prey fluctuation, increase prey biomass, and
which also corresponds to the gradient of increasing reduce bird energy requirements, enabling urban birds to
urbanization.123 attain higher densities, breed for longer periods of the year,
produce more young, and have higher annual survival.134
Wildlife Use of Urban Habitats The granivorous and omnivorous guilds make up
Most of the ecological research on wildlife in urban areas nearly 93% of the total avian biomass of urban areas.17, 73,
215
has been conducted in the eastern and southwestern U.S. Insectivorous species are common in the low-density
and Europe, and most of the data and references presented zone, but are nearly absent from the high- and medium-
in this chapter are results from those studies. Little research density zones, especially in winter.43, 51 Bark drillers and
has been done in the urban areas of the Northwest, but gleaners are reduced in the high-density zone as well.17
the basic impacts of urbanization are consistent.43, 199 Differences in nesting guilds are also evident. Species
Clergeau et al.,43 in a study comparing Quebec, Canada nesting on tree branches or buildings are more abundant
with Rennes, France, found similar patterns for species in the high-density zone, whereas ground, cavity, shrub,
abundance and compositions in both cities. They and tree-twig nesters either are absent or occur at low
concluded that the physical setting of a city does not densities in this same zone.56 Scavenging omnivores (e.g.,
greatly influence the structure of the urban bird gulls, corvids, blackbirds, and European starlings) benefit
community.43 Most of the results from these urban studies from spilled waste.134 Emlen73 characterized native species
are likely to be true for the Northwest, and if not, can at in urban areas as often having wide geographic ranges
least serve as starting points for studies in our area. and broad ecological tolerances, and by definition are
Overall, as urbanization increases, species diversity opportunistic species (e.g., mourning dove and house
declines and species densities increase, primarily because finch).
of high numbers of a few exotic species.53, 123 These exotic In Canada and France, Clergeau et al.43 found that the
species may make total abundance and biomass actually pattern of breeding bird abundance, unlike species
higher in urban areas than in adjacent, more natural diversity, is not directly associated with the urbanization
areas. 73, 123 The decline in species diversity has been gradient. Savard172 found a similar pattern in Toronto, with
 322 Wildlife–Habitat Relationships in Oregon and Washington

Table 1. Environmental conditions and constraints of urban areas and the ecological consequences.

Environmental conditions Ecological consequences

and constraints

Altered disturbance regime—

Landscaping with non-natives Alteration of type and available timing of food resource. Change in cover type. Constant or
varied maintenance.
Suppression of fire Altered succession, with unpredictable seral stages. Alteration of equilibrium.
Altered phenology Different pollination schedules. Provides extended duration of season.
Concentrated human populations Human and associated pet predation. Reduction in large predators. Large amount of human
waste. High potential for human disturbance.
Elimination of snags Elimination of habitat for primary and secondary cavity nesters, and food for many
insectivores.
Extended photoperiod— Increased day length—earlier breeding cycles, multiple broods. Lights act as effective barriers
artificial lighting to movements and dispersal.
Extensive lawn or “savannah” habitats Improved foraging for some species like robins and starlings. Elimination of foraging and nest
sites for ground- and shrub- dependent wildlife.
Fragmented landscapes Small isolated patches of habitat. Small, isolated, and high-density populations, more vulnerable
to extinction. Decreased mobility and dispersal potential.
Higher temperatures Lower energetic requirements. Allows multiple broods. May allow some birds to “winter”
over.
Human byproducts—garbage Increased food sources. Concentration of feeding animals. This dependence may interrupt
seed dispersal and pollination.
Human byproducts—sewage ponds Increased habitat and extended ranges for wetland-associated wildlife.
Human structures—barns, bridges, Increased nesting and roosting structures. Increased danger for flying animals.
buildings, chimneys, eaves, culverts
Human structures—bird boxes Temporary supplement or substitute for snags and associated cavities. Also supports
aggressive exotic birds.
Human structures—bird feeders Sustains or supplements birds and food. Seed eaters concentrated in these areas. Provide prey
concentrations (e.g., for accipiters).
Increased impervious surfaces Concentration of pollutants, decrease in stormwater percolation, increased sedimentation,
etc.
Increased fertilizers High NPK, increased algal blooms impact other beneficial vegetative growth. Low oxygen, can
cause fish kill.
Mobility barriers—lights, roads, Wildlife forced to disperse across unsuitable and dangerous habitats. High predator risk.
vehicles, structures
Power poles, towers, and power lines Increased nesting and roosting. Provides travel routes (e.g., squirrels). Increased mortality.
Presence of human pets Increased predation, particularly ground nesters and feeders
Roads Decreased mobility, increased mortality, isolation. Increase in roadside habitats.
Simplification of habitat—little or no Decrease of ground- and shrub-nesting species. Increase in shrub nesting height.
understory or mid layer (conversion
to lawn)

Table 2. Ecological adjustments made by urban wildlife.

Physiological Behavioral Life history

Greater longevity Changes in nesting habits— Reduced migrations, winter residency in areas formerly
higher nests. vacated.
Prolonged breeding season Changes in feeding and foraging Changes in food/diet—human waste products.
behavior.
More and/or multiple Tolerance of humans. Increased Changes in nesting habits.
broods intraspecific tolerance. Change in
aggressiveness—increased and
decreased.
Prolonged circadian cycles Higher population densities. Higher population densities.
Increased nocturnal activity, and
extended mating and breeding.
 CHAPTER 12: URBAN ENVIRONS 323

a higher bird abundance in residential areas adjacent to Table 3. Total number of species by association in
downtown than in the downtown area itself, a decrease each urban zone. The number in parenthsis is the
in abundance in old residential neighborhoods with tall percent of non-native species in each category.
trees, and an increase in more open and young residential
neighborhoods, followed by a decrease in the least High- Medium- Low-
urbanized sectors. density density density
It is the type and volume of vegetative cover,
particularly the shrub and canopy layer, and habitat Closely associated 12 (67) 13 (62) 15 (53)
patchiness, that influences avian diversity and numbers Associated 10 (0) 68 (3) 145 (6)
rather than species composition.16, 73 In urban areas, the Present 32 (3) 77 (7) 104 (2)
amount of woody vegetation has been singled out as the
most important factor for promoting a diverse native bird especially merlins, and accipiters seem to be taking
assemblage.199 Numerous studies, mostly from the eastern advantage of the increased prey base.48 The urbanization
U.S., have shown a positive linear relationship between of the merlin in Saskatoon, Saskatchewan, has been well-
bird species diversity and foliage height diversity across documented.110
the urban gradient.53, 123 The diversity of birds in urban Prey of free-ranging domestic cats was approximately
areas was also influenced by the age of the 20% birds in several studies.40, 78 Coleman et al.44 estimated
neighborhood, 127 type of housing, 89 and degree of between 8 and 217 million birds were killed each year by
urbanization.13 However, in California, Blair18 found that cats in Wisconsin. Other avian predators that seem to
bird diversity, bird density, and bird biomass peaked at increase in urban areas are sharp-shinned hawks, Cooper’s
intermediate levels of urbanization compared to the most hawks, and merlins.133 Human predators can also seriously
natural sites. McDonnell et al.137 predicted that species deplete local songbird populations.133 Yet Martin131 believes
richness should peak at an intermediate level of that nest predation is probably the most limiting factor
development because biotic limitations are high at the for songbirds, and that nest predators may be more
rural end, whereas physical limitations are high at the abundant in urban areas than in native habitats, having
urban end. increased dramatically in the western U.S. during the last
Unlike in forested regions, urban development in century.133 In Arizona, Emlen73 concluded that 2 ground-
deserts,73, 96 shrub, or steppe typically results in increased nesting species had been eliminated by disturbances in
insectivorous species because of the growth and an urban area.
proliferation of planted and artificially watered woody
vegetation. This increase seems to be true only for the Mammals
short-term; if urban development becomes denser over Urban mammals have been studied much less than urban
time, diversity declines. In the arid southwest, Soulé et birds. Mammalian species richness seemed to decline
al.186 found that the xeric chaparral-obligate species began rapidly with increasing human disturbance and increasing
disappearing a few decades after being isolated by urban barren ground,148 and decreasing distance to human
development. habitation. Species richness increased with increasing
The heights of birds’ nests in urban areas in the eastern density of vegetation per patch, especially in the shrub
U.S. are higher than in rural areas in response to increased mid-layer.60 Small mammal populations in urban areas,
predation, human disturbance, or to decreased amounts like birds, seem to be positively correlated with density
of lower-level vegetation.32, 52, 173 DeGraaf et al.54 found that of vegetation—residual patches of natural and semi-
various species in the eastern U.S. differed in nesting natural vegetation support the most dense populations.61,
62, 197
height in response to housing density, which may be In Oxford, Dickman and Doncaster61 found that
interpreted as a measure of disturbance. They also found remnant patches of natural or semi-natural vegetation
that shrub-nesting species respond to available vegetation contained the highest density of small mammals.
rather than to the level of disturbance in an area. Intensively managed and maintained patches, like parks
In Colorado, Cringan and Horak 48 found that and golf courses, contained fewer small mammals than
urbanization had a complex impact on raptors. Solitary expected.62
species with large home ranges were negatively impacted, Whereas many specialized native mammalian species
whereas small-mammal eaters were positively impacted are excluded from urban areas, a few native omnivorous
at first, but ultimately their populations decreased as and scavenging species benefit from the food and shelter
urbanization increased. Bird-eating raptors benefited from provided by humans. Several non-native species thrive
the large prey populations in cities, if minimum habitat in the urban setting and are more abundant in urban areas
resources and sufficient protection were available.48 than in more natural habitats.29 As Obara et al.152 noted,
Cringan and Horak48 further speculated that as biomass the urban environment is often characterized by the
of the urban bird population increased, a behavioral replacement of native by non-native species (Table 3).
response would probably first occur, with migrant and Small native mammals that do survive seem to show a
vagrant raptors feeding on the abundant city birds, low degree of territoriality and high intraspecific tolerance.
particularly during winter. If nesting habitat was suitable, These behavioral characteristics facilitate aggregations of
this might be followed by a nesting response. Falcons, individuals in favorable microhabitats and movements
 324 Wildlife–Habitat Relationships in Oregon and Washington

between habitat patches (buildings) that help the small movements between these upland patches and breeding
mammals exploit the urban environment. Yet any sites (e.g., wetlands), and the resultant isolation of
disruption of these small remnant urban patches will likely individuals from these environments and each other.162 In
affect these concentrated species more than the larger, less watersheds with urban development, amphibian
concentrated species that move frequently between abundance decreases and the quality of the remaining
patches.61, 197 Soulé et al.,185 in their study of urban San breeding habitat is severely impacted by runoff.162, 163
Diego, found that rodent populations seemed to be Clearly, reptiles or amphibians restricted to water for
particularly susceptible to local extinctions in a fragmented breeding, egg laying, and development are harmed by
urban environment. wetland loss.97, 140 Even the smallest wetlands are important
A limiting factor of small mammals in urban areas may habitats for some amphibians.91, 163 Another complicating
be the presence of predators, such as coyotes,158, 159 cats,35 factor in the conservation of amphibians is their strong
dogs,15 and foxes.65, 98 Small mammals may comprise up to philopatry to breeding site wetlands—adults may return
70% of the total prey mass of free-ranging domestic cats.78 to breeding sites several years after the sites have been
These urban predators may depress the distribution and destroyed.162, 189 Females of some species of snakes are more
abundance of many small mammal populations and prone to be road-killed than are males, because females
further reduce the availability of prey for small native travel more.36 Burrowing snakes are better able to survive
carnivores.3 in urban areas than are surface foragers, whereas larger
In many urban areas of the eastern U.S., deer are so or venomous snakes suffer great declines from
common they often become pests.204 High deer densities urbanization.36
have been attributed to the availability of abundant forage, There seems to be a strong correlation of terrestrial-
lack of large predators, and hunting restrictions within breeding amphibian distribution with large woody debris,
urban areas. In addition, these large ungulates have dead and decaying wood and organic matter, and other
become habituated to urban disturbances and tolerate habitat conditions favorable to thermoregulation,
human presence and disturbance. 204 Deer may foraging, and resting.162 However, reptiles and amphibians
occasionally attract large predators, like cougars or bears, are not as closely associated with vegetative complexity
into the urban area, which can cause human-wildlife as are birds and mammals. Instead, thin-stemmed
conflicts. emergents, including rushes, sedges, herbs, and grasses
There are basically 2 types of bats found in urban seem more important for lentic breeding species.164 Richter
environments in Oregon and Washington: the colonial bats et al.164 also found a strong inverse correlation between
including the little brown (probably the most abundant), species richness and water-level fluctuation and degree
big brown, pallid, Brazilian free-tailed, and western of urbanization. Increases in the duration and frequency
pipistrelle, and the solitary types including the hoary and of flooding, and the changes in discharge rates resulting
silver-haired bats. In urban areas, most bat species are from the magnification and increasing frequency of
found in fewer numbers and at fewer locations when existing storm peaks in urban watersheds decreased the
compared with rural sites.86 In Michigan, species diversity success of lentic breeding amphibians.27
was twice as high in rural areas as in the city.122 In this Urban development often cuts off and isolates wetlands
same study, big brown bats were the most common bats or riparian areas from formerly connected terrestrial
of the urban areas, but their numbers dropped environments. Amphibians are particularly vulnerable to
significantly in the high-density commercial zone. This this type of urban habitat fragmentation.6, 162 An example
may be the result of the lower insect numbers and diversity in Washington is the western toad, which was recently
found in urban areas as compared to rural areas.82, 122, 187 listed as a candidate species by Washington Department
Although increased foraging activity around artificial of Fish and Wildlife. Culverts may block fish and wildlife
lighting in urban areas has been reported by some,11, 77 in passage, whereas road crossings and other developments
Michigan no difference in foraging activity was detected.122 may pose barriers to wildlife following riparian
Bat foraging behavior may change in urban areas as well.84 corridors.145, 162 Streams in urban areas sometimes are
Urban animals may spend more time foraging than rural
ones, perhaps reflecting differences in prey density.86 The Table 4. Total number of species occurring in high-,
biggest problem bats may face in urban areas is medium-, and low-density urban areas
disturbance of day and night roost sites, maternity sites, in Oregon and Washington taken from
and hibernacula by humans and cats. Wildlife Habitats Data Matrix.154

Reptiles and Amphibians High- Medium- Low-density

Modification of habitat is probably the chief threat of density density
urbanization to reptiles and amphibians.36, 189 Urbanization
may destroy or severely alter upland habitats that are Amphibians 0 5 18
essential for land-dwelling stages or species, and Reptiles 1 12 21 (+1 unsure)
consequently, feeding and refuge (e.g., estivation and Birds 39 100 149 (+1 unsure)
hibernation) patches may be destroyed or fragmented.162 Mammals 14 41 76 (+1 unsure)
Such impacts increase the risk of extinction by hindering Total species 54 158 264 (+3 unsure)
 CHAPTER 12: URBAN ENVIRONS 325

completely buried in pipes or bisected by roads, making buildings.123 The English house sparrow and the European
them unavailable or unsuitable for both aquatic and starling nest in almost any cavity they can find,
terrestrial species. aggressively chasing off other cavity nesters. Savard and
Falls173 found that house sparrows preferred human
Wildlife Profiles of the structures to natural sites for nests. They seemed to prefer
crevices, especially holes in eaves, to open nests.173
Urban Zones Gulls feed at landfills and trash bins, and use parking
lots, schoolyards, and tops of buildings for roosting.
Wildlife in the High-density Zone Mallards are often seen in parks with ponds or a river,
Fifty-four wildlife species could potentially occur within
and gulls often ring the shoreline. Nectar (hummingbird)
this high-density zone (Table 4), with 12 species classified
feeders in this zone may attract rufous hummingbirds.
as closely associated, 10 as associated, and another 32
Bridges in the high-density zone often are used as nesting
species as present.154 These numbers are derived from a
structures for several bird species including white-
list of all species that may occur in any high-density zone
throated swifts, cliff swallows, crows, ravens, and
anywhere in either Oregon or Washington. Furthermore,
peregrine falcons. The American robin, house finch, and
each species will only be present if and when the
black-capped chickadee tend to avoid commercial and
appropriate habitat is available.
industrial areas found in this zone.123 Both killdeer and
Large, wide-ranging or ecologically specialized species
common nighthawks have historically nested on rooftops
and those that cannot tolerate high levels of human activity
in urban areas, but there are no recent records of rooftop
and disturbance are excluded from this high-density urban
nesting for the Northwest.154
zone. Species that do well are gregarious, weakly
Unusual as it may seem, an observer in this high-
territorial, human disturbance tolerant, and dietary
density zone may observe a merlin in winter110 or a
generalists.39, 73, 98 Exotic wildlife species often are most
peregrine falcon in summer. These two raptors have
abundant and conspicuous of all wildlife in this zone.199
adjusted to the conditions of this high-density zone, and
See Table 3 for a summary of non-native species in all
take advantage of the modified food sources, and in the
density zones.
case of the peregrine falcon, nesting structures of
Some ecological features faced by wildlife in this
downtown urban areas. 48 Merlins feed on house
densest zone are the presence of more concrete than soil,
sparrows153, 183 whereas many urban peregrine falcons may
an extensive stormwater system, a multitude of roads and
be targeting rock doves.9, 113
associated vehicles, ever-present background noise,
The high-rise buildings in this high-density zone have
artificial lighting, and a highly maintained and manicured
killed millions of birds; Mesure 139 estimated that
landscaping of non-native vegetation (Figure 3). Wildlife
approximately 100 million birds are killed each year in
dispersal or movement is limited and highly restricted,
North America. Klem116 reported the annual mortality to
and often dangerous.169 As a result, most of the species
be between 98 and 976 million birds. Why birds are unable
found in this zone are either birds, or small mammals with
to avoid urban skyscrapers is unknown, but it seems that
small home ranges.
mortality can be significantly reduced if the lights of these
Most native vegetation has been eliminated, and much
urban skyscrapers are extinguished at night.116, 117
of the wildlife habitats consist of human-made structures.
Rooftops, cornices, eaves, and arches—analogs of cavities, Mammals. Extreme urbanization has heavily impacted
rocky flats, rocky outcrops, cliffs and caves 23—may mammals; there are no native species of mammals listed
provide roosting and nesting habitats for a few species of as either “closely associated” or “ generally associated”
birds and bats, species adapted to such habitat features. within the high-density zone.154 Instead, the black rat,
In addition, most of the species found in this zone have Norway rat, and house mouse are the most common
adjusted their behavior and foraging habits or are already
adapted to take advantage of the waste products of
humans.
Birds. Seedeaters and ground-foraging omnivorous
species—starlings, house sparrows, and rock doves—are
the most common birds found in the high-density zone.73,
111
These exotics are tolerant of human activity and other
environmental disturbances found in urban areas, which
allows them to exploit artificial urban structures and out-
compete less aggressive native species. All 3 of these exotic
species are known to feed on human handouts. The rock
dove is a cliff dweller that uses buildings in the downtown
area for roosting and nesting.111 Rock doves are more
common in this high-density zone than any other, favoring
the industrial and commercial areas where there is an Figure 3. High-density urban zone. Neighborhoods in
abundance of roosting and nesting sites, especially in older Spokane, Washington. (Photograph by Howard L. Ferguson)
 326 Wildlife–Habitat Relationships in Oregon and Washington

mammals of the high-density urban areas; they are more Wildlife in the Medium-density Zone
abundant here than in more natural habitats.29 They are Larger remnant patches of native vegetation and less
all Old World exotics that have coexisted with humans fragmentation result in a 3-fold increase in the number of
for centuries. Only in coastal areas may the black rat wildlife species potentially occurring in this zone
outnumber the Norway rat.135, 210 These small mammals compared to the high-density zone. A total of 158 wildlife
use small cavities associated with the plumbing and species potentially occur within this medium-density zone
electrical pathways of buildings or stormwater pipes as (Table 4); 13 species are classified as closely associated, 68
burrows, and power and telephone lines as travel as generally associated, and another 77 species as
corridors, 135 and they have been observed feeding at potentially present. These numbers are derived from a list
dumpsters.95 These exotic urban dwelling species are well of all species that may occur in any medium-density zone
adapted for urban areas by having small or reduced home anywhere in Oregon or Washington.154 Each species will
ranges.39, 98 The house mouse may have a total home range only be present if and when the appropriate habitat is
of 10.8 yard2 (9 m2).135 Another common behavior of urban- available.
dwelling species is their ability to tolerate extremely high Wildlife diversity is greater in this zone than in the high-
population densities, densities that may be caused by density zone. Most non-native generalists that were
inhibited dispersal, 120 or because of widespread present in the inner city are still present here. However,
availability of building debris and rubbish that provides with increased vegetation and patch size, many native
abundant shelter and protection from predators and species appear in this zone. Instead of only ecological
weather.61, 62 Newton147 remarked that adequate forage and generalists of the inner city, now seed- (grass-) eaters, and
protection from predation might permit urban animals to a few insectivorous species are present, especially in older
use habitats that would be inferior under natural neighborhoods.203
conditions. Small mammals in this zone escape extinction
or genetic attrition because habitat patches are probably Birds. House sparrows and starlings often reach
close enough to allow limited colonization and gene flow.60 maximum densities in this zone. 123 Starlings occur
However, disruption of these small habitat patches will wherever cover ranges between 35 and 85%, especially
likely affect these small species more than larger species near lawns. Rock dove densities are considerably less in
because their small home ranges keep them isolated in this zone than in the high-density zone, but they may occur
one patch, unlike larger species that move more frequently in such high densities that action is required to reduce
between patches.61 their numbers. One solution is to design buildings without
Squirrels are common inhabitants of parks or older holes, cavities, or ledges, and to reduce them on existing
neighborhoods where there is an open canopy composed structures. Improved methods for food waste disposal and
of large deciduous trees. In urban parks, squirrels take regulation of feeders will also help in their control.123
advantage of human handouts.95 Raccoons and opossums With more snags,7 a small amount of woody debris, and
may be seen in this zone if there is some greenspace or increased ground cover, many new species can be found in
corridor (e.g., underground sewer or stormwater channel) this zone compared to the high-density zone.134 Primary
that allows them safe travel to and from the city. cavity nesters begin to appear in this zone, as well as an
Big brown bats often form maternity colonies inside increased variety of secondary cavity nesters. Northern
structures such as houses or churches in downtown urban flickers might be seen feeding on ants, and swallows on
areas.10 Brazilian free-tailed bats, which occur primarily insects shortly after dawn or before dusk. With high
in southwestern Oregon, nest under tile roofs and densities of house sparrows and starlings (both cavity
hibernate inside buildings.135 Bats often use bridges in this nesters) and a limited number of snags, there is often
high-density zone, if the structure contains appropriate intense competition between these aggressive exotics and
crevices. Hoary bats and little brown bats also have been native cavity nesters.108, 109
observed foraging around artificial lights and roosting Lawns are commonly found in this zone. Flock-feeding
under bridges in this zone. Bat habitat often can be species like American robins, Brewer’s blackbirds, and
enhanced in urban areas by ensuring that bridges are starlings may be seen foraging on lawns anytime during
designed to accommodate bats. For more information, visit the day. Robins are common in this zone—lawns with
Bat Conservation International’s bridge web page at http:/ shade trees seem to provide ideal habitat.143 In addition,
/www.batcon.org/bridge/ambatsbridges/index.html. watering of lawns mimics precipitation, causing
earthworms to rise to the soil surface. 143 Older
Amphibians and Reptiles. Amphibians generally are neighborhoods in this zone, with expansive lawns and
absent from high-density urban areas because of the lack large older-age shade trees, typically support more
of wetlands. According to Stebbins and Cohen189 and insectivores and more shrub and tree branch nesters than
Campbell,36 modification of habitat (i.e., wetland loss) is do younger neighborhoods.56, 127 The occasional presence
probably the chief threat of urbanization to reptiles or of towhees, juncos, and sparrows are evidence of a more
amphibians restricted to water for breeding, egg laying, dense and diverse understory compared to the high-
and development.97, 140 No native amphibians or reptiles density zone. Mourning doves may appear if there is
are listed as either “closely associated” or “generally enough woody vegetation for them to nest in and some
associated” within the high-density zone.154 Only 1 exotic unmowed lawns or grassy edges to forage in.
reptile—the red-eared slider—is listed in this zone. It is
often inappropriately released in urban ponds or lakes.
 CHAPTER 12: URBAN ENVIRONS 327

With the many individual human residences or lots in of larger mammalian omnivores recorded in this zone.
this zone, most remnant patches of native habitat are Eastern gray squirrels, raccoons, skunks, and opossums
isolated, and the amount of edge is often at a maximum. (west of the Cascades) can be quite abundant. Higher
This increased edge now attracts brown-headed cowbirds densities of these mammals seem to occur because their
(nest parasites) and more nest predators, such as jays, home ranges are smaller in urban areas than in more rural
crows, skunks, and raccoons. These nest parasites and areas. This social behavioral change has been noted in
predators, operating near the edge of an urban area, may raccoons,6, 39, 98, 174 whose densities in urban areas may be
have enough impact on birds of small habitat patches to twice as high as in comparable rural areas. Raccoons are
create population sinks (specific areas of population versatile and take advantage of urban areas; they use
declines) where local reproduction is insufficient to offset sewers and human-made structures for travel while
adult mortality.166 Additionally, cat predation is high in this searching for human-supplied foods. Buildings and
zone and may negatively impact bird populations to the culverts provide additional den sites when natural dens
point where these urban residential zones might be are unavailable in urban areas. A successful omnivore, the
considered ecological traps.41, 90, 106 raccoon is known to eat >46 different species of plants,
Crows are human tolerant and omnivorous, and do insects, crustaceans, birds, bird eggs, and several small
well in all zones: >656 different food items have been mammal species (including rabbits and Norway rats),103
identified from stomach samples of the American crow.160 and can easily open trash containers.
Bird feeders in this zone are common. Many seed-eating Another dietary generalist taking advantage of the
birds are attracted to these feeders in summer and in medium-density environments is the opossum. A native
winter, when snow may be on the ground and food is of the southeastern U.S., it was introduced in Oregon in
scarce. Jays may be present, along with white-crowned the early twentieth century, and now has spread to all areas
sparrows, dark-eyed juncos, and evening grosbeaks. The west of the Cascades in both Oregon and Washington, with
house finch, a new closely associated species of this zone, a few being recorded east of the Cascades.135 One was
often monopolizes sunflower seed feeders. Sharp-shinned reported as far east as Spokane, Washington, in 1993.112
and Cooper’s hawks may be frequent visitors, preying on The coyote is present in all medium- and low-density
the concentrated passerines, especially in the winter.71 urban zones throughout the U.S..5, 129 Although coyotes
Anna’s hummingbirds seem to take advantage of the seem to prefer relatively undisturbed habitats in urban
increased urban temperatures and the abundance of environments,159 they are able to adapt and survive in the
artificial feeders in this zone;218 they can now be found in highly modified medium-density zone. Coyotes, and other
Portland and Seattle in midwinter. native carnivores, may even help reduce the abundance
Because of the increased prey base found in this zone, of house cats and other small mammalian carnivores that
several raptor species may be observed. Red-tailed hawks, prey on songbirds, thereby indirectly contributing to the
kestrels, and great-horned owls all tolerate human maintenance of a native avifauna. 158, 186 Ecological
disturbance, allowing them to exploit the available small- specialists such as shrews, moles, and mice may persist in
mammal prey base.28, 48 Red-tailed hawks and great- small urban patches, if human disturbance is limited and
horned owls are generalist predators that seem (if enough appropriate habitat components are retained.61 However,
habitat is available)to benefit from forest fragmentation shrews and voles often are absent or rare in urban areas,
and urbanization.28, 47 Screech owls, also tolerant of people, perhaps because their diurnal behavior may inhibit
show up in this zone in forested parks or older residential movements across open spaces.217 Perhaps this is why only
neighborhoods that have large deciduous trees.87, 89 one species, the montane vole, is listed as potentially being
Kestrels are able to survive in this zone as well, using the present in this zone.154 The Townsend’s mole may be found
extensive array of telephone wires as hunting perches,48 in lawns, moist meadows, and fields.
and will breed, if cavities are available for nesting. With the presence of larger and older trees, more insects,
In this zone, collisions with human-made objects were and more water, there is a corresponding increase in the
the greatest indirect cause of avian deaths, accounting for number of bat species. Several additional Myotis species
approximately 32% of deaths.8 The Cornell Laboratory of (Yuma, long-eared, little brown, and the solitary silver-
Ornithology’s Feeder Watch Program suggested that haired bat) may occupy this medium-density zone, as well
approximately 100 million birds are killed annually by as the big brown and Brazilian free-tailed bats that are
striking windows in urban residential areas in the U.S.70 present in the high-density zone. Bat boxes in this zone
may help to supplement natural roosts and cavities. Refer
Mammals. With less impervious surface in the medium-
to Bat Conservation International’s bat box project for
density zone, more earth is available for vegetative growth
more information: http://www.batcon.org.
and for the digging and burrowing of small mammals,
such as moles (Townsend’s coast, and broad-footed), Amphibians and Reptiles. Some remnant wetlands
rabbits, and gophers. In western Oregon and Washington, persist in this zone. In some cases, reptiles and amphibians
a predator of these small mammals is the red fox, a dietary are facilitated by the human infrastructure; snakes, frogs,145
generalist that seems to have adjusted well to human and salamanders100 may travel in sewers, mains, and pipes.
development. Reservoirs and artificial ponds can be beneficial for turtles
More available habitat, larger patch size, or a and toads.145 Salamanders and turtles—both native and
combination of the two is evidenced by the wider array non-native—may be found in this zone. Reflecting the
 328 Wildlife–Habitat Relationships in Oregon and Washington

increased small-mammal prey base, snakes are more

plentiful, with garter snakes being the most likely to be
seen. The many roads and widespread use of culverts in
this zone has had significant detrimental impacts on
amphibians and reptile.162 The ubiquitous introduced
bullfrog has taken over many small ponds and lakes in
this zone. It requires warm-water ponds, marshes, and
backwaters for breeding. The Pacific tree frog may be
heard in this zone if small ponds, seasonal pools,
temporarily filled depressions, or slow streams are present
for it to breed in.
The many raccoons in this zone may negatively impact
reptiles and amphibians;12 great blue herons also help
Figure 4. Low-density urban zone. Benton County, Oregon.
regulate their numbers in this zone. The collecting of
(Photograph by Kelly A. Bettinger)
herpetofauna by humans can severely impact small
isolated populations. The impact of cat and dog predation
on reptile and amphibian populations in this zone is English house sparrow populations decrease sharply
largely unknown and needs to be investigated. when total vegetation cover reaches 60%.123 This is the zone
where the ground-nesting guilds of birds begin to
Wildlife in the Low-density Zone approach predevelopment numbers, not only for foraging,
This zone most closely approaches predevelopment but for breeding as well. Thrushes and towhees are
conditions of the native flora (Figure 4). As expected, abundant in many areas. In dormant agricultural fields
species diversity (but not density) also approaches or active hay fields, savannah, vesper, and song sparrows
predevelopment levels. A total of 264 wildlife species may be seen and heard, with juncos, golden-crowned, and
potentially occur within this low-density zone (Table 4), white-crowned sparrows living on the edge of these fields
with 15 species classified as closely associated, 145 as or in the bordering hedgerows.
generally associated, and another 104 species as potentially Bird feeders are quite common, and bird variety seen
present. These numbers are derived from a list of all at these feeders can be much greater than in the medium-
species that may occur in any low-density zone anywhere density zone: purple finches, pine siskins, black-capped
in either Oregon or Washington; each species will only be or chestnut-sided chickadees, red- and white-breasted
present if and when the appropriate habitat is available.154 nuthatches, and even red-winged blackbirds may appear,
The actual number of species will vary according to the especially in the winter. At nectar feeders all of the local
native background matrix, be it forest, shrub-steppe, or hummingbirds are likely to occur, with some, like Anna’s,
grassland, more than in either of the other two urban even over-wintering.218 Nest boxes are quite common in
zones. Most of the wildlife species found in this zone are this zone, and help supplement natural cavities, attracting
native species (Table 3). western and mountain bluebirds,101 swallows, chickadees,
nuthatches, wrens, flickers, and even pygmy and screech
Birds. Numbers of avian specialists, including
owls.
insectivores, may approximate those in undisturbed areas.
At dawn and dusk, swallows (violet-green, tree, cliff,
Swallows, woodpeckers, warblers, and chickadees are
bank, and barn) take to the air. Near the coast, purple
numerous. House sparrows may be uncommon, and
martins may be present. In wetland areas with dense
instead, chickadees, nuthatches, flickers, robins, juncos,
shrubs, diversity will be high, and birds present may
and native sparrows are the most common birds observed.
include common yellowthroats, Wilson’s, orange-
Few rock doves are present except around farms or
crowned, and sometimes yellow warblers, great blue
granaries. A more natural ground or forb layer is apparent
herons, killdeers, osprey, and rails. Song sparrows occur
by the presence of quail and sparrows feeding. Thrushes,
in most brushy spots, whereas northern orioles favor large
indicators of nearly natural ground cover and shrub layers,
open-canopied deciduous trees, along with warbling and
may be widespread. Gray catbirds may be found in this
red-eyed vireos, black-throated gray warblers, and
zone on the eastside and indicate the presence of a well-
Hutton’s vireo.
developed dense understory as well. Western tanagers are
Brown-headed cowbird populations may be quite high
present where there is forest cover of large trees with a
in this zone because of abundant livestock, and high nest
full canopy.168 Large trees in this zone also account for the
parasitism may occur. However, nest predation may be
presence of brown creepers and golden-crowned kinglets.
less common in this zone than in other zones, probably
With the retention of snags, most of the woodpecker
because of better habitat for the host species, more dense
species occur in this zone, including white-headed and
cover, and less edge.181, 212 In this zone, ravens begin to
Lewis’s woodpeckers. Because of the presence of these
outnumber crows, and in the winter, large groups of
primary cavity nesters in this zone, new cavities are
ravens may be seen together at road-kills, along with the
created each year, promoting use by secondary cavity
occasional bald eagle.
nesters, e.g., chickadees, nuthatches, bluebirds, and wrens.
 CHAPTER 12: URBAN ENVIRONS 329

With the native prey base at near natural levels, so too diversity is consistently rated “poor” when watershed
will be the predator populations. Hawks and owls are imperviousness exceeds 10-15%.176 Species that employ
present and occur in this zone for foraging as well as for feeding strategies such as filtering organic matter or
nesting. Northern harriers may also occur here, especially preying on insects are especially sensitive to changes
in and around agricultural or hay fields. triggered by increases in impervious surfaces. Fish
diversity also drops sharply at this level of impervious
Mammals. Most mammal species present before urban
surface.27, 176 Abundance and spawning success of sensitive
growth occurred will be present in this low-density zone,
species are also affected at >10% impervious surface. Trout
except large carnivores. All of the large ungulates occur
and other salmonids are particularly susceptible to
in this zone, including deer, elk, and even moose (e.g.,
changes triggered by increases in impervious surfaces.176
Spokane, Washington). Deer are often most abundant in
Even with complete retention of streamside buffers, basins
this zone because of the mix of agricultural fields and
with >10% impervious cover show measurable levels of
reduced hunting pressures. Because of high prey densities,
stream degradation.190
cougars may occur in this zone.
Mustelids recorded in this zone include the long-tailed Wildlife in the Urban Fringe Areas
weasel and mink. The deer mouse is now the most The urban fringe area may be defined as the region of
common mouse, replacing the house mouse. Raccoons, transition between the rural area and the low-density
skunks and opossums are still present but in lower, more urban zone.30, 171 Nilon and VanDruff148 characterize the
natural densities than in the inner zones. Now, with the urban fringe as being predominately vacant with
presence of downed wood, chipmunks and native agricultural lands dominated by large amounts of growing
squirrels occur. Rarely will the non-native Eastern gray space. Only higher use of rural roads may distinguish this
squirrels be found in this zone, outside of parks and golf area from the neighboring rural zone. Unlike the rural
courses. Instead, Douglas and red squirrels become more zone, neither agriculture nor the timber industry is the
common. Ground squirrels may be visible along the road primary employer, and as a result, the use and
banks. Perhaps, reflecting the amount of soil available to management of these lands is dissimilar to the intensely
diggers in this zone compared to the two other zones, 2 farmed fields or harvested woodlots of the rural areas. In
shrews, 2 voles, and 3 moles may occur. Pocket gophers this urban fringe, compared to rural areas, hedgerows are
are regionally common. often more lush, fields more often idle, pastures
With the increase in prey, coyotes also become unoccupied, timber rotations longer, and tree harvests
increasingly common in this zone. More large areas of more selective. In the urban fringe, farmland left idle
native vegetation are now present that can be used as cover before its urban development may temporarily improve
by coyotes, who travel in and around the scattered homes the forage base available to small mammals, which can
and search for food.158, 159 Black bears may be found subsequently support raptors and other small mammal
foraging. Populations of bobcats will be present unless predators.48, 148 Most important, it is this area that is in
pets are overabundant or overgrazing is prevalent. imminent danger of being developed and subdivided.
With more wetlands, muskrats, beavers, and mink may Land in this fringe area is less fragmented and often
occur. Bats depend on these small wetlands for food and less expensive than in any of the three urban zones.
water, especially the mosquito loving little brown and Protection and conservation of wildlife habitat can often
Yuma myotis bats. Preservation of maternity and be achieved at a lower cost, and often with less impact to
hibernacula sites is a high management priority for these the citizens of the region. Typically many large tracts of
animals. private and public land remain that can be identified and
Amphibians and Reptiles. Instead of seeing only the preserved as important wildlife corridors or core areas.
ubiquitous tree frog or the aggressive non-native bullfrog, Although planners and biologists often overlook the urban
both the spotted frog or western toad may be found in fringe area, it may offer the greatest opportunities for
wet areas. Most of the expected native salamanders and conservation and landscape-level management.
lizards may also be found in this zone in appropriate
habitats. Toads may be seen in and around the many lawns Key Urban Planning Issues
of residential areas where landscaping and irrigation
practices provide adequate food and shelter for survival. Growth Management
They rely heavily on Hymenoptera found in the lawns.12 Throughout the U.S., urban growth and development is
Within the urban environs, this is the zone most suited one of the largest threats to wildlife diversity. Oregon and
for amphibians and reptiles; Richter162 pointed out that Washington are two of only about a dozen states with
areas with <15% impervious surfaces are best. Concern comprehensive statewide “growth management”
here should be to limit runoff, maintain large buffers legislation. Oregon’s Comprehensive Land Use Planning
around wetlands, stabilize water level fluctuations of Program has its roots in laws passed in 1973. Oregon
wetlands, and protect shorelines where spawning occurs, measures its land use planning successes by attempting
particularly northwestern shorelines that are preferred. to meet 19 statewide planning goals. Washington began
Faunal diversity declines sharply in streams in basins taking a proactive approach to managing growth with the
with >10-15% impervious surface. Macroinvertebrate passage of the Growth Management Act in 1990. Growth
 330 Wildlife–Habitat Relationships in Oregon and Washington

management has changed local planning in many ways,

and provides opportunities for managing fish and wildlife
resources.
Controlling Urban Sprawl. Growth management requires
local jurisdictions to control urban sprawl. Urban sprawl
uses land inefficiently, is typically damaging to natural
resources, and is expensive to provide for public services.
Urban sprawl has been defined as:
Unplanned, uncontrolled, and uncoordinated single-
use development that does not provide for an attractive
and functional mix of uses and/or is not functionally
related to the surrounding land use and which
variously appears as low density, ribbon or strip,
scattered, leapfrog, or isolated development.146
Oregon and Washington both address controlling urban
sprawl in their statewide goals under their growth Figure 5. Interim urban growth boundaries for Spokane
management legislation. The intent of Oregon’s Statewide County, Washington, 1998.
Planning Goal 14 is “To provide for an orderly and efficient
transition from rural to urban land use”. In Washington, County that includes a Wildlife Area Combining Zone. It
two of the planning goals are to “encourage development applies to areas identified in Deschutes County’s
in urban areas where adequate public facilities and comprehensive plan as deer winter range, significant elk
services exist or can be provided in an efficient manner habitat, antelope range, or deer migration corridor. The
“and to “reduce the inappropriate conversion of wildlife area zone functions as an overlay zone that
undeveloped land into sprawling, low-density changes the permitted uses in the underlying zone. For
development”. Both states attempt to do this by requiring example, golf courses and dude ranches, while permitted
local jurisdictions to identify Urban Growth Boundaries conditionally in the rural residential zone, are not
(Oregon) and Urban Growth Areas (Washington). These permitted in the wildlife area zone. The zoning code also
are essentially the same thing, an administrative boundary includes an overlay zone that protects sensitive birds,
that marks where urban development ends and rural areas mammals, and wildlife areas like great blue heron
begin. Urban Growth Boundaries/Areas (UGB/A) are rookeries and Townsend’s big-eared bat sites.
included as part of a local jurisdiction’s comprehensive Washington’s Growth Management Act (GMA)
plan and are designated using population growth requires all local jurisdictions to identify and protect
projections (Figure 5). For the purposes of the Urban critical areas, and identify natural resource lands. Critical
Wildlife Manager or Urban Planner, the UGB/A marks areas include wetlands, aquifer recharge areas, fish and
where the high- and medium-density zones end and the wildlife habitat conservation areas, frequently flooded
low-density zone begins. areas, and geologically hazardous areas. The GMA directs
the State Department of Community Trade and Economic
Protecting Fish and Wildlife Habitat. Before the passage Development (DCTED) to develop minimum guidelines
of growth management legislation, fish and wildlife for identifying critical areas. Local jurisdictions must then
habitat was given a backseat when local jurisdictions identify their critical areas at least to the minimum
planned for various developments. Notable exceptions to standards identified by DCTED. Then they must draft and
this were habitats for species protected under the Federal implement regulations that protect critical areas. These
Endangered Species Act, and wetland resources protected protective regulations are often referred to as Critical Areas
under the Federal Clean Water Act. Oregon and Ordinances (CAOs).
Washington now require local jurisdictions to consider and Spokane County’s CAO provides additional protection
protect habitat areas for non-listed species as well as for wetlands, geologically hazardous areas, and fish and
special habitat areas including riparian areas. wildlife habitat conservation areas. The county has
Oregon’s Statewide Planning Goal 5 seeks “to protect adopted the Washington Department of Fish and
natural resources and conserve scenic and historic areas Wildlife’s (WDFW) Priority Habitats and Species (PHS)
and open spaces.” Local governments are required to go program to designate fish and wildlife habitat
through a process in which they inventory and evaluate conservation areas. 205 The PHS program includes
Goal 5 resources, which include riparian corridors, distribution maps of species and habitats considered to
wetlands, and other wildlife habitat. After the inventory be priorities for conservation and management. When a
and evaluation process, the local jurisdiction, in proposed development occurs within a priority habitat
consultation with the appropriate state agencies, must or within a certain distance of a priority species nest site,
decide whether to protect the resource, fully allow the developer is required to submit a Habitat Management
conflicting uses, or limit conflicting uses in some fashion. Plan that documents how the development will avoid and
An example from Oregon is the Zoning Code in Deschutes mitigate the habitat or species in question. Mitigation
 CHAPTER 12: URBAN ENVIRONS 331

measures can include establishing buffer areas, clustering Purchase of the land or easement rights, either outright,
development and preserving open space, seasonal partial, or fee simple is often the most realistic way to
restriction of construction activities, and preservation of protect valuable wildlife habitat. This can be achieved
critically important trees and plants. Habitat Management through outright acquisition, or with conservation
Plans are reviewed by WDFW. Spokane County grants easements, which allow the landowner to retain
final approval. ownership of the land while prohibiting certain activities
that would be harmful to wildlife. Easements can be
Planning for Open Space. Growth management
written in many different ways. Activities that are typically
legislation in Washington and Oregon requires local
prohibited in conservation easements include building,
jurisdictions to consider and plan for open space.
grazing, and clearing of vegetation. Public access can often
Historically, policies discussing the need for open space
be written into an easement but is not a requirement. Local
for recreation and wildlife habitat were included in local
governments often seek the help of land trusts or
comprehensive plans. However, these policy statements
organizations with land trust skills.
were typically not implemented. Planning for open space
is required in both Oregon and Washington. In Oregon, Clustering is often used in planned developments to
“open space” is included as one of the Goal 5 resources integrate open space and/or wildlife habitat. Buildings
that must be inventoried and evaluated. Washington’s or homes are grouped, or clustered, in a small area of a
GMA requires that local jurisdictions planning under the development while the remaining land is retained in
act include a Land Use Element in their comprehensive common ownership. Local jurisdictions occasionally allow
plan. Among others, “open space” is identified as one of for density bonuses when developers use this technique.
the land uses to be designated and described. There may
be some disadvantages to open space systems. They can Open Space, Greenways, and Corridors
be expensive to develop and local tax revenues (or One of the most politically acceptable ways to preserve
potential tax revenues) may be reduced. However, the and protect wildlife habitat in and around urban areas is
benefits derived from open space by both wildlife and through an open space program. Greenspaces, open
humans normally greatly outweigh any disadvantages. spaces, wildlife corridors, natural parks, and trails—all of
these terms refer to parcels of open land scattered
Land-use Planning Tools. Usually, growth management throughout a metropolitan region. Greenspace not only
makes use of existing land-use planning tools like benefits wildlife but is also recognized by the public as
comprehensive plans and development regulations. providing important aesthetic and recreational benefits to
However, under growth management legislation, policies the community. As a result, communities are often very
identified in comprehensive plans are required to be supportive of greenspaces and open spaces. As Ludwig128
consistent with the ordinances that implement them. noted, properly planned open spaces help reduce the
Wildlife habitat goals can be achieved using many of the isolation and fragmentation of habitat patches,
planning tools currently in use by local jurisdictions—both communities, and ecosystems. They also help to ensure
regulatory and voluntary. Traditional land-use planning and enhance the movement and dispersal of animals,
involves the use of different land-use zones. Zoning which promotes genetic exchange and population
regulations dictate permissible uses in the zones and stability.128
building densities.
Selection and Design Basics. When defining the vision
Transfer of Development Rights programs allow a and priorities for a green or open space program, it is
developer or landowner to transfer their “right” to develop important to look at the structure and use of the
property from an area where development is not wanted surrounding landscape and how each natural area fits
to a more appropriate area. These areas are often referred within the region as a whole. Several authors have
to as “sending” and “receiving” zones. contributed guidelines for the design of open spaces.58, 59,
214
Current Use Taxation programs give landowners the Noss 149 admitted that in the absence of detailed
ability to reduce the amount of property tax they pay on a autecological, species-specific information, some empirical
parcel if they agree to keep it in open space or wildlife generalizations for reserve design stand out. The following
habitat. Local jurisdictions can develop rating systems is a compendium of ideas from Thomas et al.,195 Noss,149
where a landowner would receive a greater tax break for and Noss and Cooperrider:150
more valuable wildlife habitat, e.g., riparian areas or 1. Maintain large, intact patches of native vegetation by
wetlands. preventing fragmentation of those patches by
development. Remember that blocks of habitat close
Educational Programs can protect or enhance urban together are better than blocks far apart.
wildlife habitat through voluntary efforts on the part of 2. Maintain connections among wildlife habitats by
the landowner. The National Wildlife Federation’s identifying and protecting corridors for movement,
Backyard Wildlife Habitat Program provides information remembering that habitat in contiguous blocks is
for urban homeowners on bird feeding, landscaping for better than fragmented blocks and that
wildlife, and other small projects that enhance habitat. The interconnected blocks are better than isolated blocks.
WDFW’s Backyard Wildlife Sanctuary Program provides Focus on the idea of connectivity.
similar information specific to the Pacific Northwest.
 332 Wildlife–Habitat Relationships in Oregon and Washington

3. Minimize edge. Round patches are better than linear Unfragmented aquatic systems with large (>656 feet
patches. [>200 m]) buffers will contain many wildlife species
4. If possible, establish priorities for species protection typically found in these habitats in non-urbanized
and protect habitats that constrain the distribution watersheds, such as kingfishers, eagles, waterfowl,
and abundance of those species. beavers, salmon, otters, and amphibians.
5. Protect rare landscape features or ecological hotspots.
Guide development toward areas of landscape Direct Impacts
containing “common” widespread features. Direct impacts of urbanization on water systems include
6. Maintain significant ecological processes in protected (1) loss or reduction of riparian habitat or buffers, (2) loss
areas. Provide enough land for processes to continue. of woody debris and other instream structures, (3)
7. Contribute to the regional persistence of rare species degradation or loss of stream channels and wetlands, (4)
by protecting some of their habitat locally. reduction in water quality, (5) habitat fragmentation, and
8. Balance the opportunity for recreation by the public (6) introduction of non-native species and pets. Many of
with the habitat needs of wildlife. Remember open these direct impacts will result in secondary changes in
spaces that are roadless or otherwise inaccessible to the aquatic environment.
humans are better than roaded and accessible habitat Loss or Reduction of Riparian Habitat and Buffers. In
blocks for wildlife. Washington, an estimated 50-90% of the riparian habitat
Open space selection should also be done on a has been lost or modified.119 Remnant riparian habitat may
landscape-level scale. An open space system should be provide the only remaining connectivity, refuges, and
planned on a regional, multi-county scale, ensuring reserves of habitat within an urban area. The function and
connection of high-quality habitats with all adjoining value of urban riparian zones can vary widely depending
counties and protection of these habitats throughout the on their width and connectivity to habitat areas outside
region. Take advantage of all lands, state or federal, already the urban zone.
protected, incorporating them into the open space system. Urbanization often results in a direct loss of riparian
A tool to help design and carry out an open space plan is zones and aquatic buffers as the natural vegetation is either
the Geographical Information System (GIS). Most states trimmed or removed and converted to impervious
have already completed a GAP analysis—a GIS-based surfaces and human structures. Bank stabilization
mapping inventory of land ownership and protection activities also contribute to the loss of aquatic buffers. The
assessment.38, 177 This analysis and the associated data can loss of riparian vegetation results in a direct reduction in
be used for a more detailed analysis required for a habitat for terrestrial wildlife species. Removal of riparian
localized open space plan. vegetation, as well as alteration of organic inputs and
hydrologic regimes by expanding urbanization, generally
Riparian, Wetland, and Stormwater Systems results in increased erosion, increased algal production,
Urbanization affects the natural drainage systems of an changes to temperature regimes, and reduced
area, its streams, wetlands, and associated riparian areas concentrations of dissolved oxygen.209 Sedimentation may
and buffers. This section is broken into three main parts: be increased as well as nutrients tied to sedimentation.
direct impacts, indirect impacts, and management. Smaller buffers result in less filtering of pollutants, such
Stormwater control structures are included in the as artificial fertilizers and pesticides from runoff, resulting
management section, as these systems are the primary in increased nutrients and other pollutants in the aquatic
tools available to mitigate the impacts of urbanization on system.119
aquatic and riparian systems. In some cases, zoning and
local ordinances, for example, those specifying buffers, can Loss of Woody Debris and Other Instream Structures.
also be instrumental in protecting local water resources. Typically, large woody debris on the ground and standing
Urbanization results in two basic types of impacts on snags and other large trees are removed. Hazard tree
drainage systems—direct and indirect. Direct impacts removal and reduction of the riparian zone directly
include dredging, straightening, bank stabilization, removes potential large woody debris from both the buffer
piping, filling, buffer removal and reduction, and zone and from the aquatic system; this results in the loss
fragmentation or isolation from other components of the of pools, riffles, backwaters, and off-channel habitats. In
landscape. Indirect impacts result when the pervious to addition, large woody debris is removed directly from
impervious surface ratios of the watershed are altered. As streams and rivers for navigation enhancement and flood
little as 10% impervious surface in a watershed can change control.
the hydroperiod of the natural drainage systems.26, 27, 176 Degradation or Loss of Stream Channels and Wetlands.
Altered hydroperiods result in habitat changes that affect Direct impacts include channelization, dredging, filling,
the fish and wildlife in these systems. Impervious cover bank stabilization, culverts, and piping, which may all
has especially important impacts on storm surges and result in stream degradation. Wetlands are frequently filled
concentrations of pollutants in streams.192, 209 The direct and in urban areas, resulting in habitat losses and further
indirect impacts are interrelated and rarely occur alteration of the water regime in a basin. Rivers and
separately. This distinction forms a useful way to frame streams are dredged and straightened, which removes and
the discussion. alters the stream bottom conditions and habitats.
 CHAPTER 12: URBAN ENVIRONS 333

Straightening results in greater stream velocities and Urbanization alters the volume, velocity, duration, and
increased stream channel instability. Streams may also be frequency of runoff events that can alter the structural
buried completely in pipes or bisected by roads, affecting integrity of aquatic systems that may not be otherwise
the connectivity of the whole system. Bank stabilization, impacted by urban developments.
loss of riparian buffers, dredging, channelization, and
Channel Instability and Runoff. Increased runoff in
filling can all increase the amount of fine sediments present
urban areas results in additional soil erosion, stream
in a stream system. Fine sediments can smother bottom
channel cutting, and water quality deterioration. Channel
invertebrates and salmon spawning gravels.
instability triggers streambank erosion and habitat
Many of our regulatory systems assume that small
degradation. Channel instability also expresses itself in
wetlands are expendable. However, most natural wetlands
the loss of instream habitat structures such as pools and
are small and are vital to maintaining ecological
riffles, large woody debris, and overhead cover.27, 119
diversity.178 Richter and Azous163 found no correlation
Channel cutting and increased erosion can result in
between wetland size and amphibian diversity. The
channels becoming straightened, with meanders, back
authors concluded that the loss of most small wetlands
channels, and overflow channels being eliminated or cut
could severely impede source-sink processes and place
off from the main channel. As channel scouring increases
remaining wetlands at increased probabilities of local
there is loss of substrates needed for salmonid spawning
amphibian population extinction.
and aquatic invertebrates. Stream channel erosion
Reduction in Water Quality. Impervious surfaces collect downstream of culverts (plunge pools) can create barriers
pollutants that wash off during storm events and may be to fish passage where they did not exist when the
deposited in large pulses directly into aquatic systems structures were installed. Sedimentation and deposition
unless there is some sort of filtration or other water quality upstream of culverts similarly impacts aquatic fauna by
control structure present. Improperly applied herbicides burying spawning gravel beds and destroying pools and
and pesticides may directly affect riparian and aquatic riffles.
flora and fauna. Reduction of riparian buffers reduces the
Water Table Alterations. More rapid runoff of stormwater
capacity of the natural system to filter out these pollutants
results in decreased infiltration and groundwater recharge.
and nutrients and exacerbates the problem. The size of
Wetlands and streams may transform to drier, more
the pollutant load is directly related to the level of
intermittent, or seasonally flooded conditions, impacting
impervious surfaces in the watershed.119 Fertilizers applied
native plant and animal species adapted to a longer wet
to lawns, gardens, golf courses, and other urban
season. Increased runoff may also result in wetlands
landscaping are carried to surface waters in stormwater
holding more water for a shorter period than under
runoff. Land clearing, which exposes soil, can result in
undisturbed conditions. These changes in the water
high levels of phosphorus in stormwater runoff. These
regime with urbanization alter the vegetative community
excess nutrients can enhance the growth of invasive
of wetland areas.6, 45, 75, 105
aquatic plant species. Decomposition of excess plant
growth can rob the water of dissolved oxygen, further Water Level Fluctuations. Increased runoff changes the
reducing its suitability for aquatic fauna. natural pattern of water level fluctuations. In the greater
Runoff from parking lots and other impervious surfaces Seattle area, water level fluctuations >7.9 inches (20 cm)
tends to be warmer, which will increase the overall are commonly seen in basins with >50% total urbanization.
temperature of the receiving waters regardless of buffer These alterations in the natural pattern of water level
size. Combined with the loss of trees that would have fluctuations are correlated with decreased amphibian
provided shade, temperatures in impervious surface areas species diversity. Amphibian species that attach their egg
may be 10-12oF higher than in the forests and fields that masses to vegetation are extremely vulnerable to water
they replace.176 Increased summer temperatures in urban level changes. Some researchers believe that such species
streams are directly related to the level of impervious will not be able to survive in basins with moderate to high
surfaces found in the basin. Lack of riparian cover and levels of urbanization.6, 162, 163
stormwater detention ponds that warm water before
releasing it into streams and wetlands can amplify the Management
observed increases in water temperatures. Management of urban riparian and aquatic systems
should take a three-pronged approach. First, manage the
Indirect Impacts amount of impervious surfaces and vegetation clearing
With urbanization, runoff rates increase, resulting in throughout the basin. Second, work to keep the riparian
significant increases in the frequency and the duration and systems as natural as possible through maintenance of
frequency of flood events. Changes in runoff rates and connectivity and adequate buffer widths. Third, manage
volumes may be more severe in western Washington and stormwater runoff to control the impacts of altered
Oregon, where stormwater is normally stored and carried hydroperiods.
off with subsurface flows, than in eastern towns where The amount of impervious surface in a basin is closely
stormwater in more typically carried across the land linked to the health of the aquatic systems in the basin.
surface. Peak discharges increase and urbanized basins Whereas maintaining <10% impervious surface is not a
may experience entirely new peak runoff events. 25 realistic goal for most urban areas, urban managers should
 334 Wildlife–Habitat Relationships in Oregon and Washington

try to minimize impervious surfaces as much as possible although commonly attempted, have had limited success.
through such techniques as landscaping requirements, Of 40 wetland mitigation projects monitored in King
clearing restrictions, use of pervious surfacing materials, County, 11 were not installed and 79% of the rest were
and under-structure parking. Basin-wide clearing unsuccessful by current performance standards. Only 1
restrictions that control both the amount and the timing site successfully replaced the functions of the impacted
of clearing can significantly mitigate the impacts to water watershed.141
quality and hydroperiod. Management of stormwater runoff in urban basins is
Maintaining connectivity and adequate buffer widths critical to maintaining the functions of natural wetland
will help riparian systems retain natural functions and and riparian systems. Stormwater detention systems must
values. Connectivity should include continuity between be big enough to accommodate the increased runoff that
instream structures, the connectivity of the riparian is generated from urban basins. They also need to be
corridor throughout the urban area, and the connectivity designed to moderate the changes in velocity, volume,
of wetlands to protected upland habitats as well as other duration, and frequency of discharges to natural systems.
aquatic systems. Landscape-level planning should account Stormwater systems in urban areas should also be
for these habitats before development. Strategies might designed to filter pollutants and oils and to remove excess
include using bridges rather than culverts, daylighting sediments from stormwater. The ideal stormwater control
impacted stream corridors, targeted acquisitions and system would also provide some method to moderate
incentive programs for riparian protection, and restrictions discharge temperatures and dissolved oxygen levels.
on development in riparian areas. Although McGuckin Stormwater detention facilities should not be confused
and Brown138 suggested integrating municipal stormwater with natural wetland systems. Stormwater detention
management facilities into urban greenways, this is not a ponds that are functioning properly will catch and
good solution. These stormwater detention ponds experience the severe water level fluctuations generated
experience extreme water level fluctuations and in urbanized basins. If the stormwater system is catching
consequently become amphibian mortality sinks.6, 163 To sediments properly, it will need to be dredged on a regular
minimize this effect: (1) separate stormwater detention basis, which will prevent more than herbaceous vegetation
ponds from natural areas, (2) do not allow vegetation with from becoming established. Stormwater basin
stem diameters of 0.12-0.20 inches (3-5 mm) to grow in maintenance schedules should be timed to have the least
detention ponds, (3) construct sides of ponds to be steep impact on wildlife species that may be using the limited
to minimize areas of appropriate water depth for breeding, habitats provided by the facility. Stormwater control
and (4) construct curbing or fencing around the ponds to structures are not intended to provide or replace wildlife
discourage use by amphibians.6, 162, 163, 164 habitat; they are intended to protect existing wildlife
Typical regulatory buffer widths of 50-100 feet (15.2- habitat from the impacts generated by urban development.
30.5 m) are designed for maintenance of water quality.119
Vegetated buffers of 75 feet (23 m) do a good job at filtering Urban Parks, Golf Courses,
pollutants and nutrients from stormwater runoff and Cemeteries, and Open Space
moderately good job of temperature regulation.119 Buffers Urban parks and open spaces are rapidly assuming a
100 feet (30.5 m), however, do a poor job of providing central role in the protection of native wildlife from urban-
wildlife habitat, large woody debris recruitment, sediment related disturbances.42 In addition, the popularity of golf
filtration, or maintenance of natural hydroperiods.119 has increased greatly in the past 20 years resulting in many
Buffers should also be wide enough to allow for natural new and expanded golf courses.144 Many of these are at
channel migration. the interface between expanding population centers and
Most urban buffers will benefit from active vegetation diminishing rural lands142 (Argyle et al., Greater Vancouver
management. Noxious and invasive weed species should Regional District, Development Services Department,
be removed or controlled. Management may include long- Burnaby, British Columbia, Canada, unpublished data).
term replacement of non-native vegetation with natives Cemeteries increase as the urban population increases. In
to lessen erosion or impact on wildlife. A strong many urban settings, parks, golf courses, and cemeteries
component of conifer trees should be maintained to often contain the last remaining patches of native
provide the best source for long-lasting large woody debris vegetation. Unfortunately, not all wildlife species can exist
in both the aquatic and terrestrial portions of the system. within these small islands. The diversity of species present
“Hazard” tree removal should be carefully monitored. in these areas is generally lower than in the original
Where trees are determined to be hazards, every attempt habitat.60, 92, 184, 186 Little information is available concerning
should be made to shorten them to a safe height while the effects of urbanization on small mammals and the
still maintaining a standing wood component. Portions ability of parks to maintain community structure in urban
of “hazard” trees that are removed should be placed either areas.1, 148 Dickman and Doncaster61 found that intensively
in the aquatic or the terrestrial portion of the system. Large managed and maintained patches of habitat, like parks
trees of any type should be maintained in the buffers for and golf courses, contained fewer small mammals than
bank stabilization and temperature regulation. expected. However, with design changes and different
The primary emphasis should be on protection of management approaches, golf courses, parks, and
existing natural systems. Replacement and restoration, cemeteries can be significantly improved in order to
provide habitat for many native wildlife species.
 CHAPTER 12: URBAN ENVIRONS 335

The following factors are important in determining the of the same flowering plants that attract hummingbirds
richness, diversity, and abundance of bird species in an will also attract butterflies.57, 121 These gardens will generate
urban park setting:42 (1) size of area, (2) amount of edge, much public interest and can serve as education points.
(3) presence of natural vegetation, (4) amount of shrub Wildlife use, especially by wading birds, waterfowl,
cover, (5) average canopy height and canopy cover, (6) and shorebirds, can be greatly enhanced by the presence
density of adjacent buildings, (7) presence of water, and of a pond or lake. Especially important for shorebirds is
(8) availability of snags. the practice of timing drawdowns with the spring and
When designing these areas and during construction, fall migration flights of these birds. During migration,
it is important to minimize the impact on the natural these birds search for food in areas of shallow water (<7
vegetation of the area. Large spatial scale or landscape- inches deep [17.8 cm]), especially where there are exposed
level planning is essential. Try to protect the largest habitat mudflats. Drawdowns should occur in spring roughly
patches possible, especially those containing landscape- from late March through early June, while fall migration
level habitat elements like rivers, lakes, and woodlots. peaks during August and September.92 Other management
Riparian corridors should always remain undisturbed and considerations include: (1) winter flooding protects the
protected with appropriate buffers. Attempt to make golf invertebrate eggs and larvae (food for birds), (2) slow
course fairways as narrow as possible leaving undisturbed drawdowns maintain soil moisture and encourage
native vegetation on the sides for “roughs”. Route the establishment of vegetation, and (3) working vegetation
course through existing openings.92 In urban parks, leave into the soil releases nutrients for invertebrates. Drawing
as much native vegetation as possible around the down the water level gradually will continually expose
perimeter. Better yet, design in connecting corridors to new areas of saturated soil and keep a steady supply of
larger native patches offsite, even if they are small. If only invertebrates available to birds.92 To protect wildlife
small native patches are available, an array of small nesting and roosting in these aquatic environments, good
patches can potentially serve the function similar to a management practices dictate that trails and roads should
single, large patch, by providing the same habitat not be built completely around these natural features.
components. The key is that they be close together and Continual disturbance by humans and their pets will chase
connected with similar habitat. Consider clustering of away all but the most tolerant wildlife from the area.
facilities, buildings, roads, and parking lots; a similar Snags are used by >85 North American bird species,
technique is being used in residential development. In including many owls, woodpeckers, chickadees,
addition, position these areas of disturbance in low-quality swallows, and other songbirds as places to find food,
habitat patches, leaving high-quality patches undisturbed. watch for prey, build homes, and attract mates.193 Snags
Pesticide and insecticide use in these areas can be are so valuable to wildlife that their protection needs to
greatly reduced with the adoption of Integrated Pest be an integral part of any forest habitat. Snags to preserve
Management (IPM) practices. IPM is a management should be >6inches (15.25 cm) in diameter—the larger the
strategy for dealing with insects, weeds, and plant diseases snag the more opportunities for nesting and foraging.
that establishes acceptable levels of infestation or damage. Snags can be made more attractive by trimming back the
Treatment only occurs if these levels are exceeded. IPM branches; however, it is best to leave 1-3 feet (0.3-0.9 m)
treatment uses cultural practices and biological controls on each limb for perching. If a snag is close enough to be
that start with the least harmful to the environment and a threat to a building or structure, it can be topped to a
progresses to harmful chemicals only as a last resort.92 height to eliminate the threat. Snags are more beneficial
Where pesticide levels are low, birds become a natural ally when left in clumps of >3. Recommended snag densities
for managers, helping to maintain insect levels below are 3-16/acre (7.4-39.5/ha). Eventually a snag will rot to
outbreak levels. the point that it will fall. Let fallen snags lie, they are still
Another factor that can contribute to increased wildlife valuable to wildlife and the environment.
use, while at the same time fit into IPM practices, is the Gavareski85 emphasized the importance of size and
use and propagation of native plants rather than exotics. vegetative condition of parks for attracting and sustaining
Exotic plants typically require more water, are less resistant wildlife; the larger the park and the more natural the
to insect infestations, and are often sterile, providing no vegetation (fewer exotics, natural understory), the more
pollen or seeds for wildlife. By using natives, the escape wildlife species will be found. Cicero42 also pointed out
and spread of exotics are also minimized. The removal of the importance of buffers in isolating species from the
native shrubs and ground covers in parklands alters the negative influences of urbanization, and in providing
nesting habits of birds.32 Altering the native forest in Seattle avenues of faunal interchange among ponds and
through planting of formal gardens, or clearing natural peripheral habitats. The number of trails should be limited
brush and reducing park size, were associated with in these areas and judiciously placed away from sensitive
decreased overall bird diversity, fewer regularly occurring breeding or roosting areas.
species, and a greater proportion of species typically
associated with urban environments.85
Specialized habitats like butterfly or hummingbird
gardens can be established even in the smallest patches of
habitat in our parks, golf courses, and cemeteries. Many
 336 Wildlife–Habitat Relationships in Oregon and Washington

Managing Urban Habitat for Wildlife Research Needs

There are many questions that need to be answered for
Backyard Wildlife in Residential Areas urban areas of the Northwest. How has growth
The key to attracting wildlife is providing proper habitat. management affected wildlife? Are urban
Generally, this entails increasing the abundance and metapopulations sources or sinks, and for what types of
variety of native vegetation on your property. The basic animals and why? Do urban areas serve as sources of
requirement for supporting wildlife in your backyard is exotic plants and animals, spreading these species to rural
to provide food, water, shelter, and places to protect young. areas? How effective are wildlife corridors in sustaining
To determine management activities that can either target populations? How can they be designed to be more
enhance, or in some cases, discourage or repel a particular effective? How does clustered housing impact wildlife?
species, consult the management activity links matrix.154 How do wildlife communities (e.g., birds, small mammals,
Using this matrix, you can determine the habitat elements bats) change with increasing urbanization? What impact
that you can affect. Or when planning, this database can does dispersed settlement (urban fringe) have on wildlife?
be used to predict potential losses and gains of species Can we accurately predict the impact that future
when different land use or management actions are urbanization will have on wildlife; can we adequately
proposed. assess different alternatives?
Research is needed to determine the impact that
Food. Imitate nature. Provide a variety of grasses, forbs,
fragmentation has on wildlife in urban areas of the Pacific
shrubs, and trees that provide fruits, nuts, and seeds
Northwest. Is fragmentation different in urban areas
through the year. Plant a variety of annuals and perennials,
because of exotics, size of patches, composition of urban
and evergreen and deciduous trees of varying height. Use
zones? How does edge impact wildlife in the urban area?
plants that naturally occur in your area and are not
Does it increase brood parasitism and nest predation? Are
introduced by humans. Native plants are adapted to your
cowbird populations increasing in urban areas of the
local environment and the local wildlife species have
Northwest?
evolved along with these plants. Benefits include less
What is the best measure of urbanization? Is the “urban
maintenance, less water, and less disease, with better
gradient” concept the best and most appropriate paradigm
survival over time and food that wildlife can use. Keep
to use when conducting research?137 How far from the
lawns to a minimum.
urban center do the impacts of urbanization extend into
Water. A year-round supply of water will help to initially the exurban area? Quantify the rural-urban exchange
attract and retain wildlife in your local area. The source of (through corridors). Determine the ease through which
water can vary from a pie plate to a stream, a birdbath, or wildlife can travel in urban areas. How far from the urban
a pond. centers do the positive and negative effects of development
extend into surrounding wildlands? How does the amount
Shelter. Wildlife need a place to rest, escape from enemies,
of juxtaposition of varying types of settlement affect birds?
and seek shelter from inclement weather. Evergreen trees
Determine the foraging difference along the urban rural
and shrubs or thorny bushes add year-round protective
gradient for different bird species and guilds.
cover from predators. Deciduous shrubs offer effective
See Marzluff et al.134 for a thorough discussion on avian
summer cover. Rock, log, slash, and mulch piles offer other
research needs in an urban environment. They recommend
cover. Clumping your plantings will help provide cover.
that research should share some of the following features:
Vines on a fence or wall or snag can provide other forms
(1) long-term investigation, (2) use of rigorous
of shelter. Don’t cut down dead trees; leave them standing,
experimental design allowing simultaneous comparison
and when they do fall, let them lie. Not only do they
of population viability along several points of the urban-
provide shelter, but food and places to raise young as well.
wildland continuum, (3) identification of mechanisms
Work with your neighbors to provide as large as possible
responsible for the effects of urbanization on birds (all
parcels of native habitat.
wildlife), and (4) quantification of avian (all wildlife)
Places to Raise Young. Snags provide places to raise demography.
young for birds, snakes, small mammals, and bats. Rock,
log, and slash piles provide nursery space for rabbits, Literature Cited
snakes, birds, mice, and shrews. A variety of heights and 1. Adams, L. W., and L. E. Dove. 1989. Wildlife reserves and corridors in
density of vegetation provides the diversity required for the urban environment. National Institute of Urban Wildlife, Columbia,
MD.
diverse wildlife. Wetlands and ponds provide proper
2. Airola, T. M., and K. Buchholz. 1984. Species structure and soil
nesting habitat for frogs, salamanders, dragonflies, and characteristics of five urban sites along the New Jersey Palisades.
other insects. Urban Ecology 8: 149-164.
For more information, join WDFW’s Backyard Wildlife 3. Alcock, I., and P. Warsop. 1982. Diet, distribution and habitat
Program: http://www.wa.gov/wdfw/wlm/ preferences of stoats and weasels in Sheffield. Sorby Record 20: 5-10.
byw_prog.htm, Oregon’s Naturescaping Program: http:/ 4. Andre, H. 1994. Effects of habitat fragmentation on birds and mammals
in landscapes with different proportions of suitable habitat: a review.
/ w w w. d f w. s t a t e . o r. u s / O D F W h t m l / E d u c a t i o n /
Oikos 71: 355-366.
Naturescaping.html, or the National Wildlife Federation’s 5. Atkinson, K.T., and D. M. Shackleton. 1991. Coyote, Canis latrans,
Backyard Wildlife Habitat Program: http:// ecology in a rural-urban environment. Canadian Field-Naturalist 105:
www.nwf.org/. 49-54.
 CHAPTER 12: URBAN ENVIRONS 337

6. Azous, A. L., and K. O. Richter. 1995. Amphibian and plant community 29. Brooks, J. E. 1973. A review of commensal rodents and their control.
responses to changing hydrology in urban wetlands. Pages 156-162 in E. Critical Reviews in Environmental Control 3: 405-453.
Robichaud, editor. Puget Sound Research ?95 Proceedings. Puget Sound 30. Bryant, C. 1982. The rural real estate market. Department of
Water Quality Authority, Olympia, WA. Geography Publication Series No. 18, University of Waterloo, Ontario,
7. Balda, R. P. 1975. The relationships of secondary cavity-nesters to snag Canada.
densities in western coniferous forests. U.S. Forest Service Wildlife 31. Burke, D. M., and E. Nol. 1998. Influence of food abundance, nest-site
Habitat Technical Bulletin 1, Albuquerque, NM. habitat, and forest fragmentation on breeding ovenbirds. The Auk 115:
8. Banks, R. C. 1979. Human-related mortality of birds in the United 96-104.
States. U.S. Fish and Wildlife Service Special Scientific Report 215. 32. Burr, R. M., and R. E. Jones. 1968. Influence of parkland habitat
9. Barber, J. C., and M. M. Barber. 1983. Prey of an urban peregrine falcon. management on birds in Delaware. Transactions North American
Maryland Birdlife 39: 108-110. Wildlife and Natural Resources Conference 33: 299-306.
10. Barbour, R. W., and W. H. Davis. 1969. Bats of America. University 33. Busnel, R. G., and John Fletcher, editors. 1978. Effects of noise on
Press of Kentucky, Lexington, KY. wildlife. Academic Press, New York, NY.
11. Barclay, R. M. R. 1984. Observations on the migration, ecology, and 34. Butcher, G. S., W. A. Niering, W. J. Barry, and R. H. Goodwin. 1981.
behavior of bats at Delta Marsh, Manitoba. Canadian Field-Naturalist Equilibrium biogeography and the size of nature preserves; an avian
98: 331-336. case study. Oecologia (Berlin) 49: 29-37.
12. Barrentine, C. D. 1991. Food habits of western toads (Bufo boreas 35. Calhoon, R. E., and C. Haspel. 1989. Urban cat populations compared
halophilus) foraging from a residential lawn. Herpetological Review 22: by season, subhabitat and supplemental feeding. Journal of Animal
84-87. Ecology 58: 321-328.
13. Batten, L. 1972. Breeding birds species diversity in relation to 36. Campbell, C. A. 1973. Survival of reptiles and amphibians in urban
increasing urbanization. Bird Study 19: 157-166. environments. Pages 61-66 in J. Noyes and D. Progulske, editors. Wildlife
14. Batten, L. 1974. Blackbird boom in suburbia. Wildlife 16: 274-277. in an urbanizing environment. The Wildlife Society, Springfield, MA.
15. Beck, A. M. 1974. The ecology of urban dogs. Pages 57-59 in J. Noyes 37. Canning, D. J., and M. Stevens. 1989. Wetlands of Washington: a
and D. Progulske, editors. Wildlife in an urbanizing environment. The resource characterization. Environmental 2010 Project, Washington
Wildlife Society, Bethesda, MD. State Department of Ecology, Olympia, WA.
16. Beissinger, S. R. 1978. Avian community organization and habitat 38. Cassidy, K. M., M. R. Smith, C. E. Grue, K. M. Dvornich, J. E. Cassady, K.
structure in a residential area and a virgin beech-maple forest. M.S. R. McAllister, and R.E. Johnson. 1997. Gap analysis of Washington State:
Thesis. Miami University, Oxford, OH. an evaluation of the protection of biodiversity.Volume 5 in K. M.
17. Beissinger, S. R. and D. R. Osborne. 1982. Effects of urbanization on Cassidy, C. E. Grue, M. R. Smith, and K. M. Dvornich, editors.
avian community organization. Condor 84: 75-83. Washington State Gap Analysis—Final Report. Washington
18. Blair, R. B. 1996. Land use and avian species diversity along an urban Cooperative Fish and Wildlife Research Unit, University of Washington,
gradient. Ecological Applications 6: 506-519. Seattle, WA.
19. Blondel, J. 1985. Mediterranean bird faunas in the light of anthropic 39. Cauley, D. L. 1974. Urban habitat requirements of four wildlife species.
pressures since the Neolithic. Pages 594-607 in V. D. Ilyichev and V.M. Pages 143-147 in J. Noyes and D. Progulske, editors. Wildlife in an
Gavrilov, editors. Acta 18th International Ornithological Congress, urbanizing environment. The Wildlife Society, Springfield, MA.
Moscow, Russia. 40. Christian, D. P. 1975.Vulnerability of meadow voles, Microtus
20. Boersma, P. D., and K. N. Almasi. 1991. Can birds tell the difference pennsylvanicus, to predation by domestic cats.American Midland
between native and introduced conifers in the arboretum? Washington Naturalist 93: 498-502.
Park Arboretum Bulletin 54: 7-9. 41. Churcher, P. B., and J. H. Lawton. 1987. Predation by domestic cats in
21. Bolger, D. T. 1999. Fragmentation effects on birds in southern an English village. Journal of Zoology 212: 439-455.
California: contrast to the paradigm. Abstract from 69th Annual 42. Cicero, C. 1989. Avian community structure in a large urban park:
Meeting of the Cooper Ornithological Society, March 29-April 3, controls of local richness and diversity. Landscape Urban Planning 17:
Portland, OR. 221-240.
22. Bogumile, O. 1986. Breeding ecology of goshawk in forest-agricultural 43. Clergeau, P., J. L. Savard, G. Mennechez, and G. Falardeau. 1998. Bird
habitats of central Poland. Pages 7-9 in H. Ouellet, editor. Proceedings abundance and diversity along an urban-rural gradient: a comparative
of XIX Congressus Internationalis Ornithologici, Ottawa, Canada. study between two cities on different continents. The Condor 100:
23. Bolen, E. G. 1991. Analogs: a concept for the research and 413-425.
management of urban wildlife. Landscape and Urban Planning 20: 285- 44. Coleman, J. S., S. A. Temple, and S. R. Craven. 1997. Cats and wildlife: a
289. conservation dilemma. Extension Publications, Madison, WI.
24. Bolsinger, C. L. 1973. Changes in commercial forest area in Oregon 45. Cooke, S. S., and A. L. Azous. 1993. Effects of urban stormwater runoff
and Washington, 1945-70. U.S. Forest Service Resource Bulletin PNW- on palustrine wetland vegetation. A Report to the U.S. Environmental
46, Portland, OR. Protection Agency, Region 10. Seattle, WA.
25. Booth, D. B. 1991. Urbanization and the natural drainage system - 46. Cousins, S. H. 1982. Species size distributions of birds and snails in an
impacts, solutions, and prognoses. Northwest Environmental Journal 7: urban area. Pages 99-109 in R. Bornkamm, J. A. Lee, and M. R. D.
93-118. Seaward, editors. Urban Ecology. Blackwell Scientific Publications,
26. Booth, D. B. and C. R. Jackson. 1994. Urbanization of aquatic systems- Oxford, England.
degradation thresholds and limits of mitigation. Pages 425-434 in R. A. 47. Craighead, J. J., and F. C. Craighead, Jr. 1956. Hawks, owls and wildlife.
Marston and V. R. Hasfurther, editors. Proceedings of symposium on Stackpole Publishing, Harrisburg, PA.
effects of human induced changes on hydrologic systems. American 48. Cringan, A. T., and G. C. Horak. 1989. Effects of urbanization on
Water Resources Association, Jackson Hole, WY. raptors in the western United States. Pages 219-228 in Proceedings of
27. Booth, D. B. and L. E. Reinelt. 1993. Consequences of urbanization on western raptor management symposium and workshop. National
aquatic systems – measured effects, degradation thresholds, and Wildlife Federation, Washington, D.C.
corrective strategies. Pages 545-550 in Watershed 93, Proceedings of a 49. Cronon, W. 1983. Changes in the land. Hill and Wang, New York, NY.
national conference on watershed management, Alexandria, VA. 50. Dansereau, P., and G. Par. 1977. Ecological grading and classification of
28. Bosakowski, T., and D. G. Smith. 1997. Distribution and species land-occupation and land-use mosaics. Geography Paper No. 58.
richness of a forest raptor community in relation to urbanization. Fisheries and Environment Canada, Ottawa, Ontario, Canada.
Journal of Raptor Research 31: 26-33.
 338 Wildlife–Habitat Relationships in Oregon and Washington

51. DeGraaf, R. M. 1978. Avian communities and habitat associations in 76. Falk, J. H. 1976. Energetics of a suburban lawn ecosystem. Ecology 57:
cities and suburbs. Pages 7-24 in C. M. Kirkpatrick, editor. Proceedings 141-150.
of the John S. Wright Forestry Conference, Wildlife and People. Purdue 77. Fenton, M. B., H. G. Merriam, and G. L. Holryod. 1983. Bats of
University, West Lafayette, IN. Kootenay, Glacier, and Mount Revelstoke national parks in Canada:
52. DeGraaf, R. M. 1985. Residential forest structure in urban and identification by echolocation calls, distribution, and biology. Canadian
suburban environments: some wildlife implications in New England, Journal of Zoology 61: 2503-2508.
USA. Journal of Arboriculture 11: 236-241. 78. Fitzgerald, B. M. 1988. Diet of domestic cats and their impact on prey
53. DeGraaf, R. M. 1986. Urban bird habitat relationships: application to populations. Pages 123-147 in D. C. Turner, and P. Bateson, editors. The
landscape design. Transactions North American Wildlife and Natural domestic cat: the biology of its behaviour. Cambridge University Press,
Resources Conference 51: 232-248. Cambridge, England.
54. DeGraaf, R. M., H. R. Pywell, and J. W. Thomas. 1975. Relationships 79. Fletcher, J. L. 1990. Review of noise and terrestrial species: 1983-1988.
between nest heights, vegetation, and housing density in New England Pages 181-188 in B. Berglund and T. Lindvall, editors. Noise as a public
suburbs. Transactions of Northeast Fish and Wildlife Conference 32: health problem.Volume 5: new advances in noise research part II.
130-150. Swedish Council For Building Research, Stockholm, Sweden.
55. DeGraaf, R. M. and J. W. Thomas. 1976. Wildlife habitat in or near 80. Forman, R. T. 1995. Land mosaics: the ecology of landscapes and
human settlements. Pages 54-62 in J. W. Anderson, editor. Trees and regions. Cambridge University Press, Cambridge, England.
forests for human settlements. Proceedings of Pl.05-00 Symposia in 81. Fox, R. 1987. Population images. United Nations Fund for Population
Vancouver, BC, Canada. Activities.
56. DeGraaf, R. M. and J. M. Wentworth. 1981. Urban bird communities 82. Frankie, G. W., and L. E. Ehler. 1978. Ecology of insects in urban
and habitats in New England. Transactions North American Wildlife and environments. Annual Review Entomology 23: 367-387.
Natural Resources Conference 46: 396-413. 83. Frey, H. T. 1984. Expansion of urban area in the United States: 1960-
57. Dennis, J. V, and M. Tekulsky. 1991. How to attract hummingbirds and 1980. U.S. Department of Agriculture Economic Research Service Staff
butterflies. Ortho Books, San Ramon, CA. Report AGES830615, Washington, D.C.
58. Diamond, J. M. 1976. Island biogeography and conservation: strategy 84. Furlonger, C. L., H. J. Dewar, and M. B. Fenton. 1987. Habitat use by
and limitations. Science 193: 1027-1029. foraging insectivorous bats. Canadian Journal Zoology 65: 284-288.
59. Diamond, J. M. and R. M. May. 1976. Island biogeography and the design 85. Gavareski, C. A. 1976. Relation of park size and vegetation to urban
of natural reserves. Pages 163-186 in R. M. May, editor. Theoretical bird populations in Seattle, WA. The Condor 78: 375-382.
ecology: principles and applications. W. B. Saunders, Philadelphia, PA. 86. Geggie, J. F., and M. B. Fenton. 1985. A comparison of foraging by
60. Dickman, C. R. 1987. Habitat fragmentation and vertebrate species Eptesicus fuscus (Chiropter:Vespertilionidae) in urban and rural
richness in an urban environment. Journal of Applied Ecology 24: 337- environments. Canadian Journal Zoology 63: 263-266.
351. 87. Gehlbach, F. 1986. Odd couples of suburbia. Natural History 95: 56-
61. Dickman, C. R. and C. P. Doncaster. 1987. The ecology of small 66.
mammals in urban habitats. Volume I. Populations in a patchy 88. Gehlbach, F. 1988. Population and environmental features that
environment. Journal of Animal Ecology 56: 629-640. promote adaptation to urban ecosystems: the case of eastern screech-
62. Dickman, C. R. and C. P. Doncaster. 1989. The ecology of small owls (Otus asio) in Texas. Pages 1809-1813 in H. Ouellet, editor.
mammals in urban habitats.Volume II. Demography and dispersal. Proceedings of XIX Congressus Internationalis Ornithologici, Ottawa,
Journal of Animal Ecology 58: 119-127. Canada.
63. Dietrich, J. 1980. Der habicht (Accipiter gentilis) im stadtverband 89. Geis, A. D. 1974. The new town bird quadrille. Natural History 83: 53-
Saarbrucken.Verhandlungen der Gesellschaft für Ökologie. 10: 301-305. 61.
64. Dietrich, J. and H. Ellenberg. 1981. Aspects of goshawk urban ecology. 90. George, W. G. 1974. Domestic cats as predators and factors in winter
Pages 163-175 in R. E. Kenward and I. M. Lindsay, editors. Understanding shortages of raptor prey. Wilson Bulletin 86: 384-396.
the goshawk. International Association of Falconry and Conservation of 91. Gibbs, J. P. 1993. Importance of small wetlands for the persistence of
Birds of Prey, Oxford, England. local populations of wetland-associated animals. Wetlands 13: 25-31.
65. Doncaster, C. P. 1985. The spatial organization of urban foxes (Vulpes 92. Gillihan, S. W. 2000. Bird conservation on golf courses: a design and
vulpes) in Oxford. Unpublished Thesis, University of Oxford, England. management manual. Sleeping Bear Press, Chelsea, MI.
66. Dorney, R. S. 1985. Urban wildlife populations: a look at uptown, 93. Goldstein, E. L., M. Gross, and R. M. DeGraaf. 1986. Breeding birds and
downtown, and suburban residents. In Wildlife survivors in the human vegetation: a quantitative assessment. Urban Ecology 9: 377-385.
niche. National Zoological Park, Smithsonian Institute, Washington, D.C. 94. Green, R. J. 1984. Native and exotic birds in a suburban habitat.
67. Dorney, R. S. 1986. Bringing wildlife back to the cities. Technology Australian Wildlife Research 11: 181-190.
Review October: 48-54. 95. Guth, R. W. 1979. The junk food guild: birds and mammals on picnic
68. Dorney, R. S. and P. W. McClellan. 1984. The urban ecosystem: its grounds and in residential areas. Illinois Audubon Bulletin 189: 3-7.
spatial structure, its scale relationships, and its subsystems attributes. 96. Guthrie, D. A. 1974. Suburban bird populations in southern California.
Environments 16: 9-20. American Midland Naturalist 92: 461-466.
69. Duckworth, F. S., and J. S. Sandberg. 1954. The effect of cities upon 97. Hall, R. J. 1980. Effects of environmental contaminants on reptiles: a
horizontal and vertical temperature gradients. Bulletin American review. U.S. Fish and Wildlife Service, Washington, D.C., Special
Meteorological Society 35: 198-207. Scientific Report—Wildlife 228: 1-12.
70. Dunn, E. H. 1993. Bird mortality from striking residential windows in 98. Harris, S. 1977. Distribution, habitat utilization and age structure of a
winter. Journal of Field Ornithology 64: 02-309. suburban fox (Vulpes vulpes) population. Mammal Review 7: 25-39.
71. Dunn, E. H. and D. L. Tessaglia. 1994. Predation of birds at feeders in 99. Haub, C., and M. M. Kent. 1989. 1989 world population data sheet.
winter. Journal of Field Ornithology 65: 8-16. Population Reference Bureau, Washington, D.C.
72. Ehrenfeld, D. W. 1970. Biological conservation. Holt, Rienhart and 100. Heck, O. A. 1971. Eastern tiger salamanders found in a window well.
Winston, New York, NY. Engelhardtia 4: 35.
73. Emlen, J. T. 1974. An urban bird community of Tucson, Arizona: 101. Herlugson, C. J. 1978. Comments on the status and distribution of
derivation, structure, and regulation. The Condor 76: 184-197. western and mountain bluebirds in Washington. Western Birds 9: 21-32.
74. Erz, W. 1966. Ecological principles in the urbanization of birds. Ostrich 102. Hobbs, E. 1988. Using ordination to analyze the composition and
Supplement 6: 357-363. structure of urban forest islands. Forest Ecology and Management 23:
75. Ewing, K. 1996. Tolerance of four wetland plant species to flooding 139-158.
and sediment deposition. Environmental and Experimental Biology 36:
131-146.
 CHAPTER 12: URBAN ENVIRONS 339

103. Hoffmann, C. O., and J. L. Gottschang. 1977. Numbers, distribution 126. Leedy, D. L. and L. W. Adams. 1986. Wildlife in urban and developing
and movements of a raccoon population in a suburban residential areas: an overview and historical perspective. Pages 8-20 in: K. Stenberg,
community. Journal of Mammalogy 58: 623-636. and W. W. Shaw, editors. Wildlife conservation and new residential
104. Hooper, R. G., E. F. Smith, H. S. Crawford, B. S. McGinnes, and V. J. developments, Tucson, AZ.
Walker. 1975. Nesting bird populations in a new town. Wildlife Society 127. Lucid,V. J. 1974. Bird utilization of habitat in residential areas. M.S.
Bulletin 3: 111-118. Thesis.Virginia Polytechnic Institute, Blacksburg, VA.
105. Houck, C. A. 1996. The distribution and abundance of invasive plant 128. Ludwig, D. R. 1995. Assessment and management of wildlife diversity
species in freshwater wetlands of the Puget Sound lowlands, King in an urban setting. Natural Areas Journal 15: 353-361.
County, Washington. M.S. Thesis. University of Washington, Seattle, WA. 129. MacCracken, J. G. 1982. Coyote food in a southern California
106. Hubbs, E. L. 1951. Food habits of feral house cats in the Sacramento suburb. Wildlife Society Bulletin 10: 80-281.
Valley. California Fish and Game 37: 177-189. 130. MacLean, C. D., and C. L. Bolsinger. 1997. Urban expansion in the
107. Huff, F. A. 1977. Effects of the urban environment on heavy rainfall forests of the Puget Sound region. U.S. Forest Service, Resource
distribution. Water Resources Bulletin 13: 807-816. Bulletin PNW-RB-225. Portland, OR.
108. Ingold, D. J. 1996. Delayed nesting decreased reproductive success in 131. Martin, T. E. 1988. Habitat and area effects on forest bird
northern flickers: implications for competition with European starlings. assemblages: is nest predation an influence? Ecology 69: 74-84.
Journal of Field Ornithology 67: 321-326. 132. Martin, T. E. 1993. Nest predation and nest sites. Bioscience 43: 523-
109. Ingold, D. J. and R. J. Densmore. 1992. Competition between 532.
European starlings and native woodpeckers for nest cavities in Ohio. 133. Marzluff, J. M. 1997. Effects of urbanization and recreation on
Sialia 14: 43-48. songbirds. Pages 89-102 in W. M. Block, and D. M. Finch, editors.
110. James, P. C., A. R. Smith, L. W. Oliphant, and L. G. Warkentin. 1987. Songbird ecology in southwestern ponderosa pine forests: a literature
Northward expansion of the wintering range of Richardson’s merlin. review. U.S. Forest Service General Technical Report RM-292, Fort
Journal of Field Ornithology 58: 112-117. Collins, CO.
111. Johnsen, A. M., and L. W. VanDruff. 1987. Summer and winter 134. Marzluff, J. M., F. R. Gehlbach, and D. A. Manuwal. 1998. Urban
distribution of introduced bird species and native bird species richness environments: influences on avifauna and challenges for the avian
within a complex urban environment. Pages 123-127 in L. W. Adams and conservationist. Pages 283-299 in: J. M. Marzluff, and R. Salabanks,
D. L. Leedy, editors. Integrating man and nature in the metropolitan editors. Avian conservation, research, and management. Island Press,
environment. Proceedings of a National Symposium on Urban Wildlife, Washington, D.C.
Chevy Chase, MD. 135. Maser, C. 1998. Mammals of the Pacific Northwest: from the coast
112. Johnson, R. E., and K. M. Cassidy. 1997. Terrestrial mammals of to the high Cascades. Oregon State University Press, Corvallis, OR.
Washington State: location data and predicted distributions. Volume 3 136. Mason, P. 1985. The impact of urban development on bird
in: K. M. Cassidy, C. E. Grue, M. R. Smith, and K. M. Dvornich, editors. communities of three Victorian towns, Lilydale, Coldstream, and Mr.
Washington State Gap Analysis—Final Report. Washington Evelyn, Victoria, Australia. Corella 9: 14-21.
Cooperative Fish and Wildlife Research Unit, University of Washington, 137. McDonnell, M. J., and S. T. A. Pickett. 1990. Ecosystem structure and
Seattle, WA. function along urban-rural gradients: an unexploited opportunity for
113. Jordheim, S. 1966. Recollection of a peregrine falcon-pigeon pursuit. ecology. Ecology 7: 1232-1237.
Blue Jay 24: 20. 138. McGuckin, C., and R. Brown. 1995. Stormwater management and
114. Jureck, R. M. 1994. A bibliography of feral, stray, and free-ranging urban greenways. Landscape and Urban Planning 33: 227-246.
domestic cats in relation to wildlife conservation. California 139. Mesure, M. 1996. Bird collisions. World Wildlife Fund, Canada,
Department of Fish and Game, Wildlife Management Division, Ontario, Canada.
Sacramento, CA. 140. Minton, S. A., Jr. 1968. The fate of amphibians and reptiles in a
115. Klein, M. L. 1993. Waterbird behavioral responses to human suburban area. Journal of Herpetology 2: 113-116.
disturbances. Wildlife Society Bulletin 21: 31-39. 141. Mockler, A. 1998. Results of monitoring King County wetland
116. Klem, D., Jr. 1989. Bird-window collisions. Wilson Bulletin 101: mitigations. King County Department of development and
606-620. environmental sciences, Renton, WA.
117. Klem, D., Jr. 1990. Collisions between birds and windows: mortality 142. Moore, K. E. 1990. Urbanization in the lower Fraser Valley, 1980-
and prevention. Journal Field Ornithology 61: 120-128. 1987.Technical Report 120. Canadian Wildlife Service, Delta, BC,
118. Knight, R. L., and K. J. Gutzwiller. 1995. Wildlife and recreationists: Canada.
coexistence through management and research. Island Press, 143. Morneau, F., C. Lépine, R. Décarie, M.Villard, and F. DesGranges. 1995.
Washington, D.C. Reproduction of American robin (Turdus migratorius) in a suburban
119. Knutson, K. L., and V. L. Naef. 1997. Management recommendations environment. Landscape and Urban Planning 32: 55-62.
for Washington’s priority habitats: riparian. Washington Department of 144. Moul, I. E., and J. E. Elliott. 1994. The bird community found on golf
Fish and Wildlife, Olympia, WA. courses in British Columbia. Northwestern Naturalist 75: 88-96.
120. Krebs, C. J., M. S. Gaines, B. L. Keller, J. H. Myers, and R. H. Tamarin. 145. Neill, W. T. 1950. Reptiles and amphibians in urban areas of Georgia.
1973. Population cycles in small rodents. Science 179: 35-41. Herpetologica 6: 113-116.
121. Kruckeberg, A. 1982. Gardening with native plants of the Pacific 146. Nelson, A. C., and J. B. Duncan. 1995. Growth management principles
Northwest; an illustrated guide. University of Washington Press, Seattle, and practices. American Planning Association, Chicago, IL.
WA. 147. Newton, I. 1986. The sparrowhawk. T. and A. D. Poyser, Ltd., Calton,
122. Kurta, A., and J. A. Teramino. 1992. Bat community structure in an Staffordshire, England.
urban park. Ecography 15: 257-261. 148. Nilon, C. H., and L. W. VanDruff. 1987. Analysis of small mammal
123. Lancaster, R. K., and W. E. Rees. 1979. Bird communities and the community data and applications to management of urban greenspaces.
structure of urban habitats. Canadian Journal of Zoology 57: 2358- Pages 53-59 in L. W. Adams and D. L. Leedy, editors. Integrating man and
2368. nature in the metropolitan environment. National Institute Urban
124. Leedy, D. L. 1975. Highway-wildlife relationships, Volume 1. A state-of- Wildlife, Columbia, MD.
the-art report. U.S. Department of Transportation, Federal Highway 149. Noss, R. F. 1992. The Wildlands Project: land conservation strategy.
Administration; FHWA-RD-76-4. Washington, D.C. Wild Earth (Special Issue): 10-25.
125. Leedy, D. L. 1975. Highway-wildlife relationships, Volume 2. An 150. Noss, R. F. and A. Cooperrider. 1994. Saving nature’s legacy:
annotated bibliography. U.S. Department of Transportation, Federal protecting and restoring biodiversity. Defenders of Wildlife and Island
Highway Administration; FHWA-RD-76-5. Washington, D.C. Press, Washington, D.C.
 340 Wildlife–Habitat Relationships in Oregon and Washington

151. Nowakowski, E. 1982. Influence of urbanization on the structure of 173. Savard, J. P. and J. B. Falls. 1981. Influence of habitat structure on the
wireworm communities (Coleoptera, Elateridae). Pages 79-87 in M. nesting heights of birds in urban areas. Canadian Journal of Zoology 59:
Luniak and B. Pisarski, editors. Animals in urban environment. Polish 924-932.
Academy of Science, Institute of Zoology, Warsaw, Poland. 174. Schinner, J. R. 1969. Ecology and life history of the raccoon (Procyon
152. Obara, H., H. Hirata, and M. Okuzaki. 1977. An aspect of the lotor) within the Clifton suburb of Cincinnati. M.S.Thesis. University of
existence of mammals in urban ecosystems. Pages 173-187 in M. Cincinnati, Cincinnati, OH.
Numata, editor. Tokyo project: interdisciplinary studies of urban 175. Schlauch, F. L. 1975. Agonistic behavior in a suburban Long Island
ecosystems in the metropolis of Tokyo, Tokyo, Japan. population of the smooth green snake (Opheodrys vernalis). Engelhardtia
153. Oliphant, L. W., and S. McTaggart. 1977. Prey utilized by urban 6: 25-26.
Merlins. Canadian Field-Naturalist 91: 190-192. 176. Schueler, T. 1994. The importance of imperviousness. Watershed
154. O’Neil, T. A., D. H. Johnson, C. Barrett, M. Trevithick, K. A. Bettinger, C. Protection Techniques 1: 100-111.
Kiilsgaard, M.Vander Heyden, E. L. Greda, B. G. Marcot, P. J. Doran, L. 177. Scott, J. M., F. Davis, B. Csuti, R. Noss, B. Butterfield, C. Groves, H.
Wunder, and S. Tank. 2001. Matrixes for wildlife-habitat relationships in Anderson, S. Caicco, F. D’Erchia, T. C. Edwards Jr., J. Ulliman, and R.
Oregon and Washington. In: D. H. Johnson and T. A. O’Neil, managing Wright. 1993. Gap Analysis: a geographic approach to protection of
directors. Wildlife-habitat relationships in Oregon and Washington. biological diversity. Wildlife Monographs 123: 1-41.
Oregon State University Press, Corvallis, OR. 178. Semlitsch, R. D., and J. R. Bodie. 1998. Are small, isolated wetlands
155. Orser, P. N., and D. J. Shure. 1972. Effects of urbanization on the expendable? Conservation Biology 12: 5.
salamander Desmognathus fuscus fuscus. Ecology 53: 1148-1154. 179. Sharpe, D. M., F. Stearns, L. A. Leitner, and J. R. Dorney. 1986. Fate of
156. Paton, P. W. C. 1994. The effect of edge on avian nest success: how natural vegetation during urban development of rural landscapes in
strong is the evidence? Conservation Biology 8: 17-26. northeastern Wisconsin. Urban Ecology 9: 267-287.
157. Petraitis, P. S., R. E. Latham, and R. A. Niesenbaum. 1989. The 180. Skagen, S. K., R. L. Knight, and G. H. Orions. 1991. Human
maintenance of species diversity by disturbance. Quarterly Review of disturbance of an avian scavenging guild. Ecological Applications 1: 215-
Biology 64: 393-418. 225.
158. Quinn, T. 1997. Coyote (Canis latrans) food habits in three urban 181. Small, M. F., and M. L. Hunter. 1988. Forest fragmentation and avian
habitat types of western Washington. Northwest Science 71: 1-5. nest predation in forested landscapes. Oecologia 76: 62-64.
159. Quinn, T. 1997. Coyote (Canis latrans) habitat selection in urban areas 182. Smith, K. G. 1978. Range extension of the blue jay into Western
of western Washington via analysis of routine movements. Northwest North America. Bird Banding 49: 208-214.
Science 71: 289-297. 183. Sodhi, N. S., and L. W. Oliphant. 1993. Prey selection by urban-
160. Reaume, T. 1987. Selective feeding by the American crow. Ontario breeding merlins. The Auk 110: 727-735.
Birds 5: 71-72. 184. Soulé, M. 1987. Viable populations for conservation. Cambridge
161. Reijnen, R. 1995. Disturbance by car traffic as a threat to breeding University Press, Cambridge, England.
birds in the Netherlands. M.S. Thesis. DLO-Institute for Forestry and 185. Soulé, M. A. C. Alberts, and D. T. Bolger. 1991. The effects of habitat
Nature Research, Wageningen, The Netherlands. fragmentation on chaparral plants and vertebrates. Oikos 63: 39-47.
162. Richter, K. O. 1997. Criteria for the restoration and creation of 186. Soulé, M. D. T. Bolger, A. C. Alberts, J. Wright, M. Sorice, and S. Hill.
wetland habitats of lentic-breeding amphibians of the Pacific 1988. Reconstructed dynamics of rapid extinctions of chaparral-
Northwest. Pages 72-94 in K. B. MacDonald and F. Weinmann, editors. requiring birds in urban habitat islands. Conservation Biology 2: 75-92.
Wetland and riparian restoration: taking a broader view. U.S. 187. Southwood, T. R. E. 1961. The number of species of insects
Environmental Protection Agency, Publication 910-R-97-007, Region 10, associated with various trees. Journal of Animal Ecology 30: 1-8.
Seattle, WA. 188. Spencer, L. 1994. Fastest growing hobby in U.S. is for the birds.
163. Richter, K. O., and A. L. Azous. 1995. Amphibian occurrence and Seattle Times, January 10, 1994.
wetland characteristics in the Puget Sound Basin. Wetlands 15:305-312. 189. Stebbins, R. C., and N. W. Cohen. 1995. A natural history of
164. Richter, K. O., and A. L. Azous, S. S. Cooke, R. W. Wisseman, and R. R. amphibians. Princeton University Press, Princeton, NJ.
Horner. 1991. Effects of stormwater runoff on wetland zoology and 190. Steedman, R. J. 1988. Modification and assessment of an index of
wetlands soils characterization and analysis. Washington State biotic integrity to quantify stream quality in Southern Ontario.
Department of Ecology Report G0091019, Olympia, WA. Canadian Journal of Fisheries and Aquatic Sciences 45: 492-501.
165. Robbins, C. S. 1979. Effects of forest fragmentation on bird 191. Sukopp, H., H. Elvers, and H. Mattes. 1982. Studies in urban ecology
populations. Pages 198-212 in Management of northcentral and of Berlin (West). Pages 115-130 in: M. Luniak and B. Pisarski, editors.
northeastern forests for nongame birds. U.S. Forest Service General Animals in Urban Environment. Polish Academy of Science, Institute of
Technical Report NC-51, Minneapolis, MN. Zoology, Warsaw, Poland.
166. Robinson, S. K., F. R. Thompson III, T. M. Donovan, D. R. Whitehead, 192. Swank, W. T., and P.V. Bolstad. 1994. Cumulative effects of land use
and J. Faaborg. 1995. Regional forest fragmentation and the nesting practices on water quality. Pages 409-421 in N. E. Peters, R. J. Allan, and
success of migratory birds. Science 267:1987-1990. V.V. Tsirkunov, editors. Proceedings of the Rostov-on-don Symposium
167. Rochelle, J. A. 1998. Forest fragmentation: wildlife and management on Hydrochemistry 1993: hydrological, chemical, and biological
implications. Conference Summary Statement, 18-19 November 1998. processes affecting the transformations and the transport of
Portland, OR. contaminants in aquatic environments. IAHS Publication 219,
168. Rosenberg, K.V., A. A. Dhondt, and J. D. Lowe. 1996. Lessons from the Oxfordshire, England
landscape. Birdscope 10: 3-5. 193. Thomas, J. W., R. B. Andersen, C. Maser, and E. L. Bull. 1979. Snags.
169. Rost, G. R., and J. A. Bailey. 1979. Distribution of mule deer and elk in Pages 60-70 in J. W. Thomas, editor. Wildlife habitats in managed forests,
relation to roads. Journal of Wildlife Management 43: 634-641. the Blue Mountains of Oregon and Washington. U.S. Forest Service
170. Rudnicky, J. L., and M. J. McDonnell. 1989. Forty-eight years of canopy Agriculture Handbook 553, Washington, D.C.
change in a hardwood-hemlock forest in New York City. Bulletin of the 194. Thomas, J. W., R. M. DeGraaf, and J. C. Mawson. 1977. Determination
Torrey Botanical Club 116: 52-64. of habitat requirements for birds in suburban areas. U.S. Forest Service
171. Russwurm, L. H. 1977. The surroundings of our cities. Community Research Paper NE-357.
Planning Associated Press, Ottawa, Ontario, Canada. 195. Thomas, J. W., C. Maser, E. D. Forsman, J. B. Lint, E. C. Meslow, B. R.
172. Savard, J. P. L. 1978. Birds in metropolitan Toronto: distribution, Noon, and J.Verner. 1990. A conservation strategy for the northern
relationships with habitat features, and nesting sites. Ph.D. Dissertation, spotted owl. Report of the interagency committee to address the
University of Toronto, Ontario, Canada. conservation strategy of the northern spotted owl. U.S. Forest Service,
Portland, OR.
 CHAPTER 12: URBAN ENVIRONS 341

196. Tomialoj‰, L. 1985. Urbanization as test of adaptive potentials in

birds. Pages 608-614 in: XVIII Congresses Internationalist Ornithologici,
Moscow, Russia.
197.Valden, D. W. 1980. Urban small mammals. Journal of Zoology 191:
403-406.
198. Van Druff, L. W., and R. N. Rowse. 1986. Habitat association of
mammals in Syracuse, New York. Urban Ecology 9: 413-434.
199. Van Druff, E. G. Bolen, and G. J. San Julian. 1994. Management of
urban wildlife. Pages 507-530 in: T. A. Bookout, editor. Research and
management techniques for wildlife and habitats. The Wildlife Society,
Bethesda, MD.
200. Vasic,V., and V. Stevanovic. 1970. Characteristics of the avifauna of the
inner city area of Belgrade. Larus 24: 107-118.
201.Vepsalainen, K., and B. Pisarski. 1982. The structure of urban ant
communities along a geographical gradient from north Finland to
Poland. Pages 155-168 in: M. Luniak and B. Pisarski, editors. Animals in
urban environment. Polish Academy of Science, Institute of Zoology,
Warsaw, Poland.
202. Vizyova, A. 1986. Urban woodlots as islands for land vertebrates: a
preliminary attempt on estimating the barrier effects of urban
structural units. Ekologia (CSSR) 5: 407-419.
203. Walcott, C. F. 1974. Changes in bird life in Cambridge, Massachusetts
from 1960 to 1964. The Auk 91: 151-160.
204. Warren, J. W. 1997. Deer overabundance. Wildlife Society Bulletin
Special Issue 25: 2.
205. Washington Department of Fish and Wildlife. 1999. Priority habitats
and species list. Washington Department of Fish and Wildlife, Olympia,
WA.
206. Washington State Department of Natural Resources. 1998. Our
changing nature. Washington Department of Natural Resources,
Olympia,WA.
207. Watson, J. W., and D. J. Pierce. 1998. Bald eagle ecology in western
Washington with an emphasis on the effects of human activity. Final
Report. Washington Department of Fish and Wildlife, Olympia, WA.
208. Wear, D. N., M. G. Turner, and R. J. Naiman. 1998. Land cover along an
urban-rural gradient: implications for water quality. Ecological
Applications 8: 619-630.
209. Welch, E. B., J. M. Jacoby, and C. W. May. 1998. Stream quality. Pages
69-94 in: R. J. Naiman and R. E. Bilby, editors. River ecology and
management: lesson from the Pacific Coastal Ecoregion. Springer-Verlag,
New York, NY.
210. Whitaker, J. O., Jr. 1996. National Audubon Society field guide to
North American mammals. Alfred A. Knopf, New York, NY.
211. Whitcomb, R. F., D. S. Robins, J. F. Lynch, B. L. Bystrak, M. K. Limkiewitz,
and D. Bystrak. 1981. Effects of forest fragmentation on avifaunas of the
eastern deciduous forest. Pages 125-205 in: R. L. Burgess and D. M.
Sharpe, editors. Forest island dynamics in man-dominated landscapes.
Springer-Verlag, New York, NY.
212. Wilcove, D. S. 1985. Nest predation in forest tracts and the decline
of migratory songbirds. Ecology 66: 1211-1214.
213. Wilcove, D. S., C. P. McLellan, and A. P. Dobson. 1986. Habitat
fragmentation in the temperate zone. Pages 237-256 in: M. E. Soulé,
editor. Conservation biology: the science of scarcity and diversity.
Sinauer Associates, Sunderland, MA.
214. Wilson, E. O., and E. O. Willis. 1975. Applied biogeography. Pages 522-
534 in: M. L. Cody, and J. M. Diamond, editors. Ecology and evolution of
communities. Harvard University Press, Cambridge, MA.
215. Woolfenden, G. E., and S. A. Rohwer. 1969. Breeding birds in a Florida
suburb. Florida State Museum Bulletin 13.
216.Yahner, R. H. 1988. Changes in wildlife communities near edges.
Conservation Biology 2: 333-339.
217.Yalden, D. W. 1980. Urban small mammals. Journal of Zoology 191:
403-406.
218. Zimmerman, D. A. 1973. Range expansion of Anna’s hummingbird.
American Birds 27: 827-835.
 13
Wildlife of Agriculture, Pastures, and
Mixed Environs
W. Daniel Edge

Composition, Structure,
and Distribution
The Agriculture, Pasture, and Mixed Environs habitat habitats for wildlife. The most common characteristic is a
(hereafter agricultural habitat) in Oregon and Washington regular pattern of disturbance. Depending on the crop
is highly diverse in part because of the large number of produced and the cultural practices employed in these
agricultural commodities produced in the two states, but areas, agricultural habitats may be substantially modified
also because this habitat classification includes other linear (e.g., cultivated, grazed, harvested, hayed, mowed,
landscape components that could not be classified pruned, tilled, etc.) at least once per year. Because of these
individually (e.g., road sides, shelterbelts, etc.). This disturbances, many agricultural habitats are important for
habitat includes managed and unmanaged pasture, wildlife only on a seasonal basis, whereas others may be
cultivated cropland, orchards, vineyards, nurseries, and ecological traps during the breeding season.7 Unlike most
associated scattered dwellings and intervening areas of of the other wildlife habitats covered in this book, almost
vegetation. There are >410 agricultural crops produced in all agricultural habitats are privately owned. Only the
the two states. This habitat is maintained across a wide urban habitat classification contains as much privately
range of ecoregions and climatic conditions typical of both owned property as agricultural habitat. As a consequence,
states. Climate constrains crop production at upper management benefitting wildlife in agricultural habitats
elevations where there are <90 frost-free days. Agricultural occurs predominately because of personal interest of the
habitat in arid regions east of the Cascades with <10 inches landowner, stewardship, other conservation benefits, the
(25 cm) of rainfall requires supplemental irrigation or recreational value associated with wildlife in these areas,
fields lying fallow for 1-2 years to accumulate sufficient or because of financial disincentives of U.S. Department
soil moisture. This habitat is found from 0 to 6,000 feet (0- of Agriculture (USDA) programs under the Farm Bill.30, 53
1,829 m) in elevation. Soil types are variable, but soils Because of the wide distribution and regular
usually have a well-developed A horizon. disturbance of agricultural habitats, wildlife that occur in
Agricultural habitat is widely distributed at low- to these areas display a number of adaptations. Many of the
mid-elevations (<6,000 feet [<1,829 m]) throughout both species that use agricultural habitats are habitat
states. This habitat is most abundant in broad river valleys generalists, adapted for using several cover types for both
in both states and on gentle rolling terrain east of the feeding and breeding. In many areas of both states,
Cascades. Agricultural habitat occurs within a matrix of pastures and cultivated cropland have replaced native
other habitat types at low- to mid-elevations, including prairie habitats. However, many of these areas (e.g., annual
Eastside (Interior) Grasslands, Shrub-steppe, Westside and perennial grass seed fields, wheat, barley,
Lowlands Conifer-Hardwood Forest, and other low- to Conservation Reserve Program fields, and hay pastures)
mid-elevation forest and woodland habitats. Agricultural are structurally similar to the native prairie habitats they
habitat often dominates the landscape where terrain is flat replaced, and often are used by habitat specialists of native
or gently rolling, on well-developed soils, broad river prairie habitats (e.g., western meadowlark, grasshopper
valleys, and areas with access to abundant irrigation water. sparrow, bobolink).8, 70
Unlike other habitat types, agricultural habitat is often
characterized by regular landscape patterns (squares, Important Habitat Elements
rectangles, and circles) and straight borders because of Agricultural habitats in Oregon and Washington include
ownership boundaries and multiple crops within a region. several habitat elements, all of which are anthropogenic
Edges can be abrupt along the habitat borders within in origin. Because this landscape is human-dominated,
agricultural habitat and with adjacent habitats. there are no habitat features that are considered unique—
Agricultural habitats have two dominant characteristics those habitat features that cannot be created. However,
that separate them from the other wildlife habitats some agricultural habitat is adjacent to unique features
described in this book: regular disturbance and ownership such as rimrocks with cliffs and caves. Important habitat
pattern, both of which are important in managing these elements in these areas include ephemeral wetlands, wells
342
 CHAPTER 13: AGRICULTURAL ENVIRONS 343

Table 1. Wildlife benefits provided by habitat elements in agricultural habitats in Oregon and Washington.

Habitat feature Wildlife benefits

Water Food Shelter Breeding Structural Edge Corridors
sites diversity habitat

Farmed or ephemeral wetland X X X X

Water developments X X X ?a
Buildings and farm structures X X X
Shelterbelts and windbreaks X X X X X X
Hedge and fence rows X X X X X X
Roadsides X X X X X X
Field borders X X X X X X
Odd areas X X X X X ?a
Unknown or uncertain benefit.
a

and water developments, deserted dwellings and other Wells and Water Developments
buildings, shelterbelts, hedge and fence rows, roadsides, Wells and water developments are habitat elements that
and field borders (Table 1). are common in agricultural areas, especially east of the
Several habitat elements, including shelterbelts, Cascade Mountains in Oregon and Washington. Water is
windbreaks, hedge and fence rows, field borders, and scarce in many of these areas, and water developments
roadsides are important components of the agricultural become an important habitat feature for wildlife. Intensive
landscapes because they provide permanent cover and livestock management, characteristic of many agricultural
structural diversity and may function as corridors.5 areas in the arid regions of both states, requires a reliable,
Furthermore, these habitat features, because of their uniform distribution of water in grazing pastures.44, 89
narrow, linear nature, are composed entirely of edge Spring developments, reservoirs, and precipitation
habitat and typically have high species diversity. 10 catchments or guzzlers are water developments used for
However, depending on their abundance and distribution livestock production that also benefit wildlife. These
in the agricultural landscape, these elements may function structures can be designed or modified to increase wildlife
as ecological traps because of high rates of predation or use and to prevent accidental drowning in steep-sided
brood parasitism.6, 70, 100 structures. 17, 22, 72, 101 Especially when protected from
livestock grazing, these sites usually develop vegetation
Ephemeral or Farmed Wetlands characteristic of riparian areas and become oases for
Agricultural habitats include numerous ephemeral or wildlife.15, 43, 44, 88
farmed wetlands. In many areas, especially along broad
floodplains, native wetlands were drained for agriculture, Buildings and Farm Structures
but most of these sites still retain two of the three Deserted dwellings and other buildings are also habitat
characteristics that define wetlands (hydrology and hydric elements in agriculture habitats. Buildings provide
soils).21, 86 Consequently, these areas have standing water structure in agricultural areas often characterized by
on them •1 month of the year. Depending on the residual homogeneous landscapes of low vegetation stature; these
vegetation or the crop from the previous growing season, structures provide habitat features that may otherwise be
these sites can be extremely important wildlife habitats.55 unavailable.20, 50 These structures are used as nest sites for
Ephemeral and farmed wetlands that were used to swallows and barn owls, roosts and hibernacula for several
produce small grains (i.e., sorghum, rice, wheat, barley, species of bats, and shelter for many small mammals,
oats, grass seed) during the previous growing season are reptiles, and amphibians.3, 46, 50
critically important as stop-over and wintering habitat for
millions of migratory waterfowl in the Pacific Flyway. Shelterbelts and Windbreaks
These sites provide both wetland habitat and abundant Shelterbelts are vital habitat features of agricultural
food sources because of the waste grain typical of small habitats. Shelterbelts and windbreaks are terms that have
grain production practices. During the northward been used interchangeably for human-made habitats
migration in the spring, the abundant food in these created by planting rows of trees and shrubs along the
wetlands is especially important for energy conservation border of agricultural fields (Figure 1). Shelterbelts have
and nutritional conditioning for egg production.45 been promoted for a number of conservation benefits
Ephemeral or farmed wetlands may also be important including (1) reducing wind erosion; (2) shelter for
breeding sites for amphibians. For example, many native growing plants, farmsteads, and livestock; (3) snow fences
hay pastures east of the Cascade Mountains in both states and living screens; (4) wood and fiber production; and (5)
are flood-irrigated during the spring months, resulting in aesthetics. They are also excellent wildlife habitat and have
large areas of grass vegetation with 2-8 in (5-20 cm) of been promoted as such by both state and federal
standing water—conditions that provide ideal breeding conservation agencies. In large areas of eastern Oregon
sites for some amphibians.
 344 Wildlife–Habitat Relationships in Oregon and Washington

Figure 1. Shelterbelts and windbreaks, created by planting Figure 2. Hedge and fence rows are rows of shrubs and trees
rows of trees and shrubs, are important wildlife habitat in bordering agricultural fields that provide food, cover, and
agricultural areas. (Photograph by W. Daniel Edge) dispersal habitat for wildlife. (Photograph by W. Daniel
Edge)
and Washington, shelterbelts and riparian areas account
for most of the tree habitat over hundreds of square miles.
Shelterbelts are disproportionally important wildlife
habitats, especially for neotropical migratory landbirds,9,
70, 75, 99
reptiles, and small mammals.98 Shelterbelts are used
for year-round cover and feeding sites, for nesting,9, 70 and
as dispersal corridors32 by birds and mammals. Shelterbelts
that incorporate several plant species and provide ground,
mid-story, and canopy-level vegetation structure have
higher wildlife species diversity than single-species,
single-structure shelterbelts.75, 99

Hedge and Fence Rows

Hedge and fence rows are habitat elements of agricultural
habitats that enhance these areas for wildlife; they are
continuous or near-continuous shrub and tree vegetation
communities that border agricultural fields (Figure 2).
Fence lines are common in agricultural landscapes,
although not as common as they used to be. Historically,
when most farms ran some livestock, fences were much Figure 3. Roadsides can provide important year-round cover
for wildlife in agricultural areas. (Photograph by W. Daniel
more common than they are now; they have declined in
Edge)
agricultural landscapes since the advent of “clean
farming.” 30, 70 Nevertheless, many fields, especially
pastures used for grazing, are separated or surrounded
by fences. Fence and hedge rows provide shelter, food, farming costs. If used adjacent to riparian areas, wetlands,
and structural complexity to agricultural landscapes (Table etc., field borders also function as filter strips and improve
1) and typically have high species richness and wildlife water quality. Few studies have looked at wildlife
abundance relative to the agricultural fields they border.9, community responses to different field borders, but
59, 70
wildlife use is likely to be dependent on the width, length,
and the structural diversity of the borders.9, 59, 70
Field Borders
Field borders are narrow strips of permanent vegetation Roadsides
planted between fields and around the perimeters of fields. Roadsides are another important habitat feature of
Field borders benefit wildlife by providing permanent agricultural habitats, and are a common attribute of all
cover, forage, increased edge habitat, and increased terrestrial ecosystems except in wilderness areas. Roads
horizontal and vertical diversity (Table 1). Field borders abound in agricultural landscapes because the culture and
also control erosion and can protect field edges that are harvest of crops typically require producers to visit fields
used as “turn-arounds” or travel lanes for farm equipment. multiple times, with different pieces of equipment (Figure
Another farmer benefit includes reduced planting and 3). Roads in agricultural ecosystems vary from simple dirt
 CHAPTER 13: AGRICULTURAL ENVIRONS 345

tracks between fields managed by the landowner to four- Wildlife Diversity in

lane Interstate Highways managed by the state
Department of Transportation (DOT). High-standard
Agricultural Habitat
roads such as county roads and state and interstate Agricultural habitats in Oregon and Washington support
highways are typically constructed by borrowing dirt from diverse wildlife communities. Agricultural habitat in the
the roadside to raise the roadbed between the borrow pits. two states is used by 342 species—more than any other
Thus, a road has the roadbed, the borrow pit, and the fore habitat. This high species richness is a function of the broad
and back slopes of the borrow pits. The fore and back slope distribution of the habitat type across the two states, but
and the borrow pits are potentially good wildlife habitat more importantly, it is a function of the wide variety of
in agricultural areas, especially compared to the adjacent habitat conditions, land uses, and crops that are included
crop fields. The borrow pits act as drainage systems and within this habitat classification. For example, agricultural
often function as ephemeral or permanent wetlands. The habitat in eastern Oregon or Washington might include
slopes are almost always covered with permanent grass unimproved and improved pastures, orchards, row crops,
or shrub vegetation, but may be gravel or bare soil. annual grains, and shelterbelts all relatively close to one
The amount of habitat provided by roadsides depends another. However, any one of these crops or cover types
on the road standard and ownership. Rights-of-way, the alone is likely to support a much more depauperate
areas set aside in the road easement, vary in width from wildlife community than a combination of types. For
jurisdiction to jurisdiction. Roads on private land are not example, unimproved pastures on the eastside of the two
required to set aside areas and many roads on farms are states might be expected to support a wildlife community
immediately bordered by the field (i.e., the field is plowed similar in diversity to the eastside grasslands, which are
right up to the roadside). Roadsides adjacent to county used by only 172 species.
roads and state and federal highways have rights-of-way Because of their high levels of disturbance, agricultural
that are •16 feet (5 m) wide on either side of the road, and habitats are prone to invasion by exotic organisms. Regular
for some Interstate highways, may be >328 feet (100 m) soil disturbance typical of many agricultural practices
wide in places. Thus, roadsides managed by the county provides opportunities for colonization by invasive weeds.
road departments or the state DOT represent more Intensive grazing may reduce the competitiveness of some
potential habitat than is managed by any other state plant species in pastures, allowing invasion of exotic plant
agency (i.e., state or county parks, state wildlife areas). species. Wildlife communities can be affected by
For example, DOT manages 7,500 miles (12,060 km) in conversion of native habitats to non-native plant
Oregon and 7,035 miles (11,320 km) in Washington of state communities. 96 However, in most areas intensively
and federal highways. If we apply the minimum of 16 feet managed for agricultural production, invasive weeds are
(5 m) on either side of the road as right-of-way, roadsides aggressively controlled through the use of herbicides or
managed by the DOT represent a minimum of 45 and 43 integrated pest management. Invasive plant species more
miles 2 (121 and 113 km 2) of habitat in Oregon and commonly become a problem in special habitats within
Washington, respectively. County road departments agricultural areas such as road sides, field borders, fence
manage almost three times the amount of habitat as the rows, and shelterbelts. Scotch broom (Cytisus scoparius)
DOT in the two states. and Himalaya blackberry (Rubus procerus) are two
Most data on wildlife use of roadsides or road rights- examples of invasive weedy species that regularly colonize
of-way come from higher-standard roads (county road up these special habitats in both states.
to interstates). Roadsides provide shelter, food, and nesting Agricultural habitats are commonly inhabited by exotic
sites in agricultural landscapes (Table 1). Adams and Geis1 wildlife species. Nutria, Virginia opossum, and bullfrogs
reported that small mammal communities along roadsides commonly occur in agricultural habitats along major river
differ depending on the standard of road—interstate valleys on the westside of both states. The rock dove, house
highways had greater species richness than did county sparrow, European starling, black rat, Norway rat, and
roads. Camp and Best13 reported higher avian species house mouse occur in many agricultural areas throughout
richness along roadsides than in adjacent row crop fields, both states. Furthermore, several species, including the
and noted that the number of species was greater than chukar, gray partridge, ring-necked pheasant, wild turkey,
the number of species reported for most grasslands. They and eastern cottontail, have been introduced for sport in
also reported that bird abundance was related to the both states. The ring-necked pheasant and gray partridge
structural development (height and density) of roadside are closely associated with agricultural habitats and have
vegetation. Camp and Best 14 found more nests in been widely introduced for the specific purpose of
backslope areas than in foreslope areas. Roadsides in establishing gamebird populations in these non-native
intensively farmed areas may be particularly good habitat habitats because agricultural areas do not support native
for some species. Warner et al.92 found a greater density gamebird populations in huntable numbers.
of pheasant nests in roadsides than in any other cover type, Because of the non-natural condition of agricultural
and reported that nest success was 29%, which was habitats, they are typically not critical habitats for
comparable to hayfields. threatened, endangered, or sensitive species. However,
conversion to agriculture has directly caused the decline
of some native ecosystems, including wetlands, riparian
areas, native prairie, and oak woodlands, and the species
 346 Wildlife–Habitat Relationships in Oregon and Washington

that depend on these systems. Several federally listed plant to a lesser extent as breeding habitat. 70 Typically,
species occur in these declining ecosystems, and neotropical migrants use habitat elements within
restoration of these ecosystems will likely require the agricultural areas such as shelterbelts, windbreaks, fence
conversion of agricultural habitats to native vegetation rows, and field borders for stop-over and breeding habitat.
types. Only one federally listed wildlife species (Columbia Waterfowl make extensive use of cropland during spring
white-tailed deer) is associated with agricultural habitats and fall migrations and as winter habitat because of waste
in the two states. Ten mammal species of concern, mostly grain in many crop fields. Canada geese extensively use
bats, occur in agricultural areas. grass seed fields and pastures during winter stop-overs
in both states. Although many large mammals such as
Wildlife Use of deer, elk, and pronghorn antelope may use agricultural
habitats year round, use is most pronounced during fall
Agricultural Habitat and winter.
Most wildlife species using agricultural habitat are either Wildlife use of agricultural habitat is dependent on the
seasonal migrants or use these areas in conjunction with crop being produced and the tillage method used to
other habitat types. Most amphibians, reptiles, birds, and cultivate the crop. Crops vary substantially in respect to
mammals are only partially associated with or present in the amount of food, shelter, structure, and frequency of
agricultural habitats (Figure 4). No amphibian or reptile disturbance. In general, unimproved pastures that are
species is closely associated with agricultural habitats, but grazed rather than hayed offer some of the best habitat
53 bird and 15 mammal species are closely associated with among agricultural crops, especially if fall regrowth is
these areas. The most common activity for all wildlife in sufficient to offer winter and early spring cover. Fruit and
agricultural habitats is breeding and feeding (Figure 5). nut orchards provide the most long-term habitat structure
However, most of these species breed and feed in other among agricultural crops, and are used for nesting and
habitats as well, and are not exclusively found in foraging by many songbird species. A few wildlife species
agricultural habitats. Most breeding and feeding activity will nest in most row crops, but these habitats are
in agricultural habitat occurs in habitat elements such as ephemeral and may be harvested during the nesting
shelterbelts, windbreaks, fence rows, and field borders season (see section on ecological traps below). Most
rather than in crop fields. Although many species do breed vegetable and root crops offer little habitat for wildlife,
or nest in crop fields, feeding is the most common activity especially after the crop has been harvested. Depending
of wildlife in cropland.9, 10, 70 Some closely associated on how crop residues are handled, many small grains may
breeders, such as the sandhill crane, short-eared owl, offer both food (waste grain) and shelter (stubble) for
western meadowlark, bobolink, grasshopper sparrow, and several months after harvest. The fallow system used for
savannah sparrow breed in agricultural habitats because producing wheat on large areas in the eastern portion of
they are structurally similar to the native grassland both states may provide stubble and waste grain for up to
ecosystems that they replaced.9 a year every other year.
Many wildlife species make only seasonal use of The tillage practice used also determines the residue
agricultural habitats. Neotropical migratory landbirds available for winter and early spring cover and the
make extensive use of agricultural habitats, primarily as availability of food. Conventional tillage is used to produce
stop-over habitat during spring and fall migrations and

80
Close associate
Figure 4. Percent of species by vertebrate 70 Associated
(11)
group closely associated, associated, or Present
present in agricultural habitats. Number
in parenthesis above bars is the number of 60
species. (8) (8) (107)
Percent of species

50 (44)

40 (6) (34)

30
(51) (51)

20 (17)

10

0
Amphibians Reptiles Birds Mammals
 CHAPTER 13: AGRICULTURAL ENVIRONS 347

100 (16)
Breeding & feeding
(81) Figure 5. Percent of species by
Feeding only vertebrate group using
80 Breeding only agricultural habitats for breeding,
feeding, or other activities.
Number in parenthesis above bars
(10) is the number of species.
Percent of species

(124)
60

(84)
40 (6)

20
(14)

(1)
(2)
0
Amphibians Reptiles Birds Mammals

many crops in both states. Under conventional tillage till uses special planters to plant the crop in the residue
practices, crop residues from the previous crop are plowed from the previous season, and crop residues are therefore
under in the late summer or fall, followed by disking and present year round. No-till provides overwinter cover,
harrowing in the spring to prepare the seed bed for waste grain is not incorporated into the soil, and arthropod
planting (Figure 6). Usually at least one pass with communities associated with plant residues are not
cultivating equipment is used to control weeds while the destroyed. However, because no-till substitutes herbicides
new crop is growing. Conventional tillage provides few for cultivation to reduce weeds, seeds from annual plants,
wildlife benefits from the time the fields are plowed until which are important food sources for some wildlife,
the new crop is established. Waste grain is typically become unavailable. Crop production under all tillage
plowed under and there are no crop residues to provide practices may be an ecological trap during the nesting
cover for -6 months of the year. However, arthropods and season.7
small mammals may be temporarily available for wildlife
following plowing and disking. Many crows, gulls, and Status, Threat, and Trends in
even coyotes are often seen following plows or using fields
shortly after plowing.
Agricultural Habitats
Conservation tillage includes a number of practices in Agricultural habitats have changed substantially over the
which •30% of the crop residues protect the soil surface past two centuries. The most pronounced change has been
between harvest and planting. Reduced till and no-till are the steady increase in the conversion to agricultural habitat
two of the more common conservation tillage practices. until the 1950s (Figure 7). Before European settlement,
Under reduced tillage, a disk is used for the primary tillage native American tribes cultivated some areas along major
in the spring; residues from the previous crop provide streams and rivers, but the total area under cultivation
overwinter cover and waste grain is not plowed under was minute compared to the extent of modern agriculture.
until disking. Reduced tillage relies on cultivation to Although trade in agricultural products was important
control weeds once the crop is established. Arthropods for some tribes, agriculture was practiced on a subsistence
associated with the previous crop’s residue may be scale by native Americans. European and Asian
available for wildlife under reduced tillage practices. No- immigrants introduced the horse-drawn plow beginning

Figure 6. Conventional tillage practices

remove over-winter cover and bury waste
grain, providing few benefits for wildlife.
 348 Wildlife–Habitat Relationships in Oregon and Washington

Landscape and Farm Diversity

Since the beginning of the twentieth century agricultural
habitats have changed dramatically in respect to diversity
and spatial configuration, because of changes in
agricultural markets and technology, food choice,
urbanization and economic development, tax policy, farm
policy, and agricultural research, etc.30, 56 Farm diversity
has declined dramatically since the 1960s. Before this, most
farms produced multiple products and many were
diversified crop-livestock operations. Many farmers
produced forage and feed grains for their livestock in
addition to other crops. Forage and hay pastures, which
required longer crop rotations and in general were better
habitat for wildlife than row crops, were more abundant
than they are now. Since the 1960s the trend in agricultural
Figure 7. The number of farms and amount of farmland in the
production has been toward specialization in crop
United States, 1900-1997.84
production on individual farms as well as regionally.56 The
result is extensive agricultural habitat that is spatially and
in the early 1800s, and throughout the rest of the century structurally homogeneous. This decline in habitat
much of the land on the flood plains of the broad river diversity has resulted in declining wildlife diversity in
valleys in both states was converted to agriculture. This agricultural habitats.9, 30, 70
expansion of agriculture resulted directly in substantial Changes in farm and field size also have resulted in
declines in natural habitats, especially wetlands, native less diverse agricultural habitats. Although the number
prairies, and riparian woodlands. The advent of steam and of farms has declined by >50% since the 1950s, the number
gasoline-powered machinery accelerated the rate of of harvested acres has remained relatively constant,84 and
conversion of native habitats to agriculture, and by the the average farm size has almost tripled. Coupled with
1930s the land best suited for agriculture had already been increased product specialization and operation scale,
converted. Between the 1950s and 1980s improvements average field size has also increased substantially. This
in irrigation technology and USDA programs encouraged increase in field size decreases the amount of edge habitat
conversion of less suitable areas to farmland, primarily around fields, which are important sources of wildlife
arid areas in the eastern portions of both states. However, diversity in agricultural habitats.9, 10, 66, 100 Furthermore,
recessions in the 1950s, 1960s, and early 1980s resulted in increased field size also decreases the abundance of habitat
a decline in the number of farms and total harvested acres. elements such as shelterbelts, fence rows, and field
Since 1985, U.S. agricultural policy has changed the borders. Wildlife response to field size and adjacent
amount and character of agricultural habitat in both habitats is species-specific,10, 35, 66, 100 and dependent on land-
Oregon and Washington. The Food Security Act of 1985 use patterns.27, 74, 81 Heckert35 characterized grassland bird
(the Farm Bill) contained the first important conservation species as edge species, area-sensitive, and vegetation-
provisions from a wildlife standpoint.30 The 1985 Farm restricted. Edge species such as the song sparrow, red-
Bill contained disincentives for further conversion of winged blackbird, and American goldfinch might be
wetlands and highly erodible soils to agriculture under expected to decline as field size increased. However, larger
the Swampbuster and Sodbuster programs. More field sizes will be beneficial for area-sensitive species such
importantly, the 1985 Farm Bill introduced the as the western meadowlark, grasshopper sparrow,
Conservation Reserve Program (CRP), which paid farmers savannah sparrow, and bobolink 35 (see Chapter 20,
to take highly erodible soils out of production under 10- Westside Grasslands and Chaparral).
year contracts. The CRP has resulted in significant Changes in farming practices also have resulted in
improvements of wildlife habitat in farmlands, especially declines in habitat diversity. The advent of “clean farming”
for grassland-dependent species.8, 30, 37, 52, 94 Building on the promoted by the Land Grant universities in both states
success of the 1985 Farm Bill, the 1995 Farm Bill enhanced caused further declines in the availability of habitat
the CRP with a continuous program and introduced elements in agricultural habitats. Field borders, fence rows,
several other significant conservation programs including and road sides were removed in an effort to eliminate
the Wetland Reserve, Environmental Quality Incentives, refugia for weeds and pest species. Thus “clean farming”
and Waterbank programs (see McKenzie53 for a description has resulted in further simplification of agricultural
of these programs). The net result of these programs has landscapes.
been a decline in the total land area under crop production
and enhancement of wildlife habitat on farms and ranches, Ecological Traps
especially wetlands and riparian areas. Agricultural habitats have a high potential to become
ecological traps because of farm operations and the
abundance and distribution of habitat features in
agricultural landscapes. Ecological traps are human-made
 CHAPTER 13: AGRICULTURAL ENVIRONS 349

areas that, based on physical or vegetation characteristics, Conventional Tillage

appear suitable for nesting but which, by virtue of some Primary tillage— plow in fall
confounding factor(s), result in population sinks rather
than sources for species that use those sites.7, 29 Disk
As previously described, most agricultural habitats are Harrow
characterized by a regular pattern of disturbance, which
varies with the crop being produced and the production Plant
methods used to grow, cultivate, and harvest the crop. Fertilize
Production of most crops requires multiple field
operations that may include plowing, disking, harrowing, Rotary hoe
planting, cultivating, applying herbicides and pesticides, Cultivate 2X
as well as harvesting (Figure 8). The production of some
crops may require •10 field operations during the growing
season, and most agricultural crops produced in Oregon Reduced Tillage
and Washington require at least four field operations. The Primary tillage—disk in spring
number of field operations depend on the tillage method
being used to produce the crop. Reduced or no-till Disk
production methods substitute at least one herbicide
application for ground-disturbing cultivation (Figure 8). Plant
The natural history characteristics of wildlife nesting
in agricultural fields determine the level of disturbance
from field operations. According to Best,7 severity of nest Fertilize
disturbance by field operations depends on the (1) nest
position in relation to the crop rows, (2) length of the Cultivate 2X
nesting cycle, (3) duration of the breeding season and the
species’ propensity to renest after nest failure, and (4)
timing of the breeding season. Nest position within a crop No Tillage
field is important because field operations can destroy
nests; destruction is more likely for between-row nesters Plant
such as ring-necked pheasants, killdeer, grasshopper
sparrows, and western meadowlarks than for within row Fertilize
nesters such as mourning doves and vesper sparrows.
Duration of nesting season and the propensity to renest is Spray herbicides
important for some species; mourning doves have a long
nesting season and will renest multiple times. Ring-necked Cultivate (optional)
pheasants have a long nest cycle and typically only raise
one brood per season (but some will renest). Other species
will renest •1 time. Timing of the breeding season is also
Mar Apr May Jun Jul Aug Sep
important. Horned larks begin nesting almost 1 month
before field operations begin whereas mourning dove and
vesper sparrow nesting extends beyond when fields are Figure 8. Sequence of field operations used in the production
cultivated. Best7 also points out that another form of of a crop is dependent on the tillage method used for crop
ecological trap occurs when hayfields and pastures are production. This diagram illustrates the production of corn
converted into no-till row crops. Species that are strongly and soybeans (after Best,7 used with permission).
philopatric and nest in grasslands (western meadowlarks
and grasshopper sparrows) may not perceive the extent previously mentioned, agricultural landscapes have
of habitat alteration until later in the nesting season and changed over the past quarter-century, resulting in larger
are unlikely to establish new territories elsewhere. field sizes and fewer edges and habitat elements such as
Mammal populations are also susceptible to ecological shelterbelts, fence rows, and field borders. As the
entrapment. Edge et al.26 reported that mowing of alfalfa abundance of edge and special habitats declines, those that
caused an approximately 50% decline in gray-tailed vole remain in the landscape become fragmented and
populations, regardless of the initial density. successively isolated. Survival and reproductive success
Ecological traps in agricultural habitats also may occur of wildlife using these remaining habitats may be low
because of the amount and distribution of edge habitat because of depredation from omnivorous predators.74, 79,
100
and habitat elements. Edges as ecological traps have Populations of many omnivorous predators including
received a substantial amount of research.66, 100 However, corvids, gulls, canids, raccoons, opossums, skunks, and
most of this work is related to forest-edge habitats, and rodents are enhanced by the interspersion of cropland food
may only be relevant to agricultural ecosystems where sources (i.e., crops, waste grains, arthropods, and fruits in
woodlots of varying size are intermixed in farmlands. As fence rows, hedgerows, or shelterbelts). Many of these
 350 Wildlife–Habitat Relationships in Oregon and Washington

predators are highly mobile and preferentially search edge of field testing a pesticide, which often exceeds $1 million.
habitats, and as a result, nests in or near predator travel However, without field tests, greater uncertainty exists
lanes may have a low likelihood of success. regarding the potential impact of pesticides on nontarget
Brood parasitism by brown-headed cowbirds may be species.42, 65, 82 Furthermore, the models the EPA uses to
pronounced in agricultural landscapes. Cowbirds are well predict effects of a chemical on wildlife often perform
adapted to agricultural landscapes51 and exploit a variety poorly under field conditions25, 73, 90 because of assumptions
of host species.69 Agricultural landscapes with abundant inherent in the use of these models.82
edge habitats composed of shrub and tree communities A pesticide’s capacity to harm wildlife is dependent
are well suited to cowbirds because of the species’ practice on several factors, including the chemical’s characteristics
of spatially partitioning their breeding and feeding and toxicity, and the timing, duration, and dose of
activities.23, 24, 71 Cowbird feeding areas typically are open exposure.62: 133, 80 The characteristics of a pesticide include
fields, livestock facilities and corrals, and other human- its propensity to bioaccumulate and its degradation rate.
altered areas,2, 23 whereas breeding sites are host-rich areas Toxicity of a pesticide refers to how poisonous the chemical
such as riparian areas,2 sparse forests,71 and edge habitats is to wildlife. Pesticides vary substantially in their toxicity
between forests and meadows.29 Thus, the distribution and to wildlife; some are relatively nontoxic even at high
pattern of shrub and woody edge habitats in agricultural application rates whereas others are extremely toxic at
landscapes may increase brood parasitism and create normal application rates. Typically, insecticides are much
ecological traps for species nesting in those areas.11, 38, 69 more toxic to wildlife than are herbicides or fungicides.
Furthermore, toxicity of a chemical varies substantially
Agricultural Chemicals among wildlife species and even within a species
Modern agriculture typically requires extensive chemical depending on the age, sex, reproductive condition, and
inputs in the form of pesticides and fertilizers. These health of the exposed animals. The timing of a chemical
chemicals used in agricultural areas have both direct and application may determine its impact on wildlife—some
indirect effects on wildlife living in farm landscapes, and wildlife are more susceptible during nesting, migration,
have been a concern of wildlife biologists.16, 41, 63 The impact or periods of low food availability. The duration of
of chemical pollutants on nontarget species, primarily exposure can be important because some species are
birds and mammals, is of special concern for threatened affected by a pesticide only if the exposure is extended.
or endangered species85 or species that exist in small Finally, the effect of a pesticide is dependent on the dose
isolated populations.54, 93 or concentration of the chemical in the environment. Some
pesticides can be applied within a range of applications
Pesticides. Pesticides, including herbicides, insecticides,
rates for a pest in a particular crop. A pesticide applied at
fungicides, and rodenticides, are widely used to control
a higher application rate will almost always have a greater
agricultural pests. For example, the 1997 fall potato crop
potential to impact wildlife than the same chemical
alone required 333 tons (302 metric tons) of pesticides
applied at a low application rate.
(herbicides, insecticides, and fungicides) in Oregon and
Wildlife exposure to pesticides occurs in several ways.
634 tons (575 metric tons) in Washington.84 Depending on
Primary routes of exposure are through consumption,
the crop being produced, pesticides often are applied
absorption through the skin, and inhalation of pesticide
several times during the year; some crops may require
vapors when the chemicals are applied. Wildlife that nest
the use of pesticides every 10 days to 2 weeks throughout
or feed in recently sprayed crops may be subject to primary
the growing season. Furthermore, pesticides are often
routes of exposure. Animals may consume pesticides
applied in mixtures or cocktails of •2 chemicals. Because
directly, e.g., through consumption of contaminated
they are widespread and toxic, pesticides have the
vegetation, rodenticide baits, or pesticides granules as grit.
potential to adversely affect wildlife throughout both
Secondary exposure to pesticides occurs when wildlife
states. The environmental impact of pesticides is an
consume other animals that have been exposed. Birds and
extensive subject that would require an entire book for
other wildlife are attracted to insects dying on the soil
complete coverage. I provide only a cursory review of
surface after insecticide treatments, and may be exposed
some of the main points in this section.
when they eat these insects or feed them to their young.
The U.S. Environmental Protection Agency (EPA)
Secondary poisoning can be a concern for some predators
attempts to minimize the effects of pesticides on wildlife
that consume rodents poisoned by rodenticides.
through implementation of the Federal Insecticide,
Effects of pesticides on wildlife can be both direct and
Fungicide, and Rodenticide Act of 1988 (FIFRA), which
indirect. Direct effects include both lethal and sublethal
sets regulations on the acceptance and use of chemical
exposures. Not all wildlife exposed to pesticides receive a
pesticides. Even with these regulations, some chemicals
lethal dose and immediately die; smaller sublethal doses
are used that result in detrimental effects to nontarget birds
may be more common, but typically are difficult to verify.
and mammals.41 One criterion originally required by
Sublethal doses can lead to changes in behavior, weight
FIFRA was to field test certain pesticides prior to
loss, impaired reproduction or predator avoidance, and
registration. However, in 1992 the regulations were revised
inability to thermoregulate.31 Pesticides can indirectly
and field tests were no longer required58, 87 (L. Fisher, U.S.
affect wildlife by altering habitat and reducing food
Environmental Protection Agency, administrative
resources. Although less toxic to wildlife, herbicides can
memorandum). One reason for this revision was the cost
 CHAPTER 13: AGRICULTURAL ENVIRONS 351

have profound impacts on wildlife by altering habitat. Some wildlife damage occurs in almost all agricultural
Many seed-producing “weed” species are important crops and in facilities, production equipment, and systems.
components of wildlife diets. Furthermore, weed control A thorough review of the types of damage caused by
can destroy insect habitats and result in less diverse insect wildlife is beyond the scope of this chapter, but is
communities, which in turn may reduce the availability presented in Hygnstrom et al.36 Big game cause substantial
of some important foods for some wildlife.80 Herbicides damage in hay, pasture, alfalfa, and grain crops, especially
may also modify wildlife cover and reduce vegetation in the eastern half of both states. Waterfowl, particularly
complexity. Widespread use of herbicides in fence rows, Canada geese, cause widespread losses to pastures and
field borders, roadsides, and shelterbelts to control weedy grass seed fields during winter. Passerines damage a wide
vegetation has reduced the value of these habitat areas variety of fruit, nut, and grain crops in both states. Rodents,
for wildlife. primarily voles, house mice, ground squirrels, and pocket
gophers, damage alfalfa, hay, grain, fruit, and nut crops.
Fertilizers and Livestock Waste. Modern agricultural
Mice and rats cause significant damage to stored
practices depend heavily on the application of fertilizers
agricultural products, especially grains, in both states.
to maintain soil productivity and to grow many of the
Rodents also damage irrigation systems, and mounds of
crop varieties that have high nutrient demands. Hundreds
dirt from burrowing rodents can damage harvesting
of million pounds of fertilizer are used in Oregon and
equipment. Predators, primarily coyotes and mountain
Washington annually. For example, in 1997, Oregon wheat
lions, cause losses to livestock operators in both states.
farmers used 37,650 tons (34,150 metric tons) of nitrogen
Managing animal damage problems is a difficult and
and 2,350 tons (2,130 metric tons) of phosphate fertilizers,
controversial subject. Biologists from the wildlife agencies
while Washington wheat farmers used 72,700 tons (65,940
in both states and the USDA Wildlife Services spend
metric tons) of nitrogen and 8,250 tons (7,480 metric tons)
substantial amounts of time and agency resources in
of phosphate.84 Fertilizers cause few direct impacts on
resolving these problems. Animal damage specialists have
wildlife, but indirectly impact aquatic organisms because
a wide array of options in their tool kits that generally fall
of reduced water quality. Nonpoint source (NPS) pollution
in the categories of habitat modification, alternative crop
from fertilizers may adversely impact amphibian
production systems, hazing and scaring, repellents, and
reproduction, but that has not been documented.
population control. Specific control techniques for each
Phosphorus and nitrogen are the primary elements in NPS
wildlife species are presented in Hygnstrom et al.36
pollution in agricultural areas. Phosphorus pollution
Although often discussed, birth control is generally not
causes eutrophication of lakes and streams. The rapid
an option because of expense and ineffective delivery
growth of algae in response to phosphorus can limit
systems.28 Lethal forms of control receive increased
oxygen availability in aquatic systems and cause fish die-
scrutiny in the highly urbanized areas in both states, and
offs, and presumably kills developing eggs and larvae of
management agencies are continually searching for and
amphibians. Nitrogen is primarily a human health
testing alternative control methods. However, regardless
concern.
of the public’s perception of animal damage management,
Livestock waste in agricultural habitats also may result
it will always be a necessary component of wildlife
in water quality problems. Livestock confinement
management in Oregon and Washington, where we have
operations such as dairies and feedlots produce massive
both intensive agricultural production and abundant and
amounts of waste that can result in reduced water quality
diverse wildlife communities.
where wastes are improperly stored or recycled. Livestock
waste disposal during the past 20 years has moved from
being primarily a problem with lack of or improper storage Managing Agricultural
to one of inadequate land base for efficient reuse of manure Habitat for Wildlife
nutrients.49 Although agricultural habitats have high wildlife diversity
on a statewide basis, diversity is typically low at the farm
Animal Damage level, and opportunities abound for enhancing wildlife
Wildlife in agricultural habitats cause significant damage habitats and increasing wildlife diversity on farms and
to agricultural crops and production systems. This damage ranches. State and federal agencies in both states have
may strain relationships between landowners and wildlife numerous assistance programs (Table 2) to help
and habitat management agencies, and may cause some landowners manage their properties for improved
landowners to be reluctant to improve wildlife habitat on environmental quality. Interested landowners should
their properties. Loss to wildlife damage represents a contact the agency in charge of the program. All of these
significant portion of the agricultural production in programs are voluntary and contain varying levels of
Oregon and Washington. According to a 1997 survey incentives or disincentives to encourage participation.
conducted by the Oregon Agricultural Statistics Service,83 However, biologists working with private landowners
wildlife caused Oregon farmers and ranchers $158 million must recognize that successful programs for enhancing
in damage to crops, livestock, injury, and prevention wildlife habitats on private lands will depend on the
expenses. Approximately 47% of Oregon’s farms reported landowners’ personal interest and sense of stewardship.48
some type of wildlife damage; this loss represents 3.3% of The level of comfort and participation in habitat
the total value of agricultural production in Oregon.83
 352 Wildlife–Habitat Relationships in Oregon and Washington

Table 2. Landowner assistance programs Table 3. Suggestions for managing odd areas on
administered in 1999 by federal agencies in Oregon farms or ranches.61, 78
and Washington that aid private landowners in
managing properties for improved environmental Protect from fire and grazing, or if grazed, control timing and
quality. intensity to prevent damage or loss of shrubs and trees.
Develop food plots adjacent to odd areas.
Plant fruit or mast-producting shrubs and trees—cherry,
USDA Natural Resources Conservation Service and
hawthorne, wild plum, wild pear, serviceberry, crabapple, and
Farm Services Agency
oaks are good candidates depending on the site or region.
The Conservation Farm Option—a pilot program intended to
Enlarge odd areas during crop rotation by adjusting field
foster innovative conservation in whole-farm management.
boundaries.
Conservation of Private Grazing Land—provides technical
Plan for connectivity among odd areas and other habitat patches.
assistance for conservation needs on pasture, hayland, and
Manipulate vegetation with controlled fire, mowing, or a brush-
range.
hog to stimulate new growth of understory and enhance
Conservation Reserve Program—a land retirement program
structural diversity.
that establishes permanent grass or tree cover on
In areas where snow drift or high winds may be a problem,
environmentally valuable cropland or pasture.
develop the site as a shelterbelt or windbreak to provide
Environmental Quality Incentives Program—provides incentives
cover needed.
for addressing conservation needs on cropland or areas used
Add brush piles from pruning of shrubs and trees.
for livestock operations.
Flood Risk Reduction Program—encourages less intensive uses potential sites for habitat development. Many odd areas
of frequently flooded cropland. may already be good habitat and need little in the way of
Highly Erodible Land Conservation Compliance Program— enhancement. Management of odd areas and plant species
requires participants who receive USDA assistance to used to enhance wildlife habitat depends on the region
implement a soil conservation system on highly erodible
(i.e., what plants are adapted to soil and climate, and what
farmed land.
is available) and the type of site. Snyder78 and Payne and
Resource Conservation and Development Program—helps local
Bryant61 suggested a number of practices that landowners
groups plan and conduct rural development and natural
resource conservation activities.
might consider implementing on odd areas (Table 3).
Water Bank Program—compensates landowners for protecting Typically, the landowner’s desire for particular wildlife
regionally important migratory bird wetlands. species and other habitats available on a farm would guide
Watershed Protection and Flood Prevention Program—a cost- what practices are used.
share program to implement watershed work plans that
protect watershed functions. Food Plots
Wetland Conservation Compliance Program—a disincentive Food plots are sites cultivated specifically to provide food
program that links wetland conservation to receipt of USDA for wildlife species, and have been widely recommended
farm program benefits. by wildlife agencies around the country. Food plots can
Wetland Reserve Program—a land-retirement program for be used to supplement natural food supplies where they
restoring and protecting wetlands. are likely to be limited, or designed to attract wildlife to
Wildlife Habitat Incentives Program—assists landowners to plan specific areas for viewing. They are also commonly used
and pay for habitat improvements associated with farming in conjunction with recreational hunting or fee hunting.
operations. Food plots may lead to better over-winter survival and
U.S. Fish and Wildlife Service spring conditioning for egg laying. For example, Robel et
Partners for Wildlife—a cost-share program for enhancing al.68 showed that bobwhite quail with access to food plots
wildlife habitats, especially wetlands. during winter in Kansas maintained higher body weights
and fat reserves, both of which are assumed to lead to
enhancement programs will vary from landowner to better over-winter survival and early conditioning for egg
landowner. Significant changes in wildlife habitats on production.
private lands may take many years and will depend on Food plots are not preferred to other forms of habitat
local biologists building trust with private landowners. management, but are typically used by landowners who
wish to rapidly enhance wildlife habitats on their
Odd Areas properties. Payne and Bryant 61:436 listed several
Odd areas are areas on farms that are not cultivated disadvantages to food plots: (1) they concentrate wildlife,
because of location, topography, erosion, geomorphology, thereby increasing vulnerability to disease and predation;
or soil conditions. Odd areas typically include sites such (2) food plots attract undesirable wildlife and feral animals
as rock piles, rock outcrops, borrow and gravel pits, eroded (i.e., hogs, starlings, or other damage-causing species); (3)
areas in fields, bare knobs, sinkholes, gullies, and good they represent a direct expense to the farmer; (4) food plots
cropland cut off from the rest of the field by a stream, ditch, may result in high concentrations of wildlife or
gully or other obstruction. 61 Center-pivot irrigation populations above carrying capacity; and (5) in arid
systems typically leave odd areas between the irrigated regions plantings are likely to fail during drought years,
fields. Most farms have some odd areas, which represent and are probably unneeded during wet years.
 CHAPTER 13: AGRICULTURAL ENVIRONS 353

Table 4. Metabolizable energy in seeds of plants Food plants recommended for wildlife vary from region
commonly used for food plots to region like the wildlife species the plots are intended to
for seed-eating birds. benefit, but grain sorghum, corn, legumes, and cereal
grains are the most common. Payne and Bryant61 listed
Seed Metabolizable energy content (kcal/g) plants commonly used for food plots for several wildlife
species. Sorghum is typically ranked over corn because it
is easier to establish in areas without irrigation, the smaller
Excellent (>4.14 kcal/g)
seed is acceptable to more wildlife species, and because it
Giant ragweed 4.32
has energy content similar to corn. Energy content of the
Good (3.49-4.14 kcal/g) more common wildlife food plants varies substantially
Western ragweed 3.88 (Table 4).
Corn 3.87 The size, shape, location, and distribution of food plots
Soybean 3.78 are as important as the particular plant foods that are
Sunflower 3.65 cultivated. Size depends on the target wildlife species. For
Sorghum 3.59 most birds and small game species, plots that are 0.5-5
Low (2.61-3.48 kcal/g) acres (0.2-2.0 ha) are usually sufficient. Ungulates may
German millet 3.47 require 10- to 20-acre (4- to 8-ha) plots or larger, depending
Prostrate lespedeza 3.42 on the size of the herd being fed. In general, food plots are
Korean lespedeza 3.14 linear, which maximizes edge, access, and interspersion
Wheat 3.06 with other habitat types, and distributes animals to reduce
Thistle 2.70 vulnerability to disease and predation. Width of these
Shrub lespedeza 2.69 linear strips also depends on the target species. For most
small animals 10-20 feet (3-6 m) is wide enough. Often,
Poor (<2.61 kcal/g)
this is limited more by the equipment used to cultivate a
Partridgepea 2.42
Smartweed 2.30 food plot. Leaving 4-5 rows of a crop unharvested is a
Multiflora rose hips 2.02 commonly recommended practice that leaves a strip
Switchgrass 1.86 approximately 10-20 feet (3-6 m) wide depending on the
Smooth sumac 1.48 crop and harvesting equipment. Payne and Bryant61
suggested that plots should not exceed 328 feet (100 m)
After Robel et al.,67 used with permission. for ungulates and 164 feet (50 m) for some medium to
small animals.
The spacing and distribution of food plots is another
consideration. Usually only 1-3% of an area might need
to be intensively managed for food plots. 61: 434 The
distribution of food plots depends on the mobility of the
target species and constraints imposed by normal farm
operations. Typically, recommendations have been
developed only for game species. For pheasants, one 3.7-
acre (1.5-ha) plot for every 123 acres (50 ha) would insure

Figure 9. Deer use

of food plots will be
greatest in the
optimal use area
within 410 feet
(125 m) of cover;
food plots >655 feet
(200 m) from cover
will receive little
use by deer (after
Leckenby et al.47).
 354 Wildlife–Habitat Relationships in Oregon and Washington

that a food plot was always within 1,640 feet (500 m). For Yahner99 suggested that rows should be 16 feet (5 m) apart
deer, two 15- to 20-acre plots per 1,235 acres (6-8 ha per to allow for herbaceous and shrub development in the
500 ha) has been recommended.61: 434 understory.
Location of food plots relative to other habitat attributes Planting configuration may also be important for
is most important; proximity to cover is a prime wildlife. Plantings that are irregularly spaced or that are
consideration. For example, deer will be unlikely to use not straight would increase the amount of edge habitat
food plots >738 feet (225 m) from cover (Figure 9). Geese and would reduce line-of-sight distances from predators
and waterfowl, conversely, will be less likely to use food and other sources of disturbance.57
plots that are within 82-164 feet (25-50 m) of cover that
might conceal predators. In areas with high winds, food Hedge and Fence Rows
plots are more likely to be used if they are on the leeward Plant species used in fence and hedge rows vary from
side of cover. region to region and should be adapted to local climate
and soil conditions. Multiple species within fence and
Shelterbelts and Windbreaks hedge rows will increase wildlife diversity.59 Species that
The management of shelterbelts and windbreaks will vary, produce fruit or berries and that have dense growth forms
depending on the location and the purpose of the are good candidates for fence rows. In Oregon and
shelterbelt. Typically, shelterbelts are designed and planted Washington, wildrose (Rosa spp.), wild pear, hawthorn
for specific conservation purposes other than wildlife (Crataegus spp.), blackberry, and cherry (Prunus spp.) are
habitat enhancement (i.e., reduced soil erosion, visual or commonly used. However, in some areas these species
noise screening, snow fences, livestock protection). may be considered invasive weeds.
However, because shelterbelts provide important Hedge and fence rows can be planted or allowed to
structural and cover components in otherwise structurally grow from natural regeneration. If planted, success is best
simple landscapes, these habitat features can greatly where high-quality planting stock is used. Where possible,
enhance wildlife diversity and abundance on a farm. hedge and fence rows should be at least 10 feet (3 m) wide.
Wildlife managers and landowners should expect about Creating mosaics through selective mowing and
20 years before a shelterbelt begins to function in the woodcutting, and retaining 1.7-2.4 snags (>18 inches
manner for which it was designed. However, these sites diameter at breast height [dbh], 33-49 feet tall)/mile (1-2
will be used by wildlife immediately. snags [>46 cm dbh, 10-15 m tall]/km) of fence row have
Numerous plant species have been used for shelterbelt also been suggested for hedge and fence row
plantings; they should be suitable for the purpose and management.76 These sites, like shelterbelts, will need to
adapted to soil and climate. The USDA maintains a website be protected from fire and grazing. O’Connor and Shrubb59
(http://www2.nrcs.usda.gov/Netdynamics/Vegspec/ suggested that the least amount of hedge row
pages/HomeVegspec.htm) with software that helps users management as possible will make the most favorable
identify plant materials for use in restoration efforts. The conditions for wildlife. Hedge and fence rows can be
species that should be planted will vary from region to created relatively easily by plowing or disking the site
region depending on availability of seed or stock. Species where they are to be created, placing fence posts in a
used should be tolerant of herbicides that may be used in staggered configuration, and running a wire or string for
areas being protected. Planting a variety of species is also perching by birds. Seeds in the bird droppings rapidly
important because it reduces the likelihood of losing the establish preferred plants in the plowed or disked areas.
shelterbelt to a species-specific pest (e.g., Dutch elm
disease), and the greater structural diversity will increase Field Borders
wildlife diversity.75, 99 Payne and Bryant61 developed a list Field borders are typically seeded with fast growing
of plants commonly used in shelterbelts. Because many grasses or legumes that are adapted to the site. Field
sites where shelterbelts are desired have severe climatic borders should be at least 16 feet (5 m) wide. In areas where
conditions (low precipitation, high winds), only high- they function as filter strips around wetlands and riparian
quality planting stock should be used. It may be necessary areas, they should be 20-98 feet (6-30 m) wide. Field
to reduce competition from other plants during the borders may be hayed for forage, but as in all haying
establishment phase by using herbicides or mechanical operations where wildlife are a concern, haying should
means. In arid regions, supplemental water may also be be delayed as late as possible to avoid the breeding season
required during the establishment phase. Shelterbelts also (usually after 15 July). Field borders, like most grass or
need to be protected from fire and grazing. grass/legume mixtures, occasionally need to be mowed
Shelterbelt width depends on the number of rows of to maintain plant vigor or to prevent invasion by woody
plantings, and typically the number of rows used in plants. Maintenance mowing of this type can be done once
conservation practices depends on the thermal protection every two or three years. Another consideration for both
required and the need to control snow drift. Podol64 haying and mowing is to allow some regrowth before the
suggested that any windbreak <108 feet (33 m) wide in fall/winter dormant period. Weeds are a common concern
northern areas would not create snow-free areas in winter among farmers that have herbaceous field borders. Many
and would have little value for winter cover. For wildlife of the “weed” species are actually good sources of food
purposes, the wider the better. When planting rows, for wildlife. However, most farmers will want to control
 CHAPTER 13: AGRICULTURAL ENVIRONS 355

invasive weeds that are likely to interfere with crop where roads intersect traditional migration routes or big
production. Weeds in field borders can usually be game winter ranges. For some less mobile species such as
controlled with spot application of herbicides. reptiles and amphibians, roads may be a substantial barrier
to movement. Construction of wildlife-proof fences or
Roadsides underpasses have proven to be effective for reducing the
Because of the prevalence of roadsides in both states, number of wildlife-vehicle collisions. In cases where the
management of these habitats can profoundly affect road is a barrier to movement or along a migration route,
wildlife. Roadside management varies a great deal underpasses allow these movements to occur relatively
depending on jurisdiction. The DOTs in Oregon and unhindered if they are correctly located.77, 91
Washington are charged with management of interstate,
U.S., and state highways. County governments typically Reducing Pesticide Impacts
manage county roads, while farmers manage the roads Reducing the effects of pesticides on wildlife can be
on their own property. Most roadsides managed under accomplished in a number of ways. Reducing the total
the first two jurisdictions are managed with safety as a amount of pesticides is a general goal of alternative or
primary concern. Roadside vegetation is mowed to reduce sustainable agriculture56 that would benefit wildlife.
visual obstructions, and in some cases to discourage Adopting integrated pest management (IPM) strategies
wildlife, especially deer, use adjacent to the road. Many has been widely promoted as a means of reducing the
roads on private lands do not have roadside vegetation. reliance on pesticides.56 IPM is a crop management system
Rather, the field is plowed or cultivated right up to the that includes all aspects of crop production in developing
road. Encouraging private landowners to leave a 16-feet strategies for reducing crop damage from pests and
(5-m) uncultivated strip of vegetation on each side of a disease. These strategies might include but are not limited
road will provide wildlife habitat as well as other to genetic engineering of the crop, residue management,
conservation benefits including reducing sedimentation tillage practices, fertilizer management, pest monitoring,
in streams and off-road movement of petrochemicals. pesticide applications, and introduction of pest predators.
Warner et al.92 noted the most attractive roadsides for In most cases, IPM will result in a reduction in the amount
pheasant nesting were well established areas seeded with of pesticides applied.56 In some cases, wildlife predators
a grass-legume mix. Roadsides on private lands are might be part of the IPM strategy. For example,
typically managed to prevent encroachment of woody encouraging raptor use of crop fields may help reduce
species and to control invasion by noxious weeds. rodent populations.34, 40, 96
Because of the linear nature of roadsides and the regular Chemical selection is another way to reduce pesticide
mowing, these areas may become ecological traps.6, 14 Birds impacts on wildlife. Several pesticides are usually
nesting in roadsides, as in other linear habitats, may suffer available for controlling a particular pest in a particular
high rates of predation. Bergin et al.6 reported 23% nest crop. Most farmers select the cheapest chemical that gets
loss overall on artificial nests, but predation varied from the job done, but many are willing to pay more for a
area to area. Furthermore, they reported significantly chemical that would cause fewer environmental impacts.
greater predation on the backslope than the foreslope (30% Pesticides vary considerably in their toxicity (Table 5) and
vs. 16%). Backslopes may either have more predators persistence. Selecting the chemical with the lowest toxicity
moving through them, or predators preferentially search to the group of animals that might be exposed to it and
backslopes. Foreslopes in their study were more likely to chemicals with shorter half-lives would help reduce
be mowed (12% vs. 0%), and the shorter vegetation or pesticide impacts on wildlife. The pesticide information
mowing activity might make predators wary of searching profile system (http://ace.orst.edu/info/extoxnet/pips/
these habitats. Mowing also directly destroys nests. ghindex.html) is an easy way to determine the relative
Because of these concerns mowing should be delayed until toxicity and the persistence of pesticides that a farmer may
mid-July where feasible, to allow successful nesting by consider using.
ground-nesting birds and small mammals. In most cases, Controlling the timing and location of a pesticide
one-pass management is sufficient to address safety and application is another means by which impacts on wildlife
woody vegetation encroachment concerns. One-pass can be reduced. Where flexibility is possible, applying a
management involves a single-mower-width pass on the pesticide when wildlife are not in the area would reduce
foreslope adjacent to the road each year, beginning after wildlife exposure to the toxicant. For example, waterfowl
mid-July. Woody vegetation on the remainder of the commonly use croplands during the short spring
foreslope and the backslope can be controlled by mowing migration period. If a pesticide application can be delayed
once every two to three years. Invasive weeds along the until after the birds have moved through the area, there
roadsides in all jurisdictions can usually be controlled with would be fewer potential impacts. This type of flexibility
spot application of herbicides. is often not possible. An alternative is to avoid the use of
Vehicle collisions with deer cause over $1 billion in pesticides where wildlife are known to occur. Habitat
damage and >200 human fatalities annually in the U.S.,19 features such as shelterbelts, fence rows, field borders, and
and can be an important source of mortality for some odd areas should be protected from pesticides with a no-
wildlife populations. Although such collisions occur spray buffer zone of at least 16 feet (5 m) in the crop field
throughout both states, problems are most pronounced adjacent to the habitat (Figure 10). Wetlands and riparian
 356 Wildlife–Habitat Relationships in Oregon and Washington

Table 5. Toxicities to birds, mammals, and fish of insecticides and miticides

used on fruit crops.

Pesticide (brand name) Birdsa Mammals a Fishb

azinphos-methyl (Guthion) H H EH
carbaryl (Sevin) L L H
chlorpyrifos (Lorsban) H L EH
diazinon (Diazinon) Hc M EH
dicofol (Kelthane) L L EH
dimethoate (Defend) Hc M M
dinocap (Karathane) M L EH
endosulfan (Thiodan) M H EH
esfenvalerate (Asana) L L EH
formetanate (Carzol) M H M
fenbutatin-oxide (Vendex) L L EH
malathion L L EH
methidathion (Supracide) H M EH
methomyl (Lannate) H H H
methyl parathion (Penncap-M) Hc H H
oxamyi (Vydate) H H M
oxythioquinox (Morestan) — L EH
permethrin (Ambush, Pounce) L L EH
phosmet (Limidan) L M EH
propargite (Omite) L L EH
After Palmer and Bromley,60 used with permission.
Wildife hazard is based on the following toxicities: H (highly toxic) = LD50 <30 mg/kg and LC50
a

<500 ppm; M (moderately toxic) = LD50 >30 and <100 mg/kg and/or LC50 >500 and <1,000
ppm; L (low toxicity) = LD50 >100 mg/kg and LC50 >1,000 ppm; NT (not toxic).
b
Fish 96-hour LC50 toxicities are as follows: EH (extremely toxic) <0.1 ppm; H (highly toxic)
0.1-1.0 ppm; M (moderately toxic) 1.0-10.0 ppm; L (low toxicity) >10.0 ppm.
c
Active ingredient (not necessarily a specific product) has caused wildlife deaths.

Figure 10. Protect

sensitive habitat such as
wetlands, shelterbelts,
fencerows, etc., from
pesticides with a no-
spray buffer (dashed
line). A better
alternative would also
include a permanent
grass field border that
would prevent
movement of chemicals
during runoff.
(Photograph by W.
Daniel Edge)
 CHAPTER 13: AGRICULTURAL ENVIRONS 357

areas should be protected with buffer zones composed of farmland to native grassland habitat, or vice versa. This
grass and herbaceous vegetation to reduce the amount of query generated a list of 241 species that are associated
chemicals (pesticides and fertilizers) and sediment that with the two habitats in the two-county area.
reaches the water during periods of runoff. Depending Approximately 42% (101) of the species occur in
on the steepness of the slope adjacent to wetlands and agricultural habitat and not in Westside Grasslands, but
riparian areas, buffer zones of up to 98 feet (30 m) may be only 4 species are exclusive to Westside Grasslands (Table
required. Grassed waterways in crop fields are 6). Without consideration of the abundance of these two
recommended for preventing chemical runoff from habitats and the ecological role of individual species
reaching water sources. associated with the habitats, one might easily conclude
Controlling chemical drift is another means of reducing that conversion of Westside Grasslands to agricultural
the impacts of pesticides on wildlife. Chemical drift is the habitat would generally be beneficial to more species of
movement of a chemical off the intended site of wildlife. It is difficult to separate Westside Grasslands from
application, and is a principal cause of wildlife exposure unimproved pastures with remote sensing data, and
to pesticides. Both physical (i.e., chemical droplets) and therefore, an accurate estimate of the land area covered
vapor drift may occur when a pesticide is applied, by these two habitats in Linn and Benton counties is not
depending primarily on wind velocity. Pesticides should possible. However, agricultural habitat makes up >640
not be applied when wind speeds exceed 5 mph (8 kph) miles2 (1,657 km2) of the land area in these two counties,
or when there are gusts of up to 10 mph (16 kph), and whereas unimproved pastures account for about 80 miles2
never when sensitive habitats or wildlife are down wind. (207 km2); Westside Grasslands are only a small proportion
Droplet size is another major factor in chemical drift. of that (<5%) (T. O’Neil, Northwest Habitat Institute,
Modern spray nozzles are designed to produce specific personal communication). Furthermore, the species
droplet sizes with a particular pressure. Using the most exclusively associated with agricultural habitats are exotic
modern spray equipment that is properly calibrated will species (e.g., house sparrow, house mouse, Norway rat),
help reduce the amount of drift. or more often are associated with some special habitat
features, such as ephemeral wetlands (e.g., shorebirds and
waterfowl) or shelterbelts and fence or hedge rows (e.g.,
Case Study vireos and warblers).
The data matrixes in this book provide a powerful tool
for assessing the impacts of management practices and
landuse policies on wildlife populations. However, Research Needs in
predicting changes in wildlife communities that result Agricultural Habitats
from changes in agricultural habitats is problematic A review of the literature cited section of this chapter
because of the widespread distribution of agricultural reveals that little research related to wildlife in agricultural
habitats throughout the two states and because many ecosystems has been conducted in Oregon or Washington;
wildlife species are only associated with agricultural most research on this topic has been conducted in the
habitats for feeding or occur in those habitats because of Midwest. Although the relationships I report in this
habitat features (e.g., shelterbelts, field borders, ephemeral chapter are probably valid in our area, there are many
wetlands, etc.). Such predictions should be considered research questions that will need to be addressed in order
only for relatively small areas within the two states (i.e., to effectively manage wildlife on farms and ranches. These
one to a few counties) and should consider the ecological research questions include responses of wildlife to crop
roles of the species in those habitats. management practices, integrated pest management,
As an example, I queried the database for the species landscape design, human dimensions, and the impacts of
that occur in Agricultural and Westside Grassland habitats agricultural policy on wildlife resources.
in Linn and Benton counties, Oregon. Such a query might Most crops can be produced with several alternative
be used to assess the impact on wildlife communities of crop management systems, and each system will provide
agricultural policies encouraging farmers to convert a different array of wildlife benefits and costs. The growing

Table 6. Wildlife habitat association matrix for wildlife species that are found in western grassland and
agricultural habitats in Linn and Benton counties, Oregon.

Westside Grasslands
Closely associated Associated Present Does not occur

Agricultural
Closely associated 8 21 7 15
Associated 2 58 13 48
Present 0 13 14 38
Does not occur 0 2 2
 358 Wildlife–Habitat Relationships in Oregon and Washington

interest in sustainable or alternative agriculture among Human dimensions research is needed to identify a
farmers will likely produce additional management broad range of social and economic factors that affect how
systems providing additional opportunities for wildlife farmers manage their properties. Research needs in this
to exist on farms. Residue management and tillage area cover the gamut of the people side of wildlife
practices are two areas of crop management that directly management. Farmers’ perceptions regarding wildlife
affect the year-round value of croplands to wildlife, and damage97 and habitat enhancement practices have been
additional research is needed to identify those practices documented,18 but research is also needed to identify
that are most beneficial to wildlife.9, 70 Because of the many alternative management systems that might be adopted
agricultural commodities produced in the two states, we on broad scales. As farmers and ranchers move to diversify
will be unable to document wildlife responses to operations, there is likely to be an increasing interest in
alternative practices for most crops. Therefore, research wildlife-based alternative income opportunities such as
should focus on the most broadly produced crops, such fee hunting and ecotourism.4, 12 Although some aspects of
as wheat, grass seed, hay pastures, and orchard fruits and fee hunting have been documented in Oregon and
vineyards. Washington,12, 39 the demand and willingness to pay for
Integrated pest management is another component of ecotourism in the two states has not been determined.
crop management systems where little research has been Finally, human dimensions research is needed to identify
done on the direct effects on wildlife. Methods for processes and mechanisms by which successful
managing animal damage to crops, ways of reducing conservation practices are implemented at the watershed
impacts of pest management on wildlife, and a better scale.33
understanding of how wildlife can be used to control crop The final area of research is the impact of agricultural
pests all need to be examined. Wildlife damage to crops is policy on wildlife resources. USDA farm programs may
a perennial problem that will become more complicated directly affect how farmers manage their properties from
with the increasing concern for animal welfare. Alternative local to regional scales.30 Land retirement programs such
management systems that discourage wildlife use of as the Conservation Reserve Program can affect wildlife
commonly damaged crops need to be researched. For diversity and abundance on broad spatial scales.8, 52
example, living mulches in orchards, vineyards, and other Continued support of these programs in the next Farm
crops help to control runoff and invasive weeds, but Bill and when the 10-year contracts expire will depend on
depending on which grass species is used and how the documenting their wildlife and ecological benefits.
mulch is managed, damage by voles may vary
substantially. IPM also includes the use of pesticides, and Literature Cited
although a great deal is known about wildlife response to 1. Adams, W. L., and A. D. Geis. 1979. Roads and roadside habitat in
individual pesticides based on laboratory studies, relation to small mammal distribution and abundance. Pages 54-1 to
54-17 in: D. Arner and R. E. Tillman, editors. Environmental concerns in
additional studies are needed to determine population-
rights-of-way management. EPRI WS-78-141, Electric Power Research
level effects under field conditions.25, 82 The effects of using Institute, Palo Alto, CA.
mixtures of pesticides on wildlife are virtually unknown. 2. Airola, D. A. 1986. Brown-headed cowbird parasitism and habitat
Finally, wildlife eat many arthropod and vertebrate pest disturbance in the Sierra Nevada. Journal of Wildlife Management 50:
species and can be part of the pest management strategy. 571-575.
Encouraging predators to use crop fields, and the 3. Barbour, R. W., and W. H. Davis. 1969. Bats of America. University Press
effectiveness of predators in controlling crop pests need of Kentucky, Lexington, KY.
4. Benson, D. E. 1998. Enfranchise landowners for land and wildlife
additional research. For example, Wolff et al.96 documented stewardship: examples from the western United States. Human
an 11-fold increase in American kestrel use of grass fields Dimensions of Wildlife 3: 59-68.
with supplemental perches, but increased use by kestrels 5. Beier, P., and R. F. Noss. 1998. Do habitat corridors provide
did not result in a decline in vole populations at high connectivity? Conservation Biology 12: 1241-1252.
densities. 6. Bergin, T. M., L. B. Best, and K. E. Freemark. 1997. An experimental study
Understanding how agricultural production affects of predation on artificial nests in roadsides adjacent to agricultural
habitats in Iowa. Wilson Bulletin 109: 437-448.
wildlife at the landscape scale is another important
7. Best, L. B. 1986. Conservation tillage: ecological traps for nesting birds?
research need. The abundance and distribution of crops Wildlife Society Bulletin 14: 308-317.
and habitat elements such as shelterbelts, riparian buffers, 8. ———, H. Campa III, K. E. Kemp, R. J. Robel, M. R. Ryan, J. A. Savidge, H.
fence rows, and field borders at watershed and regional P. Weeks, Jr., and S. R. Winterstein. 1997. Bird abundance and nesting in
scales have important implications for the long-term CRP fields and cropland in the Midwest: a regional approach. Wildlife
persistence of wildlife in agricultural landscapes. How and Society Bulletin 25: 864-877.
to what extent some of these habitat features are used for 9. ———, K. E. Freemark, J. J. Dinsmore, and M. Camp. 1995. A review and
synthesis of habitat use by breeding birds in agricultural landscapes of
dispersal is poorly understood.5 Habitat elements may
Iowa. American Midland Naturalist 134: 1-29.
become ecological traps and function as population sinks 10. ———, R. C. Whitmore, and G. M. Booth. 1990. Use of cornfields by
depending on their abundance and distribution within a birds during the breeding season: the importance of edge habitat.
landscape. Determining how predation and brood American Midland Naturalist 123: 84-99.
parasitism are influenced by the spatial and temporal 11. Brittingham, M. C., and S. A. Temple. 1983. Have cowbirds caused
distribution of habitat components is a vital research need forest songbirds to decline? BioScience 33: 31-35.
in agricultural habitats.
 CHAPTER 13: AGRICULTURAL ENVIRONS 359

12. Butler, L. D., and J. P. Workman. 1990. A comparison of fee hunting on 38. Johnson, R. G., and S. A. Temple. 1990. Nest predation and brood
private rangelands in central Oregon and the Texas Trans Pecos. Pages parasitism of tallgrass prairie birds. Journal of Wildlife Management 54:
269-276 in: L. A. Renecker and R. J. Hudson, editors.Wildlife production: 106-111.
conservation and sustainable development. University of Alaska 39. Johnson, R. L. 1989. Market fee hunting opportunities in the presence
Extension Service Miscellaneous Publication 91-6, Fairbanks, AK. of abundant public land. Western Journal of Applied Forestry 4: 24-26.
13. Camp, M., and L. B. Best. 1993. Bird abundance and species richness in 40. Kay, B. J., L. E. Twigg, T. J. Korn, and H. I. Nicol. 1994. The use of artificial
roadsides adjacent to Iowa rowcrop fields. Wildlife Society Bulletin 21: perches to increase predation on house mice (Mus domesticus) by
315-325. raptors. Wildlife Research 21: 95-106.
14. ———, and ———. 1994. Nest density and nesting success of birds 41. Kendall, R. J., and J. Ackerman. 1992. Terrestrial wildlife exposed to
in roadsides adjacent to rowcrop fields. American Midland Naturalist agrochemicals: an ecological risk assessment perspective.
131: 347-358. Environmental Toxicology and Chemistry 11: 1727-1749.
15. Candelaria, L. M., and M. K. Wood. 1981. Wildlife use of stockwatering 42. ———, and T. E. Lacher, Jr. 1994. Wildlife toxicology and population
facilities. Rangelands 3: 194-196. modeling: integrated studies of agroecosystems. Lewis Publishers, Boca
16. Carson, R. 1962. Silent spring. Houghton-Mifflin, Boston, MA. Raton, FL.
17. Chilgren, J. D. 1979. Drowning of grassland birds in stock tanks. 43. Kindschy, R. R. 1986. Rangeland vegetative succession: implications to
Wilson Bulletin 91: 345-346. wildlife. Rangelands 8: 157-159.
18. Conover, M. R. 1998. Perceptions of American agricultural producers 44. ———. 1996. Fences, waterholes, and other range improvements.
about wildlife on their farms and ranches. Wildlife Society Bulletin 26: Pages 369-381 in: P. R. Krausman, editor. Rangeland Wildlife. Society for
597-604. Range Management, Denver, CO.
19. ———, W. C. Pitt, K. K. Kessler, T. J. DuBow, and W. A. Sanborn. 1995. 45. Krapu, G. L., and K. J. Reineke. 1992. Foraging ecology and nutrition.
Review of human injuries, illnesses, and economic losses caused by Pages 1-29 in: B. D. J. Batt, A. D. Afton, M. G. Anderson, C. D. Ankney, D.
wildlife in the United States. Wildlife Society Bulletin 23: 407-414. H. Johnson, J. A. Kadlec, and G. L. Krapu, editors. Ecology and
20. Cooperrider, A. 1986. Terrestrial physical features. Pages 587-601 in: A. management of breeding waterfowl and wetlands. University of
Y. Cooperrider, R. J. Boyd, and H. R. Stuart, editors. Inventory and Minnesota Press, Minneapolis, MN.
monitoring of wildlife habitat. U.S. Department of Interior, Bureau of 46. Kunz, T. H. 1982. Roosting ecology. Pages 1-46 in: T. H. Kunz, editor.
Land Management, U.S. Government Printing Office, Washington, D.C. Ecology of bats. Plenum Publication Corporation, New York, NY.
21. Cowardin, L. M., V. Carter, F. C. Golet, and E. T. LaRoe. 1979. 47. Leckenby, D. A., D. P. Sheehy, C. H. Nellis, R. J. Scherzinger, I. D. Luman,
Classification of wetlands and deepwater habitats of the United States. W. Elmore, J. C. Lemos, L. Doughty, and C. E. Trainer. 1982. Wildlife
U.S. Fish and Wildlife Service. FWS/OBS-79/31. Washington, D.C. habitats in managed rangelands—the Great Basin of southeastern
22. Craig, T. H., and L. R. Powers. 1976. Raptor mortality due to drowning Oregon—mule deer. U.S. Forest Service and Bureau of Land
in a livestock watering tank. Condor 78: 412. Management, General Technical Report PNW-139.
23. Darley, J. A. 1982. Territoriality and mating behavior of the male 48. Leopold, A. 1945. The outlook for farm wildlife. Transactions of the
brown-headed cowbird. Condor 84: 15-21. North American Wildlife Conference 10: 165-168.
24. Dufty, A. M., Jr. 1982. Movements and activities of radio-tracked 49. Logan, T. J. 1993. Agricultural best management practices for water
brown-headed cowbirds. Auk 99: 316-327. pollution control: current issues. Agriculture, Ecosystems and
25. Edge, W. D., R. L. Carey, J. O. Wolff, L. M. Ganio, and T. Manning. 1996. Environment 46: 223-231.
Effects of Guthion 2S on Microtus canicaudus: a risk assessment 50. Maser, C., J. W. Thomas, I. D. Luman, and R. Anderson. 1979. Wildlife
validation. Journal of Applied Ecology 33: 269-278. habitats in managed rangelands—the Great Basin of southeastern
26. ———, J. O. Wolff, and R. L. Carey. 1995. Density-dependent Oregon—man-made habitats. U.S. Forest Service and Bureau of Land
responses of gray-tailed voles to mowing. Journal of Wildlife Management, General Technical Report PNW-86.
Management 59: 245-251. 51. Mayfield, H. F. 1965. The brown-headed cowbird with old and new
27. Fitzgibbon, C. D. 1997. Small mammals in farm woodlands: the effects hosts. Living Bird 4: 128.
of habitat, isolation and surrounding land-use patterns. Journal of 52. McCoy, T. D., M. R. Ryan, E. W. Kurzejeski, and L. W. Burger, Jr. 1999.
Applied Ecology 34: 530-539. Conservation Reserve Program: source or sink habitat for grassland
28. Garrott, R. A. 1995. Effective management of free-ranging ungulate birds in Missouri? Journal of Wildlife Management 63: 530-538.
populations using contraception. Wildlife Society Bulletin 23: 445-452. 53. McKenzie, D. F. 1997. A wildlife manager’s field guide to the Farm Bill.
29. Gates, J. E., and L. W. Gysel. 1978. Avian nest dispersion and fledging Wildlife Management Institute, Washington, D.C.
success in field-forest ecotones. Ecology 59: 871-883. 54. National Academy of Sciences. 1981. Testing for effects of chemicals
30. Gerard, P. W. 1995. Agricultural practices, farm policy, and the on ecosystems. National Academy Press, Washington, D.C.
conservation of biological diversity. U.S. Department of the Interior, 55. National Audubon Society. 1996. Oases for wildlife: small and farmed
National Biological Service, Biological Science Report 4. wetlands. National Audubon Society, Washington, D.C.
31. Grue, C. E., W. J. Fleming, D. J. Busby, and E. F. Hill. 1983. Assessing the 56. National Research Council. 1989. Alternative agriculture. National
hazards of organophosphate pesticides to wildlife. North American Academy Press, Washington, D.C.
Wildlife and Natural Resources Conference 48: 200-219. 57. Norrgard, R. 1989. Woody cover plantings for wildlife. Minnesota
32. Haas, C. A. 1995. Dispersal and use of corridors by birds in wooded Department of Natural Resources, St. Paul, MN.
patches on an agricultural landscape. Conservation Biology 9: 845-854. 58. Norton, S. B., D. J. Rodier, J. H. Gentile, W. H. van der Schalie, W. P.
33. Habron, G. 1999. An assessment of community-based adaptive Wood, and M. W. Slimak. 1992. A framework for ecological risk
watershed management in three Umpqua Basin watersheds. assessment at the EPA. Environmental Toxicology and Chemistry 11:
Dissertation, Oregon State University, Corvallis, OR. 1663-1672.
34. Hall, T. R., W. E. Howard, and R. E. Marsh. 1981. Raptor use of artificial 59. O’Connor, R. J., and M. Shrubb. 1986. Farming and birds. Cambridge
perches. Wildlife Society Bulletin 9: 296-298. University Press, Cambridge, England.
35. Heckert, J. R. 1994. The effects of habitat fragmentation on mid- 60. Palmer, W. E., and P. T. Bromley. 1992. Wildlife and pesticides: fruit trees.
western grassland bird communities. Ecological Applications 4: 461-471. North Carolina Cooperative Extension Service, North Carolina State
36. Hygnstrom, S. E., R. M. Timm, and G. E. Larson, editors. 1994. University AG-463-7, Raleigh, NC.
Prevention and control of wildlife damage. University of Nebraska 61. Payne, N. F., and F. C. Bryant. 1994. Techniques for wildlife habitat
Cooperative Extension, University of Nebraska, Lincoln, NE. management of uplands. McGraw-Hill, Inc., New York, NY.
37. Johnson, D. H., and L. D. Igl. 1995. Contributions of the Conservation 62. Peterle, T. J. 1991. Wildlife toxicology.Van Nostrand Reinhold, New
Reserve Program to populations of breeding birds in North Dakota. York, NY.
Wilson Bulletin 107: 709-718.
 360 Wildlife–Habitat Relationships in Oregon and Washington

63. Pimentel, D. S., H. Acquay, M. Biltonen, P. Rice, M. Silva, J. Nelson, V. 85. U.S. Environmental Protection Agency. 1991. Endangered species
Lipner, S. Giordano, A. Horowitz, and M. D’Amore. 1992. Environmental protection program as it relates to pesticide regulatory activities.
and economic costs of pesticide use. Bioscience 42: 750-760. Report to Congress. EPA 540-09-91-120. EPA Office of Pesticides and
64. Podol, E. B. 1979. Utilization of windbreaks by wildlife. Pages 121-127 Toxic Substances, Washington, D.C.
in: Windbreak management workshop proceedings. Great Plains 86. ———. 1991. Federal manual for identifying jurisdictional wetlands,
Agriculture Council Publication 92. Lincoln, NE. proposed revisions. U.S. Environmental Protection Agency, U.S. Army
65. Rattner, B., and A. Fairbrother. 1991. Biological variability and the Corps of Engineers, U.S. Fish and Wildlife Service, and U.S. Soil
influence of stress on cholinesterase activity. Pages 89-108 in: P. Mineau, Conservation Service.Washington, D.C.
editor. Chemicals in agriculture,Volume 2. Cholinesterase inhibiting 87. ———. 1992. Framework for ecological risk assessment. EPA/630/R-
insecticides, their impact on wildlife and the environment. Elsevier, New 92/001. Washington, D.C.
York, NY. 88. Uresk, D. W., and K. Severson. 1988. Waterfowl and shorebird use of
66. Reese, K. P., and J. T. Ratti. 1988. Edge effect: a concept under scrutiny. surface-mined and unfenced livestock water impoundments on the
Transactions of the North American Wildlife and Natural Resources northern Great Plains. Great Basin Naturalist 48: 353-357.
Conference 53: 127-136. 89. Vallentine, J. F. 1971. Range development and improvements. Brigham
67. Robel, R. J., A. R. Bisset, T. M. Clement, Jr., A. D. Dayton, and K. L. Young University Press, Provo, UT.
Morgan. 1979. Metabolizable energy of important foods of bobwhites 90. Wang, G., W. D. Edge, and J. O. Wolff. 1999. A field test of the quotient
in Kansas. Journal of Wildlife Management 43: 982-987. method for predicting risk to Microtus canicaudus in grasslands.Archives
68. ———, R. M. Case, and A. R. Bisset. 1974. Energetics of food plots in of Environmental Contamination and Toxicology 36: 207-212.
bobwhite management. Journal of Wildlife Management 38: 563-564. 91. Ward, A. L. 1982. Mule deer behavior in relation to fencing and
69. Robinson, S. K., F. R. Thompson III, T. M. Donovan, D. R. Whitehead, and underpasses on Interstate 80 in Wyoming. Transportation Research
J. Faaborg. 1995. Regional forest fragmentation and the nesting success Record 859: 8-10.
of migratory birds. Science 267: 1987-1990. 92. Warner, R. E., G. B. Joselyn, and S. L. Etter. 1987. Factors affecting
70. Rodenhouse, N. L., L. B. Best, R. J. O’Conner, and E. K. Bollinger. 1993. roadside nesting by pheasants in Illinois. Wildlife Society Bulletin 15:
Effects of temperate agriculture on neotropical migrant landbirds. Pages 221-228.
280-295 in: D. M. Finch and P. W. Stangel, editors. Status and 93. Weinstein, D. A., and E. M. Birk. 1989. The effects of chemicals on the
management of neotropical migratory birds. U.S. Forest Service, structure of terrestrial ecosystems: mechanisms and patterns of
General Technical Report RM-229. change. Pages 181-209 in: S. A. Levin, M. A. Harwell, J. R. Kelly, and K. D.
71. Rothstein, S. I., J. Verner, and E. Stevens. 1984. Radio-tracking confirms Kimball, editors. Ecotoxicology: Problems and approaches. Springer-
a unique diurnal pattern of spatial occurrence in the parasitic brown- Verlag, New York, NY.
headed cowbird. Ecology 65: 77-88. 94. Wildlife Management Institute. 1994. The Conservation Reserve
72. Sanderson, H. R., T. M. Quigley, E. E. Swan, and L. R. Spink. 1990. Program: a wildlife conservation legacy. Wildlife Management Institute,
Specifications for structural range improvements. U.S. Forest Service, Washington, D.C.
General Technical Report PNW-250. 95. Wilson, S. D., and J. W. Belcher. 1989. Plant and bird communities of
73. Schauber, E. M., W. D. Edge, and J. O. Wolff. 1996. Insecticide effects on native prairie and introduced Eurasian vegetation in Manitoba, Canada.
small mammals: influence of vegetation structure and diet. Ecological Conservation Biology 3: 39-44.
Applications 7: 136-150. 96. Wolff, J. O., T. Fox, R. R. Skillen, and G. Wang. 1999. The effects of
74. Schmitz, R. A., and W. R. Clark. 1999. Survival of ring-necked pheasant supplemental perch sites on avian predation and demography of vole
hens during spring in relation to landscape features. Journal of Wildlife populations. Canadian Journal of Zoology 77: 535-541.
Management 63: 147-154. 97. Wywialowski, A. P. 1994. Agricultural producers’ perceptions of
75. Schroeder, R. L., T. T. Cable, and S. L. Haire. 1992. Wildlife species wildlife-caused losses. Wildlife Society Bulletin 22: 370-382.
richness in shelterbelts: test of a habitat model. Wildlife Society Bulletin 98. Yahner, R. H. 1982. Microhabitat use by small mammals in farmstead
20: 264-273. shelterbelts. Journal of Mammalogy 63: 440-445.
76. Shalaway, S. D. 1985. Fencerow management for nesting birds in 99. ———. 1983. Small mammals in farmstead shelterbelts: habitat
Michigan. Wildlife Society Bulletin 13: 302-306. correlates of seasonal abundance and community structure. Journal of
77. Singer, F. J., and J. L. Doherty. 1985. Managing mountain goats at a Wildlife Management 47: 74-84.
highway crossing. Wildlife Society Bulletin 13: 469-477. 100. ———. 1988. Changes in wildlife communities near edges.
78. Snyder, W. D. 1984. Ring-necked pheasant nesting ecology and wheat Conservation Biology 2: 333-339.
farming on the high plains. Journal of Wildlife Management 48: 878-888. 101.Yoakum, J., W. P. Dasmann, H. R. Sanderson, C. M. Nixon, and H. S.
79. Stallman, H., and L. B. Best. 1996. Bird use of an experimental strip Crawford. 1980. Habitat improvement techniques. Pages 329-410 in: S.
intercropping system in northeast Iowa. Journal of Wildlife D. Schemnitz, editor. Wildlife management techniques manual, fourth
Management 60: 354-362. edition. The Wildlife Society, Inc., Washington, D.C.
80. Stinson, E. R., and P. T. Bromley. 1991. Pesticides and wildlife: a guide to
reducing impacts on animals and their habitat. Virginia Cooperative
Extension Communications, Blackburg, VA.
81. Tiebout, H. M., III, and R. A. Anderson. 1996. A comparison of
corridors and intrinsic connectivity to promote dispersal in transient
successional landscapes. Conservation Biology 11: 620-627.
82. ———, and K. E. Brugger. 1995. Ecological risk assessment of
pesticides for terrrestrial vertebrates: evaluation and application of the
U.S. Environmental Protection Agency’s quotient model. Conservation
Biology 9: 1605-1618.
83. U.S. Department of Agriculture. 1998. 1997 Oregon Wildlife damage
survey. U.S. Department of Agriculture, Oregon Agricultural Statistics
Service, Portland, OR.
84. ———. 1999. Agricultural statistics 1999. U.S. Department of
Agriculture, National Agricultural Statistics Service, U.S. Government
Printing Office, Washington, D.C.
 15
Wildlife of Coastal and Marine Habitats
Joseph B. Buchanan, David H. Johnson, Eva L. Greda,
Gregory A. Green,Terence R.Wahl, & Steven J. Jeffries

Introduction
The marine environment contains perhaps the most Bay, Netarts Bay); blind estuaries occur where sediments
dynamic collection of habitats in the Pacific Northwest. close the outlet to the ocean, primarily during summer
The habitats included in this chapter—coastal dunes and months (e.g., Elk River, Sixes River); drowned river
beaches, bays and estuaries, coastal headlands and islets, estuaries occur where a rise in sea-level floods a former
nearshore, inland marine deeper waters, marine shelf, and river valley (e.g. Coos Bay, Yaquina Bay); many Puget
oceanic waters—are uniquely moderated by movement Sound estuaries are considered fjord estuaries due to the
of marine water. Coastal upwellings, tides, currents, and glacial origin of Puget Sound.158 The major estuaries in
waves all influence habitat use by wildlife. Washington and Oregon are indicated in Figure 1.
The total area of individual habitats covered in this Salt marsh is associated with many of the estuaries,
chapter varies substantially. Although estuaries, beaches, although tremendous amounts of salt marsh habitat have
dunes, and headlands occupy comparatively smaller areas been either destroyed or degraded through diking and
than other habitats, the area of marine water (out to 200 conversion to agriculture or urbanization, primarily
miles offshore) is obviously immense (about 65.5 million during the late nineteenth and twentieth centuries.19, 50 For
acres [26.5 million ha]). Many of the habitats are remote example, approximately 90% of the salt marsh in Coos
areas—coastal headlands, marine shelf, and oceanic Bay was destroyed between about 1880 and 1980.158
waters—with low levels of human activity, whereas others Similarly, about 60% of the salt marsh in Puget Sound was
(beaches, estuaries, inland marine deeper waters) are lost following European settlement of that area.158 To a
important commercial areas or popular recreation sites. large extent, high quality (native) salt marsh is currently
At least 284 regularly occurring species (219 birds, 58 found only in very limited areas.
mammals, 7 reptiles, and 3 amphibians) are associated Vast intertidal mudflats are present in the protected
with these habitats. bays and inlets where sediment carried by the riverine
In this chapter we present information useful to those system is deposited. Some bays, particularly Coos Bay,
with an interest in the ecology, conservation, and Grays Harbor and sites in Puget Sound, are important
management of the marine environment. We begin by shipping ports, and shipping channels in some of these
generally describing the habitats and features of the areas are maintained by regular dredging of accumulated
habitats important to wildlife. After a summary of sediments; small islands within the Columbia River
information on species richness, adaptations of marine mouth, Grays Harbor, and Coos Bay, for example, have
wildlife, and threats to the wildlife community, we been created or altered by deposited dredge spoils.
describe the distribution and abundance of representative The conditions in bays and estuaries, and along their
species in the different habitats. Acknowledging the shorelines, are quite variable, typically reflecting each site’s
importance of oceanic warming, we describe the unique geological and disturbance history. Akins and
phenomenon of El Niño/Southern Oscillation events (and Jefferson1 recognized eight types of salt marsh in Oregon
oceanic warming in general), and conclude with a section estuaries. These types vary as a function of marsh
on conservation issues in the marine environment. elevation, slope, substrate, drainage, and vegetation cover.
These conditions influence the species composition
associated with a particular salt marsh. Highly disturbed
Habitat Descriptions
sites often lack or have much reduced salt marsh areas,
Estuaries represent the dynamic interface of riverine and
an important habitat component of estuarine ecosystems.
marine systems. The complex ecological processes that
Some salt marshes are important habitat areas for roosting
occur in estuaries are beyond the scope of this chapter
shorebirds and gulls, and are also used as haul-out areas
and are described elsewhere.54, 118, 158, 161 There are four
(e.g. in Washington’s Hood Canal) by harbor seals.
commonly recognized types of estuaries in the coastal
Estuarine tide flats and salt marshes are important hunting
Pacific Northwest: bar-built estuaries develop where
grounds for a variety of raptor species including falcons,
accumulated sand restricts the flow of water to the ocean
buteos, harriers, and owls. These birds benefit from the
and thereby influences sediment transport (e.g. Willapa
presence of drift logs, used as hunting or resting perches.
389
 390 Wildlife–Habitat Relationships in Oregon and Washington

Figure 1. Location of
estuaries in the marine
regions of Washington
and Oregon. Note that
some estuaries (Elliott
Bay, Commencement Bay,
and Budd Inlet in
Washington) have been
largely converted to
industrial or other
human-development
purposes and have
therefore lost most
estuarine function.
 CHAPTER 15: COASTAL/MARINE 391

Beaches can vary from broad, relatively flat expanses 26 in Oregon (see Figure 2). Some dunes are associated
of sand to narrow, fairly steep slopes dominated by with rivers or bays whereas others are on broad coastal
pebbles or larger rocks. Pacific beaches are strongly plains or terraces.192 Many dunes are vegetated with
influenced by long-shore drift, the seasonal currents that European beach grass, an exotic species used to stabilize
transport sand and sediment that physically alters the dune movement.194 Vegetation associated with dunes can
width or other characteristics of a beach.103 The dominant be quite diverse, with 21 plant communities identified.192
feature of the beach is the sand or pebble substrate. Grain Most dunes in Washington rise no more than 4-6 m,
size can be quite variable within and among beaches; the whereas some dune systems in Oregon attain heights over
most important areas for nesting, roosting, and foraging 25 m above the interdune base.192 In some areas, lower
shorebirds are typified by fine-grained sand. In many foredunes develop in the areas between the main body of
areas, the top several cm of sand is obviously rich in dunes and beaches (seaward) or salt marsh (bayward).
invertebrate life and sustains large concentrations of With the exception of Clatsop Spit, Oregon, where rapid
overwintering and migrant sanderlings.29, 120 Other than progradation was responsible for the creation of foredunes
the actual substrate, the most prominent features of coastal (A. Wiedemann, pers. comm.), the foredunes along the
beaches are the streams (or rivers) and drift logs. The coast are associated with stabilizing vegetation. In fact,
streams and rivers contribute to the dynamic movement Cooper47 found no foredunes along the coast prior to the
of sand and silt in this environment. Larger rivers carry establishment of European beachgrass. Very small dune
huge amounts of silt from uplands and deposit the formations (embryo dunes) occur seaward of larger dunes
material in embayments or offshore. Much of this material (e.g. Leadbetter Point, Washington) although it should be
is eventually transported ashore by currents and wave noted that these dunes are not persistent (i.e., they are
action.103 Small watercourses carry less sediment but often regularly destroyed by wave action) and are not
create new channels through low dunes and influence considered foredunes.193
local erosion. These small streams appear to influence the The dominant features of the coastal dunes, other than
distribution of shorebirds along the beach29 by providing the shape and function (i.e., amount and direction of
a source of fresh water important for bathing. movement 192), vary depending on the vegetation
Large numbers of drift logs wash up on beaches, community present. Typically, the vegetation has a strong
particularly during storms. These logs often persist on herbaceous, grass or shrub component.76, 192 Shrubs are
beaches for years until they are carried away or moved to important habitats for overwintering passerines such as
higher dunes by strong storms; buried in sand; or used as fox sparrow and yellow-rumped warbler. Raptors such
firewood, either on site or after being cut and removed. as the northern harrier use trees as perch sites. Small
Current amounts of drift logs on beaches tend to be far mammals and amphibians and reptiles associated with
lower than were present decades ago.79 Drift logs on coastal dunes find protective cover for foraging and
beaches are used as foraging and resting perches by denning in and under large drift logs deposited by winter
falcons. Very small pieces of wood are occasionally used storms. Such logs also are used as resting and feeding sites
as wind breaks by roosting shorebirds, particularly the by falcons.
snowy plover, in windy weather. Another common feature of coastal dune systems,
Situated immediately landward of the beaches, sand particularly those influenced by the introduction of
dunes are found along 307 of the 764 km (40%) of outer European beachgrass, is the presence of very small, often
coastline from Cape Flattery, Washington, to the California secluded pools formed by storm- or rainwaters. The use
border.192 There are 3 dune locations in Washington and of these pools by wildlife has not been well documented,
although they are known to be used by waterfowl and
certain shorebirds (e.g. greater yellowlegs; J. Buchanan,
unpubl. data).
Each of the habitats above is influenced by the
movement of water, either through periodic storms or the
regular pattern of the tidal cycle. Beaches and dunes can
be scoured or displaced by regular currents or wave action
during particularly severe storms. Similarly, mudflats can
also be scoured by storm runoff or regular strong tidal
currents. Sand beaches can recede or accrete depending
on the placement of new sediments in the system and the
strength of currents. 103 Salt marsh progradation is
occurring in some coastal estuaries, but in the region as a
whole has occurred at a lesser rate than salt marsh loss.158
Beaches and mudflats are alternately covered or
exposed for hours at a time due to daily fluctuations in
the region’s tidal regime, which has both diurnal and semi-
California Gulls at Fort Canby State Park, Washington. diurnal characteristics (there are generally two high and
(Photo by Terence R. Wahl) two low tides, of varying height, each day). In larger
392 Wildlife–Habitat Relationships in Oregon and Washington

Figure 2. Distribution of dune locations in Washington and

Oregon (map from Wiedemann;192 see Cooper47).

estuaries such as Grays Harbor and Willapa Bay, >50 km2

of tide flats may be exposed during low tides.
A number of human-made structures are used by
wildlife associated with coastal beaches, dunes, bays, and
estuaries. Perhaps the most permanent of these structures
are the jetties used to maintain shorelines and harbor
entrances. These sites are often used by roosting gulls and
cormorants, and are important sites for rock sandpiper,
surfbird, black turnstone, and ruddy turnstone. Shellfish
beds in some areas are also used by wildlife such as
turnstones, although the value of the areas is not clear. In
fact, a recent study indicated that dunlins and western
sandpipers were less common in bivalve management
areas than in unmodified areas.99
Other structures used by a wide variety of species
include buoys, channel markers, pilings, and docks.
Buoys, channel markers, and docks are often used as haul-
out sites by harbor seals or sea lions. These structures are
also used as resting or foraging perches by a variety of
birds including gulls, peregrine falcons, shorebirds, and
pelagic cormorants (occasionally used for nesting). Pilings
are used in some areas as high tide roost sites by black-
bellied plovers and are used for nest sites by cavity-
dependent species such as the purple martin.
The Coastal Headlands and Islets habitat includes the
offshore rocks and islands, and coastal headlands and
 CHAPTER 15: COASTAL/MARINE 393

bluffs dominated by salal or evergreen huckleberry. The Nearshore Marine habitats include those marine
Coastal headlands are defined as rocky cliffs of non-glacial water areas (high tide line to a depth of 66 ft [20 m]) along
origin on mainlands facing marine waters. Examples of shorelines that are not significantly affected by freshwater
coastal headlands include some of the San Juan Islands, inputs (i.e., excludes Bays and Estuaries). Adjacent to
the area between Cape Flattery and Point Grenville, and Marine Shelf, Inland Marine Deeper Waters, Bays and
much of the coastline of Oregon. Some headlands are Estuaries, and a number of terrestrial-based habitats,
fronted by sand beaches, whereas others rise directly from Nearshore Marine habitat occurs in a mosaic with Coastal
open water. Most bluffs in Washington are fronted by Headlands and Islets. A wide range of benthic habitats
cobble beach (areas of Puget Sound), while bluffs in support marine organisms capable of withstanding short-
Oregon rise from open water or beach. Headlands tend to term exposure to air. Larger kelps, such as Macrocystis
be much taller (up to hundreds of meters) whereas bluffs integrifolia and Nereocystis leutkeana can be found in rocky-
tend to be smaller than about 20-50 m. bottom subtidal areas, whereas rocky-bottom intertidal
There are several key habitat features associated with areas support smaller marine vegetation such as brown
coastal headlands. Most coastal headlands and the larger rockweed (Pelvetiopsis limitata) and surfgrass (Phyllospadix
offshore islands have at least a thin layer of soil that scouleri). Strongly influenced by tidal rhythms, wave
supports grasses and evergreen shrubs, and in many areas, action, storm events, and light penetration, the Nearshore
evergreen forest. It is these areas, primarily on offshore Marine habitat is characterized by a high degree of
islands, but occasionally on mainland headlands, that patchiness resulting in differences in its faunal makeup
support great breeding colonies of seabirds. Consequently, and use. Examples of succession may be found in kelp
soil and vegetation properties that allow seabirds to dig forests where herbivory or the scouring action of tidal
long but shallow burrows are essential for many of these currents removes vegetation, and on rocky intertidal
birds.78 Most of the headlands and bluffs have very steep shores where wave action periodically disturbs established
or vertical cliff faces. The areas are used by ledge-nesting communities.
seabirds and species such as the peregrine falcon. Other The Inland Marine Deeper Water habitat occurs in
species, such as belted kingfisher and pigeon guillemot, northwestern portions of Washington State and British
nest in coastal bluffs where burrows can be excavated in Columbia, and does not occur in Oregon. This habitat
the face of the escarpment.66 Snags and large trees present includes waters (>66 ft [>20 m] deep) of the Strait of
on these headland faces are used as perch locations and Georgia, Puget Sound, Hood Canal, and the Strait of Juan
nest sites by species such as the bald eagle. The shrub de Fuca. Bays and Estuaries, Coastal Headlands and Islets,
vegetation provides habitat conditions for ground- and and Marine Nearshore habitats adjoin Inland Marine
shrub-nesting species such as winter wren and song Waters Habitat. This habitat further merges with the
sparrow. Marine Shelf habitat in the Strait of Juan de Fuca and is
Along much of the coastline there are rocky benches or used extensively for navigation, commercial transport of
accumulations of large boulders at the base of the goods, recreation, tourism, and fishery operations. Varying
headlands. Low-lying bench areas are used as foraging in specific composition and distribution with fluctuating
sites by rock-shoreline shorebirds and as haul-out sites physical factors, organics, silt, and sand are the primary
by harbor seals. Higher elevation sites (those above the substrate components of this habitat. Inland marine deeper
wave-splash zone) are used as nesting areas by the black waters do not support a significant number of plants due
oystercatcher. to light diffusion resulting from the depth of water
involved. Seasonal variation in tidal regimes, precipitation
and riverine discharges (winter highs), as well as periodic
storm events cause changes in temperature, salinity,
energy level, and erosion and accretion that effectively
shape this habitat.
The Marine Shelf and Oceanic habitats are generally
described topographically as the shelf (waters 20-200 m
in depth), slope (200-2,000 m), and offshore or rise (greater
than 2,000 m in depth) (Figure 3). The shelf can be further
subdivided into inner (20-100 m) and outer 100-200 m)
regions, especially since the oceanography and biology
can change considerably closer to the shelf edge. Off
Oregon and Washington the continental shelf varies in
width from 15 to 40 km, and is relatively uniform
compared to other coastal regions of North America.
Nevertheless, there are numerous geomorphological
features that influence the physical and biological
processes. The continental shelf along Oregon is
Harbor Seals hauling out at low tide along Oregon coast. characterized by a series of oceanic banks, including Daisy,
(Photo by Steven J. Jeffries) Stonewall, Perpetua, Heceta, and Siltcoos Banks, and a
 394 Wildlife–Habitat Relationships in Oregon and Washington

Current waters with shelf edge geomorphological features.

Coastal upwelling occurs most frequently in summer and
fall, promoted by northerly and northwesterly winds.
Upwelling intensity is typically greatest along the
southern Oregon coast (the Cape Blanco upwelling zone).
The Columbia River effluent amounts to 60% to 90% of
the freshwater entering the Pacific Ocean between the
Strait of Juan de Fuca and San Francisco Bay.
Marine wildlife occupying the outer shelf, slope, and
offshore waters concentrate where deep and shallow
waters mix. Some of the highest concentrations occur
along the shelf slope influenced by the California Current,
and the very edge of the outer shelf where coastal and
current waters mix. These mixing zones are especially
prominent at Heceta Bank where its unique bending shape
and proximity to the Cape Blanco upwelling zone infuse
the Bank with deepwater nutrients. Other important
mixing zones occur along the headwalls of Washington’s
submarine canyons.
Additional oceanographic features include the
Columbia River plume on the upper layers of the shelf
and offshore waters (contributing to the “dilute domain”68)
but while the effects of this on bird distribution are
unknown,23 the high concentration of freshwater within
this plume may limit its attraction to marine life. This
outflow extends northward along the Washington coast
in winter and shifts to offshore and south in summer. More
dramatic when encountered but not always present over
the outer shelf or slope, the “blue water” boundary
suggests oceanic water mass boundaries like the Subarctic
Boundary across the North Pacific,68 where water color,
salinity, sea surface temperatures, and seabird species
composition and abundance change dramatically;180 these
conditions should not be confused with warm water
incursions. One noticeable change in seabird abundance
at this boundary area, where albacore are often found, is
the replacement of foraging fork-tailed storm-petrels by
Leach’s storm-petrels on the warmer, offshore side.177 In
Figure 3. Major bathymetric features of the offshore region of addition, patterns of marine bird occurrence off
Washington and Oregon. Washington and Oregon probably reflect events or
conditions in the Arctic, the southern hemisphere or
major promontory, Cape Blanco. In contrast, the elsewhere; reproductive success or failure and feeding
Washington shelf is furrowed by the Nitinat, Juan de Fuca, opportunities at staging areas or during migration may
Quinalt, Grays, Guide, Willapa, and Astoria submarine determine populations of visiting species here.
canyons. The 20- to 25-km-wide Strait of Juan de Fuca,
separating Vancouver Island from mainland Washington,
is a glacially excavated channel that is the primary avenue Species Richness and Adapatations of
for vigorous estuarine exchange between the coastal ocean Marine Wildlife
and the inland marine waters of Washington and British The marine habitats differ markedly in their attributes and
Columbia. The Columbia River discharges through a their associated species. Evaluation of the information
relatively narrow and deep river mouth that defines the presented in Table 1 reveals a number of patterns of
coastal boundary between Oregon and Washington. interest. Species richness was greatest in the bays and
Unlike California, and except for the southern part of estuaries, followed by dunes and beaches, the marine
Oregon, most of the region is not directly affected by the nearshore zone, the marine shelf area, and the oceanic
California Current system away from the more oceanic zone. Consistent with this pattern, there was a strong
areas. 68, 108 Consequently, the nearshore biological inverse relationship between species richness and the
productivity is primarily due to wind-driven coastal proportion of closely associated species found in the
upwelling. In contrast, offshore nutrification results largely habitat. Not surprisingly, the oceanic habitat, with only
from water mixing due to the interaction of the California 42 species, had the highest proportion (69%) of closely
 CHAPTER 15: COASTAL/MARINE 395

Table 1. Species richness in the different marine habitats based on the degree of association. Because some
species are associated with more than one habitat, totals within rows are not additive. Species of uncertain
association (n = 9) and of historical occurrence (n = 1) were not included in the table.

Species Groups Habitats

Dunes/ Headlands Bays/ Inland Marine Marine Oceanic
Beaches Estuaries Marine Nearshore Shelf

Closely Associateda
Marine Mammals 0 3 2 5 6 11 10
Terrestrial Mammals 1 0 4 0 0 0 0
Birds 29 25 60 21 32 28 18
Reptiles 1 0 0 0 0 0 1
Amphibians 0 0 0 0 0 0 0
Total for Group 31 28 66 26 38 39 29
Generally Associated b
Marine Mammals 2 0 1 2 1 5 1
Terrestrial Mammals 11 20 5 0 1 0 0
Birds 61 38 67 18 31 15 7
Reptiles 4 4 1 0 0 1 0
Amphibians 1 1 0 0 0 0 0
Total for Group 79 63 74 20 33 21 8
Present c
Marine Mammals 0 0 2 2 2 1 2
Terrestrial Mammals 14 7 2 1 0 0 0
Birds 27 17 30 12 12 10 3
Reptiles 0 0 0 0 0 0 0
Amphibians 0 0 0 0 0 0 0
Total for Group 41 24 34 15 14 11 5
Total for Habitat 151 115 174 61 85 67 42
a
Closely associated species: A species is widely known to depend on a habitat for part or all of its life history requirements.
Identifying this association implies that the species has an essential need for this habitat for its maintenance and viability.
Generally associated species: A species exhibits a high degree of adaptability and may be supported by a number of habitats. In other
b

words, the habitats play a supportive role for its maintenance and viability.
Present species: A species demonstrates occasional use of a habitat. The habitat provides marginal support to the species for its
c

maintenance and viability.

associated species. Conversely, the bays/estuaries and Threats to Marine Habitats and Wildlife
dunes/beaches had the lowest proportions of closely Communities
associated species (38% and 20%, respectively). The two There are many threats to the species and habitats
latter habitats also had the highest proportions of generally associated with the marine environment. Because an in-
associated species (42% and 51%, respectively). These depth discussion of this topic would be too lengthy for
numbers make sense intuitively; generalists are less this chapter, we have summarized the major issues and
common in the oceanic and predominantly marine zones provided references so readers can locate additional
and are more prevalent in the more diverse habitats pertinent information (Table 3).
associated with the marine/upland interface. In addition to the threats we summarized, a noteworthy
The wildlife species associated with the marine threat is now emerging as a result of decades of human
environment have adapted to prevalent conditions along inability to co-exist with wildlife populations on a
the coasts of Washington and Oregon. Certainly, the sustained basis. This threat is best described as the
characteristics of migrant species are also uniquely presence of conflicting management objectives. As efforts
influenced by conditions throughout the range of each to manage wildlife populations become more difficult due
species. All species are constrained in some way by biotic to reduced or eliminated options, coupled with ever-
or abiotic conditions. Some of these constraints, and the increasing pressures from an expanding human
associated ecological consequences and adaptations, are population, the potential for management conflict
presented in Table 2. becomes more likely and more significant. A current
example is the issue of Caspian tern predation on salmon
 396 Wildlife–Habitat Relationships in Oregon and Washington

Table 2. Examples of constraints, ecological consequences and animal adaptations

in the marine environment.

Environmental Constraints Ecological Consequences Adaptations

Physiological/Structural Behavioral

Temporal changes in prey Higher energetic costs Long-legged birds can forage Nocturnal foraging allows greater
availability (daily or seasonal); associated with avoiding in varying water depths to access to tidal areas; periods of
depleted prey populations predators, moving between extend the foraging period; inactivity (roosting) to minimize
foraging and roosting locations, utilize lipid deposits to support energy expenditure; prey
searching for prey; density- emigration; fat storage as switching, habitat switching;
dependent competition; blubber; replenish lipid minimize energetic costs by
mortality reserves and store as fat; use reducing time spent flying (e.g.
high-energy intake to fuel western grebes)
migration; body shape (and
wing structure) adapted to
avoid predators (e.g. shorebirds)
and sustain long migratory
movements
Adverse weather conditions Increased energetic demands; Utilize lipid deposits to Temporal flexibility in
(flooding of foraging, roosting, less time spent roosting, support emigration; insulating reproductive timing
or nesting areas; high winds) foraging, and other plumage, pelage, and blubber
maintenance requirements;
reduced foraging efficiency;
failed reproduction
Spatially limited habitats Greater physiological costs Strong seasonality in habitat use;
associated with migration; excellent dispersal/ migration
density-dependent competition abilities; colonial breeding;
breeding rookeries
Predation or predator density Disturbance; increased Body shape and wing structure Balancing food intake and vigilance
energetic costs; increased adapted to avoid predators; with predation risk; predator
vigilance and predator cryptic plumage avoidance strategies effective in
avoidance behavior suited to open habitat; flocking behavior
the habitat; mortality enhances foraging behavior and
predator detection; nest
placement (on islands, burrow
into soil); colonial breeding;
breeding rookeries
Cold marine water Warming adaptations required Insulating plumage, pelage and
temperatures blubber; small ears and robust
body parts
Marine waters as habitat Required ability to subsist in Tolerance of saline conditions; Ability of waterbirds to swim and
marine environment webbed feet and wing shape obtain food under water surface
facilitate swimming under water
by waterbirds; flippers

smolts near the tern nesting colony in the lower Columbia Wildlife Communities
River estuary. The terns’ appetite for salmon smolts The species groups associated with marine habitats exhibit
concerns fisheries biologists and others engaged in a great range of seasonal occurrence patterns in the region.
management and recovery of salmon stocks. Potential With the exception of the leatherback sea turtle, all reptiles,
solutions to the problem proposed by various parties have amphibians, and terrestrial mammals are considered year-
included tern population control and colony relocation. round residents. Similarly, most marine mammals are
Certainly there is hope for successful colony relocation, year-round residents; however, seven species (e.g.
given the species’ known ability to respond to dynamic humpback whale, northern fur seal) occur only or
habitat changes. However, the take-home message is that primarily during spring/summer or fall/winter (Table 4).
a naturally established Caspian tern colony (although It is among the birds that the greatest variability in seasonal
located on a dredge spoil island), one of the largest in the occurrence exists (Table 4). Although year-round residents
world, and therefore of great conservation significance, is are well represented (31% of the birds), they are
in conflict with salmon conservation efforts. The solution, outnumbered by winter residents (37%). Other species
unfortunately, is active engagement and intervention. breed only or are exclusively migrants. A small number
 CHAPTER 15: COASTAL/MARINE 397

Table 3. Summary of actual or potential threats to habitats or species in the marine environment.

Major Habitat Associations References

Threat Oceanic Marine Marine Inland Bay/ Coastal Beach/
shelf nearshore marine Estuary headland Dune

Habitat Lossa
Habitat Loss Xb Xb X 19, 20

Habitat Degradation
Habitat alteration X X 27, 194

Climate change X X X X X X X 187

Exotic vegetation X X X 27, 99, 125

Exotic invertebrates ? ? ? X X ? ? 27

Exotic vertebrates X X 78, 94

Pollution
Agricultural X 34, 49, 157,

Industrial/Municipal X X 34, 38, 100, 157,

Oil X X X X X X X 27, 32, 73, 165, 100

Heavy metal X ? 27, 34, 49, 3100

Plastic particle X X X X X X 27, 55, 149

Human Disturbance
Pedestrian X X X 27

Motorized vehicles X X X X 27

Non-motorized watercraft X X 27

Pets X X X 27, 62

Hunting X 27

Military activities X 166

Mortality Factors
Collisions with utility lines X 27

Market hunting X X 46, 71, 83

Collisions with X X X 27

vehicles or watercraft
Gillnet and longline X X X X 57, 117, 126

fisheries activities
entanglement
Changes in predator X 144

or predation levels
a
Habitat loss is here defined as permanent conversion of suitable habitat. Habitat degradation refers to a range of conditions, from
slightly to completely degraded; by definition, it is possible to restore degraded habitats.
b
Loss of reef structure on ocean bottom due to trawling (J. Hodder, personal communication).

of far ranging, non-breeding seabirds and marine Many of the species found in these habitats engage in
mammals occur at various seasons, but do not occur year- lengthy migrations or dispersal movements. With the
round. exception of the leatherback sea turtle, all reptiles,
Habitat-associated patterns of seasonal occurrence are amphibians, and terrestrial mammals generally move less
evident as well. Higher proportions of species are found than 1,000 km (Table 4). On the other hand, 69% of the
in coastal dunes and coastal headlands during summer marine mammals and 85% of the birds regularly travel
(year-round residents or summer only in Table 5) than in distances greater than 1,000 km. In fact, the majority of
other habitats. With the exception of oceanic waters, in strictly migrant species travel over 10,000 km (16% of the
which 48% of the species occur in winter (year-round and marine mammals and 23% of the birds travel over 10,000
winter columns in Table 5), all habitats support high km) (Table 4).
proportions of species in winter. Bays/estuaries, inland
marine deeper waters, and nearshore waters support the Coastal Dunes, Beaches, Bays, and Estuaries
highest proportions of species which overwinter but do The beach habitat supports very large numbers of
not breed locally (Table 5). vertebrates, and several areas in the region are recognized
 398 Wildlife–Habitat Relationships in Oregon and Washington

Table 4. Summary of seasonal occurrence and migration/dispersal capability of birds and marine mammals in
coastal and marine habitats of Washington and Oregon.

Seasonal Status Migration/Dispersal Capability Number of Species/

<10 km & sedentary <100 km <1,000 km <10,000 km >10,000 km Seasonal Category

Birds
Year-round 7 (10%) 3 (4%) 16 (24%) 41 (61%) 0 67 (31%)
Summer only 0 0 2 (8%) 16 (64%) 7 (28%) 25 (12%)
Winter only 0 0 5 (6%) 65 (82%) 9 (11%) 79 (37%)
Transient 0 0 0 12 (33%) 24 (67%) 36 (17%)
Other 0 0 0 0 9 (100%) 9 (4%)
Total 7 (3%) 3 (1%) 23 (11%) 134 (62%) 49 (23%) 216
Marine Mammals
Year-round 0 1 (8%) 5 6 0 12 (63%)
Summer only 0 0 0 1 3 4 (21%)
Winter only 0 0 0 3 0 3 (16%)
Transient 0 0 0 0 0 0
Other 0 0 0 0 0 0
Total 0 1 (5%) 5 (26%) 10 (53%) 3 (16%) 19

Table 5. Summary of seasonal residency of species associated with marine habitats in the Pacific Northwest.

Habitat Number & Percentage of Species in each Residency Category Notes

Year-round Summer only Winter only Spring/ Other
Autumn only

Coastal Dunes and Beaches 80 (52%) 16 (10%) 36 (23%) 22 (14%) 0 Does not include 4 uncertain, one
extirpated species
Coastal Headlands and Islets 68 (58%) 22 (19%) 19 (16%) 8 (7%) 0 Does not include 3 uncertain
status species
Bays and Estuaries 64 (36%) 16 (9%) 65 (37%) 32 (18%) 1 Does not include 1 uncertain
status species
Inland marine deeper waters 23 (37%) 5 (8%) 25 (40%) 9 (14%) 1
Marine nearshore 27 (30%) 8 (9%) 37 (42%) 15 (17) 2 Does not include 1 uncertain
status species
Marine shelf 25 (35%) 5 (7%) 20 (28%) 13 (18%) 9 (13%)
Oceanic 17 (39%) 4 (9%) 4 (9%) 10 (23%) 9 (20%)

as internationally important sites. For example, the tides (J. Gilligan, pers. comm.). Additionally, the area
northern coast of Oregon and the beaches adjacent to between Coquille Point and Cape Blanco also supports
Grays Harbor and Willapa Bay support some of the flocks of roosting shorebirds, primarily during brief
highest densities of spring (472/km120) and autumn (214/ migratory stopovers (D. Lauten, pers. comm.). Ocean
km31) migrant sanderlings in North America. The three Shores/Copalis Beach, North Beach, and Clatsop/Sunset
Washington beaches (Ocean Shores/Copalis Beach, Beach (Oregon) qualify as sites of regional importance62
Grayland Beach, North Beach [also known as Long Beach]) in the Western Hemisphere Shorebird Reserve Network84
supported an average of 50.2, 43.3, and 46.6 sanderlings/ (WHSRN). Eight beaches in the region are considered
km during winter between 1982-83 and 1989-90.29 These critical habitat for the threatened snowy plover.172 Sand
beaches are also important areas for roosting shorebirds islands in large estuaries (e.g. Grays Harbor, Willapa Bay,
during daily high tides that inundate the nearby Columbia River mouth) also support large breeding
embayments. Winter counts of dunlins averaged 408.7, colonies of gulls and terns,142, 167, 173 and are used as haulout
464.4, and 618.7/km on the three beaches. North Beach is sites by harbor seals (see below).
also an important area for roosting black-bellied plovers29 Estuarine sites in the region support substantial winter
(high count of 3848 in winter 1994-95).31 Although many populations of Arctic- and sub-arctic-nesting shorebirds.31,
141
Oregon beaches support sanderlings, only a few (e.g., Important wintering sites for shorebirds include Willapa
those adjacent to Bandon Marsh and Tillamook Bay) Bay (>70,000 birds),31 Grays Harbor (mean=about 20,000
support other species of roosting shorebirds during higher between 1979 and 198), 141 Columbia River estuary
 CHAPTER 15: COASTAL/MARINE 399

(20,000), 141 Port Susan Bay (>30,000), 65 Skagit Bay

(>29,000),65 and Coos Bay (11,000).62, 141 The most abundant
winter resident is the dunlin, which typically comprises
about 90-95% of the winter shorebird community in
western Washington.31, 65 Other common winter resident
shorebirds of estuaries (although often local in
distribution) include black-bellied plover, greater
yellowlegs, and western sandpiper.30, 31, 65, 124, 141
The region’s estuaries are crucial links in the Pacific
Flyway for migrating shorebirds, waterfowl, and other
waterbirds. Grays Harbor is one of the three most
important shorebird stopover sites on the Pacific Flyway
south of Alaska; estimates of about 1 million birds were
made in spring 1981.88 High counts of >100,000 migrant
shorebirds have been made at Willapa Bay31 and the
Columbia River estuary.132, 133, 134, 135, 136, 137 Other WHSRN Harbor Seals hauled out at low tide, Gertrude Island,
sites for shorebirds include Port Susan Bay (>50,000 in both Washington. (Photo by Steven J. Jeffries)
spring and autumn migration), Skagit Bay (>20,000 in
spring), Padilla Bay (>30,000 in spring65), and Tillamook zone nearest shore (Table 6), and density can vary
Bay (nearly 100,000 in autumn 1979, although the site now substantially over broad areas such as Puget Sound
apparently supports far fewer birds) and Coos Bay (Figures 5, 6).
(perhaps 20,000 birds).62, 84 The western sandpiper is by Because of their shallow waters and extensive intertidal
far the most abundant migrant in the region; other sand bars and mud flats, estuaries provide excellent
abundant species include dunlin and short-billed habitat for harbor seals. This habitat provides numerous
dowitcher. Black-bellied plover, semipalmated plover, red low tide haulout areas along channels and exposed tide
knot (primarily in Washington), sanderling, and greater flats which are used year round as resting areas. Harbor
yellowlegs are common during migration.31, 65, 88, 139, 141, 191 seals also select many of the larger estuaries (e.g. Tillamook
At least 26 estuarine sites in Washington and Oregon have Bay, Willapa Bay, Grays Harbor and Padilla Bay) as
supported at least 4,000 shorebirds during the non- preferred haulout locations during pupping season.
breeding period (see Figure 4) and 22 additional sites have Estuaries provide a variety of important prey for harbor
supported at least 1000 shorebirds.62 seals ranging from flatfishes to salmon. In addition to
The most abundant waterfowl restricted primarily to harbor seals, some estuaries such as Grays Harbor provide
marine water habitats are diving ducks. Based on excellent shallow water feeding areas for gray whales
systematic aerial surveys in Puget Sound, these include migrating through Oregon and Washington coastal waters.
scoters (primarily surf scoter and white-winged scoter),
bufflehead, and goldeneyes (Barrow’s goldeneye and Inland Marine Deeper Water
common goldeneye).127 Densities (number of birds/km2) Marine Birds. A relatively low number of species utilize
of these Arctic- and sub-arctic-nesting species in the the deeper inland marine waters. These are mostly mid-
nearshore zone can be quite high and vary substantially water or surface feeders, often found in large flocks which
among years (Table 6). In fact, annual variability in concentrate at convergences or “tide rips” in channels and
abundance (density) is evident in all marine birds in the passages, in offshore, deeper waters of large embayments,
and near baitfish schools in the Strait of Juan de Fuca.
Opportunistic gulls make little apparent distinction
between deep water (>20 m depth) and nearshore (<20
m) waters (see CD-ROM in this book [Wildlife Habitats
matrix]).
Some birds using deep waters often commute
considerable distances between night roosts on rocks and
islands or water “roost” areas. Cormorants and common
murres do this in large numbers. Cormorants must roost
ashore nocturnally and for periods during daylight hours.
Daily movements between roosts along the Strait of Juan
de Fuca and Rosario Strait and the interior channels and
bays in the San Juan Islands and Whatcom, Skagit and
Island counties, for example, are consistently observed.
Cormorants are limited by nesting colony site
Spring migrant shorebirds, primarily Western Sandpipers, at requirements during summer, and are not found much
high tide roost, Bowerman Basin, Grays Harbor, Washington. south of Admiralty Inlet, unlike the winter pattern when
(Photo by Joseph B. Buchanan) numerous cormorants move into all portions of Puget
Sound.
400 Wildlife–Habitat Relationships in Oregon and Washington

Figure 4. Location of estuarine and beach sites that have

supported >4,000 shorebirds in at least one season.62 Sites
referenced by numbers on map are: 1) Drayton Harbor, 2)
Bellingham Bay, 3) Lummi Bay, 4) Chuckanut Bay, 5)
Samish Bay, 6) Padilla Bay, 7) Fidalgo Bay, 8) Skagit Bay, 9)
Port Susan Bay, 10) Snohomish River delta, 11) Dungeness
Bay, 12) Sequim Bay, 13) Totten Inlet, 14) Ocean Shores/
Copalis Beach, 15) Grays Harbor, 16) Grayland Beach, 17)
Willapa Bay, 18) North Beach, 19) Columbia River estuary,
20) Clatsop Beach, 21) Sunset Beach, 22) Tillamook Bay, 23)
Netarts Bay, 24) Oregon Dunes Beach, 25) Coos Bay, and 26)
Coquille River estuary and vicinity, and beach south to Cape
Blanco.

Table 6. Densities (birds/km2) of various waterfowl

species, species groups, and all marine waterbird
species combined, in the Bays and Estuaries of Puget
Sound during winter (December to February, 1992-
93 to 1998-99).*

Species or Species Group Median Density Range

Goldeneye (2 species) 21.97 17.27-38.34

Scaup (2 species) 20.88 13.62-29.05
Scoter (3 species) 56.34 53.56-70.43
Bufflehead 44.38 34.12-64.32
Diving Ducks 165.11 144.45-239.86
Dabbling Ducks 233.78 69.15-722.48
All Marine Birds 621.82 335.84-1284.93
* Data from Nysewander and Evenson128)

Species diversity and populations are larger in winter

than in summer. Resident species are joined in winter by
large numbers of birds from coastal areas to the north and
south and from inland North America. Briggs et al.23 point
out that relatively high densities of wintering common
murres in inland marine waters of Washington may reflect
response to seasonally rougher sea conditions (and poorer
foraging possibilities) over the shelf. Seasonal numbers
vary between years, with populations of species breeding
on the ocean coast—Brandt’s cormorant, common murre
and ancient murrelet—particularly subject to
oceanographic conditions. Likewise, winter feeding
conditions here can affect populations returning to
breeding areas.
Population status and change in inland waters is
incompletely known and apparent changes lack
explanation. Aerial surveys comparing present-day
populations with 1978-79 characterizations of waters north
of Puget Sound183 and surveys of areas south of that in the
early 1980s (Washington Department of Fish and Wildlife,
unpubl. data) indicate that scoters and western grebes
have declined as much as 50% or more in some areas.146
Data from Christmas Bird Counts and unquantified
observations suggest similar changes and also apparent
increased seasonal abundances of phalaropes and above-
average occurrences of offshore species like storm-petrels
and auklets. These apparent patterns could reflect ocean
productivity failure in recent years.
 CHAPTER 15: COASTAL/MARINE 401

Figure 5. Density of
scaups in Puget
Sound, Washington,
derived from aerial
transect counts. Data
and figure from
Nysewander and
Evenson.128

In summer, overall populations are relatively low in Brandt’s cormorants and common murres comprise nearly
the inland marine deeper waters. Cormorants, gulls, and all of the seabird community in the area during summer.
alcids concentrate in active tidal convergence areas like There are no sizable “natural” marine bird colony sites
northern Admiralty Inlet, off Point-No-Point, in the south of the eastern end of the Strait of Juan de Fuca.
Narrows near Tacoma, Case and Carr inlets, in Rosario, Human populations and development preclude the
Haro and Georgia straits, Deception Pass, and in the San establishment of gull and tern colonies except at a few
Juan Islands passages like San Juan and Spieden human-built locations, including abandoned piers and
channels.183 Numbers of birds in Puget Sound, Hood building roofs. There are relatively few scattered
Canal, and waters east of Whidbey Island in summer 1982 cormorant and pigeon guillemot nests.66, 167 Nesting “deep
were very low.185 Deep waters are especially important in water ” species are confined almost exclusively to
the eastern Strait of Juan de Fuca and Admiralty Inlet Protection and Smith Islands in the eastern Strait of Juan
where a significant proportion of the Washington-Oregon de Fuca. Population estimates prior to 1989 included
breeding population of rhinoceros auklets forages.186 The 35,000 rhinoceros auklets and a small number of tufted
auklet concentration in summer results in Admiralty Inlet puffins. Pre-1990s numbers of other nesting species which
numbers peaking then,183 unlike other inland locations forage in both nearshore and deeper waters were 10,600
where summer populations are the lowest of the year. cormorants, 20,000 gulls and 2,400 pigeon guillemots.167
Rhinoceros auklets and variable numbers of non-breeding Except for rhinoceros auklets, inland marine nesting
 402 Wildlife–Habitat Relationships in Oregon and Washington

Figure 6. Density of
rhinoceros auklets in
Puget Sound,
Washington, derived
from aerial transect
counts. Data and figure
from Nysewander and
Evenson.128

populations of “deep-water” foragers are essentially non- observed several times in the San Juan Islands and Georgia
existent. The regional auklet population may benefit in Strait areas179 (T. Wahl unpubl. data). Though inside the
their use of the area because it appears that inland marine 20 m depth contour, up to 25,000 scoters at herring spawn
colonies are less susceptible to oceanic food shortages and events in the eastern Georgia Strait region (in 1979)
breeding failure. indicate the importance of this prey source and why large
Maximum seasonal populations in northern inland flocks of Pacific loons were present in deep water just
marine waters were not estimated for 1978-79.183 Projected offshore (T. Wahl, unpubl. data).
numbers for Hood Canal-Puget Sound of wintering
Marine Mammals. Washington’s inland water with
deepwater alcids (almost all common murres) in
depths greater than 20 m include most of Puget Sound,
December 1982 and February 1983 were 14,000-19,000; the
Hood Canal, Strait of Georgia, and eastern Strait of Juan
estimate for grebes, mostly western grebes, was 27,000-
de Fuca. These deeper marine waters are used primarily
40,000.184 One indication of the importance of the inland
as foraging areas for 6 species of marine mammals
area is a rough calculation of about 200,000 common
including the harbor seal, California sea lion, harbor
murres in the eastern Strait of Juan de Fuca in early fall
porpoise, Dall’s porpoise, minke whale and killer whale.
migrating into inland waters to the south and north in
The harbor seal population in Washington’s inland
1978-79 (T. Wahl, unpubl. data). Similarly, wintering flocks
waters currently exceeds 17,000 animals, and forages on
of 1,600 Pacific loons and 2,000 Brandt’s cormorants were
a variety of seasonally abundant marine fish found in the
 CHAPTER 15: COASTAL/MARINE 403

Strait of Juan de Fuca, Strait of Georgia, San Juan Islands, The killer whale, or orca, is the best known and most
Hood Canal and Puget Sound. Primary prey of harbor spectacular cetacean found in Washington’s inland marine
seals includes Pacific whiting, herring, salmon, Pacific deeper waters. Studied in San Juan Islands, Strait of Juan
sandlance, tomcod, plainfin midshipman, various flatfish, de Fuca, and Strait of Georgia since the early 1970s using
and sculpins. Harbor seals are considered year-round photo identification, the population status, natural history
residents in these waters and appear to select prey such and behavior patterns of the southern resident killer
as Pacific whiting, herring, or salmon, which concentrate whales (pods J, K and L) are well known. Currently
in local areas to spawn. In the other cases they feed on the numbering a total of 90 individuals, these pods move
variety of less abundant but widely distributed prey such throughout the inland waters, feeding primarily on the
as Pacific midshipman, flatfish, or sculpins.97, 123, 130, 155 various runs of salmon returning to rivers which flow into
Dispersing into the inland waters of Washington and Puget Sound and the Strait of Georgia.13, 72, 131, 138 In addition
British Columbia in late summer from their breeding to the southern resident pods, which are fish eaters,
grounds in California, California sea lions forage primarily additional “transient” pods of killer whales occasionally
on returning adult salmon at river mouths in the fall, enter Washington’s inland waters from the Pacific Ocean.
spawning stocks of Pacific whiting in Port Susan and the Although very little is known about these “transient”
Strait of Georgia in fall and winter, and herring staging pods, they appear to forage primarily on other marine
and spawning areas off Cherry Point and in the Strait of mammals including harbor seals and gray whales.7, 8
Georgia. Other important prey of California sea lions in
Puget Sound includes market squid and dogfish. In many Coastal Headlands, Islets,
cases relatively large groups of California sea lions (500- and Marine Nearshore
1,000 animals) use haulout areas in close proximity to Marine Birds. Although most species associated with
deeper waters used as spawning grounds for preferred headlands and islets are found in low densities along much
prey (i.e., use of log-booms at Everett for feeding on of the coast, high concentrations of seabirds are found in
spawning Pacific whiting stocks in Port Susan).121, 122, 123 localized nesting colonies (Figures 7, 8, 9). Estimates of
As with the pinnipeds, the common cetaceans (harbor total breeding populations are available for some species,
porpoise, Dall’s porpoise, minke whale and killer whale) but for many there is considerable uncertainty. Moreover,
of Washington’s deeper marine waters use these areas for estimates from the period prior to documented population
foraging as well as for reproduction and social interactions. declines in the 1990s may no longer be accurate (e.g.
Harbor porpoise, while once considered common common murre populations in the region have varied
throughout Washington’s inland waters, are rarely seen substantially over the past two decades).11, 43, 195
today outside of the San Juan Islands, Strait of Georgia, In Oregon, the most abundant breeding species are
and Strait of Juan de Fuca.105, 138, 156 Causes suggested for Leach’s storm-petrel and common murre. Leach’s storm-
the disappearance of harbor porpoise from Puget Sound petrel is found at several major colonies (1966-1967
include mortality in fishing gear, effects of environmental surveys): Goat Island (high estimate of 535,800 birds),
contaminants, and increased vessel activity.138 Within Hunter’s Island (208,000), Whalehead Island (118,300),
Washington inland waters (Strait of Juan de Fuca, Strait Crook Point Island (101,500), and Saddle Rock (47,900).173
of Georgia, and San Juan Islands), the harbor porpoise The common murre is also abundant in Oregon but has a
stock was recently estimated at just over 3,500 animals.105 wider distribution: Since 1966, 23 sites have supported at
Important prey of harbor porpoise includes squid, shrimp, least 5,000 birds (131,481 at Three Arch Rocks, 28,531 at
and small schooling fish, such as herring and Pacific Cascade Head complex, 27,658 at Bird Rocks complex,
sandlance. 24,316 at Whalehead Island, 24,057 at Gull Rock [Curry
Dall’s porpoise are the small black and white cetacean County], 23,667 at Orford Reef, 23,220 at Cat and Kittens,
distributed throughout the inland waters of Washington. 19,146 at Yaquina Head, 15,117 at Hubbard Mound Reef,
This is the species frequently encountered riding the bow 14,377 at Gull Rock [Lincoln County], 14,018 at Redfish
wave of vessels in northern Puget Sound and the San Juan Rock, 13,839 at Mack Arch, 12,865 at Island Rock, 10,512
Islands. In open water this species leaves a characteristic at Twin Rocks, 8,031 at Cape Meares [south face], 7,679 at
“rooster tail” as it cavorts about. No comprehensive Castle Rock [Clatsop County], 7,488 at Tillamook Rock,
population assessment has been conducted for the species 7,079 at Pillar Rocks, 6,647 at North Coquille Point Rock,
in Washington’s inland waters, but the species is 6,000 at both Tower Rock and Mack Reef, 5,940 at Pyramid
considered common.138 Dall’s porpoise feed on small Rock, 5,342 at Gull Rock [Clatsop County], and 7 sites have
schooling fish such as herring and Pacific sandlance. supported at least 1,000 birds (3,198 at Face Rock, 2,968 at
Minke whales are the smallest and most common Goat Island, 2,855 at Rogue River Reef, 2,694 at Sea Lion
baleen whale found in northern Puget Sound, San Juan Rock, 2,506 at Brown Rock, 2,091 at an unnamed rock near
Islands, Strait of Juan de Fuca, and Strait of Georgia.138 Island Rock, 1,011 at Cape Blanco area).43, 112, 173 It should
No population estimate is available for this species in be noted that the common murre experienced a dramatic
Washington’s inland waters. As with the other cetaceans population decline along the Washington coast associated
found in inland waters, the minke whale’s diet consists with oceanic warming in the northeastern Pacific Ocean.195
primarily of small schooling fish. Although murre numbers have increased in the last few
years, their abundance remains below that prior to the
 404 Wildlife–Habitat Relationships in Oregon and Washington

Figure 7. Location of seabird

nesting colonies in
Washington (see text for
details). Sites referenced by
numbers on map are: 1) Smith
Island, 2) Protection Island,
3) Seal and Sail Rocks, 4)
Tatoosh Island, 5) Bodelteh
Island, 6) Carroll Island, 7)
Jagged Island, 8) Petrel Rock,
9) Cakesosta, 10) Rounded
Island, 11) Alexander Island,
12) Destruction Island, 13)
Willoughby Rock, 14) Split
Rock, 15) Point Grenville, and
16) Big Stack and Grenville
Arch.

1982-83 El Niño event. In Oregon, the population of Huntington Island, 2,200 at Rounded Island, 2035 at Big
common murres has been stable in recent years, although Stack [Grenville Pillar], 1135 at Cakesota).140, 167, 196, 197
colony attendance levels have fluctuated and productivity Other species, primarily associated with cliff faces, are
has been generally poor43 (R. Lowe, pers. comm.). All other generally less common. Surveys conducted in greater
seabird species associated with coastal headlands and Puget Sound in 1999 found at least 10,633 breeding pigeon
nearshore habitats in Oregon are found in far lower guillemots at 367 locations, primarily in southern Puget
numbers than murres.173 Sound, the Admiralty Inlet/eastern Strait of Juan de Fuca
The available data indicate that seabird colonies in region, and the San Juan Islands and vicinity.66 Belted
Washington support comparatively fewer birds than those kingfishers also nest in burrows in coastal bluffs but are
in Oregon. The most abundant breeding seabird species found in individual territories. A notable resident of this
in Washington (from baseline data 1978-1982) is the habitat is the peregrine falcon, which nests along the
Cassin’s auklet (estimated population is 87,600 birds, of northern outer coast of Washington (26 known sites of
which an estimated 55,000 occur at Alexander Island; the which 18 were reproductive at least one year between 1997
species’ breeding range is limited to 8 currently known and 1999198 (Washington Department of Fish and Wildlife
locations between Point Grenville and Seal and Sail database), the northern Puget Sound and San Juan Islands
Rocks).167 Other abundant species include fork-tailed area (18 sites of which 14 were reproductive between 1997
storm-petrel (1,600 at Carroll Island, 1,900 on two islands and 1999; WDFW database), and along the Oregon coast
in the Bodelteh Islands group), Leach’s storm-petrel (20 sites, of which 7 were reproductive between 1997 and
(20,000 at Jagged Island, 10,000 at Carroll Island, 2,600 at 1999; Oregon Department of Fish and Wildlife database;
Petrel Rock, 2,000 at Alexander Island), double-crested J. Pagel, pers. comm.).
cormorant (about 1,100 at Protection Island), pelagic Densities of birds foraging in nearshore and shelf
cormorant (7 of 61 colonies support a total of about 2,000 habitats usually are higher than those in oceanic waters
birds), western gull (perhaps 6,000-8,000 occur south of and include at least 30 species which are abundant at one
Destruction Island), glaucous-winged gull (over 10,000 at or more seasons during the year (see Life History matrix).
Protection Island), rhinoceros auklet (34,216 at Protection Depth associations of birds may be more pronounced or
Island, 23,600 at Destruction Island, 2,588 at Smith Island), more obvious in Washington, where the shelf is broad,
tufted puffin (7,800 at Jagged Island, 4,000 at Alexander than in Oregon, where the shelf is relatively narrow. Many
Island), and common murre, 1996-1998 (10,400 at Split species concentrate to forage at coastal outflows of major
Rock, 5,300 at Willoughby Rock, 5,000 at Grenville Arch, rivers, embayments and the mouth of the Strait of Juan
4,214 at Tatoosh Island, 3,610 at Carroll Island, 2,355 at de Fuca. Depending on tides, weather and season,
distribution in these habitats varies greatly.
 CHAPTER 15: COASTAL/MARINE 405

Figure 9. Location of seabird nesting colonies on the southern

Oregon coast (see text for details). Sites referenced by
numbers on map are: 13) North Coquille Point Rock, 14) Face
Rock, 15) Cat and Kittens, 16) Tower Rock, 17) Gull Rock,
18) Cape Blanco, 19) Orford Reef, 20) Redfish Rock, 21)
Island Rock, 22) Hubbard Mound Reef, 23) Rogue River Reef,
24) Hunter’s Island, 25) Crook Point Island, 26) Saddle Rock,
27) Mack Reef, 28) Mack Arch, 29) Whalehead Island, 30)
Twin Rocks, and 31) Goat Island.

Patterns of marine bird occurrence off the Washington

and Oregon coasts likely reflect conditions in the Arctic,
the southern hemisphere or elsewhere; reproductive
success or failure and feeding opportunities at staging
Figure 8. Location of seabird nesting colonies on the northern areas during migration or winter residence may determine
Oregon coast (see text for details). Sites referenced by populations of species in the region. There is great
numbers on map are: 1) Tillamook Rock, 2) Sea Lion Rock, 3) interannual variation in seasonal populations, some of
Bird Rocks, 4) Castle Rock, 5) Gull Rock, 6) Pyramid Rock, 7)
which may be due to variations in oceanic productivity
Cape Meares, 8) Pillar Rocks, 9) Three Arch Rocks, 10)
within the region, or to large-scale factors such as El Niño
Cascade Head, 11) Gull Rock, and 12) Yaquina Head.
or general oceanic warming.187 Much of the variation noted
is not fully explained, as in the case of fluctuating numbers
of regular transoceanic migrants like phalaropes and
jaegers in nearshore waters.
Brandt’s cormorants, brown pelicans, Heermann’s gulls
and Caspian terns concentrate in embayments or at
estuarine outflows and harbor entrances but also forage
in nearshore waters. In addition to these, large numbers
of gulls and alcids utilize nearshore areas. In particular
 406 Wildlife–Habitat Relationships in Oregon and Washington

waters farther offshore. Variations due to events like El

Niño may mask “natural” variation, and subsequent
analyses of cause and effect are extremely complicated.
Common murres historically have been widely dispersed
over the shelf, but flocks in Washington concentrate over
the inner shelf in less than 100 m depth177, 178 (to 150 m
depth).23 Similarly, summer-fall flocks of sooty shearwaters
often are found in the same area. These species are
attracted in to nearshore waters seasonally.
At present, breeding populations of a number of species
are incompletely known. The following estimates (prior
to reproductive failures in the 1990s) may no longer
accurately reflect the number of birds present.11, 43, 195
Washington and Oregon breeding numbers were
estimated to be 400,000 for Leach’s storm-petrel and 5,000
Brandt’s Cormorants at roost on Whale Rocks off Lopez for fork-tailed storm-petrel; 15 5,700 double-crested
Island, Washington. (Photo by Terence R. Wahl) cormorants, 16,600 Brandt’s cormorants and 12,000 pelagic
cormorants; 160 25,000 glaucous-winged and western
gulls; 176 up to 20,000 Caspian terns (in southwest
common murres and rhinoceros auklets may concentrate Washington and the lower Columbia River);111 7,660
in very large numbers seasonally, especially in summer pigeon guillemots67, 162 (although a recent estimate for
when anchovies are abundant. This season also draws very Puget Sound was 10,633);66 457,000 common murres;33
large numbers of sooty shearwaters to the nearshore, with 8,000-12,000 marbled murrelets; 67 87,600+ Cassin’s
historical estimates of 100,000 to 1 million present in Grays auklets;167, 168 25,000 rhinoceros auklets;33 and 28,000 tufted
Harbor and Willapa Bay and at the mouth of the Columbia puffins.33
River.9 As in other marine habitats, piscivores forage in
multi-species flocks, with gulls typically locating prey Marine Mammals. A total of 31 marine mammal species
schools, and then, depending on the season, other species have been reported from the marine waters of Washington
like shearwaters, cormorants, pelicans, alcids and jaegers and Oregon. 12, 138, 156 A number of these species are
arrive to forage until prey schools are dispersed or considered pelagic offshore species, which are rare visitors
decimated. to nearshore waters during El Niño events, or come ashore
The nearshore area is an important migration corridor as strandings along coastal beaches. The species
for a number of common seabirds (e.g., phalaropes, other composition of marine mammals found at coastal
shorebirds, and waterfowl), which nest in the Arctic, headlands, islets and nearshore waters (<20 m) of Puget
coastally, and inland in western North America. Sound, Hood Canal, San Juan Islands, Strait of Georgia,
Additionally, brown pelicans and Heermann’s gulls move Strait of Juan de Fuca, and along the outer coasts of
north in early summer and return south in the fall. In late Washington and Oregon is dominated by 3 species of
summer-early fall Brandt’s cormorants and common pinnipeds: the harbor seal, California sea lion and Steller
murres move north for the winter, many enroute to the sea lion. Additionally, elephant seals migrate into
Strait of Juan de Fuca and inland marine waters of Washington and Oregon coastal waters with a small
Washington and British Columbia; these species return breeding rookery recently reported on the southern
south in the spring to colonies in Oregon and California Oregon coast at Cape Arago.90 Two species of cetaceans
(T. Wahl, unpubl. data). In winter, the nearshore waters (gray whale and harbor porpoise) are commonly found
are utilized by loons, some grebes, cormorants, and gulls distributed in Washington and Oregon waters from just
and especially by large numbers of diving ducks, with outside breaker line to the inner continental shelf. In
scoters representing 50% of total numbers in November.23 addition, the sea otter is an important component of the
Large winter and migrant populations of 14 species nearshore marine environment of the northern
concentrate at deep water tidal fronts,183, 184 and lower Washington (Olympic Peninsula) coast as well.
numbers of other nearshore species forage As of 1996, the harbor seal population in Washington
opportunistically in the deeper waters offshore. Almost was estimated at over 34,000 animals. These animals use
all regions have highest numbers in winter, with loons, 319 haulout areas, primarily on intertidal islets, rock, reefs,
western grebes, Brandt’s cormorants, several species of sandbars and shoals.123 Areas of high harbor seal densities
gulls, 5 alcids (especially common murre) dominating the include Willapa Bay, Grays Harbor, Destruction Island,
seabird community. One diving duck, the oldsquaw, Giants Graveyard, Cape Alava, San Juan Islands, Skagit
forages locally over deep-water banks, often in flocks of Bay, Gertrude Island and Hood Canal. The largest haulout
hundreds.179 Three planktivore specialists—red-necked sites include Cape Alava/Bodelteh Island area for harbor
phalarope, Bonaparte’s gull, and common tern—are seals, California sea lions and Steller sea lions; Carroll
widespread during migrations. Island for Steller and California sea lions; and Giant’s
Nearshore waters are occasionally visited by large Graveyard/Toleak Point area, Grays Harbor, Willapa Bay,
concentrations of species normally associated with deeper and Columbia River for harbor seals.98 An example of the
 CHAPTER 15: COASTAL/MARINE 407

Oregon coastal estuaries (Coos Bay, Alsea Bay, Tillamook

Bay and Columbia River) also provide important habitat
for harbor seals. These estuaries provide protected areas
on intertidal sandbars and shoals used for pupping and
breeding, with pups present from early April through
June.12, 24, 107, 170 The largest haulout sites include Tillamook
Bay for harbor seals; Cascade Head for California sea lions
and Steller sea lion; Sea Lion Caves (Florence, OR) for
Steller and California sea lions; Cape Arago for harbor
seals, elephant seals, Steller sea lions and California sea
lions; Orford Reef for Steller sea lions (breeding rookery)
and California sea lions; and Rogue Reef for Steller sea
lions (breeding rookery), California sea lions, and harbor
seals.24
The nearshore waters of Oregon provide a variety of
food for harbor seals with important prey items in their
diet consisting of schooling fishes (Pacific whiting, smelts,
herring and Pacific mackerel), various flatfish, lamprey,
and salmonids.123, 148 During the winter, eulachon smelt
provide a seasonally abundant prey for harbor seals as
well.
California sea lions are seasonal migrants into
nearshore marine waters of Washington and Oregon that
results from a northward dispersal of subadult and adult
males from their rookeries in California and Baja Mexico
following their annual breeding season in June.12, 24, 114, 123
A wave of 5,000-6,000 animals move northward into
Oregon, Washington and British Columbia nearshore
waters in late summer and early fall, remaining in
Northwest waters until late spring, when the majority of
these animals return south to their breeding rookeries.123
California sea lions use haulout sites at or near the mouths
of most of the major rivers along the Oregon and
Washington coasts. In Oregon, haulout areas are located
at Rogue Reef, Orford Reef, Cape Arago, Sea Lion Caves,
Cascade Head, and Columbia River. In Washington,
haulout areas are located at Split Rock, Carroll Island,
Figure 10. Distribution of harbor seal haulout sites in Puget Bodelteh Island, Tatoosh Island, Waadah Island and near
Sound, Washington (small circles: 100-500 seals, large Everett in Puget Sound. The diet of California sea lion in
circles: >500 seals; data from Washington Department of Fish Washington and Oregon consists primarily of seasonally
and Wildlife). abundant schooling species such as Pacific whiting,
herring, Pacific mackerel, eulachon, salmon, and squid.123,
148
distribution of harbor seal haulout sites is shown in Figure Movements and distribution of California sea lions
10. through nearshore waters of Oregon, Washington and
Bays and estuaries along the outer coast and inland British Columbia has been correlated with spawning
waters of Washington provide important pupping and aggregations of their prey, and indicate the ability of
breeding areas for harbor seals. Harbor seal pups are California sea lions to cue into locally abundant
present along the outer Washington coast from late April concentrations of these species.123 This species has been
through July; pups are present in the inland waters of reported to move up the Columbia and Willamette rivers
Washington from late June through September.12, 107, 155, 170 following spawning runs of salmon and eulachon as
Important prey of harbor seals in Washington waters well.12, 123
include Pacific whiting, herring, salmonids, smelts, Steller sea lions occur year round in Washington and
plainfin midshipmen and various flatfishes.12, 123, 130 Oregon nearshore waters, and include both breeding and
Harbor seals are the most abundant pinnipeds in non-breeding animals. The main breeding rookeries for
nearshore marine waters of Oregon, with a population of Steller sea lions along the Oregon coast are located at
over 9,000 animals using 101 haulout areas along the Rogue and Orford reefs; relatively small breeding
Oregon coast.123 Areas of high harbor seal densities in rookeries are reported at Sea Lion Caves and Three Arch
Oregon include coastal headlands, reefs and islets at Rogue Rocks.24, 121 Additional haulout locations in Oregon are
Reef, Orford Reef, Cape Arago, and Tillamook Head. located at Cape Arago, Cascade Head, and Columbia
 408 Wildlife–Habitat Relationships in Oregon and Washington

River. Seasonal abundances in Oregon nearshore waters breaker line to out over the continental shelf.36, 81, 106 The
range from 1,000-4,000 animals, with peak counts Oregon and Washington coastal stock was estimated at
occurring during the breeding season in early summer. over 44,000 animals.106
Although both adult males and females are present in In the nearshore waters off the coast of Washington
Washington, no breeding rookeries occur. Haulout from Destruction Island to Neah Bay is found a growing
locations are found along the outer Washington coast at population of sea otters. Extirpated from the coast of
Split Rock, Carroll Island, Bodelteh Island, Cape Alava Washington in the early 1900s, the sea otter was
and Tatoosh Island. Seasonal abundances range from 500- translocated to the Olympic Peninsula coast in 1969 and
1,500 animals along the outer Washington coast. Relatively 1970 from Amchitka Island, Alaska.96 The species has
small numbers of Steller sea lions occur at haulout reestablished itself off the coast with a growing population,
locations in the inland waters of Washington, although which (as of July 1999) is estimated at 605 animals. Sea
500-1,000 animals move through the Strait of Juan de Fuca otters can be expected to play a significant role in shaping
and into British Columbia waters annually to feed on the nearshore urchin/kelp community along the Olympic
herring spawning in the Strait of Geogia north of Nanaimo. Peninsula coast, with a diet consisting of sea urchins,
Similar to California sea lions, the primary diet of Steller clams, Dungeness crab, octopus, and other benthic
sea lions in the nearshore waters of Washington and invertebrates.104
Oregon consists of Pacific whiting, herring, salmon,
lamprey, rockfish, flatfish, and squid.148 Marine Shelf and Oceanic
The gray whale is one of the best known cetaceans Marine Birds. Seabird populations are characterized by
because of its annual migration along the west coast of great seasonality as well as distribution by general depth
North America. Every year, the Eastern North Pacific stock “habitats” over shelf, slope, and offshore/oceanic waters.
of gray whales migrates through the nearshore and coastal Off Washington, there is a general change in species
waters of Washington and Oregon traveling from breeding composition over the shelf at about 100 m depth, with
grounds in Baja Mexico to feeding areas to the north along albatrosses, fulmars, shearwaters (other than sooty
the west coast of North America and Alaska. This stock of shearwater), and storm-petrels becoming more numerous
gray whales has recovered from whaling and over- offshore over the deeper “outer” shelf. Off the Oregon
exploitation in the late nineteenth century, and is estimated coast these changes may be less apparent where the
in excess of 26,000 animals. Because of this recovery, the continental shelf is relatively narrower. The spring
Eastern North Pacific stock of gray whales was removed population includes a large number of coastal breeders,
from the List of Endangered Species in 1994. 153 The such as fork-tailed storm-petrel, Leach’s storm-petrel,
southward migration takes the gray whale population common murre, and Cassin’s auklet. In late spring, large
from their feeding grounds to breeding lagoons in Baja numbers of shearwaters arrive from the southern
Mexico, with whales passing along the Oregon and hemisphere for the summer, and birds from Alaska and
Washington coasts from late November to early January.89, Canada arrive as well. Diversity increases during fall
153
The northward dispersal and migration of gray whales migration when migrant phalaropes, jaegers, terns and
from their breeding ground brings them into the nearshore alcids, in addition to summer residents, not only
waters of Oregon and Washington waters from late substantially increases the diversity of abundant species
February to late May.89, 153 Although the majority of the to 19 but also accounts for the largest overall seabird
population migrates to feeding grounds in the Bering and populations of the year.23, 182, 184 By winter, many of the
Chukchi Seas, a small proportion of the gray whale northern birds have moved south, with large numbers of
population remains in nearshore areas along the Oregon, northern-breeding gulls and fulmars remaining. In
Washington, and British Columbia coasts to feed during general, the bird community associated with shallower
summer months.51, 153 Foraging aggregations usually are water is dominated by diving or pursuit plunging seabirds
found along the Oregon coast from near Yaquina Head to that prey on pelagic fish such as herring, anchovy,
Depoe Bay. Along the Washington coast, feeding areas are eulachon, and smelt (e.g., murres, auklets, puffins, and
located in shallow water areas of Grays Harbor, Cape sooty shearwaters), while the deeper oceanic waters
Alava, Cape Flattery, western Strait of Juan de Fuca, and support surface-feeding birds (e.g., gulls, albatrosses,
along Whidbey Island (Saratoga Passage). Gray whales fulmars, other shearwaters, and storm-petrels) that feed
are benthic feeders, and usually forage in areas where they largely on squid and plankton (including jellyfish).6, 145, 180,
189
find dense concentrations of amphipod crustaceans, Jaegers and skuas are generally found in deeper waters,
polychaete worms, sand shrimp, and herring spawn where they pirate prey from offshore gulls.
(roe).37, 53, 138, 153, 190 Individual gray whales in nearshore For offshore populations especially, variability is one
waters of Washington and British Columbia have been of the most consistent features at temporal scales ranging
reported returning to the same areas to feed year after from the short-term187 to long-term cycles, which may
year.37, 53, 153 persist for decades. Even more so than in nearshore waters,
The smallish, olive gray harbor porpoise is the most there is great inter-annual variation in seasonal
common small cetacean found in nearshore marine waters populations present in offshore waters. Some of this is due
of Oregon and Washington. Shy by nature, this species is to variations in oceanic productivity within the region and
found along the Pacific Ocean coast from just outside the some may be due to large-scale variations (El Niño, Pacific
 CHAPTER 15: COASTAL/MARINE 409

Decadal Oscillation). Some variations are due presumably

to smaller-scale variations outside the region, such as at
Arctic breeding grounds. Much of the variation is not well
understood.
The shelf area, like the nearshore coastal waters, is a
regular migration route for a number of species, which,
though they do not forage in numbers in the habitats, are
noted resting or foraging on occasion. Loons, particularly
Pacific loons, many geese, dabbling and diving ducks,
shorebirds and lost landbirds transit the shelf area. Pigeon
guillemots, apparently migrating north for the winter, are
noted in August (T. Wahl unpubl. data), and small
numbers of Xantus’ murrelets dispersing from southern
breeding areas to Oregon-Washington offshore waters177
are examples of less-obvious occurrences. Sabine’s Gulls migrating off Grays Harbor, Washington.
Birds concentrate at current boundaries and (Photo by Terence R. Wahl)
topographically induced upwellings.23 These oceanic
fronts or ‘rips’ at the edges of coastal upwellings
concentrate small zooplankton and their larger predators. also to be dominated by rhinoceros auklets, tufted puffins,
These attract planktivorous storm-petrels, phalaropes and and 4 species of gulls (western, herring, glaucous-winged,
terns and, subsequently, jaegers and gulls. Nocturnal and California).
foraging, which is known for a number of species, is Compared to oceanic waters, seabird densities are
apparently more important to birds over the shelf and considerably higher23 along the shelf edge and slope, with
oceanic waters than to birds inshore. Oceanic birds are seasonal densities ranging from about 2 to 7 birds/km2.
characterized by patchy distribution, which reflects prey Seasonal densities were highest during the summer (when
distribution and prey detection, and a response to activities again tubenoses dominated), but relatively high densities
like commercial fishing, which attracts birds. were also found winter (when gulls, fulmars, and Cassin’s
The water mixing zones of the shelf slope and outer auklets dominated) and spring (when Leach’s storm-
shelf edge support a high diversity of seabirds capitalizing petrels were prevalent). Seabirds here tended to
on the biological richness of this area. Over 20 species of concentrate into locally high densities, especially along
birds characterize these regions. Five breed along the the seaward edges of Heceta and Swiftsure banks, the
Oregon and Washington coasts (fork-tailed storm-petrel, Cape Blanco upwelling center, and behind shrimp and
Leach’s storm-petrel, Cassin’s auklet, rhinoceros auklet, factory trawlers.23, 181 In these waters, Briggs et al.23 found
and tufted puffin); while 3 other breeders (western gull, local densities of sooty shearwaters exceeding 2,000 birds/
glaucous-winged gulls and common murre) are found km2.
here, they occur in much smaller numbers than closer to Farther offshore over oceanic waters (>2000 m deep),
shore.23, 177 Conspicuous southern hemisphere breeders Sanger154 found the seabird composition to change further.
(from Australia, New Zealand, and the Antarctica region) Black-footed albatrosses predominated, except during the
found here include the pink-footed shearwater, Buller’s winter when most of the population of this species had
shearwater, sooty shearwater, and south polar skua. returned to its Hawaiian breeding grounds. Fork-tailed
Hawaiian breeders include both the black-footed albatross storm-petrel and Leach’s storm-petrels are common
and Laysan albatross. The California gull breeds inland. during the summer months. By winter, northern fulmars,
All the remaining dominant species breed in Alaska and herring gulls, glaucous-winged gulls, and black-legged
Canada, and include the northern fulmar, red phalarope, kittiwakes comprise the bulk of the population.
red-necked phalarope, Pomerine jaeger, long-tailed jaeger, Few estimates have been made of seasonal offshore-
Arctic tern, and 5 gulls (Bonaparte’s, herring, Thayer’s, shelf populations. Briggs et al.23 estimated 1.8 million birds
Sabine’s, and black-legged kittiwake). present at one time in late summer for an oceanic, shelf
The highest seabird densities are found in shelf waters, and nearshore waters study area off Washington and
due to the high numbers of local breeding birds and Oregon. Wahl178 roughly estimated a seasonal total of 1.2
nearshore migrants. Briggs et al.23 found shelf water million off Washington. Maximum densities were in
densities of between 40 and 50 birds/km2 during the August-September, with highest densities over the
summer months, due mainly to the nearly 500,000 continental shelf and lower numbers offshore in oceanic
common murres that nest in Oregon 23 and the waters.23, 178 Foraging breeding birds aggregated to within
approximately 88,000 Cassin’s auklets nesting along the about a 50 km radius from colonies.23 Estimates of breeding
outer coast of Washington.167 Also, tremendous flocks (up population size of a number of offshore species may not
to 100,000) of sooty shearwaters invade the shelf waters be very accurate, particularly following the widespread
to feed on northern anchovies during the summer months, reproductive failures in the 1990s (see “Coastal Headlands,
with local densities23 exceeding 1,000 birds/km2. Wahl177 Islets, and Nearshore” for other population information).
found the inner shelf (20-100 m deep) seabird community
 410 Wildlife–Habitat Relationships in Oregon and Washington

Marine Mammals. At least 24 species of cetaceans and 5 The larger whales found near the shelf edge or over
pinnipeds are known to occur in the marine waters off slope waters include the humpback, fin, Baird’s beaked,
Oregon and Washington.18, 81, 115, 156 However, some species and killer whales. Green et al.81 found all 4 species most
are rarely seen, or at least rarely identified. These include common during the summer months, although fins were
offshore species such as Sei whales, pygmy sperm whales, absent during the spring; humpbacks were absent during
and Stejneger’s, Hubb’s, and Cuvier’s beaked whales. the winter, and Baird’s beaked whales were missing
Most beaked whale records are from strandings. Other during the fall and winter. Killer whales were found year-
species of beaked whales (especially mesoplodonts) may round. Most of the humpback whale sightings by Green
also occur in the deeper offshore waters of Oregon and et al.81 occurred immediately south of Hecata Bank, on
Washington. Sei whale records derive mostly from old Swiftsure Bank near the Canadian border, or off the mouth
whaling logs. Blue and northern right whales are also very of the Columbia River. Nearly all the fin whales were
rare, but both have been observed off Oregon and observed in a lower to mid-slope region called Newport
Washington in recent years.82, 152 Others, such as false killer Valley approximately 85 to 90 km west of Newport,
whales, short-finned pilot whales, striped dolphins, and Oregon.81 Similarly, most of the killer whale sightings (3
common dolphins, may intrude into Oregon and pods) occurred near the shelf edge about 35 to 40 km off
Washington waters during years of very warm water, Newport. Other sightings of pods (2) occurred near the
although pilot whales may have been much more common shelf edge off Washington. Four of the 5 groups of Baird’s
off the coasts than they are now. A record of a single beaked whales observed by Green et al.81 occurred near
belukha whale156 is obviously extralimital. the lower edge of the slope. A single group of 7
Extralimital strandings of ringed and ribbon seals have mesoplodont whales was observed by Green et al.81 in
been reported in California.115 Although neither has been slope waters of Washington.
recorded for Oregon or Washington, it is likely that these Photographic identification studies on humpback
Arctic seals passed through Oregon and Washington whales have revealed that the animals seen off Oregon
waters before reaching California. and southern Washington are part of a distinct feeding
Gray whales, harbor porpoise, harbor seals, California aggregation that spends the spring, summer, and fall along
sea lions, and Steller sea lions are largely denizens of the the coast of California, Oregon, and Washington, and
coastal and inner shelf waters. All but the gray whale are migrates primarily to Mexico and Central America in
occasionally observed in shelf edge waters, though most winter.39, 40 This population numbers just over 900 and has
of the at-sea sightings of pinnipeds occur near coastal been increasing at about 7-8% per year.35 Humpback
rookeries or haulouts.18 whales identified off northern Washington near the British
Eleven species of marine mammals are character- Columbia border show a low rate of interchange with
istically found in the shelf edge and slope waters. The three locations to the north and south and consist of a relatively
most common species are the Pacific white-sided, Risso’s, small number of whales that show strong seasonal site
and northern right whale dolphins. Green et al.81 estimated fidelity and migrate in winter to breeding grounds in
the spring Pacific white-sided dolphin population to be Mexico, Hawaii, and off Japan.41, 52, 169
nearly 40,000 animals (including offshore animals), a Most (68%) of the 19 elephant seals observed at-sea by
number far exceeding any other population of marine Bonnell et al.18 were found in shelf edge or slope waters.
mammal occurring off Oregon and Washington. Green et These deep diving animals spend little time at the surface
al.’s81 estimate for spring and summer Risso’s dolphins (thus, are rarely seen) when occurring in deeper waters,
was nearly 8,000. Both species were most common during and may be more common than they appear off Oregon
the spring and summer, less so during the fall, and rare and Washington. Elephant seals equipped with radio
during the winter months. Northern right whale dolphins, transmitters have been tracked by satellite from California,
on the other hand, were found to be most common during through Oregon and Washington, to the Gulf of Alaska.60
the fall and absent during the winter.81 Risso’s dolphins In contrast, migrating fur seals are easily seen as they rest
were far more common over the shelf slope than the shelf at the surface. When standardized for differences in survey
edge (and entirely absent in offshore waters). These effort, Bonnell et al.18 estimated that about 27% of the
dolphins feed exclusively on the squid found in northern fur seals occurred in outer shelf or slope waters.
abundance in the slope currents. Interestingly, all three In total, Bonnell et al.18 recorded 172 sightings of fur seals,
species of southern-originating dolphins were rarely seen most (87%) between January and May, with an estimated
off southern Oregon in the vicinity of Cape Blanco. It is April peak abundance of about 7,000 animals.
likely these temperate-water dolphins avoid the colder, Only 5 species of marine mammals dominate the
upwelled water found off Cape Blanco. The other smaller deeper offshore waters: Pacific white-sided dolphins,
cetacean using shelf edge and slope waters is the Dall’s northern right whale dolphins, Dall’s porpoise, sperm
porpoise. Overall, these porpoise seem to be equally whales, and northern fur seals.18, 81 However, the rarely-
common in outer shelf and slope waters, although they seen mesoplodont whales are thought to prefer these
shift onto the shelf during the summer and fall, following waters, and Risso’s dolphins have been seen far offshore
schooling fish and squid,70 and move offshore during the during NOAA ship-board surveys (M. Dahlheim, NOAA/
winter.81 Although they are easily observed, Green et al.81 National Marine Mammal Laboratory, pers. comm.).
estimated that a little over 2,000 were found year-round.
 CHAPTER 15: COASTAL/MARINE 411

As with seabirds, the distribution of marine mammals scientists believe that the oceanic warming is due to
is linked to their diet and foraging habitats. Marine decadal oscillations and that the current conditions will
mammals that feed on schooling pelagic fish or shallow- not persist indefinitely.92 Significant temperature changes
water squid (e.g., seals, sea lions, harbor porpoise, and of as much as 10°C in a relatively short period (several
Dall’s porpoise) are found closer to shore, while squid- years) have been documented in some parts of the
eating marine mammals (e.g., dolphins, Dall’s porpoise, world.101 Significantly, the Pacific Decadal Oscillation
sperm whales, beaked whales, and fur seals) are found in (PDO) has closely interacted with the major El Niño
the deeper waters. Humpback whales are generally found episodes of the past two decades. Every few decades, the
where plankton or herring concentrate, regardless of water North Pacific basin shifts from being unusually cold water
depth, and elephant seals feed on deep-water fish, temperatures in western and central parts of the basin and
especially hagfish and squid.45 warm in the east to an essentially opposite pattern. The
Since 1985, 4 species of sea turtles have been found PDO phase in effect since the 1970s, which corresponds
stranded along the Oregon and Washington coasts: to the recent severe El Niño events, is characterized by
leatherback, loggerhead, green, and olive ridley (J. warmer waters in the eastern North Pacific basin. The
Scordino, NOAA, pers. comm.). It is likely that records of relationship between El Niño episodes and the PDO is
the latter three warm-water species represent extralimital not well-understood.101, 113, 143
sightings, possibly of disoriented animals, or those The conditions that influence the occurrence of El Niño
sickened by the cooler temperate waters. Leatherback are best described in relation to typical weather and
turtles, however, appear to be a regular summer visitor to oceanic conditions. Under normal conditions, cooler sea
shelf and slope waters. Bowlby et al.21 reported on 16 surface temperatures, a shallower thermocline depth, and
leatherback turtles sighted during the marine mammal a lower sea level prevail in the eastern tropical Pacific
and seabird surveys conducted under Brueggeman et al.25 Ocean relative to areas further east. This unequal state is
Turtles were sighted during the months of June, July, and influenced and maintained by the Pacific trade winds. The
September, with the 9 July sightings occurring farther western Pacific Ocean absorbs more heat and this warmer
inshore (shelf waters). air rises and moves east, where it sinks. The resulting
easterly trade winds occur at lower altitudes than westerly
El Niño and La Niña Events winds, and they push sea water (which is warmed by the
The effects of El Niño have been well documented and sun) westward via the Equatorial Current. Cold marine
publicized, perhaps due to the severity and frequency of waters in the eastern tropical Pacific Ocean are maintained
these events in the past two decades. The phenomenon by the Peru Current and equatorial as well as coastal
influences weather conditions in marine environments, upwellings, which transport cold subsurface water to the
which in turn produce powerful storms and adverse surface. The resulting temperature and pressure gradients
weather conditions that impact humans and wildlife between the east and the west are maintained and further
populations over vast areas of the planet. Although first strengthened by a positive feedback.10
documented in 1726, El Niño has occurred for tens of
thousands of years. The ocean warming events vary in
their intensity and timing, although in recent years the
events have occurred, on average, every 3.8 years, with
very strong episodes occurring every 9.9 years.147 The two
greatest El Niño events in recorded history occurred in
1982-83 and 1997-98.102
The cold water counterpart of El Niño is La Niña. The
conditions in the Pacific region during La Niña episodes
are exactly opposite those encountered during an El Niño
event. Sea surface temperatures are anomalously high in
the eastern equatorial Pacific and the east-to-west trade
winds are very strong. In the Pacific Northwest, winters
are colder and wetter in a La Niña year than in other years.
The fluctuations between El Niño and La Niña states are
often called the El Niño Southern Oscillation (ENSO).
A recent long-term warming of oceanic waters in the
northeast Pacific may have intensified the severity of El
Niño episodes. Some scientists believe this change in
temperature is related to an increase in global surface
temperatures. In the last century, average temperatures
have increased by about 0.5°C and are expected to rise
another 1.0 to 3.5°C by 2100.116 Some argue that the increase Figure 11. Schematic of the normal and El Niño conditions
in temperature is the result of anthropogenic factors, prevailing in the equatorial Pacific; from Frankignoul.75
especially greenhouse gases, which trap rising heat. Other
 412 Wildlife–Habitat Relationships in Oregon and Washington

During El Niño, the pool of warm water in the western narrower in March 1983.10 Sea surface temperatures, which
Pacific expands eastward, changing wind patterns. normally range from 7°C to 11ºC, reached a maximum of
Normally strong, easterly winds may weaken due to 18ºC in July 1983, extending out to at least 200 km from
change in air pressure, and the normally weak westerly the Oregon coast, and reaching depths of over 100 m.80
winds strengthen. With the decrease in easterly wind
Marine Birds. The most common responses by marine
speed, the accumulated warm water on the west is
birds to El Niño or other oceanic warming conditions have
released in the form of a Kelvin wave, readjusting sea level.
been altered species distribution, and changes in species
As a result, warm water is transported eastward, the
abundance, reproductive output, and behavior. For marine
equatorial upwelling process decreases, sea surface
birds off the Washington and Oregon coasts, these changes
temperatures and thermocline differences diminish, and
likely reflect reductions in the availability of suitable prey.
sea level in the eastern Pacific rises by 0.2-0.4 m. As higher
Ocean temperatures off the California coast during the
temperatures shift to the east, so do patterns of
1982-83 El Niño were 1-4°C higher than normal.
precipitation. Usually arid middle Pacific islands and
Concomitant with this temperature increase, zooplankton
coastal regions of Ecuador and Peru are drenched with
densities declined by 90% below normal during spring
rains and tormented by winds10, 75, 150 (Figure 11). The
1983; phytoplankton declined as well and was distributed
magnitude of each El Niño episode depends on many
much deeper in the water column. In addition, rockfish
factors, including timing, location, magnitude of oceanic
and northern anchovy populations, which support large
warming, and phase of interdecadal climatic oscillations.
and diverse populations of seabirds, experienced lowered
Consequently, it is not yet possible to predict the onset of
reproductive success.22
such an episode. Once El Niño starts, however, it is
Perhaps the most significant and well-documented
possible to determine its spatial and temporal distribution
avian response to El Niño is population change.
with current climate models.159
Populations of tufted puffins and Cassin’s auklets in
How a given animal is affected by El Niño depends on
central coastal California were reduced by 50% in response
a variety of factors. The most important factors are the
to El Niño conditions.4, 63 Over the period 1972-1998, and
severity of a particular warming episode, and its
particularly during the 1990s, a decade of prolonged
geographical extent. Not all El Niños influence conditions
decline in ocean productivity, Wahl and Tweit187 found
as far north and east as the Oregon and Washington coasts.
significant decreases in abundance of a number of species
During less intense episodes, for example, coastal waters
off southwest Washington, apparently reflecting a decline
off California are the northern limit of El Niño conditions.
in biological productivity. Most dramatic was the 90%
Only a few, severe and long-lasting El Niños, particularly
decline in sooty shearwaters shown for Washington and
the 1940-41, 1957-58, 1982-83, and 1997-98 events, widely
California (see also Veit et al.).174 Also decreasing were
affected many marine species across the Pacific Ocean and
flesh-footed and Buller’s shearwaters and regionally
along the coasts of the Americas.10
breeding Cassin’s auklet, tufted puffin, and common
Of all the changes related to El Niño, the deepening of
murres. Especially noticeable were changes in bird
the thermocline has by far the greatest impact on the
densities before and after the “step,” a widespread, subtle
marine fauna of the eastern and central Pacific Ocean. The
change in environmental variables that occurred in the
eastern Pacific Ocean contains some of the richest waters
1970s,64 and before and after about 1990 when ocean
on the planet in terms of biological productivity. Nutrients
productivity decreased and stayed low for years. Habitat
from the breakdown of organic materials sink and
switching was evident during the 1990s off Grays Harbor,
accumulate in the cold waters below the thermocline.
with rhinoceros auklets moving more inshore where prey
Phytoplankton has ready access to nutrients and takes
abundance was maintained, while offshore productivity
advantage of sunlight for photosynthesis when the
declined. Common murres also moved inshore, though
thermocline is shallow. The Kelvin wave causes the mixed
to a lesser extent, while sooty shearwaters declined in
layer of water above the thermocline to increase and the
numbers there as well as offshore.187
productivity to decrease, thereby affecting the entire food
Populations of rock sandpipers demonstrated an
web of the eastern Pacific. Upwelling, if it occurs at all,
unusual trend following the 1982-83 El Niño episode. Their
can no longer replenish cold, saline, and nutrient-rich
abundance in coastal Christmas Bird Count circles was
water from subsurface layers, because the thermocline is
higher prior to the 1982-83 El Niño than it was after that
lower, extending the warm layers of water too deeply.
event.28 It was speculated that the species underwent a
During upwelling, water is transported from depths of
range contraction during a prolonged period of oceanic
between 40 and 80 m, but during severe El Niño episodes,
warming.28
the thermocline deepens by another 100-150 m, effectively
Brandt’s and pelagic cormorants, western gulls,
negating the upwelling process.10 At the peak of the
common murres, pigeon guillemots, and Cassin’s auklets
1982-83 El Niño in March there was a 5-fold reduction in
experienced low nesting success during El Niño years.63
absolute productivity at a cross-equatorial transect, and a
In 1983, the hatching success of common murre chicks on
20-fold decrease near the Galapagos Islands. A normally
Farallon Islands, California, was 31.7% lower than in other
400 km wide nutrient-rich section of the Pacific ocean
years (i.e., 1972-1983) and the fledging success rate was
along the coast of Peru and Ecuador was diminished to a
only 15%, compared to 94% in other years.14 Brandt’s
30-50 km wide band in November 1982, and became even
 CHAPTER 15: COASTAL/MARINE 413

cormorants abandoned their nests in large numbers

during the 1983 breeding season along the Oregon coast
from Coquille Point Rocks to Sea Lion Caves. Some
cormorants managed successfully to raise chicks by
breeding earlier in the season.91 Chick production by
murres on the Oregon coast was lower than during pre-El
Niño years. 80 Pelagic cormorants suffered low
reproductive success all along the Oregon coast from Cape
Meares to Coos Bay. Brandt’s cormorants had lower
reproductive success during 1982-83 and during the El
Niño of 1987-88, but recovered in subsequent years.
The diet of cormorants and gulls at the Farallones,
California during the 1976 and 1978 warm water episodes Figure 12. The number of breeding common murres in 5
and the 1982-83 El Niño was more diverse and differed in California colonies and 28 colonies on the outer coast of
composition from the diet in non-El Niño years. Seabirds Washington, 1979-1990. California counts are reduced by a
were exploring different locations in search of food.2 The factor of 10 to complement the y-axis scale, fitted to the
very few cormorants, common murres, and gulls that were Washington counts (from Ainley et al.3).
seen at the Farallones during 1983 spent more time in
search for food than during normal years. As a result, the During the 1982-83 El Niño, a number of warm-water
feeding frequencies decreased and adults brought fewer species, such as black-vented shearwater (Puffinus
fish to chicks. With a decrease in the abundance of juvenile opisthomelas), black storm petrel (Oceanodroma melania),
rockfish (Sebastes spp.) during the warm-water episodes Craveri’s murrelet (Endomychura craveri), and brown
in 1973, 1976, 1978, and 1983, common murres and pigeon booby (Sula leucogaster) were present near the Farallon
guillemots switched to other prey species, such as Islands in California. Unusually high numbers of Laysan
anchovies, squid and smelt.5 The change in diet and an albatrosses and black-footed albatrosses were seen off the
increase in foraging time likely decreased the caloric intake coast of British Columbia during the 1992-93 El Niño
of nestlings, contributing to the lowered reproductive episode. 77 A breeding range expansion occurred in
success of the seabirds. California when the Heerman’s Gull expanded north into
Egg laying among common murres during the 1983 southern California, apparently in response to large
breeding season was not as synchronized as it was in numbers of red crabs (Pleuroncodes planipes) that migrated
normal years, exposing chicks to predators such as western to that region from the south during El Niño.56
gulls.14, 163 Food shortage during the 1982-83 episode Populations of some seabirds have been slow to recover
contributed to conspecific predation on western gulls from the impacts of El Niño events. Prior to the 1982-83
chicks.164 Some of the birds arrived at their breeding event, Brandt’s cormorant numbers at the Farallon Islands,
grounds later in the season than usual (e.g. pigeon California, ranged between 15,000 and 25,000 birds. The
Guillemots arrived in mid-July instead of June and early population crashed, and had increased to only 3,000 -
July at Farallon Islands, California) and did not breed at 10,000 by 19943 (Figure 13). The common murre population
all,2 or in much lower numbers.163 During the 1982-83 in California has increased by only 1-2% per year since
event, double-crested cormorants produced less than half 1982-83 in comparison to the 7.8% annual increase prior
the normal number of nestlings. to that El Niño event3 (Figure 12). Common murres were
Habitat switching was evident during the 1990s off not as fortunate in Washington. It took until 1987 to recover
Grays Harbor, presumably in response to declining prey from a significant drop in population (from 18,335-31,520
abundance offshore. Common murres, as one conspicuous birds breeding during 1979-1982 to only 3,190 in 1983),
example, have historically been widely dispersed over the and then the population was decimated to levels similar
shelf with flocks concentrating over the inner shelf (100 to those in the 1982-83 episode by the 1987-88 El Niño3, 110,
m to 150 m depth).23 Numbers of murres decreased during 195
(Figures 12, 13). On the other hand, rhinoceros auklets
years of low productivity in the 1990s, and birds were recovered quickly after the 1982-83 and 1986 El Niños
relatively closer inshore, usually in nearshore waters.187 during which they experienced only a minor decrease in
Even more noticeable was the shift inshore of rhinoceros numbers. 3 Double-crested cormorants along the
auklets; their numbers increased throughout the time Washington coast appear to have recovered from El Niño
period of 1972-1998. Sooty shearwaters declined in overall episodes195 (Figure 13).
numbers nearshore as well as offshore.174, 187 Observations Some resident marine species do not appear to be
and anecdotal reports in the 1990s suggested that a number impacted by oceanic warming. Boersma and
of species may have concentrated in inland marine waters Wheelwright16 suggested that storm petrels nesting along
(Strait of Juan de Fuca) during migrations or, in the case the Washington and Oregon coasts were better able to cope
of regional breeders, during failed breeding seasons. These with the fluctuating environmental conditions through
included storm-petrels, brown pelicans, phalaropes, their ability to leave their eggs unattended. Resistant to
jaegers and gulls (T. Wahl, unpubl. data). chilling and largely protected in underground burrows,
petrel eggs may be left unattended for 31 days in
 414 Wildlife–Habitat Relationships in Oregon and Washington

they may have benefited from salmon hatchery smolts

released in the Yaquina estuary the summer of 1983.11
Although adult mortality of pigeon guillemots increased
substantially and resulted in a nesting rate 19% below
normal, there were no changes in the number of chicks
that hatched and fledged per nest.91 In contrast to common
murres and cormorants, pigeon guillemots feed more on
epibenthic fish so they were less affected by a decrease in
primary productivity.
Marine Mammals. Despite a number of studies conducted
on marine mammal responses to El Niño conditions in
California58, 59, 109 and Alaska,199 very little information is
available pertaining to marine mammals in the coastal
areas of Washington and Oregon. The summary below is
drawn from information relating to California sea lions at
their California breeding areas. The California sea lion is
distributed along the Pacific coast of North America from
British Columbia to the southern tip of Baja California and
Gulf of California, Mexico. The primary breeding areas
are in the Channel Islands off California. There are no
breeding areas for this species north of California.
Although it is not clear whether the magnitude of impacts
to sea lion populations noted in California occur in marine
mammal populations in Washington and Oregon, this
seems possible during severe El Niño events when oceanic
productivity is altered off the coasts of Washington and
Oregon.
Birth rates at the breeding rookeries in the Channel
Islands declined substantially following the 1982-83 El
Niño. The decrease in number of births at all rookeries
there ranged from 30-62%. By 1984, birth rates had
returned to pre-El Niño levels at San Nicolas Island, but
lower birth rates prevailed at other islands through 1986.59
The reduced birth rates were at least partially due to a
lower number of females present at breeding sites.74
There was no apparent increase in female mortality
during the 1982-83 El Niño. Females have high fidelity to
breeding grounds, but the very strong El Niño of 1982-83,
in combination with human disturbances, apparently
forced them to disperse.74 Indeed, counts of sea lions were
Figure 13. Seabird fluctuations and the occurrence of warm
much higher at their non-breeding rookeries, the Farallon
water events on the Washington outer coast, 1979-1990 (from
Wilson195). Islands, during the summers of 1983 and 1984. Adult
males, which fast during the breeding season and migrate
north afterward, were not affected.93
southeastern Alaska16 and up to 17 days on the Farallon Prenatal growth of California sea lion pups did not
Islands, California.2 This adaptation enables the adult birds appear to be affected. Females were already pregnant at
to spend more time in search of food. Similarly, Leach’s the onset of El Niño, and the mass of pups born that year
storm petrel feed further offshore, in areas not influenced did not differ significantly from non-El Niño years.17
by upwelling. 80 In fact, numbers of breeding petrels Growth during the first month of life did not differ
increased off southern California during periods of significantly either, but pups began to loose mass when
warmer ocean temperatures. 175 Of 5 species which they reached two months of age.17, 95 The number of
increased in abundance (birds/km) off the Washington foraging trips made by females from San Miguel Island
coast, only albatrosses, fulmar, and large gulls, species increased on average by 37% in 1983, and pups received
readily attracted to fishing vessels, increased offshore. In less milk than normal (female pups 3%; male pups 12%).
nearshore waters, however, two species that feed on Lower milk intake resulted in a 23% reduction in growth
anchovies and other fish increased significantly: brown rates of pups and a decrease in body mass for pups 2
pelican in Washington in the 1990s, and rhinoceros auklet months of age and older. 95 “Sneak suckling”, not
since at least 1983.187 Western gulls were able to switch to commonly seen in California sea lions, increased, as did
alternate food sources during the 1982-83 El Niño, and the proportion of yearlings suckling. Females that lost
 CHAPTER 15: COASTAL/MARINE 415

pups or did not breed that season shifted their attention in abundance (H. Allen, pers. comm.) following successful
to raising older offspring. Body mass of female and male efforts to limit the use of harmful pollutants that formerly
pups declined by 25-35%, resulting in increased mortality limited populations. Other species with stable or
at San Nicolas and San Clemente (but not on San Miguel).59 increasing populations include double-crested cormorant
The greater investment in foraging effort, which was and California gull.128, 187
already considerable,48 therefore appeared to be related
to the increase in pup mortality. The individual trips were Conservation Mechanisms
not longer, but dives were deeper than normal as sea lions Continued human population growth and the
tried to compensate for more dispersed or deeper localized concomitant demands on natural resources for public
prey.69 It appears that food habits changed dramatically commodities and recreation are challenges that require
during and shortly after the El Niño.59, 85 dedicated management attention. For example, recent
estimates indicate that the population of Washington could
Population Status double by 2045.188 Despite the successes in recovering
The habitats of the marine-terrestrial interface support a populations of endangered species, many other species
substantial number of threatened, at-risk, vulnerable, or are at risk or will become so in the next several decades.
rare species. Threatened species include the snowy plover The reasons are many (Table 2) and the “solutions” are
and marbled murrelet, the former occurring on sand becoming more complicated and expensive.
beaches and low foredunes and the latter in coastal marine The primary means for addressing these threats to
waters where it forages for fish. The snowy plover was species and habitats are 1) more proactive and innovative
listed as a threatened species because of habitat destruction management (including growth management) that is
and human disturbance in nesting areas.172 At-risk species designed to provide long-term solutions to protect wildlife
include the common murre, western grebe, scaups, and and habitats and allow greater flexibility and options, 2)
scoters, species that have experienced population declines restoration and recovery of degraded habitats and wildlife
in all or part of the region over the past two decades.127, 195 populations, and 3) protection of the remaining important
Some of the species (e.g. western grebe, surf scoter) are habitats. Although a detailed discussion of the first two
known to carry high concentrations of environmental topics is beyond the scope of this chapter, a few relevant
pollutants.86, 87 Vulnerable species are those that meet points are in order.
certain life requisites only in certain habitats or where key The development of conservation initiatives in marine
habitat attributes are present; the destruction or habitats has increased dramatically in recent years.
degradation of those habitats could pose a substantial Recognizing that many species must be managed and
threat to such species. An example is the sanderling, which protected at very large spatial scales, management and
forages and roosts primarily on sand beaches and is conservation plans have been developed for waterfowl42,
171
vulnerable to the harmful effects of oil spills in that habitat. and shorebirds. 62 A conservation plan is being
Another way to define vulnerability is in terms of the developed for colonially nesting birds. Important aspects
proportion of the species’ population that uses an area; of the shorebird plan include public education and
areas that represent a substantial part of a species’ range identification of voluntary conservation measures (e.g.
or support a significant proportion of the population might conservation easements) that could be implemented on
be considered vulnerable. 44, 62 Sanderlings, western private lands. Such actions may be crucial given that many
sandpipers, dunlins, and Leach’s storm-petrel, despite important estuarine sites in Washington and Oregon
being abundant in coastal Washington and/or Oregon, contain areas of privately owned land.62
easily qualify in this category because such large Human activities in the region have greatly influenced
proportions of the species’ overall or regional populations the extent and quality of important wildlife habitats.19, 50
depend on habitats along our coasts.120, 139, 167, 173 Finally, rare With the tremendous growth in the field of wildlife
species include those that are numerically uncommon or ecology in recent decades, particularly in our
rare in the region, although their populations may be understanding of species-habitat relationships, there now
stable. Examples of such species include the gyrfalcon and appear to be opportunities to restore destroyed or
snowy owl. degraded habitats. Although restoration ecology should
Many other species associated with these habitats have not be viewed as a panacea for unwise or unplanned
maintained high levels of abundance or are increasing over growth and development, it does offer hope that degraded
previous population levels. For example, snow geese, habitats can be recovered to the extent that they can
brant, many “puddle” ducks, and loons appear to be function for wildlife. A variety of restoration activities that
common and stable in Washington estuaries (D. Kraege, would benefit shorebirds have been described.27, 62
pers. comm., D. Nysewander, pers. comm.). Although
trend data are lacking, it appears that dunlin and Marine Protected Areas
sanderling populations may be stable over the past few Protection of marine habitats is an important component
decades. 141 Sea otter populations have increased of marine conservation efforts. Certain marine areas in
substantially on the Washington coast following re- Washington and Oregon are now recognized as Marine
establishment efforts there in 1969-1970.96 Bald eagle and Protected Areas (MPAs). The concept of the MPA is
peregrine falcon populations in the region have increased relatively recent, being defined, as “any area of intertidal
 416 Wildlife–Habitat Relationships in Oregon and Washington

or subtidal terrain, together with its overlying water and (UNESCO; Man and the Biosphere Reserve), National
associated flora, fauna, historical and cultural features, Oceanic and Atmospheric Administration (Olympic Coast
which has been reserved by law or other effective means National Marine Sanctuary), U.S. Bureau of Land
to protect part or all of the enclosed environment.”119 The Management (various protected natural areas and
number of MPAs has grown from 118 worldwide in 1970 reserves), Environmental Protection Agency (National
to over 1,300 in the 1990s. 119 Important underlying Estuary Plan sites), and local governments, Native
principles of the MPA program are the restoration of American tribes, and private organizations (e.g., The
depleted stocks of fish through the establishment of no- Nature Conservancy, San Juan Preservation Trust).
harvest areas, and protection of both endangered species Various state laws (e.g. ORS 196 and ORS 197),
and critical habitats. statewide Planning Goals, the Territorial Sea Plan, other
MPAs in Washington and Oregon are managed in a agency rules and statutes, as well as federal laws, apply
variety of administrative contexts. Parks, marine to Oregon ocean resources. Oregon Policy Advisory
sanctuaries, wildlife refuges, wilderness areas, and Council,129 state agencies, local governments, and federal
conservation areas have been established and agencies implement the above policies and laws. One of
administered by state or federal government, local and the elements of the ocean-resource management is the
tribal governments, and private organizations. The MPAs Rocky Shores Strategy program, which addresses
vary in size, management intent, and the degree to which shoreline and offshore rocks and reefs. Oregon rocky
they are protected. Some MPAs are non-contiguous, shores belong to the public, with the Oregon Division of
composed of many elements such as rocks, reefs, or State Lands (ODSL) being the trustee on the public’s
islands. Although there are no standard regulations used behalf. ODSL shares management of the rocky shores with
to manage the body of MPAs, a number of MPAs were Oregon Parks and Recreation Department. Marine life is
established and are managed as a result of cooperation under the jurisdiction of the ODFW, and the “dry” part of
among multiple agencies and organizations.119, 129, 151 the offshore rocks and islands are managed by the U. S.
The need to establish and manage areas to protect Fish and Wildlife Service as National Wildlife Refuges.
marine habitats and species was recognized in Washington Designated sites are managed by numerous state and
only recently. Increasing awareness of degradation of federal agencies or private non-profit organizations. MPAs
marine habitats led to research and monitoring in the are found along much of the Oregon coast (Figure 14b).
shared inland marine waters of Washington and British The primary goals of the MPAs are varied, but they
Columbia and eventually to the signing of an typically emphasize protection of rare or sensitive marine
Environmental Cooperation Agreement between British species and/or habitats. Some of the MPAs also place great
Columbia and Washington State in 1992. The goal of the importance on the scientific, educational, and recreational
agreement was to address habitat degradation and values unique to the site. Species protection ranges from
declines in fish and shellfish populations in Puget Sound, bottom fish, shellfish, and marine invertebrates, to wildlife
Hood Canal, Strait of Juan de Fuca and the Strait of such as harbor seal, California and Steller’s sea lions,
Georgia. The recognition of the importance of MPAs was waterfowl, seabirds, shorebirds, bald eagle, and peregrine
one of the outcomes of the agreement.119 Since the founding falcon. Harvest of certain food species and full access are
of the Washington State Seashore Conservation Areas in allowed in most MPAs, although access may be difficult
1969 and the Washington Marine Protected Areas Work in some areas due to terrain features or the remote nature
Group in 1995, the entire intertidal coast of Washington of the site; access is prohibited (e.g., some Natural Area
has become a single MPA.119, 151 There are 33 additional Preserves) or restricted to particular areas (e.g., Padilla
MPAs on the outer coast and 102 in Puget Sound (Figure Bay National Estuarine Research Reserve, breeding sites
14a).61, 119, 151 on refuges) at very few MPAs. Further details regarding
MPAs in Washington and Oregon are owned and MPAs in Washington and Oregon are available
managed by a diverse group of organizations. These elsewhere.61, 119, 129, 151
include Washington Department of Natural Resources
(Natural Area Preserves and Natural Resources Research Needs
Conservation Areas), Washington Department of Fish and There is a great need for additional research regarding the
Wildlife (Wildlife Areas), Washington State Parks and species and habitats of the marine environment. Some of
Recreation Commission (State Parks), Washington the research topics are species or habitat-specific, whereas
Department of Ecology (Padilla Bay National Estuarine many other topics apply to multiple species and habitats.
Research Reserve [this site is cooperatively managed with General research topics that apply to multiple species and
other agencies]), University of Washington (Marine habitats include: 1) response of marine bird and mammal
Biological Preserve in waters of San Juan County and populations to the effects of oil and other pollutants
waters surrounding Cypress Island in Skagit County), (essentially all habitats), various fisheries activities
Oregon Department of Fish and Wildlife (Marine Gardens, (estuaries, inland marine, nearshore, and shelf), and El
Research Reserves), Oregon Parks and Recreation Niño (essentially all habitats), 2) population demography
Department (State Parks, Habitat Refuges), U.S. Fish and and monitoring of status or trends (relates to species or
Wildlife Service (National Wildlife Refuges), United communities in all habitats), 3) development of methods
Nations Educational, Scientific and Cultural Organization to reduce impacts of fisheries activities on non-target
CHAPTER 15: COASTAL/MARINE 417

Figure 14. Distribution of selected Marine Protected Areas in

Washington and Oregon.

National Estuary Program

National Estuarine Research Preserve
National Wildlife Refuge
State Marine Protected Areas
National Marine Sanctuary

species (estuaries, inland marine), and 4) responses to

human disturbances (beaches, estuaries, headlands).
Examples of species- or habitat-specific research needs
include: behavioral, physiological, or other responses to
restored habitats (e.g. snowy plover on beaches after
beachgrass removal, shorebirds at restored roost sites in
an estuary such as Tillamook Bay, shorebird or waterfowl
foraging behavior in cordgrass removal areas); more
general studies of habitat use (foraging habitat preferences
of waterbirds in estuaries as a function of tidal phase);
studies that enable inference to be made about habitat
quality (e.g. studies of food habits and roost site selection
by waterfowl, waterbirds, shorebirds, colonially-nesting
gulls in estuaries); and the effects of sedimentation on
estuarine productivity.

Acknowledgments
We thank Janet Hodder and Dave Nysewander for making
comments that improved the manuscript; John
Calambokidis, Al Didier, Jr., Roy Lowe, and Ken Warheit
for providing important information and references; and
Jeff Gilligan and Dave Lauten for sharing their expertise.

Literature Cited
1. Akins, G. J., and C. A. Jefferson. 1973. Coastal wetlands of Oregon.
Oregon Coastal Conservation and Development Commission. Salem,
OR.
2. Ainley, D. G., R. J. Boekelheide, S. H. Morrell, and C. S. Strong. 1990.
Pigeon Guillemot. Pages 277-305 in: D. G. Ainley, and R. J. Boekelheide,
editors. Seabirds of the Farallon Islands: Ecology, dynamics, and
structure of an upwelling-system community. Stanford University Press,
CA.
3. ———, W. J. Sydeman, S. A. Hatch, and U. W. Wilson. 1994. Seabird
population trends along the west coast of North America: cause and
the extent of regional concordance. Studies in Avian Biology 15:
119-133.
4. ———, L. B., Spear, and S. G. Allen. 1996. Variation in the diet of the
Cassin’s auklet reveals spatial, temporal, seasonal, and decadal
occurrence patterns of euphasids off California, USA. Marine Ecology
Progress Series 137: 1-10.
5. ———, L. B. Spear, S. G. Allen and C. A. Ribic. 1996. Temporal and
spatial patterns in the diet of the common murre in California waters.
Condor 98: 691-705.
 418 Wildlife–Habitat Relationships in Oregon and Washington

6. Ashmole, N. P. 1971. Seabird ecology and the marine environment. 24. Brown, R. F. 1988. Assessment of pinniped populations in Oregon,
Pages 112-286 in: D. S. Farner and J. R. King, editors. Avian biology,Vol. 1. April 1984 to April 1985. NMFS-NWAFC Processed Rept. 88-05, 44 p.
Academic Press, New York, NY. Alaska Fisheries Science Center, NMFS, NOAA, 7600 Sand Point Way
7. Baird, R. W., and L. M. Dill. 1995. Occurrence and behavior of transient NE, Seattle, WA.
killer whales: seasonal and pod-specific variability, foraging behavior, and 25. Brueggeman, J. J. ed. 1992. Oregon and Washington Marine Mammal
prey handling. Canadian Journal of Zoology 73: 1300-1311. and Seabird Surveys—Final Report. OCS Study MMS 91-0093. Ebasco
8. ———, and ———. 1996. Ecological and social determinants of group Environmental, Bellevue, WA and Ecological Consulting, Inc., Portland,
size in transient killer whales. Behavioral Ecology 7: 408-416. OR.
9. Baldridge, A., and J. B. Crowell, Jr. 1966. The fall migration: northern 26. Reference deleted in proof.
Pacific coast region. Audubon Field Notes 20: 82. 27. Buchanan, J. B. In press. Shorebirds: plovers, oystercatchers, avocets
10. Barber, R. T., and F. P. Chavez. 1983. Biological consequences of El Niño. and stilts, sandpipers, snipes, and phalaropes. In E. M. Larsen, and N.
Science 222: 1204-1210. Nordstrom, editors. Management recommendations for priority bird
11. Bayer, R. D. 1986. Breeding success of seabirds along the mid-Oregon species. Washington Department of Fish and Wildlife, Olympia, WA.
coast concurrent with the 1983 El Niño. Murrelet 67: 23-26. 28. ———. 1999. Recent changes in the winter distribution and
12. Beach, R. J., A. C. Geiger, S. J. Jeffries, S. D. Treacy, and B. L. Troutman. abundance of rock sandpipers in North America. Western Birds 30:
1985. Marine mammals and their interactions with fisheries of the 193-199.
Columbia River and adjacent waters, 1980-1982. NMFS-AFSC 29. ———. 1992. Winter abundance of shorebirds at coastal beaches of
Processed Rept. 85-04, 316 pp. Alaska Fisheries Science Center, NMFS, Washington. Washington Birds 2: 12-19.
NOAA, 7600 Sand Point Way NE, Seattle, WA. 30. ———. 1988. Migration and winter populations of greater yellowlegs,
13. Bigg, M. A., P. F. Olesiuk, G. M. Ellis, J. K. B. Ford, and K. C. Balcomb. Tringa melanoleuca, in western Washington. Canadian Field-Naturalist
1990. Social organization and genealogy of resident killer whales 102: 611-616.
(Orcinus orca) in the coastal waters of British Columbia and Washington 31. ———, and J. R. Evenson. 1997. Abundance of shorebirds at Willapa
State. Reports to the International Whaling Commission, Special Issue Bay, Washington. Western Birds 28: 158-168.
12: 383-405. 32. Burger, A. E., and D. M. Fry. 1993. Effects of oil pollution on seabirds in
14. Boekelheide, R. J., D. G. Ainley, S. H. Morrell, H. R. Huber, and T. J. Lewis. the northeast Pacific. Pages 254-263 in: K.Vermeer, K. T. Briggs, K. H.
1990. Common murre. Pages 245-275 in: D. G. Ainley, and R. J. Morgan, and D. Siegel-Causey, editors. The status, ecology and
Boekelheide, editors. Seabirds of the Farallon Islands: Ecology, dynamics, conservation of marine birds of the North Pacific. Canadian Wildlife
and structure of an upwelling-system community. Stanford University Service Special Publication, Ottawa, Canada.
Press, CA. 33. Byrd, G.V., E. C. Murphy, G. W. Kaiser, A. Y. Kondratyev, and Y.V. Shibaev.
15. Boersma, P. D., and M. J. Green. 1993. Conservation of storm-petrels 1993. Status and ecology of offshore fish-feeding alcids (murres and
in the North Pacific. Pages 112-121 in: K.Vermeer, K. T. Briggs, K. H. puffins) in the North Pacific. Pages 176-198 in: K.Vermeer, K. T. Briggs, K.
Morgan, and D. Siegel-Causey, editors. The status, ecology, and H. Morgan, and D. Siegel-Causey, editors. The status, ecology and
conservation of marine birds of the North Pacific. Canadian Wildlife conservation of marine birds of the North Pacific. Canadian Wildlife
Service Special Publication, Ottawa. Service Special Publication, Ottawa, Canada.
16. ———, and N. T. Wheelwright. 1979. Egg neglect in the 34. Calambokidis, J., J. B. Buchanan, G. H. Steiger, and J. R. Evenson. 1991.
procellariiformes: reproductive adaptations in the fork-tailed storm- Toxic contaminants in Puget Sound wildlife: Literature review and
petrel. Condor 81: 157-165. recommendations for research and monitoring. Contract report 68-
17. Boness, D. J., O. T. Oftedal, and K. A. Ono. 1991. The effect of El Niño D8-0085 to U.S. Environmental Protection Agency, Seattle, WA.
on pup development in the California sea lion (Zalophus californianus). 35. ———, T. Chandler, K. Rasmussen, G. H. Steiger, and L. Schlender.
Early postnatal growth. Pages 173-179 in: F.Trillmich, and K.A. Ono, 1999. Humpback and blue whale photographic identification research
editors. Pinnipeds and El Niño: Responses to environmental stress. off California, Oregon, and Washington in 1998. Final report to
Springer-Verlag, Berlin, Germany. Southwest Fisheries Science Center, Olympic Coast National Marine
18. Bonnell, M. L., C. E. Bowlby, and G. A. Green. 1992. Pinniped Sanctuaries, and University of California at Santa Cruz. Cascadia
distribution and abundance off Oregon and Washington, 1989-1990. Research Collective, Olympia, WA.
Chapter 2 in: J. J. Brueggeman, editor. Oregon and Washington marine 36. ——— J. C. Cubbage, J. R. Evenson, S. D. Osmek, J. L. Laake, P. J. Gearin,
mammal and seabird surveys. OCS Study MMS 91-0093. Ebasco B. J. Turnock, S. J. Jeffries, and R. F. Brown. 1993. Abundance estimates of
Environmental, Bellevue, WA and Ecological Consulting, Inc., Portland, harbor porpoise in Washington and Oregon waters. Final Report to
OR. National Marine Mammal Laboratory, AFSC, NMFS, Seattle, WA.
19. Boule, M., N. Olmsted, and T. Miller. 1983. Inventory of wetland Cascadia Research Collective, Olympia, WA.
resources and an evaluation of wetland management in western 37. ———, J. R. Evenson, G. H. Steiger, and S. J. Jeffries. 1994. Gray whales
Washington. Washington Department of Ecology, Olympia, WA. of Washington State: Natural history and photographic catalog.
20. ———, and K. F. Bierly. 1987. History of estuarine wetland Cascadia Research Collective, Olympia, WA.
development and alteration: What have we wrought? Northwest 38. ———, J. Peard, G. H. Steiger, J. C. Cubbage, and R. L. DeLong. 1984.
Environmental Journal 3: 43-61. Chemical contaminants in marine mammals from Washington State.
21. Bowlby, C. E., G. A. Green, and M. L. Bonnell. 1994. Observations of NOAA Technical Memorandum NOS OMS 6, Rockville, MD.
leatherback turtles offshore of Washington and Oregon. Northwestern 39. ———, G. H. Steiger, J. R. Evenson, K. R. Flynn, K. C. Balcomb, D. E.
Naturalist 75: 33-35. Claridge, P. Bloedel, J. M. Straley, C. S. Baker, O. von Ziegesar, M. E.
22. Briggs, K. T., and E. W. Chu. 1987. Trophic relationships and food Dahlheim, J. M. Waite, J. D. Darling, G. Ellis, and G. A. Green. 1996.
requirements of California seabirds: updating models of trophic impact. Interchange and isolation of humpback whales off California and other
Pages 279-304 in: J. P. Croxall, editor. Seabird feeding ecology and role in North Pacific feeding grounds. Marine Mammal Science 12: 215-226.
marine ecosystems. Cambridge University Press, Cambridge, England. 40. ———, ——— K. Rasmussen, J. Urbán R., K. C. Balcomb, P. Ladrón de
23. ———, D. H. Varoujean, W. W. Williams, R. G. Ford, M. L. Bonnell, and J. Guevara P., M. Salinas Z., J. K. Jacobsen, C. S. Baker, L. M. Herman, S.
L. Casey. 1992. Seabirds of the Oregon and Washington OCS, 1989- Cerchio, and J. D. Darling. 2000. Migratory destinations of humpback
1990. Chapter 3 in: J. J. Brueggeman, editor. Oregon and Washington whales that feed off California, Oregon and Washington. Marine
marine mammal and seabird surveys. OCS Study MMS 91-0093. Ebasco Ecology Progress Series 192: 295-304
Environmental, Bellevue, WA and Ecological Consulting, Inc., Portland, 41. ———, ———, J. M. Straley, L. M. Herman, S. Cerchio, D. R. Salden, R.
OR. J. Urbán, J. K. Jacobsen, O. von Ziegesar, K. C. Balcomb, C. M. Gabriele,
M. E. Dahlheim, S. Uchida, G. Ellis, Y. Miyamura, P. Ladron de Guevara,
 CHAPTER 15: COASTAL/MARINE 419

M.Yamaguchi, F. Sato, S. A. Mizroch, L. Schlender, K. Rasmussen, and J. 60. ———, B.S. Stewart, and R.D. Hill. 1992. Documenting migrations of
Barlow. In press. Movements and population structure if humpback northern elephant seals using day length. Marine Mammal Science
whales in the North Pacific Basin. Marine Mammal Science. 8:155-159.
42. Canadian Wildlife Service and U.S. Fish and Wildlife Service. 1998. 61. Didier, A. J., Jr. 1998. Marine Protected Areas of Washington and
North American waterfowl management plan (1998 update). Oregon. Pacific Fishery Management Council. Contract No. 98-08.
43. Carter, H. R., U. W. Wilson, R. W. Lowe, M. S. Rodway, D. A. Manuwal, J. 62. Drut, M. S. and J. B. Buchanan. 2000. U.S. National shorebird
E. Takekawa, and J. L. Yee. In press. Population trends of the common conservation plan: Northern Pacific coast working group regional
murre (Uria aalge californica). In: D.A. Manuwal, H. R. Carter, and T. management plan.
Zimmerman, editors. Biology and conservation of the common murre 63. Duffy, D. C. 1990. Seabirds and the 1982-1983 El Niño-Southern
in California, Oregon, Washington, and British Columbia. U.S. Geological Oscillation. Pages 395-415 in: P. W. Glynn, editor. Global ecological
Survey, Information and Technology Report, Washington, D.C. consequences of the 1982-83 El Niño Southern Oscillation. Elsevier
44. Carter, M. F., W. C. Hunter, D. N. Pashley, and K.V. Rosenberg. 2000. Oceanography Series, 52. University of Miami, FL.
Setting conservation priorities for landbirds in the United States: The 64. Ebbesmeyer, C. C., D. R. Cayan, D. R. McLain, F. H. Nichols, D. H.
Partners in Flight approach. Auk 117:541-548. Peterson, and K. T. Redmond. 1991. 1976 step in the Pacific climate:
45. Condit, R., and B. J. Le Boeuf. 1984. Feeding habits and feeding grounds forty environmental changes between 1968-1975 and 1977-1984. Pages
of the northern elephant seal. Journal of Mammalogy 65: 281-290. 129-141 in: J. L. Betancourt, and V. L. Tharp, editors. Proceedings of the
46. Cooke, W. W. 1910. Distribution and migration of North American Seventieth Annual Pacific Climate (PACLIM) Workshop. California
shorebirds. United States Biological Survey Bulletin No. 35. Department of Water Resources. Interagency Ecological Studies
47. Cooper, W. S. 1958. Coastal sand dunes of Oregon and Washington. Program Technical Report 26.
Geological Society of America. Memoir 72. 65. Evenson, J. R., and J. B. Buchanan. 1997. Seasonal abundance of
48. Costa, D. P., G. A. Antonelis, and R. L. DeLong. 1991. Effects of El Niño shorebirds at Puget Sound estuaries. Washington Birds 6:34-62.
on the foraging energetics of the California sea lion. Pages 156-165 in: F. 66. ———, D. R. Nysewander, and M. Mahaffy. 1999. Progress report of
Trillmich, and K.A. Ono, editors. Pinnipeds and El Niño. Responses to the 1999 pigeon guillemot breeding colony census of Washington
environmental stress. Springer-Verlag, Berlin, Germany. inland marine waters. Washington Department of Fish and Wildlife.
49. Custer, T. W., and J. P. Myers. 1990. Organochlorines, mercury, and 67. Ewins, P. J., H. R. Carter, and Y.V. Shibaev. 1993. The status, distribution,
selenium in wintering shorebirds from Washington and California. and ecology of inshore fish-feeding alcids (Cepphus guillemots and
California Fish and Game 76: 118-125. Brachyramphus murrelets in the North Pacific. Pages 164-175 in: K.
50. Dahl, T. E. 1990. Wetlands of the United States 1780s to 1980s. U.S. Vermeer, K. T. Briggs, K. H. Morgan, and D. Siegel-Causey, editors. The
Fish and Wildlife Service, Washington, D.C. status, ecology and conservation of marine birds of the North Pacific.
51. Darling, J. D. 1984. Gray whales off Vancouver Island, British Columbia. Canadian Wildlife Service Special Publication, Ottawa, Canada.
Pages 267-287 in: M. L. Jones, S. L. Swartz, and S. Leatherwood, editors. 68. Favorite, F., A. J. Dodimead, and K. Nasu. 1976. Oceanography of the
The Gray Whale, Eschrictius robustus.Academic Press, Inc., Orlando, FL. subarctic Pacific region 1960-71. International Northern Pacific
52. ———, J. Calambokidis, K. C. Balcomb, P. Bloedel, K. R. Flynn, A. Fisheries Commission Bulletin No. 33.
Mochizuki, K. Mori, F. Sato, H. Suganuma, and M.Yamaguchi. 1996. 69. Feldkamp, S. D., R. L. DeLong, and G. A. Antonelis. 1991. Effects of El
Movement of a humpback whale (Megaptera novawangliae) from Japan Niño 1983 on the foraging patterns of California sea lions (Zalophus
to British Columbia and return. Marine Mammal Science 12: 281-287. californianus) near San Miguel Island, California. Pages 146-155 in: F.
53. ———, K.E. Keogh, and T. E. Steeves. 1998. Gray whale (Eschrichtius Trillmich, and K. A. Ono, editors. Pinnipeds and El Niño. Responses to
robustus) habitat utilization and prey species off Vancouver Island, environmental stress. Springer-Verlag, Berlin, Germany.
British Columbia. Marine Mammal Science 14: 692-720. 70. Fiscus, C., and K. Niggol. 1965. Observations of cetaceans off
54. Day, J. W., Jr., C. A. S. Hall, W. M. Kemp, and A. Yáñez-Arancibia. 1989. California, Oregon, and Washington. U.S. Fish and Wildlife Service,
Estuarine ecology. John Wiley and Sons, New York, NY. Special Science Report-Fisheries No. 498.
55. Day, R. D., H. S. Wehle, and F. C. Coleman. 1985. Ingestion of plastic 71. Forbush, E. H. 1912. A history of the game birds, wildfowl, and shore
pollutants by marine birds. Pages 344-386 in: R. S. Shomura, and H. O. birds of Massachusetts and adjacent states. Massachusetts Board of
Yoshida, editors. Proceedings of the workshop on the fate and impact Agriculture.
of marine debris. U.S. Department Committee, NOAA Technical Memo, 72. Ford, J. K. B., G. M. Ellis, L. G. Barrett-Lennard, A. B. Morton, R. S. Palm,
NMFS, NOAA-TM-NMFS-SWFC-54. and K. C. Balcomb. 1999. Dietary specialization of two sympatric
56. Dayton, P. K., and M. J. Tegner. 1990. Bottoms beneath troubled waters: populations of killer whales (Orcinus orca) in coastal British Columbia
benthic impacts of the 1982-1984 El Niño in the temperate zone. Pages and adjacent waters. Canadian Journal of Zoology 76: 1456-1471.
433-472 in: P. W. Glynn, editor. Global ecological consequences of the 73. Ford, R. G., D. H.Varoujean, D. R. Warrick, W. A. Williams, D. B. Lewis,
1982-1983 El Niño-Southern Oscillation. Elsevier Oceanography Series, C. L. Hewitt, and J. L. Casey. 1991. Final report: seabird mortality
52. University of Miami, Miami, FL. resulting from the Nestucca oil spill incident, winter 1988-89. Report
57. DeGange, A. R., R. H. Day, J. E. Takekawa, and V. M. Mendenhall. 1993. for Washington Dept. of Wildlife.
Losses of seabirds to gill nets in the North Pacific. Pages 204-211 in: K. 74. Francis, J. M., and C. B. Heath. 1991. The effects of El Niño on the
Vermeer, K. T. Briggs, K. H. Morgan, and D. Siegel-Causey, editors. The frequency and sex ratio of suckling yearlings in the California sea lion.
status, ecology, and conservation of marine birds of the North Pacific. Pages 193-210 in: F. Trillmich, and K. A. Ono, editors. Pinnipeds and El
Canadian Wildlife Service Special Publication, Ottawa, Canada. Niño. Responses to environmental stress. Springer-Verlag, Berlin,
58. DeLong, R. L., and G. A. Antonelis. 1991. Impact of the 1982-1983 El Germany.
Niño on the northern fur seal population at San Miguel Island, 75. Frankignoul, C. 1994. Equatorial ocean dynamics: observations and
California. Pages 75-83 in: F. Trillmich, and K. A. Ono, editors. Pinnipeds modeling. Pages 157-236 in: P. Malanotte Rizzoli, editor. The central
and El Niño. Responses to environmental stress. Springer-Verlag, Berlin, circulation of the Oceans. Instituto Veneto Di Scienze, Lettere Ed Arti,
Germany. Venice, Italy.
59. ———, ———, C. W. Oliver, B. S. Stewart, M. C. Lowry, and P. K. 76. Franklin, J. F., and C.T. Dyrness. 1973. Natural vegetation of Oregon
Yochem. 1991. Effects of the 1982-83 El Niño on several population and Washington. U.S. Forest Service Gen. Tech. Rep. PNW-8. Reprinted
parameters and diet of California sea lions on the California Channel 1988, Oregon State University Press, Corvallis, OR.
Islands. Pages 166-172 in: F. Trillmich, and K. A. Ono, editors. Pinnipeds 77. Gaston, A. J. 1994. Unusual number of Laysan albatrosses, Diomedea
and El Niño. Responses to environmental stress. Springer-Verlag, Berlin, immutabilis, off the west coast of Haida Gwaii, Queen Charlotte Islands,
Germany. British Columbia. Canadian Field-Naturalist. 108: 373.
 420 Wildlife–Habitat Relationships in Oregon and Washington

78. ———, and S. B. C. Dechesne. 1996. Rhinoceros auklet (Cerorhinca Milk intake. Pages 180-184 in: F. Trillmich, and K. A. Ono, editors.
monocerata). Pages 1-20 in: A. Poole, and F. Gill, editors.The birds of Pinnipeds and El Niño. Responses to environmental stress.
North America. The Academy of Natural Sciences, Philadelphia, PA, and Springer-Verlag, Berlin, Germany.
The American Ornithologists’ Union, Washington, D.C. 96. Jameson, R. J., K. W. Kenyon, S. Jeffries, and G. A.Vanblaricom. 1986.
79. Gonor, J. J., J. R. Sedell, and P. A. Benner. 1988. What we know about Status of a translocated sea otter population and its habitat in
large trees in estuaries, in the sea, and on coastal beaches. Pages 83-112 Washington. Murrelet 67: 84-87.
in: C. Maser, R. F. Tarrant, J. M. Trappe, and J. F. Franklin, editors. From the 97. Jeffries, S. J., R. F. Brown, H. R. Huber, and R. L. DeLong. 1997.
forest to the sea: A story of fallen trees. USDA General Technical Assessment of harbor seals in Washington and Oregon, 1996. Pages 83-
Report PNW-GTR-229. Portland, OR. 94 in: P. S. Hill, and D. P. DeMaster, editors. MMPA and ESA
80. Graybill, R., and J. Hodder. 1985. Effects of the 1982-83 El Niño on Implementation Program, 1996. AFSC Processed Rept. 97-10. Alaska
reproduction of six species of seabirds in Oregon. Pages 205-210 in: W. Fisheries Science Center, NMFS, NOAA, 7600 Sand Point Way NE,
S. Wooster, and D. L. Fluharty, editors. El Niño North: El Niño effects in Seattle, WA.
the eastern subarctic Pacific Ocean. Washington Sea Grant Program, 98. ———, P. J. Gearin, H. R. Huber, D. L. Saul, and D. A. Pruett. 2000. Atlas
University of Washington, Seattle, WA. of seal and sea lion haulout sites in Washington. Washington
81. Green, G. A., J. J. Brueggeman, R. A. Grotefendt, C. E. Bowlby, M. L. Department of Fish and Wildlife, Olympia, WA.
Bonnell, and K. T. Balcomb, III. 1992. Cetacean distribution and 99. Kelly, J. P., J. G. Evens, R. W. Stallcup, and D. Wimpfheimer. 1996. Effects
abundance off Oregon and Washington, 1989-1990. Chapter 1 in: J. J. of aquaculture on habitat use by wintering shorebirds in Tomales Bay,
Brueggeman, editor. Oregon and Washington Marine Mammal and California. California Fish and Game 82: 160-174.
Seabird Surveys. OCS Study MMS 91-0093. Ebasco Environmental, 100. Kennish, M. J. 1992. Ecology of estuaries: Anthropogenic effects. CRC
Bellevue, WA and Ecological Consulting, Inc., Portland, OR. Press, Boca Raton, FL.
82. ———, R. A. Grotefendt, M. A. Smultea, C. E. Bowlby, and R. A. 101. Kerr, R. A. 1998. Warming’s unpleasant surprise: Shivering in the
Rowlett. 1993. Delphinid aerial surveys in Oregon and Washington greenhouse? Science. 281: 156-158.
offshore waters. Final Report to National Marine Mammal Laboratory, 102. ———. 1999. Big El Niños ride the back of slower climate change.
National Marine Fisheries Service, Seattle, WA. Ebasco Environmental, Science 283: 1108-1109.
Bellevue, WA. 103. Komar, P. D. 1997. The Pacific Northwest Coast: Living with the
83. Grinnell, J., H. C. Bryant, and T. I. Storer. 1918. The game birds of shores of Oregon and Washington. Duke University Press, Durham,
California. University of California Press, Berkeley, CA. NC.
84. Harrington, B., and E. Perry. 1995. Important shorebird staging sites 104. Kvitek, R. G., D. Shull, D. Canestro, E. C. Bowlby, and B. L. Troutman.
meeting Western Hemisphere Shorebird Reserve Network criteria in 1989. Sea otters and benthic prey communities in Washington State.
the United States. USDI Fish and Wildlife Service. Marine Mammal Science 5: 266-280.
85. Heath, C. B., K. A. Ono, D. J. Bonese, and J. M. Francis. 1991. The 105. Laake, J. L., R. L. DeLong, J. Calambokidis, and S. Osmek. 1997.
influence of El Niño on female attendance patterns in the California Abundance and distribution of marine mammals in Washington and
sea lion. Pages 138-145 in: F.Trillmich, and K. A. Ono, editors. Pinnipeds British Columbia inside waters, 1996. Pages 67-73 in: P. S. Hill, and D. P.
and El Niño. Responses to environmental stress. Springer-Verlag, Berlin, DeMaster, editors. MMPA and ESA Implementation Program, 1996.
Germany. AFSC Processed Report 97-10. Alaska Fisheries Science Center, NMFS,
86. Henny, C. J., L. J. Blus, and R. A. Grove. 1990. Western grebe, NOAA, 7600 Sand Point Way NE, Seattle, WA.
Aechmophorus occidentalis, wintering biology and contaminant 106. ——— J. Calambokidis, and S. Osmek. 1998. Survey report for the
accumulation in Commencement Bay, Puget Sound, Washington. 1997 aerial surveys for harbor porpoise and other marine mammals of
Canadian Field-Naturalist 104: 460-472. Oregon, Washington and British Columbia outside waters. Pages 77-97
87. ———, ———, ———, and S. P. Thompson. 1991. Accumulation of in: P. S. Hill, and D. P. DeMaster, editors. MMPA and ESA Implementation
trace elements and organochlorines by surf scoters wintering in the Program, 1997. AFSC Processed Report 98-10. Alaska Fisheries Science
Pacific Northwest. Northwestern Naturalist 72: 43-60. Center, NMFS, NOAA, 7600 Sand Point Way NE, Seattle, WA.
88. Herman, S. G., and J. B. Bulger. 1981. Distribution and abundance of 107. Lamont, M. M., J. T. Vida, J. T. Harvey, S. Jeffries, R. Brown, H. R. Huber,
shorebirds during the 1981 spring migration at Grays Harbor, R. DeLong, and W. K. Thomas. 1996. Genetic substructure of the Pacific
Washington. U.S. Army Corps of Engineers, Contract Report harbor seal (Phoca vitulina richardsi) off Washington, Oregon, and
DACW67-81-0936. Seattle, WA. California. Marine Mammal Science 12: 402-413.
89. Herzing, D. L., and B. R. Mate. 1984. Gray whale migrations along the 108. Landry, M. L. and B. M. Hickey, editors. 1989. Coastal oceanography
Oregon coast, 1978-1981. Pages. 289-307 in M. L. Jones, S. L. Swartz, and of Washington and Oregon. Elsevier Press, Amsterdam, Netherlands.
S. Leatherwood, editors.The Gray Whale, Eschrictius robustus.Academic 109. Le Boeuf, B. J., and J. Reiter. 1991. Biological effects associated with El
Press, Inc., Orlando, FL. Niño-Southern Oscillation, 1982-83, on northern elephant seals
90. Hodder, J., R. F. Brown, and C. Cziesla. 1998. The northern elephant breeding at Año Nuevo, California. Pages 206-218 in: F. Trillmich, and K.
seal in Oregon: A pupping range extension and onshore occurrence. Ono, editors. Pinnipeds and El Niño. Responses to environmental
Marine Mammal Science 14: 873-880. stress. Springer-Verlag, Berlin, Germany.
91. ———, and M. R. Graybill. 1985. Reproduction and survival of 110. Lowe, R. 1993. El Niño hard on Oregon seabirds. Pacific Seabird
seabirds in Oregon during the 1982-83 El Niño. Condor 535-541. Group Bulletin 20: 62.
92. Holden, C. 1999. Pumping up the greenhouse. Science 285: 2057. 111. ———. 1998. Regional reports, Washington and Oregon. Pacific
93. Huber, H. R., C. Beckham, and J. Nisbet. 1991. Effects of the 1982-83 El Seabirds 25: 84.
Niño on northern elephant seals on the South Farallon Islands, 112. ———, and D. S. Pitkin. 1996. Replicate aerial photographic censuses
California. Pages 219-233 in: F. Trillmich, and K. A. Ono, editors. of Oregon Common Murre colonies 1995. USFWS, Newport, OR.
Pinnipeds and El Niño. Responses to environmental stress. 113. McPhaden, M. J. 1999. Genesis and evolution of the 1997-98 El Niño.
Springer-Verlag, Berlin, Germany. Science 283: 950-954.
94. Huntington, C. E., R. G. Butler, and R. A. Mauck. 1996. Leach’s storm- 114. Mate, B. R. 1975. Annual migrations of the sea lions Eumetopias
petrel (Oceanodroma leucorhoa). Pages 1-32 in:A. Poole, and F. Gill, jubatus and Zalophus californianus along the Oregon coast. Rapports et
editors. The birds of North America. The Academy of Natural Sciences, Proces-Verbaux des Reunions. Conseil International pour l’Exploration
Philadelphia, PA, and The American Ornithologists’ Union, Washington, Scientifique de la Mer Meditarranee (Monaco) 169: 455-461.
D.C. 115. ———. 1981. Marine mammals. Pages 372-492 in: C. Maser, B. R.
95. Iverson, S. J., O. T. Oftedal, and D. J. Boness. 1991. The effect of El Niño Mate, J. F. Franklin, and C. T. Dyrness, editors. Natural history of Oregon
on pup development in the California sea lion (Zalophus californianus). coast mammals. USDA Forest Service, General Technical Report PNW-
133.
 CHAPTER 15: COASTAL/MARINE 421

116. Melillo, J. M. 1999. Warm, warm on the range. Science 283: 183-4. 137. ———. 1994d. Shorebird survey of the Lower Columbia Estuary:
117. Melvin, E. F., J. K. Parrish, and L. L. Conquest. 1999. Novel tools to August 1994. Unpublished report to U.S. fish and Wildlife Service,
reduce seabird bycatch in coastal gillnet fisheries. Conservation Biology Portland, OR.
13: 1386-1397. 138. Osbourne, R., J. Calambokidis, and E. M. Dorsey. 1988. A guide to
118. Mitsch, W. J., and J. G. Gosselink. 1993. Wetlands (second edition). Van marine mammals of Greater Puget Sound. Island Publishers, Anacortes,
Nostrand Reinhold, New York, NY. WA.
119. Murray, M. R. 1998. The status of Marine Protected Areas in Puget 139. Page, G. W., L. E. Stenzel, and J. E. Kjelmyr. 1999. Overview of
Sound. Puget Sound/Georgia Basin Environmental Report Series No. 8. shorebird abundance and distribution in wetlands of the Pacific coast
Olympia, WA. of the contiguous United States. Condor 101: 461-471.
120. Myers, J. P., C. T. Schick, and C. J. Hohenberger. 1984. Notes on the 140. Parrish, J. K. 1998. Attendance and reproductive success of Tatoosh
1983 distribution of sanderlings along the United States’ Pacific coast. Island common murres: Draft final report 1997. University of
Wader Study Group Bulletin 40: 22-26. Washington, Seattle,WA.
121. National Marine Fisheries Service (NMFS). 1992. Final recovery plan 141. Paulson, D. 1993. Shorebirds of the Pacific Northwest. University of
for Steller sea lions (Eumetopias jubatus). Prepared by the Steller Sea Washington Press, Seattle, WA.
Lion Recovery Team for the National Marine Fisheries Service, Office 142. Penland, S. 1981. Natural history of the Caspian tern in Grays
of Protected Resources, Silver Springs, MD. Harbor, Washington. Murrelet 62: 66-72.
122. ———. 1995. Environmental assessment on protecting winter-run 143. Philander, S. G. H. 1990. El Niño, La Niña and the Southern
wild steelhead from predation by California sea lions in the Lake Oscillation. Academic Press, San Diego, CA.
Washington ship canal. NMFS Environmental Assessment Report. 144. Pitkin, D. S., and R. W. Lowe. 2000. Replicate aerial photographic
Alaska Fisheries Science Center, NMFS, NOAA, 7600 Sand Point Way censuses of Oregon common murre colonies, 1996-1997. Unpublished
NE, Seattle, WA. report to the Tenyo Maru Trustee Council. U.S. Fish and Wildlife
123. ———. 1997. Investigation of scientific information on the impacts Service, Newport, OR.
of California sea lions and Pacific harbor seals on salmonids and on the 145. Prince, P. A., and R. A. Morgan. 1987. Diet and feeding ecology of
coastal ecosystems of Washington, Oregon, and California. NOAA Tech. Procellariiformes. Pages 135-171 in: J. P. Croxall, editor. Seabirds: feeding
Memo. NMFS-NWAFSC-28. Northwest Region, NMFS, NOAA, 7600 ecology and role in marine ecosystems. Cambridge University Press,
Sand Point Way NE, Seattle, WA. Cambridge, England.
124. Nehls, H. B. 1994. Oregon shorebirds: Their status and movements. 146. Puget Sound Water Quality Action Team. 1999. 2000 Puget Sound
Oregon Department of Fish and Wildlife, Technical Report 94-1-02. update: seventh report of the Puget Sound Ambient Monitoring
125. Nichols, F. H., J. K. Thompson, and L. E. Schemel. 1990. Remarkable Program. Second draft.
invasion of San Francisco Bay (California, USA) by the Asian clam 147. Quinn, W. H., and V. T. Neal. 1987. El Niño occurrences over the past
Potamo-corbula amurensis. II. Displacement of a former community. four and a half centuries. Journal of Geophysical Research 92 (C13):
Marine Ecology Progress Series 66: 95-101. 14449-14461.
126. Northridge, S. P. 1991. Driftnet fisheries and their impacts on 148. Reimer, S. D., and R. F. Brown. 1997. Prey of pinnipeds at selected
nontarget species: A worldwide review. Fisheries Technical Paper 320. sites in Oregon identified by scat (fecal) analysis, 1983-1996. Oregon
Food and Agriculture Organization of the United Nations, Rome, Italy. Dept. of Fish and Wildlife, Wildlife Diversity Program, Tech. Rept. 97-6-
127. Nysewander, D. R., and J. R. Evenson. 1998. Status and trends for 02.
selected diving duck species examined by the marine bird component, 149. Ribic, C. A., S. W. Johnson, and C. A. Cole. 1997. Distribution, type,
Puget Sound Ambient Monitoring Program (PSAMP), Washington accumulation, and source of marine debris in the United States, 1989-
Department of Fish and Wildlife. Pages 847-867 in: R. Strickland, editor. 1993. Pages 35-47 in: J. M. Coe, and D. B. Rogers, editors. Marine debris:
Puget Sound Research ’98. Puget Sound Water Quality Action Team, sources, impacts, and solutions. Springer-Verlag, New York, NY.
Olympia, WA. 150. Richmond, R. H. 1989. The effects of the El Niño/Southern
128. ———, and ———. 2000. Puget Sound Ambient Monitoring Oscillation on the dispersal of corals and other marine organisms.
Program: Report of the marine bird, waterfowl, and marine mammal Pages 127-140 In: P. W. Glynn, editor. Global Ecological Consequences
monitoring project for July 1992 to December 1999. Washington of the 1982-83 El Niño Southern Oscillation. Elsevier Oceanography
Department of Fish and Wildlife, Olympia, WA. Series, 52. University of Miami, FL.
129. Ocean Policy Advisory Council. 1994. State of Oregon Territorial Sea 151. Robinson, M. K. 1999. The Status of Washington’s coastal Marine
Plan. Ocean Policy Advisory Council, Portland, OR. Protected Areas. Washington Department of Fish and Wildlife.
130. Olesiuk, P. F. 1993. Annual prey consumption by harbor seals (Phoca Interjurisdictional Resource Management. Olympia, WA.
vitulina) in the Strait of Georgia, British Columbia. Fishery Bull., U.S. 91: 152. Rowlett, R. A., G. A. Green, C. E. Bowlby, and M. A. Smultea. 1994. The
491-515. first photographic documentation of a northern right whale off
131. ———, M.A. Bigg, and G. M. Ellis. 1990. Life history and population Washington State. Northwestern Naturalist 75: 102-104.
dynamics of resident killer whales (Orcinus orca) in the coastal waters 153. Rugh, D. J., M. M. Muto, S. E. Moore, and D. P. DeMaster. 1999. Status
of British Columbia and Washington State. Reports to the International review of the Eastern North Pacific stock of gray whales. NOAA Tech.
Whaling Commission, Special Issue 12: 209-243. Memo. NMFS-AFSC-103. Alaska Fisheries Science Center, NMFS,
132. Oregon Department of Fish and Wildlife. 1992. Shorebird survey of NOAA, 7600 Sand Point Way NE, Seattle, WA.
the Lower Columbia Estuary: August 1992. Unpublished table of survey 154. Sanger, G. A. 1970. The seasonal distributions of some seabirds off
results on file with U.S. Fish and Wildlife Service, Portland, OR. Washington and Oregon, with notes on their ecology and behavior.
133. ———. 1993. Shorebird survey of the Lower Columbia Estuary: Condor 72: 339-357.
August 22, 1993. Unpublished report to U.S. Fish and Wildlife Service, 155. Scheffer,V. B., and J. W. Slipp. 1944. The harbor seal in Washington
Portland, OR. State. American Midland Naturalist 32: 373-416.
134. ———. 1994a. Shorebird survey of the Lower Columbia Estuary: 156. ———, and ———. 1948. The whales and dolphins of Washington
April 24, 1994. Unpublished report to U.S. Fish and Wildlife Service, State with a key to the cetaceans of the west coast of North America.
Portland, OR. American Midland Naturalist 39: 257-337.
135. ——— 1994b. Shorebird survey of the Lower Columbia Estuary: 157. Schick, C. T., L. A. Brennan, J. B. Buchanan, M. A. Finger, T. M. Johnson,
August 21, 1994. Unpublished report to U.S. Fish and Wildlife Service, and S. G. Herman. 1987. Organochlorine contamination in shorebirds
Portland, OR. from Washington State and the significance for their falcon predators.
136. ———. 1994c. Shorebird survey of the Lower Columbia Estuary: Environmental Monitoring and Assessment 8: 1-17.
April 1994. Unpublished report to U.S. Fish and Wildlife Service,
Portland, OR.
 422 Wildlife–Habitat Relationships in Oregon and Washington

158. Seliskar, D. M., and J. L. Gallagher. 1983. The ecology of tidal marshes 178. ———. 1984. Distribution and abundance of seabirds over the
of the Pacific Northwest coast: A community profile. USDI Fish and continental shelf off Washington (including information on selected
Wildlife Service, FWS/OBS-82/32. marine mammals). Washington State Department of Ecology.
159. Shukla, J. 1998. Predictability in the midst of chaos: A scientific basis 179. ———. 1996. Waterbirds in Washington’s inland marine waters:
for climate forecasting. Science 282:728-731. some high counts from systematic censusing. Washington Birds 5: 29-
160. Siegel-Causey, D., and N. M. Litvinenko. 1993. Status, ecology, and 50.
conservation of shags and cormorants of the temperate North Pacific. 180. ———, D. G. Ainley, A. H. Benedict, and A.R. DeGange. 1989.
Pages 122-130 in: K.Vermeer,, K. T. Briggs, K. H. Morgan, and D. Siegel- Associations between seabirds and water masses in the northern
Causey, editors. The status, ecology and conservation of marine birds of Pacific Ocean in summer. Marine Biology 103: 1-17.
the North Pacific. Canadian Wildlife Service Special Publication, Ottawa, 181. ———, and D. Heinemann. 1979. Seabirds and fishing vessels: co-
Canada. occurrence and attraction. Condor 8: 390-396.
161. Simenstad, C. A. 1983. The ecology of estuarine channels of the 182. ———, K. H. Morgan, and K. Vermeer. 1993. Seabird distribution off
Pacific Northwest coast: A community profile. U.S. Fish and Wildlife British Columbia and Washington. Pages 39-47 in: K.Vermeer, K. T.
Service, FWS/OBS-83-05. Briggs, K. H. Morgan, D. Siegel-Causey, editors. The status, ecology and
162. Sowls, A. L., A. R. DeGange, J. W. Nelson, and G. S. Lester. 1980. conservation of marine birds of the North Pacific. Canadian Wildlife
Catalog of California seabird colonies. U.S. Fish and Wildlife Service. Service Special Publication, Ottawa, Canada.
FWS/OBS 80/37. 183. ———, S. M. Speich, D. A. Mauwal, K.V. Hirsch, and C. Miller. 1981.
163. Spear, L. B. 1993. Dynamics and effects of western gull feeding in a Marine bird populations of the Strait of Juan de Fuca, Strait of Georgia
colony of guillemots and Brandt’s cormorants. Journal of Animal and adjacent waters in 1978 and 1979. U.S. Environmental Protection
Ecology 62: 399-414. Agency, DOC/EPA interagency Energy/Environment R&D Program
164. ——— T. M. Penniman, J. F. Penniman, H. R. Carter, and D. G. Ainley. Report EPA/600/f-81/156.
1987. Survivorship and mortality factors in a population of western 184. ———, and ———. 1983. First winter survey of marine birds in
gulls. Studies in Avian Biology 10: 44-56. Puget Sound and Hood Canal, December 1982 and February 1983.
165. Speich, S. M., and S. P. Thompson. 1987. Impacts on waterbirds from Report for the Washington Department of Game. Nongame Wildlife
the 1984 Columbia River and Whidbey Island, Washington, oil spills. Program.
Western Birds 18: 109-116. 185. ———, and ———. 1984. Survey of marine birds in Puget Sound,
166. ——— B. L. Troutman, A. C. Geiger, P. J. Meehan-Martin, and S. L. Hood Canal and waters east of Whidbey Island, Washington, in summer
Jeffries. 1987. Evaluation of military flight operations on wildlife of the 1982. Western Birds 15: 1-14.
Copalis National Wildlife Refuge, 1984-1985. Department of the Navy, 186. ———, and ———. 1994. Distribution of foraging Rhinoceros
West. Div., Naval Facilities Engineering Command, San Bruno, CA. Auklets in the Strait of Juan de Fuca, Washington. Northwestern
167. ———, and T. R. Wahl. 1989. Catalog of Washington seabird colonies. Naturalist 75: 63-69.
U.S. Dept. Interior, Fish and Wildl. Serv., Biol. Rept. 88(6). MMS 89-0054. 187. ———, and B. Tweit. In press. Seabird abundances off Washington,
168. Springer, A. M., A. Y. Kondratayev, H. Ogi, Y. V. Shibaev, and G. B. van 1972-1998.Western Birds.
Vliet. 1993. Status, ecology, and conservation of Synthliboramphus 188. Washington Department of Natural Resources. 1998. Our changing
murrelets and auklets. Pages 187-203 in: K.Vermeer, K. T. Briggs, K. H. nature: Natural resource trends in Washington state. Olympia, WA.
Morgan, and D. Siegel-Causey, editors. The status, ecology and 189. Weins, J. A., and J. M. Scott. 1975. Model estimation of energy flow in
conservation of marine birds of the North Pacific. Canadian Wildlife Oregon coastal seabird populations. Condor 77: 439-452.
Service Special Publication, Ottawa, Canada. 190. Weitkamp, L. A., R. C. Wissman, C. A. Simenstad, K. L. Fresh, and J. G.
169. Steiger, G. H., J. Calambokidis, D. K. Ellifrit, K. C. Balcomb, J. D. Darling, Odell. 1992. Gray whale foraging on ghost shrimp (Callianasa
and G. A. Green. 1999. Distribution and population structure of californiensis) in littoral sand flats of Puget Sound, USA. Canadian Journal
humpback whales off Oregon and Washington. in: Abstracts Thirteenth of Zoology 70: 2275-2280.
Biennial Conference on the Biology of Marine Mammals, Maui, HI 28 191. Widrig, R. S. 1979. The shorebirds of Leadbetter Point. Independently
November—3 December 1999. Society for Marine Mammalogy, published.
Lawrence, KS. 192. Wiedemann, A. M. 1984. The ecology of Pacific Northwest coastal
170. Temte, J. L. 1993. Population differentiation of the Pacific harbor seal: sand dunes: A community profile. USDI Fish and Wildlife Service, FWS/
cranial morphology parallels birth timing. Ph.D. dissertation, University OBS-84/04.
of Wisconsin, Madison, WI. 193. ———. 1998. Coastal foredune development, Oregon, USA. Journal
171. U.S. Fish and Wildlife Service. 1997. Pacific Coast Joint Venture of Coastal Research, Special Issue 26: 45-51.
strategic plan. Portland, OR. 194. ———, and A. Pickart. 1996.The Ammophila problem on the
172. ———. 1999. Endangered and threatened wildlife and plants; northwest coast of North America. Landscape and Urban Planning 34:
Designation of critical habitat for the Pacific coast population of the 287-299.
Western Snowy Plover; Final rule. Federal Register 64: 68507-68544. 195. Wilson, U. W. 1991. Responses of three seabird species to El Niño
173. Varoujean, D. H. 1979. Seabird colony catalog: Washington, Oregon events and other warm episodes on the Washington coast, 1979-1990.
and California. U.S. Fish and Wildlife Service, Portland, OR. Condor 93: 853-858.
174.Veit, R. R., J. A. McGowan, D. G. Ainley, T. R. Wahl, and P. Pyle. 1997. 196. ———. 1996. Washington common murre colony surveys 1996.
Apex marine predator declines ninety percent in association with USFWS, Sequim, WA.
changing oceanic climate. Global Change Biology 3: 23-28. 197. ———. 1999. Common murre colony surveys, Washington Islands
175. ———, P. Pyle, and J. A. McGovan. 1996. Ocean warming and National Wildlife Refuge, 1997-1998. USFWS, Port Angeles, WA.
long-term change in pelagic bird abundance within the California 198. ———, A. McMillan, and F. C. Dobler. 2000. Nesting, population
current system. Marine Ecology Progress Series 139: 11-18. trend and breeding success of peregrine falcons on the Washington
176.Vermeer, K., D. B. Irons, E.Velarde, and Y. Watanuki. 1993. Status, outer coast, 1980-98. Journal of Raptor Research 34:67-74.
conservation and management of nesting Larus gulls in the North 199.York, A. E. 1995. The relationship of several environmental indices to
Pacific. Pages 131-139 in: K.Vermeer, K. T. Briggs, K. H. Morgan, and D. the survival of juvenile male northern fur seals (Callorhinus ursinus)
Siegel-Causey, editors. The status, ecology and conservation of marine from the Pribilof Islands. Pages 317-327 in: R. J. Beamish, editor. Climate
birds of the North Pacific. Canadian Wildlife Service Special Publication, change and northern fish populations. Canadian Special Publication for
Ottawa, Canada. the Fisheries and Aquatic Science No. 121.
177. Wahl, T. R. 1975. Seabirds in Washington’s offshore zone. Western
Birds 6: 117-134.
 16
Introduced Wildlife of Oregon and Washington
Gary W.Witmer & Jeffrey C. Lewis

Introduction
Each species of wildlife occurs as part of an ecosystem, In this chapter, we present information on wildlife
interacting in many ways with other plant and animal introduced to Oregon and Washington. While other terms
species in that system as well as with the abiotic could be used, we will refer to these relatively new
components such as soil, air, water, and other substrates. members of the fauna of Oregon and Washington as
The array of wildlife species around the globe has been “introduced” species. Occasionally, the term “exotic” will
shaped by geological and climatological events as well as be used, especially in the context of legal terminology, such
by eons of evolution and natural selection. Species have as state or federal laws and regulations. We will not include
come and gone and those remaining have, in most cases, species that are expanding their range on their own
co-evolved or co-adapted with many other species so that without the direct intervention of humans; examples of
relatively stable, and often complex, relationships exist. these species include the cattle egret and the barred owl.
Usually, a great many niches have been carved out and Also, we will not include the reintroduction or population
occupied, creating a distinct flora and fauna in each region augmentation of native species, although much of this
of the globe that is maintained under conditions of relative activity is occurring in the Pacific Northwest for
stability over time. Natural disturbances (wind, fire) and conservation and biodiversity purposes.
large-scale events (volcanic eruptions, drought) may Additionally, we will only include introduced species
occasionally alter that stability and the relationships of birds, mammals, amphibians, and reptiles. In additional
between species, but an overall homeostatis “a return to to at least 125 species of vertebrates, it has been estimated
the climatic community steady state” usually prevails. that over 2,000 species of plants and over 1,100 species of
These and other concepts of biogeography have been invertebrates have been introduced into the U.S., along
discussed in detail.31, 162 with 111 species of fish and over 50 plant pathogens.116
Species we refer to as “native” or “indigenous” There are many introduced species of plants, invertebrates,
naturally occur in a particular area and have been there and fish that occur in the Pacific Northwest, and many
for a very long time. However, events can occur that bring are of major concern with regard to ecosystem integrity,
individuals of a new species into a region where they come natural resource management, crop protection, or human
into contact with many species with which they are neither health and safety. Detailed discussions on the rapid and
co-adapted nor co-evolved. In most cases, these newly destructive spread of various noxious plant species that
arrived individuals soon succumb, but some may survive have been introduced to the Pacific Northwest have been
long enough to interact with, or disturb, normal presented by Peck,120 Stein and Flack,139 and Toney et al.146
relationships in the community. In a few cases, the Most plant species introduced to the Pacific Northwest
newcomers may survive, reproduce, and become have been perennial forbs originating from Eurasia,
established in the ecosystem, permanently altering although there has been a trend towards woody species
relationships among or between species. These newcomers introductions in more recent years.146 It is ironic that many
are usually referred to as “introduced,” “exotic,” “non- plant species were purposefully introduced for wildlife
native,” or “non-indigenous.” Species that are very habitat enhancement.130 Introduced invertebrate species
successful at this are sometimes called “invasive” species. and their impacts have been discussed,68, 139 as well as
These species are often capable of spreading unchecked, introduced fish species and their impacts to aquatic
increasing to high population levels, and comprising a systems.30, 139
significant portion of the total biota. In cases where the In this chapter we will discuss why wildlife
species has thrived in the new location for a relatively long introductions occur; the benefits and problems associated
period of time (in terms of human generations), they are with introductions; regulation of introductions; the
considered “naturalized” and are essentially considered introductions that have occurred in Oregon and
a regular part of the local flora and fauna (species Washington; the known or potential interactions between
complex). introduced species and native species; and the

423
 424 Wildlife–Habitat Relationships in Oregon and Washington

management of introduced species. We will also include Table 1. Some reasons why wildlife introductions
several case histories that characterize favorable and occur.
adverse aspects of wildlife introductions.

How Introductions Occur A. Accidental introductions

Wildlife species can be introduced to new areas through a 1. Escaped captivity
variety of mechanisms, both accidental and purposeful 2. Stowaways
(Table 1). Accidental introductions can result from animals 3. Expanded range of species after introduction elsewhere
escaping captivity, as has occurred with fox, mink, monk B. Purposeful introductions
parakeet (Myiopsitta monachus), various livestock species, 4. Aesthetics
and an array of wild ungulates such as fallow (Dama dama) 5. Economics
and axis (Axis axis) deer. As stowaways on ships, trains or 6. Recreation
other vehicles, some rodent species (Norway rat, black 7. Source of food
rat, house mouse) and bird species (house sparrow) have 8. Filling “vacant” niches
achieved worldwide distribution. Finally, human 9. Biological control
alteration of habitats or native species ranges, after an 10. Released from captive population
initial introduction elsewhere, has resulted in the altered 11. Release of rehabilitated or problem animals
and often expanded range of number of species such as 12. Whimsy: “what the heck, let’s see what happens”
the opossum into regions of the country in which they 13. Gifts
did not historically occur.
Purposeful introductions have occurred for many captivity on a whim: “what the heck, let’s see what
reasons (Table 1). The desire to have bird species from the happens.” This may have occurred with some parakeet/
countries of their European heritage, hence aesthetics, led parrot species as well as with some reptile and amphibian
the Portland Song Bird Club to attempt many songbird species. Finally, persons have given animals (wild or
introductions, including the starling.46, 76 Similarly, eastern domestic) away as gifts, which later escaped or were
gray and fox squirrels have been released in many urban/ released and established free-ranging populations.
suburban areas of the west.154
Many “game” species, as well as some domestic What Makes Introductions
livestock species, have been introduced to provide some Succeed or Fail?
combination of recreational hunting or economic benefit Most wildlife introductions, whether accidental or
(fur, food, or clothing). These species include many species purposeful, fail to establish free-ranging and sustained
of upland game birds, turkeys (Meleagris gallopavo), foxes, populations.59, 163 There are many reasons why this is the
eastern cottontail rabbits, nutria, and various species of most likely outcome of an introduction: inadequate
deer. Domestic species have been released to provide a numbers of animals, poor health or genetic quality of
future source of food or transportation (pig, Sus scrofa; animals, predation, disease or parasites, inadequate
goat, Capra hircus; horse, Equus caballus; burro, Equus habitat, competition with native species, poor planning,
asinus). and others.59, 163
In some cases, species were introduced to fill a On the other hand, certain characteristics of a species
perceived vacant niche, as with upland game birds, or population make it more likely to be successful at
carnivores on islands, and herbivores on islands. In “invading” a new area and becoming established.44, 112
actuality, in some of those cases, the populations of native These include a large native range, high mobility, broad
species occupying those niches had been greatly reduced diet, short generation time, high genetic variability,
by over-harvest or by human-induced changes in habitats gregariousness, larger size than most closely related
or predator-prey relationships (e.g., many native grouse species, few predators, association with humans,
species). Carnivores (such as fox; mongoose, Herpestes association with freshwater habitats, and ability to
spp.; European ferret, Mustela putorius; and domestic cat, function under a wide range of physical conditions. Often
Felis catus) have been introduced, especially on islands, to these species are “habitat generalists” and have a “broad
help control rodent or rabbit populations, many of which ecological amplitude.”44
were also introduced accidentally or purposefully at an It is important to recognize that many factors are
earlier date. This form of biological control has rarely, if involved in the success or failure of a wildlife introduction.
ever, been successful in its intended purpose. Even chance and timing play a role.33 Disturbance of a
In some cases, animals kept in captivity were released site or community, often human-induced, may make the
because the owners no longer cared or could afford to area more susceptible to invasion.51, 117, 125 As such, it is
maintain the animals, or because the economic incentive difficult to predict whether or not a given introduction
to raise the animals had declined, as with bullfrogs, nutria, will succeed or fail.129, 136, 137 There have been some efforts
mink, fox, exotic deer, “road-side zoo” animals, and some to construct predictive models of the likelihood of
species of livestock. In a few cases, the animals released successful establishment of an introduced species.136, 137
from captivity may have been rehabilitated animals or Unfortunately, there is much to be learned in this area of
problem animals. Some persons may release animals from ecology. In terms of regulation of wildlife introductions,
 CHAPTER 16: INTRODUCED SPECIES 425

Table 2. Potential adverse ecological consequences of

introduced wildlife species.

A. Effects on physical environment

1. Water quality, quantity
2. Soil compaction
3. Soil erosion
4. Nutrient balance
B. Effects on flora
5. Species composition
6. Species abundance
7.Vegetative structure
8. Plant succession
9. Species endangerment
Red fox. (Photograph by Jeffrey C. Lewis)
C. Effects on fauna
10. Competition
this situation has historically resulted in an “innocent until a. food
proven guilty” attitude, and species introductions are not b. habitats
prohibited until it is known that they are likely to cause c. interference
adverse effects; and once these occur, they may be 11. Predation
impossible to reverse. 12. Disease/parasite transmission
13. Hybridization
Potential Benefits and Adverse Effects of 14. Species endangerment
Introductions D. Direct effects to humans
A large number of introductions of plants and animals 15. Disease/parasites to humans, livestock, pets
has already occurred, and continues to occur, in the United 16. Crop damage
States.116 Some past introductions have benefited the public 17. Structural damage
for the reasons listed in Table 1. Consider, for example, 18. Livestock predation
domestic livestock and upland game birds. In Oregon, the 19. Livestock forage competition
introduction of ring-necked pheasant in 1881 resulted in 20. Human food consumption and contamination
large economic and recreational benefits—so much so, that 21. Human safety
captive-reared birds were soon being exported to many 22. Aesthetics
other states.35 Upland game species (both bird and small E. Major ecosystem disruption or alteration
mammal) continue to provide large revenues and 23. Combinations of the above effects
extensive recreation to many states.73
There are many potential or realized ecological in Washington. Major disruptions are most common on
consequences of wildlife introductions (Table 2). To date, islands where rats,165 carnivores,164 or feral livestock148 have
the most visible effects of introductions to Pacific been introduced. Erosion and community changes (species
Northwest ecosystems appear to be from plant and composition, abundances, biodiversity, and species loss)
invertebrate species introductions, although overgrazing have occurred in these situations. On the North American
by domestic livestock has affected some dryland areas. mainland, similar effects have occurred in the Great Smoky
We note that the effects of an introduced wildlife species, Mountains National Park from the introduction of feral
however, may take hundreds of years to become evident: pigs.10 Feral horses and burros have had substantial
the “blink of an eye” in ecological time. The effects can be impacts on some southwestern ecosystems.41 Species-
to the physical environment, the flora, the fauna, and specific examples of realized or (more often) potential
humans directly, or more often, to a combination of these ecological effects of wildlife introductions in Oregon and
ecosystem elements. Perhaps the most common effects are Washington are presented later in this chapter.
from herbivory, competition, or predation. However,
many other types of effects can occur, such as Regulation of Wildlife Introductions
hybridization with native species 133 and disease The regulation, policies, and guidelines for wildlife
transmission.52 Numerous examples of ecological effects introductions in the United States have had a long and
were presented by MacDonald et al.107 and Simberloff.134 convoluted history. A large number of governmental
In some cases, a major disruption of the ecosystem can agencies—at federal, state, and local levels—have played
occur, but this has not yet been well documented from roles; the net effect often being inconsistent, inadequate,
wildlife introductions in terrestrial ecosystems in Oregon or contradictory efforts among agencies, or policies that
and Washington, with the possible exceptions of San Juan changed dramatically over time within an agency.116, 131
Island (European rabbit),28 Destruction Island (European There is a strong need for not only more regulation of
rabbit),6 and the Olympic Peninsula (mountain goat),67 all species introductions, but also better coordination of
 426 Wildlife–Habitat Relationships in Oregon and Washington

regulation across jurisdictional boundaries and under a permit and a commitment from the state wildlife
governmental levels.88, 116 agency, 7) treatment of exotics leaving federal land as
Regulations and practices have evolved from trespassing livestock with the responsible party held liable,
encouraging the importation and release of animals to and 8) periodic review of public policy regarding exotic
improve agricultural resources, hunting opportunities, or species.
local economies to restricting importations because of The Wildlife Society published a policy statement on
disease hazards, threats to agricultural resources or human species introductions in 1975. This policy included the
health and safety, or potential disruption of natural following three considerations:91
ecosystems. As early as 1923, Taylor142 discussed benefits, 1. Support the introduction of exotic species only after
adverse effects, methods, and regulations for the competent scientists have thoroughly studied the
introduction of upland game birds in the Pacific situation and potential effects and quarantine
Northwest. requirements have been met.
The U. S. Department of Interior’s (USDI) Federal— 2. Urge that no state, provincial, or national agency
State Cooperative Foreign Game Program of 1948 added introduce an exotic species or permit such an
an element of central authority at a time when the introduction unless that species can be contained
importation of game species into the United States was within its jurisdiction, or unless surrounding
being strongly pursued. This program was guided by three jurisdictions have sanctioned the introduction.
objectives: to provide an ecological and life history data 3. Exclude from the provisions of this policy the
base to individuals or agencies, to discourage importation of exotic species by officially recognized
introductions when the data base suggested an scientific and educational organizations, and the
introduction was unwise, and to fill vacant or institutional exchange of such species provided that
understocked habitats with foreign species as an alternate the exotics are maintained in captivity at all times.
course of action following appropriate testing and trial President Carter signed Executive Order 11987 in 1977.
introductions.128 This document, in part, restricted federal agencies from
In 1966, the USDI published guidelines and introducing species to lands they administer, encouraged
recommendations for the importation of wildlife.128 These the prevention of introductions by other levels of
guidelines incorporated eight conditions: government and by private citizens, and restricted federal
1. Critically determine that a need exists, with desirable support of introductions outside the United States.128 These
ecological, recreational, and economic impacts. limitations applied unless either the Secretary of Interior
2. A definite niche is available and unsuited for a native or the Secretary of Agriculture determined that the
species. introduction would not have an adverse effect on natural
3. Introductions should not be considered if they ecosystems.
threaten the reduction or displacement of native An international position statement, containing policies
populations; nor should existing or proposed land and guidelines similar to those above, was developed and
uses be in conflict with an exotic species transplant. approved by the International Union for Conservation of
4. Introductions should be preceded by ecological Nature and Natural Resources (IUCN) in 1987.116 The
studies of both the animal and the habitat proposed IUCN is an organization comprised of scientific experts
at the release site. and government officials involved in conservation around
5. Disease relationships require special study as well as the world.
the steps assuring appropriate quarantine leading to The concern about “invasive alien” species continues
disease-free stock. to generate activity by the federal government. On June
6. Exotic species with close relatives in the United States 17, 1997, Vice President Gore directed the preparation of
should be avoided, to preclude hybridization with a strategy to combat the introduction and spread of non-
native wildlife. native plants and animals in the United States that are
7. Small-scale experiments and a thorough evaluation of causing great economic and ecological harm to the nation.
these should precede larger introductions. A draft document has been prepared that reviews the
8. Before an exotic species is released, methods for situation, makes recommendations, and provides an
controlling its abundance and expansion must be action plan. 1 Based on the results of the Task Force,
available. President Clinton signed an Executive Order on Invasive
These guidelines resulted in eight recommend-ations Species on February 3, 1999. Its goals are to prevent the
from the USDI. These were meant to apply to federal lands introduction of invasive species, to provide for their
or federal actions and included: 1) no decisions until a control, and to minimize the economic, ecological, and
full assessment is at hand, 2) no exotic species placed in human health impacts that they cause. It was estimated
national parks or lands devoted to the preservation of that invasive species cost the U.S. economy about $123
native biota, 3) no exotic species placed in the vicinity of billion per year. The Order establishes Invasive Species
rare or uncommon native species, 4) no exotic grazers Council assigned the task of setting up an Advisory
placed on federal lands devoted to domestic grazers, 5) Committee and preparing and implementing an Invasive
no exotic big game placed in areas devoted to intensive Species Management Plan. The Plan will 1) detail and
land uses, 6) no introductions on federal lands except recommend performance-oriented goals and objectives,
 CHAPTER 16: INTRODUCED SPECIES 427

2) review existing and prospective approaches and

authorities, 3) identify pathways of introductions and
ways to minimize risks of introduction, 4) identify research
needs, 5) be science-based, 6) recommend and implement
measures to reduce introductions and control those that
have occurred, 7) identify requirements to achieve goals
and objectives, and 8) evaluate and report on the success
in achieving goals and objectives.
We have reviewed some of the long history of
introduced wildlife concerns, policies, and
recommendations. The groundwork has been set for a
vigorous effort to reduce introductions and to manage
existing introductions. It remains to be seen what level of
success will be achieved towards this goal.
Bullfrog. (Photograph by Jeffrey C. Lewis)
Current Federal and State Regulations
The two main federal agencies regulating wildlife species
introductions in the United States are the U.S. Department For many decades, many states, including Oregon and
of Agriculture’s (USDA) Animal and Plant Health Washington, had few regulations regarding the
Inspection Service (APHIS) and the USDI Fish and Wildlife importation or keeping of “exotic” wildlife or the
Service (FWS).116, 131 A major function of APHIS is to protect protection of native biodiversity from exotics.21, 116 Many
United States agriculture (both plants and animals) from states in the late 1800s and early 1900s, including Oregon
diseases or plant and animal “pests” that might gain access and Washington, encouraged—or were directly involved
to the country or be transported between states. Border with—the propagation or release of many game species,
inspections, quarantines, disease testing, and eradication including exotic species. These practices have largely been
programs are some of its routine functions, and most of curtailed in recent decades, with notable exceptions such
APHIS’s pest exclusion occurs at ports of entry. APHIS as with wild turkey and Sichuan pheasant, P. c.
does not expressly prohibit species-specific imports, but suehschanensis.116 The Southeastern Cooperative Wildlife
requires adequate quarantine and veterinarian inspection Disease Study provided a “model law” in 1988 to help
before such imports or transportations are allowed (9 CFR guide states in regulating animal imports that addressed
Ch. 1). The agency is particularly strict regarding veterinary, humane, public safety, ecological, and other
hedgehogs (Erinaceus spp.) and the brush-tailed possum concerns.116 It recommended a permit requirement for
(Trichosurus vulpecula). A major concern of APHIS is to introduced species, that certain common domestic and
prevent the entry of Newcastle’s disease, chlamydiosis, naturalized species be exempt from the regulations, that
foot-and-mouth disease, rinderpest, bovine tuberculosis, criteria and a list be developed for “environmentally
and other communicable diseases of livestock and wildlife. injurious animals,” and that a technical advisory
APHIS is also active in management and research to committee be formed to provide advice. Both Oregon and
prevent entry of the brown tree snake (Boiga irregularis) Washington legislatures and wildlife agencies have
into Hawaii and the mainland United States.17 enacted detailed and specific regulations on the
The FWS protects threatened and endangered species importation and keeping of introduced wildlife.
by, among other activities, restricting the importation and Oregon Administrative Rules (OAR 635-056-0000 to -
exportation of federally listed species under the 0150) prohibit the importation or keeping of numerous
Endangered Species Act and the Convention on vertebrate species, including hedgehog, tri-colored
International Trade in Endangered Species (CITES). Except squirrel (Callosciurus spp.), brush-tailed possum, bats of
under permit and various restrictions, the FWS expressly any species, mongoose, wild pig, chamois (Rupicapra spp.),
prohibits the importation and release of individuals, non-domestic goat (Capra spp., except C. hircus),
progeny, or eggs of many species of vertebrates into the wildebeest (Connochaetes spp.), gazelles (Procapra spp.),
United States, to protect national resources (50 CFR Ch. capybara (Hydrochaeris hydrochaeris), prairie dogs (Cynomys
1). These species include flying fox (Pteropus spp.), spp.), any species of wild canid (except fox), Egyptian
mongoose, European rabbit, wild dog (genus Cuon), multi- goose (Alopochen aegyptiacus), African clawed frog
mammate rat (Mastomys spp.), raccoon dog (Nyctereutes (Xenopus spp.), bullfrogs (Pyxicephalus spp.), alpine newt
procyonoides), starling, quelea (Quelea qualea), Java sparrow (Triturus spp.), brown tree snakes, snapping turtle
(Padda oryzivora), red-whiskered bul-bul (Pycnonotus (Chelydridae, all spp.), pond sliders (Pseudemys spp.),
jocosus), all species of amphibians, and all species of Chinese pond turtle (Chinemys spp.), pond turtle
reptiles. Additionally, the importation and transportation (Chrysemys spp.), painted turtle (Chrysemys spp.), map
of birds of the family Psittacidae (parrots, parakeets, turtle (Graptemys spp.), North American (Apalone spp.) and
macaws, etc.) are regulated by the U.S. Public Health African (Trionyx spp.) soft shell turtles, European pond
Service because of disease hazards (42 CFR Parts 71 and turtle (Emys orbicularis), Blanding’s turtle (Emydoidea
72). blandingii), common mud turtle (Kinosteron subrunum),
 428 Wildlife–Habitat Relationships in Oregon and Washington

common musk turtle (K. odoratum), and Asian pond turtle important to increase public awareness of the risks posed
(Mauremys spp.). Many fish species are also prohibited. by wildlife introductions. A good public education
Exceptions, by permit, are made for zoos and research program on this subject should lead not only to more
facilities if they are escape-proof and are staffed and public support for the prevention of future introductions
equipped to provide adequate care. There is also a long and the management or eradication of past introductions
list of domestic or otherwise exempt species, including (in terms of supporting appropriate legislation,
dogs (Canis familiaris), cats, burros, horses, swine, management practices, and requisite budgets), but also
European rabbits, ferrets, and parrots and parakeets for better public compliance with federal and state
(Psittacidae, all spp.). The State has specific requirements regulations.
involving the sale, transportation, and holding of exotic
animals, to help prevent escape of, or disease transmission Introduced Wildlife Species in Oregon and
by, introduced animals. There are strict reporting rules that Washington
apply when an introduced species escapes captivity. There have been a large number of wildlife introductions
Washington has similar laws (Chapter 77.08 RCW) and to Oregon and Washington, dating back to the 1700s (e.g.,
prohibits the importation or keeping of mute swan, horses and burros). Most attempted introductions,
mongoose, wild pig, collared peccary (Tayassu tajacu), and whether accidental or purposeful, have failed. For
various species of exotic bovids and cervids. Many species example, Portland Bird Club attempted but failed between
of fish are also prohibited. 1889 and 190746, 76 to introduce many species of songbirds,
Much of the importation of introduced wildlife into the including Eurasian skylark, Alauda arvensis; wood lark,
United States is because of the enormous pet industry.116 Lullula arborea; blackcap, Sylvia atricapilla; European robin,
About 23 percent of the vertebrate species of foreign origin Erithacus rubecula; nightingale, Luscinia megarhynchos;
that currently live in the wild in the United States were Eurasian blackbird, Turdus merula; song thrush, T.
originally imported as cage birds or other wildlife pets.116 philomelos; parrot crossbill, Loxia pytyopsittacus; Eurasian
There is a growing concern about the depletion of animals siskin, Carduelis spinus; Eurasian goldfinch, C. carduelis;
from exportation countries (hence the species listings in linnet, Acanthis cannabina; Eurasian bullfinch, Pyrrhula
CITES, including its appendices), but also about the pyrrhula; chaffinch, Fringilla coelebs; house sparrow, and
significant hazard that introduced species pose to native European starling.
species, ecosystems, agriculture, and forestry.1, 116, 124 It has We have compiled a list of 42 wildlife species
even been speculated that the liberalization of introductions to Oregon and Washington that have
international free trade may increase species introductions established free-ranging populations at least on a localized
around the globe.74 The probability of accidental release scale (Table 3).7, 20, 27, 34, 37, 46, 54, 70, 75, 76, 94, 97, 98, 99, 104, 152 The
of non-native wildlife as well as disease transfer and other information we provide on the 42 species includes
hazards can be reduced by the improvement of existing common and scientific names, the date, and location of
programs and the implementation of specific actions as the introduction(s), the reason for the introduction, the
presented by the Ad Hoc Federal Invasive Alien Species current status and distribution (in general terms), and the
Task Force1 and the Office of Technology Assessment.116 country of origin (Table 3). We note that many of the
Their recommendations include: acknowledge that the species were introduced over a period of time and at
prevention of introductions is paramount, encourage several locations. The list includes 18 birds, 19 mammals,
governments to take an active role by establishing national 3 reptiles, and 2 amphibians. About half (19 of 42) of the
and regional councils, develop new scientific expertise for species listed have achieved widespread distribution in
dealing with introductions, develop a white-black-gray Oregon or Washington.
list to assist in regulating exotic species, develop a Most bird species were introduced for hunting or
comprehensive program to prevent unintentional aesthetic purposes, although several arrived by range
introductions by identifying major pathways and methods expansion after being introduced elsewhere. Many of the
to interdict and reduce impacts, develop and implement mammal introductions were escapees or animals released
an international regime for control and support when no longer needed or economically valuable. Several
cooperation through development assistance, develop a were introduced for hunting or fur farming. The Old
Web-based network of information, convene educational World rodents arrived as stowaways. Most amphibian and
workshops, and consult with the United States Congress reptile introductions were for aesthetic, pet, or food
regarding new regulations and funding authority. The purposes.
white-black-gray list would delineate species that are Additionally, there are many other non-native wildlife
automatically allowed, never allowed, or allowed only species that have been observed or reported in Oregon or
after thorough investigation, respectively. Washington, but information on their occurrences is very
The implementation of adequate programs to prevent limited and we cannot be sure whether or not those species
the accidental, or purposeful but prohibited, release of are established (Table 4). We have included this species
introduced diseases, plants, invertebrates, and vertebrates list because of the potential ecological consequences if they
is especially important because of the difficulty and do become established and more widespread in
expense of eradication once an introduced species becomes distribution.
dispersed and established. 36, 135 Furthermore, it is
 CHAPTER 16: INTRODUCED SPECIES 429

Table 3. Wildlife species introduced to Oregon or Washington.7, 20, 27, 34, 37, 46, 54, 70, 75, 76, 94, 97, 98, 99, 104, 152

Species Place/Date Reason Status Origin

Trumpeter swan, Harney Co. OR 1939-58; aesthetics; hunting very limited, small numbers SE Alaska, NW Canada
Cygnus buccinator Spokane Co. WA 1963 in OR and WA and somewhat south

Mute swan, Cygnus olor Lincoln Co. OR 1950s; aesthetics, escapees? very limited, small numbers Eurasia
Deschutes Co. OR 1960s in OR and WA
American black duck, Snohomish Co.WA, hunting small localized popn in E United States
Anas rubripes date unknown Puget Sound
Chukar, Alectoris chukar Lake Co. OR 1951; hunting, brood stock sale scattered popns in E OR Eurasia
Deschutes Co. OR 1952; and E WA
Klamath Co. OR 1960s;
E. WA 1930s
Gray partridge, Linn Co. OR 1900; hunting, brood stock sale scattered popns NE OR Eurasia
Perdix perdix 23 counties OR 1913-14; and E WA
Spokane Co.WA 1906;
Columbia Co. WA 1908
Ring-necked pheasant, Linn Co. OR 1881-82; hunting, brood stock sale widespread, common Eurasia
Phasianus colchicus Protection Is. WA 1883
White-tailed ptarmigan, Wallowa Co. OR aesthetics?, hunting?
localized, small numbers in SE Alaska, W Canada
Lagopus leucurus 1967-69 OR; native to WA into WA
Wild turkey, OR 1899 (failed); manyhunting widespread, moderate E United States,
Meleagris gallopavo OR counties 1961-83; numbers in E OR and Southcentral United
E. WA 1970s E WA; some on San Juan Is. States
California quail, Thurston Co.WA 1857; hunting, brood stock sale widespread, common; native SW United States just
Callipepla californica many OR counties 1914 to S OR into OR
Scaled quail, Yakima Co. WA 1913 hunting, brood stock localized, small numbers; Southcentral United
Callipepla squamata sale, escapees extirpated? States
Northern bobwhite quail, Walla Walla WA 1865 & hunting, brood stock sale localized, small numbers E United States
Colinus virginianus 1920; Whidbey Is. WA
1871; Linn Co. OR 1882
Mountain quail, W WA 1880 hunting, brood stock sale localized in WA; native in SW United States into
Oreortyx pictus OR SE WA
Rock dove, Columba livia many OR counties aesthetics, racing, widespread, common Eurasia
<1900; W WA <1940 messengers, then range
expansion
Monk parakeet, Multnomah Co. OR escapees small numbers, Portland South America
Myiopsitta monachus 1969 area; some in WA?
Skylark, Alauda arvensis Portland OR 1889 aesthetics, then range small numbers on San Juan Eurasia
(failed?);Vanc. Is. BC expansion Is.
1903
European starling, Portland OR 1889 aesthetics, then range widespread, common OR Eurasia
Sturnus vulgaris (failed); arr. on own expansion after introd. and WA
1940s OR & WA to E United States
Crested mynah, Vancouver BC 1894 aesthetics, then range localized, small numbers in SE Asia
Acridotheres cristatellus expansion Seattle, Bellingham areas
House sparrow, Spokane Co.WA 1895; aesthetics and insect widespread, common OR Eurasia
Passer domesticus King Co.WA 1897; control to E United and WA
Portland OR 1889 States, range expansion
Virginia opossum, Umatilla Co. OR 1910; aesthetics, pets escapees, locally common, esp. W WA, E United States
Didelphis virginiana WA <1941 fur trapping W OR & NE OR
Red fox, Vulpes vulpes many places W WA by fox hunting, fur farming, widespread W OR & WA; Holarctic
1909, W OR by 1915 escapees less so in E OR & WA
European ferret, San Juan Is. WA rabbit control small population remains? Europe
Mustela putorius
 430 Wildlife–Habitat Relationships in Oregon and Washington

Species Place/Date Reason Status Origin

House cat, Felis catus <1800 escapees, pest control widespread, OR & WA? Eurasia, Africa
Domestic dog, <1800 escapees occasional occurrences Eurasia
Canis familiaris
Burro, Equus asinus E OR late 1700s escapees or released small population in SE OR Africa
when no longer needed
Horse, Equus caballus E OR late 1700s escapees or released moderate population in Asia
when no longer needed SE OR
Feral pig, Sus scrofa SW OR late 1800s; hunting, escapees? Very small, localized Eurasia
Skagit Co. WA 1981 populations or extirpated
Axis deer, Axis axis Pierce Co. WA >1980 aesthetics or escapees small, localized population India
Fallow deer, Dama dama King Co. WA >1980 aesthetics or escapees small, localized population Europe
Mountain goat, NE OR 1950 & aesthetics, hunting moderately abundant in Alaska to WA, Cascade
Oreamos americanus Columbia Gorge OR Olympic mtns., native to and Rocky Mtns.
1969 (failed); Olympic N Cascade & Rocky Mtns.
Mtns. WA early 1900s
Eastern gray squirrel, King Co.WA 1925; aesthetics localized, urban/suburban E United States
Sciurus carolinensis W OR 1919 areas of W & NE OR and
W WA
Fox squirrel, Sciuris niger W OR & WA <1940; aesthetics localized, urban/suburban E United States
Baker Co. OR 1950s areas of E WA, W and NE
OR
House mouse, OR and WA late 1700s stowaway, then range widespread, urban/suburban Europe
Mus musculus expansion areas OR & WA
Norway rat, OR and WA <1850 stowaway, then range localized, urban/suburban Asia
Rattus norvegicus expansion areas OR & WA
Black rat, Rattus rattus OR and WA <1800 stowaway, then range localized, urban/suburban Asia
expansion areas OR & WA
Nutria, Myocastor coypus King Co.WA 1930s; fur farming, escapees, localized, mostly W OR South America
Lincoln and Tillamook vegetation control? & WA
Cos. OR 1937
European rabbit, San Juan Co.. WA 1929; aesthetics, hunting Island populations persist, Europe
Oryctolagus cuniculus Destruction Is. WA 1970 other localized populations
in WA?
Eastern cottontail, Whitman Co. WA hunting widespread, locally abundant E United States
Sylvilagus floridanus 1926; Linn and Benton
Cos. OR 1940s
Bullfrog, Rana catesbeiana many places OR & WA insect control, aesthetics, widespread, locally abundant E and Central United
1914 on hunting, food, then range States
expansion
Green frog, King, Stevens and aesthetics, hunting? very localized, small E. United States
Rana clamitans Whatcom Cos. WA populations
Snapping turtle, many places W OR aesthetics, hunting?, localized, small populations E and Central United
Chelydra serpentina & W WA 1950s on pets, food States
Red-eared slider turtle, many places OR aesthetics?, pets, Locally common in W and SE United States
Trachemys scripta elegans and WA escapees Central OR
Plateau striped whiptail, Jefferson Co. OR aesthetics? Localized, small population SW United States
Cnemidophorus velox

Habitat Use by Introduced freshwater/riparian systems. Most species (71%) can be

Wildlife Species considered habitat generalists, using several general
All general habitat categories that occur in Oregon or habitat categories.
Washington are used by at least one of the 42 introduced Human-disturbed areas (agriculture lands, urban/
species, although few introduced species use alpine or suburban areas) are used by a large number of the
marine habitats (Table 5). Only 12 of 42 (29%) introduced introduced wildlife species, 19 and 18 species, respectively
species are affiliated with only one general habitat (Table 5). This group of species includes most introduced
category; most of those 12 species are restricted to upland game birds, songbirds, and mammals. Forests
 CHAPTER 16: INTRODUCED SPECIES 431

Table 4. Other introduced wildlife species that have been occasionally observed or reported in Oregon or
Washington.*

Birds Mammals Amphibians/reptiles

Domestic goose, Anser cygnoides Wolf-dog hybrid, Canis lupus x familiaris Eastern mud turtle, Kinosternon subrubrum
Egyptian goose, Alopochen aegyptiacus Domestic cow, Bos taurus Stinkpot, Sternotherus odoratus
Graylag goose, Anser anser Domestic goat, Capra hircus Painted turtle (non-natives), Chrysemys picta
Domestic mallards, Anas platyrhynchos Domestic sheep, Ovis aries Eastern box turtle, Terrapene carolina
Muscovy duck, Cairina moschata Barbary sheep, Ammotragus lervia Ornate box turtle, Terrapene ornata
Red-legged partridge, Alectoris rufa Mouflon sheep, Ovis musimon Malayan box turtle, Cuora amboinensis
Sichuan pheasant, Phasianus colchicus Desert tortoise, Gopherus agassizii
suehschanensis Texas tortoise, Gopherus berlandieri
Golden pheasant, Chrysolophus pictus Gopher tortoise, Gopherus polyphemus
Peafowl, Pavo cristatus Hermann’s tortoise, Testudo hermanni
Guineafowl, Numida meleagris Reeve’s turtle, Chinemys reevesi
Psittacines (misc. Parrots, Spiny softshell turtle, Apalone spinifera
cockatoos, macaws) Florida softshell turtle, Trionyx ferox
Big-headed turtle, Platysternon megacephalum
Caiman, Caiman crocodilus
* Little is known about the status of most of these species; most probably do not comprise free-ranging, self-sustaining populations
and have not expanded their range beyond the release site(s) in either state, however, these events could occur in the future.

Table 5. Use of general habitat categories by 42 wildlife species introduced to Oregon or Washington.

No. (% of Group) of Species by General Habitat Categorya

Wildlife Group Forest Shrub/grass Agriculture Urb/suburb Freshw/rip Marine Alpine

Birds (18 spp.) 8 (44%) 8 (44%) 11 (61%) 7 (39%) 4 (22%) 1 (6%) 1 (6%)
Mammals (19 spp.) 11 (58%) 4 (21%) 8 (42%) 11 (58%) 5 (26%) 0 (0%) 1 (5%)
Amphibians/reptiles 1 (20%) 1 (20%) 0 (0%) 0 (0%) 5 (100%) 0 (0%) 0 (0%)
(5 spp.)
Total: (42 spp.) 20 (48%) 13 (31%) 19 (45%) 18 (43%) 14 (33%) 1 (2%) 2 (5%)
a
Most species use more than one general habitat category.

(especially open, deciduous or mixed forests) are used by habitats, and grass-forb dominated areas. On the other
about half (48%) of the introduced species. Freshwater/ hand, there is little potential for competition between
riparian habitats are used by approximately equal white-tailed ptarmigans and ruffed grouse. Competition
numbers of introduced bird, mammal, and amphibian/ may also occur between introduced upland game birds
reptile species groups, with all introduced reptiles and and native sage and sharp-tailed grouse; all of these
amphibians using those habitats. Shrub/grass habitats are species use shrub-steppe and sagebrush-steppe habitats,
used by introduced bird species and eastern cottontail as well as grass-dominated areas and riparian areas. Most
rabbits, but especially upland game birds. of these species use agricultural lands, too. Likewise, in
Specific habitat associations for many of the introduced some situations, the introduced eastern cottontail rabbit
wildlife species have not been well defined. Some insight may compete with native rabbits (pygmy, and brush
for some species can be gained from Brown,12 Guenther rabbits; Nuttall’s cottontail) in grass/sedge meadows and
and Kucera60 and Thomas.144 That information, along with alder bottomlands as well as riparian areas, agricultural
species-specific literature and expert opinion, has been lands, brushpiles (including downed woody materials),
used to complete the wildlife habitat matrixes of this book. and burrows. For many introduced species, we do not
These matrixes can be used, to some extent, to project the know enough about what specific habitats they could use,
potential competition between native species and given the opportunity and time for populations to occupy
introduced species. For example, the introduced eastern those habitats. For example, axis and fallow deer could
gray and fox squirrels use oak woodlands as do native potentially occupy many of the same habitats as the native
western gray squirrels; all three species use variously-aged black-tailed deer and the endangered Columbia white-
forest stands and all use snags. White-tailed ptarmigans, tailed deer. Interested persons are referred to the matrixes
introduced to Oregon, may compete with native blue and for further investigation of potential habitat competition
spruce grouse in the use of alpine meadows, subalpine fir between native and introduced species.
 432 Wildlife–Habitat Relationships in Oregon and Washington

Ecological Consequences of
Wildlife Introductions
It is very important to recognize the potential ecological
consequences of wildlife introductions. While
considerable effort and expense have been invested in
dealing with introduced plants and insects in the Pacific
Northwest, introduced wildlife also has caused, or has the
potential to cause, substantial harm to Pacific Northwest
ecosystems and agricultural resources (Table 2). There are
many potential or realized ecological consequences for
each of the 42 wildlife species introductions that have
occurred in Oregon and Washington (Table 6).
Several points need to be emphasized. The code “NK”
(none known) appears frequently in Table 6. With wildlife
introductions, we are often presented with ecological Wild burro. (Photograph by Jeffrey C. Lewis)
“situations” with which we have little or no experience;
hence, our predictive powers are very limited.
Additionally, serious effects may occur long after the introduced rodent species cause many types of structural
introductions. The rather cavalier attitude of the past (“let’s damage to human dwellings, livestock facilities, and
do it and see what happens”) is no longer acceptable, given constructed features such as dikes, dams, levees,
the many legal mandates and policies for species and transmission lines, and irrigation systems.69 Considerable
biodiversity protection, healthy ecosystem maintenance, effort is expended each year to reduce the negative effects
and the protection of human health and agricultural of these species. On the other hand, the introduction of
resources. We must not only deal with existing upland game birds may have increased the prey base for
introduction problems, but must strive to prevent future native predators, both avian and mammalian, as well as
introductions that have significant potential for adverse having provided recreation as intended.
consequences. Both require a greater ecological and Most introduced mammal species have the potential
managerial knowledge base than we now possess. On the to adversely affect various species of native plants and
other hand, great strides in agriculture, recreation, local animals, as well as humans and their resources, through
economies, biological control of pests, and even in the herbivory or predation. Species such as red foxes, ferrets,
medical profession, have been made as a result of species and feral dogs and cats can inflict high levels of mortality
introductions. Obviously, a careful and deliberate analysis, on ground nesting birds and have been implicated in the
on a case-by-case basis, must be made before proceeding endangerment of numerous species, such as snowy
with any wildlife introduction. plovers, least terns (Sterna antillarum), and clapper rails
Introduced wildlife species have the potential for (Rallus longirostris).164 Similar results have been reported
several, if not numerous, adverse ecological consequences for introduced carnivores in Europe.82 Feral or free-ranging
(Table 6). Most introduced birds have the potential to cats kill large numbers of songbirds every year in urban/
adversely affect native birds, especially through forage and suburban settings.25 Essentially, all introduced herbivorous
nest site competition. A classic example is the ability of mammals can cause plant damage and may even impede
starlings to usurp nest sites from wood ducks, bluebirds, regeneration of some plant species. Usually the amount
woodpeckers of many species, and many other of damage is related to the density of introduced
songbirds.57, 155 Additionally, some hybridization problems mammals; hence, existing small, introduced populations
exist; for example, black ducks hybridize with mallards,133 of axis and fallow deer are probably not causing significant
some upland game species hybridize with native species8 impact to the native flora. On the other hand, high
and eastern cottontail rabbits hybridize with the brush densities or concentrated use can cause substantial impacts
rabbit.151 Avian diseases, such as avian tuberculosis, on native flora, as occurs with European rabbits, mountain
Newcastle’s disease, salmonellosis, and chlamydiosis, can goats, and feral pigs, horses, and burros. In some cases,
be transmitted to native species.52 Members of the parrot endangerment of native plant species may occur, as with
family make popular pets, but they have the potential to herbivory by introduced mountain goats on the Olympic
spread avian tuberculosis, a disease transmissible to not Peninsula.67 Many of the introduced mammal species,
only other wildlife, but to pets, livestock, and humans.52 especially carnivores and Old World rodents, have been
Congregations of introduced bird species such as house implicated in the transmission of disease to native wildlife,
sparrows, rock doves, and starlings at roosts or feeding livestock, pets, or humans; these diseases include rabies,
stations have produced significant disease hazards (e.g., plague, distemper, relapsing fever, and leptospirosis.38
histoplasmosis, ornithosis, salmonellosis).52 Several of the Introduced amphibians and reptiles have been
introduced bird species (starling, house sparrow, rock implicated in the decline of many native aquatic fauna
dove, skylark, crested mynah, Acridotheres cristatellus) have through predation or competition.29, 111 The introduced
a great potential to damage crops, contaminate foodstuffs, bullfrog is a classic example and will be considered in more
and cause aesthetic problems. 52, 55 Additionally, the detail in the case histories at the end of the chapter.
 CHAPTER 16: INTRODUCED SPECIES 433

Table 6. Potential or realized ecological consequences of wildlife species introduced to Oregon or Washington.

Species Physical Flora Fauna Human

Environment

Trumpeter swan, Cygnus buccinator NK NK FC, NC, AB AB

Mute swan, Cygnus olor NK NK FC, NC, AB AB
American black duck, Anas rubripes NK NK H NK
Chukar, Alectoris chukar NK NK D, FC, NC, IP CD
Gray partridge, Perdix perdix NK NK D, FC, NC, IP CD
Ring-necked pheasant, Phasianus colchicus NK NK D, FC, NC, IP, H CD
White-tailed ptarmigan, Lagopus leucurus NK NK FC, NC, AB NK
Wild turkey, Meleagris gallopavo NK PR FC, IP NK
California quail, Callipepla californica NK NK D, FC, NC, IP, H CD
Scaled quail, Callipepla squamata NK NK D, FC, NC, IP, H CD
Northern bobwhite quail, Colinus virginianus NK NK D, FC, NC, IP, H CD
Mountain quail, Oreortyx pictus NK NK D, FC, NC, IP, H CD
Rock dove, Columba livia NK NK FC, NC, D A, HD, CF
Monk parakeet, Myiopsitta monachus NK PD? FC, NC, D CD
Skylark, Alauda arvensis NK PD? FC, NC, D? CD?
European starling, Sturnus vulgaris NK PD? FC, NC, D CF, A, HD
Crested mynah, Acridotheres cristatellus NK PD? FC, NC, D? CF, A, HD?
House sparrow, Passer domesticus NK PD? PC, FC, NC, D CF, A, HD
Virginia opossum, Didelphis virginiana NK NK P, D A, HD
Red fox, Vulpes vulpes NK PD? P, PC, E, H, D CD, LP, HD
European ferret, Mustela putorius NK NK P, E, H?, D? AB, HD?
House cat, Felis catus NK NK P, PC, E, H?, D HD
Domestic dog, Canis familiaris NK NK P, PC, E, H, D AB, LP, HD
Burro, Equus asinus S, W PD, E FC, D LC, HD
Horse, Equus caballus S, W PD, E FC, D LC, HD
Feral pig, Sus scrofa S, W PD, PR FC, D, IP AB, LC
Axis deer, Axis axis NK PD, PR H?, D? NK
Fallow deer, Dama dama NK PD, PR H?, D? NK
Mountain goat, Oreamos americanus S PD, PR, E FC, IP NK
Eastern gray squirrel, Sciurus carolinensis NK PD, PR FC, NC, AB, D SD
Fox squirrel, Sciuris niger NK PD, PR FC, NC, AB, D SD
House mouse, Mus musculus NK NK FC?, D HD, A, CF, SD
Norway rat, Rattus norvegicus NK NK FC?, D HD, A, CF, SD
Black rat, Rattus rattus NK PD? FC?, D HD, A, CF, SD
Nutria, Myocastor coypus S, W PD, PR FC?, D HD, SD
European rabbit, Oryctolagus cuniculus S PD, PR, E FC, NC?, D, IP? CD, LC, HD
Eastern cottontail, Sylvilagus floridanus NK PD, PR?, E? FC, NC?, D, IP CD, LC, HD
Bullfrog, Rana catesbeiana NK NK PC, P, E NK
Green frog, Rana clamitans NK NK PC, P?, E? NK
Snapping turtle, Chelydra serpentina NK NK PC, P, E?, AB, D AB
Red-eared slider turtle, Trachemys scripta NK PD? P?, PC?, FC?, D, AB NK
Plateau striped whiptail, Cnemidophorus velox NK NK PC? NK
A=aesthetics, AB=aggressive behavior, CD=crop damage, CF=contamination of foods, D=disease/parasites, E=species endangerment,
FC=forage competition, H=hybridization, HD=human/livestock/pet disease/parasites, IP=increase prey base, LC=livestock forage
competition, LP=livestock predation, NC=nest competition, NK=none known, P=predation, PC=prey competition, PD=plant damage,
PR=plant regeneration, S=soil erosion, SD=structural damage, W=water quality/quantity.
 434 Wildlife–Habitat Relationships in Oregon and Washington

Management of Introduced
Wildlife Species
It is better to prevent the introduction of an unwanted
species rather than deal with the management or
attempted eradication of the species once it becomes
established.36, 135 On the other hand, many introduced
wildlife species (upland birds, cottontail rabbits, nutria,
red fox) are managed as “game” species by state wildlife
agencies, using traditional methods of harvestable wildlife
management. Usually, a harvest license is required;
seasons, bag limits, methods of take, and other regulation
are set each year; and, in some cases, populations and
harvests are monitored.
There are many wildlife management methods
English (house) sparrow. (Photograph by Jeffrey C. Lewis) available to assist us in the management of “undesirable”
introduced wildlife species (Table 7). These physical,
mechanical, chemical, biological, and cultural methods are
used to reduce the carrying capacity of the area for the
species, to reduce population density, or to keep animals
out of certain areas. Reduction of populations by lethal
Certain introduced species have the potential for means may only provide a temporary “fix” unless habitats
substantial ecosystem disruption, the final and highly can be modified to reduce their carrying capacity for the
significant category in Table 2. Of the 42 wildlife species introduced species.149 On the other hand, commercial
already introduced into Oregon or Washington, we would exploitation or bounties on introduced wildlife has been
include feral livestock (pigs, horses, burros), mountain used in some situations as a way to keep population levels
goats (in areas where they are not native), nutria, and down while generating local income.22, 118 Pathogenic
European rabbits in this category. Other species (such as agents are rarely used to control vertebrate populations
the Old World rodents) cause substantial ecological because of the need for specificity, start-up costs and
disruption in tropical ecosystems, but are not as damaging potential hazards, although efforts continue in Australia.114
to temperate ecosystems. Major ecosystem disruption can Often a variety of methods are employed, as in an
occur when these species seriously impact the physical integrated pest management (IPM) approach.13, 17, 118, 165 If
environment (soil parameters, erosion, water quantity and a new species is released in the area, it is important to
quality), achieve relatively high densities, exert heavy restrict its spread as soon as possible. 9 Research is
grazing pressure, or successfully compete with native underway on chemosensory and reproductive inhibition
fauna. Mountain goat impacts on the Olympic Peninsula devices that may provide valuable tools in the future
and the difficulties of resolution have been described,18, 67 management of introduced species.48
and so have impacts and management of feral horse and Eradication of an introduced species is often the
burro.19, 41, 98 Feral pigs have been studied extensively management goal, but is difficult to achieve. 9, 24, 127
around the world because of their very significant impacts Nonetheless, eradication has been achieved in some
to ecosystems. 10, 98 The pros and cons of nutria places, especially on islands.127, 148, 165 An entirely different
introductions in the Pacific Northwest and elsewhere have philosophy is to “let nature take its course” and assume
also been discussed.89, 93, 98 Similar discussions for European that eventually introduced species will drop out on their
rabbits were presented as well.6, 28, 63, 98 Because of the own, will fit in satisfactorily, or will result in a worldwide
significant potential for ecological disruption by these homogenization of the planet’s flora and fauna. While this
species, there have been extensive efforts to eradicate them may be the ultimate fate of the global flora and fauna, we,
after introduction, especially from islands, and most are as resource managers and concerned citizens, should not
banned from import at the federal or state level. take such a defeatist attitude.88

Table 7. Examples of methods for the management of introduced wildlife species by category.

Cultural/Habitat Physical Chemical Biological Other

Crop selection Barriers Repellents Predators Bounties

Cover reduction Traps Toxicants Disease/Parasites Insurance
Water removal Electrocution Reproductive inhibition Resistant plants Harvest
Sanitation Flooding Aversive conditioning Lethal genes Acclimation
Buffer crops Shooting/Frightening Tranquilizers, other drugs Biosonics Acceptance
devices
 CHAPTER 16: INTRODUCED SPECIES 435

In most cases, an integrated management approach will subspecies, the Sichuan pheasant, that is better adapted
be required to control most introduced species, using a to wooded or shrubby habitats.138 Pheasants are currently
problem assessment, action plan, several methods, and widespread in Oregon and Washington. The natural
monitoring. Adequate surveillance and control at the point history of the pheasant in the United States and the Pacific
of origin are important. Additionally, adequate budgets, Northwest has been recounted.2, 99, 104, 138
public support, and access to private lands will be essential
Ecological Implications. Although the introduction of
to the successful management or eradication of most
pheasants and other upland game birds (see Table 3) has
introduced wildlife species. 9, 36, 165 Using introduced
largely been considered positive, they are not without
rodents as an example; Witmer et al.164, 165 discussed the
some adverse ecological effects. Concerns were expressed
many considerations of introduced species management
as early as 1923 in Washington that pheasants may damage
and eradication.
crops (sprouting corn, potatoes) and gardens.142 They
There are many socio-political, economic, and
compete for food and nest sites with native grouse species,
ecological issues associated with introduced species.88, 143
especially because they are more adaptable and tolerant
Realizing this, and involving the appropriate and
of disturbed landscapes and because they may be released
interested parties in the decision-making process, will be
in large numbers on a regular basis. 138, 150, 159 Their
essential to the successful resolution of current and future
aggressive behavior can displace other birds. Nest
wildlife introductions.
parasitism with blue grouse, ruffed grouse, and other
upland game bird species has been reported.83, 132, 142, 150, 158,
Case Histories 159

We conclude this chapter with several case histories of There is the potential for hybridization between
wildlife introductions in Oregon and Washington. As we pheasants, native grouse, and other upland game bird
have mentioned, some introductions can be considered species,8, 78 but the extent and seriousness of this effect is
“positive” while many are considered “negative” for a not known. The potential transfer of diseases, such as
number of reasons. In reality, most introductions have the avian tuberculosis, to other bird species and even humans,
potential for both positive and negative effects. Only time pets, and livestock has been noted.52
and our concerted efforts will determine the future status Management and Research Needs. In Oregon and
of native and introduced wildlife species and of Washington, pheasants are classified as upland game birds
ecosystems in Oregon and Washington. In many cases, and are managed with season lengths and bag limits to
we will continue to live with these naturalized species. regulate the number of harvested birds. During most
years, female pheasants have been protected from harvest
1. Ring-necked Pheasant
to increase recruitment. Wild bird populations have been
History, Distribution, and Status. Ring-necked pheasants
supplemented with pen-reared birds to increase harvest,
are native from the Caucasus Mountains of Eurasia
although the use of this strategy has greatly declined.
through Southeast Asia to Northern Japan where they are
There have often been state and federal efforts to
closely associated with river valleys, bamboo stands, and
encourage agricultural crop producers to manage their
agricultural lands. They have been widely introduced
lands for the benefit of pheasants and other upland game
throughout the world, primarily for upland game hunting,
species. The Conservation Reserve Program is an example
but also for viewing.99, 104 Substantial revenues have been
of one such program. Activities involve establishing or
generated for state wildlife agencies and for local
maintaining areas of woody or herbaceous vegetation; in
economies from upland game seasons. 73 The first
drier areas, water sources (“guzzlers”) may be provided.
successful introduction to the United States occurred in
It is likely that pheasants will be a less significant
1882 when Owen Denney, an Oregon attorney and judge,
element of the Oregon and Washington introduced
had 28 birds from China delivered to Portland.138 He began
avifauna in the future. There are many reasons for this,
breeding the birds, and they did so well that he was soon
including long-term declines in wild populations, the
shipping them to other parts of Oregon, into Washington
reduced emphasis on pen-rearing and release of birds,
in about 1883, and, eventually, to other states. The first
human encroachment on pheasant habitats, clean farming
pheasant hunting in the United States occurred in Oregon
practices, and an increased interest in improving
in 1891. In 1911, the State of Oregon opened the first large-
conditions for native upland game bird species. That being
scale, state-operated game bird farm.95 The facility in the
said, there is still substantial interest in this naturalized
Willamette Valley achieved peak production in 1950 when
member of the Pacific Northwest fauna, and a wish to
over 70,000 pheasants were reared and released. The State
assure that its regional presence will continue.
of Washington followed with the development of
extensive game bird farms.142 It has been estimated that 2. European Starling
about 100,000 pheasants were harvested in Washington History, Distribution, and Status. The European starling
in 1922,142 and that number tripled by 1950.95 More recently, is a palearctic species that originally ranged throughout
however, many of the rearing facilities have been shut Europe and east to Lake Baikal, Siberia, and the Middle
down because of increased costs, low survival rate of pen- East.99 It has since become naturalized, via numerous
reared birds, and other problems.32 Interestingly, this introductions, to most of North America, South Africa,
situation has resulted in attempts to introduce another Australia, and New Zealand.99 Starlings were purposefully
 436 Wildlife–Habitat Relationships in Oregon and Washington

and successfully introduced to Central Park, New York and roost availability; the use of frightening devices based
City, in 1890-91, although the Portland Songbird Club on chemicals, sounds, or objects; the use of repellents and
attempted an introduction in Portland, Oregon, in 1889 sticky substances; the use of toxicants; shooting; and
that failed. 77 The species range expansion in North trapping, with or without live bird decoys.80 Although
America is nothing less than amazing, reaching Mexico some of these methods have been moderately successful
in the 1940s and Alaska in the 1970s.16, 85 While numbers for a while, most are of limited effectiveness and must be
of starlings appear to have stabilized over much of North repeated on a regular and long-term basis. 16, 49 The
America, they are now one of the most numerous bird difficulty of dealing with starlings at high density roosts
species in North America.16 The species was introduced has been documented by Glahn et al.56 In all likelihood, a
for aesthetic purposes, to bring a little of the Old World to combination of methods and the alteration of crop and
the New World. Starlings first appeared in Oregon and livestock production practices would be most likely to
Washington in 1943.77 They are now abundant throughout provide damage reduction or population reduction.49
most of Oregon and Washington, especially in urban/ Starlings are here to stay and can be expected to
suburban and agricultural settings. The natural history of continue to impact some native bird species. We need to
the species has been described.16, 49, 104 better understand the interactions of starlings with food
sources, habitats, and other species and with the control
Ecological Implications. Few benefits have been
measures that we employ. We also need to develop more
attributed to the introduction of starlings in North
effective damage management methods; research is
America. Cabe16 noted, however, that much basic research
underway on avian repellents and on
on avian biology has been done using starlings. Starlings
immunocontraception. There has been some effort to
have also been attributed with high levels of insect
develop specific methods to reduce the ability of starlings
consumption, may be hunted, and may provide food for
to usurp nest sites from other species; Grabill57 attempted
humans in some situations.49 Finally, starlings occur in
to increase wood duck nesting success by placing starling
highly disturbed settings that might otherwise have few
nest boxes near wood duck nest boxes. He relied on the
birds present.
agonistic behavior of starlings during nesting to keep other
Feare49 reviewed the many adverse effects of starlings,
starlings from using the wood duck nest boxes. Fielder et
including plant damage; food and nest competition with
al.50 reduced starling use of wood duck nest boxes by
native bird species; disease and parasite transfer to
covering the opening from the end of the wood duck
wildlife, livestock, pets, and humans; fruit consumption
nesting season until just before the initiation of the next
and damage; livestock food consumption and
wood duck nesting season. Lumsden105 and McGilvrey
contamination; aircraft strikes; and aesthetic problems
and Uhler109 also presented designs to reduce nest box use
(droppings, odors, noise). Much of the concern about
by starlings. Knowledge of starling flocking behavior was
ecological effects of starlings seems to involve their highly
instrumental in the development of Avitrol, a chemical
competitive ability to usurp nest cavities (both natural and
frightening agent. A few birds are allowed to feed on
man-made) and thus contribute to the declines in
treated bait. They become sick, fly erratically, and give
populations of native cavity nesters. Adverse effects have
warning cries that frighten other starlings from the area.80
been noted for bluebirds, 39, 122 purple martins, 11 tree
Most starlings ingesting Avitrol will eventually die;
swallows, 122 northern flickers, 72 various species of
therefore, the chemical must be used carefully to minimize
woodpeckers,71, 84, 147, 156 and various cavity nesting duck
secondary poisoning hazards.66 Geis53 noted that starling
species.50, 57, 105 Not only is it difficult for these species to
and house sparrow numbers could be kept at lower
find and hold nest cavities in the presence of starlings,
densities by the careful design of urban structures;
but starlings may also parasitize the nests of other species
latticework on apartment buildings, for example, was very
by destroying eggs or hatchlings.50, 57, 122 Brush15 noted,
attractive to starlings and house sparrows. These examples
however, that significant cavity competition probably only
illustrate the value of a thorough knowledge of the biology
occurs where natural cavities are very limited.
and ecology, including behavior, of a species to assist in
Economic losses and damage to planted crops (corn,
resolving conflicts.
winter wheat), fruits (grapes, peaches, blueberries,
strawberries, figs, apples, and cherries) and livestock 3. Nutria
feedlots have been described,49, 80 as well as the disease History, Distribution, and Status. The nutria is a large
problems caused by starlings.49, 52, 80 Constantin and Floyd26 semi-aquatic rodent native to South America, which has
discussed the hazards of starlings and other birds at been introduced into a number of areas in North America
airports. Typically, starling problems are quite localized. since the 1930s. Oregon and Washington are among 15 or
Management and Research Needs. The manage-ment of more states with feral nutria populations.161 Nutrias were
starlings is problematic at best because of their exceptional first brought to the Northwest in the 1930s in the
ability to exploit human-altered landscapes. 49 Many expectation that nutria farming would become a lucrative
methods are used to reduce starling numbers, the damage endeavor.61, 89, 94 Inflated breeding stock prices, poor
they cause, or to disperse aggregations. These include reproduction, large farming expenses, and little economic
attempts at exclusion from buildings, ledges, and trees; return for nutria pelts (~$1.00 per pelt) resulted in the
cultural and habitat modifications that reduce food, water, collapse of an industry whose boom was short-lived.47, 87,
 CHAPTER 16: INTRODUCED SPECIES 437

89, 161
More than 600 nutria farms existed in Oregon from in damage to agriculture, drainage systems, earthen
the 1930s to the 1950s,89 and a number of farms existed in structures (dikes, levees, embankments), and vegetative
Washington at this time.61, 93 Flooding and storms damaged communities. 47, 89, 96 Burrowing can disintegrate and
holding structures and allowed some nutrias to escape weaken these structures, and may cause them to fail.96, 161
from fur farms, however, farmers often released their stock Ironically, nutrias were introduced in some areas to help
when farming became uneconomical. By the 1940s, nutrias control marsh vegetation.47 Kuhn and Peloquin89 reported
had been captured by trappers or collected on both sides nutria damage to agricultural crops in the Willamette
of the Cascades in Oregon and Washington, but most Valley and estimated losses of thousands of dollars per
nutrias were found in the Puget Sound area, the Willamette year. Humans, livestock and pets are vulnerable to a
Valley, along coastal Oregon rivers, and along the number of diseases and parasites carried by nutrias,
Columbia River.70, 79, 89, 93, 108 Only the Yakima River drainage including equine encephalomyelitis, leptospirosis,
in southcentral Washington supports substantial numbers hemorragic septicemia, paratyphoid, salmonellosis,
east of the Cascade Mountains. giardiasis, tapeworms, and liver flukes.96, 161 The aggressive
The nutria is an unclassified wildlife species in Oregon behavior of nutria also poses a hazard to pets that
and Washington, and it can be harvested in unlimited approach them too closely ( J. Tabor, Washington
numbers at any time of the year. The records indicate Department of Fish and Wildlife, pers. comm.).
fluctuating harvest levels of nutrias, which may reflect Personnel with the USDA Wildlife Services and state
fluctuating pelt prices152 rather than fluctuating population wildlife officers respond to nutria damage complaints.
densities. Nutria harvest data also indicate a relatively Although a number of damage prevention and control
stable population, in that nutrias are consistently captured methods exist for nutrias,96 commercial trapping appears
in the same counties (i.e., nutrias do not appear to be to be the most common method used in Oregon and
spreading to previously unoccupied counties in Washington. Some trappers have certainly benefited from
appreciable numbers). Short-term stability, however, does the introduction of nutrias, although the monetary benefits
not necessarily mean that all habitats suitable for nutrias appear limited as nutria pelts are not highly valued for
have been colonized or that a range expansion will not fur.152 Low pelt prices offer little incentive to most trappers
occur in the future. The natural history of the nutria has and consequently, commercial trapping may be limited
been described in detail.47, 87, 161 as a management tool for nutria populations. Conversely,
control of pest nutria can be a source of income for some
Ecological Implications. Through foraging, nutrias can
trappers and pest control professionals.
denude expanses of vegetation, eliminating vegetative
structure.87, 157 While nutrias are generally opportunistic Management and Research Needs. Trapping and
vegetarians, Wentz157 found that broadleaf arrowhead localized control efforts have been used to manage nutria
(Saggittaria latifolia) and smartweed (Polygonum spp.) were populations since they were first introduced, and these
selected by nutrias in Oregon, and these plants may be techniques will likely continue to provide for nutria
locally reduced or extirpated by foraging nutrias. Nutrias management in the future. Trapping records indicate a
construct resting and feeding platforms of compacted relative stable nutria population in the Pacific Northwest.
vegetation in wet areas, form trails between these Until new information indicates that nutria impacts are
platforms through vegetation, and also create grooming particularly severe to certain species, ecological
areas, dens, and runs or slides at the water’s edge.47, 87 communities, or geographic areas, it is unlikely that
These activities can significantly impact vegetative current management methods will be altered or replaced.
communities.87, 157 The clearing of vegetation by nutrias Lobbying efforts to ban trapping or outcries for nutria
may alter plant succession, and convert marsh ecosystems eradication could alter the status quo, but these do not
to more open-water environments. appear to be immediate issues in Oregon or Washington.
In Louisiana, increasing nutria harvests in the mid- With the exception of research by Peloquin121 on growth
1900s coincided with decreasing muskrat harvests.87 The and reproduction and Wentz157 on nutria density and
apparent decline in the muskrat population could have impacts to marsh vegetation, little study of the nutria has
been the result of many factors, but the nutria irruption been conducted in the Pacific Northwest, and none has
was considered among the most significant. Apparent been published from Washington. Future research should
declines in muskrat numbers have also been observed in focus on how the nutria’s alteration of aquatic
areas where nutrias are abundant on the Finley National environments and its physical presence (i.e., potential
Wildlife Refuge in western Oregon (H. Brunkal, U. S. Fish competition and disease transmission) could impact
and Wildlife Service, pers. comm.). Alteration of the sensitive fauna and vegetative communities. This research
vegetative community would be expected to have a may also prompt study into alternative management
significant influence on native fauna, especially sensitive techniques for nutrias.
amphibians and species that have niches similar to the
nutria (e.g., muskrat, some waterfowl). Unfortunately, 4. Red Fox
little information is available on the direct or indirect History, Distribution, and Status. Native to North
impacts of nutrias on other fauna. America, Europe, Asia, and northern Africa,103 the red fox
Nutrias cause direct and indirect impacts to humans has the largest geographic range of any terrestrial
by their foraging and burrowing activities, which result carnivore with the possible exception of the domestic cat.
 438 Wildlife–Habitat Relationships in Oregon and Washington

European red foxes were introduced to the East Coast in incidentally with other foods. Foxes may be important
the 1600s and 1700s23, 126 and Australia in the mid 1800s106 seed dispersers of both native and non-native plants. Den-
for fox hunting; actions that confused the taxonomy of site excavation and other digging could minimally disrupt
red foxes in eastern North America and greatly expanded flora, but would also expose a medium for seed
the range of the red fox. Non-native red foxes were brought germination.
to Oregon and Washington in the early 1900s for fur Red foxes commonly prey on insects, earthworms,
farming40 and fox hunting or trapping.4, 108, 160 In the 1910s, small- to medium-sized birds and mammals, and
the fur industry was rapidly spreading west across the herpetofauna; predation on crustaceans and fish and the
continent, when choice breeding stock and pelts from red use of carrion has also been documented.64, 100, 103 Red foxes
foxes (predominantly the silver phase foxes) were sold are noted predators of species valued by humans as pets,64
for thousands of dollars.40 By 1915, the first fox farms were livestock, 58, 115 game birds and mammals, 45, 160 and
established in Oregon and Washington. 40 Many endangered species.164 Conservation of the snowy plover
introductions occurred when foxes were released or in Oregon and Washington could be hindered by red foxes
escaped from farms3, 4, 40 or evaded hounds and hunters, should they become established near nesting colonies
forming free-roaming populations of non-native red foxes along the coast, as has happened in California. Interference
in both states. More recent introductions of red foxes in with the reproductive behavior of native fauna, especially
North America have included the release of pet foxes, the ground-nesting birds, can be significant.166 Aubry4, 5
illegal release of farm foxes by animal rights activists, and suggested that non-native red foxes might not be
the translocation of non-native foxes into previously physiologically or behaviorally capable of surviving in
unoccupied areas by pet owners, wildlife rehabilitators, high-elevation habitats. However, an introduction of non-
and animal control officials.102 native red foxes within the historical ranges of native red
Non-native red foxes occur throughout many of the foxes could result in resource competition, interbreeding
lowland areas in western Oregon and Washington4, 5 and and disease transmission. 5, 101 It is unknown if
in several disjunct populations in eastern Oregon and interbreeding with non-native foxes would reduce the
Washington. The distribution of non-native red foxes has fitness of native red fox populations. The transmission of
largely been determined as those areas where red foxes diseases, including sarcoptic mange, rabies, canine
occur outside the historical ranges of native Cascade (V. distemper, parvovirus, and leptospirosis, is a threat that
v. cascadensis) and Rocky Mountain red foxes (V. v. red foxes pose to other mammals.103 Additionally, resource
macroura).4, 5, 7, 62 The Cascade red fox historically occurred competition,81 disease transmission, and interbreeding145
in the high-elevation meadows and parklands of the would be expected to negatively affect native kit foxes
Cascade Range, whereas the Rocky Mountain red fox should red foxes become established in southeastern
occupied similar habitats in northeastern Washington, and Oregon.
in the Blue and Wallowa Mountains. 7, 37, 62 Because Foxes may negatively impact humans in several ways,
relatively little information is available on the locations including livestock depredation, crop damage, disease
and operation of fox farms, especially in Oregon, and transmission to humans and their pets, and predation or
because there are no known means of visually injury of pets. More indirectly, non-native red foxes may
distinguishing native from non-native red foxes, it is not negatively impact humans by affecting species valued by
known if introductions have occurred within the ranges the public (e.g., game or protected species). Positive
of the native red foxes. impacts to the public include recreational and economical
Red foxes are considered a furbearing species in Oregon opportunities of trapping, hunting, and fur farming, and
and a furbearing game animal in Washington with no the recreational opportunities of feeding and watching
administrative distinctions made between native and non- wild foxes, along with the enjoyment from having foxes
native red foxes. Red foxes can be trapped in most areas as pets; the latter is strongly discouraged by wildlife
of Oregon during a regulated season. In Washington, red professionals.
foxes can be trapped during a regulated season except in
Management and Research Needs. Relatively little is
Whatcom, Skagit, and Island Counties, and a portion of
known about the populations of non-native red foxes in
Cowlitz County. Because market prices strongly influence
Oregon and Washington. Non-native red foxes provide
the harvest of most furbearer species, harvest data are not
additional harvest opportunities for trappers, and
a good indication of fox population trends. While little
management in this regard comprises season restrictions
information is available on population trends, there are
and harvest regulations. Management has also involved
no indications that fox populations in either state are
communication with the public about occasional livestock
increasing or decreasing dramatically. The natural history
kills and concerns about fox predation on domestic cats.
of the red fox has been described in detail.103, 106, 141, 152, 153
Damage prevention and control methods for red foxes
Ecological Implications. With the exception of excavating were reviewed by Phillips and Schmidt.123 The wide
holes for dens and prey items, and leaving some uneaten distribution of red foxes in western Oregon and
prey remains scattered about, red foxes probably have little Washington reflects the potential for disease transmission
effect on their physical environment. Non-native red foxes to pets, livestock, other wildlife, and humans, with rabies
feed on a variety of fruit-producing plants.100 They also being of particular concern. Research on non-native red
eat leafy vegetation, some of which may be ingested foxes to determine distribution and densities, identify
 CHAPTER 16: INTRODUCED SPECIES 439

disease prevalence, and characterize food habits is needed a substantial contribution to nutrient cycling in aquatic
to better understand populations and potential impacts ecosystems.
in Oregon and Washington. Much of the literature on the bullfrog in western North
America has been concerned with the effect of bullfrog
5. Bullfrog predation on native fauna, especially other ranids.42, 65, 86,
History, Distribution, and Status. Bullfrogs are native to 111
Kupferberg90 demonstrated that bullfrogs negatively
North America east of the Rocky Mountains; however, affected the growth of developing yellow-legged frogs by
their range has greatly expanded due to introductions by outcompeting them for food resources. Although many
humans in western North America, South America, reports have implicated the bullfrog as a major cause of
Europe, and Asia.14 Bullfrogs were first introduced into declines in some native species, Hayes and Jennings65
the Northwest in the early 1900s to provide food (i.e., frog argued that this has not been clearly proven and a number
legs), opportunities for frog hunting, and stock for frog of other factors may be at work, such as predation by
farms.92, 140 Being the largest frog in North America, introduced fish, habitat alteration, commercial
bullfrog legs were a highly prized food, and frog farming exploitation, and the effects of toxicants. However, until
to supply the demand for bullfrog legs was undertaken proven otherwise, it may be wise to consider bullfrog
but rarely succeeded.14 Lampman92 states that bullfrogs predation and competition as detrimental to a number of
from Idaho were first brought to eastern Oregon in 1914, vulnerable, sensitive, or listed species in Oregon and
and subsequent introductions in western Oregon in 1921 Washington, including the Oregon spotted frog, leopard
involved releasing additional Idaho bullfrogs and frog, red-legged frog, foothill yellow-legged frog, and
bullfrogs from the previously introduced populations in western pond turtle. Bullfrog predation on hatchling
eastern Oregon. Nussbaum et al.113 reported the release of western pond turtles has prompted management efforts
18 bullfrogs in the Grant’s Pass area in 1931 and that to protect remaining populations in Oregon and
bullfrogs were soon well established in the upper Rogue Washington. Management efforts involve collecting
River Valley. In Washington, Dvornich et al.43 reported the hatchling turtles from western pond turtle nests and
first specimens collected in the 1930s, which suggests that placing them in captivity until they are too large to be
the first successful introductions occurred in the 1920s and eaten by bullfrogs (R. Goggans, Oregon Department of
early 1930s. Fish and Wildlife, and K. Slavens, Washington Department
Bullfrogs are largely aquatic and occur in lower- of Fish and Wildlife, pers. comm.). These efforts, which
elevation freshwater habitats on both sides of the Cascades started in the early 1990s in Oregon and Washington, have
in Oregon and Washington and along much of the been successful at recruiting young turtles into resident
Columbia River.97 Within this range, the bullfrog has populations, and some female recruits from the first “head-
become widely established and locally abundant because started” cohorts are expected to be large enough to breed
it is a capable colonizer of a wide variety of habitats and a in 1999.
prolific breeder.14 They are classified as a game fish in Bullfrogs are beneficial to some people as a source of
Oregon and as a game species in Washington; however, food, sport, and economic gain. Universities and schools
these classifications may soon change in both states. have created a significant demand for bullfrogs for use in
Fishing or hunting license (in Oregon and Washington, classroom and laboratory study. Others simply enjoy the
respectively) is required to harvest bullfrogs and there are sound of bullfrogs or stock them for the pleasure of having
season restrictions, but no bag limits. Bullfrogs may still bullfrogs on their property; the latter is strongly
be expanding their range as suitable habitats are colonized discouraged by wildlife professionals.
by invading or introduced individuals. Humans continue Bullfrog predation on and competition with native
to introduce bullfrogs into new, previously unoccupied species are impacts that cause concern among many
areas. Water-garden and pond stores in Vancouver and people. This concern may prompt a modification of the
Portland recently sold bullfrog tadpoles from California legal status of the bullfrog to allow for more effective
and North Carolina (at $3.00 per tadpole) to individuals bullfrog management and protection of native species
interested in stocking their ponds with bullfrogs (selling vulnerable to bullfrog predation and competition.
or possessing live bullfrogs is illegal in Washington
without a permit). Also, some summer festivals include a Management and Research Needs. The predation and
frog-jumping contest where captured frogs (often competition threats posed by bullfrogs to native species
bullfrogs) may be released after the contest into previously have prompted consideration of bullfrog eradication in
unoccupied habitats. The natural history of bullfrogs has some localized areas. Removing egg masses, killing adults,
been described in detail.14, 113 or promoting harvests of bullfrogs may act to reduce their
impacts on native species. However, their local abundance,
Ecological Implications. Both tadpoles and adults feed widespread distribution, ability to disperse and recolonize
on vegetation,14, 113 although plant consumption by adults habitats, and the tendency for people to transplant
is likely the result of incidental ingestion while capturing bullfrogs, makes eradication difficult at best.110 Perpetual
a prey item. Consumption of vegetation by bullfrog bullfrog control may be required where management is
tadpoles could constitute an impact on local flora and a important to protect or restore native species. Research
significant indirect impact on native species that use this efforts that determine the degree to which bullfrogs
flora for food or cover,90 but this consumption could make threaten native species, relative to other causes of species
 440 Wildlife–Habitat Relationships in Oregon and Washington

decline, will help us focus our management actions on 7. Bailey,V. 1936. The mammals and life zones of Oregon. Bureau of
the most critical problems. Research into ways of Biological Survey. North American Fauna No. 55. U.S. Department of
controlling or eradicating bullfrogs without harming Agriculture,Washington, D.C.
8. Blackburn, D. 1977. An apparent ring-necked pheasant X blue grouse
native species would also be valuable.110 Studies that focus hybrid. Murrelet 58:78.
on single species or communities that may be impacted 9. Bomford, M. and P. O’Brien. 1995. Eradication or control for vertebrate
by bullfrog presence should also focus on obtaining data pests. Wildlife Society Bulletin 23:249-255.
to credibly address the impacts of bullfrogs. 10. Bratton, S. P. 1975. The effect of the European wild boar, Sus scrofa, on
gray beech forest in the Great Smoky Mountains. Thesis, Cornell
University, Ithaca, New York, NY.
Conclusions 11. Brown, C. R. 1981. The impact of Starlings on Purple Martin
At least 42 introduced species of wildlife (birds, mammals, populations in undamaged colonies. American Birds 35: 266-268.
amphibians, reptiles) occur in Oregon and Washington. 12. Brown, E. R., editor. 1985. Management of wildlife and fish habitats in
Introductions have occurred for many reasons, both forests of western Oregon and Washington. U.S. Department of
accidental and purposeful. Some have greatly contributed Agriculture, Forest Service, Pacific Northwest Region. Portland, OR.
13. Bruggers, R. L., E. Rodriguez, and M. Zaccagnini. 1998. Planning for bird
to outdoor recreation, local economies, and state wildlife
pest problem resolution: a case study. International Biodeterioration
agency revenues, while others have had adverse ecological and Biodegradation 42:173-184.
consequences through direct or indirect mechanisms (e.g., 14. Bury, R. B., and J. A. Whelan. 1984. Ecology and management of the
resource competition, displacement, predation, bullfrog. U.S. Fish and Wildlife Service. Resource Publ. No. 155.
hybridization, and disease transmission). Economic losses Washington, D.C.
to human-valued resources and public health hazards 15. Brush, T. 1983. Cavity use by secondary cavity-nesting birds and
have been documented. There has been more stringent responses to manipulations. Condor 85:461-466.
16. Cabe, P. R. 1993. European Starling. The Birds of North America 48: 1-
regulation of introduced species at the federal and state
24.
levels in recent years, in part due to the increased concern 17. Campbell, E. W., G. Rodda, T. Fritts, and R. Bruggers. 1999. An
about potential harm to native flora and fauna. Options integrated management plan for the brown tree snake on Pacific
for resource managers include prevention of entry, islands. Pages 423-433 in G. Rodda,Y. Sawai, D. Chiszar, and H. Tanaka,
eradication, and management after dispersal and editors. Problem snake management. Comstock, Ithaca, New York, NY.
establishment; the first is perhaps the most practical, while 18. Carlquist, B. 1990. An effective management plan for the exotic
mountain goats in Olympic National Park. Natural Areas Journal 10:12-
the latter is the most commonly employed option.
18.
Eradication is difficult and expensive in most situations. 19. Carothers, S. W., M. Stitt, and R. Johnson. 1976. Feral asses on public
More surveillance and control at the point of origin is lands: an analysis of biotic impact, legal considerations and management
needed. A sustained effort, using a variety of methods and alternatives. Proceedings of North American Wildlife Conference
the principles of integrated pest management, is needed 41:396-406.
to limit adverse effects. Each situation must be assessed 20. Cassidy, K. M., C. Grue, M. Smith, and K. Dvornich, editors. 1997.
on a species- and site-specific basis. New methods are Washington State Gap Analysis—Final Report. Volumes 1-5. Washington
Cooperative Fish and Wildlife Research Unit, University of Washington,
needed to improve the monitoring and management of
Seattle, WA.
introduced wildlife. 21. Center for Wildlife Law. 1996. Saving biodiversity: a status report on
state laws, policies and programs. Defenders of Wildlife, Albuquerque,
Acknowledgments New Mexico.
We would like to acknowledge the many people that 22. Choquenot, D., P. O’Brien, and J. Hone. 1995. Commercial use of
provided information or review comments on a portion pests: can it contribute to conservation objectives? Pages 251-258 in G.
or all of this chapter: Keith Aubry, Charles Bruce, Richard C. Grigg, P. Hale, and D. Lunney, editors. Conservation through
Bruggers, Alan Clark, Bruce Coblentz, Larry Cooper, John sustainable use of wildlife. Centre for Conservation Biology, The
University of Queensland, Brisbane, Australia.
Crawford, Richard Dolbeer, Joe Engles, James Glahn, 23. Churcher, C. S. 1959. The specific status of the New World red fox.
Rebecca Goggans, Marc Hayes, David Johnson, Kelly Journal of Mammalogy 40:513-520.
McAllister, Charles Meslow, Thomas O’Neil, Michael 24. Coblentz, B. E. 1990. Exotic orgainisms: a dilemma for conservation
Pipas, and Kate Slavens. biology. Conservation Biology 4:261-265.
25. Coleman, J. S., S. Temple, and S. Craven. 1997. Cats and wildlife: a
Literature Cited conservation dilemma. University of Wisconsin Cooperative Extension
1. Ad Hoc Federal Invasive Alien Species Task Force. 1997. Campaign Service, Madison, WI.
against invasive alien species: an action plan for the nation. Washington, 26. Constantin, B., and J. Floyd. 1992. An avian/airport study for Standiford
D.C. Airport, Louisville, Kentucky: results and management implications.
2. Allen, D. L. 1956. Pheasants of North America. Stackpole Books, Proceedings Eastern Wildlife Damage Control Conference 5:166-170.
Harrisburg, PA. 27. Corkran, C. C., and C. Thoms. 1996. Amphibians of Oregon,
3. Ashbrook, F. G. 1923. Silver fox farming. U. S. Dept. Agric. Bull. No. 1151. Washington, and British Columbia. Lone Pine, Renton, WA.
Washington, D.C. 28. Couch, L. K. 1929. Introduced European rabbits in the San Juan
4. Aubry, K. B. 1983. The Cascade red fox: distribution, morphology, Islands, Washington. Journal of Mammalogy 10:334-336.
zoogeography and ecology. Ph.D. Dissertation. University of 29. Coulter, M. W. 1957. Predation by snapping turtles upon aquatic birds
Washington, Seattle,WA. in Maine marshes. Journal of Wildlife Management 21:17-21.
5. ———. 1984. The recent history and present distribution of the red 30. Courtenay, Jr., W. R., and C. Kohler. 1986. Exotic fishes in North
fox in Washington. Northwest Science 58:69-79. American fisheries management. Pages 401-413 in Richard H. Stroud,
6. ———, and S. D. West. 1984. The status of native and introduced editor. Proceedings of The Role of Fish Culture in Fisheries
mammals on Destruction Island, Washington. The Murrelet 65:80-83. Management at Lake Ozark, Missouri. American Fisheries Society,
Bethesda, MD.
 CHAPTER 16: INTRODUCED SPECIES 441

31. Cox, C. B., I. N. Healy, and P. D. Moore. 1976. Biogeography: an 54. Gilligan, J., editor. 1994. Birds of Oregon. Cinclus, McMinnville, OR.
ecological and evolutionary approach. Second edition. Blackwell 55. Glahn, J. F., and D. L. Otis. 1986. Factors influencing blackbird and
Scientific, Oxford, England. European starling damage at livestock feeding operations. Journal of
32. Crawford, J. A., and S. Meyers. 1980. Survival of ring-necked pheasant Wildlife Management 50:15-19.
hens in the Willamette Valley, Oregon. Unpublished report. Oregon 56. ———, A. R. Stickley, Jr., J. F. Heisterberg, and D. F. Mott. 1991. Impact
Department of Fish and Wildlife, Portland, OR. of roost control on local urban and agricultural blackbird problems.
33. Crawley, M. J. 1989. Chance and timing in biological invasions. Pages Wildlife Society Bulletin 19:511-522.
407-423 in J. A. Drake, editor. Biological invasions: a global perspective. 57. Grabill, B. A., 1977. Reducing starling use of wood-duck boxes. Wildlife
John Wiley & Sons, New York, NY. Society Bulletin 5:69-70.
34. Csuti, B., A. J. Kimerling, T. A. O’Neil, M. M. Shaughnessy, E. P. Gaines, 58. Graf, W. 1947. Mouse populations in relation to predation by foxes
and M. M. P. Huso. 1997. Atlas of Oregon wildlife: distribution, habitat, and hawks. The Murrelet 28:18-21.
and natural history. Oregon State University Press, Corvallis, OR. 59. Griffith, B., J. M. Scott, J. W. Carpenter, and C. Reed. 1989. Translocation
35. Dahlgren, R. B. 1987. The Ring-Necked Pheasant. Pages 305-311 in as a species conservation tool: status and strategy. Science 245:477-
Harmon Kallman, editor. Restoring America’s wildlife- 1937-1987: the 480.
first 50 years of the Federal Aid in Wildlife Restoration (Pittman- 60. Guenther, K., and T. Kucera. 1978. Wildlife of the Pacific Northwest:
Robertson) Act. U. S. Department of the Interior, Fish and Wildlife occurrence and distribution by habitat, BLM district, and National
Service, Washington D.C. Forest. U.S. Department of Agriculture, Forest Service, Portland, OR.
36. Dahlsten, D. L. 1986. Control of invaders. Pages 275-302 in Harold A. 61. Guenther, S. E. 1950. Nutria. Washington State Game Department,
Mooney, editor. Ecology of biological invasions of North America and Olympia, Washington, USA. Game Bulletin 2:5.
Hawaii. Springer-Verlag, New York, NY. 62. Hall, E. R. 1981. Mammals of North America. J. Wiley and Sons, New
37. Dalquest, W. W. 1948. Mammals of Washington. University of Kansas York, NY.
Publ. Natural History No. 2. University of Kansas Press, Lawrence, KS.p 63. Hall, L. 1977. Feral rabbits on San Juan Island, Washington. Northwest
38. Davis, J. W., L. Karstad, and D. Trainer. 1981. Infectious diseases of wild Science 51:293-297.
mammals. Iowa State University Press, Ames, IA. 64. Harris, S. 1981. On the food of suburban foxes (Vulpes vulpes), with
39. Davis, W. H., and W. C. McComb, 1989. Bluebirds and Starlings: special reference to London. Mammal Review 11:151-168.
competition for nest sites. Sialia 11:124-126,138. 65. Hayes, M. P., and M. R. Jennings. 1986. Decline of ranid frog species in
40. Dearborn, N. 1915. Silver fox farming in eastern North America. western North America: are bullfrogs responsible? Journal of
Bulletin No. 301. U. S. Department of Agriculture. Washington, D.C. Herpetology 20:490-509.
41. Douglas, C. L., and D. Leslie, Jr. 1996. Feral animals on rangelands. 66. Holler, N. R., and E. W. Schafer, Jr. 1982. Potential secondary hazards of
Pages 281-292 in P. Krausman, editor. Rangeland wildlife. Society for avitrol baits to sharp-shinned hawks and American kestrels. Journal of
Range Management. Denver, CO. Wildlife Management 46:457-462.
42. Dumas, P. C. 1966. Studies of the Rana species complex in the Pacific 67. Houston, D. B., E. G. Schreiner, and B. B. Moorhead. 1994. Mountain
Northwest. Copeia 1966:60-74. goats in Olympic National Park: biology and management of an
43. Dvornich, K. M., K. R. McAllister, and K. B. Aubry. 1997. Amphibians introduced species. Scientific Monograph NPS/NROLYM/NRSM-94/25.
and reptiles of Washington State: location data and predicted U.S. Department of Interior, National Park Service, Washington, D.C..
distributions.Volume 2, Washington State Gap Analysis, Final Report. 68. Human, K. G., and D. M. Gordon, 1997. Effects of Argentine ants on
Washington Cooperative Fish and Wildlife Research Unit, University of vertebrate biodiversity in Northern California. Conservation Biology
Washington, Seattle, WA. 11:1242-1248.
44. Ehrlich, P. R., 1989. Attributes of invaders and the invading process: 69. Hygnstrom, S. E., R. M. Timm, and G. E. Larson. 1994. Prevention and
vertebrates. Pages 315-328 in J. A. Drake, editor. Biological invasions: a control of wildlife damage. University of Nebraska Cooperative
global perspective. John Wiley & Sons, New York, NY. Extension, Lincoln, NE.
45. Epstein, M. B., G. A. Feldhamer, and R. L. Joyner. 1983. Predation on 70. Ingles, L. G. 1965. Mammals of the Pacific States: California, Oregon,
white-tailed deer fawns by bobcats, foxes, and alligators: predator Washington. Stanford University Press, Stanford, CA.
assessment. Proceedings of the Annual Conference of the Southeastern 71. Ingold, D. J. 1994. Influence of nest-site competition between
Association of Fish and Wildlife Agencies 37:161-172 European starlings and woodpeckers. Wilson Bulletin 106: 227-241.
46. Evanich, J. 1986. Introduced birds of Oregon. Oregon Birds 12:156- 72. ———. 1996. Delayed nesting decreases reproductive success in
186. northern flickers: implications for competition with European starlings.
47. Evans, J. 1970. About nutria and their control. Resource Publ. No. 86. Journal of Field Ornithology 67:321-326.
U. S. Department of Interior, Fish and Wildlife Service, Washington, D.C. 73. International Association of Fish and Wildlife Agencies. 1997. The
48. Fall, M. W., and W. B. Jackson. 1998. A new era of vertebrate pest economic importance of hunting: economic data on hunting
control? International Biodeterioration and Biodegradation 42:85-91. throughout the entire United States. International Association of Fish
49. Feare, C. J. 1984. The starling. Oxford University Press, New York, NY. and Wildlife Agencies, Washington, D.C.
50. Fielder, P. C., B. G. Keesee, and P. A. Popushinsky. 1990. Wood duck use 74. Jenkins, P. T. 1996. Free trade and exotic species introductions.
of nesting structures in central Washington. Pages 265-267 in L. H. Conservation Biology 10:300-302.
Fredrickson, G. V. Burger, S. P. Havera, D. A. Graber, R. E. Kirby, and T. S. 75. Jewett, S. G., W. Taylor, W. Shaw, and J. Aldrich. 1953. Birds of
Taylor, editors. Proceedings of North American Wood Duck Washington State. University of Washington Press, Seattle, WA.
Symposium, St. Louis, MI. 76. Jobanek, G. A. 1987. Bringing the old world to the new: the
51. Fox, M. D., and B. J. Fox. 1986. The susceptibility of natural introduction of foreign songbirds into Oregon. Oregon Birds 13:59-74.
communities to invasion. Pages 57-66 in R. H. Groves and J. J. Burdon, 77. ——— 1993. The European starling in Oregon. Oregon Birds 19:93-
editors. Ecology of biological invasions. Cambridge University Press, 96.
Cambridge, England. 78. Johnsgard, P. A. 1973. Grouse and quail of North America. University
52. Friend, M., editor. 1987. Field guide to wildlife diseases. Resource of Nebraska Press, Lincoln, NE.
Publication No. 167. U. S. Department of the Interior, Fish and Wildlife 79. Johnson, R. E., and K. M. Cassidy. 1997. Terrestrial mammals of
Service,Washington, D.C. Washington State: location data and predicted distributions. Volume 3,
53. Geis, A. D. 1973. Effects of urbanization and type of urban Washington State Gap Analysis- Final Report. Washington Cooperative
development on bird populations. Pages 97-105 in J. H. Noyes and D. R. Fish and Wildlife Research Unit, University of Washington, Seattle, WA.
Progulske, editors. Proceedings of a symposium on wildlife in an
urbanizing environment. University of Massachusetts, Amherst, MA.
 442 Wildlife–Habitat Relationships in Oregon and Washington

80. Johnson, R. J., and J. F. Glahn. 1994. European starlings. Pages E109- 105. Lumsden, H. G. 1986. Choice of nest boxes by tree swallows, house
E120 in S. Hygnstrom, R. Timm, and G. Larsen, editors. Prevention and wrens, eastern bluebirds, and European starlings. Canadian Field-
control of wildlife damage. University of Nebraska Cooperative Naturalist 100:343-349.
Extension Service, Lincoln, NE. 106. Macdonald, D. W. 1987. Running with the fox. Facts On File, New
81. Johnson, W. E., T. K. Fuller, and W. L. Franklin. 1996. Sympatry in canids: York, NY.
a review and assessment. Pages 189-218 in J. L. Gittleman, ed. Carnivore 107. MacDonald, I. A.W., L. Loope, M. Usher, and O. Hamann. 1989.
behavior, ecology, and evolution.Volume 2. Cornell University Press, Wildlife conservation and the invasion of nature reserves by
Ithaca, NY. introduced species: a global perspective. Pages 215-255 in J.A. Drake,
82. Kauhala, K. 1996. Introduced carnivores in Europe with special editor. Scope 37: Biological invasions, a global perspective. John Wiley &
reference to central and northern Europe. Wildlife Biology 2:197-204. Sons, New York, NY.
83. Kebbe, C. E. 1958. Deposition of pheasant eggs in ruffed grouse nest. 108. Mace, R. U. 1970. Oregon’s furbearing animals. Oregon State Game
The Murrelet 39:10. Commission Wildlife Bulletin Number 6. Portland, OR.
84. Kerpez, T. A., and N. S. Smith. 1990. Competition between European 109. McGilvrey, F. B., and F. M. Uhler. 1971. A starling-deterrent wood duck
starlings and native woodpeckers for nest cavities in Saguaros. Auk 107: nest box. Journal of Wildlife Management 35:793-797.
367-375. 110. Moler, P. E. 1994. Frogs and toads. Pages F-9 - F-11 in S. E. Hygnstrom,
85. Kessel, B. 1953. Distribution and migration of the European starling in R. M. Timm, and G. E. Larsen, editors. Prevention and control of wildlife
North America. Condor 55:49-67. damage. University of Nebraska Cooperative Extension Service,
86. Kiesecker, J. M., and A. R. Blaustein. 1998. Effects of introduced Lincoln, NE.
bullfrogs and smallmouth bass on microhabitat use, growth, and survival 111. Moyle, P. B. 1973. Effects of introduced Bullfrogs, Rana catesbeiana, on
of native red-legged frogs. Conservation Biology 12:776-787. the native frogs of the San Joaquin Valley, CA. Copeia 1973:18-22.
87. Kinler, N. W., G. Linscombe, and P. R. Ramsey. 1987. Nutria. Pages 326- 112. Newsome, A. E., and I. R. Noble. 1986. Ecological and physiological
343 in M. Nowak, J. A. Baker, M. E. Obbard, and B. Malloch, edsitors. Wild characters of invading species. In: R. H. Groves and J. J. Burdon, editors.
furbearer management and conservation in North America. Ontario Ecology of biological invasions. Cambridge University Press, Cambridge,
Ministry of Natural Resources, Toronto, Ontario, Canada. England.
88. Knopf, F. L. 1992. Faunal mixing, faunal integrity, and the biopolitical 113. Nussbaum, R. A., E. D. Brodie, Jr., and R. M. Storm. 1983. Amphibians
template for diversity conservation. Transactions of North American and reptiles of the Pacific Northwest. University of Idaho Press,
Wildlife and Natural Resources Conference 57:331-343. Moscow, ID.
89. Kuhn, L. W., and E. P. Peloquin. 1974. Oregon’s nutria problem. 114. O’Brien, P., and S. Thomas. 1998. Rabbit calicivirus: update on a new
Proceedings of the Vertebrate Pest Conference 6:101-105. biological control for pest rabbits in Australia. Proceedings of the
90. Kupferberg, S. J. 1997. Bullfrog (Rana catesbeiana) invasion of a Vertebrate Pest Conference 18:397-405.
California river: the role of larval competition. Ecology 78:1736-1751. 115. O’Gara, B. W., K. C. Brawley, J. R. Munoz, and D. R. Henne. 1983.
91. Labisky, R. F. 1975. Ecopolicies of The Wildlife Society. Wildlife Society Predation on domestic sheep on a western Montana ranch. Wildlife
Bulletin 3:36-43. Society Bulletin 11:253-264.
92. Lampman, B. H. 1946. The coming of the pond fishes. Binfords and 116. Office of Technology Assessment. 1993. Harmful non-indigenous
Mort, Portland, OR. species in the United States. OTA-F-565. U.S. Government Printing
93. Larrison, E. J. 1943. Feral coypus in the Pacific Northwest. The Office, Washington, D.C.
Murrelet 24:3-9. 117. Orians, G. H., 1986. Site characteristics favoring invasions. Pages 133-
94. ———. 1976. Mammals of the northwest: Washington, Oregon, Idaho, 148 in Harold A. Mooney and James A. Drake, editors. Ecology of
and British Columbia. Seattle Audubon Society. Seattle, WA. biological invasions of North America and Hawaii. Springer-Verlag, New
95. Lauckhart, J. B., and J. W. McKean. 1956. Chinese pheasants in the York, NY.
Northwest. Pages 43-89 in Durward L. Allen, editor. Pheasants of North 118. Parkes, J. P. 1996. Integrating the management of introduced mammal
America. Stackpole Company, Harrisburg, PA. pests of conservation values in New Zealand. Wildlife Biology 2:179-
96. LeBlanc, D. J. 1994. Nutria. Pages B-71 - B-80 in S. E. Hygnstrom, R. M. 184.
Timm, and G. E. Larsen, editors. Prevention and control of wildlife 119. ———, G. Nugent, and B. Warburton. 1996. Commercial
damage. University of Nebraska Cooperative Extension Service, exploitation as a pest control tool for introduced mammals in New
Lincoln, NE. Zealand. Wildlife Biology 2:171-177.
97. Leonard, W. P., H. A. Brown, L. L. C. Jones, K. R. McAllister, and R. M. 120. Peck, M. E., 1948. Invasion of exotic plants and their economic
Storm. 1993. Amphibians of Washington and Oregon. Seattle Audubon significance in Oregon. Northwest Science 22:126-130.
Society. Seattle, WA. 121. Peloquin, E. P. 1969. Growth and reproduction of the feral nutria
98. Lever, C. 1985. Naturalized mammals of the world. Longman Scientific Myocator coypus (Molina) near Corvallis, Oregon. M. S.Thesis, Oregon
and Technical, NY. State University, Corvallis, OR.
99. ———. 1987. Naturalized birds of the world. Longman Scientific and 122. Peterson, B., and G. Gauthier. 1985. Nest site use by cavity-nesting
Technical, New York, NY. birds of the Cariboo Parkland, British Columbia. Wilson Bulletin 97:
100. Lewis, J. C., K. L. Sallee, and R. T. Golightly, Jr. 1993. Introduced red fox 319-331.
in California. Non-game Bird and Mammal Section Report 93-10. 123. Phillips, R.L., and R.H. Schmidt. 1994. Foxes. Pages C-83 - C-88 in S.E.
California Department of Fish and Game, Sacramento, CA. Hygnstrom, R.M. Timm, and G.E. Larsen, editors. Prevention and control
101. ———, R. T. Golightly, Jr., and R. M. Jurek. 1995. Introduction of non- of wildlife damage. University of Nebraska Cooperative Extension
native red foxes in California: implications for the Sierra Nevada red Service, Lincoln, NE.
fox. Transactions of the Western Section of The Wildlife Society 31:29- 124. Pimentel, D. 1986. Biological invasions of plant and animals in
32. agriculture and forestry. Pages 149-162 in Harold A. Mooney, editor.
102. ———, K. L. Sallee, R. T. Golightly, Jr., and R. M. Jurek. 1998. Social and Ecology of biological invasions of North America and Hawaii. Springer-
biological aspects of non-native red fox management in California. Verlag, New York, NY.
Proceedings of the Vertebrate Pest Conference 18:150-155. 125. Pimm, S. L. 1989. Theories of predicting success and impact of
103. Lloyd, H. G. 1980. The red fox. B. T. Batsford, London, England. introduced species. Pages 351-367 in J.A. Drake, editor. Biological
104. Long, J. L. 1981. Introduced birds of the world: the worldwide invasions: a global perspective. John Wiley & Sons, New York, NY.
history, distribution, and influence of birds introduced to new 126. Presnall, C. C. 1958. The present status of exotic mammals in the
environments. Universe Books, New York, NY. United States. Journal of Wildlife Management 22:45-50.
127. Rainbolt, R. E. and B. E. Coblentz. 1997. A different perspective on
eradication of vertebrate pests. Wildlife Society Bulletin 25:189-191.
 CHAPTER 16: INTRODUCED SPECIES 443

128. Robinson, W. L., and E. G. Bolen. 1989. Wildlife ecology and 154. Washington Department of Wildlife. 1993. Status of the western
management. Second edition. Macmillan, New York, NY. gray squirrel, Sciurus griseus, in Washington. Unpublished report.
129. Roughgarden, J., 1986. Predicting invasions and rates of spread. Pages Washington Department of Wildlife, Olympia, WA.
179-188 in Harold A. Mooney and James A. Drake, editors. Ecology of 155. Weitzel, N. H. 1988. Nest-site competition between the European
biological invasions of North America and Hawaii. Springer-Verlag, New starling and native breeding birds in Northwestern Nevada. Condor
York, NY. 90:515-517.
130. Scheffer, T. H. 1935. Introduced food plants for waterfowl. The 156. Welsh, J.E., and R. A. Howard, Jr. 1983. Characteristics of snags
Murrelet 16:66-69. influencing their selection by cavity-nesting birds. Transactions of the
131. Schmitz, D. C., and R. G. Westbrooks. 1997. The federal government’s Northeast Fish and Wildlife Conference 40:177.
role. Pages 329-337 in Daniel Simberloff, editor. Strangers in paradise: 157.Wentz,W.A. 1971.The impact of nutria (Myocastor coypus) on marsh
impact and management of nonindigenous species in Florida. Island vegetation in the Willamette Valley, Oregon. M. S. Thesis. Oregon State
Press, Washington, D.C. University, Corvallis, OR.
132. Schmutz, J. A. 1988. Ring-necked pheasant parasitism of wild turkey 158. Westemeier, R. L., and T. L. Esker. 1989. An unsuccessful clutch of
nests. Wilson Bulletin 100:508-509. northern bobwhites with hatched pheasant eggs. Wilson Bulletin
133. Simberloff, D. 1996. Hybridization between native and introduced 101:640-642.
wildlife species: importance for conservation. Wildlife Biology 2:143- 159. Westerskov, K. 1990. Partridges and pheasants: competitors or
150. sharers? Pages 183-201 in Kevin E. Church, Richard E. Warner, and
134. ———. 1997a. The biology of invasions. Pages 3-17 in Strangers in Stephen J. Brady, editors. Gray partridge and ring-necked pheasant
paradise: impact and management of nonindigenous species in Florida. workshop. Kansas Department of Wildlife and Parks, Topeka, KS.
D. Simblerloff, editor. Island Press, Washington, D.C. 160. Wilcomb, M. J. 1948. Fox populations and food habits in relation to
135. ———. 1997b. Eradication. Pages 221-228 in Strangers in paradise: game birds in the Willamette Valley, Oregon. M. S. Thesis. Oregon State
impact and management of nonindigenous species in Florida. D. University, Corvallis, OR.
Simberloff, editor. Island Press, Washington, D.C. 161. Willner, G. R. 1982. Nutria. Pages 1059-1076 in J. A. Chapman and G.
136. Smallwood, K. S., and T. P. Salmon. 1992. A rating system for potential A. Feldhamer, editors. Wild mammals of North America: biology,
exotic bird and mammal pests. Biological Conservation 62:149-159. management, and economics. Johns Hopkins University Press,
137. ———. 1994. Site invasibility by exotic birds and mammals. Baltimore, MD.
Biological Conservation 69:251-259. 162. Wilson, E.O., and F.M. Peters, editors. 1988. Biodiversity. National
138. Spreyer, M. 1995. Pheasant obsession. Birder’s World 9:44-48. Academy Press, Washington, D.C.
139. Stein, B. A., and S. R. Flack, editors. 1996. Americas least wanted: alien 163. Witmer, G. W. 1990. Re-introduction of elk in the United States.
species invasion of U.S. ecosystems. The Nature Conservancy, Journal of the Pennsylvania Academy of Science 54:131-135.
Arlington, VA. 164. ———, J. L. Bucknall, T. H. Fritts, and D. G. Moreno. 1996. Predator
140. Storer, T. I. 1933. Frogs and their commercial use. California Fish and management to protect endangered avian species. Transactions of the
Game 19:203-213. North American Wildlife and Natural Resources Conference 61:102-
141. Storm, G. L., R. D. Andrews, R. L. Phillips, R. A. Bishop, D. B. Siniff, and J. 108.
R. Tester. 1976. Morphology, reproduction, and mortality of mid- 165. ———, E. W. Campbell, and F. Boyd. 1998. Rat management for
western fox populations. Wildlife Monographs 49. endangered species protection in the United States Virgin Islands.
142. Taylor, W. P. 1923. Upland game birds in the state of Washington. The Proceedings of the Vertebrate Pest Conference 18:281-286.
Murrelet 4:3-15. 166. Zembal, R. 1992. Status and management of light-footed clapper rails
143. Temple, S. A. 1990. The nasty necessity: eradicating exotics. in coastal southern California. Transactions of the Western Section of
Conservation Biology 4:113-115. The Wildlife Society 28:1-5.
144. Thomas, J. W., editor. 1979. Wildlife habitats in managed forests; the
Blue Mountains of Oregon and Washington. Agricultural Handbook No.
553. U.S. Department of Agriculture, Forest Service, Portland, OR.
145. Thornton, W. A., G. C. Creel, and R. E. Trimble. 1971. Hybridization in
the fox genus Vulpes in west Texas. Southwest Naturalist 25:423-434.
146. Toney, J. C., P. M. Rice, and F. Forcella. 1998. Exotic plant records in
the Northwest United States 1950-1996: an ecological assessment.
Northwest Science 72:198-209.
147. Troetschler, R. G., 1976. Acorn woodpecker breeding strategy as
affected by starling nest-hole competition. Condor 78:151-165.
148. Van Vuren, D. 1992. Eradication of feral goats and sheep from island
ecosystems. Proceedings of the Vertebrate Pest Conference 15:377-
381.
149. Van Vuren, D. 1998. Manipulating habitat quality to manage vertebrate
pests. Proceedings of the Vertebrate Pest Conference 18:383-390.
150.Vance, D. R., and R. L. Westemeier. 1979. Interactions of pheasants
and prairie chickens in Illinois. Wildlife Society Bulletin 7:221-225.
151.Verts, B. J., and L. Carraway. 1980. Natural hybridization of Sylvilagus
bachmani and introduced S. floridanus in Oregon.The Murrelet 61:95-
98.
152.———, and ———. 1998. Land mammals of Oregon. University of
California Press, Berkeley, CA.
153.Voigt, D. R. 1987. Red Fox. Pages 379-392 in M. Nowak, J. A. Baker, M.
E. Obbard, and B. Malloch, editors. Wild furbearer management and
conservation in North America. Ontario Ministry of Natural
Resources.Toronto, Ontario, Canada.
 17
Genetic Considerations for Introduced and
Augmented Populations
Susan M. Haig & R. Steven Wagner

Introduction
Habitat fragmentation has occurred at a record pace the design of new populations. Molecular techniques,
throughout many areas of the Pacific Northwest. As a pedigree analyses, or viability models are tools used to
consequence, wildlife and land managers in this region evaluate “genetic factors.”
are forced to deal with recovery of small populations. In the broadest sense, small populations should be
Frequently, these efforts have focused on politically managed to avoid short-term demographic disasters and
charged issues related to spotted owls, marbled murrelets, to insure long-term adaptability (i.e., maximize genetic
or salmon (Oncorhynchus spp.). Increasingly, however, diversity). Loss of genetic diversity diminishes the
there are more species facing extirpation that receive little flexibility of populations to adapt to changing conditions,
attention and fall outside the protection provided by more resulting in a decreased ability to cope with parasites,
recognized species. disease, competitors, changing climatic conditions, etc.
Direct conservation of the land is usually the best There has been much debate over prioritization of genetic
approach to dealing with habitat loss for most species. and demographic concerns. Some suggest that over the
However, habitat conservation efforts often come too late short term, concern for demographic factors should take
for natural recovery of specific populations or species and precedence over genetic issues.37 However, the interactions
more drastic actions are needed to assure their survival between genetic and demographic factors are not mutually
and viability. To achieve this goal, individuals may need exclusive. The demographic structure of a population
to be translocated to/from other populations or, if a species affects the genetic structure and vice versa. Consequently,
or population has gone extinct in one locale, a simple management decisions can be made when
reintroduction from captivity or another wild population establishing a new population (either via reintroduction
may be necessary. or augmentation) that will improve both demographic and
This seemingly simple concept, of moving individuals genetic structure of the population.20 Thus, both factors
among populations or establishing new ones, has had should be considered from the start of a program.
variable success and is quite controversial. For example,
the success rate was only 11% (16 cases) among 145
reintroduction programs that released captive animals 1. Factors Influencing Population Viability
back to the wild.6 The success rate among 80 translocation Demographic factors—factors that affect individual
programs was better, but less than half (46%) resulted in birth and death rates. Demography is most critical
establishing self-sustaining populations of mammals and to consider if a population is very small
birds.19, 60 Similar efforts for reptiles and amphibians were (approximately less than 40 individuals), because
even less successful—only 5 of 25 programs were stochastic events can be severe in small
successful. 9 Thus, reintroduction and translocation populations making them susceptible to Allee
programs should only be carried out as a last resort.56 effects.2
There are many explanations for the lack of success of Genetic factors—factors that influence small
these programs (reviewed in Chapter 18), however, in this populations through random genetic drift
chapter we will outline genetic factors that need to be (random loss of alleles) and inbreeding which can
carefully considered if a decision is reached to establish a reduce genetic diversity.
new population or augment an existing population. As in Environmental factors—factors that can impact a
any population recovery effort, there are numerous population of any size. They include variations in
overlapping factors to consider 51 which include: weather, food availability, and other factors that
Demographic factors, environmental factors, catastrophic events, affect survival, reproduction and abundance.
and genetic factors (see Box 1). The least understood and/ Catastrophic factors—significant random events
or addressed concern in new population management is that can affect a population of any size. They
the consideration of genetic factors. There are numerous include flood, fire, hurricanes, etc.
ways in which “genetic factors” are defined7 and used in

444
 CHAPTER 17: REINTRODUCED SPECIES 445

A contrary view is that some amount of inbreeding

2. Factors Related to allows for the maintenance of co-adapted genes52, 53, 59 (i.e.,
Random Genetic Drift adaptation to the local environment). Especially in new
Random Genetic Drift61, 62—mechanism by which populations, this potential benefit is generally outweighed
populations lose genetic diversity. It is the result of by the importance of establishing populations with
random changes in gene frequency from unrelated individuals so as to increase genetic diversity
generation to generation. as quickly as possible. Consequently, there is some concern
Effective Population Size (Ne,61)—is inversely that managers, in an effort to mitigate the effects of drift,
proportional to the rate of loss of genetic diversity. may introduce genetically-distant individuals into the
It is equal to the size of an ideal population. population. This type of introduction could lead to the
Ideal Population—defined as a population meeting disruption of co-adapted gene complexes or the ability to
the following assumptions: random mating (e.g. adapt to the local environment. The result would be
no inbreeding), 1:1 breeding sex ratio, equal outbreeding depression which could have consequences as
family sizes, large population size, and severe as inbreeding depression59 (Box 3). Populations
non-overlapping generations. should be monitored beyond the first generation (F1) for
Population Bottleneck—a drastic decline in Ne the effects of outbreeding. Usually the first generation
resulting in a small number of survivors passing shows increased fitness but this can quickly erode in
on a fraction of genes from the original subsequent generations as co-adapted complexes are
population. broken up.41 Overall, breeding strategies should mitigate
between the effects of inbreeding and outbreeding.
An important measure for determining the rate of loss
of genetic diversity in a population is effective population
size61 (Ne, Box 2). Effective population size is inversely
The most critical genetic threats facing small or new proportional to the loss of genetic diversity. The concept
populations are the loss of diversity and the potential for of Ne can be confusing, but it is an important concept for
inbreeding depression.3 Genetic diversity is lost primarily managing genetic and demographic factors in a
through random genetic drift and, as a consequence, this population—especially as new populations are
increases inbreeding and decreases effective population size established. Most simply, effective population size is equal
(see Box 2). In general, populations lose genetic diversity to the size of an ideal population, which is defined based
over time, with the rate of loss largely dependent on the on the following assumptions: random breeding (e.g., no
number of individuals in the population. This loss can be inbreeding), 1:1 breeding sex ratio, equal family sizes, large
balanced by the number of founders or migrants into the population size, non-overlapping generations, etc. The
population—factors that are most affected early on in a effective size is usually smaller than the censused population
new population. Random genetic drift is the mechanism by size (N) because most natural populations do not meet all
which small populations lose genetic diversity, while of the assumptions of an ideal population. The ratio Ne/
effective population size is a way to measure this rate of loss. N, the comparison of effective to censused population size,
Thus, the appropriate strategy for new/small population gives an indication of how different the censused size is
management is to minimize random genetic drift and from the ideal size. Thus, it can be used by population
maximize the effective size of the population. managers to assess the status of a population and to judge
Random genetic drift reduces two components of progress toward achieving a more genetically diverse and
genetic diversity in small populations: allelic diversity and demographically stable (i.e., birth rates at least equal death
heterozygosity1 (see Box 3). Maintaining both factors rates and emigration) population. Moreover, it can be used
becomes increasingly difficult as populations become
smaller, because the effect of drift becomes greater due to
the random sampling of fewer genes, resulting in greater
fluctuations in gene frequency. Therefore, while a number 3. Population Genetic Terminology
of factors can affect genetic diversity, random genetic drift Allelic diversity—number of unique alleles that
is an overriding factor in controlling loss of genetic occur in a population.
variation.36 Heterozygosity—frequency of alleles per locus.
Substantial genetic drift can result in increased Inbreeding—mating of close relatives.
inbreeding. Inbreeding can be simply defined as the mating Inbreeding depression—loss of fitness resulting from
of close relatives, but is often confused with inbreeding inbreeding.
depression, which is the loss of fitness resulting from Outbreeding—mating of genetically distant
inbreeding. Generally, demographic factors, such as individuals.
fecundity, fertility, development rate, age of sexual Outbreeding depression—loss of fitness resulting
maturity, and litter size can be affected by increased from the mating of genetically distant individuals,
inbreeding. Thus, from a conservation perspective, usually associated with the disruption of
inbreeding is typically considered detrimental and to be co-adapted gene complexes.
avoided.49
 446 Wildlife–Habitat Relationships in Oregon and Washington

as a means of assessing the relative effects of factors that a short-term management goal of Ne=50 and a long-term
might contribute to the viability of a new population. management goal of N e=500 in order to maintain
A sudden or drastic decline in Ne or a population population viability. Long-term goals proposing an
bottleneck results when a small number of survivors pass optimum effective population size are controversial:
on a fraction of genes from the original population. By recently Franklin and Frankham15 revised the target to
definition, bottlenecks occur when populations suffer include a range of Ne between 500-1000, while others
drastic declines, such as when animals are brought into contend the Ne should be higher, between 1000-5000.38, 42
captivity, or a new population is established. Taken to an However, as Lande and Barrowclough38 point out, these
extreme, this could result in population extinction. Thus, goals are unrealistic for many populations because they
it is critical to consider problems associated with are too general (e.g., an Ne of 50 for many rodents would
bottlenecks in planning new populations. That is, selection be much easier to attain than a Ne of 50 for grizzly bears)
of individuals to establish or enhance a population must and do not take into account population-specific life
be undertaken with extreme caution. Also, managers history characters and demography. Thus, population
should try to minimize the effect of a bottleneck by viability goals should be viewed with respect to effective
increasing the number of individuals as quickly as population size but specific goals should be developed
possible. This can be achieved by introducing more for each situation in order to maintain genetic diversity in
breeding age individuals—with consideration to mating a realistic fashion.
system and current sex ratio; providing extra-protective
How many/which individuals to translocate. While more
measures for eggs, nests, or juveniles; creating artificial
is usually better, often there are limited choices regarding
nest sites or breeding areas; considering some type of
the number of individuals available to start a population.
predator management measures, etc.
Also a balance must be struck between offsetting the effects
Establishing Viable Populations of drift, preserving local adaptation and providing enough
If a decision is made to establish a new population or individuals to overcome genetic and demographic threats.
enhance an existing population, some practical decisions This is illustrated by a small population of red-cockaded
should be made in view of the above discussion. Often woodpeckers (Picoides borealis) in South Carolina, where
genetic considerations are quite simple and can managers wanted to increase the population of 5-6 birds
profoundly affect a program’s success, yet they are to a viable population.23 Pre-translocation modelling
frequently ignored. For example, in a world-wide review indicated that annual translocation of three females and
of 94 translocation or reintroduction programs, Kalmer35 two males for a ten-year period would achieve the same
found genetic factors were only considered in 47 cases. genetic and demographic goals as moving many more
Listed below are a few questions that should be considered individuals—which were not available from donor
for both the new and donor population: populations anyway. Thus, the modelling was able to give
1. What is the population viability goal we are trying to managers some realistic goals. Some general guidelines
reach in the new population and maintained in the for increasing N e and assuring a genetically fit and
donor population? demographically stable population include the
2. How many individuals are needed/could be given following:39, 48
up? 1. Begin with at least 20-30 unrelated founders.
3. What is the relatedness of individuals to be 2. Increase carrying capacity quickly.
introduced/to be lost? 3. Equalize founder contribution/mean kinship.
4. Minimize inbreeding.
Viability goals. As with any recovery planning effort, an 5. Equalize family size.
assessment of needs and risks must be carried out prior 6. Establish a demographically stable population.
to moving individuals. Managers need a realistic idea of Meeting any or all of these management objectives may
what they have (genetically and demographically) and be difficult or impossible for many species or situations;
what they can hope for, given careful planning. Unrealistic however, effectively addressing any of them will increase
goals will insure the failure of any program. Molecular Ne and result in a more stable population.
methods can be used to assess genetic diversity, the genetic
distance among various populations, and identify Monitor progress towards recovery. Genetic diversity
relatedness of individuals in a population.20 Pedigree should be measured during various phases of population
analyses can also be used to assess the genetic status of growth as a means of diagnosing progress toward
donor populations or evaluate breeding options.21, 22, 24 viability.17, 58 For example, during the founder phase, the
Finally, population viability models can be used to number of founders and their relatedness has a critical
evaluate various strategies for long and short-term effect on genetic diversity. As the population begins to
survival.23 increase, a slow population growth rate, an extended
Several short-term and long-term population goals duration of growth phase, or a small carrying capacity
have been proposed in the conservation literature. can reduce genetic diversity. Once the carrying capacity
Primarily, the maintenance of genetic diversity will has been reached, a small carrying capacity or short
increase with more individuals in a population and an duration of the capacity phase can also result in loss of
increased Ne.11, 12, 13 More specifically, Franklin14 suggested diversity.
 CHAPTER 17: REINTRODUCED SPECIES 447

During any phase of population establishment, Hedrick and Parker31 evaluated stocking options for the
inbreeding, outbreeding, or the loss of founder lines can endangered Gila topminnow (Poeciliopsis o. occidentalis) by
become a problem. Thus, managers need to carefully examining genes from the major histocompatibility
monitor productivity of adults and survivorship of young. complex (MHC). MHC genes are thought to have adaptive
Inbreeding can be examined in a variety of ways, but significance, particularly for resistance to parasites and
fundamental to any analysis will be establishment of a pathogens, hence their diversity within populations may
pedigree from the beginning of the population. If reduced be informative relative to their adaptive potential. They
survivorship is observed and appears to have a genetic found significant differences among individuals from the
basis, setting up test pairings may confirm the problem. four watersheds they sampled, and recommended that
Even if this is not possible, separating pairs of suspected individuals not be mixed among watersheds.
relatives may be advised. If new populations are too small As new populations are established, consideration
or it is too intrusive to alter pairings, introduction of new should be given to the overall metapopulation structure.
individuals may help. In general, inbreeding can best be While many strategies have been debated, it usually comes
avoided by increasing the population size quickly and down to consideration of the SLOSS model 55 (i.e.,
selecting pairs to minimize mean kinship and equalize managing for a single large or several small populations).
founder contribution. That is, how do we balance the benefits of a single large
population with those of several small? Genetically and
Donor Populations and Metapopulation demographically, a large population is generally more
Management viable over a longer period of time, whereas small
Establishment of new populations cannot be carried out populations face greater risk of immediate extinction.
in a vacuum; that is, without regard to the effect on the However, small populations can serve as reservoirs for
structure of donor populations and the overall unique alleles, provide a buffer if catastrophe hits the large
metapopulation. One of the first factors to consider is the population, and provide links to other populations.40 Thus,
source of donors to establish a new population. In general, a mix of both large and inter-linked small populations may
choosing from populations that are genetically and be the most effective scenario. In translocation/
ecologically similar will result in individuals that have reintroduction programs, maintenance of the
adapted to the area’s particular characteristics. For metapopulation may be carried out via natural or human-
example, in re-establishing a viable population of red- induced movements of individuals among populations.
cockaded woodpeckers in South Carolina, molecular Since the 1930s, the rule of thumb has been that movement
analyses were performed on samples from throughout the into a population of one individual per generation would
species range to determine the most genetically similar offset the negative effects of drift.61 Reconsideration of this
donor population.24, 26 Donor birds were then chosen from rule led Mills and Allendorf44 to recommend that one
populations that were genetically and geographically individual was a minimum and movement of up to ten
close, would not suffer as a result of loss of individuals, individuals per local population per generation would
and had similar habitat conditions. better assure maintenance of a viable metapopulation.
The recommendation to move individuals from the
most similar population is not always an alternative. In Genetic Tools to Assess and Monitor New
the case of the Florida panther (Felis concolor coryi), the Populations
genetically closest individuals were from a Texas (F. c. Molecular techniques are increasingly employed to aid in
stanleyana) subspecies. In this case, outbreeding depression designing population enhancement strategies.20 These
might have been an issue, however, there were no other analyses are most helpful in defining founder
options. Modelling the potential impact on the Florida relationships, monitoring genetic diversity, and
population led biologists to conclude that the benefits determining relationships among populations in which
would outweigh the risks.29 In another unique situation, translocations and augmentations are being considered.

Table 1. Application of molecular techniques in resolving issues of small population conservation.

Application Allozymes DNA fingerprints Microsatellites RAPDs mtDNA MHC

(RFLPs)

Parentage (X)1 X X (X) X

Mating system X X X (X) X
Pedigree definition X X X
Population structure X (X) X X X X
Population differentiation X X X X X
Hybridization X X X X X
Phylogeny (X) X
1. () indicates that while appropriate, another technique would most likely yield more helpful information.
 448 Wildlife–Habitat Relationships in Oregon and Washington

Each technique is best suited for a limited range of This technique can be very quick and cost effective if
questions, hence several may be necessary in order to get primers have been previously designed. However, these
a complete picture of a population structure46 (Table 1). primers are often species-specific and can be extremely
For example, DNA fingerprints or restriction fragment difficult to elucidate, especially in birds.47 Thus, without
length polymorphisms (RFLPs) have a very high mutation pre-designed primers this technique can be one of the most
rate and can show substantial variability among closely- costly and time consuming techniques to develop.
related individuals. However, this variability reaches Use of randomly amplified polymorphic DNA
saturation among genetically distant individuals. (RAPDs) is increasingly applied to problems of population
Therefore, they are better at resolving questions related structure, differentiation, and hybridization.24, 25, 26, 27 There
to parentage, pedigree, and population structure than for have also been a limited number of studies that have
resolving higher level analyses such as addressing employed RAPDs to infer parentage and pedigrees.
questions regarding population differentiation or RAPDs are one of the easiest, fastest and most cost effective
phylogeny. of all the molecular techniques. The development phase
Protein electrophoresis or allozyme analyses are one of screening for variable primers is minimal and straight
of the most widely used techniques used to assay genetic forward. It is also possible to screen a large number of
diversity34 as they can be used to address a number of individuals (20-30) per population for a number of loci
questions ranging from parentage to phylogeny. Even with (15-20) relatively quickly. RAPDs are also a PCR based
the advent of new techniques, they remain one of the most technique so almost any type of tissue may be used. One
powerful, quick, and inexpensive techniques for certain drawback of RAPDs is that they are a dominant marker,
types of questions. Typically, any type of tissue can be used therefore heterozygotes can not be directly scored. RAPDs
(i.e., blood, muscle, heart, liver, feather pulp), and 5-10 have been used to evaluate translocation strategies for the
variable loci are usually screened. While historically Australian woody perennial Corrigan grevillea (Grevillea
allozymes have been used for parentage, population scapigera), a plant that has been reduced to four
structure, and population differentiation studies, populations (total of 27 plants) by habitat destruction.50
depending upon the taxa and scale of the question, there The RAPD data indicated little difference among
might not be adequate variability to resolve some of these populations, hence they recommended that translocations
questions. For example, for many avian species, allozyme would not have negative genetic effects.
variability is too low to address many population-level Mitochondrial DNA (mtDNA) markers are some of the
questions.5 most widely-used in conservation biology.4 Their best use
Often in new populations, relatedness of individuals is in addressing questions of population structure,
in the wild donor population is not known, making it hybridization, and phylogeny. For small populations, this
difficult to choose individuals to translocate. DNA marker can be useful for identifying populations from
fingerprinting (minisatellites) and microsatellite loci are which to translocate individuals. MtDNA is a maternally-
two techniques increasingly employed to aid in this inherited, extrachromosomal small molecule of DNA
question. For example, DNA fingerprinting was used to (~16kb for vertebrates) found in the mitochondria. The
determine the relatedness of founders to the captive molecule is useful because it evolves at a rapid rate and
population of Guam rails (Rallus owstoni) prior to their different regions have variable mutation rates.
reintroduction to the wild.21 These fingerprinting profiles Furthermore, since it is uniparentally inherited and non-
tested five hypothesis proposed to explain relatedness recombining there is little sequence heterogeneity within
among the nine founders. Band-sharing among the an individual. MtDNA studies involve two kinds of
profiles was correlated with relatedness calculated from analyses: DNA sequencing or restriction site analysis
hypothesized pedigrees. This relationship was used to (RFLPs). DNA sequencing usually involves sequencing
determine the most plausible founder hypothesis, and to of 400-1000 bases of various genes for intraspecific
construct a breeding/management scheme that phylogeny studies. For example, the cytochrome b, COI,
maximized genetic diversity in the new population. and NADH genes have been sequenced for intraspecific
Microsatellites have a broader range of application phylogeny studies and the highly variable d-loop region
than DNA fingerprints as they can address questions has been used to explore both intraspecific phylogeny and
related to population differentiation. For example, population stucture. Given the availability of universal
microsatellites were used to determine the primers for many mtDNA genes, these kinds of studies
appropriateness of moving Marianas crows (Corvus are moderately priced. Phylogenetic analysis among
kubaryi) from Rota to Guam.10 Microsatellites also have an populations of the wet forest-restricted frog Litoria
advantage over DNA fingerprinting and random pearsoniana using the COI gene (460bp), indicated that
amplified polymorphic DNA (RAPDs) because conservation efforts should be made at the scale of major
heterozygotes can be scored directly. Since the rainforest isolates. Furthermore, investigators suggested
microsatellite technique is based upon the polymerase that the conservation status of the species should be
chain reaction45 (PCR), any type of tissue may be used with considered independently for northern and southern
small amounts of tissue (e.g., blood, hair, feathers, scat, populations due to significant differentiation.43
bone, tissue from museum specimens, etc.). These studies The major histocompatibility complex (MHC) is
usually involve screening 5-15 variable loci per individual. increasingly used as a marker to assess genetic variability
 CHAPTER 17: REINTRODUCED SPECIES 449

in captive populations. 28, 30 MHC has a number of among these populations and help determine if
advantages over other markers. First, MHC is suggested translocating among them is advisable. For example,
to have alleles of selective significance, unlike the other because origin of the Puget Sound turtles is not clear,
the markers which we assume to be neutral. Second, MHC augmentation of the Gorge sites with Puget Sound turtles
has important immune functions and is studied widely could break up co-adapted gene complexes and lead to
to understand autoimmune diseases, tissue trans- outbreeding depression. Currently, the Washington
plantation, and immune response to diseases. A study of Department of Fish and Wildlife has been “headstarting”
two populations of pocket gophers found low variability juveniles in captivity. They have mixed juveniles across
at the DQα locus and each fixed for unique alleles.54 These the Gorge sites but not with Puget Sound or other sites.
results were also consistent with the rejection of reciprocal This approach of increasing the population size but not
skin grafts, therefore, the authors suggested that these mixing populations appears prudent until further
populations should not be mixed. molecular analysis can be completed.
In addition to choosing a technique, the feasibility of
obtaining adequate samples for each type of study must Upland Sandpipers
be considered in the planning stages. For example, Upland sandpipers are a shorebird species suffering from
parentage/pedigree studies require comparisons of highly drastic population decline and fragmentation in the
variable genetic loci between individuals. For these types Northwest due to habitat deterioration, often as a result
of questions sampling complete families (i.e., mother, of overgrazing and agriculture.8, 16, 32 They have been
father, all offspring, other potential fathers, and helpers; extirpated from Washington and less than 100 remain in
n = ~20 families) and multiple generations will be eastern Oregon and Idaho. The nearest viable populations
necessary. Population structure/differentiation studies are in the prairies of the Dakotas and Manitoba, but even
involve relative comparisons among populations, so there these are declining. Thus some of the genetic issues include
must be adequate sampling of both individuals per whether translocating birds from the Midwest would
population (~20-40 per population). Hybridization and result in an outbreeding problem, whether the Midwest
phylogeny studies generally involve comparisons at populations would be further jeopardized if they donated
higher taxonomic levels and markers that have a slower individuals, and what population structure would be most
rate of mutation, therefore, depending on the marker, viable for the new populations. As with pond turtles,
fewer individuals per group (~1-5) need to be sampled. mtDNA or microsatellite analyses of Northwest and
Midwest birds will help assess differences between
Genetic Considerations for populations. Some population viability modelling will
help determine the effect of loss of individuals on the
Northwest Populations donor populations as well as what will be needed for the
The information presented here attains the greatest new or enhanced populations.
significance when we consider issues close to home. Game
animals have been translocated for years within and Pygmy Rabbits
among sites in the Northwest. More recently, however, Pygmy rabbits are a Columbia Basin species whose native
population restoration strategies have been warranted for sagebrush habitat is now so reduced that only five
small populations in the region. The following examples populations remain in southeastern Washington
illustrate species where restoration efforts will be (Washington Department of Fish and Wildlife, unpub.
enhanced if genetic factors are considered along with the data). With less than 100, if not less than 50, individuals
other issues. per population, biologists suspect that there is little genetic
or demographic interchange among populations.
Western Pond Turtles Furthermore, the dynamic nature of the Columbia Basin
Western pond turtles have been rapidly declining often renders these small populations susceptible to
throughout their Pacific Coast range for nearly a century catastrophic events such as floods, fires, and disease. Thus,
and are extirpated from most of Washington and Oregon.8, since all populations are small, a decision to manipulate
33, 57
There are numerous hypotheses for their decline which any population may result in its extinction, yet no action
include disease, human disturbance, and predation by may have the same result. Therefore, a starting point for
introduced largemouth bass (Micropterus salmoides) and species recovery would be to assess the genetic distance
bullfrogs. In the Northwest, there are less than 250 among populations with mtDNA or microsatellites. If
individuals among six sites on the Washington side of the genetic distances were not significant, consideration of
Columbia River Gorge and one site on the Oregon side of moving several individuals among populations may be
the Gorge. Another site, in Puget Sound, is located at an warranted given concern about loss of genetic diversity
artificial pond and originated from 3 founders.18 It is not due to drift. A more complex decision could be then be
known if these individuals are native to the Puget Sound considered regarding population consolidation. If some
basin because each founder was found in a semi-rural populations are too small to survive and there are no donor
environment where there were no known populations. populations outside Washington and Oregon, it may be
From a genetic perspective, phylogenetic studies using best to move the remaining individuals to a nearby
mitochondrial DNA would help define the relationships population with enough suitable habitat to sustain them.
 450 Wildlife–Habitat Relationships in Oregon and Washington

Clearly, this would be a difficult decision as moving an 6. Beck, B.B., L.G. Rapaport, M.S. Price, and A. Wilson. 1994.
entire population may result in loss of protective efforts Reintroduction of captive born animals. Pages 265-284 in P.J.S. Olney,
for the habitat they occupied. However, if it stabilized G.M. Mace, and A.T.C. Feistner, editors. Creative Conservation:
Interactive Management of Wild and Captive Animals. Chapman and
another population on the brink of extinction, it may be a Hall, London.
worthwhile endeavor. Revisiting the pros and cons 7. Beissinger, S.R, J.R. Walters, D.G. Catanzaro, K.G. Smith, J.B. Dunning, Jr.,
suggested by the SLOSS model would be in order prior to S.M. Haig, B.R. Noon, and B.M. Smith. The use of models in avian
undertaking such an action. conservation. Current Ornithology (in press).
8. Csuti, B., A.J. Kimmerling, T.A. O’Neil, M.M. Shaughnessy, E.P. Gaines, and
Oregon Silverspot Butterfly M.M.P. Huso. 1997. Atlas of Oregon Wildlife. Oregon State University
Oregon silverspot butterfly (Speyeria zerene hippolyta) is a Press, Corvallis, OR.
9. Dodd, C.K., and R.A. Seigel. 1991. Relocation, repatriation, and
species extirpated from Washington and has only three
translocation of amphibians and reptiles: are they conservation
remnant populations in Oregon. To restore the butterflies strategies that work? Herpetologia 47: 336-350.
to their native Washington habitat, translocated 10. Duckworth, W.D., S.R. Beissinger, S.R. Derrickson, T.H. Fritts, S.M. Haig,
individuals from Oregon would have to be chosen from F.C. James, J.M. Marzluff, and B.A. Rideout. 1997. Scientific Bases for
areas that are geographically closest and occur in habitats Preservation of the Marianas Crow. National Academy of Science.
most similar to what the original Washington populations Washington D.C.
11. Frankel, O.H., and M.E. Soule. 1981. Conservation and Evolution.
occupied. Prior to any movements, Oregon populations
Cambridge University Press, New York, NY.
need to be assessed to determine if they could sustain loss 12. Frankham. R. 1996. Relationship of genetic variation to population
of a few individuals without significant loss of viability. size in wildlife. Conservation Biology 10:1500-1508.
Next, genetic distance should be assessed. If there is no 13. ———, and I.R. Franklin. 1998. Response to Lynch and Lande. Animal
significant genetic difference among populations, it might Conservation 1:73.
be best to choose individuals from habitats as similar to 14. Franklin, I.R. 1980. Evolutionary change in small populations. Pages
the new habitats as possible. This will speed their 135-149 in M. E. Soule and B. A. Wilcox, editors. Conservation
Biology—An Evolutionary Perspective. Sinauer Assoc. Sunderland MA.
adaptation to the new site and more quickly establish the
15. ———, and R. Frankham. 1998. How large must populations be to
new populations. retain evolutionary potential? Animal Conservation 1: 69-73.
16. Gilligan, J., M. Smith, D. Rogers, and A. Contreras. 1994. Birds of
Conclusions Oregon: Status and Distribution. Cinclus Publications, McMinnville, OR.
17. Gompper, M.E., P.B. Stacey, and J. Berger. 1997. Conservation
In this chapter we have reviewed the significance of
implications of the natural loss of lineages in wild mammals and birds.
considering genetic factors in augmenting or establishing Conservation Biology 11: 857-867.
new populations. Because single individuals can have a 18. Gray, E.M. 1995. DNA fingerprinting reveals a lack of genetic variation
significant impact on population structure and genetic in northern populations of the Western Pond Turtle (Clemmys
diversity can be lost quickly in small populations, new marmorata). Conservation Biology 9:1244-1255.
populations can be helped most effectively if factors such 19. Griffith, B., J.M. Scott, J.W. Carpenter, and C. Reed. 1989.
as the number of founders, relatedness of founders, and Translocations as a species conservation tool: status and strategy.
Science 245: 477-480.
genetic distances between donor and recipient populations
20. Haig, S.M. 1998. Molecular contributions to conservation. Ecology 79:
are considered at the beginning of recovery efforts, not as 413-425.
an afterthought. These factors affect both the genetic and 21. ———, J.D. Ballou, and N.J. Casna. 1994. Identification of kin structure
demographic stability of new populations, and can often among Guam Rail founders: a comparison of pedigrees and DNA
be measured most efficiently with genetic techniques such profiles. Molecular Ecology 5:109-119.
as molecular tools. Overall, genetic management is often 22. ———, ——— and S.R. Derrickson. 1990. Management options for
overlooked due to managers unfamiliarity with terms or preserving genetic diversity: reintroduction of the Guam Rail to the
wild. Conservation Biology 4: 290-300; 464.
techniques, yet it can have a significant impact on overall 23. ———, J.R. Belthoff, and D.H. Allen. 1993. Population viability analysis
population viability. for a small population of Red-cockaded Woodpeckers and an
evaluation of population enhancement strategies. Conservation Biology
L iterature Cited 7:289-301.
1. Allendorf, F.W., and R.F. Leary. 1986. Heterozygosity and fitness in 24. ———., R. Bowman, and T.D. Mullins. 1996. Population structure of
natural popualtions of birds and mammals. Pages 57-76 in M.E. Soule, Red-cockaded Woodpeckers in south Florida: RAPDs revisited.
ed. Conservation Biology: the Science of Scarcity and Diversity. Sinauer Molecular Ecology 5: 725-734.
Assoc., Sunderland, MA. 25. ———, C.L. Gratto-Trevor, T.D. Mullins, and M.A. Colwell. 1997.
2. Andrewartha, H.G., and L.C. Birch. 1954. The Distribution and Population identification of western hemisphere shorebirds throughout
Abundance of Animals. University of Chicago Press, Chicago, IL. the annual cycle. Molecular Ecology 6: 413-427.
3. Avise, J.C. 1994. Molecular Markers, Natural History, and Evolution. 26. ———, J.M. Rhymer, and D.G. Heckel. 1994. Population differentiation
Chapman and Hall, New York, NY. in randomly amplified polymorphic DNA of Red-cockaded
4. ———. 1995. Mitochondrial DNA polymorphism and a connection Woodpeckers. Molecular Ecology 3: 581-595.
between genetics and demography of relevance to conservation. 27. ———, R.S. Wagner, E.D. Forsman, and T.D. Mullins. Geographic
Conservation Biology 9: 686-690. variation and genetic structure in Spotted Owls. Conservation
5. Barrowclough, G.F. 1983. Biochemical studies of microevolutionary Genetics (in review).
processes. Pages 223-261 in A.H. Brush and G.A. Clark, editors. 28. Hedrick, P.W. 1994. Evolutionary genetics of the major
Perspectives in Ornithology. Cambridge University Press, New York, histocompatibility complex. American Naturalist 143: 945-964.
NY. 29. ———. 1995. Gene flow and genetic restoration: the Florida Panther
as a case study. Conservation Biology 9:996-1007.
 CHAPTER 17: REINTRODUCED SPECIES 451

30. ———, and P.S. Miller. 1996. Rare alleles, MHC and captive breeding. 53. ———. 1993. The natural and unnatural history of inbreeding and
Pages 187-204 in V. Loeschcke, J.Tomiuk, and S.K. Jain, editors. outbreeding. Pages 143-172 in N. W Thornhill, editor. The Natural
Conservation Genetics. Birkhauser, Basel, Switzerland. History of Inbreeding and Outbreeding. University of Chicago Press,
31. ———, and K.M. Parker. 1998. MHC variation in the endangered Gila Chicago, IL.
Topminnow. Evolution 52: 194- 199. 54. Sanjayan, M.A., K. Crooks, G. Zegers, and D. Foran. 1996. Genetic
32. Herman, S.G., J.W. Scoville, and S.G. Waltcher. 1984. The Upland variation and the immune response in natural populations of pocket
Sandpiper in Bear Valley and Logan Valley, Grant County, Oregon. gophers. Conservation Biology 10:1519-1527.
Report to Oregon Department of Fish and Wildlife. 55. Simberloff, D.S. and L.G. Abele 1976. Island biogeography theory and
33. Holland, D.C. 1993. A synopsis of the distribution and current status conservation practice. Science 191: 285-286.
of the Western Pond Turtle (Clemmys marmorata). Report to Oregon 56. Snyder, N.R., S.R. Derrickson, S.R. Beissinger, J.W. Wiley, T.B. Smith,
Department of Fish and Wildlife. W.D. Toone, and B. Miller. 1996. Limitations of captive breeding in
34. Hubby, J.L. and R.C. Lewontin. 1966. A molecular approach to the endangered species recovery. Conservation Biology 10: 338-348.
study of genetic heterozygosity in natural populations, I: The number of 57. Stebbins, R.C. 1985. A Field Guide to Western Reptiles and
alleles at different loci in Drosophila psuedoobscura. Genetics 54:577-94. Amphibians. Second Edition, Houghton Mifflin Co. Boston, MA.
35. Kalmer, A. 1995. Genetic assessments in animal translocation 58. Stockwell, C.A., M. Mulvey, and G.L. Vinyard. 1996. Translocations and
programmes: re-introductions, re-stocking, and conservation the preservation of allelic diversity. Conservation Biology 10: 1133-
introductions. M.S. Thesis, University of Kent, Canterbury, England. 1141.
36. Lacy, R.C. 1987. Loss of genetic diversity from managed populations: 59. Templeton, A.R. 1986. Coadaptation and outbreeding depression.
interacting effects of drift, mutation, immigration, selection, and Pages105-116 in M.E. Soule, editor. Conservation Biology: the science of
population subdivision. Conservation Biology 1:143-158. scarcity and diversity. Sinauer Assoc., Sunderland, MA.
37. Lande, R. 1995. Mutation and conservation. Conservation Biology 9: 60. Wolf, C.M., B. Griffith, C. Reed, and S.A. Temple. 1996. Avian and
782-791. mammalian translocations: update and reanalysis of 1987 survey data.
38. ———, and G.F. Barrowclough. 1995. Effective population size, Conservation Biology 10: 1142-1154.
genetic variation, and their use in population management. Pages 87- 61. Wright, S. 1931. Evolution in Mendelian populations. Genetics 16: 97-
123 in Viable Populations for Conservation. M.E. Soule, editor. 159.
Cambridge University Press, Cambridge, England. 62. ———. 1951. The genetical structure of populations. Annals of
39. Leberg, P. L. 1990. Genetic considerations in the design of Eugenics 15: 323-354.
introduction programs. Trans. 55th N.A. Wildl.& National Research
Conference: 609-619.
40. Lesica, P. and F.W. Allendorf. 1995. When are peripheral populations
valuable for conservation? Conservation Biology 9: 753-760.
41. Lynch, M. 1996. A quantitative-genetic perspective on conservation
issues. Pages 471-501 in J. C. Avise and J. L. Hamrick, editors.
Conservation Genetics: Case Histories from Nature. Chapman Hall,
New York, NY.
42. ———, and R. Lande. 1998. The critically effective size for a
genetically secure population. Animal Conservation 1: 70-72.
43. McGuigan, K., K. McDonald, K. Parris, and C. Moritz. 1998.
Mitochondrial DNA diversity and historical biogeography of a wet
forest-restricted frog (Litoria pearsoniana) from mid-east Australia.
Molecular Ecology 7: 175-186.
44. Mills, L.S., and F.W. Allendorf. 1996. The one-migrant-per-generation
rule in conservation and management. Conservation Biology 10:1509-
1518.
45. Mullis, K., F. Faloona, S. Scharf, R. Saiki, G. Horn, and H. Erlich. 1986.
Specific enzymatic amplification of DNA in vitro: the polymerase chain
reaction. Cold Springs Harbor Symposium on Quantitative Biology 51:
263-273.
46. Parker, P.G., A.A. Snow, M.D. Schug, G.C. Booton, and P.A. Fuerst. 1998.
What molecules can tell us about populations: choosing and using a
molecular marker. Ecology 79: 361-382.
47. Primmer, C.R., T. Raudsepp, B.P. Chowdhary, A.P. Moller, and H.
Ellegren. 1997. Low frequency of microsatellites in the avian genome.
Genome Research 7: 471-482.
48. Ralls, K. and J.D. Ballou. 1992. Managing genetic diversity in captive
breeding and reintroduction programs. Trans. 57th N.A. Wildl.& Nat.
Res. Conf.: 263-282
49. ———, P.H. Harvey, and A.M. Lyles. 1986. Inbreeding in natural
populations of birds and mammals. Pages 35-36 in M. E. Soule, editor.
Conservation Biology: The Science of Scarcity and Diversity. Sinauer
Assoc. Sunderland, MA.
50. Rossetto, M., P.K. Weaver, and K.W. Dixon. 1995. Use of RAPD
analysis in devising conservation strategies for the rare and endangered
Grevillea scapigera (Proteaceae). Molecular Ecology 4:321-329.
51. Shaffer, M.L. 1981. Minimum viable population sizes for conservation.
BioScience 31: 131-134.
52. Shields, W.M. 1982. Philopatry, inbreeding, and the evolution of sex.
State University of New York Press, Albany, NY.
 18
Extirpated Species of Oregon and
Washington
Constance Iten,Thomas A. O’Neil, Kelly A. Bettinger,
& David H. Johnson

Introduction
Extinction is forever, but extirpation is not! Extinction is although a few individuals may occasionally still be found
an incremental process of loss from local to regional and, during the breeding season. No efforts have been made
ultimately, to global disappearance of a species. Since all to reintroduce or augment populations of the latter three
extinctions are incremental, the best place to try and species. Thus, most of the species listed here can be
conserve a species is at the local level. Causes of local considered a “short list” for either Oregon and/or
extinctions have been proposed to be the result of small Washington, each having different potential for being
to catastrophic random events that occur in populations, reintroduced to their former range.
genetics, or our environment. Events that create a loss of One additional species that was nearly extinct in the
integrity within our ecosystems have also been surmised 1940s, the short-tailed albatross, will not be covered in this
to enhance local extinction.120, 147 One to many stochastic chapter, as it breeds only on one or two islands off the
events may actually affect a species. We felt it would be of coast of Japan. Formerly, the albatross was abundant
value to give short historical reviews of those species that enough in shallow coastal waters of the North Pacific
have undergone local extirpation from Oregon and/or (probably as post-breeding foragers) that it regularly
Washington. We have provided as best as possible occurred in coastal Indian middens.49 It is unknown if it
estimates of historical ranges and abundances. But the was ever completely absent from Oregon and Washington
reader should realize that levels of interest in and waters, and in recent years as the breeding population has
knowledge of different species varied greatly in the past, increased in Japan, there have been sightings off our coasts.
and therefore the historical information on some species
may be incomplete.
What follows are short descriptions of 13 wildlife Table 1. A list of 14 species extirpated from Oregon
species that are (or formerly were) extirpated from Oregon and/or Washington.
and/or Washington. Table 1 was developed after
reviewing 593 wildlife species that occur within the Wildlife Species Oregon Washington
terrestrial, freshwater, and marine environments of the two
states. Of the 14 species listed, only one, the California Bighorn sheep Xa Xa
condor, was near the brink of global extinction before Bison X X
substantial efforts were initiated to rescue the species. Four California condor X X
species, the bighorn sheep, sea otter, sharp-tailed grouse, Sea otter X Xa
Trumpeter swan Xa Xb
and trumpeter swan have undergone reintroduction
Yellow-billed cuckoo X X
effort(s) into the region with varying levels of success.
Gray wolf X
Efforts to reintroduce the trumpeter swan have been
Grizzly bear X
limited to breeding populations, as wintering populations
Merlin Xc
have rebounded due to protection measures. The Sharp-tailed grouse Xd
pronghorn has been reduced in Oregon and extirpated Wyoming ground squirrel X
from Washington. Efforts to reintroduce them in Pronghorn antelope X
Washington have been unsuccessful. The bison has been Upland sandpiper X
brought back in limited numbers by private ranching Short-tailed albatross X X
ventures. The gray wolf, grizzly bear, and Wyoming
ground squirrel have undergone regional or statewide a = successfully reintroduced
extirpations without efforts to reintroduce them. The b = reintroduced breeding population but not re-established;
breeding population of the merlin in Oregon is considered winter population is more secure
nearly extirpated but wintering populations continue to c = breeding population likely extirpated; winter population
be present. The final two species, upland sandpiper and more secure
yellow-billed cuckoo, are also considered extirpated, d = reintroduced but not re-established

452
 CHAPTER 18: EXTIRPATED SPECIES 453

Extirpation of these species has occurred for a variety

of reasons. Some were pursued and killed for bounty
because they endangered humans and their livestock,
some were shot for their meat and hides, and others were
poisoned and trapped because they were considered
agricultural pests. Extirpation of several of the species that
were initially abundant was influenced by multiple
factors, while other species had small populations that
quietly slipped from view. Loss of habitat was a key
determinant in the declines of most of the species. In most
cases, the loss of these 13 species was in some manner or
another human caused. Conversely, re-establishing viable
populations of any of these species will require active
human management.

Species Extirpated from

Oregon and Washington
Bighorn Sheep (Ovis canadensis)
Other Common Names: Mountain sheep, California
bighorn, rimrock bighorn, Rocky Mountain bighorn.

Former Distribution
Bighorn sheep were formerly found from southeastern
British Columbia and southwestern Alberta south along northeastern Oregon.23 The first transplant of California
the Cascade and Sierra Nevada ranges into Baja California. bighorn sheep at Hart Mountain occurred in 1954, between
In Oregon, they occurred throughout much of the non- 1954 and 1996 a total of 976 California bighorn sheep have
forested region east of the Cascade Range, with Rocky been moved within, into, or out of Oregon. Currently, there
Mountain bighorn occupying the northeastern corner of are approximately 2500 California bighorn sheep in
the state and California bighorn the remainder of the Oregon in 26 herd ranges throughout the state (Van Dyke,
range. Rocky Mountain bighorn sheep were numerous in Oregon Department of Fish and Wildlife, pers. comm.).
the Wallowa Mountains, Snake River, and Grande Ronde In 1957, California bighorn sheep were re-introduced
River Canyons.8 California bighorn ranged over southeast into Washington State. Over the next 16 years, California
and south-central Oregon and through much of the John bighorns were released into 10 areas of eastern Washington
Day and Deschutes River drainages in the north-central that had been within the ranges of the original native
part of the state.24 Bighorns were extirpated from Oregon populations. In 1999, the California bighorn population
by the mid-1940s.22 was estimated at 786 in 7 herds located throughout central,
In Washington, California bighorns were probably north central, northeastern, and southeastern Washington
found scattered on the eastern slopes of the Cascade (Rice, Washington Department of Fish and Wildlife, pers.
Mountains. The last surviving populations of the comm.).
California bighorn were seen in the north-central part of Rocky Mountain bighorn sheep were reintroduced in
the state and the Blue Mountains of southeast Washington Washington beginning in 1972.65 In 1999 there were a total
in the early 1900s. Rocky Mountain bighorns were found of 237 Rocky Mountain bighorns in 3 herds in southeastern
in the northeastern and southeastern corners of Washington and 1 herd in northeastern Washington (Cliff
Washington at the edges of the major bighorn sheep Rice, Washington Department of Fish and Wildlife, pers.
populations of Oregon and Idaho. Bighorns were comm.).
extirpated from Washington by 1935.65
Discussion
Current Status and Distribution Although many bighorn sheep were probably extirpated
Bighorn sheep have been successfully re-introduced into due to indiscriminate hunting practices prior to
both Oregon and Washington. The first attempt to restore established regulations, disease is considered the principal
bighorn sheep in Oregon was the release of 23 Rocky cause of bighorn declines. Additionally, unregulated
Mountain bighorns at Hart Mountain in 1939. The grazing by domestic livestock is believed to have
transplant failed, however, with the last survivor seen in contributed to the overall decline of bighorn sheep.141
1947. Although the cause of the failure has not been Bighorn sheep are very susceptible to a variety of
established, the subspecies used may have been a factor.24 diseases. The introduction of and proximity to domestic
Successful restoration of Rocky Mountain bighorns began livestock as European settlement expanded provided the
in 1971. As of 1996, there were approximately 500 Rocky vector for many of these diseases. Bighorns have no
Mountain Bighorn sheep in 10 established herds in natural resistance to bacteria and parasites carried by
domestic stock.65
 454 Wildlife–Habitat Relationships in Oregon and Washington

Pasturella spp. is the most common type of bacteria Under the Initiative, state and federal agencies will
found in the respiratory tracts of bighorns. Certain types increase efforts to reintroduce bighorn sheep and manage
of Pasturella produce an acute bacterial pneumonia, which habitat and populations to establish new herds and
in many cases is stress-related and brought on by factors increase the size of existing herds. Data will be collected
such as severe winter weather.36, 117, 127 Domestic sheep have on bighorn sheep ecology and factors limiting population
been implicated in the transmission of Pasturella spp. to size to be analyzed and incorporated into management
bighorns.144 strategies. The area will serve as a landscape level model
Lungworm-pneumonia complex is another major for bighorn sheep restoration.51
source of mortality in bighorn sheep. Areas of high sheep
densities and poor forage can predispose animals with Bison (Bos bison)
heavy lungworm infestations to a bacterial infection which Other Common Names: buffalo, American bison, plains
causes pneumonia.144 bison, prairie bison, tatanka.
Scabies is a contagious skin disease of bighorn sheep.
Infected sheep stop eating, which leads to weight loss and Former Distribution
death from environmental influences or other disease. Prehistorically, the bison was found in the central
Outbreaks of scabies have resulted in significant grasslands and northern parklands of North America, in
population declines of both California and Rocky habitats that ranged from semidesert to boreal forest with
Mountain bighorns in California, Colorado, Idaho, suitable grazing. Bison have been known to occur
Montana, Oregon, and Wyoming. 19, 121 Scabies was throughout North America and as far north as Alaska and
responsible for a die-off of more than half of the south to Mexico and from the Cascades and Sierra Nevada
Cottonwood Creek herd in Washington state in 1988.144 ranges to the Atlantic Coast.46, 84, 141 Native bison that were
Other infectious diseases important to bighorn free roaming have never been observed in Oregon or
populations include contagious ecthyma, paratuberculosis Washington and were initially thought to be present in
(Johne’s disease), actinomycosis (lumpy jaw), foot rot, low numbers and a rare visitor to each state. However,
necrobacillosis, keratoconjunctivitis, leptospirosis, caseous according to Verts and Carraway,141 “records in Idaho,
lymphadenitis, and listeriosis.144 Oregon and Nevada indicate that the bison were
The bighorn sheep management goal for Washington widespread, although likely not abundant.” The Snake
State is to “increase bighorn sheep populations of both River valley probably served as a corridor to lead them
subspecies statewide to a level where all available habitat westward. By crossing the Snake River or by following its
within their historic distribution is filled and each herd is southern bank, they could enter Oregon and establish
self-sustaining.”144 Management in Washington centers on populations in the eastern and central region of Oregon.
several issues. Noxious weed control is important for Archeological evidence of bison occurring in the Columbia
maintaining quality forage habitat; restoration and Plateau of Oregon and Washington has been provided by
reintroduction are priorities, as several herds may require Osborne93 and Van Vuren and Bray140 who recap which
augmentation; herds in the Blue Mountains require museums contain bison remains (i.e., several bison skulls
monitoring after a Pasturella die-off in 1995; and are housed at Oregon State University’s Horner Museum).
coordination and cooperation with tribes will be necessary Other written and verbal accounts of bison occurring
to manage potential tribal hunting impacts.145 in Oregon and Washington have been found in the
Oregon is attempting to increase bighorn sheep Washington Historical Quarterly as reported by Kingston68
numbers as well as broaden the genetic diversity of the and Pacific Northwest Quarterly as reported by Haines,45
California bighorn sheep in the state and throughout its and are as follows:
range, through cooperative efforts with other states and • In February 1875, Professor O.C. Marsh noted that “the
provinces. Trapping and transplanting are used to control most western point at which I have myself observed
population size in established herds thought to be large remains of buffalo was in 1873 on Willow Creek, Eastern
enough to allow removal without impacting population Oregon among the foothills on the eastern side of the
health. A ram to ewe ratio of 1:1 is maintained in those mountains.”
herds through controlled hunting.24
The Hells Canyon Initiative is a long-term project to • The drying of Malheur Lake in 1930 disclosed the
restore Rocky Mountain bighorn sheep to the over 5 skeletal remains of more than 40 buffalo scattered over an
million acres of the Snake River Drainage of Washington, area of 2,000 acres. It was in this region that Peter Skeene
Oregon, and Idaho. 51 The Initiative represents a Ogden saw evidence of buffalo in 1826.
cooperative effort among agencies, organizations, and • Regarding the Columbia River region, David Thompson
many private individuals with formal agreement between in 1811 noted he was told by Indians that he met at the
the state fish and wildlife agencies, the U.S. Forest Service, confluence of the Snake and Columbia Rivers that they
the Bureau of Land Management, and the Foundation for could go 3 days to where the buffalo were. This would
North American Wild Sheep. The overall goal of the mean that the buffalo came within 75 to 100 miles of the
Initiative is to “restore self-sustaining bighorn sheep herds Snake River.
to suitable habitat in the Hells Canyon area.”
 CHAPTER 18: EXTIRPATED SPECIES 455

Discussion
The decline of the bison in Oregon and Washington
appears to have been caused by a combination of events,
but primarily drought, hunting, and overexploitation.
Other sources of bison mortality include severe weather
and on occasion drowning. Today, there are only a few
instances of bison hunting. In Montana, along the northern
boundary of Yellowstone Park, bison are viewed as
problematic and can be shot when they leave the park. In
Utah, bison are considered a game species and can be
legally hunted.
Currently, bison are making a comeback primarily with
the support of private interests. Game ranching of bison
is profitable, and thus a viable commodity that helps
expand their numbers and locations. Reintroducing bison
to the wild, however, may be somewhat problematic
because they show little respect for traditional livestock
fencing and are known to carry brucellosis. Brucellosis
may cause abortions in bison and cattle, which is against
economic interests of the cattle industry.84 However, Native
American people have a strong connection with bison, and
reintroduction for cultural reasons is given careful
consideration. Thus, the subject of reintroduction is likely
to continue. Finally, bison can coexist with other ungulates
• In Washington, George Gibbs, who accompanied the (like deer and elk) on the same range because there appears
Stevens surveying party of 1853 as a geologist and to be very little interspecific competition between them
ethnologist, said that an Indian hunter told him that a lost due to different habitat use and food habits.83 Therefore,
buffalo bull had been killed 25 years before in the Grand we may see more bison in the future in Washington and
Coulee. Further, Dr. Charles Pickering who accompanied Oregon.
the party of Lieutenant Johnson of the Wilkes expedition
in 1841, which crossed the Cascades and visited Colville, California Condor (Gymnogyps californianus)
Lapwai, and other places mentioned that “a single instance Other Common Names: condor, California condor, spirit
was on record of a stray animal having been seen in the bird.
vicinity of Colville.” Mr. W. W. Lewis stated that years
ago older Spokane Indians told him that their fathers Former Distribution
surrounded and killed the last buffalo in the Spokane California condors, although never abundant in recent
Valley somewhere near the Idaho border. He figured that centuries, were historically reported as year-round
the time of reference was about 1810 to 1820. Also, the residents from British Columbia south to Baja California,96
Spokane Indians told Duncan McDonald that older and were thought to have nested as far north as southern
members of their tribe killed a number of buffalo north of British Columbia.146 The condor is considered a relic of
Moses Lake or the Grand Coulee. the ice age, and formerly was very widespread—bones
were found from New York and Florida—but well prior
Current Status and Distribution to European discovery of the continent its range had begun
The bison had been extirpated from Oregon and to shrink westward 31.
Washington prior to European settlement of the region. Bones of at least 63 California condors were found at
Today, bison occur in isolated populations usually within the “Five Mile Rapids” archeological site along the
a park, preserve, or on private ranches. Most free ranging Columbia River, five miles east of the Dalles, Oregon.
herds occur in Canada in either Mackenzie Sanctuary or Carbon-14 dating made it possible to date the remains
Wood Buffalo National Park, and in the United States in between 10,000 and 7,500 years before present. According
Yellowstone National Park or the National Bison Range.84 to Miller,84a the birds were attracted to the site by the
By 1903, there were thought to be less than 1,700 bison presence of abundant living and carcasses of salmon and
in all of North America and most of these were in zoos or human refuse resulting from fishing. Another bone
private herds. 42 By 1983, they had flourished from fragment positively identified as that of California condor
relocation and other conservation efforts to more than was found in an Indian shell mound, 6 miles north of
75,000.61 As of 1995, there were about 2,000 bison in 15 Brookings, Coos County, Oregon.84b
private ranches in Oregon.78 As of 2000, there were 750- Some of the earliest records of condors found in the
800 bison on 10 ranches in Washington (Jones, Northwest Pacific Northwest were those in the published journals of
Bison Assoc., pers. comm.). Lewis and Clark and of Patrick Gass, made while on their
expedition to the lower Columbia River. A photostat of
the original journal of Meriwether Lewis for February 17,
 456 Wildlife–Habitat Relationships in Oregon and Washington

The spring of 1827 was severe, and much snow had

fallen. The consequence was that many horses died at
Fort Vancouver, and we were visited by the various
species of beasts and birds of prey that abound in that
country. Most conspicuous among these were the
California vulture. This magnate of the air was ever
hovering around, wheeling in successive circles for a
time, then changing the wing as if wishing to describe
the figure 8; the ends of the pinions, when near enough
to be seen, having a bend waving upwards, all his
movements, whether soaring or floating, ascending or
descending, were lines of beauty. In flight he is the most
majestic bird I have seen. One morning a large
specimen was brought into our square, and we had all a
hearty laugh at the eagerness with which the Botanist
[Douglas] pounced upon it. In a very short time he had
it almost in his embraces fathoming its stretch of wings,
which not being able to compass, a measure was
brought, and he found it full nine feet from tip to tip.35
Roselle Putnam,102 a young woman who immigrated
to the Oregon territories in 1843, wrote to her mother and
sister in 1852 describing her surroundings in the Umpqua
Valley. In her letter she includes a description of the
condor:
1806, published by Harris, 48 includes an excellent . . . the largest wild bird in the country is the vulture,
description of a living condor and a sketch of the head, which is only an overgrown buzzard—it only preys on
leaving no doubt as to the identity of the bird. In addition the dead carcasses—I saw one measured which I think
to sightings of the birds, several condors were captured was between ten & eleven feet from the point of one
or killed by the Lewis and Clark expedition, as shown in wing to the point of the other.
their journal entries: Suckley129 noted that:
76:vol 3 p.174
1805 October 30: mouth Wind River, some seen The Californian vulture [=California condor] visits the
1805 November 18: mouth Columbia River, one Columbia River in fall, when its shores are lined with
killed 76:vol 3 p. 232 great numbers of dead salmon, on which this and other
1805 November 30: mouth Columbia River, seen 76:vol 3 p. 259 vultures, besides crows, ravens, and many quadrupeds,
feast for a couple of months.
1806 January 3: Fort Clatsop, near mouth Columbia
River, seen 76:vol 3 p. 309 Swainson133 made specific reference to the black vulture
[=California condor] as common on “the plains of the
1806 February 17: near Fort Clatsop, one wounded and
Multnomah” [=Willamette River]. By 1850 they were
captured 76:vol 4 p. 81
rarely found north of California.69
1806 March 15: near Fort Clatsop, two killed 43:p. 203 Finley33 mentions sightings of condors in 1903 and
1806 March 28: Deer Island, Columbia River, hunters said again in 1904 by George and Henry Peck at Drain in
eagles and vultures devoured four deer 76:vol 4 p. 211 Douglas County, Oregon as well as in the report by Henry
Northwest Company fur trader Alexander Henry made Peck of a condor killed on the southern Oregon coast a
record in his journal of seeing a condor in the Willamette “number of years” earlier. Finley concluded:
Valley on January 25, 1814.63:Elliott Coues, p. 112 David Douglas,29 These records seem to show that if the California condor
a Scottish botanist, referred to the range of the California was formerly found in the region of the Columbia River
condor north of California as follows: the numbers have decreased and the last of these
I have met with them as far to the north as 49°N northern birds seem to have taken refuge in the rough
latitude, in the summer and autumn months, but mountains regions of southern Oregon.
nowhere so abundant as in the Columbia Valley This represents the last published sighting of California
between the Grand Rapids [=The Dalles, Wasco condor in Oregon and Washington.
county] and the sea. For many years precise population numbers were
Douglas30 noted in 1826 that “the Large Buzzard, so unknown, although they were considered to be a declining
common on the shores of the Columbia, is also plentiful species since the 1890s. No longer found outside of
here [=eastern Linn County, OR69]: saw nine in one flock.” California, the estimated total population dropped to 25
George Barnston, a close friend of Douglas, recounted to 30 birds by the late 1970s.139
an event which took place at Fort Vancouver:
 CHAPTER 18: EXTIRPATED SPECIES 457

Current Distribution Sea Otter (Enhydea lutris)

California condors are not currently found in Oregon and Other Common Names: southern sea otter, otter.
Washington. As of 1999, there are only 160 California
condors in the world, including 47 in the wild and 113 in Former Distribution
captive breeding facilities.97 The birds in the wild were Sea otters once occupied a nearly continuous range along
liberated at release sites in California and Arizona. the coastlines bordering the North Pacific Ocean, from Baja
California to the northern islands of Japan.14 Except for a
Discussion small remnant population in Central California, sea otters
Factors in the decline of the California condor include were extirpated from the majority of their former eastern
climatic as well as human-related activities.96 Although Pacific Ocean range between Prince William Sound,
scientists cannot pinpoint the exact reason for the Alaska and Baja California by the early 1900s.67, 104
reduction in the condor population, random, wanton In Washington, they were once found in abundance
shooting has generally been considered the single most along the central portion of the outer coast between Grays
serious cause of the condor’s decline.139 Other factors such Harbor to Point Grenville.14 Major concentrations were
as lead contamination from the ingestion of lead bullet found between Point Grenville and Grays Harbor, and at
fragments in carcasses,13 egg and specimen collection, North Head.16
poisoning (both intentional and inadvertent), eggshell Along the Oregon coast sea otters were originally found
thinning from DDT,31 and habitat modifications96 have also in areas of exploitable abundance. In 1857 Baird wrote:
contributed to their decline. The birds in the Pacific “They [sea otters] are found so abundantly near Cape
Northwest also may have been members of an isolated Mendocino [California] to Port Orford [Oregon] that
population that suffered from human exploitation. Such several companies have been organized and equipped in
exploitation occurred between 1805 and 1850, during San Francisco expressly for their capture.”129
which time at least 13 condors were killed.21, 29, 35, 69, 102
Condors do not reach breeding capability until at least Current Distribution
6 years of age. That fact, coupled with an extremely low Despite re-introduction attempts, the sea otter population
reproductive rate, laying only one egg per clutch, and not in Oregon became extinct in the mid-1970s, and otters are
always laying in consecutive years, makes it difficult for no longer found in the state.104
the condor to recover from a significant reduction in In Washington, 1999 surveys found a total population
population.146 If, as proposed by Koford,69 the Pacific of 605 ranging from Tatoosh Island off Cape Flattery south
Northwest population was a discrete one of only two or to Destruction Island. The survey results indicate that the
three-dozen individuals, the known mortality in the early population in Washington has been growing at a finite
1800s would have been enough to jeopardize it seriously.146 rate of about 11% since 1989, and is expanding
The U.S. Fish and Wildlife Service listed the California southward.60
condor as endangered in 1967. In an effort to study and
preserve the few remaining condors, birds were captured
and fitted with tags or radio transmitters to learn about
their feeding, mating, and rearing habits. After confirming
condors would lay a second or third time after losing an
egg, in 1983 biologists began removing eggs laid in the
wild to hatch in captivity to increase production. Young
birds were also caught and bred in captivity. Despite these
efforts, the numbers of condors in the wild continued to
decline precipitously to a point in 1987, when the last wild
condor was captured and brought into the captive
breeding program to preserve the species.139 In 1992, the
first condors were reintroduced into the Los Padres
National Forest’s Sespe Condor Sanctuary.
From 1992 to 1999, a total of 55 birds have been released
in California and Arizona as a “non-essential/
experimental population” under section 10j of the
Endangered Species Act. The recovery plan for condors
includes continuing releases and the investigation of
additional release sites in areas within the historical range
of the California condor.97
 458 Wildlife–Habitat Relationships in Oregon and Washington

Discussion fisheries. Potential management solutions include:

Sea otters were once plentiful throughout their range. In establishing zonal management units and the creation of
the early 1600s the Chinese were already receiving sea otter take guidelines or closures for net fisheries.75
pelts from the Kuril Islands, harvested by Ainu hunters.32
Discovered by the Russians in the 1740s at the Pribilof Trumpeter Swan (Cygnus buccinator)
Islands of Alaska, they were hunted southward until they Other Common Names: white swan, wild swan, bugler.
were all but exterminated in most of their former range.
By 1870, hundreds of thousands had been harvested for Former Distribution
their luxuriant fur, considered the finest and most valuable Historically, the trumpeter swan’s breeding range was
of any fur-bearing animal.15 The purchase of Alaska by extensive. The swan occurred from central Alaska east to
the United States in 1867 was based largely on the Nova Scotia, New Brunswick, and Newfoundland south
anticipated revenue from further fur harvests of both sea to the Carolinas, western Tennessee, northwestern
otters and fur seals. Russia, realizing that these resources Mississippi, eastern Arkansas, and Missouri, and west to
were spent, sold the Alaska territory for a mere 7.2 million Nebraska, Wyoming, southern Idaho, northeastern
dollars.104 After the purchase, sea otter harvest intensified Oregon, and possibly California.85 The breeding range
and by the early 1900s the species was nearly extinct. In shown by Banko9 is more conservative and does not
1911, when unregulated killing was finally stopped, there include Oregon, eastern Canada, or the southern United
were only a few remaining animals in 13 known States. In winter, the trumpeter swan was found from
locations.67 southern Alaska south to southern California, possibly east
Between 1965 and 1972, 708 sea otters were translocated through Arizona and New Mexico98 to the lower Rio
to unoccupied habitats in Alaska, British Columbia, Grande Valley and the Gulf Coast of Texas,10 east along
Washington, and Oregon. In Washington, despite the 55% the Gulf Coast to central Florida.105
mortality of the initial transplant, sea otters became The first and perhaps only evidence of breeding in
established and have begun to expand their range along Oregon appears to be a report in the journal Oologist of
the outer coast.59, 60 trumpeter swans at Malheur Lake in Harney County
In Oregon, the translocated population declined between 25 May and 15 June 1921.63 Apparently no swans
drastically throughout the 1970s and is now considered were known to breed in Oregon again until attempts to
extinct.59, 104 Failure of the Oregon population has been establish a breeding population at Malheur National
attributed largely to emigration and mortality, resulting Wildlife Refuge began in 1939, and the first nesting
in a population too small to assure continued population occurred in 1958.58
growth.59 In Oregon, this species was noted by multiple authors
Sea otters require habitat along exposed saltwater as common in migration and/or winter along the lower
coastlines with rocky islands and points of rock to provide Columbia River, in the Willamette Valley, and east of the
shelter. Shallow (less than 180 feet of water) shorelines Cascades in the 1800s.44, 63 A writer by the name of John
with underwater reef formations and extensive kelp beds M. Murphy, who visited Oregon in the late 1800s,
are preferred, particularly those with abundant
invertebrate bottom fauna. Areas such as these are
available along the coastline of both Oregon and
Washington.
Successful reintroduction of sea otters in Oregon may
require consideration of the source of the translocated
animals. Otters captured in Alaska are within the Aleutian
zoogeographical province, while Oregon lies within a
transitional zoogeographic province that separates the
Californian and Aleutian provinces. It has been suggested
that the more southerly habitat found in Oregon may have
been less suitable for sea otters from northern
populations. 59 Additionally, the number of otters
transplanted must be capable of reproducing at a rate
greater than that of the combined rate of emigration and
mortality to establish a successful population.59
In Washington, with an expanding sea otter
population,60 management considerations may soon
involve the need for the resolution of conflicts with
commercial, tribal, and recreational fisheries. Populations
of sea otter prey items form the basis of many commercial
and recreational fisheries, including sea urchins, razor
clams, mussels, and crabs.75 Another commercial fishery
interaction involves the by-catch of sea otters in net
 CHAPTER 18: EXTIRPATED SPECIES 459

described both whistling (=tundra swan) and trumpeter in Montana, but the first nesting did not occur until 1958.58
swans as so abundant at the mouth of the Columbia River The breeding population at Malheur NWR peaked in the
that they appeared as “cumulus clouds” in late fall.118 He early 1980s at 19 breeding pairs and a total of 77
also noted that both swans were “exceedingly common” individuals. Between 1990 and 1999, the number of young
in the Klamath Lake region, and along the Willamette, fledged each year ranged from 3 to 18, and counts of fall
Columbia, and Snake Rivers. In 1891, Thomas G. Farrell adults ranged from 19 to 75 birds (the count of 75 was
wrote of an interview with an old resident of Seavies Island inflated by the release of 52 birds in 1992). Numbers have
(=Sauvie Island) who described “immense flocks” of been steadily dropping since 1992. In 1991, an effort was
tundra and trumpeter swans that formerly visited begun to establish a separate breeding population at
Columbia and Multnomah Counties.63 He goes on to say Summer Lake Wildlife Area in Lake County with birds
that “Of late years very few of these birds [trumpeter from Montana, Idaho, and Malheur NWR.58 In 1991, birds
swans] visit the valley of the Columbia, and….it has never occurred only in Harney County in the summer, whereas
been my good fortune to meet with a member of the in 1999, trumpeter swans were summering in Harney,
species.”63:T. G. Farrell p. 132 This decline in abundance is echoed Lake, Klamath, Crook, Grant, and Baker Counties. In
again more than a decade later by John Minto, a pioneer addition, 4 juvenile swans were recently released on the
who arrived in the Willamette Valley in the 1840s and Deschutes River in Bend as part of a plan to replace
noted that waterfowl populations in general had breeding exotic mute swans with the native trumpeter
plummeted in numbers by the early 1900s, such that (Ivey, U.S. Fish and Wildlife Service, pers. comm.). The
newcomers to the area hardly believed accounts of the birds in eastern Oregon winter locally, sometimes
earlier abundance.118 By the time Gabrielson and Jewett wandering elsewhere in Oregon, northeastern California,
published Birds of Oregon in 1940, they considered the or western Nevada.57 The most regular winter location for
species gone from Oregon and stated that there had been trumpeter swans in western Oregon is in western Polk
“no authentic records for many years.” County, where a small flock has occurred since the mid-
The only evidence that the trumpeter swan may have 1980s.44 In recent winters, small numbers have also been
formerly nested in the state of Washington is a report by reported between late October and March along the
C. F. Yokum of an account by an “old settler” of breeding Columbia River in Multnomah, Columbia, and Clatsop
at Cherry Lake in Whitman County until about 1918.9 No Counties.
nesting was confirmed until birds were introduced at In Washington, 6 trumpeter swan cygnets were
Turnbull National Wildlife Refuge in Spokane County in introduced to Turnbull National Wildlife Refuge in
1963 and successful nesting occurred from 1967 to 1969.122 Spokane County in 1963.122 Nesting was attempted in 1966
In migration and winter, trumpeter swans were and successful from 1967 to 1969. No mated pairs or
reported by Suckley in 1860 to be more abundant on the nesting occurred again until 1994 when a pair was again
Columbia River than in Puget Sound.62 He describes seen, but as of 1997 that pair had not successfully bred.
immense flocks of swans along the shores of the river and The only other recent report of a trumpeter swan in
spread out along the margin of the water for a distance Washington in summer was of an adult seen 17 June 1991
varying from an eighth to a quarter of a mile. By the 1920s, at Calispell Lake in Pend Oreille County. 122 Trumpeter
the only known trumpeter swans in the state were a small swans winter in Skagit and Snohomish counties (north
flock of about 18 birds that for several years moved from Puget Sound) where high counts include 100 at Skagit Flats
wintering areas in British Columbia into Okanogan on 31 October 1996, 160 at Stanwood in Snohomish County
County to winter somewhere in eastern Washington or on 16 November 1996, and 480 at Swan Reserve in Skagit
Oregon.62 County on 14 February 1998.106, 111 Outside of the typical
wintering areas, individual birds or groups of up to 10
Current Status and Distribution are on occasion reported from Island, Grant, Clallam,
Currently, the natural breeding range of the trumpeter Clark, Benton, Jefferson, Walla Walla, Klickitat, Kittitas,
swan is much reduced. The Pacific Coast population, Pacific, Cowlitz, Grays Harbor, and Thurston Counties.106,
which breeds in Alaska and winters from southern Alaska 107, 108, 111, 113, 114, 115, 116

south to Washington and Oregon, numbered about 13,000 The trumpeter swan is not listed as an endangered,
individuals in 1990.85 The Rocky Mountain population threatened, or sensitive species in Oregon or Washington.
which breeds in the Rocky Mountains from Yukon and In 1989, the U.S. Fish and Wildlife Service was petitioned
Northwest Territories south through eastern British to list the Rocky Mountain population (which includes
Columbia, Alberta, Montana, Wyoming, and Idaho restored flocks in Oregon) because of low numbers and
numbered about 2,200 in 1992. It includes restored flocks winter vulnerability, but the petition was denied.85
in Oregon and Nevada. The Interior population was
estimated at 629 birds in 1993 and includes restored flocks Discussion
in South Dakota, Minnesota, Wisconsin, Michigan, and The initial cause of the trumpeter swan decline was over-
Ontario. hunting for food, feathers, and skins.9, 85 Factors negatively
Attempts to re-establish a breeding population at affecting trumpeter swans include lead poisoning from
Malheur National Wildlife Refuge in Oregon began in 1939 consuming lead shot or fishing weights, collisions with
with birds from Red Rocks Lake National Wildlife Refuge power lines and barbed wire fences, accidental shooting
 460 Wildlife–Habitat Relationships in Oregon and Washington

by hunters mistaking the birds for other species of swans as they were described by John K. Townsend as abundant
during hunting season, illegal shooting, and competition in the area of present day Vancouver, Washington in the
with the exotic and more aggressive mute swan.9, 85 In 1830s.64 In 1916 R. Bruce Horsfall described yellow-billed
Oregon, the greatest cause of mortality between 1958 and cuckoos as “rare birds throughout most parts of Oregon.”54
1999 was due to power line collisions.58 The second most In June 1924 Ira N. Gabrielson described encountering
important mortality factor during that time in Oregon was “Twelve or fifteen birds” in an area along the Columbia
predation, but losses may be exaggerated, as many swans River near Portland, Multnomah County, where only one
were wing-clipped and flightless. Once flying age is bird had been noted in the previous 5 years.39 He further
reached, adult trumpeter swans have few natural stated that they “continued to be found there in numbers
predators, though they are occasionally taken by golden all summer.” By the 1940s, Gabrielson and Jewett 40
eagles and very rarely by coyotes.9 considered the yellow-billed cuckoo “not a common bird
Most breeding habitat is considered secure, though anywhere in Oregon” but did consider it fairly common
there are still local problems with pollution, recreation, in some past years (1920s) along the Columbia and
mining, and development.85 Swans nest in freshwater Willamette Rivers. Jewett et al.62 described it as a “rare
areas characterized by shallow, stable levels of unpolluted, summer resident in the humid Transition Zone of western
fresh water; accessible forage; room for take off Washington” and noted earlier reports from the 1920s and
(approximately 100 m); emergent vegetation, muskrat or 1930s that described the cuckoo as “fairly plentiful” near
other platforms for nest sites; and low human Lake Washington and “a regular summer resident,
disturbance.85 Similar areas of ice-free water are used in although not numerous” in Whatcom County.
fall and winter by migrants and wintering birds. The In Washington, the yellow-billed cuckoo was formerly
inadequate quality and quantity of winter habitat is an uncommon breeder throughout the Puget Trough and
considered one of the most important limiting factors in along major river systems in eastern Washington122 and
trumpeter swan restoration. Most winter habitat has been considered extirpated as a breeder by 1934.56 In Oregon, it
lost to urban development. was formerly an abundant or common breeder along the
The trumpeter swan often responds well to Columbia River west of the Cascades, and in the
reintroduction efforts, and active restoration projects to Willamette and Rogue River Valleys.44, 56, 81 It also occurred
former breeding areas in Minnesota, Wisconsin, Michigan, along major river systems such as the Grande Ronde,
and Ontario, as well as Oregon, are ongoing. 85 An Umatilla, Columbia, Owyhee, and Snake in eastern
additional challenge related to reintroduction that needs Oregon, and was considered extirpated as a breeder by
to be addressed is the loss of migratory traditions in some 1945.56
flocks.
Trumpeter swans are long-lived—up to 24 years in the Current Status and Distribution
wild and 32 years in captivity—and they do not typically Since the 1930s, individual birds have been reported in
begin to breed until 4-7 years old.85 Washington from Grays Harbor, King, Snohomish,
Whatcom, Okanogan, Douglas, Grant, Franklin, Benton,
Yellow-billed Cuckoo (Coccyzus americanus)
Other Common Names: rain bird, rain cuckoo, kow-kow.

Former Distribution
Some authorities recognize two subspecies; others
consider the species to be monotypic.56 Coccyzus americanus
occidentalis, the California cuckoo, formerly bred from
southwest British Columbia, Washington, Oregon,
northern Utah, central Colorado, and west Texas south
and west to southern Baja California, Sinaloa, and
Chihuahua in Mexico. Coccyzus americanus americanus is
found breeding in the remainder of the range throughout
eastern North America south to eastern Mexico, the
Greater Antilles, and possibly Central America and
northwestern South America. The northern range limit is
roughly equivalent to the northern tier of U.S. states, with
populations extending into some areas of extreme
southern Canada. Yellow-billed cuckoos winter in South
America as far south as northern Argentina.2
Though common and widespread in the east, western
populations of yellow-billed cuckoos probably always had
a limited distribution, as they were mostly restricted to
deciduous forested lowland riparian habitats. They may,
however, have been locally abundant in favorable habitat,
 CHAPTER 18: EXTIRPATED SPECIES 461

and Walla Walla counties. The most recent confirmed 1950s was the proverbial final nail in the coffin.41 The
sighting is from Elma, Grays Harbor County on 3 August yellow-billed cuckoo is a 100% insectivorous bird that
1996 (Rogers, Washington Ornithological Society, pers. feeds mainly on large insects such as caterpillars
comm.). No breeding has been confirmed in Washington (including forest tent caterpillars), katydids, cicadas, gypsy
for more than 60 years. moths, grasshoppers, and crickets.56 These larger insects
In Oregon, there were only two records between 1940 typically have cycles of abundance from year to year and
and 1970, both from La Grande, Union County.44 Since 1970 as a result, cuckoos are thought to be nomadic just prior
there have been more than 40 records of individual birds to breeding as they appraise local food resources. Areas
or pairs from Union, Harney, Malheur, Klamath, which one year had numerous nesting pairs may have
Deschutes, Jefferson, Umatilla, Wallowa, Baker, Lake, few or none the next, depending on food resources. This
Linn, Clackamas, and Multnomah counties. All records ties back to the available amount of habitat, in that
have occurred between 20 May and 12 September, with protecting a limited area of riparian forest may not be
most in June and July. The most recent records are from adequate to sustain cuckoo populations in a region, since
Malheur National Wildlife Refuge in May and June 1999.126, they need to be able to wander and find not only areas of
131
Since the 1940s, no breeding was confirmed in Oregon suitable habitat, but habitat with an abundance of cyclical
until 1992, when a nesting pair was found in La Grande, insects that season.
Union County. A female with a brood patch hit a window Changes in habitat on the wintering grounds are not
in Bend, Deschutes County on 18 June 1990. considered an issue in the yellow-billed cuckoo’s decline
The western population was formerly listed as a because their preferred winter habitat is the shrubby
Category 2 (candidate for listing), but as of 1990 the second-growth that results from cutting mature and old-
western population was listed as a Category 3B (former growth forests.
federal candidate for listing) because it was not considered No specific measures are being taken to improve habitat
a valid subspecies.37, 56 The U.S. Fish and Wildlife Service conditions for cuckoos in Oregon and Washington at this
(USFWS) was petitioned again in 1999 to consider listing time, although they may benefit from efforts to restore
the cuckoo as endangered, and made a determination in riparian areas along major lowland river systems for other
February 2000 that listing the western subspecies may be resource goals if such projects result in the formation of
warranted (Harvey, U.S. Fish and Wildlife Service, pers. extensive stands of cottonwood and willow, and the
comm.). A status review was begun and the USFWS is preservation or restoration of seasonal flooding patterns
currently funding DNA research to determine the validity that promote sloughs and oxbows.
of the subspecies. As of 1999, the yellow-billed cuckoo is
listed as a Species of Special Concern in Washington, and Species Extirpated from Oregon
as a Species of Special Concern—Critical in Oregon.
Gray Wolf (Canis lupus)
Discussion
Other Common Names: grey wolf, timber wolf.
Several factors have been implicated in the decline of the
yellow-billed cuckoo in the western U.S., but the most Former Distribution
important one is habitat loss. Cuckoos in the western U.S. Wolves were once found in all habitats of the northern
breed in extensive deciduous riparian thickets or in hemisphere, with the exception of tropical forests and arid
cottonwood and willow forests. Areas near backwaters deserts.90 In western North America, wolves ranged from
such as sloughs and oxbows are particularly important. Alaska south through Canada to northern California.141
Size of the habitat patch appears to be crucial as well, as The northern Rocky Mountain gray wolf subspecies (C. l.
very few cuckoos were found in areas where suitable irremotus) was listed as endangered by the USFWS in 1973;
habitat was less than 100 m wide and less than 10 ha in the entire species, C. lupus, was listed as endangered in
size in California.41 Laymon and Halterman72 speculate the conterminous United States, except Minnesota in 1978.1
that cuckoos may require intact riparian woodlands of In 1857 Newberry reported:
more than 40 ha in size for breeding in California. The
Though less common than the “coyote,” the large gray
strict ties to river bottoms, sloughs, damp thickets and
wolf is found in all the uninhabited parts of California
swampy places in the west appears to be tied to a
and Oregon. Very few were seen by members of our
requirement of high humidity needed to keep cuckoo eggs
party, none were killed; and we had everywhere
from desiccating; deciduous forests in the eastern U.S.
evidence that this species was much less numerously
where the species probably evolved are consistently
represented on the Pacific Coast than on the Upper
humid in the summer.41 Activities that have contributed
Missouri. In the Cascade Mountains we saw tracks of
to the loss of floodplain forests include channelization of
some of these wolves of most portentous size. All the
river systems, water withdrawal for irrigation, dam
large wolves seen by any of our party were gray, and all
building, flood control projects, livestock grazing, and
the skins, which I saw in possession of Indians or
clearing of areas for firewood, agriculture, and
whites were also gray, and it is probable that the white
development. It is thought that once populations were
and black varieties are never found in California. On
stressed because of lack of suitable habitat, the widespread
the upper Columbia, in Oregon and Washington
spraying of DDT and other pesticides in the 1940s and
Territories, where the wolves are more numerous and
 462 Wildlife–Habitat Relationships in Oregon and Washington

Currently, the only known wolf packs in the greater

Pacific Northwest region are in British Columbia in the
Elaho River drainage, near Squamish, B.C., in central
Idaho, and in Yellowstone National Park in Montana.71

Discussion
Prior to the introduction of the horse in the 1700s, the
human population was considered to have been “too low”
to exert much pressure on wildlife populations.101 Some
native peoples of the Pacific Northwest region revered the
wolf and had a taboo against killing them. Other tribes
hunted wolves for their pelts for use in robes or blankets.
In total, pressure on wildlife populations in the area was
minimal. With the arrival of the horse, the impact on
wildlife increased significantly, as native hunters were able
to travel longer distances and transport more game.71 In
addition to intensive settlement by Europeans, the
establishment of the Hudson’s Bay Company in the area
to “. . . make a profit for the stockholders by means of
exploitation of the fur trade. . . .” initiated the decline of
the wolf population. Between 1827 and 1859, 7,761 wolf
pelts were traded at four posts in the Cascades area, from
Thompson River, B.C. south to Fort Vancouver on the
Columbia River, and east to Fort Colville. An additional
8,234 wolf pelts were traded at Fort Nez Perce at the
the winters are colder, the same variations occur which
confluence of the Walla Walla and the Columbia Rivers.71
are common on the upper Missouri.88
As settlers moved into the area, livestock depredation
The “Cascades wolf” Canis lupus fucus was historically by wolves became a major concern. In 1841, Admiral
found throughout the Cascade Range in both Washington Charles Wilkes wrote about a wolf problem at Fort
and Oregon. It was described as a medium-sized wolf, Vancouver, which necessitated bringing in from the field
with “a dark cinnamon or cinnamon-buffy” coloration “large numbers of cattle. . . for in consequence of the
with the dorsal area “profusely overlaid with black” and numerous wolves that are prowling about; in some places
was referred to by early white settlers as “red legs.” Lewis it becomes necessary for the keeper to protect his beasts
and Clark were the first white men to record the Cascades even in the daytime.”150 In 1843, the first “wolf meeting”
wolf, in 1805-06. They referred to it as the “large brown was called and a $5.00 bounty on wolves was established.71
wolf” said to inhabit “California and the banks of the By 1871, the Washington Territorial Predator Control
Columbia” river.150 Act provided bounties on most predators in the
The Cascades wolf is considered to have become extinct Washington territory. In 1915, the Department of
throughout its former range in the northwestern United Agriculture designated federal funds for “predator
States by 1977.38, 99 In Oregon, wolves were considered to control” on national forests and other public lands. In 1916,
have been extirpated by 1972.92 the U.S. Biological Service expanded the program to
include private lands. Records from Washington State in
Current Status and Distribution
1924 described the program:
Today, wolves are found in less than one percent of their
former range in the contiguous United States. In during September, 23 federal, State and Cooperative
Washington and Oregon, there have been a number of hunters worked a total of 653 days. We expect every
sightings of wolf-like canids in the last 25 years. In man to put out all the poison baits possible. . . to cover
Washington, no recent live sightings have been confirmed the greatest possible area. . . to use it in all sections
as a wild wolf, and to date neither an identified pack nor where the work can be carried on without endangering
mating pair has been found. Two wolves found dead in dogs and domestic animals. . . our first duty is to get
the Selkirk Mountains of northeastern Washington were rid of all the predatory animals possible.7
confirmed to have strayed from the re-introduced 9-mile The last bountied wolf in Oregon was recorded in 1946.
and Magic packs of Montana (John Almack, Washington Since that time only two specimens have been collected,
Department of Fish and Wildlife, pers. comm.). However, one in 1974 in southern Baker County and one in 1978 in
there have been several documented family units of eastern Douglas County. In Washington, the U.S. Biological
wolves in Washington in the last 12 years, indicating some Service records indicate that only 2 wolves were taken by
level of reproduction.1 In 1999, a wolf was captured in hunters between 1915 and 1929.137 The most recent
Oregon after traveling hundreds of miles from the Nez confirmed wolf sighting in Washington was recorded in
Perce Indian reservation, where efforts are being made to 1975. Identified as a British Columbia wolf Canis lupus
establish packs of Canadian wolves in central Idaho. columbianus, it was killed in Douglas County, after it had
 CHAPTER 18: EXTIRPATED SPECIES 463

killed three calves. The Washington State Department of Current Status and Distribution
Game concluded in their 1975 Status Report that: Grizzly populations are not threatened in Canada or
wolves may still be present in Washington, but that Alaska, but in September 1975 the grizzly bear was
presence is represented by very small numbers, if at all. designated as “threatened” south of the Canada border.
The wolf killed in Douglas county may have been a As of 1999, grizzly bear populations could be found in 6
wild animal, but Canadian authorities cannot verify the disparate ecosystems in Idaho, Montana, Washington, and
possibility of its coming from adjacent ranges in British Wyoming.95 More specifically, they are found in the North
Columbia.142 Cascades of Washington, Selkirk Mountains of Idaho,
Cabinet-Yaak of Montana, North Continental Divide of
Currently, there are no efforts being made to re-
Montana and Canada, Bitterroot Mountains of Idaho and
introduce wolves to Washington or Oregon. Both
Montana, and the Greater Yellowstone of Idaho, Montana,
Washington and Oregon are considered within the
and Wyoming. Critical habitat for the bear in the lower 48
Northern Rocky Mountain wolf recovery plan area. The
states was designated in November 1976 and includes all
plan is currently focusing on re-introduction efforts in
of the current bear population centers except the Bitterroot
Yellowstone National Park. Budget constraints allow for
Mountains.
little more than investigation of sightings outside the
Grizzly bears have persisted in the lower 48 states only
recovery area (Almack, Washington Department of Fish
in areas where extensive remote habitat has prevented or
and Wildlife, pers. comm.).
reduced human-caused mortality. Typically, these areas
are protected by the Wilderness Act of 1964.26 Population
Grizzly Bear (Ursus arctos)
estimates have been very difficult to obtain, but based on
Other Common Names: brown bear, big brown bear, griz,
a long-term study conducted by Craighead et al.27 the
grisly, silvertip, white bear, Moccasin Joe.
Yellowstone population varied from 172 to 309 (1 bear per
Former Distribution 80.3 km2).
Grizzlies once roamed most of western and central North
Discussion
America and could be found from the Arctic Ocean to
Grizzly bears occur in a variety of habitats from alpine to
Mexico and from the prairies of the Great Plains to the
lowland valleys. They use early seral stages, riparian
Pacific coast.95 There is also some evidence that the earlier
habitats, grasslands, and shrublands, but mostly live in
range of grizzlies may have extended through parts of
rugged, largely inaccessible areas. Travel corridors are
eastern North America.141 The historical distribution of the
important to grizzly bear conservation because the current
grizzly bear is best summarized by Rausch.103 Since the
population centers are disjunct. The primary factors
settlement of Europeans, the range of the grizzly bear has
affecting population declines are loss of habitat, loss of
been steadily retreating, especially from the south and
food sources, and direct mortality. Historically, grizzlies
eastern portions of the United States.26 The greatest
were killed by humans defending their property and
number of bears could be found in the western portions
personal safety. They were also trapped and hunted.
of the United States and Canada. Along the Pacific coast,
the grizzly bear used to range from Alaska to Mexico with
viable populations still existing only in Alaska, Canada,
and a small population in the interior of Washington. In
the southern portion of the Pacific coast, the grizzly once
roamed through Mexico, California, and Oregon. Most of
the grizzly population in Mexico was thought to have been
removed early, suggesting that the population dynamics
in that area were weak. However, grizzly bear populations
in California appeared viable, with a population estimated
at 10,000; nonetheless, all had vanished by 1924.26
The grizzly bear has been extirpated from Oregon for
almost 70 years, with the last verified animal shot on
Chesnimnus Creek in Wallowa County on 14 September,
1931. Earlier accounts of the grizzly bear in Oregon suggest
that the species occurred throughout the state prior to
European settlement in the Cascade Range, Siskiyou, Blue,
Steens, and Wallowa mountains; the Klamath, Rogue,
Umpqua, and Willamette valleys; and some of the high
desert east and south of Bend.8, 141 The subspecies that used
to occur in Oregon was Ursus arctos horribilis.103 Grizzly
bears also ranged over most of Washington. Today, a small
population remains in the Northern Cascades, but its
viability is unknown.
 464 Wildlife–Habitat Relationships in Oregon and Washington

Sullivan130 discussed early trapping activities and noted

that 382 grizzly bear hides passed through Fort Colville
in 1849. However, Verts and Carraway’s141 review of the
species in Oregon found no population estimates prior to
their extirpation from the state. Only anecdotal
information exists that suggests that populations were
scarce by the early 1900s.8
The principal challenge in managing the grizzly bear
is that large areas are required that involve multi-agency
jurisdictions. Therefore, different mandates, laws, and
concerns arising from various federal, state, and private
organizations transcend the local politics of an area. Often
the biggest concern revolves around personal safety,
specifically when there is a potential for bears to interact
with humans. The bears are long lived, have a low
reproductive rate, and the cubs can spend several years
with their mothers. Thus, direct mortality of a small
number of bears can greatly influence the population
dynamics of an area. Reintroducing grizzlies to Oregon is
not without its own unique challenges, but is possible, as
the state does have some large areas that are still remote.
Likely places to reintroduce the grizzly bear in Oregon
include the John Day Wilderness, or Hells Canyon
Wilderness and Recreational Area—Eagle Cap Wilderness
ecosystems. these nests suggested that at least one was not in a cavity;
Gleason, pers. comm.). One nest was located in the
Merlin (Falco columbarius) Puyallup Valley of Pierce County; no location was given
Other Common Names: pigeon hawk, pigeon falcon. for the other. Black merlins were reported on several
occasions hunting house sparrows and harassing pigeons
Former distribution in the cities of Portland and Seattle.40, 62
The merlin is a circumboreal species breeding in Alaska, The taiga merlin was first reported as nesting in Oregon
most of Canada, parts of the northwestern and in 1857 near Klamath Lake.40 Gabrielson and Jewett40
northeastern U.S., Britain, Scandinavia, Finland, Iceland, describe the taiga merlin as “Very rare breeding bird and
Siberia, and north central Russia.123 North American uncommon migrant and winter resident in eastern
populations winter in southwestern Alaska, coastal Oregon. Very rare straggler west of Cascades.” They
western Canada, western and southern U.S., south into hypothesized that the species was more common than
Panama, West Indies, Central America, northern South reports indicate because of the potential difficulty in
America, Venezuela, Colombia, Ecuador, and northern distinguishing it from young sharp-shinned hawks. In
Peru. Washington, Jewett et al.62 describe the taiga merlin as a
Of the three subspecies that occur in North America, “Permanent resident in eastern Washington, rare migrant
two breed in Washington and one formerly bred in west of the Cascades.”
Oregon.44, 123 Falco columbarius columbarius (taiga merlin)
is a rare breeder in high elevation forests of the north Current Status and Distribution
Cascades and northeastern Washington, and formerly The merlin is still found throughout its former range (see
bred in eastern Oregon; F. c. suckleyi (black merlin) is a above) and in past decades actually expanded its range in
rare breeder along the outer coast, Hood Canal, and Puget the northern Great Plains and in New England.123
Sound areas of Washington. 44, 122 The black merlin is In Oregon, there are no confirmed breeding records
considered mostly sedentary while the taiga merlin is from recent decades, but there are regular reports of
mostly migratory. merlins in the summer (June-July) from the 1970s, 1980s,
In Oregon, Gabrielson and Jewett40 considered the black and 1990s, including at least 33 sighting reports during
merlin a “Rare winter resident of [the] coast” and “only the 1995-1999 Breeding Bird Atlas effort44 (Adamus,
of casual occurrence inland.” In Washington, Jewett et Oregon State University, pers. comm.). The breeding
al.62 considered the black merlin a “permanent resident season reports come from throughout most of western and
coastwise and in tide flats and open prairie country of the eastern Oregon, lending uncertainty to the theory that only
Sound region of western Washington; in fall and winter the taiga merlin breeds or bred in the state. One of the
some individuals wandering to points east of the Cascade strongest pieces of evidence of possible breeding in recent
Mountains.” Only 2 confirmed nests were known at the years comes from Potamus Point in Morrow County where
time in Washington, both in cavities high in dead trees a bird appeared to be on territory 10-11 July 1998.125 In
(although a subsequent review of the original reports on addition, there are 6 unconfirmed reports by different
 CHAPTER 18: EXTIRPATED SPECIES 465

observers of possible nests: a cavity nest near Klamath Canada, and tend to be easier to see as they occupy more
Falls, date unknown; a stick nest and a cavity nest on the habitats than would be used for breeding.
upper Metolius River, date unknown; a nest under loose As with the peregrine falcon, DDT use caused eggshell
bark near Reedsport in 1966; a cavity nest near Cottage thinning in the merlin, and studies in the 1960s in the Great
Grove in 1972; and a cavity nest near Roseburg in 1988 or Plains and Canada documented a 30% decline in merlin
1989 (Fenske, pers. comm.). Merlins are more common in reproductive success when compared to pre-1950 data.123
Oregon in winter, as migrants and wintering birds move Though most merlin populations are currently thought
into the state. They can be found throughout the state, to be reproducing well, a study of eggs collected between
but are most regular near major estuaries, lakes, reservoirs, 1980-1988 in the Canadian Prairie Provinces showed that
and occasionally in cities where food supplies are reliable.25 35.7% of the eggs still contained levels of DDT likely to
Unusually high counts were reported from two Audubon negatively affect reproductive success.89 Though no merlin
Christmas Birds Counts (CBC) in December 1997 when 6 eggs have been tested in Oregon or Washington, eggshells
birds were tallied on the Sauvie Island CBC in Multnomah from some peregrine falcon nests in the region still contain
County and 4 birds were tallied on the Dallas CBC in Polk levels of DDT that warrant concern, and some pairs
County.136 continue to lose their clutches by sitting on and breaking
In Washington, up to 13 nests and an additional 5 the eggs (Pagel, U.S. Forest Service, pers. comm.). A sample
territorial pairs of the black merlin were found in the 1980s of 350 eggshell fragments collected in recent years in
and 1990s in Skagit, Snohomish, Clallam, Grays Harbor, Oregon, Washington, and northern California were a mean
San Juan, and Jefferson Counties122 (Gleason, pers. comm.). 17.4% thinner than eggshells from pre-DDT use (Pagel,
Of 9 nests monitored in the late 1990s, all were in old corvid U.S. Forest Service, unpubl. data). The greatest thinning
nests and 8 were in Douglas fir or western hemlock trees occurred in southwestern Oregon and northwestern
with diameters ranging from 24 inches to 13 feet (Gleason, California where eggshells were a mean 24% thinner.
pers. comm.). There are two records of possible breeding Given that merlins and peregrines have somewhat similar
by taiga merlins during the Washington Breeding Bird ranges, migratory habitats, and diets (both are bird eaters)
Atlas period (1987-1996): an adult seen north of Mt. Adams in Oregon and Washington, it is possible that merlins also
in Yakima County flying over old-growth forest and a pair could still be experiencing eggshell thinning.
seen in Stevens County122 (Tom Gleason, pers. comm.). A In most areas of their range, merlin populations
third record was reported of an adult at Sanpoil, Ferry currently appear to be increasing, with the possible
County on 19 June 1997.109 As in Oregon, merlins are more exception of the northeastern U.S., where acid rain is
common and widespread in Washington during winter thought to be a factor.123 In the prairie regions, merlins are
when birds from the north move into the region. declining or have disappeared from areas where nest trees
As of 1999, the merlin has no federal or Oregon State are being lost from woodlots and shelterbelts and where
status, but in Washington the breeding population is vegetation is cleared from around prairie potholes, but
considered a Species of Concern—Candidate. the decline may be offset by increases in nesting
populations in some cities in the region.123 It is unknown
Discussion what the effects might be from habitat loss and change on
Merlins hunt a variety of small to medium-sized birds, wintering grounds in Central and South America.
and also will feed to a lesser extent on larger flying insects Documented threats to breeding populations in Oregon
such as dragonflies, and small mammals and reptiles.123 and Washington—at least for those birds not nesting in
They do not build nests of their own; instead they use old protected areas such as the Olympic National Park—
crow, raven, hawk, or magpie nests, tree cavities, or cliff include direct persecution and nest tree loss. Of the 9 nests
ledges. At least in western Washington, merlins seem to monitored by Tom Gleason in western Washington in the
prefer “waterfront property” and nest near lakes, rivers, 1990s, adults at two nests were shot and an additional two
and along the Puget Sound, perhaps because these areas nest trees were cut down by landowners. All 9 nests were
provide breaks in otherwise continuous forest and/or on private land and many were near homes in low-density
because prey species are more abundant. waterfront neighborhoods with mature conifer trees.
Washington and Oregon are at the edge of the species’
breeding range and the density of breeding pairs has Sharp-tailed Grouse
probably always been less than in core areas of their range. (Tympanuchus phasianellus)
But there are probably more breeding pairs of merlins in Other Common Names: Kennicott’s grouse, western
western and eastern Oregon and Washington than the prairie chicken, prairie sharp-tailed grouse, willow grouse.
numbers of confirmed nests indicate, particularly given
the number of sightings of birds during the breeding Former Distribution
season. One explanation for the paucity of confirmed nest At the time of European settlement, Columbian sharp-
sites is the effort it requires to locate a pair and find the tailed grouse were found throughout much of the inland
nest—it took Tom Gleason five years and many hours of Pacific Northwest, as well as locations in western
patient searching and watching to find just 8 nests (Tom Colorado, northern Utah, northern New Mexico, western
Gleason, pers. comm.). Wintering populations are more Montana, and western Wyoming.77
abundant as birds move into the region from Alaska and
 466 Wildlife–Habitat Relationships in Oregon and Washington

In the late 1800s, sharp-tailed grouse were still

abundant in both Washington and Oregon. Early settlers
in eastern Washington found the Columbian sharp-tailed
grouse to be the most abundant and best-known game
bird in the area.149 They were also abundant in the Grande
Ronde and Powder River valleys in Baker and Union
Counties of Oregon. In 1881, an article on hunting in that
area recounted:
This is the only part of Oregon where prairie chicken
[=sharp-tailed grouse] may be found, and this fall I saw
my first specimen of this beautiful bird, and as I have
always wished, this, the first chicken I ever beheld, in
its peculiar flight fell clean to my gun, and as I picked
up the noble grouse it seemed a shame to kill so
handsome a bird. . . . Great numbers of birds could be
killed here daily, but with me sport loses its charm
when it approaches slaughter, so what might have been
scores I was content to make dozens.119
Settlers in Washington harvested wagonloads of sharp-
tails in a single day in the 1880s and 1890s, presumably in
high-concentration areas such as the shrub/meadow
steppe bordering river tributaries of eastern Washington.70
By 1895, sharp-tailed grouse were declining from just
such hunting practices. In 1895, an article in the Oregon
Sharp-tailed grouse were once abundant in eastern Naturalist stated:
Washington and Oregon. Early European settlement,
this bird [sharp-tailed grouse] once so abundant on the
farming practices, and hunting pressure contributed to
plains of eastern Oregon, Washington and ‘Big bend of
their decline. By the late 1920s they had been extirpated
the Columbia,’ is slowly and surely repeating the
from at least six counties in Washington. In 1940
history of the pinnated grouse [=lesser prairie chicken]
Columbian sharp-tailed grouse were described as: “now
of the Western states. Where 15 years ago they could be
being scarce and apparently in danger of early extinction
seen in winter in flocks of fifty or more; flocks of a
in Oregon.”28 The last sighting of Columbian sharp-tails
dozen are now uncommon. Then their great enemy was
in Oregon was in the late 1960s in Baker County. They
the coyote, who robbed their nest and caught their
were considered extirpated from Oregon by the early 1970s
young. Now it is man with his traps and guns.6
(Sands, The Nature Conservancy, pers. comm.).
In 1899, L. B. Quimby in an unpublished report for the
Current Status and Distribution Oregon Department of Fish and Wildlife recommended
Sharp-tailed grouse persist in 8 scattered remnant hunting restrictions in light of the rapidly declining
populations in Douglas, Lincoln, and Okanogan counties numbers of sharp-tailed grouse attributed to over-harvest,
in Washington State. They are confined to small, severely particularly in the winter months. After the establishment
fragmented populations, some of which are located on of hunting seasons and bag limits for upland game birds,
degraded habitat. Much of the remaining potential sharp- sharp-tailed grouse experienced a short-lived recovery. But
tail habitat in Washington is threatened by further by the turn of the century populations were once again
alteration and fragmentation.50 In 1997 the total population decreasing (Crawford, Oregon State University, pers.
was estimated to be 1,000 birds (Mike Schroeder, comm.).
Washington Department of Fish and Wildlife, pers. Additionally, farming practices were acknowledged as
comm.). adding to the decline. The 1914 Oregon Sportsman stated:
In Oregon, a small, stable sharp-tail population has The method of farming in this section [Umatilla
been re-established in Wallowa County. Three leks County], which is almost entirely summer fallowing
currently support approximately 100 birds (Vic Coggins, and burning the stubble in the spring, has nearly
Oregon Department of Fish and Wildlife, pers. comm.). destroyed the prairie chicken [=sharp-tailed grouse]. . . .
100
Discussion
Once plentiful throughout their range, sharp-tailed grouse There were early warnings that efforts would need to
were harvested in abundance by early explorers and be undertaken to protect the grouse, such as the account
settlers. Populations were further impacted by the of the Columbian sharp-tailed grouse in 1915, which
conversion of grassland and sagebrush habitats to stated:
agriculture, livestock grazing in shrub/meadow steppe This grouse is one of our best game birds, but it has
and increased settlement in their former range. decreased very rapidly with the change of conditions
 CHAPTER 18: EXTIRPATED SPECIES 467

because it does not prosper in the vicinity of man. It has Wyoming Ground Squirrel
always held its own in the sagebrush with the coyotes (Spermophilus elegans)
and other natural enemies, but like the sage grouse, it Other Common Name: Richardson’s ground squirrel.
has suffered a great deal on account of the extensive
pasturing of sheep. . . . The only way this grouse can be Former Distribution
saved from extinction in many of the regions where it Fossil remains of the Spermophilus richardsonnii complex—
lives is by the setting aside of certain areas not fit for of which the Wyoming ground squirrel is a member—are
agricultural purposes as wild bird refuges where known from Alberta to Kansas, Texas, and Oklahoma, and
hunting is forbidden and the birds are well protected. as far east as Minnesota.46, 141 The oldest fossils of the
Otherwise, the history of this western grouse will be the Spermophilus richardsonni complex were found in Southern
same as that of its cousin, the pinnate grouse or heath Alberta, Canada. 47 Of Washington and Oregon, the
hen [=greater prairie chicken].34 Wyoming ground squirrel only occurred in southeastern
By the mid-1920s sightings of the sharp-tailed grouse Oregon.141 Trying to be more exact as to its previous
in Oregon were reduced to “a very few” in Wallowa distribution is problematic because of taxonomic
county, considered one of the leading counties in the state classification issues. In 1863, this species was first
as to the number of game birds.4 identified as a subspecies (Spermophilus richardsonni
At that same time in Washington, hunting was elegans) by Kennicott;66 in 1928 Howell55 classified it as a
restricted in all counties to a 4-month sharp-tail season species (Citellus elegans nevadensis); in 1938 it was
(August-November) with a daily bag limit of 10, replacing reclassified by Howell (Citellus richardsonnii nevadensis);
the previous 6-month season and daily bag limit of 20 of and in 1971 Nadler et al.87 finally classified the species as
the late 1800s. In 1909, Whitman County reduced the Spermophilus elegans nevandensis.
season to 3 months and a daily bag limit of only 5, and
Current Status and Distribution
closed the season entirely in 1919. Between 1933 and 1952,
As of 1999, the Wyoming ground squirrel occurs in 3
a moratorium was placed on sharp-tailed grouse hunting
distinct areas with separate subspecies: nevadensis in
in Washington. In 1953, hunting for sharp-tailed grouse
southeastern Oregon, southwestern Idaho and northern
was re-opened until 1988, when population declines
Nevada; aureus in southwestern Montana and eastern
resulted in a statewide closure.50 Sharp-tailed grouse were
Idaho; and elegans in the southern half of Wyoming and
placed on the state list as a threatened species in
northwestern third of Colorado with small extensions into
Washington in 1998 (Schroeder, Washington Department
Utah and Nebraska.151 In Oregon, Spermophilus elegans
of Fish and Wildlife, pers. comm.).
nevadensis is only known from 7 specimens collected at 2
In Oregon, Columbian sharp-tailed grouse were listed
sites: (1) in Malheur County, Oregon and (2) near
as an endangered species in 1969.79 The last known
McDermitt, Nevada. In the 1970s, the Wyoming ground
sighting in Oregon occurred in 1968 or 1969 in Baker
squirrel in Oregon was viewed as endangered if present
County.91 It is now considered extirpated throughout the
state of Oregon (Crawford, Oregon State University, pers.
comm.). The U.S. Fish and Wildlife Service considered the
sharp-tailed grouse in the Northwest a sensitive species
in 1982.138
The Washington Department of Fish and Wildlife has
been coordinating the acquisition of upland habitat for
sharp-tailed grouse through the Bonneville Power
Administration and the Washington Wildlife and
Recreation Coalition. Approximately 22,250 acres have
been purchased for sharp-tailed grouse. Additionally,
WDFW has been working with private landowners to
increase the benefits of CRP lands for sharp-tailed grouse.
Ongoing research, coordination, and partnerships are
focused on improving sharp-tailed grouse habitat, and
evaluating potential sites for possible reintroduction or
augmentation of populations.50
In Oregon, re-introduction efforts began in 1991 in
Wallowa County near Enterprise. As a result of successive
annual transplants with birds from southeastern Idaho,
two small populations have been successfully established
in areas of former distribution. In 1999, it was estimated
that the existing population consisted of 100 birds. Future
plans include re-introductions into other areas of the state
formerly occupied by sharp-tailed grouse (Sands, The
Nature Conservancy; Coggins, Oregon Department of Fish
and Wildlife, pers. comm.).
 468 Wildlife–Habitat Relationships in Oregon and Washington

in the state,141 because there had been no records for the suspected them to be “nearly, if not quite extinct there. . .”
species since 1927. Efforts in the 1980s to confirm presence in 1857, having never “met them.” He observed that
of the Wyoming ground squirrel within southwestern pronghorn were found sparingly in Oregon; but felt,
Idaho proved unsuccessful. In 1986, the first Oregon Non- unless stragglers occurred on the Spokane plains, it was
game Wildlife Management Plan was developed, and this doubtful whether they entered the Washington Territory.
species was listed as extirpated, but identified as the They were said by the Indians to have been formerly quite
Richardson’s ground squirrel.80 plentiful at the Dalles of the Columbia, but were nearly
exterminated in that locality by 1857.129
Discussion Pronghorn antelopes occurred on the Columbia Plateau
The Wyoming ground squirrel is a member of the from north central Washington through southeastern
unspotted ground squirrels and lives underground in Washington from about 8000 B.C. They were extirpated
burrows for most of the year. In areas where it is more in Washington prior to most European settlement,
common, like northern Nevada, the Wyoming ground although archeological and ethnographic records
squirrel can be found in meadows or habitats that emulate substantiate the presence of a few pronghorn into the early
meadow characteristics, as well as on occasion in 1800s.17, 93 The spatial distribution of the archeological and
sagebrush or bottom lands.46 ethnographic records corresponds with the semi-arid
The Wyoming ground squirrel can occupy mountain sagebrush habitat preferred by pronghorns.17
meadows from about 5000 feet (~1500 m) elevation to
above timberline.5, 20, 73 This ground squirrel is preyed upon Current Distribution
by snakes, coyotes, foxes, weasels, badgers, and hawks. After suffering declines through the early 1900s,
No one really knows the reason(s) for the decline of pronghorn numbers have steadily increased in Oregon,
the Wyoming ground squirrel in Oregon. The Oregon in part due to the establishment of the Sheldon-Hart
population is at the northwest extent of its range; hence, Mountain National Wildlife refuges in Oregon and
the Wyoming ground squirrel population may not ever Nevada.141 The current population of pronghorn in Oregon
have been very great. Competition with other ground is approximately 20,000 animals (Van Dyke, Oregon
squirrels may have played an initial role in limiting their Department of Fish and Wildlife, pers. comm.).
distribution, but the disjunct population of Spermophilus In Washington, re-introductions were attempted
elegans nevadensis is considered a relict one that has been beginning in 1938. Initially, 38 of 138 fawns brought into
out-competed by the Belding’s ground squirrel in more the state survived to be released in various areas including
mesic habitats, and by the Merriam’s and Piute ground the Squaw Creek Refuge (the present day Yakima Training
squirrels in arid habitats. The viability of the Wyoming Center), Adams County, the Colockum Game Range in
ground squirrel has been negatively affected by shooting, Kittitas County, and Grant County. 86 By 1950 the
poisoning, trapping, and conversion or loss of habitat.3 population had reached 200. Additional transplants of
pronghorns were made in 1968. Seven adults and 15 fawns
Species Extirpated from Washington were released in Grant and Kittitas Counties. Despite these

Pronghorn Antelope (Antilocarpa americana)

Other Common Names: pronghorn, antelope, prongbuck,
berrendos, little pale deer, small caribou.

Former Distribution
Pronghorns formerly occurred from southwestern
Manitoba, western Minnesota, eastern Texas, and Hidalgo
west to southwestern Alberta, eastern Washington,
southwestern Oregon, California (except coastal regions),
and Baja California.141
In Oregon, early reports indicated pronghorns were
abundant and widely distributed.8 Pronghorns originally
occurred throughout the area east of the Cascade Range
except for the Blue and Wallowa mountains. They also
occurred west of the Cascade Range in the Rogue River
valley and at the headwaters of the North Umpqua River.
From the beginning of European settlement through the
mid-nineteenth century, pronghorns rapidly declined and
became less widely distributed, although they were never
completely extirpated from Oregon.141
Lewis and Clark mention antelope occurring on the
great plains of the Columbia, though by no means as
abundantly as east of the Rocky Mountains. Baird
 CHAPTER 18: EXTIRPATED SPECIES 469

introductions, the population continued to decline, to 125 are: “to exchange information and encourage the
in 1971,94 and to 60 in 1978.143 None of the introduced perpetuation of sustainable wild herds of pronghorn as
animals survived to establish self-sustaining herds, and an ecological, aesthetic, and recreational natural resource
at present, pronghorn are not found in Washington (R. on public and private western rangelands at their most
Johnson, Washington Department of Fish and Wildlife, productive levels consistent with other land uses.”74
pers. comm.). Wildlife managers from both Oregon and Washington
participate in the Workshop, and have contributed to the
Discussion development of the Pronghorn Management Guides, a
Colonization of western rangelands by European settlers “compendium of suggested practices and techniques for
caused a more than a 99% decline in pronghorn managing pronghorn and their habitat.”74
numbers. 148 Fencing, habitat loss, competition with
domestic livestock, and un-regulated hunting all Upland Sandpiper (Bartramia longicauda)
contributed to the decline.74 Osborne93 suggested that Other Common Names: upland plover, Bartram’s
climatic changes throughout the Columbia Plateau were sandpiper, highland plover, field plover.
primarily responsible for displacing pronghorn from their
former pre- and proto-historic range in Washington. He Former Distribution
suggested: “colder winters, deeper, longer lasting snows, The original status of upland sandpipers in Oregon—and
and an influx of northern predators” as aspects of a likely Washington as well—before the introduction of
climatic pulsation beginning as much as 4,000 years ago livestock is unknown.81 It is thought that the potential
might well have upset the balance that made the eastern breeding range of the upland sandpiper included much
part of Washington habitable for the antelope. of eastern Oregon and Washington, as scattered sightings
Archeological sites where concentrations of pronghorn have occurred from throughout the area. Experience from
remains were found indicate a large number of individuals Oregon, however, has shown that it takes intensive
were acquired near those sites.17 Ethnographic records surveys by experienced observers in order to confirm
confirm group hunting methods; pronghorn driven into nesting.82
coulees toward waiting hunters134 as well as the use of Though a number of references from the early 1900s
group surrounds and natural enclosures.132 C. L. Brown18 included Oregon in descriptions of the range of the upland
felt the introduction of the horse and gun to the Plateau sandpiper, they all appear to be based on J. C. Merrill’s
Indians accelerated the extirpation of the pronghorn 1888 “Notes on the birds of Fort Klamath, Oregon” and a
antelope. Large number of furs and hides were traded to few scattered other specimens and sightings.63 Citing
obtain horses, while both guns and horses made it easier uncertainty in Merrill’s report, S. G. Jewett placed the
for hunters to acquire antelopes from greater distances. species on Oregon’s hypothetical list in 1929. He appeared
Economic reasons rather than subsistence needs began to to reverse himself by 1931 after breeding was confirmed
dictate hunting choices, making certain hunted species in Umatilla County, and more birds were seen in Grant
greater targets.18 County. By 1940, Gabrielson and Jewett list the upland
Although it is unknown how important a role disease sandpiper as a rare summer resident in the vicinity of
may have played in early pronghorn declines, losses from Ukiah, Umatilla County, and Bear and Logan Valleys,
disease have had serious impacts on present day Grant County.40
populations. Bluetongue, carried by cattle, caused the Upland sandpipers were first reported in Washington
death of at least 3,200 pronghorn in eastern Wyoming in in 1905, but no other evidence of their occurrence was
1976, and another 300 deaths in northeastern Wyoming found until 1928 when it was confirmed as breeding in
in 1984.135 Other serious disease vectors include parasitic Spokane County.62 In 1953, Jewett et al. describe the species
worms, which can be the cause of high fawn mortality.11 as an “uncommon and local summer resident in eastern
Bever12 found that areas over-grazed by domestic sheep Washington” with at least one small nesting population
resulted in high parasitic loads in pronghorn. in Spokane County.
Management of pronghorn in Oregon includes
managing the Sheldon-Hart Mountain National Wildlife Current Status and Distribution
refuges for pronghorn, in efforts to improve range The breeding range of the upland sandpiper stretches from
conditions and hunting regulations that promote herd north-central Alaska, northern Yukon, central and
health and vigor.141 In Washington, after unsuccessful northeastern British Columbia, southwestern Mackenzie
attempts to re-establish herds in their former range on the through the Canadian prairies to southern Quebec, central
Columbia Plateau, pronghorn management efforts have Maine, and New Brunswick, southeast to West Virginia,
not been pursued (R. Johnson, Washington Department central Virginia, and Maryland, and west throughout the
of Fish and Wildlife, pers. comm.). American Midwest to northeastern Oregon and eastern
The Pronghorn Antelope Workshop meets biennially; Washington.2, 53 Though the range covers a broad area,
attendees represent western state and provincial wildlife upland sandpipers are restricted to scattered local areas
agencies, federal land and wildlife agencies, universities of suitable habitat, and are particularly scarce west of the
and colleges, consultants, and private interests from Rocky Mountains. Populations in Oregon, Washington,
Canada, Mexico, and United States. The Workshop’s goals and Idaho are considered disjunct and peripheral to the
main range. Upland sandpipers winter in eastern South
 470 Wildlife–Habitat Relationships in Oregon and Washington

or Endangered in 10 other states, and as a candidate for

designation as Threatened or Endangered in British
Columbia, as of 1999 the species currently has no special
classification under the federal Endangered Species Act.53,
82
A recovery plan for the upland sandpiper in Washington
State was published in February 1995.

Discussion
The first threat to upland sandpiper populations occurred
in the late 1800s and early 1900s when market hunters
turned their attention from the disappearing passenger
pigeon to the upland sandpiper.53 With no bag limits or
closed seasons, and birds being shipped by boxcar loads
to market, declines were noticed by the 1870s. Birds were
also hunted on their South American wintering grounds.
Legal hunting of the upland sandpiper in North America
ended with the Migratory Birds Convention Act of 1916,
and the birds are now also protected by law in Argentina,
though illegal harvest likely still occurs in South America.53
The biggest threats to populations now are habitat loss
and alteration. Upland sandpipers are a grassland species,
requiring a juxtaposition of tall, dense vegetation for
nesting and shorter, more open grassy areas for resting,
feeding, and brood rearing.53 Platforms for courtship and
America from Surinam and northern Brazil south to territorial displays such as rocks or fence posts and a lack
central Argentina and Uruguay.2 of rugged topography appear to be important too. In
Although it is very small, Oregon has the largest known Oregon, nests have been found on the borders of
nesting population of upland sandpipers west of the seasonally wet mountain meadows, and near forest-
Rockies, and the status of the species in Oregon today is grassland or sagebrush-grassland edges.82 No descriptions
likely similar to what it was in the 1930s and 1940s. Surveys are available for the two known Washington nest sites.
conducted in 1984 estimated a population of about 100 Development of residential housing, commercial
birds in Oregon, with over half occurring in Bear and business areas, and gravel pits has contributed to
Logan valleys in Grant County.52 Subsequent surveys up permanent habitat loss. Conversion of native grasslands
through 1993 suggest a significant decline in numbers, to agricultural crop fields has likely resulted in “sink
however, from the mid-1980s. 82 Breeding has been populations,” because although upland sandpipers nest
confirmed in Bear and Logan Valleys, Grant County; in such areas, there is probably no reproductive gain, due
Bridge Creek Wildlife Area and Ukiah Meadows, Umatilla to repeated disturbances associated with crop
County; Campbell Flat on the Starkey Experimental Forest, management practices. Hayfield and pasture mowing
Union County; and Sycan Marsh, Lake County. 81, 128 have been implicated in chick mortality. Exotic plant
Regular sightings of upland sandpipers, including invasion by species such as the spotted knapweed has also
territorial behavior and display flights and song, were contributed to habitat loss and alteration. Heavy grazing
reported up through summer 1999 from Grant County.124, results in the reduction of grass cover to the point that
125, 126
upland sandpipers no longer use such areas for nesting,
In Washington, the last known record from the breeding and chicks and eggs are trampled and destroyed by cattle.
area near Spokane was of two adults seen on 22 September It may be possible, however, to carefully control stocking,
1993, after intensive surveys turned up no birds during timing, and duration of grazing in order to provide the
the breeding season82 (R. Rogers, Washington Ornitho- required juxtaposition of all cover conditions from short
logical Society, pers. comm.). Since then, individual to tall grasses.82
sightings have been reported from Pacific, Douglas, Grays While predators are an expected source of mortality,
Harbor, King, Okanogan, and Clallam Counties, all in agricultural areas likely attract more or different predators
August or September 110, 112 (R. Rogers, Washington than would be found in large intact areas of native
Ornithological Society, pers. comm.). No sightings were grasslands (foxes, skunks, American crows, coyotes,
reported for the state of Washington in 1994, 1996, or 1999, raccoons). Studies have also shown that the upland
even though intensive surveys were conducted in the east sandpiper is a loosely colonial nester where populations
Spokane Valley in 1994 82 (R. Rogers, Washington are healthy, and mobbing behavior by nesting adults helps
Ornithological Society, pers. comm.). reduce losses to predators.82
The upland sandpiper was listed as Endangered by the Upland sandpipers are largely insectivorous, and over
state of Washington in 1981 82 and is a Species of Special half of their diet is comprised of crickets, locusts,
Concern—Critical in Oregon. Though listed as Threatened grasshoppers, and weevils. 53 Heavy and widespread
 CHAPTER 18: EXTIRPATED SPECIES 471

insecticide use both on the wintering grounds and 23. ———, and P. E. Matthews. 1996. Rocky Mountain bighorn sheep in
breeding grounds, particularly in the 1940s, is also thought Oregon, history, and present status. Biennial Symposium North. Wild
to be partly responsible for population declines. Sheep and Goat Council 10:87-92.
24. ———, Matthews, P. E., and W.Van Dyke. 1996. History of
transplanting mountain goats and mountain sheep - Oregon. Biennial
Literature Cited Symposium North. Wild Sheep and Goat Council 10:190-195.
1. Almack, J. A., and S. H. Fitkin. 1998. Grizzly bear and gray wolf 25. Contreras, A. 1997. Northwest birds in winter. Oregon State
investigations in Washington State 1994-1995, final progress report. University Press, Corvallis, OR.
Washington Department of Fish and Wildlife, Olympia, WA. 26. Craighead, J. J., and J. Mitchell. 1982. Grizzly bear. In J. Chapman and G.
2. American Ornithologists’ Union. 1998. Check-list of North American Feldhamer, editors. Wild mammals of North America - biology,
Birds. 7th edition. American Ornithologists’ Union, Washington, D.C. management, and economics. John Hopkins University Press, Baltimore,
3. Andelt, W. F. and S. N. Hopper. 1998. Managing Wyoming ground MD.
squirrels. Colorado State University Cooperative Extension, No. 6.505. 27. ———, J. R. Varney, and F. C. Craighead. 1974. A population analysis of
Colorado State University, Fort Collins, CO. the Yellowstone grizzly bears. Bulletin 40, Forest and Conservation
4. Anonymous. 1926. Wallowa County rich in birds. Oregon Sportsman. Experimental Station, School of Forestry, University of Montana,
2(3):22. Missoula, MT.
5. Armstrong, D. M. 1972. Distribution of mammals in Colorado. 28. Cushing, J. E. Jr. 1941. The Columbian sharp-tailed grouse in Lake
Monographs of the Museum of Natural History, University of Kansas County, Oregon. Condor 43:75.
3:1-415. 29. Douglas, D. 1829. Observations on the Vultur californianus of Shaw.
6. Averill, A. B. 1895. The Sharp-tailed grouse. Oregon Naturalist 2:81-82. Zoological Journal 4:328-330.
7. Bach, G. R., and H. W. Dobyns. 1924. Hunter’s newsletter—Washington 30. Douglas, D. 1914. Journal kept by David Douglas during his travels in
District. Washington State Department of Agriculture, Bureau of North America 1823-1827. Antiquarian Press, New York, NY.
Biological Survey. Olympia, WA. 31. Ehrlich, P. R., D. S. Dobkin, and D. Wheye. 1988. The birder’s handbook.
8. Bailey,V. 1936. The mammals and lifezones of Oregon. North American Simon and Schuster, New York, NY.
Fauna, No. 55. U.S. Department of Agriculture, Bureau of Biological 32. Farr, A. C. M., and F. L. Bunnell. 1980. The sea otter in British Columbia
Survey, Washington, D.C. - a problem or opportunity. Pages 110-128 in R. Stace-Smith, L. Johns,
9. Banko, W. E. 1960. The trumpeter swan. University of Nebraska Press, and P. Joslin, editors. Threatened and endangered species and habitats in
Lincoln, NE. British Columbia and the Yukon. British Columbia Ministry of
10. Banko, W. E., and A. W. Schorger. 1976. Trumpeter swan. Pages 5-71 in Environment, Fish and Wildlife Branch, Victoria, BC.
R. S. Palmer, editor. Handbook of North American birds.Volume 2, 33. Finley, W. L. 1908. Life history of the California condor: Part II-
Waterfowl. Yale University Press, New Haven, CT. historical data and range of the condor. Condor 10:5-10.
11. Bever, W. 1950. Parasites and diseases of South Dakota antelope. 34. Finley, W. L. 1915. Columbian sharp-tailed grouse. Oregon Sportsman.
Pages 12-16 in Pittman-Robertson Job Completion Report Project 12- 3:138-140.
R-7. South Dakota Fish and Game Department, Pierre, SD. 35. Fleming, J. H. 1924. The California condor in Washington: another
12. ———. 1957. The incidence and degree of the parasitic load among version of an old record. Condor 26:111-112.
antelope and the development of field techniques to measure such 36. Foreyt, W. J. and D. A. Jessup. 1982. Fatal pneumonia of bighorn sheep
parasitism. Pittman-Robertson Project 12-R-14. No. 1-5, 2. South following association with domestic sheep. Journal of Wildlife Diseases
Dakota Fish and Game Department, Pierre, SD. 18:163-168.
13. Bloom, P. H., J. M. Scott, O. H. Pattee, and M. R. Smith. 1989. Lead 37. Franzreb, K. E., and S. A. Laymon. 1993. A reassessment of the
contamination of golden eagle Aquila chrysaetos within the range of taxonomic status of the yellow-billed cuckoo. Western Birds 24:17-28.
California condor Gymnogyps californianus. In B. U. Meyburg and R. D. 38. Friss, L. K. 1985. An investigation of subspecific relationships of the
Chancellor, editors. Raptors of the modern world. WWGBP: Berlin, gray wolf, Canis lupus, in British Columbia. M.S.Thesis. University of
London, and Paris. Victoria, British Columbia, Canada.
14. Bowlby, C. E., B. L. Troutman, and S. J. Jefferies. 1988. Sea otters in 39. Gabrielson, I. N. 1924. The season: Portland (Oregon) region. Bird-
Washington: distribution, abundance, and activity patterns. Washington Lore 26:343-344.
Department of Wildlife. Marine Mammal Investigations, Final Report for 40. Gabrielson, I. N., and S. G. Jewett. 1940. Birds of Oregon. Oregon
National Coastal Resources, Research and Development Institute, State College, Corvallis, OR.
Newport, OR. 41. Gaines, D. and S. A. Laymon. 1984. Decline, status and preservation of
15. Bonnot, P. 1951. The sea lions, seals, and sea otter of the California the yellow-billed cuckoo in California. Western Birds 15:49-80.
coast. California Fish and Game 37:371-389. 42. Garrietson, M. S. 1938. The American bison. New York Zoological
16. Brittell, J. D., Brown, J. M., and R. L. Easton. 1976. Marine shoreline Society, New York, NY.
fauna of Washington. Volume 2, Habitat requirements of selected 43. Gass, P. 1904. Gass’s journal of the Lewis and Clark expedition.
species. Washington Departments of Game and Ecology. Olympia, WA. Reprint of edition of 1811. A. C. McClurg, Chicago, IL.
17. Brown, C. L. 1977. An examination and evaluation of the archeological 44. Gilligan, J., M. Smith, D. Rogers, and A. Contreras. 1993. Birds of
occurrence of the American pronghorn antelope in the Columbia Oregon. Cinclus Publications, McMinnville, OR.
Plateau. M.A. Thesis. Washington State University, Pullman, WA. 45. Haines, F. D. 1940. The western limits of the buffalo range. Pacific
18.———, 1981. Note to Washington Department of Game 7 April Northwest Quarterly 31: 389-398.
1981. 46. Hall, R. E. 1981. The mammals of North America. Second edition. John
19. Buechner, H. K. 1960. The bighorn sheep in the United States, its past, Wiley & Sons, New York, NY.
present and future. Wildlife Monographs Number 4. 47. Harrington, C. R. 1978. Quaternary vertebrate faunas of Canada and
20. Burnett, W. L. 1931. Life history studies of the Wyoming ground Alaska and their suggested chronological sequence. Syllogeus 15:1-105.
squirrel (Citellus elegans) in Colorado. Bulletin of Colorado Agriculture 48. Harris, H. 1941. The annals of Gymnops to 1900. Condor 43: 3-55.
Experiment Station 373:1-23. 49. Hasegawa, H. and A. R. DeGange. 1982. The short-tailed albatross,
21. Burroughs, R. D. 1961. The natural history of the Lewis and Clark Diomedea albatrus, its status, distribution and natural history.American
Expedition. Michigan State University Press, East Lansing, MI. Birds 36: 806-814.
22. Coggins, V. L. 1980. Present status of Rocky Mountain bighorn sheep 50. Hays, D., M. Tirhi, and D. Stinson. 1988. Washington state status report
in Northeast Oregon. Biennial Symposium North. Wild Sheep and Goat for the sharp-tailed grouse. Washington Department of Fish and
Council 2:90-105. Wildlife, Olympia, WA.
 472 Wildlife–Habitat Relationships in Oregon and Washington

51. Hells Canyon Bighorn Sheep Restoration Committee. 1997. and habitat from Canada to Mexico. 18th Biennial Pronghorn Antelope
Restoration of Bighorn Sheep. The Hells Canyon Initiative. Idaho Workshop, Prescott, AZ.
Department of Fish and Game, Lewiston, ID. 75. Lentfer, J. W. 1988. Selected marine mammals of Alaska. Species
52. Herman, S. G., J. W. Scoville, and S. G. Waltcher. 1985. The upland accounts with research and management recommendations. Marine
sandpiper in Bear Valley and Logan Valley, Grant County, Oregon. Mammal Commission, Washington, D.C.
Unpublished report to Oregon Department of Fish and Wildlife, 76. Lewis, M., and W. Clark. 1905. Original journals of the Lewis and Clark
Portland, OR. expedition 1804-1806. R. G. Thwaites, ed.Volume 3, part 2;Volume 4,
53. Hooper, T. D. 1997. Status of the upland sandpiper in the Chilcotin- parts 1 and 2. Dodd, Mead, New York, NY.
Caribou Region, in British Columbia. Wildlife Working Report WR-86, 77. Linn, R. M., editor. 1979. Proceedings of the 1st Conference on
Ministry of Environment, Lands and Parks, Williams Lake, BC, Canada. Scientific Research in the National Parks. National Park Service
54. Horsfall, R. B. 1916. California cuckoo. Oregon Sportsman 4:195-196. Transactions and Proceedings Series No. 5.
55. Howell, A. H. 1938. Revision of the North American ground squirrels, 78. Manning, R. 1995. The buffalo is coming back. Defenders Magazine
with a classification of the North American Sciuridae. North American 71:6-15.
Fauna 56:1-256. 79. Marshall, D. B. 1969. Endangered plants and animals of Oregon. III.
56. Hughes, J. M. 1999.Yellow-billed cuckoo (Coccyzus americanus). In A. Birds. Oregon Agricultural Experiment Station Special Report 278.
Poole and F. Gill, editors. The Birds of North America, Number 418. The 80. Marshall, D. B. 1986. Oregon Nongame Wildlife Management Plan.
Birds of North America, Philadelphia, PA. Oregon Department of Fish and Wildlife, Portland, OR.
57. Ivey, G. L. 1990. Population status of trumpeter swans in southeastern 81. Marshall, D. B., M. W. Chilcote, and H. Weeks. 1996. Species at risk:
Oregon. Pages 118-122 in D. Compton, editor. Proceedings and Papers sensitive, threatened and endangered vertebrates of Oregon. 2nd
of the Eleventh Trumpeter Swan Society Conference. The Trumpeter edition. Oregon Department of Fish and Wildlife, Portland, OR. .
Swan Society, Maple Plain, MN. 82. McAllister, K. R. 1995. Washington state recovery plan for the upland
58. Ivey, G. L., M. J. St. Louis, and B. D. Bales. 2000 (In press). The status of sandpiper. Washington Department of Fish and Wildlife, Olympia, WA.
the Oregon trumpeter swan program. In R. Shea, editor. Proceedings 83. McCullough,Y. B. 1980. Niche separation of seven North American
and Papers of the Seventeenth Trumpeter Swan Society Conference. ungulates on the National Bison Range, Montana. Ph.D. Dissertation,
The Trumpeter Swan Society, Maple Plain, MN. University of Michigan, Ann Arbor, MI.
59. Jameson, R. J., K. W. Kenyon, A. M. Johnson, and H. M. Wright. 1982. 84. Meagher, M. 1986. Bison bison. Mammalian Species Number 266.The
History and status of translocated sea otter populations in North American Society of Mammalogists.
America. Wildlife Society Bulletin 10:100-107. 84a. Miller, L. 1957. Bird remains from an Oregon Indian Midden. Condor
60. Jameson, R. J., and S. Jeffries. 1999. Results of the 1999 survey of the 59:59-63.
reintroduced sea otter population in Washington State. Unpublished 84b. Miller, A. H. 1942. California condor bone form the coast of
report. southern Oregon. Murrelet 23:77.
61. Jennings, D. C., and J. Hebbring. 1983. Buffalo management and 85. Mitchell, C. D. 1994.Trumpeter swan (Cygnus buccinator). In A. Poole
marketing. National Buffalo Association, Custer, SD. and F. Gill, editors. The Birds of North America, Number 105. The Birds
62. Jewett, S. A., W. P. Taylor, W. T. Shaw, and J. W. Aldrich. 1953. Birds of of North America, Philadelphia, PA.
Washington State. University of Washington Press, Seattle, WA. 86. Moreland, R. 1969. Special Report. Antelope introductions.
63. Jobanek, G. A. 1997. An annotated bibliography of Oregon bird Washington Department of Game Status Report 1969. Olympia, WA.
literature published before 1935. Oregon State University Press, 87. Nadler, C. F., R. S. Hoffman, and K. R. Greer. 1971. Chromosomal
Corvallis, OR. divergence during evolution of ground squirrel populations (Rodentia:
64. Jobanek, G. A., and D. B. Marshall. 1992. John K. Townsend’s 1836 Spermophilus). Systematic Zoology 20:298-305.
report of the birds of the lower Columbia River region, Oregon and 88. Newberry, J. S. 1857. Report on the zoology of the route. Chapter 2.
Washington. Northwest Naturalist 73:1-14. Report upon the birds. Pages 73-110 in Reports of explorations and
65. Johnson, R. L. 1983. Mountain goats and mountain sheep of surveys, to ascertain the most practicable and economical route for a
Washington. Biological Bulletin Number 18. Washington State Game railroad from the Mississippi River to the Pacific Ocean…1854-55.
Department, Olympia,WA. Volume 6. Part 4. Zoological report. Number 2. Beverly Tucker, printer,
66. Kennicott, R. 1863. Descriptions of four new species of Spermophilus, Washington D.C.
in the collection of the Smithsonian Institution. Proceedings of the 89. Noble, D. G., and J. E. Elliot. 1990. Levels of contaminants in Canadian
Academy of Natural Sciences of Philadelphia 15:157-158. raptors, 1966 to 1988, effects, and temporal trends. Canadian Field-
67. Kenyon, K. W. 1969. The sea otter in the eastern Pacific Ocean. North Naturalist 104:222-243.
American Fauna 68:1-352. 90. Nowak, R. M., and J. L. Pardiso. 1983. Walker’s mammals of the world.
68. Kingston, C. S. 1932. Buffalo in the Pacific Northwest. The Washington John Hopkins University Press, Baltimore, MD.
Historical Quarterly 23:163-173. 91. Olson, B. 1976. Status report of the Columbian sharp-tailed grouse.
69. Koford, C. B. 1953. The California condor. National Audubon Society Oregon Wildlife 31(4):10.
Research Report Number 4. 92. Olterman, J. H., and B. J. Verts. 1972. Endangered plants and animals of
70. Larrison, E. J., and K. G. Sonnenberg. 1968. Washington birds: their Oregon IV. Mammals. Oregon State University, Agricultural Experiment
location and identification. The Seattle Audubon Society, Seattle, WA. Station, Special Report, 364:1-47.
71. Laufer, J. R., and P. T. Jenkins. 1989. Historical and present status of the 93. Osborne, D. 1953. Archeological occurrences of the pronghorn, bison,
gray wolf in the Cascade Mountains of Washington. Northwest and horse in the Columbia Plateau. Scientific Monthly 77:5 260-269.
Environmental Journal 5:313-327. 94. Parsons, Z. 1971. Unpublished report on pronghorn antelope
72. Laymon, S. A., and M. D. Halterman. 1987. Can the western subspecies transplants in Washington. Washington Department of Game, Olympia,
of yellow-billed cuckoo be saved from extinction? Western Birds 18:19- WA.
25. 95. Pasitschniak-Arts, M. 1993. Ursus arctos. Mammalian Species, Number
73. Lechleitner, R. R. 1969. Wild mammals of Colorado: their appearance, 439. The American society of Mammalogists.
habits, distribution, and abundance. Pruett Publication Company, 96. Pattee, O. H. and S. R. Wilbur. 1989. Turkey vulture and California
Boulder, CO. condor. In B. G. Pendelton, editor. Proceedings of the Western Raptor
74. Lee, R. M., J. D. Yoakum, B. W. O’Gara, T. M. Pojar, and R. A. Ockenfels, Management Symposium and Workshop.
editors. 1998. Pronghorn Management Guides. Biological and 97. The Peregrine Fund. 1999. California Condors to be released on
management principles and practices to sustain pronghorn populations December 7, 1999. The Peregrine Fund. http://www.peregrinefund.org/
press/con1299.html.
 CHAPTER 18: EXTIRPATED SPECIES 473

98. Phillips, A., J. Marshall, and G. Monson. 1964. The birds of Arizona. on the survey of the Northern Pacific Railroad route, relating to the
University of Arizona Press, Tucson, AZ. natural history of the regions explored, with full catalogues and
99. Pisano, R. 1977. Status of the wolf in North America. International descriptions of the plants and animals collected from 1853 to 1860.
Union for Wildlife Conservation. Baillière Brothers, New York, NY.
100. Plamondon, J. B. 1914. Chinese pheasants in Umatilla County. 130. Sullivan, P. T. 1983. A preliminary study of historic and recent reports
Oregon Sportsman 2(9):20. of grizzly bears, Ursus arctos, in the north Cascades area of Washington.
101. Presnall, C. C. 1946. Wildlife conservation as affected by American Washington Department of Fish and Wildlife, Olympia, WA.
Indian and Caucasian concepts. Journal of Mammalogy 27:458. 131. Sullivan, P. T. 1999. Field notes: eastern Oregon spring 1999. Oregon
102. Putnam, R. 1928. The letters of Roselle Putnam. Oregon Historical Birds 25:101-106.
Quarterly 29:242-264. 132. Suphan, R. J. 1974. Ethnological report on the Wasco and Tenino
103. Rausch, R. L. 1963. Geographic variation in size of North American relative to socio-political organization and land use. Garland, New York,
brown bears, Ursus arctos Linnaeus, as indicated by condylobasal length. NY.
Canadian Journal of Zoology 41:33-45. 133. Swainson, W., and J. Richardson. 1831. Fauna boreali-Americana; or
104. Riedman, M. L., and J. A. Estes. 1987. A review of the history, the zoology of the northern parts of British America. Part II. The birds.
distribution, and foraging ecology of sea otters. In G. R. VanBlaricom J. Murray, London.
and J. A. Estes, editors. The community ecology of sea otters. Springer- 134. Teit, J. A. 1928. The Salishan tribes of the western Plateau. Bureau of
Verlag, Berlin. American Ethnology Annual Report 45.
105. Rogers, P. M., and D. A. Hammer. 1978. Ancestral breeding and 135. Thorne, E. T., E. S. Williams, T. R. Spraker, W. Helms, and T. Segerstrom.
wintering ranges of the Trumpeter Swan (Cygnus buccinator) in eastern 1988. Bluetongue in free-ranging pronghorn antelope (Antilocapra
United States. Tennessee Valley Authority, Knoxville,TN. americana) in Wyoming 1976 and 1984. Journal of Wildlife Diseases
106. Rogers, R. 1997a. Washington field notes. WOSNews 48:8-10. 24:113-119.
107. Rogers, R. 1997b. Washington field notes. WOSNews 50:7-10. 136. Tice, B. 1998. Field notes: Western Oregon winter 1997-1998.
108. Rogers, R. 1997c. Washington field notes. WOSNews 51:8-10. Oregon Birds 24:96-99.
109 Rogers, R. 1998a. Washington field notes. WOSNews 52:8-11. 137. United States Congress. 1929. Control of predatory animals. 70th
110. Rogers, R. 1998b. Washington field notes. WOSNews 53:8-10. Congress, 2nd session. House Document No. 497. Washington D.C.,
111. Rogers, R. 1998c. Washington field notes. WOSNews 56:8-10. U.S. Government Printing Office.
112. Rogers, R. 1999a. Washington field notes. WOSNews 59:9-15. 138. U.S. Fish and Wildlife Service. 1982. Sensitive bird species, Region
113. Rogers, R. 1999b. Washington field notes. WOSNews 60:11-15. One. U.S. Fish and Wildlife Service, Portland, OR.
114. Rogers, R. 1999c. Washington field notes. WOSNews 61:8-11. 139. U.S. Fish and Wildlife Service. 1998. California Condor: Gymnogyps
115. Rogers, R. 1999d. Washington field notes. WOSNews 62:8-11. californianus.Wildlife fact sheet series 1. California condor. USFWS,
116. Rogers, R. 1999e. Washington field notes. WOSNews 63:8-14. Arlington, VA. http://www.fws.gov.
117. Ryder, T. J., E. S. Williams, K. W. Mills, K. H. Bowles, and E. T. Thorne. 140. Van Vuren, D., and M. Bray. 1985. The recent geographic distribution
1992. Effect of pneumonia on population size and lamb recruitment in of Bison bison in Oregon. Murrelet 66:56-58.
Whiskey Mountain on bighorn sheep. Proceedings of the Biennial 141.Verts, B. J., and L. Carraway. 1998. Land mammals of Oregon.
Symposium of the North American Wild Sheep and Goat Council University of California Press, Berkeley, CA.
8:136-146. 142. Washington Department of Game. 1975. 1974-1975 big game status
118. Rymon, L. M. 1969. A critical analysis of wildlife conservation in report, wolf section. Washington Department of Game. Olympia, WA.
Oregon. Ph.D. Dissertation, Oregon State University, Corvallis, OR. . 143. Washington Department of Game. 1978. 1978 big game status
119. S., J. 1881. Shooting in Oregon. Forest and Stream 16:9. report. Washington Department of Game, Olympia, WA.
120. Shaffer, M. L. 1981. Minimum population sizes for species 144. Washington Department of Fish and Wildlife. 1995. Washington State
conservation. BioScience 31:131-134. Management Plan for bighorn sheep. Washington Department of Fish
121. Smith, D. 1954. The bighorn sheep in Idaho. Idaho Department of and Wildlife, Olympia, WA.
Fish and Game, Wildlife. Bulletin No. 1. 145. ———, 1998. 1998 game status and trend report. Washington
122. Smith, M. R., P. W. Mattocks, Jr., and K. M. Cassidy. 1997. Breeding Department of Fish and Wildlife, Olympia, WA.
birds of Washington State.Volume 4 in K. M. Cassidy, C. E. Grue, M. R. 146. Wilbur, S. R. 1973. The California condor in the Pacific Northwest.
Smith, and K. M. Dvornich, editors. Washington State Gap Analysis— Auk 90:196-198.
Final Report. Seattle Audubon Society Publications in Zoology Number 147. Wilcox, B. A., and D. D. Murphy. 1983. Conservation strategy: the
1, Seattle,WA. effects of fragmentation on extinction. American Naturalist 125:879-
123. Sodhi, N. S., L. W. Oliphant, P. C. James, and I. G. Warkentin. 1993. 887.
Merlin (Falco columbarius). In A. Poole and F. Gill, editors.The Birds of 148.Yoakum, J. D. 1968. A review of the distribution and abundance of
North America, Number 44. The Birds of North America, Philadelphia, American pronghorn antelope. Antelope States Workshop Proceedings
PA. 3:4-14.
124. Spencer, K. T. 1998. Field notes: Eastern Oregon summer 1997. 149.Yocom, C. F. 1952. Columbian sharp-tailed grouse (Pediocetes
Oregon Birds 24:29-31. phasianellus columbianus) in the state of Washington.American Midland
125. Spencer, K. T. 1999. Field notes: Eastern Oregon June-July 1998. Naturalist 48:185-192.
Oregon Birds 25:13-20. 150. Young, S. P. 1944. The wolves of North America, Part I. American
126. Spencer, K. T. 2000. Field notes: Eastern Oregon summer 1999. Wildlife Institute, Washington D.C. Condor 43(1):75.
Oregon Birds 26:138-143.
127. Spraker, T. R., and C. P. Hibler. 1982. An overview of the clinical signs,
gross and histological lesions of the pneumonia complex of bighorn
sheep. Proceedings of the Biennial Symposium of the North American
Wild Sheep and Goat Council 3:163-172.
128. Stern, M. A., and G. A. Rosenberg. 1985. Occurrence of a breeding
upland sandpiper at Sycan Marsh, Oregon. Murrelet 66:34-35.
129. Suckley, G., and J. G. Cooper. 1860. The natural history of
Washington Territory and Oregon with much relating to Minnesota,
Nebraska, Kansas, Utah and California, between the thirty-sixth and
forty-ninth parallels of latitude, being those parts of the final reports
 19
Characterizing Species at Risk
John F. Lehmkuhl, Bruce G. Marcot, & Timothy Quinn

Introduction
For at least the past three decades, researchers and management objective will not be met. Risk is the
managers have used various forms of risk and decision likelihood of an outcome occurring multiplied by the value
analysis for assessing the status and effects of management given to an outcome. This means that to fully evaluate
practices on populations or habitat. Early methods that the “risk” of any course of action, management objectives
continue to be practiced today use habitat relationships first must be articulated in terms that can be clearly
information to assess changes in macro- or microhabitat analyzed.
area from land management practices or other human
activities.36, 147, 171 Other forms use risk analysis to identify Distinguishing Risk Analysis from
and rank species of conservation concern. 58, 77, 107, 120 Risk Management
Quantitative population viability models have received In risk analysis, researchers list possible outcomes,
extensive attention for analyzing probabilities of extinction estimate their likelihood under one or more management
for single species under current or future conditions,1, 80 alternatives, and then weight each outcome with a value.
but they are just one of many tools in the risk analysis Outcome values are assigned by managers based on social,
toolbox. Decision models have used risk analysis to economic, political, or other goals. Defining what is
compare extinction likelihood of wildlife populations to desired (outcome values) and the likelihood of achieving
the economic cost of their conservation.49 Many other it (outcome likelihood) helps determine the risk of a
examples can be found in the literature. decision (Figure 1). In risk management, the manager first
In this chapter, we attempt to describe some of those describes the values they place on possible outcomes,
risk analysis tools, but also strive to provide readers with ideally in conjunction with the risk analysis. Then the
a basic understanding of the underlying theory and criteria manager describes their risk attitude towards seeking or
for identifying species at risk. We focus on analytical tools avoiding particular kinds of outcomes. Then, they apply
that might be used with available datasets, like the CD- this attitude to the risk analysis results and choose an
ROM in this book, to meet the risk analysis challenge at acceptable course of action that best fits their risk attitude
different levels of geographic and ecological scale. We and the outcome values.
define the components of risk analysis, describe the criteria Data that fuel risk analyses can be uncertain in several
for identifying species at risk, review methods of species ways. There can be errors of measurement, sampling
risk analysis, examine the attributes of species currently errors, and natural variation. Data errors of these kinds
considered at risk, and work several examples of risk can result in greater variances in the estimation of the
analysis using the life history and habitat relationships biological factor of interest, which can add some degree
information in this book. of confusion to the risk management phase. How these
errors add up and interact is typically ignored in risk
Defining Risk analyses, but the problem of error propagation is not to
In much of the biological literature, the term “risk” really be neglected. Methods to negate the effects include
means “likelihood” of some event or condition occurring.1 stochastic simulation analysis and sensitivity analysis,144
However, formally, ecological risk analysis typically use of meta-analysis techniques,34, 38, 56 partitioning or
differentiates the estimation of outcome likelihood bootstrapping samples,111 and, when the problem is simple
(possibility of change or loss) from management values enough, outright analytic solutions.115
(or utilities) placed on those outcomes. Outcome A common erroneous view of uncertainty (variance)
likelihood can be estimated quantitatively (e.g., 90% in risk analysis outcomes is that it appears as a lack of
persistence of a nonzero population over 100 years) from evidence of adverse effects, or even as evidence of no effect.
simulation modeling (rarely from empirical study), This can be a biologically, and politically, hazardous
qualitatively from simpler habitat or life history models, approach leading, in the worst cases, to species extirpation
or from expert judgment as an ordinal scale (rank order) or extinction. What can the risk analyzer do in the face of
of outcomes. “Risk” is the likelihood that some such unknowns? True data unknowns mean that no

474
 CHAPTER 19: SPECIES AT RISK 475

statistical estimation or population projection is even exotic species will only increase with the growth of the
possible, even by using expert judgment. The risk analyzer global economy.
can skirt the issue by borrowing data from a related Landscape change (habitat loss, degradation, and
species, time period, or geographic location, or can fragmentation) is the primary cause of endangerment for
conduct a sensitivity analysis, letting an unknown variable two-thirds of the world’s vertebrates currently considered
vary over some biologically reasonable range of values at risk of extinction.128 Some major habitat types such as
and determining the degree to which the outcome (e.g., the North American tallgrass prairie have been virtually
population persistence likelihood) is influenced by that eliminated (converted to other uses) while others such as
variable. A more formal analysis of the value of additional the heathlands in the United Kingdom and thorn scrub in
information also can be conducted to determine the worth Sri Lanka have been reduced to less than 30% of their
of expending time and money in seeking values of original extent.141 Currently some of the highest rates of
unknown variables, or refining estimates of ones known. forest conversion to agricultural and urban land are found
in the humid tropics, 186 our great storehouses of
Criteria for Identifying Species at Risk biodiversity. Other habitat types are changed in less
If current trends in human behavior continue, the twenty- obvious ways. For example, while the amount of forest
first century may come to be known as a mass extinction cover is increasing in Europe and the former USSR, the
event rivaling the extinction event of 65 million years ago. clearing of virgin forest worldwide has increased
Some estimates suggest that up to one quarter of all species exponentially since the 1500s.99 Of the 4 billion hectares
now living may be lost in the next fifty years,102 mainly estimated to be in forest cover today, 1.7 billion are
through habitat alteration, but also through introduction secondary forests or plantations.185
of invasive species, pollution, and other human activities. As an area is converted from natural vegetation to some
Clearly, to avoid such a catastrophe, natural resource other use, the remaining habitat is commonly divided into
professionals must better understand the process of smaller parcels. The effects of these actions on biodiversity
extinction, how humans can affect species’ extinction risks, are diverse,32, 131, 146, 181 but often discussed in terms of three
and the attributes that make species particularly related effects: fragmentation, edge, and isolation. It is
extinction-prone. In particular, decision makers and risk important to note that although human alterations to the
analyzers need theory and predictive tools to make wise environment are often considered deleterious to most
land-use decisions in short time frames and otherwise species, the effects of landscape change including
data-poor environments, particularly in the next fifty fragmentation, isolation, and the creation of edge habitat
years, when the world’s human population is expected to are species-specific. What constitutes a patchy
peak. Below, we outline some of what we know and do
not know about the process of extinction.
The viability of populations is driven by two basic types
of processes: systematic and stochastic. Systematic
processes generally reduce the size of populations,155 and
result from the removal of something essential from the
environment, such as space, shelter, or food, or from the
insertion of something lethal, such as a new predator or
poison. Systematic processes tend to reduce populations
sizes to levels that make them susceptible to stochastic
processes that lead to extinction. Stochastic processes, on
the other hand, are random events or a series of events
that can dramatically affect birth and death rates within
populations.

Systematic Extinction Processes

Primary processes. Of the 485 animals and 584 plant
species that have been certified as extinct since 1600,102
approximately 61% of the animal extinctions and 27% of
the plants are island endemics. Most of these island
extinctions resulted from Diamond’s29 “evil quartet” of
systematic processes: habitat destruction and
fragmentation, introduction of exotic species, over-
exploitation by humans, and chains of extinction.
Although this same evil quartet affects mainland species,
island endemics, having evolved in the absence of
predators or competitors including humans, are Figure 1. Example of risk management and risk analysis as
particularly vulnerable to extirpation when exotic applied to population viability planning (from Marcot et al.
predators or competitors are introduced. The spread of 1986).
 476 Wildlife–Habitat Relationships in Oregon and Washington

environment for one species may constitute a Isolation. Isolation refers to the accessibility of habitat
homogeneous environment for another species. Likewise, patches to organisms, and mostly is related to two
degraded habitat for one species may constitute ideal measures: distance between patches and the composition
habitat for another. Moreover, some species will benefit of matrix lands surrounding those patches. All else being
from human activities either directly by the creation of equal, as distance between patches increases, or suitability
habitat (edge for example) or through indirect effects such of the matrix land as habitat decreases, the more isolated
as reducing competition or predation pressure. a patch becomes to a particular species. Demography
within and dispersal among patches, which is a function
Fragmentation. Habitat loss results in the reduction of
of isolation, will often determine the response of
habitat and in discontinuities in the distribution of
individual species to fragmentation.63
remaining habitat, that is, fragmentation. Fragmentation
Upon isolation, a patch is likely to have more species
can be viewed as the creation of remnant vegetation
than it can sustain in the long-term. Species may be lost
patches surrounded by a matrix composed of different
from the patch in a process called species relaxation.112 The
habitat or land/uses.146 For illustrative purposes, consider
most rapid extinctions are likely to occur in species that
a single species on a landscape where each parcel of land
depend entirely on native vegetation, require large
is either classified as habitat or non-habitat. At one end of
(relative to the patch size) territories, and those that exist
the fragmentation spectrum and in the simplest case,
at low densities and are thus vulnerable to stochastic
patches of habitat may be so small that they support no
processes.146
individuals (i.e. smaller than a single home range, or
missing some important habitat feature such as nesting Stochastic Extinction Processes
substrate). In areas that are slightly less fragmented, small Demographic stochasticity. Stochastic, or random,
populations may persist in some of the larger habitat processes are usually the proximate causes155 of extinction
patches. However, these small populations may be at for populations already small from systematic processes.
increased risk of extirpation due to stochastic processes There are four types of stochasticity: demographic,
described below. environmental, natural catastrophes, and genetic.37, 160, 161,
A collection of small populations (or subpopulations) 164
Demographic stochasticity is the result of chance
each inhabiting its own patch may constitute a variation in vital rates. This form of stochasticity affects
metapopulation. Although there are many variations on individuals randomly and increases the variance in
the metapopulation theme, the distinguishing mortality and/or natality rates and thus increases the
characteristic of metapopulations is that population probability of extinction.43, 74, 101 For example, a species
dynamics within a patch are relatively independent of consisting of very few breeding pairs by chance alone
dynamics in other patches. That is, dispersal among could produce offspring all of the same sex, effectively
patches is a relatively rare event that has little effect on reducing the rate of successful reproduction in the next
the extant population’s growth rate. In theory, persistence generation. Alternatively, consider a species where the
of a metapopulation is related to the rate at which each annual probability of mortality based on the population
patch (subpopulation) goes extinct and is recolonized by as a whole is 50%. If we represent the fate of individuals
individuals from another patch. The metapopulation as the result of a coin toss, it is easy to demonstrate that 5
persists if recolonization is greater than extinction rate. tails in row (where tails equal death) are easier to obtain
Edge effects. As patches of habitat become smaller (and than 100 tails in a row. The smaller the number of coin
the matrix expands), the ratio of edge habitat to interior tosses (smaller the population) the more likely they will
habitat increases. Edges can be thought of as areas of all come up tails. Demographic uncertainty is an important
abiotic and biotic gradients between patch habitat and the consideration usually only for relatively small
matrix. For example, in a clear-cut matrix with forest populations152—on the order of 100s.
patches, the outside edge of the forest patch has conditions Environmental stochasticity. Environmental stochasticity
(e.g. light humidity, temperature, plant species is the variation in mean fecundity and survivorship rates
composition) more like those in the clear-cut than in any across space and time, although it is most commonly
other part of the forest patch. Edges essentially reduce the thought of as temporal variation at a single site. It results
amount of habitat for species requiring interior conditions from changes of environmental or habitat quality and
of the patch. Shade tolerant species may become restricted interspecific interactions with competitors, predators,
to the interior parts of the habitat patches, with different parasites and diseases.101 In contrast to demographic
species requiring different distance from the edge.73, 136 stochasticity, environmental stochasticity can affect
Edge habitat may have different types of species individuals in the population more or less equally to
interactions than either the matrix or the patch interior. increase the variation in birth and death rates, and thus
For example, nest predators and avian brood parasites are variation in abundance.43, 74 Natural catastrophes, such as
more common along forest edges than in patch interiors.169, floods, fires, and droughts, often occur at random intervals
180
The importance of this phenomenon depends on the and can have dramatic and usually adverse effects on
scale of edge effects: some outside influences penetrate small populations. Also, because some catastrophes such
only a few meters from the boundary (e.g. sunlight),82 as hurricanes are very large, they can adversely affect even
whereas others infiltrate hundreds of meters in to the very large populations. However, as the geographic range
interior (e.g., wind,25 nest predators).180
 CHAPTER 19: SPECIES AT RISK 477

of a species increases (i.e. larger population) there is less (systematic processes). In 1907, a portion of Martha’s
likely to be spatial correlation of environmental variation Vineyard was set aside as a refuge for the bird and a
across that range. Thus, large populations may be buffered program of predator control was implemented. The
against environmental stochasticity occurring at relatively population responded, and by 1916 had reached the size
small spatial scales. of more than 800 birds. In 1916 a fire (natural catastrophe)
destroyed most of the heath hens’ nests and during the
Genetic stochasticity. Genetic stochasticity is the variation
following winter the birds suffered unusually heavy
in frequency of alleles within a gene pool. The type of
predation (environmental stochasticity) from a relatively
genetic stochasticity that conservation biologists are most
high concentration of a common heath hen predator, the
concerned with is loss of allelic diversity. This arises from
goshawk. The combined effect of these events reduced the
a reduction in the “effective population size” (that is, the
population to 100-150 individuals. In 1920, after the
effective number of breeding adults) with consequential
population had increased to about 200, disease
deleterious short-term effects of random allele fixation,
(environmental stochasticity) reduced the population to
founder effects, inbreeding or outbreeding on
below 100. Though the species persisted a while longer,
reproductive success, and long-term losses of genetic
by 1932 the last survivor was gone. In the final stages of
variation (heterozygosity) and adaptability to
the population decline, the birds appeared to have become
environmental change from genetic drift.37, 148 Inbreeding
increasingly sterile (inbreeding depression) and the
has been shown to reduce reproductive success in small
proportion of males increased (demographic stochasticity).
or isolated populations, including rainbow trout, 70
Which of these events served as the coup de grace is
ungulates,6, 133, 134 red-cockaded woodpeckers,48 and pit
unknown. Theory suggests that in general, the probability
vipers.84 In the past few years, the emphasis on genetics
of extinction for small populations is highest from
in conservation biology has decreased for a variety of
environmental effects, followed by demographic
reasons, the most important being the conclusion that
stochasticity, longer-term genetic effects, and lastly
populations large enough to be demographically self
catastrophes,42, 72 although this hierarchy itself varies by
sustaining are likely to remain genetically viable
species and condition.
indefinitely.55, 74, 110 However, the relationship between
In contrast to systematic pressures, the magnitude of
long-term viability and genetic diversity remains unclear
stochastic threats depends more on population size than
and is an active area of research.
on life history traits of a species. In part, this is because
Interactive effects on populations. The population size life history traits are generally selected traits imparting
at which stochastic processes become important is likely persistence of the species within particular bounds of
to vary over time and space according to factors affecting environmental conditions. Thus, a general theory of
successful reproduction and dispersal.44 For example, stochastic processes has been developed based on
given two populations of equal size, high rates of population size alone. Systematic processes, on the other
reproduction or immigration in one population would hand, are likely to be as different from each other as the
allow faster recovery from disturbance than where life history traits of species affected by those processes. It
reproduction or immigration are slow. Such population may not be possible to construct a universal theory of
size thresholds also are species-specific and largely population decline comparable to the stochastic theory of
determined by the interaction among the particular small populations. 24 However, understanding the
characteristics of a species life history and population mechanisms of extinction (i.e. why some species and not
structure, and environmental conditions.42 Birth and death others decline in the face of systematic extinction
models of population demography indicate that the risk processes) is fundamentally important to conservation
of extinction declines rapidly with increasing population biologists and wildlife managers.
size.44 As an example, Morrison et al.115 analyzed the
exponential rate of decline of native Hawaiian bird Criteria of Species at Risk
populations, and demonstrated that species with very Population vulnerability analysis. The basic question is:
small populations decline at a far faster rate than do larger why do some populations decline and others do not? We
populations. Extinction risk as a function of environmental can use the conceptual “population vulnerability analysis”
stochasticity and large-scale catastrophes also will framework to determine criteria and processes for
decrease with increasing population size, but at a slower assessing extinction risk.42 The risk of extinction due to
rate than with demographic effects. In reality, extinction systematic processes is likely a function of the interaction
of small populations is probably the result of interactions of: (1) the physical and biotic environment of the
among these four processes,42 although their relative individual and population; (2) the population phenotype,
contribution to extinction is largely unknown and possibly including the species life history, behavior, morphology,
quite complex. physiology, and habitat requirements; and (3) how the first
For example, consider the extinction of the heath hen two factors interact to determine population distribution
(Tympanuchus cupido cupido), a species that was fairly and individual fitness.42, 174 The relative importance of these
common in the New England States at the time of three factors is a matter of debate and probably depends
European settlement.157 By 1900, there were fewer than on the particular blend of species characteristics and
100 birds, all of which were found on Martha’s Vineyard environmental characteristics. Despite the potential
 478 Wildlife–Habitat Relationships in Oregon and Washington

complexity of understanding the mechanisms of • In populations with high rather than low variability in
extinction, a number of studies have attempted to reduce numbers (but see Tracy and George).174
the number of factors into an easily understood and
Interactive effects. While these predictions can help us
measurable suite of characteristics. In our case, we are
formulate testable hypotheses, they say nothing about the
interested in determining a priori whether a particular
processes which drive extinction, or how these life history
species is more extinction prone than is another species.
and other factors work to increase risk. Moreover, the
The question becomes: how do certain population
relationship between factors that limit population size and
distribution patterns and life history characteristics enable
distribution and chances of extinction are complex, at least
species to persist in the face of the most common threats
for species that have existed in small numbers within small
to their viability and the ones which humans can most
geographic ranges for millions of years. Burgman et al.18
control—habitat loss and fragmentation?
suggest that factors that lead to relative rarity or restricted
The probability that a population will persist with
geographic ranges may be independent of those that cause
habitat loss is largely a function of population density or
extinction. Even those species that are tolerant of a wide
abundance: the greater abundance or density the lower
range of environmental conditions may lack available
the risk of local extinction.27, 28, 119, 125, 163, 170, 182 That is, larger
habitat.
populations, and populations distributed in a
In addition to these general characteristics, Akçakaya
metapopulation pattern (with low correlation of stochastic
et al.1 identified several properties that may make species
factors and environmental conditions among population
especially susceptible to human impacts:
centers) are less vulnerable to stochastic processes than
• Habitat overlap: species that occur in the same types
are small populations or populations lacking
of habitats that people or introduced superior
metapopulation structures, all else being equal (see
competitors prefer are more at risk than other species
discussion above).
occurring in less preferred areas. Humans tend to
The interaction of population distribution and
populate areas with fertile soils and benign climates
geographic scale of management concern also is an
such as major rivers valleys and coastlines. These
important issue in determining and managing risk. For
areas are subject to waste disposal and urban and
example, species that are globally common, but regionally
agricultural development.
or locally rare, may not be of concern at the large scale,
• Harvesting: species that taste good or otherwise
but may be important for local management to preserve
valuable to humans are often at risk from hunting.
subspecies and the evolutionary potential of the species
Human exploitation may exacerbate species’
at the edge of its range.95 On the other hand, for species
properties, which put them at risk. For example, sea
that are locally common, but rare at broader scales, risk
turtles require 10-40 years to reach sexual maturity,
management might be important at both local and higher
so removal of breeding adults rapidly depletes a
geographic scales to maintain “source” populations or
population and slows the rate of recovery when
strongholds of viability.
exploitation ends (e.g., populations stay small long
In addition to population structure, life history
periods of time).71
characteristics are important attributes of the “population
• Large home range requirements: animals with
phenotype” that may affect persistence. 96, 106, 153, 156
extensive home ranges often occur at low densities
Vulnerability to extinction in isolated habitat patches may
and are susceptible to the reduction and
increase with large body size;27, 28, 125 low fecundity;125
fragmentation of habitat.
specialization on patchy resources;64, 168 behavioral patterns
• Limited adaptability and resilience: species with
requiring the formation of large groups and thus
limited dispersal capabilities, limited reproductive
concentrated resources, dependence on keystone or link
capacity (e.g. sea turtles), or narrow and inflexible
species, or occurrence at the edge of a species range;170
habitat requirements may be incapable of rapid
temporal variation in population size; 28, 43, 44, 66 and
recovery from disturbance making them prone to
metabolic rate.182 Many of these life history traits are
extinction in human dominated landscapes.
related to body size.
To this list we can add dispersal ability and factors
Based on his study of colonization and extinction rates
influencing successful dispersal and recolonization:
of birds on islands, Pimm et al. 125 summarized the
species with poor dispersal ability are subject more to local
theoretical underpinnings of extinction risk in four
stochastic influences than are species with broad dispersal
predictions. All other things being equal, extinction rate
ability; that is, dispersal ability influences the distribution
will be greater:
patterns of a species.53
• In small populations than in large populations (see
above), Lessons for management. There are two important lessons
• At low population densities, in small-bodied, fast from small-population theory. 1) managing small
growing, short-lived species than in large-bodied, populations requires enormous effort, as does conducting
slow-growing, long-lived species, research on them to determine causes for decline. The best
• At high population densities in large bodied, slow- time to ensure viability is when populations are still large.
growing, long-lived species than in small-bodied, fast 2) Managers should attempt to reverse or mollify adverse
growing, short-lived species, and effects of systematic threats to persistence and increase
 CHAPTER 19: SPECIES AT RISK 479

the effective size of threatened populations as rapidly as Habitat Relationships Assessments

possible.55 Additionally, in some cases, it is advantageous Wildlife habitat relationships (WHR) assessments or
to establish more than one population center to avoid models use a variety of procedures to estimate the effects
losing the entire population to single environmental of changes in habitat quantity or quality under
stochastic influences or catastrophes. In extreme cases, management alternatives. The objective of WHR models
captive propagation followed by reintroduction into is to assess how changes in vegetation composition or
restored habitats, although terribly expensive, may be one structure from management might result in habitat loss
of the best recourses.149 for on or more species. Multiple species typically are
analyzed in guilds or groups of species with similar habitat
Methods for Determining relationships, foraging or nesting behavior, movement, or
other life-history attributes. Habitat relationship
Species at Risk
assessments can be applied at stand, landscape, and
Methods for wildlife risk assessment vary in taxonomic
regional scales, although Raphael and Marcot90 found such
scope, procedural complexity, and objective. The scope of
models most accurate and appropriate for use with some
assessments ranges from single-species assessments to
species at landscape or larger scales.
multi-species ecosystem or biodiversity assessments in
The methodological roots of wildlife habitat
which many species are considered in the same effort.
relationships programs for vertebrates are in the work of
Procedural complexity varies from simple deterministic
Patton,122 Thomas et al.,171 Lehmkuhl and Patton,75 and
estimates of risk (e.g., consistent loss of habitat area, and
Brown,14 among others. Indexed WHR models include the
use of ordinal risk rankings) to complex probabilistic
habitat suitability models (HSI),147 the habitat evaluation
estimates for explicit time periods that incorporate
procedures (HEP),36, 178 and habitat effectiveness models
uncertainty related to demographic and environmental
(e.g., elk models by Thomas et al.,171 Wisdom et al.,184
stochasticity, or management processes (Figure 2). Risk
Thomas et al.)173 Morrison et al.115 review these models.
assessments also vary in their objectives, from estimates
Habitat is typically defined as the sum of area in plant
only of extinction risk to estimations of management risk
communities and their structural variants (macrohabitats).
that integrate extinction risk with social, economic, or other
Adding to the suitability of an environment are habitat
biological considerations. Several of the methods to assess
elements (microhabitats) used for feeding, nesting, and
risk of habitat or population loss described below, such as
resting. Analysis usually determines the change in area of
habitat relationships assessments and models, are covered
available habitats under management alternatives and
in detail elsewhere in this book; here, we give an overview
describes the implications for associated species.
of these and other methods for analyzing risk of loss.
Versatility indices of habitat use are the number, or
breadth, of macrohabitats or habitat elements used by
individual species. Versatility indices are used to assess
implications of habitat change for species that use few
habitats (low versatility) or several to many habitats
(higher versatility). Index models (HEP, HSI)
independently describe or index the value (typically scaled
1 to 10 or 0 to 1) of several macrohabitat and microhabitat
attributes (e.g., westside conifer forests and snag density
therein) then calculate a summary index of habitat value.
Risk is determined from HEP and HSI approaches by
comparing model index values against an independently
determined threshold of habitat change considered to put
the species at risk.
It is problematic, however, in HEP and HSI models to
identify such thresholds. Thus, HEP and HSI type models
are better used to qualitatively assess alternative
conditions in a rank order comparison, rather than to
estimate absolute population response. In some cases,
historical and projected trends in area of forest habitats
have been related to population density in habitat types
to depict potential trends in the abundance of vertebrate
taxa and functional groups over large areas.137, 139 Similar
trends and analyses pertain to other, non-forest
Figure 2. Methods of risk assessment for animal and plant environments.
species vary along gradients of taxonomic scope and
procedural complexity. Additionally, methods vary in
objective from estimating ecological risk of extinction to
management risk.
 480 Wildlife–Habitat Relationships in Oregon and Washington

Rank Models been developed that integrate extinction and management

Rank models work within a population vulnerability risk attributes to set state-wide or global management
framework to evaluate risk.9, 42, 150 These models provide a priorities for individual species.4, 20, 62, 81, 114, 120, 135
ranking of extirpation or management risk, using
attributes of population structure and distribution, life Population Viability Analyses and Assessments
history, habitat-use versatility, or habitat area and spatial Population viability analysis (PVA) is a formal modeling
pattern. For ordinal-rank models, each factor is usually process for quantifying the likelihood that a species will
assigned an integer score, or rank, relative to its persist for a given time into the future.1, 9, 42, 150, 152 In PVA,
contribution to extirpation or management risk, for attributes of population structure and demography, life
example, on a scale of 1 to 10. Quantitative data on habitat history, genetic structure, and habitat quality and quantity
area or population size, as well as qualitative information typically are used in models combining population
on life history attributes, can be used to develop the scores. demography and geographically referenced
In this approach, multiple species typically are evaluated metapopulation dynamics to assess viability likelihood
at the same time using the same variables. The scale of and extinction risk. A very significant innovation that
application is most often regional or larger, but could be distinguishes this form of risk assessment from that based
applied down to watershed or smaller scales. Ordinal-rank on WHR models and rank methods is a consideration of
models have been developed to assess the impacts of large- uncertainty arising from natural variation or
scale habitat loss and fragmentation on potential unpredictability in population and environmental
population persistence.51, 61, 77 processes. Sometimes, uncertainty is addressed by
The International Union for the Conservation of Nature depicting viability as likelihoods of extinction over a
and Natural Resources (IUCN) developed a categorical specified period of time, rather than as a single
rank system57 for their Red Lists58, 59 to assess worldwide deterministic estimate for an unspecified time period as
species extinction risk based on population size and with rank methods, sensitivity analysis of population
distribution (see Mace and Collar83 for an application) parameters, or estimating the variance of modeling
(Figure 3). The IUCN system has been further developed analysis outcomes. However, the formal PVA approach
for local application to assess extinction and associated seldom accounts for propagation of error terms and
management risk within individual countries or other general lack of scientific knowledge on the ecology of a
artificial management units.41, 124 species.
The Nature Conservancy (TNC) developed a global For some authors, the process of PVA has been
categorical-rank extinction and management rating associated with determining the “minimum viable
system for the Natural Heritage Program that is based population” (MVP), which is presumed to be the
primarily on qualitative estimates of plant and animal minimum number of individuals that ensure a
population abundance and distribution.98 The system also population’s persistence for a specified length of time.150
ranks “site biodiversity significance” and “site protection The value of the MVP concept has proven very limited,
urgency” on a rank scale of 1-5 for use in developing however, because the “minimum” population size is not
management priorities. Other ordinal-rank models have a simple number, but varies with population parameters,
the acceptable probability of persistence, the length of
planning period,42, 79 and how populations and their
Extinct
environments change unpredictably. Also, acceptable
Extinct in the wild
persistence likelihoods and planning period durations are
functions of the amount of risk managers are willing to
accept in trading species viability for other values; thus,
(Adequate data)
the “minimum” viable population size largely is a function
Critically endangered of conservation policy instead of pure science.
(Threatened) Typically, quantitative PVA models are developed for
Endangered
single species because of model complexity and the need
Vulnerable
Conservation for detailed data on population dynamics and habitat
(Evaluated) dependent
relationships. Several less-quantitative approaches have
Lower risk Near threatened been developed to assess viability for multiple species—
Least concern these may be termed Population Viability “Assessments”
to distinguish them from the more stochastic and
Data deficient demographic modeling-based PVA process. The scale of
analysis in population viability assessments can vary from
Not evaluated
regions for wide-ranging species with sparse populations,
such as the northern spotted owl 94, 177 to local site
Figure 3. Rank categories of ecological risk developed by the conditions for less-mobile species with locally disjunct
International Union for the Conservation of Nature and populations.
Natural Resources (IUCN 1994) for the Red List of Population viability assessment was developed to
threatened and endangered species. accommodate the varying availability of species data and
 CHAPTER 19: SPECIES AT RISK 481

the needs and constraints of scientists and managers while alternatives.79, 87, 132 Decision and knowledge-based systems
meeting the basic criterion for estimating the likelihood differ from quantitative PVA models by using rigorous
of persistence over a specified period of time. For example, steps to capture and quantify expertise, and by explicitly
expert-opinion approaches were used to estimate the incorporating management (as opposed to ecological) risk.
likelihood of persistence of the northern spotted owl and As with PVA, decision analysis models are usually too
other species associated with late-successional forests in complex to use with more than 1 species at a time, and are
the coastal Pacific Northwest.33, 61, 94, 172, 177 Subsequent useful at watershed to regional scales.
regional assessments of multiple species for the interior Decision-tree risk analysis is the most commonly used
Columbia River basin,76 and southeast Alaska154 adapted procedure, and has been done for grizzly bear,88 black-
and modified methods developed for the Pacific footed ferrets,85 tigers,86 and Sumatran rhinos.87 Expected
Northwest.33 value of perfect and sample information, and Bayesian
These approaches typically have had expert panelists statistics46, 168 are other methods that may be useful for
score the relative abundance and distribution of habitats wildlife risk analysis.93 Expert systems and other expert
or populations at particular points in time under advisory approaches are gaining acceptance as important
alternative management scenarios. The scores are tools when used to analyze risk related to specific
described by several outcome classes to depict the problems of wildlife resource management.93, 108
likelihood of persistence under management alternatives.
Uncertainty of viability likelihood in one approach76 was Toxicological risk assessments
estimated as the variance among and within viability The literature on toxicological risk assessments for wildlife
scores of expert panelists. In a current effort, however, this is relatively well established. 124, 143, 175, 176 Procedural
“black box” of expert opinion is being opened by using standards have been set by the U.S. Environmental
Bayesian belief network models of species’ interactions Protection Agency for laboratory and field risk assessment
with their key environmental correlates as a way to of environmental contaminants on wildlife.35, 176 Risk is
measure habitat or population potential across landscapes quantified175 as (1) dose-response evaluated by, (a) the ratio
over time (B. Marcot, pers. comm., USDA Forest Service, of environmental concentration over lethal concentration
Portland, Oregon). resulting in 50% mortality of test individuals (LC50), and
Model forms of PVA quantify the probability of (b) the ratio of toxicant consumption over the lethal dose
persistence for single species by using mathematical and resulting in 50% mortality of individuals (LD50); and (2)
simulation models. Those forms explicitly model the by the environmental exposure quantified as the amount
probability of persistence as a function of population of toxicant in the environment over the amount required
structure and distribution, genetic structure or processes, for LD 50 . Environmental exposure is a function of
habitat quality and quality, or life history attributes. population and life history attributes, as used for other
Uncertainty can be evaluated with sensitivity analysis that risk assessments. Risk increases as these ratios approach
varies parameter values within a range of known or 1. Those metrics of risk do not quantify, however, the
hypothesized values.1, 79, 104, 167 Random, or stochastic, interaction of toxicants with other factors or the important
effects are modeled from hypothesized (or, less commonly, population consequences of sub-lethal effects.
empirically fit) probability density functions. Several Single species are the focus of toxicological assessments.
programs have been developed to model PVA, each with Application is usually at small scales of individual
particular strengths and weaknesses.78, 79, 113 Examples of contaminant sites or treatment areas, as in the case of
PVA can be found in the literature for birds,3, 7, 11, 12, 13, 23, 45, 48, pesticides. Watershed or regional applications would sum
89, 103, 142
mammals,5, 16, 21, 22, 39, 52, 67, 69, 72, 80, 97, 121, 159, 187 reptiles,162 the lower-order effects at individual sites. Population
fish,8, 26, 31, 100, 140 invertebrates,116 and plants.10, 17, 19, 109, 117 models are increasingly being used to assess toxicological
It is our considered opinion that the quantitative PVA risk.30, 40, 50, 68, 127, 158, 179
approach is useful primarily in uncommon cases where
species have been previously screened to be at high risk, Advantages and Disadvantages of Methods
and where data on their population demography and WHR and rank models. Simple WHR habitat models or
genetics, and variations in their environment and habitats rank models that rate species risk to develop priorities for
are known and can be adequately modeled. For most cases, conservation or assess generalized risk from specific
however, the more qualitative population viability threats (e.g. forest fragmentation) have the advantages of
assessment approach is likely to be adequate for policy being broadly applicable, rapid, and incorporating
decision-making at broad scales. ecological data, theory, and local expert knowledge when
research data are few. Simple ranking techniques usually
Decision Analysis and are easily understood by managers in other disciplines,
Knowledge-Based Systems and can be applied to estimate risk for many species over
Decision analysis integrates extinction risk assessment large areas.
with management risk assessment. Biological and However, the generality of the procedure brings with
management uncertainty, in terms of probabilities and it disadvantages. Information based on expert opinion
their variation, are used to calculate expected values of may be biased by personal experience or tacit motivational
species abundance under several management bias of the contributors. When a common set of life history,
 482 Wildlife–Habitat Relationships in Oregon and Washington

population, or habitat attributes are used to examine risk Improvements Needed in

for a group of species, there is the potential that an attribute Assessing Wildlife Risk
that is critical for the persistence of one species might be Improvements in wildlife risk assessment can be made in
missing because it was generally considered not several areas. There is an enduring need for better
important. Likewise, a model developed for one place information on species’ population structure and the range
might fit poorly in other areas. Simple WHR and rank of variability and response to environments over space
models do not incorporate variation in risk associated with and time. Also needed for model development are data
variability of species populations, environmental on habitat selection and the impacts of management on
conditions, or uncertainty in species responses that is both population structure and habitat selection. Data also
necessary for application to small scales of management, are needed for verification of models—a critical and often
such as stands and sub-watersheds. overlooked aspect of model building because of the
PVA models. Quantitative risk analyses, such as single- difficulty of obtaining independent data at appropriate
species PVA models, require detailed life history and spatial or temporal scales. Long-term datasets, such as
population information, consider variation in assessing Breeding Bird Surveys, waterfowl surveys, and other data
risk, and often examine effects of management from formal monitoring programs may provide useful
alternatives. Variation in management and species data for model verification. Data for model building and
responses to risk factors is specifically estimated as verification can be gained through conventional research,
likelihood, or through sensitivity analysis. PVA models but also through better monitoring of management
can help clarify assumptions, create testable hypotheses outcomes within the framework of adaptive
relative to model parameters and processes, synthesize management.54 Both approaches aim to provide reliable
and integrate knowledge, and demand explicit and knowledge using basic scientific methods; hence, the
rigorous reasoning.9 perception that research is for scientists and adaptive
These potential advantages of PVA models are often management for managers is a false dichotomy that
offset by poor documentation, high cost, or limitations of hinders progress in the field.
the methodology, which have been amply discussed in Just as important is the need for methods to better
the literature.2, 11, 24, 47, 66, 74, 104, 105, 113, 167 At the heart, too, estimate economic and social values of wildlife, and
persistence likelihood is not strictly an empirically explicitly incorporate the costs incurred by alternative
verifiable parameter—you cannot conduct a field study management practices. 118, 165 Central to the goal of
or experiment to verify the risk levels stated in a PVA ecosystem management is defining essential ecosystem
without putting at risk the very populations you are trying components, linking processes, and endpoints or desired
to conserve. Thus, PVA is used under the presumption future conditions of those properties.118
that its mosaic of models and concepts, which individually
can be derived from sound theoretical or empirical Characterizing Risk Species and
evidence on other species, together constitute a useful Their Attributes
approach for describing changes in some scarce species Below are several examples of methods for characterizing
of interest. species at risk, determining the specific attributes that
Decision analysis. Alternatively, highly structured distinguish high-risk species from other species, and
procedures of risk assessment using decision analysis have comparing risk from several management alternatives.
the advantage of explicitly stating key assumptions and The examples are not the only ways risk analysis might
possible management alternatives in a standard format.85, be done (see the methods section of this chapter), but are
87, 90, 92
Management actions and species responses are intended to illustrate approaches to the problems. The first
expressed as probabilities with a defined range of expected two examples use different but related procedures to
values and variation. Different types, sources, and units differentiate and classify imperiled, secure, and
of management and ecological information can be intermediate-risk species, and develop a rule set for
incorporated to facilitate management decisions. classifying other species. A third example uses principles
However, there are several disadvantages of decision discussed in the section of this chapter on criteria for
analysis when used in population viability assessments.92, identifying species at risk to score and rank potential risk
118
Probabilities are often difficult to accurately assess by based on life history characteristics, when prior knowledge
use of expert opinion, and assumptions to quantitatively of risk is unavailable. That same approach could be used
estimate probabilities may inject too much additional to rank management priorities. A fourth example describes
uncertainty. Also, small changes in probabilities may a population viability assessment used to assess land
greatly affect the results of the analysis. Not all management alternatives.
management alternatives and future environmental For the first three examples, we developed a database
conditions may be foreseen, thus biasing the management for a sample of 60 species that were selected based on
decision. Also, it may be difficult to define specific criteria shown in Appendix 1. A sample of 20 threatened,
management objectives for individual species and to endangered, or sensitive (TES) species in Oregon and
quantify outcomes with respect to habitat conditions from Washington (e.g., Photo 1) were combined with a sample
biological, and especially social and political, viewpoints. of 20 species we considered secure (e.g., Photo 2) and 20
 CHAPTER 19: SPECIES AT RISK 483

Photo 1. Bobolink (Dolichonyx oryzivorous) in Shrub-steppe

habitat. An example of a “Group I” species, i.e., species
known to be imperiled. (Photo: Bruce G. Marcot)

Photo 3. Sharptail snake (Contia tenuis) in Southwest

Oregon Mixed Conifer-Hardwood Forest habitat. An example
of a “Group III” species, i.e., species with intermediate
security. (Photo: Bruce G. Marcot)

Using Classification Trees to Determine

Influences on Species Risk
Photo 2. Ensatina (Ensatina eschscholtzii) in Westside This first example shows how a classification tree
Lowlands Conifer-Hardwood Forest habitat. An example of a
procedure 183 can be used to determine the relative
“Group II” species, i.e., species very likely secure. (Photo:
contribution of life history, population distribution, and
Bruce G. Marcot)
versatility attributes in the matrixes to known risk-group
membership. The results of that analysis, then, can be used
species that had potentially intermediate viability risk to develop a rule-set to predict risk-group membership
(e.g., Photo 3). for species with unknown risk potential. Our example
For the species in the example database, we extracted concentrates on comparing the attributes of TES species
a subset of the life history and habitat relationships fields with species in secure or potential intermediate-risk
from the matrixes (CD-ROM with this book) that categories. It is important to remember that this and the
correspond to attributes associated with variation in following examples are merely illustrations of what can
viability, as discussed in the section of this chapter on be done with the larger list of species and attributes that
criteria for identifying species at risk. We combined some might be used in a real situation.
matrix values for a few fields to somewhat simplify the The classification tree procedure worked in a way
example analyses. Where field values were given as similar to a dichotomous key used for plant identification:
“unknown” we used the most likely value, based on our it started with all the species in a group, then sequentially
knowledge of species, to completely fill the data matrix split each subsample based on the most discriminating
for the example analysis. We designated primary fields in variable until no significant splits could be performed. The
the example database as those most likely to account for “proportional reduction in error” (PRE) statistic described
risk levels based on our experience, and some secondary the goodness-of-fit, or strength, of the model, similar to
life history fields that could be important attributes of the conventional R2 statistic, varying between 0 and 1, with
species at risk. Other fields were purely for information 1 indicating a perfect fit of the model. The classification
to interpret the analysis. We calculated two “versatility” tree figure showed the splits and relative number of
indices of habitat cover and structural-type use from the species within each subsample. Crosstabulation and t-tests
data given in the habitat matrixes. The indices were also were used to examine the relationship between each
calculated as the percentage of all habitat cover or life history and versatility field and risk group
structural types that the animal is known to use.
 484 Wildlife–Habitat Relationships in Oregon and Washington

Risk groups

Number of species
14
12 Imperilled
Secure
10
Intermediate
8
6
4
2
0
contiguous gaps patchy isolated scarce
Population distribution
Figure 5. Numbers of vertebrate species in the risk analysis
sample dataset by population distribution category and
viability risk group.

Other than habitat and structural versatility, population

distribution was the only other variable that differed
among risk groups (Figure 5). Imperiled species largely
had patchy, isolated, or scarce distributions; whereas,
Figure 4. Classification tree of 60 vertebrate species in Oregon secure species mostly were contiguously distributed or
and Washington known to be imperiled, secure, or of potential had gaps in their distribution. Distributions of
intermediate viability risk. The classification used life history intermediate species varied between those 2 extremes.
and habitat versatility attributes from the matrices. See text
for an explanation of the species and variable selection A Rule Induction Approach:
processes. Risk groups are represented by bars in the order of SARA—Species At Risk Advisor
imperiled, secure, and intermediate; the height of the bars This example discusses the use of a rule induction
indicates the number of species in each group and box, approach developed by B. Marcot to identify species at
starting with 20 species in each group. risk based on their life history attributes. Rule induction
is a knowledge-base modeling approach for devising an
membership. Differences among groups were reported if optimal rule to distinguish among categories based on a
significant at P<0.05. set of example cases. For example, Stockwell et al.166 used
Imperiled species in our sample dataset were best rule induction to predict different density levels of gliders
distinguished from the other species by their use of few (Petauroides volans) in Australia. In our example, the rule
structural habitat types, as reflected by low structural distinguishes among the three risk levels (imperiled,
habitat versatility (PRE=0.125; Figure 4). Structural habitat secure, and intermediate) and the examples are the 60
versatility of imperiled species (mean=41%) was almost sample species used in the data set.
half that of secure species (mean=75%), and somewhat There are many kinds of rule induction algorithms,
less than secure and intermediate species combined particularly in the area of knowledge-base and expert
(mean=55%). Intermediate-risk species when considered systems programming and statistical analysis.60 Used here
alone could not be neatly separated from imperiled or is the Quinlan ID-3 rule-induction algorithm.123, 129 This
secure species. That pattern was evident in the approach is similar to the classification tree example
classification tree (Figure 4) where intermediate-risk presented above in that it identifies an optimal division
species were split nearly equally between imperiled and of the known cases based on successively identifying
secure groups on the basis of structural versatility. variables and their values that best distinguish among
Although not appearing critical for separating risk groups, remaining cases. What results is an optimal rule, much
habitat cover-type versatility had the same pattern of like a botanical key, that leads to final categories of the
differences among groups, which might be expected with three species risk levels. The ID-3 rule-induction algorithm
a high correlation between structural and habitat and other related knowledge-base approaches, however,
versatility (r=0.76). differ from the classification tree analysis in that they can
In some cases, population distribution and habitat handle both categorical and ordinal data, as well as
cover versatility could be used to distinguish imperiled continuous (ratio-scale) data.
from other species. Imperiled and intermediate-risk The knowledge-base expert system programming
species with low structural versatility on the left side of package 1st Class (1st Class Expert Systems, Inc., Wyland,
tree appeared to have similar life history and distribution MA; Release 3.65) was used to do the rule-induction
traits and could not be easily separated by additional analysis. In general, rule induction and use of knowledge
attributes from the sample dataset (Figure 4). However, base modeling differ from classification tree analyses in
among the species with high versatility on the right side several ways. First, a knowledge base programming
of the tree, scarce population distribution and relatively approach can produce a user-friendly interactive query
low habitat cover versatility distinguished most of the few system based on the optimal rule. In this example, the
imperiled species from secure or intermediate species. query system produced is named SARA—the Species At
 CHAPTER 19: SPECIES AT RISK 485

Risk Advisor. Second, the knowledge base programming Strigiformes), and that it uses only 20% of all available
approach can account for uncertainty and unknowns that habitat types (HabVers < 34.5%). The SARA rule set, and
cannot be included in a classification tree analysis. the knowledge base expert system, would key this species
Uncertainty is handled by depicting likelihood of out to belonging to Group I, that is, a potentially imperiled
assigning each example to each outcome category. It then species. This is because at least 1 known case from the set
uses these likelihoods to calculate a probability of an of 60 example species had these conditions and was known
unknown case (e.g., a species not included in the example to be imperiled (state or federally listed), the northern
data set) belonging to each outcome category (each risk spotted owl. Whether another owl with these same
level). Third, during the interactive query, a knowledge attributes would be similarly imperiled is postulated by
base model such as produced by 1st Class can allow the the SARA analysis but would bear further evaluation. So
user to enter “do not know” or “no information” to any the purpose of the SARA model, particularly if developed
question, and from there the program determines the next among a broader set of species, is to provide a tool for
best sequence of as-yet-unasked questions to optimally initially screening species for further evaluation.
determine the outcome category. Fourth, because it is not An analysis was also done that combined the secure
limited to statistical assumptions inherent in the and intermediate categories and compared this
parametric classification tree approach, the ID-3 rule- combination to the imperiled category. This more simply
induction algorithm is not limited to a specific number of determined which life history attributes might be used to
variables given a fixed number of example cases (e.g., 5- identify imperiled species although it could not
10 example cases are needed for each variable considered differentiate between secure and intermediate categories.
in the classification tree approach). Fifth, “classification In this analysis, a somewhat different set of attributes were
trees,” being a strictly parametric statistical analysis identified as key differentiating factors: habitat versatility,
procedure, provides confidence values describing the structural versatility, clutch or litter size, summer site
degree to which each factor accounts for classification; fidelity, breeding status, population distribution, and
knowledge base approaches typically do not. Thus, rule taxonomic order. That a different set of factors resulted in
induction and classification trees type classification this analysis as compared with the above analysis is not
analyses are complementary approaches (see Conclusions unexpected when the same example species cases were
section). combined in a different way. In this example, one possible
SARA uses 29 life history attributes, 1 taxonomic combination of attributes that would lead to identifying
attribute, and 60 example species. Its optimal rule is the imperiled category is if a species had a structural
displayed in Figure 6. It should be remembered that this versatility < 54% (StrucVers<54), a mean litter or clutch
analysis is only an example of the kind of assessment that size of 3 or less (NLitt_Clut=nlit3), and documented or
can be done to determine how life history attributes might suspected breeding in both Washington and Oregon
contribute to viability risk levels. Similar analyses can be (BreedStatus=bs3). Again, this should be taken only as an
done on other species’ attributes such as population size, example of the kind of analysis possible; a fuller evaluation
trend, density, and so on, if data are available. Results of with more species would likely produce more reliable
this analysis, and the SARA model, should not be results that could be evaluated against a larger set of
interpreted as definitive analyses of the influence of life species with known population status.
history attributes on risk levels; at best, this analysis poses
testable hypotheses, and urges a fuller assessment of a A Rank-Model Example
fuller set of species. This example shows how population and life history
Analysis results suggest that 9 life history attributes attributes can be used to develop a simple rank model
and 1 taxonomic attribute can account for, and differentiate (see methods section of this chapter) to evaluate potential
among, the 3 viability risk levels for the 60 example extirpation risk when there is no a priori knowledge of
species. These attributes are population distribution, risk, as was known in the previous examples. We scored
foraging substrate, elevational range, age at first risk for each value of the primary life history and
reproduction, structural versatility, habitat versatility, versatility fields in the example dataset on a scale of 1 to
landscape use, migration distance, and taxonomic order. 10 (Appendix 2), with 10 being the highest risk, using the
It should be noted that rule induction algorithms generate principles discussed in the section of this chapter on
results whether or not they are statistically valid, so the criteria for identifying species at risk. For example, species
relative power with which each of these life history with high annual production of offspring or numbers of
attributes contributes to predicting viability risk levels is litters were scored lower risk for that attribute than species
not known. This is where the parametric statistical analysis with low productivity. The rationale was that a population
of classification trees can complement this analysis. of species with high fecundity likely would more easily
An example of how SARA can be used is for identifying recover from disturbance in their environment, i.e., be
a species of unknown risk level. For example, suppose a more resilient, and less susceptible to local extirpation than
species has a population distribution consisting of gaps species with low productivity. We then summed the scores
with habitat broadly distributed but with interruptions of each field to calculate a total score for each species.
causing some population isolation (PopnDistrb = pop2, Species with high scores would potentially be at greater
as in Figure 6 and Appendix 3), that it is an owl (Order = risk of extirpation than species with low scores. Differences
 486 Wildlife–Habitat Relationships in Oregon and Washington

A. Population distribution is contiguous.

B. The organism forages underwater or aerially, or foraging substrate is unknown.
C. The upper elevation range of typical or regular occurrence is up to 1000 ft. (no identification)
CC. The upper elevation range of typical or regular occurrence is up to 3000 ft. (no identification)
CCC. The upper elevation range of typical or regular occurrence is up to 5000 ft. Group III
CCCC. The upper elevation range of typical or regular occurrence is >5000 ft. Group I
BB. The organism does not forage underwater or aerially, and foraging substrate is not unknown.
C. The average age at first breeding (females) is <6 months. (no identification)
CC. The average age at first breeding (females) is 1 year. Group II
CCC. The average age at first breeding (females) is 2 years.
D. The structural versatility of the species is <99. Group II
DD. The structural versatility of the species is >=99. Group III
CCCC. The average age at first breeding (females) is 3 years. Group III
CCCCC. The average age at first breeding (females) is 4+ years. Group II
AA. Population distribution consists of gaps.
B. The taxonomic order is Caudata.
C. The structural versatility of the species is <90.50. Group II
CC. The structural versatility of the species is >=90.50. Group III
BB. The taxonomic order is Anura. (no identification)
BBB. The taxonomic order is Squamata. Group III
BBBB. The taxonomic order is Falconiformes.
C. The average age at first breeding (females) is <6 months. (no identification)
CC. The average age at first breeding (females) is 1 year. Group II
CCC. The average age at first breeding (females) is 2 years. Group III
CCCC. The average age at first breeding (females) is 3 years. Group I
CCCCC. The average age at first breeding (females) is 4+ years. (no identification)
BBBBB. The taxonomic order is Charadriiformes. Group II
BBBBBB. The taxonomic order is Strigiformes.
C. The habitat versatility of the species is <34.50. Group I
CC. The habitat versatility of the species is >=34.50 and < 50.00. Group III
CC. The habitat versatility of the species is >=50.00. Group II
BBBBBBB. The taxonomic order is Apodiformes.
C. The habitat versatility of the species is <53.50. Group II
CC. The habitat versatility of the species is >=53.50. Group III

Figure 6. The example, optimized rule from the SARA among species or species groups in risk score or life history
(Species At Risk Advisor) example knowledge base model that attributes were considered significant at P<0.05.
determines species’ viability risk levels based on their life The distribution of risk scores was very normally
history attributes. This rule was induced from analysis of 60 distributed, ranging from 33 to 94 points with a mean of
example wildlife species in Washington and Oregon, and is 65 (Figure 7). A preponderance of low scores would have
only an example of the type of analysis possible by using indicated few species likely to be negatively affected by
knowledge base, rule-induction analysis. The decision points management; whereas a preponderance of species with
in the rule correspond to the questions and states shown in high risk scores would indicate a potentially large risk of
Appendix 3. Group I = imperiled species; Group II = secure action. Ranking species by their risk score from high to
species; Group III = intermediate species. See text for
low would indicate priorities for further analysis or field
explanation of methods.
work to accurately determine the impacts of management.
How well does this risk scoring and ranking method
coincide with the species risk group, as defined in the
sample dataset and analyzed in earlier examples? Risk
scores for species showed a similar relationship among
risk groups to those found with the classification and
SARA procedures. Imperiled species on average had
higher risk scores (mean=72) than secure species
(mean=56) (Figure 8). Intermediate-risk species (mean=68)
did not differ from imperiled species in average risk score,
but did have higher scores than secure species. That
pattern was largely a result of similarly high values for
 CHAPTER 19: SPECIES AT RISK 487

BBBBBBBB. The taxonomic order is Piciformes.

C. The habitat versatility of the species is <50.00. Group III
CC. The habitat versatility of the species is >=50.00. Group II
BBBBBBBBB. The taxonomic order is Passeriformes.
C. It is a “patch” species, likely using only 1 homogenous habitat patch during the life cycle. Group III
CC. It is a “mosaic” species, likely using an aggregate of habitat patches but 1 structural stage.
D. The migration or seasonal movement is <100 km. (no identification)
DD. The migration or seasonal movement is 100 - 1000 km. Group I
DDD. The migration or seasonal movement is >1000 km. Group II
DDDD. The species is non-migratory. Group II
CCC. It is a “generalist” species, likely using all or many patch types, & >1 structural stage. Group III
CCCC. It is a “contrast” species, likely requiring contrast between 2 structural stages in close proximity.
(no identification)
BBBBBBBBBB. The taxonomic order is Rodentia.
C. The structural versatility of the species is <28.50. Group I
CC. The structural versatility of the species is >=28.50. Group III
BBBBBBBBBBB. The taxonomic order is Carnivora. Group II
AAA. Population distribution consists of patchily distributed populations.
B. The average age at first breeding (females) is <6 months. Group II
BB. The average age at first breeding (females) is 1 year. Group I
BBB. The average age at first breeding (females) is 2 years. Group III
BBBB. The average age at first breeding (females) is 3 years.
C. The habitat versatility of the species is <34.50. Group I
CC. The habitat versatility of the species is >=34.50. Group II
BBBBB. The average age at first breeding (females) is 4+ years. Group III
AAAA. Population distribution consists of isolated population(s).
B. The migration or seasonal movement is <100 km. Group III
BB. The migration or seasonal movement is 100 - 1000 km. Group III
BBB. The migration or seasonal movement is >1000 km. Group I
BBBB. The species is non-migratory. Group I
AAAAA. Population distribution is scarce.
B. The habitat versatility of the species is <16.00. Group III
BB. The habitat versatility of the species is >=16.00. Group I

both imperiled and intermediate species in the life history species, and some secure species. Only 9 of the 60 species
field describing population distribution, as noted in the clustered as high-risk, with equal numbers of imperiled
classification tree example. and intermediate risk species. The poor match between
Could we use risk scores to create our own, perhaps cluster and a priori risk group, in some cases, indicates
better, risk groups instead of using existing legal or that legal definitions of risk may not always be adequate
administrative definitions of risk, which might reflect to define risk, and unlisted species might also need to be
political as much as ecological conditions? We used cluster analyzed.
analysis to create three new “risk” clusters, or groups, of
species based only on our risk scores, and then we Population Viability Assessments:
compared the clusters with the previously defined risk The Interior Columbia Basin
groups to see how well they matched the old groups. A final example is taken from the population viability
Cluster analysis identified low- (mean=51), assessment of draft management alternatives for the
intermediate- (mean=68), and high-risk (mean=84) species Interior Columbia Basin Ecosystem Management Project.76
clusters (Table 1); but, cluster membership did not always The purpose of the assessment was to determine the
match the a priori-assigned risk groups. That result might degree to which habitat conditions on lands administered
be expected considering the inability to perfectly separate by the Forest Service and Bureau of Land Management
species in imperiled, intermediate-risk, and secure risk within the interior Columbia River basin contribute to
groups in the earlier examples. The low risk cluster long-term persistence (at least 100 years) of select plant
included about 60% of the secure species, 20% of the and animal species of conservation concern. Secondarily,
intermediate-risk species, and 10% of the imperiled they examined the extent to which other lands and other
species. Most species clustered as moderate-risk, with influences might affect populations.
nearly equal numbers of imperiled and intermediate-risk
 488 Wildlife–Habitat Relationships in Oregon and Washington

Figure 7. The distribution of risk scores for 60 species in the

example dataset using the methodology described for rank Figure 8. The distribution of risk scores, estimated by the rank
model procedures. Risk scores were estimated by scoring each model procedure, by viability group (imperiled [I], secure [II],
life history, population, or habitat attribute in the example intermediate [III]) determined a priori for the example dataset
dataset for its potential contribution to extirpation risk, then of 60 species. Boxes indicate the mid-range (50% quartile) of
summing the attribute scores for a single risk estimate. data values around the median; notches in the box are the
confidence interval around the median. Whiskers on the boxes
show the outside quartiles, and asterisks show outliers.
Table 1. Crosstabulation of risk group memberships
based on clustering risk scores of life history
attributes and habitat versatility, compared with risk The assessment was not a quantitative population
groups in the sample dataset based on legal or viability analysis, as it did not employ an explicit model
administrative designations of threatened, of genetic or demographic risk to species persistence.
endangered, or sensitive status and expert opinion. Rather, the qualitative assessment was a structured and
reasoned series of judgments about projected amounts and
Risk Score Cluster distributions of habitat and the likelihood that such habitat
would allow populations to persist over the long term.
Risk group 1 2 3 Total
(sample dataset) (low) (high) (moderate) Thus, it met the essential criterion of a population viability
assessment to provide an estimate of the likelihood that a
population will persist to some arbitrarily chosen future
I (Imperiled) time.
Count 2 4 14 20 Species assessments were based on expert opinion
% within Group I 10% 20% 70% 100% about the likely outcome for species and their habitats
II (Secure) under a variety of possible management alternatives.
Count 12 1 7 20 Expert judgments were solicited from eight expert panels.
% within Group II 60% 5% 35% 100% The panels were provided with information on the species
distributions, habitat relationships, and known population
III (Intermediate) trends, and with information on the effects of management
Count 4 4 12 20
alternatives on species macro- and micro-habitat elements.
% within Group III 20% 20% 60% 100%
Based on that information, the experts were asked to make
Total two judgments. The first judgment rated the species’ likely
Count 18 9 33 60 distribution based only on habitat conditions on the
% within Group 30% 15% 55% 100% federal lands and the natural history characteristics of the
species. The second was a cumulative effects analysis of
the likely condition of species populations across all
ownerships. Factors considered included demographic
characteristics, responses to varying qualities of habitat
for specific life functions, types and ranges of seasonal
and permanent movements, genetic characteristics, and
biotic interactions (e.g., competition, predation,
herbivory).
Expert judgments were registered through a process
of likelihood voting using a structured outcome scale. The
 CHAPTER 19: SPECIES AT RISK 489

Table 2. Viability outcomes used for the population weighted mean outcome was also calculated. This was
viability assessment of draft management calculated by assigning a value to each of the outcome
alternatives for the Interior Columbia Basin categories (Outcome 1 value=1, Outcome 2 value=2, etc.),
Ecosystem Management Project.76 multiplying the mean likelihood of that outcome by its
assigned value, adding these products for all outcomes,
and then dividing by 100. Outcomes were considered
Outcome 1.
improving or declining from the historical to current, or
Habitat is broadly distributed across the planning area with
from current to future, periods if they changed at least
opportunity for continuous or nearly continuous occupation by
0.5, a value corresponding to one standard deviation of
the species and little or no limitation on population interactions.
the mean outcome. The assessment did not provide a
Outcome 2. simple determination of what does and does not constitute
Habitat is broadly distributed across the planning area but gaps a “viable” population. This was considered a strength of
exist within this distribution. Disjunct patches of habitat are the process rather than a weakness, as there are no simple
typically large enough and close enough to other patches to thresholds for viability, particularly when assessments are
permit dispersal among patches and to allow species to interact done on a broad array of taxa. Rather than providing a
as a metapopulation. simple determination, this assessment described likely
Outcome 3. future conditions for species and habitats and provided
Habitat exists primarily as patches, some of which are small or for comparison of those conditions to current and
isolated to the degree that species interactions are limited. Local historical conditions. Lack of a simple determination,
subpopulations in most of the species’ range interact as a however, added complexity to the job of interpreting the
metapopulation but some patches are so disjunct that results and using them in a decision-making framework.
subpopulations in those patches are essentially isolated from The authors recommended that interpretation of the
other populations. results emphasize comparison of the projected future
Outcome 4.
conditions under the alternatives to historical and current
Habitat is typically distributed as isolated patches, with strong conditions.
limitation in interactions of populations among patches and A variety of cautions must be applied to the
limited opportunity for dispersal among patches. Some local interpretation of this form of assessment. These cautions
populations may be extirpated and rate of recolonization will fall into four areas: (1) the broad geographic and temporal
likely be slow. scale of the analysis limits local inference; (2) the resolution
of the data and planning guidance define the level of
Outcome 5. confidence in results; (3) limitations on the ability to infer
Habitat is very scarce throughout the area with little or no
population results from habitat analysis; and (4) gaps in
possibility of interactions among local populations, strong
knowledge limit confidence or geographic scope of
potential for extirpations, and little likelihood of recolonization.
inference. Lehmkuhl et al.76 give a detailed discussion of
these cautions and assumptions.
outcome scale depicted 5 distinct possible outcomes for
the habitat or population, each representing points along Conclusions Drawn from the
a gradient ranging from a broadly distributed condition Example Analyses
with high likelihood of persistence to a poorly distributed The examples were meant to illustrate a practical range of
condition with high likelihood of extirpation (Table 2). For risk analysis methods that would be relatively easy to
each judgment, each expert spread 100 likelihood votes implement using the data matrixes in this book and
across these 5 outcomes (Table 3). Placing 100 votes on a conservation theory. The examples focused on frequently
single outcome indicated certainty in that outcome; encountered situations when the taxonomic scope is fairly
whereas, spreading the votes among several outcomes broad (i.e., many species need to be analyzed), and when
indicated less certainty in any one outcome. Consensus procedural, or analytical, complexity may be limited by
among panelists was not an objective of the process; available data. The geographic scale (extent) of application
moreover, the independence of experts’ judgments was would usually be large—at watershed or larger scales.
necessary to assess the uncertainty (standard deviation) The first two examples used data in related procedures
of viability likelihood. Uncertainty included two to differentiate and classify imperiled, secure, and
components, variation of likelihood distributions among intermediate-risk species, and to develop a rule set for
panelists and the spread of likelihood points among classifying other species of unknown risk. The third
outcomes by each panelist. example used species risk theory and data to score and
There were 2 primary analyses performed on the data rank potential risk based on attributes of life history and
derived from the panels. First, the mean likelihood scores population distribution, when prior knowledge of risk was
for all experts for each of the outcomes was calculated. unavailable. That same type of model could be used to
For example, if there were 4 experts on a panel, and their rank management priorities based on criteria other than
likelihood votes for outcome 2 for a particular species were extirpation risk, e.g. social or economic value. The fourth
30, 30, 60, and 40, then the mean likelihood score was 40. example described a qualitative population viability
Next, to summarize mean likelihood for each species, a assessment that linked in the data matrixes with expert
 490 Wildlife–Habitat Relationships in Oregon and Washington

Table 3. Example of the likelihood voting system used to assess viability

outcomes for selected species of conservation concern for the Interior
Columbia Basin Ecosystem Management Project.*76

Species: Flammulated Owl Management Alternative

Outcome Historic Current Traditional Restore Reserve

1 contiguous 1 0 0 0 0
2 gaps 75 2 1 26 15
3 patchy 20 40 20 60 50
4 isolated 4 38 39 13 27
5 scarce 0 20 40 1 8
Total score 100 100 100 100 100
* One hundred likelihood points are distributed across the five outcomes for each time
period or alternative. The distribution of points reflects the certainty of a particular
outcome.

opinion to assess land management alternatives. seem to hold equally across the 4 taxonomic classes. That
Other more quantitative methods, such as demographic also was the general finding of Russell et al.,145 who found
PVA models, are valuable tools to address some of those that distributions of extinctions and threat classifications
risk issues and should not be ruled out. They were not were clustered unevenly within certain genera and
illustrated because their complexity demands information families, particularly in taxa that contain few species.
far more sophisticated than is available in the data Whether taxonomic affiliation and diversity alone
matrixes, and that is nonexistent for the vast majority of predisposes species to certain risk levels, or patterns of
species that the manager has to consider. Quantitative PVA habitat use and population distribution are themselves
models will be most appropriate when the focus of directed by life history characteristics, is unclear and needs
management is a single species suspected or known to be further analysis.
at risk, and when reliable data are available to model The first three analyses were developed mostly as
population and habitat dynamics under management examples of the kinds of evaluations that could be done
alternatives. with data to test how life history attributes might
It is first and foremost necessary to remember that the contribute to risk levels. The rank model, the third
examples were meant simply to illustrate some of the ways example, also ranked species by potential risk. More
information in the data matrixes can be used to determine specific and more thorough analyses can follow these
factors that might contribute to risk levels of species, or to examples by: (1) focusing on species within specific
rank species in terms of risk. Further and more complete individual habitats, structural conditions, physiographic
analyses, using these methods or others described in this provinces, taxonomic groups, known risk categories, or
chapter, are really needed to verify our initial findings on geographic areas; (2) rerunning such analyses for all such
how life history or taxonomic categories could predispose species within the data matrixes; (3) better determining
some species to various risk levels. Those conclusions the mechanisms by which life history traits influence
should be taken as tentative working hypotheses. habitat selection and population distribution; and (4)
The following is an example of how one might draw better determining the influence of life history traits on
conclusions from the example risk analyses. Conclusions risk levels among species within genera and families.
from these types of analyses might differ if the focus of Following more complete analysis, the results can be
such analyses shifted to specific taxonomic or functional integrated into a broader risk management framework.
groups. That given, it could be significant that the best The risk analysis portion of a decision evaluation can
distinguishing features of the three species groups include those life history traits and habitat selection
analyzed in the first three examples mostly refer to habitat behavior of species found to portend risk levels, and
selection, habitat breadth, and population distribution, thereby determine how alternative management actions
and not as much to inherent life history attributes per se. might influence the species’ habitat use and population
At the start of this analysis, we expected at least some life response. In the risk management phase, the decision-
history attributes, such as age at first reproduction, maker then would have explicit information on which
reproduction rates, site fidelity, and movement distances, species might be at greater risk.
to predict risk level. Life history attributes still may be The qualitative PVA example for the Columbia River
important to distinguish risk levels among species within Basin showed how information from the first three types
genera or families, which remains to be formally tested of analyses could be melded with matrix data and expert
with an expanded analysis of the entire species dataset. opinion to assess and score the effects of different land
However, the importance of life history attributes for management alternatives at a broad regional scale. That
characterizing risk within taxonomic groups does not information, then, would inform decision makers on how
 CHAPTER 19: SPECIES AT RISK 491

best to manage risk. The qualitative expert-opinion process Literature Cited

used in the example could be developed further by 1. Akçakaya, H. R., and J. L. Atwood. 1997. A habitat-based metapopulation
formalizing the relationships between life history, habitat, model of the California gnatcatcher. Conservation Biology 11:422-434.
and population status in a decision analysis or knowledge- 2. ———, M. A. Burgman, and L. R. Ginzburg. 1997. Applied population
based system. Then, the risk analysis process could be ecology: principles and computer exercises using RAMAS EcoLab 1.0
Applied Biomathematics, New York, NY.
easily refined or repeated as management alternatives are 3. ———, M. A. McCarthy, and J. L. Pearce. 1995. Linking landscape data
modified from public input or changing agency priorities. with population viability analysis: Management options for the helmeted
Formalizing such expert models also has the advantage honeyeater Lichenostomus melanops cassidix. Biological Conservation
of explicitly stating the relationships between species 73:169-176.
attributes, habitat, and risk, thus, opening the scientific 4. Allendorf, F. W., D. Bayles, D. L. Bottom, K. P. Currens, C. A. Frissell, D.
“black box” for review and agreement among interested Hankin, J. A. Lichatowich, W. Nehlsen, P. C. Trotter, and T. H. Williams.
1997. Prioritizing Pacific salmon stocks for conservation. Conservation
parties that the best available science is being used for
Biology 11:140-152.
management. 5. Armbruster, P., and R. Lande. 1993. A population viability analysis for
African elephant (Loxodonta africana): how big should reserves be?
Conclusions Conservation Biology 7:602-610.
6. Ballou, J., and K. Ralls. 1982. Inbreeding genetics and juvenile mortality
Wildlife managers do risk analyses that vary in spatial
in small populations of ungulates: a detailed analysis. Biological
scale, taxonomic scope, and complexity of information Conservation 24:239-72.
needs and methods—it is a daunting task. Moreover, the 7. Beissinger, S. R. 1995. Modeling extinction in periodic environments:
public is scrutinizing resource management actions that Everglades water levels and snail kite population viability. Ecological
can affect wildlife viability, and is asking for the best Applications 5:618-631.
available science to analyze the effects of those actions. 8. Botsford, L. W., and J. G. Brittnacher. 1998. Viability of Sacramento River
Armed with a basic understanding of the theory and winter-run chinook salmon. Conservation Biology 12:65-79.
9. Boyce, M. S. 1992. Population viability analysis. Annual Review of
criteria for identifying species at risk, managers and
Ecology and Systematics 23:481-506.
scientists can use available datasets and analytical tools 10. Bradstock, R. A., M. Bedward, J. Scott, and D. A. Keith. 1996. Simulation
to meet that challenge at different levels of geographic of the effect of spatial and temporal variation in fire regimes on the
and ecological scale. population viability of a Banksia species. Conservation Biology 10:776-
Species are at risk from a combination of systematic 784.
and stochastic processes that affect population size and 11. Brook, B. W., L. Lim, R. Harden, and R. Frankham. 1997a. Does
distribution. Systematic processes are usually the ultimate population viability analysis software predict the behaviour of real
populations? A retrospective study on the Lord Howe Island Woodhen
cause of extirpation, and occur when something essential (Tricholimnas sylvestris, Sclater). Biological Conservation 82:119-128.
is removed from the environment, such as habitat, or when 12. ———, ———, ———, ———. 1997b. How secure is the Lord
something lethal is inserted into the environment, such Howe Island Woodhen? A population viability analysis using VORTEX.
as a new predator or poison. Stochastic, or random, Pacific Conservation Biology 3:125-133.
processes are usually the proximate cause of extirpation 13. Brooker, L. C., and M. G. Brooker. 1994. A model for the effects of fire
in populations already made small by systematic and fragmentation on the population viability of the Splendid Fairy-
wren. Pacific Conservation Biology 1:344-358.
pressures. The vulnerability, or risk, for species is a
14. Brown, E. R., editor. 1985. Management of wildlife and fish habitats in
function of how systematic and stochastic processes forests of western Oregon and Washington. Pub No. R6-F&WL-192-
interact with habitat selection behavior, and the 1985, 2 vols. USDA Forest Service, Pacific Northwest Region, Portland,
demographic and life history attributes of the species. OR.
Habitat selection, habitat breadth, and population 15. Burgman, M., D. Cantoni, and P. Vogel. 1992. Shrews in suburbia an
distribution likely are among the most important application of Goodman’s extinction model. Biological Conservation
influences on risk level. 61:117-123.
16. ———, S. Ferson, and D. Lindenmayer. 1995. The effect of the initial
Understanding the processes underlying species risk
age-class distribution on extinction risks: implications for the
allows managers to select the right analysis tools. While reintroduction of Leadbeater’s possum. Pages 15-19 in M. Serena,
much of the conservation literature on wildlife risk editor. Reintroduction biology of Australian and New Zealand fauna.
analysis focuses on data-intensive quantitative PVA Surrey Beatty, Chipping Norton, England
models, there are many situations for which that tool is 17. Burgman, M. A. 1990. A stage-structured stochastic population model
unsuitable in terms of number of species to be analyzed, for the giant kelp Macrocystis-pyrifera. International Colloquim on
the data available for the analysis, or the time and money Dynamical Models in Biology, Lausanne, Switzerland, September 13-16,
1988. Memoires De La Societe Vaudoise Des Sciences Naturelles
available to do it. Fortunately, there are datasets and 18:355.
analytical tools that can meet the risk analysis needs of 18. ———, S. Ferson, and H. R. Akçakaya. 1993. Risk assessment in
managers for different levels of taxonomic and procedural conservation biology. Chapman and Hall, London, England.
complexity. The data matrixes included in this book are 19. ———, and B. B. Lamont. 1992. A stochastic model for the viability of
an important basic resource for risk analysis that can be Banksia-cuneata populations environmental demographic and genetic
linked with analytical methods in this chapter. Results of effects. Journal of Applied Ecology 29:719-727.
20. Burke, R. L., and S. R. Humphrey. 1987. Rarity as a criterion for
such risk assessments will provide the essential
endangerment in Florida’s fauna. Oryx 21:97-102.
information for decision-makers about the alternatives for 21. Burke, R. L., J. Tasse, C. Badgley, S. R. Jones, N. Fishkein, S. Phillips, and
managing risk. M. E. Soulé. 1991. Conservation of Stephen’s kangaroo rat (Dipodomys
stephensi): planning for persistence. Bulletin of the Southern California
Academy of Science 90:10-40.
 492 Wildlife–Habitat Relationships in Oregon and Washington

22. Burton, M. P. 1994. Alternative projections of decline of the African 45. Green, R. E., M. W. Pienkowski, and J. A. Love. 1996. Long-term viability
elephant. Biological Conservation 70:183-188. of the re-introduced population of the white-tailed eagle Haliaeetus
23. Bustamante, J. 1996. Population viability analysis of captive and albicilla in Scotland. Journal of Applied Ecology 33:357-368.
released bearded vulture populations. Conservation Biology 10:822- 46. Grubb, T. G. 1988. Pattern recognition—a simple model for evaluating
831. wildlife habitat. RM-GTR-487. USDA Forest Service, Rocky Mountain
24. Caughley, G. 1994. Directions in conservation biology. Journal of Forest and Range Experiment Station, Fort Collins, CO.
Animal Ecology 63:215-244. 47. Grumbine, R. E. 1990. Viable populations, reserve size, and federal land
25. Chen, J. 1991. Edge effects: microclimatic pattern and biological management: a critique. Conservation Biology 4:127-134.
responses in old growth Douglas-fir forests. Ph.D. dissertation, 48. Haig, S. M., J. R. Belthoff, and D. H. Allen. 1993. Population viability
University of Washington, Seattle, WA. analysis for a small population of red-cockaded woodpeckers and an
26. Cisneros-Mata, M. A., L. W. Botsford, and J. F. Quinn. 1997. Projecting evaluation of enhancement strategies. Conservation Biology 7:289-301.
viability of Totoaba macdonaldi, a population with unknown age- 49. Haight, R. G. 1995. Comparing extinction risk and economic cost in
dependent variability. Ecological Applications 7:968-980. wildlife conservation planning. Ecological Applications 5:767-775.
27. Diamond, J., K. D. Bishop, and S. Van Balen. 1987. Bird survival in an 50. Hallam, T. G., and R. R. Lassiter. 1994. Individual-based mathematical
isolated Javan woodland: island of mirror? Conservation Biology modeling approaches in ecotoxicology: a promising direction for
1:132-133. aquatic population and community ecological risk assessment. Pages
28. Diamond, J. M. 1984. “Normal” extinctions of isolated populations. 531-542 in Ronald J. Kendall, and Thomas E. Lacher, editors. Wildlife
Pages 191-246 in M. H. Nitecki, editor. Extinctions. University of toxicology and population modeling: integrated studies of
Chicago Press, Chicago, IL. agroecosystems. Lewis Publishers, Boca Raton, FL.
29. ———. 1989. The present, past and future of human-caused 51. Hansen, A. J., and D. L. Urban. 1992. Avian response to landscape
extinctions. Philosophical Transactions of the Royal Society of London patterns—the role of species’ life histories. Landscape Ecology 7:163-
B325:469-477. 180.
30. Emlen, J. M. 1989. Terrestrial population models for ecological risk 52. Harcourt, A. H. 1995. Population viability estimates: theory and
assessment: a state-of-the-art review. Environmental Toxicology and practice for a wild gorilla population. Conservation Biology 9:134-142.
Chemistry 8:831-842. 53. Hokit, D. G., B. M. Stith, and L. C. Branch. 1999. Effects of landscape
31. ———. 1995. Population viability of the Snake River chinook salmon structure in Florida scrub: a population perspective. Ecological
(Oncorhynchus tshawytscha). Canadian of Journal of Fisheries and Applications 9:124-134.
Aquatic Sciences 52:1442-1448. 54. Holling, C. S. 1984. Adaptive environmental assessment and
32. Fahrig, L., and J. Paloheimo. 1988. Effects of spatial arrangement of management. John Wiley & Sons, New York, NY.
habitat patches on local population size. Ecology 69:468-475. 55. Holtsinger, K. E., and P. Vitt. 1997. The future of conservation biology:
33. FEMAT. 1993. Forest ecosystem management: an ecological, economic, what’s a geneticist to do? Pages 202-216 in S. T. A. Pickett, R. S. Ostfeld,
and social assessment. Report of the Forest Ecosystem Management M. Shachak, and G. E. Likens, editors. The ecological basis of
Assessment Team. U.S. Government Printing Office, Washington, D.C. conservation: heterogeneity, ecosystem, and biodiversity. Chapman and
34. Fernandez-Duque, E., and C.Valeggia. 1994. Meta-analysis: a valuable Hall, New York, NY.
tool in conservation research. Conservation Biology 8:555-561. 56. Hunter, J. E., and F. L. Schmidt. 1990. Methods of meta-analysis. Sage
35. Fite, E., L. W. Turner, N. J. Cook, C. Stunkard, and R. M. Lee. 1988. Publications, Thousand Oaks, CA.
Guidance document for conducting terrestrial field studies. EPA 540/09, 57. IUCN. 1994. Red list categories. Gland, Switzerland.
88-109. U.S. Environmental Protection Agency, Office of Pesticide 58. ———. 1996a. Red list of threatened animals. Gland, Switzerland.
Programs, Washington, D.C. 59. ———. 1996b. Red list of threatened plants. Gland, Switzerland.
36. Flood, B. S., M. E. Sangster, R. D. Sparrowe, and T. S. Baskett. 1977. A 60. Jeffers, J. N. R. 1991. Rule induction methods in forestry research. AI
handbook for habitat evaluation procedure. USDI Fish and Wildlife Applications 5:37-44.
Service, Washington, D.C. 61. Johnson, K. N., J. Franklin F., J. W. Thomas, and J. Gordon. 1991.
37. Frankel, O. H., and M. E. Soulé. 1981. Conservation and evolution. Alternatives for management of late-successional forests of the Pacific
Cambridge University Press. Northwest: a report to the Agriculture Committee and The Merchant
38. Franklin, A. B., and T. M. Shenk. 1995. Meta-analysis as a tool for Marine and Fisheries Committee of the U.S. House of Representatives.
monitoring wildlife populations. Pages 484-487 in J. A. Bissonette, and P. Scientific Panel on Late-Successional Forest Ecosystems, USDA Forest
R. Krausman, editors. Integrating people and wildlife for a sustainable Service, Washington, D.C.
future. The Wildlife Society, Bethesda, MD. 62. Julin, K. R., compiler. 1997. Assessments of wildlife viability, old-growth
39. Fritts, S. H., and L. N. Carbyn. 1995. Population viability, nature timber volume estimates, forested wetlands, and slope stability. PNW
reserves, and the outlook for gray wolf conservation in North America. Gen. Tech. Rep. PNW-392. USDA Forest Service, Pacific Northwest
Restoration Ecology 3:26-38. Research Station, Portland, OR.
40. Gagne, J. A. 1994. The future application of ecological models in 63. Kareiva, P. 1987. Habitat fragmentation and the stability of predatory-
environmental risk assessment. Pages 497-499 in Ronald J. Kendall, and prey interactions. Nature 326:288-290.
Thomas E. Lacher, editors. Wildlife toxicology and population modeling: 64. ———, D. Skelly, and M. Ruckelshaus. 1997. Reevaluating the use of
integrated studies of agroecosystems. Lewis Publishers, Boca Raton, FL. models to predict the consequences of habitat loss and fragmentation.
41. Gardenfors, U. 1996. Application of Red List categories on a regional Pages 156-166 in S. T. A. Pickett, R. S. Ostfeld, M. Shachak, and G. E.
scale. In IUCN Red List of Threatened Animals. Gland, Switzerland. Likens, editors. The ecological basis of conservation: heterogeneity,
42. Gilpin, M. E., and M. E. Soulé. 1986. Minimum viable populations: ecosystems, and biodiversity. Chapman & Hall, New York, NY.
processes of species extinction. Pages 19-34 in M. E. Soulé, editor. 65. Karr, J. R. 1982a. Avian extinction on Barro Colorado Island, Panama:
Conservation biology: the science of scarcity and diversity. Sinauer a reassessment. American Naturalist 119:220-239.
Associates, Sunderland, MA. 66. ———. 1982b. Population variability and extinctions in the avifauna of
43. Goodman, D. 1987a. How do species persist? lessons for a tropical land-bridge island. Ecology 63:1975-1978.
conservation biology. Conservation Biology 1:59-62. 67. Kelly, P. A., K. D. Allred, H. P. Possingham, and D. F. Williams. 1996.
44. ———. 1987b. The demography of chance extinction. Pages 11-34 in Extinction risk assessment for the San Joaquin kit fox. Bulletin of the
M. E. Soulé, editor.Viable populations for conservation. Cambridge Ecological Society of America 77:228.
University Press, 68. Kendall, R. J., and J. Akerman. 1992. Terrestrial wildlife exposed to
agrochemicals an ecological risk assessment perspective. Environmental
Toxicology and Chemistry 11:1727-1749.
 CHAPTER 19: SPECIES AT RISK 493

69. Kenney, J. S., J. L. D. Smith, A. M. Starfield, and C. W. McDougal. 1995. 88. ———, and C. Servheen. 1992. Integrating biological and sociological
The long-term effects of tiger poaching on population viability. concerns in endangered species management: augmentation of grizzly
Conservation Biology 9:1127-1133. bear populations. Conservation Biology 6:426-434.
70. Kincaid, H. L. 1976. Effects of inbreeding on rainbow trout 89. ———, G. F. Wilhere, and Q. Dong. 1995. Population viability analysis
populations. Transactions of the American Fisheries Society 2:273-80. for red-cockaded woodpeckers in the Georgia Piedmont. Journal of
71. King, F. W. 1987. Thirteen milestones on the road to extinction. Pages Wildlife Management 59:533-542.
7-18 in R. Fitter, and M. Fitter, editors. The road to extinction. IUCN, 90. Marcot, B. G. 1986. Concepts of risk analysis as applied to viable
Gland, Switzerland. population assessment and planning. Pages 89-102 in B. A. Wilcox, P. F.
72. Kinnaird, M. F., and T. G. O’Brien. 1991. Viable populations for an Brussard, and B. G. Marcot, editors. The management of viable
endangered forest primate, the Tana River Mangabey (Cercocebus populations: theory, applications, and case studies. Center for
galeritus galerius). Conservation Biology 5:203-213. Conservation Biology, Stanford, CA.
73. Klein, B. C. 1989. Effects of forest fragmentation on dung and carrion 91. ———., R. S. Holthausen, and H. Salwasser. 1986.Viable population
beetle communities in central Amazonia. Ecology 70:1715-1725. planning. Pages 49-62 in B. A. Wilcox, P. F. Brussard, and B. G. Marcot,
74. Lande, R. 1988. Genetics and demography in biological conservation. editors. The management of viable populations: theory, applications, and
Science 241:16-241. case studies. Center for Conservation Biology, Stanford, CA.
75. Lehmkuhl, J. F., and D. R. Patton. 1984. User’s manual for the RUN 92. ———. 1987. Use of decision tree analysis for assessing wildlife-
WILD III data storage and retrieval system. USDA Forest Service, silviculture relationships. USFS Unpublished Report
Southwestern Region, Wildlife Unit, Albuquerque, NM. 93. ———. 1992. Putting data, experience and professional judgment to
76. ———, M. G. Raphael, R. S. Holthausen, J. R. Hickenbottom, R. H. work in making land management decisions. Pages 140-161 in J. B.
Naney, and J. S. Shelly. 1997. Chapter 4. Effects of planning alternatives Nyberg and W. B. Kessler, editors. Integrating timber and wildlife in
on terrestrial species in the interior Columbia River basin. Pages 539- forest landscapes: a matter of scale. B.C. Ministry of Forests, Victoria,
730 in T. M. Quigley, K. M. Lee, and S. J. Arbelbide, technical editors. B.C., Canada.
Evaluation of EIS alternatives by the science integration team. PNW- 94. ———, and R. Holthausen. 1987. Analyzing population viability of the
GTR-406. USDA, Forest Service, Pacific Northwest Research Station, spotted owl in the Pacific Northwest. Transactions of the North
Portland, OR. American Wildland and Natural Resources Conference 52:333-347.
77. ———, and L. F. Ruggiero. 1991. Forest fragmentation in the Pacific 95. ———, L. K. Croft, J. F. Lehmkuhl, R. H. Naney, C. G. Niwa, W. R.
Northwest and its potential effects on wildlife. Pages 35-46 in L. F. Owen, and R. E. Sandquist. 1998. Macroecology, paleoecology, and
Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff, editors. Wildlife and ecological integrity of terrestrial species and communities of the
vegetation of unmanaged Douglas-fir forests. PNW-GTR-285. USDA interior Columbia River Basin and portions of the Klamath and Great
Forest Service, Pacific Northwest Research Station, Portland, OR. Basins. PNW Gen. Tech. Rep. PNW-410. USDA Forest Service, Pacific
78. Lindenmayer, D. B., M. A. Burgman, H. R. Akçakaya, R. C. Lacy, and H. P. Northwest Research Station, Portland, OR.
Possingham. 1995. A review of the generic computer programs ALEX, 96. Margules, C., A. J. Higgs, and R. W. Rafe. 1982. Modern biogeographic
RAMAS-space and VORTEX for modeling the viability of wildlife theory: are there any lessons for nature reserve design? Biological
metapopulations. Ecological Modeling 82:161-174. Conservation 24:115-128.
79. ———, T. W. Clark, R. C. Lacy, and V. C. Thomas. 1993. Population 97. Marmontel, M., S. R. Humphrey, and T. J. O’Shea. 1997. Population
viability analysis as a tool in wildlife conservation policy: with reference viability analysis of the Florida manatee (Trichechus manatus latirostris),
to Australia. Environmental Management 17:745-758. 1976-1991. Conservation Biology 11:467-481.
80. ———, and H. P. Possingham. 1995. Modeling the impacts of wildfire 98. Master, L. L. 1991. Assessing threats and setting priorities in
on the viability of metapopulations of the endangered Australian conservation. Conservation Biology 5:559-563.
species of arboreal marsupial, Leadbeater’s possum. Forest Ecology and 99. Mather, A. S. 1990. Global forest resources. Belhaven Press, London,
Management 74:197-222. England.
81. ———, and ———. 1996. Ranking conservation and timber 100. Matsuda, H., T.Yahara, and Y. Uozumi. 1997. Is tuna critically
management options for Leadbeater’s possum in southeastern endangered? Extinction risk of a large and overexploited population.
Australia using population viability analysis. Conservation Biology Ecological Research 12:345-356.
10:235-251. 101. May, R. M. 1973. Stability in randomly fluctuating versus deterministic
82. Lovejoy, T. E., R. O. Bierregaard, Jr., A. B. Rylands, J. R. Malcom, C. E. environments. American Naturalist 107:621-650.
Quintels, L. H. Harper, K. S. Brown, Jr. , A. H. Powell, G. V. N. Powell, H. O. 102. ———, J. H. Lawton, and N. E. Stork. 1995. Assessing extinction
R. Schubart, and M. Hays. 1986. Edge and other effects of isolation on rates. Pages 1-24 in J. H. Lawton and R. M. May, editors. Extinction rates.
Amazon forest fragments. Pages 257-285 in M. E. Soulé, editor. Oxford University Press, New York, NY.
Conservation biology: the science of scarcity and diversity. Sinauer 103. McCarthy, M. A. 1995. Population viability analysis of the helmeted
Associates, Sunderland, MA. honeyeater: risk assessment of captive management and reintroduction.
83. Mace, G. M., and N. J. Collar. 1995. Extinction risk assessment for birds Pages 21-25 in M. Serena, editor. Reintroduction biology of Australian
through quantitative criteria. IBIS 137:240-246. and New Zealand fauna. Surrey Beatty, Chipping Norton, England.
84. Madsen, T, B. Stille, and R. Shine. 1996. Inbreeding depression in an 104. ———, and M. A. Burgman. 1995. Coping with uncertainty in forest
isolated population of adders Vipera berus. Biological Conservation wildlife planning. Forest Ecology and Management 74:23-36.
75:113-8. 105. ———, ———, and S. Ferson. 1995. Sensitivity analysis for models
85. Maquire, L. A., T. W. Clark, R. Crete, J. Cada, C. Groves, M. L. Shaffer, of population viability. Biological Conservation 73:93-100.
and U. S. Seal. 1988. Black-footed ferret recovery in Montana: a decision 106. McCoy, E. D. 1982. The application of island-biogeographic theory to
analysis. Wildlife Society Bulletin 16:111-120. forest tracts: problems in the determination of turnover rates.
86. ———, and R. C. Lacy. 1990. Allocating scarce resources for Biological Conservation 22:217-227.
conservation of endangered subspecies: partitioning zoo space for 107. McElroy, D. M., J. A. Shoemaker, and M. E. Douglas. 1997.
tigers. Conservation Biology 4:157-166. Discriminating Gila robusta and Gila cypha: risk assessment and the
87. ———, U. S. Seal, and P. F. Brussard. 1987. Managing critically Endangered Species Act. Ecological Applications 7:958-967.
endangered species: the Sumatran rhino as a case study. Pages 141-158 108. McNay, R. S., R. E. Page, and A. Campbell. 1987. Application of expert-
in M. E. Soulé, editor.Viable populations for conservation. Cambridge based decision models to promote integrated management of forests.
University Press, Cambridge, U.K. Transactions of the North American Wildland and Natural Resources
Conference 52:82-91.
 494 Wildlife–Habitat Relationships in Oregon and Washington

109. Menges, E. S. 1990. Population viability analysis for an endangered 135. Ranjit Daniels, R. J., M. Hegde, N.V. Joshi, and M. Gadgil. 1991.
plant. Conservation Biology 4:52-62. Assigning conservation value: a case study in India. Conservation
110. ———. 1991. The application of minimum viable population theory Biology 5:464-475.
to plants. Pages 45-61 in D. A. Falk and K. E. Holtsinger, editors. 136. Ranney, J. W., J. M. C. Bruner, and J. B. Levenson. 1981. The importance
Genetics and conservation of rare plants. Oxford University Press, of edge in the structure and dynamics of forest islands. Page 67-95 in R.
New York, NY. L. Burgess and D. M. Sharpe, editors. Forest island dynamics in man-
111. Meyer, J. S., C. G. Ingersoll, L. L. McDonald, and M. S. Boyce. 1986. dominated landscapes. Springer, New York, NY.
Estimating uncertainty in population growth rates: jackknife vs. 137. Raphael, M. G. 1988. Long-term trends in abundance of amphibians,
bootstrap techniques. Ecology 67:1156-1166. reptiles, and mammals in Douglas-fir forests of northwestern California.
112. Miller, R. I., and L. D. Harris. 1977. Isolation and extirpation in wildlife Pages 23-31 in Management of amphibians, reptiles, and small mammals
reserves. Biological Conservation 12:311-315. in North America. RM-GTR-166. USDA Forest Service, Flagstaff, AZ.
113. Mills, L. S., S. G. Hayes, C. Baldwin, M. J. Wisdom, J. Citta, D. J. Mattson, 138. ———, and Marcot, B. G. 1986. Validation of a wildlife-habitat-
and K. Murphy. 1996. Factors leading to different viability predictions relationships model: vertebrates in a Douglas-fir sere. Pages 129-138 in
for a grizzly bear data set. Conservation Biology 10:863-873. J. Verner, M. L. Morrison, and C. J. Ralph. editors. Wildlife 2000: modeling
114. Millsap, B. A., J. A. Gore, D. E. Runde, and S. I. Cerulean. 1990. Setting habitat relationships of terrestrial vertebrates. University of Wisconsin
priorities for the conservation of fish and wildlife species in Florida. Press, Madison, WI.
Wildlife Monographs 111:1-57. 139. ———, K.V. Rosenberg, and B. G. Marcot. 1988. Large-scale changes
115. Morrison, M. L., B. G. Marcot, and R. W. Mannan. 1998. Wildlife- in bird populations of Douglas-fir forests, northwestern California.
habitat relationships: concepts and applications. Second edition. Pages 63-83 in Bird conservation. University of Wisconsin Press,
University of Wisconsin Press, Madison, WI. Madison, WI.
116. Murphy, D.D., K.E. Freas, and S.B. Weiss. 1990. An environment- 140. Ratner, S., R. Lande, and B. B. Roper. 1997. Population viability analysis
metapopulation approach to viability analysis for a threatened of spring chinook salmon in the South Umpqua River, Oregon.
invertebrate. Conservation Biology 4:41-51. Conservation Biology 11:879-889.
117. Nantel, P., D. Gagnon, and A. Nault. 1996. Population viability analysis 141. Redford, K. H., A. Taber, and J. A. Simonetti. 1990. There is more to
of American ginseng and wild leek harvested in stochastic biodiversity than the tropical rainforests. Conservation Biology 4:328-
environments. Conservation Biology 10:608-621. 330.
118. Nash, S. 1991. What price nature? Future ecological risk assessments 142. Reed, J. M., C. S. Elphick, and L. W. Oring. 1998. Life-history and
may chart the values, and the odds. BioScience 41:677-680. viability analysis of the endangered Hawaiian stilt. Biological
119. Newmark, W. D. 1987. A land-bridge island perspective on Conservation 84:35-45.
mammalian extinctions in western North American parks. Nature 143. Rodier, D. J., and M. G. Zeeman. 1994. Ecological risk assessment.
325:430-432. Pages 581-604 in Lorris G. Cockerham and Barbara S. Shane, editors.
120. Niemi, G. J. 1982. Determining priorities in nongame management. Basic environmental toxicology. CRC Press, Boca Raton, FL.
Loon 54:28-54. 144. Rossi, R. E., P. W. Borth, and J. J. Tollefson. 1993. Stochastic simulation
121. Nolet, B. A., and J. M. Baveco. 1996. Development and viability of a for characterizing ecological spatial patterns and appraising risk.
translocated beaver Castor fiber population in the Netherlands. Ecological Applications 3:719-735.
Biological Conservation 75:125-137. 145. Russell, G. J., T. M. Brooks, M. M. McKinney, and C. G. Anderson. 1998.
122. Patton, D. R. 1978. RUN WILD: a storage and retrieval system for Present and future taxonomic selectivity in bird and mammal
wildlife habitat information. RM-GTR-51. USDA Forest Service, Rocky extinctions. Conservation Biology 12:1365-1376.
Mountain Forest and Range Experiment Station, Fort Collins, CO. 146. Saunders, D. A., R. J. Hobbs, and C. R. Margules. 1991. Biological
123. Pazzani, M., and D. Kibler. 1992. The role of prior knowledge in consequence of ecosystem fragmentation: a review. Conservation
inductive learning. Machine Learning 9:54-97. Biology 5:18-32.
124. Peterle, T. J. 1991. Wildlife toxicology. Van Nostrand Reinhold, New 147. Schamberger, M., A. H. Farmer, and J. W. Terrell. 1982. Habitat
York, NY. suitability index models: introduction. FWS/OBS-82/10. USDI Fish and
125. Pimm, S. L., H. L. Jones, and J. Diamond. 1988. On the risk of Wildlife Service, Washington, D.C.
extinction. American Naturalist 132:757-785. 148. Schonewald-Cox, C. M., S. M. Chambers, B. MacBryde, and L. Thomas,
126. Pinchera, F., L. Boitani, and F. Corsi. 1997. Application to the editors. 1983. Genetics and conservation: a reference for managing wild
terrestrial vertebrates of Italy of a system proposed by IUCN for a animal and plant populations. Benjamin/Cummings, Menlo Park, CA.
new classification of national Red List categories. Biodiversity and 149. Seal, U. S. 1986. Goals of captive propagation programmes for the
Conservation. 6:959-978. conservation of endangered species. International Zoo Yearbook 24/
127. Power, M., D. G. Dixon, and G. Power. 1994. Modeling population 25:174-179.
exposure-response functions for use in environmental risk assessment. 150. Shaffer, M. L., 1981. Minimum population sizes for species
Journal of Aquatic Ecosystem Health 3:45-58. conservation. BioScience 31:131-134.
128. Prescott-Allen, R., and C. Prescott-Allen. 1978. Sourcebook for a 151. ———. 1983. Determining minimum viable population sizes for the
world conservation strategy: threatened vertebrates. IUCN, Gland, grizzly bear. International Conference on Bear Research and
Switzerland. Management 5:133-139.
129. Quinlan, J. R. 1986a. Induction of decision trees. Machine Learning 152. ———. 1987. Minimum viable populations: coping with uncertainty.
1:81-106. Pages 69-86 in M. E. Soulé, editor.Viable populations for conservation.
130. ———. 1986b. Simplifying decision trees. International Journal of Cambridge University Press, Cambridge, U.K.
Man-Machine Studies 27:221-234. 153. ———, and F. B. Samson. 1985. Population size and extinction: a
131. Quinn, J. F., and A. Hastings. 1987. Extinction in subdivided habitats. note on determining critical population sizes. American Naturalist
Conservation Biology 1:198-209. 125:144-152.
132. Raiffa, H. 1968. Decision analysis: introductory lectures on choices 154. Shaw, C. G., III. 1999. Use of risk assessment panels during revision
under certainty. Addison-Wesley, Reading, MA. of the Tongass Land and Resource Management Plan. PNW Gen. Tech.
133. Ralls, K, J. D. Ballou, and A. Templton. 1988. Estimates of lethal Rep. PNW-460. USDA Forest Service, Pacific Northwest Research
equivalents and the cost of inbreeding in mammals. Conservation Station, Portland, OR.
Biology 2:185-93. 155. Simberloff, D. 1988. The contribution of population and community
134. ———, K. Brugger, and J. Ballou. 1979. Inbreeding and juvenile biology to conservation science. Annual Review of Ecology and
mortality in small populations of ungulates. Science 206:1101-3. Systematics 19:473-511.
 CHAPTER 19: SPECIES AT RISK 495

156. ———, and L. G. Abele. 1982. Refuge design and island 179.Veerkamp, W., and C. Wolff. 1996. Fate and exposure models in
biogeographic theory: effects of fragmentation. American Naturalist relation to risk assessment: developments and validation criteria.
120:41-50. Environmental Science and Pollution Research International 3:91-95.
157 Simon, N., and P. Geroudet. 1970. Last survivors. World Publishing 180. Wilcove, D. S. 1985. Nest predation in forest tracts and the decline
Co., New York, NY. of migratory songbirds. Ecology 66:1211-1214.
158. Skalski, J. R., and S. G. Smith. 1994. Risk assessment in avian 181. ———, C. H. McLellan, and A. P. Dobson. 1986. Habitat
toxicology using experimental and epidemiology approaches. Pages fragmentation in the temperate zone. Pages 237-256 in M. E. Soulé,
467-488 in Ronald J. Kendall and Thomas E. Lacher, editors. Wildlife editor. Conservation biology: the science of scarcity and diversity.
toxicology and population modeling: integrated studies of Sinauer Associates, Sunderland, MA.
agroecosystems. Lewis Publishers, Boca Raton, FL. 182. Wilcox, B. A. 1980. Insular ecology and conservation. Pages 95-117 in
159. Song, Y. L. 1996. Population viability analysis for two isolated M. E. Soulé and B. A. Wilcox, editors. Conservation biology: an
populations of Haianan Eld’s deer. Conservation Biology 10:1467-1472. evolutionary-ecological perspective. Sinauer Associates, Sunderland,
160. Soulé, M. E. 1986. The effects of fragmentation. Pages 233-236 in M. E. MA.
Soulé, editor. Conservation biology: the science of scarcity and diversity. 183. Wilkinson, L. 1997. Classification and regression trees. Pages 13-30 in
Sinauer Associates, Sunderland, MA. SYSTAT 7.0: new statistics. SPSS Inc., Chicago, IL.
161. ———, editor. 1987. Viable populations for conservation. Cambridge 184. Wisdom, M. J., L. R. Bright, C. G. Carey, W. W. Hines, R. J. Pedersen, D.
University Press, Cambridge, U.K. A. Smithey, J. W. Thomas, and G. W. Witmer. 1986. A model to evaluate
162. ———. 1989. Risk analysis for the concho water snake. Endangered elk habitat in western Oregon. USDA Forest Service PNW Region, and
Species Update 6:19, 22-25. USDI Bureau of Land Management and Oregon Department of Fish
163. ———, D. T. Bolger, A. C. Alberts, R. Sauvajot, J. Wright, M. Sorice, and and Wildlife, Portland, OR.
S. Hill. 1988. Reconstructed dynamics of rapid extinctions of 185. World Resources Institute (WRI). 1991. World Resources Report
chaparral-requiring birds in urban habitat islands. Conservation Biology 1991-1992. A guide to the global environment. Oxford University Press,
2:75-92. New York, NY.
164. ———, and Wilcox, B. A., editors. 1980. Conservation biology: an 186. ———. 1992. World Resources Report 1992-1993: Toward
evolutionary-ecological approach. Sinauer Associates, Sunderland, MA. sustainable development. Oxford University Press, New York, NY.
165. Starfield, A. M., and A. M. Herr. 1991. A response to Maguire. 187. Xu, H., and H. Lu. 1996. A preliminary analysis of population viability
(Letters). Conservation Biology 5:435. for Chinese water deer (Hydropotes inermis) lived in Yancheng.Acta
166. Stockwell, D. R. B., S. M. Davey, J. R. Davis, and I. R. Noble. 1990. Using Theriologica Sinica. 16:81-88.
induction of decision trees to predict greater glider density. AI
Applications in Natural Resource Management 4:33-43.
167. Taylor, B. L. 1995. The reliability of using population viability analysis
for risk classification of species. Conservation Biology 9:551-558.
168. ———, P. R. Wade, R. A. Stehn, and J. F. Cochrane. 1996. A Bayesian
approach to classification criteria for spectacled eiders. Ecological
Applications 6:1077-1089.
169. Temple, S. A., and J. R. Cary. 1988. Modeling dynamics of habitat:
interior bird populations in fragmented landscapes. Conservation
Biology 2:340-347.
170. Terborgh, J. W., and B. Winter. 1980. Some causes of extinction. Pages
119-133 in M. E. Soulé, editor. Conservation biology: the science of
scarcity and diversity. Sinauer Associates, Sunderland, MA.
171. Thomas, J. W., H. Black, Jr., R. J. Scherzinger, and R. J. Pedersen. 1979.
Deer and elk. Pages 104-127 in J. W. Thomas, editor. Wildlife habitats in
managed forests—the Blue Mountains of Oregon and Washington. U.S.
Gov. Printing Office, Washington, D.C.
172. ———, E. D. Forsman, J. B. Lint, E. C. Meslow, B. R. Noon, and J.
Verner. 1990. A conservation strategy for the spotted owl. U.S.
Government Printing Office, Portland, OR.
173. ———, D. A. Leckenby, M. Henjum, R. J. Pedersen, and L. D. Bryant.
1988. Habitat-effectiveness index for elk on Blue Mountain winter
ranges. PNW-GTR-218. USDA Forest Service, Pacific Northwest
Research Station, Portland, OR.
174. Tracy, C. R., and T. L. George. 1992. On the determinants of
extinction. The American Naturalist 139:102-121.
175. Urban, D. J. 1990. The use of terrestrial field data in the practical
application of ecological risk assessment principles. Pages 319-334 in
L. A. Somerville and C. H. Walker, editors. Pesticide effects on terrestrial
wildlife. Taylor & Francis, London, England.
176. ———, and Cook, N. J. 1986. Ecological risk assessment - hazard
evaluation division standard evaluation procedure. EPA 540/9-86-167.
U.S. Environmental Protection Agency, Office of Pesticide Programs,
Washington, D.C.
177. USDA. 1988. Final supplement to the Environmental Impact
Statement for and Amendment to the Pacific Northwest Regional
Guide. USDA Forest Service, Pacific Northwest Region, Portland, OR.
178. U.S. Fish and Wildlife Service. 1980. Habitat evaluation procedures
(HEP). Ecological Services Manual No. 102. U.S. Government Printing
Office, Washington, D.C.
 496 Wildlife–Habitat Relationships in Oregon and Washington

Appendix 1. Group I. Species Known To Be Imperiled

This group includes 20 Federally listed threatened or
Criteria and procedure for selecting species to endangered species, as well as species listed by
demonstrate viability risk analyses for this chapter. Washington or Oregon State as candidates, sensitive,
To demonstrate the several examples of species viability critical, or in decline. Where multiple species matched
risk analysis presented in this chapter, we selected a these criteria, where possible we chose the species with
representative set of species from the Oregon-Washington the distribution that included both states.
species-environment database. At the time of the analysis, Dunn’s Salamander
the databases were not completed, so we selected 20 Van Dyke’s Salamander
species in each of 3 risk groups. The 3 risk groups were: Northern Leopard Frog
Group I, species known to be imperiled; Group II, species Plateau Striped Whiptail
very likely secure; and Group III, species with intermediate Striped Whipsnake
security. Bald Eagle
To keep the comparisons at some level of parity across Peregrine Falcon
groups, in each group we included the same number of Snowy Plover
species from each selected taxonomic class and order. The Northern Spotted Owl
number of species by class and order represented in each Vaux’s Swift
of the 3 groups was as follows: White-Headed Woodpecker
Gray Flycatcher
Amphibians—3 total Western Bluebird
Salamanders—2; Frogs—1 Bobolink
Reptiles—2 total Grasshopper Sparrow
Lizards—1; Snakes—1 Gray-Tailed Vole
Gray Wolf
Mammals—5 total Wolverine
Voles—1; Canids—1; Weasels or raccoons—2; Cats—1 Fisher
Lynx
Birds—10 total
Non-owl raptors—2; Shorebirds—1; Owls—1; Group II. Species Very Likely Secure
Swifts—1; Woodpeckers—1; Flycatchers—1; Passerines— This group includes 20 species known to be widespread,
3 common, or abundant, and thus highly likely to be secure.
Roughskin Newt
To reduce the variation in these small samples, we Ensatina
excluded from these lists: Pacific Treefrog
(a) marine or pelagic species, Western Fence Lizard
(b) obligate shorebirds, and species wholly dependent Gopher Snake
on inland freshwater aquatic environments, such as Turkey Vulture
most waterfowl, although some amphibians were American Kestrel
included, Killdeer
(c) introduced or exotic species, such as wild turkey, Barn Owl
(d) species at the edge of their distributional ranges White-Throated Swift
(e.g., Tennessee warbler), Hairy Woodpecker
(e) species that occur in the WA-OR area only Cordilleran Flycatcher
sporadically or irruptively (e.g., snowy owl), Cedar Waxwing
(f) species that occur in only a small portion of the White-Crowned Sparrow
geographic area, particularly in only one state (e.g., Purple Finch
Olympic marmot); an exception is lynx, which Meadow Vole
historically ranged into Oregon and for which there Coyote
are intermittent (and largely unconfirmed) sightings Raccoon
in Oregon. Long-Tailed Weasel
We used this nonrandom selection process because our Bobcat
aim was to demonstrate selected types of risk analysis
models across as wide array of species types as possible,
and not to develop a thorough risk analysis for all species.
The species chosen for the analyses were as follows.
 CHAPTER 19: SPECIES AT RISK 497

Group III. Species With Intermediate Security

This group includes 20 species with viability security
judged to be intermediate between that of Groups I and
II.
Tiger Salamander
Larch Mountain Salamander
Columbian Spotted Frog
Western Whiptail
Sharptail Snake
Cooper’s Hawk
Merlin
Black-Bellied Plover
Northern Saw-Whet Owl
Black Swift
Pileated Woodpecker
Willow Flycatcher
Bohemian Waxwing
Sage Sparrow
Gray-Crowned Rosy Finch
White-Footed Vole
Kit Fox
Ringtail
American Marten
Mountain Lion
Appendix 2.
Life history and habitat versatility fields used in examples
of risk analysis procedures. Values for some fields were simplified from the original values in the
matrixes. Risk score for values of some fields was estimated from the literature for the examples.

SHP Field Name FieldTypea FieldValuesb Risk Scorec

Common name 3 common name na

Taxonomic class 3 A=amphibian; B=bird; M=mammal; R=reptile na
Taxonomic order 3 name of taxonomic order na
Taxonomic family 3 name of taxonomic family na
Breeding status 3 1=breeds in OR; 2=breeds in WA; 3=breeds in both na
states; 4=non-breeder
Occurrence status by state 3 0=does not occur in the state; 1=native; 2=non-native; na
3=reintroduced
Risk categoryd 1 1=Group I—species known to be imperiled; 2=Group II na
—species very likely secure; 3=Group III—species
with intermediate security
Type of seasonal activity 2 1=hibernation; 2=estivation; 3=both (hibernation & na
estivation) 4=none
Migration/seasonal movements 1 1=latitudinal 7
2=altitudinal 4
4=year round resident 1
6=both 1 and 2 10
Migration/seasonal move- 1 1=< 10 km & <100 km 4
ment distance class 2=< 1,000 km 7
3=< 10,000 & >10,000 km 10
4=none 1
Forms aggregations 1 y=yes 10
n=no 0
Juvenile dispersal distance class 1 1=<100 m & < 1 km 10
2=<10 km 5
3=< 100 km & >100 km 0
 498 Wildlife–Habitat Relationships in Oregon and Washington

SHP Field Name FieldTypea FieldValuesb Risk Scorec

Average # of offspring per 1 1=1-2 & 3-6 10

litter, or eggs per clutch 2=7-10 & 11-15 5
3=16-20 & 21-50 & >50 0
# Litters or clutches / year 1 1<1 10
2<2 7
3<3 4
4>3 1
Average age at first breeding 1 1=<6 mo. 0
(females) 2=1 yr. 1
3=2 yr. 4
4=3 yr. 7
5=4+ yr. 10
Mating system 2 1=polygamy; 2=lifetime monogamy; 3=lek ; na
4=promiscuous
Home range size class 1 1=<1 ha & 1-10 ha 1
2=11-50 ha & 51-100 ha 4
3=101-500 ha & 501-1000 ha 7
4=1001-10,000 ha & > 10,000 ha 10
Fidelity to summer range 2 1=high; 2=medium; 3=low na
Fidelity to winter range 2 1=high; 2=medium; 3=low na
Geographic range 1 1=locally endemic 10
2=regionally endemic 7
3=moderately widespread, or widespread 1
4=WA peripheral or OR peripheral 10
Population distribution 1 1=contiguous 0
2=gaps 1
3=patchy 4
4=isolated 7
5=scarce 10
Landscape use 2 1=patch; 2=mosaic; 3=generalist; 4=contrast na
Elevational range 2 1=0-1000 feet; 2=<3000 feet; 3=<5000 feet; 4=>5000 feet na
Diet 1 1=primary consumer (herbivore) 2
2=secondary or tertiary consumer (carnivore) 8
3=primary invertebrates 4
4=both 1 & 3 (largely, insectivorous and eats seeds and 0
fruits)
Foraging location 2 1 terrestrial (underground, on ground, in down wood); na
2=lower vegetation (shrub or tree bole/bark); 3=upper
vegetation (tree canopy); 4=other (unknown, in or
underwater, or aerial)
Mass (body size) 1 1=<500 gm 0
2=<1 kg 2
3=<10 kg 4
4=<100 kg 6
5=<1000 kg 8
6=>1000 kg 10
Habitat versatilityd 1 Continuous variable from 1-100 representing the
percentage of the 32 habitats used by the species
(“Y” code in occurrence field of habitat matrix)
0-25% 10
26-50% 7
51-75% 4
76-100% 0
Structural versatilityd 1 Continuous variable from 1-100 representing the
percentage of the 54 structural types used by the species
( “Y” code in occurrence field of habitat matrix)
0-25% 10
26-50% 7
51-75% 4
76-100% 0
 CHAPTER 19: SPECIES AT RISK 499

Notes to Appendix 2 Washington]

a
1=primary field; 2=secondary field; 3=information field. The bs3 [the species is documented or suspected as breeding in both
example classification tree analysis used primary fields in the states]
calculations, and the secondary and information fields to aid post bs4 [the species is documented or suspected of being a
hoc interpretations. The example rule-induction analysis used all non-breeder in both states]
3 types of fields in the analysis. The differences in which types of OccStatusOR [What is the occurrence status in
fields were used pertained to meeting the statistical assumptions Oregon?]
of the two analyses. oso1 [native]
b
Values found in the matrix for some fields were combined to oso2 [non-native (introduced accidentally or purposefully, or
simplify the example analyses. self-invader)]
oso3 [reintroduced (native species extirpated then reestablished
c
Created for this analysis and not found in the matrixes. Relative
w/ introductions)]
risk score (1-10) for each field value. Values were estimated for
oso0 [does not occur in the state]
primary fields only for use in the rank-model example.
OccStatusWA [What is the occurrence status in
d
Field created for this analysis and not found in the matrixes.
Washington?]
Appendix 3. osw1 [native]
osw2 [non-native (introduced accidentally or purposefully, or
Running the Species At Risk Advisor (SARA): self-invader)]
factors and their values used in the example rule- osw3 [reintroduced (native species extirpated then reestablished
induction knowledge base model that provides w/ introductions)]
most likely viability risk levels of wildlife species. owo0 [does not occur in the state]
OccStatusBo [What is the occurrence status in both
Welcome to SARA: Species at Risk Advisor Oregon and Washington?]
Version 0.30, 4 January 1999, author: Bruce G. Marcot osb1 [native]
This is an expert advisory model that demonstrates osb2 [non-native (introduced accidentally or purposefully, or
how life history characteristics can be used to infer self-invader)]
potential risk levels of terrestrial vertebrate wildlife TypSeaInact [What is the type of seasonal inactivity?]
species. This model is based on a selected sample of 60 tsi1 [hibernation (dormancy associated with cold period of the
species—20 species in each of 3 known risk categories year)]
(imperiled/listed, secure, and intermediate). The model tsi2 [estivation (dormancy associated with warm/dry period of
is based on an optimized rule set using the Quinlan Q3 the year)]
rule-induction algorithm, that produces the most efficient tsi3 [both hibernation & estivation]
use of the example information for inferring risk levels. tsi4 [none]
This model is ONLY A DEMONSTRATION of the kind of Mig_SeasMov [What is the type of migrational or
risk level analysis possible. It is NOT intended to provide seasonal movement?]
definitive predictions of risk levels of all species. NOTE: migs1 [latitudinal (change in latitude, north/south)]
You may respond to each question by choosing the most migs2 [altitudinal (change in elevation)]
appropriate answer, or typing “?” (without the quotes) to migs4 [year-round resident (stays in same vicinity throughout the
denote that you don’t know. Let’s begin! year)]
TaxClas [What is the taxonomic class?] migs6 [both latitudinal and altitudinal]
amphibian MigDist [What is the migration or seasonal movement
reptile distance class?]
bird migd1 [< 100 km]
mammal migd2 [< 1000 km]
TaxOrder [What is the taxonomic order?] migd3 [> 1000 km]
Caudata migd4 [none (non-migratory)]
Anura Aggreg [Does the species form aggregations or
Squamata groups?]
Falconiform [Falconiformes] yes [yes (forms concentrations of individuals during some
Charadriifo [Charadriiformes] periods or activities)]
Strigiforme [Strigiformes] no
Apodiformes JuvDispDist [What is the juvenile dispersal distance
Piciformes class?]
Passeriform [Passeriformes] juv1 [< 1 km]
Rodentia juv2 [< 10 km]
Carnivora juv3 [> 10 km]
BreedStatus [What is the breeding status?] NOffsprg [What is the average number of offspring per
bs1 [the species is documented or suspected as breeding in litter, or eggs per clutch?]
Oregon] noff1 [1-6]
bs2 [the species is documented or suspected as breeding in noff2 [7-15]
 500 Wildlife–Habitat Relationships in Oregon and Washington

noff3 [>15] lnd2 [mosaic (sp. uses aggregates of patches of habitat, but 1
NLitt_Clut [What is the number of litters or clutches structural stage)]
per year?] lnd3 [generalist (sp. uses all or many patch types, & >1 structural
nlit1 [< or = 1] stage)]
nlit2 [< or = 2] lnd4 [contrast (sp. requires contrast betw. 2 struc. stages in close
nlit3 [< or = 3] proximity)]
nlit4 [> 3] ElevRng [What is the upper elevational range of typical
Age1st [What is the average age at first breeding or regular occurrence?]
(females)?] ele1 [up to 1000 ft]
age1 [< 6 mo.] ele2 [up to 3000 ft]
age2 [1 yr] ele3 [up to 5000 ft]
age3 [2 yrs] ele4 [> 5000 ft]
age4 [3 yrs] Diet [What is the diet category?]
age5 [4+ yrs] diet1 [primary consumer (herbivore)]
MatingSys [What is the mating system?] diet2 [secondary or tertiary consumer (carnivore)]
mat1 [polygamy (seasonal monogamy, polyandry, or polygyny)] diet3 [primarily invertebrates]
mat2 [lifetime monogamy] diet4 [omnivore (both herbivore and carnivore, mostly insects &
mat3 [lek] seeds/fruits)]
mat4 [promiscuity] Forag1 [Does the organism forage on terrestrial
HRSizeClass [What is the home range size class?] substrates (underground, on ground, or on/in down
hr1 [< or = 10 ha] wood)?]
hr2 [< or = 100 ha] yes
hr3 [< or = 1000 ha] no
hr4 [> 1000 ha] Forag2 [Does the organism forage in lower vegetation
SiteFidSumm [What is the site fidelity to summer (shrub or understory, or tree bole/bark)?]
range (i.e., home range overlap among years)?] yes
sw1 [high (>50% of popn returns to same home range or no
territory the next year)] Forag3 [Does the organism forage in upper vegetation
sw2 [medium (>10% of popn returns)] (tree canopy)?]
sw3 [low (<10% of popn returns)] yes
SiteFidWint [What is the site fidelity to winter range no
(i.e., home range overlap among years)?] Forag4 [Does the organism forage underwater or
sw1 [high (>50% of popn returns to same home range or aerially; or is foraging substrate unknown?]
territory the next year)] yes
sw2 [medium (>10% of popn returns)] no
sw3 [low (<10% of popn returns)] BodyMass [What is the body mass class (weight of
GeogRng [What is the geographic range?] larger sex)?]
geog1 [locally endemic (occurs only in OR &/or WA, or a small bm1 [< 500 gm]
area thereof)] bm2 [< 1 kg]
geog2 [regionally endemic (occurs only in Pacific Northwest or bm3 [< 10 kg]
Great Basin region)] bm4 [< 100 kg]
geog3 [moderately widespread, or widespread (occurs thru bm5 [< 1000 kg]
western states/No. Amer.)] bm6 [> 1000 kg]
geog4 [WA peripheral or OR peripheral (popn in WA or OR is
HabVers<?>
<?>
on edge of its range)]
<?>
StrucVers<?>
PopnDistrb [What is the population distribution?]
pop1 [contiguous (habitat is broadly & contiguously distributed RESULT [The risk level of the species is:]
over sp. range)] GrpI [Group I—Species known to be imperiled (there is at least
pop2 [gaps (habitat broadly distributed but w/ gaps causing some one example of a species with these attributes that is Federally
popn isolation)] or State listed)]
pop3 [patchy (habitat exists mostly as disjunct patches, moderate GrpII [Group II—Species very likely secure (there is at least one
recolonization)] example of a species with these characteristics that is
pop4 [isolated (habitat occurs only as locally isolated patches, widespread or common and not in any danger)]
low recolonization)] GrpIII [Group III—Species with intermediate security (there is at
pop5 [scarce (habitat is very scarce thru OR/WA, little/no least one species with these characteristics that is intermediate
recolonization)] in security status)]
LndscpUse [What is the landscape use category?]
lnd1 [patch (sp. likely uses only 1 homogeneous habitat patch
during life cycle)]
 20
Terrestrial and Marine Management Activities:
Links to Habitat Elements and Ecological Processes
MadeleineVander Heyden & Bruce G. Marcot

Introduction
Much has been published on the effects of human activities of the activity that would be necessary to produce an effect,
on the environment,5, 9, 16 and recent efforts have attempted is linked in the Matrix. For example, one could ask: overall,
to synthesize the literature into comprehensive, easily do road management activities have the potential to affect
accessible, digital formats. For example, several state soil structure and soil organic matter? The Management
wildlife agencies have produced Web pages that include Activities Matrix can be used as a guide to depict or predict
information on the effects of human activities on influences on HEs and ecological processes, and to pose
individual wildlife species. We believe that a weakness in concepts, frameworks, and hypotheses regarding the
this approach is its inability to capture the complexity of effects of management activities. It is necessarily not a
impacts beyond the individual species level. Ecosystem definitive model that precisely predicts site-specific
management suggests that scientists and managers should impacts resulting from the described activities.
also determine the effects of management activities on
ecological processes and functions, and on the long-term The Land Use and Management Activities
sustainability, diversity, and productivity of resources and We wanted to include activities that we thought were most
environments. This chapter discusses a Washington- relevant to Oregon and Washington, and were most useful
Oregon “Management Activities Matrix” (located on the to characterize across all land ownerships. We limited the
CD-ROM) and illustrates how it can be used with the other activities addressed to those that affect inland and
data matrices described in this volume to ask more nearshore-marine HEs only. Our activity list was refined
advanced questions than suggested by existing and edited several times with the assistance of many
information bases. For example, the Management resource scientists and managers expert in particular areas
Activities Matrix allows the manager not only to (see Acknowledgements). Thirteen broad categories of
determine which species may be affected by a particular management activities were identified, ranging from
land use, but also to assess what ecological functions are nearshore marine resource management to urban
involved, and which Habitat Elements (HEs).14 In this development, and encompassing activities occurring on
chapter we present and illustrate this unique perspective all land ownerships, locations, and jurisdictions in both
and methodology. Our approach is similar to the concept Oregon and Washington. These broad categories contain
of the food web: effects of management actions are not 152 specific activities (see Table 1).
only direct and linear but also can have surprisingly
indirect and nonlinear implications. To gauge the full Information on CD-ROM
extent of an activity’s impact, it becomes necessary to Under the Data Query section on the CD-ROM, there are
examine the entire web of influences. two submenus under Management Activities; one to query
for related habitat elements, the other to query the citations
Development of the Management that support the Management Activity Matrix. The user
Activities Matrix can query these to either obtain very specific information
Producing the Management Activities Matrix presented on individual HEs or to obtain general information from
many challenges. Environmental impacts are complex and the literature about the potential effects of an activity.
defy simple categorization. Therefore we have limited the These two queries are described below.
Management Activities Matrix only to those relationships Querying for Related Habitat Elements. Also based on
that were identified either in the literature or by an expert the published literature, this table identifies which specific
panel, and for the most part, we have reported only direct HEs are potentially affected by each management activity
impacts. We did not attempt to describe in detail the exact (either positively or negatively). In addition, under the
nature of every possible influence, because they depend heading “Query for Related Habitat Elements,” users can
heavily on many factors including local site characteristics query each management activity to obtain a
and the specific nature of the proposed activity. Any comprehensive list of all HE-management activity links.
potentially affected HE, regardless of the scale or intensity This list was developed using the literature cited in the

501
 502 Wildlife–Habitat Relationships in Oregon and Washington

table and expert panels (discussed below under Data the database along with links identified from the literature
Sources). to form a comprehensive list of all HEs potentially
associated with each activity. Individual activities may be
Querying for Citations. Based on the published literature,
queried to obtain this information (see Query for Related
this table describes the effects of a specific activity, e.g.
Habitat Elements on the CD-ROM).
“thinning” or “impounding water,” on various ecological
processes such as biological diversity or ecosystem Major Influences of Activities on Wildlife
function. Throughout Washington and Oregon, management
Linking HEs to management activities allows the activities affecting the most wildlife species (>500) pertain
manager to query the various data matrixes in this volume, to conversion of habitat for development of human
asking such questions as: What species are associated with habitation, recreation, mineral extraction, forestry, water
the HEs linked to this specific management activity, and, supply development, controlling and prescribing fire, and
furthermore, what are their Key Ecological Functions livestock grazing (Table 1). The management activities
(KEFs)?11 We illustrate this approach in detail in the section affecting the least wildlife species (<100) include
“How to Use the Matrix: Example Queries.” mushroom harvesting, snow-related recreation, creation
of artificial nest sites, and providing dead and down wood
Data Sources
(Table 1). If the dead and down wood category is a
We used two sources of data to populate the Management
surprise,15 note that this activity still influences some 96
Activities Matrix: the published literature and an expert
wildlife species and 9 HEs.
panel. During the literature review, we used several
The ranked order of management activities having the
reference databases to search titles, keywords, and full
most and least influence on number of wildlife species
abstracts for information, using the management activities
and HEs varies somewhat among specific wildlife habitats.
as key words. We searched literature on wildlife,
However, the general pattern of land-development
ornithology, fisheries, mammalogy, and zoology. The
activities having the greatest effect remains mostly the
wildlife database search included all papers published
same across most terrestrial habitats, including coastal
during 1937-97 in the Journal of Wildlife Management,
environments. In addition, not surprisingly, other
Wildlife Monographs, and the Wildlife Society Bulletin (>8,400
management activities related to aquatic resource
records). The ornithology database search included all
management (Management Activity 2; see Table 1) and
papers published during 1955-97 in the Auk, Ornithological
marine activities (Management Activity 10) show up as
Monographs, Condor, Studies in Avian Biology, Wilson
potentially influencing a large number of species and HEs
Bulletin, and the Journal of Field Ornithology (>18,000
in coastal environments. For example, in Bays and
records). The mammalogy database search included all
Estuaries (Wildlife Habitat 28), the most influential
papers published during 1950-97 in the Journal of
management activity is recreational development
Mammalogy, Mammalian Species, and American Society of
(potentially influencing 173 wildlife species in that
Mammalogists Special Publications (>10,900 records). The
habitat); other activities of major influence (potentially
ecological database search included papers published
influencing >150 wildlife species) pertain to conversion
during 1945-97 in Ecology, Ecological Applications, and
of native habitats; road and building construction;
Ecological Monographs, plus some additional papers
controlling water pollution; dredging; harbor, marina, and
published in the Journal of Vegetation Science (1900-97), for
ferry terminal development; wastewater treatment; and
a total of >12,000 records. We also obtained references from
water level management.
our personal reference databases, project library, and the
Cambridge Scientific Review. How to Use the Matrix: Example Queries
Whenever a link was found in the published literature Using a relational database such as Access or Paradox,
that associated a particular management activity with an the following is an example of a query on fire management
ecological process or a HE, we coded the information in that illustrates specifically how the Matrix may be used.
our database. If the information was from research, we Let’s say we’re interested in the influence of the
briefly described the results, where the study took place, management activity “low- to moderate-intensity burns”
at what time of year, and in what habitat. When the (Management Activity 1B), as may be used during
referenced links were not the results of research, the above prescribed burning to help restore some grasslands and
information was filled out to the extent possible. We listed forests east of the crest of the Cascade Mountains in
habitats as they were described, and did not attempt to Washington and Oregon, and how such influences may
“crosswalk” the habitat classes to those used in this differ between grassland and forest environments.
volume (see Wildlife Habitats).3 We first linked the Management Activities Matrix to
To ensure that all potentially affected HEs were the Habitat Elements (HEs) Matrix and counted 83 HEs
properly identified for each activity, we convened an potentially influenced by this activity (the HE tally here
expert panel consisting of scientists and managers with refers to HE category headings and subheadings, so there
backgrounds in the various activity areas (see is some redundancy in these figures). Next, we narrowed
Acknowledgements). The panelists reviewed each activity this by linking the number of HEs affected by this activity
and indicated which HEs were potentially linked to it. to those only associated with wildlife species that occur
Links obtained from the panels were incorporated into
 CHAPTER 20: MANAGEMENT ACTIVITIES 503

Table 1. Management activities potentially affecting the greatest, and least, number of wildlife species,
and the associated number of potentially affected Habitat Elements (HEs) across all wildlife habitats
in Washington and Oregon.*

Management activity No. wildlife species No. habitat

affected elements
affected

Greatest influence (>500 species)

Conversion of native habitats 577 74
Recreational developments 570 128
Road construction and obliteration 555 70
Mineral exploration 553 123
Building houses and businesses 552 149
Surface/strip mining and processing 544 141
Forest management (in general) 532 64
Conversion of shrubland to native or non-native grassland (for livestock management) 528 51
Conversion of shrubland to native or non-native grassland (for shrubland or grassland management) 528 51
Clearcutting 525 88
Establishing/maintaining greenways and greenbelts 517 105
Suppressing wildfire 514 83
Prescribed/controlled high intensity burns 503 91
Increasing water supply 503 66
Decreasing water supply 503 66
Livestock grazing 500 70
Least influence (<100 species)
Retaining/providing dead/down wood 96 9
Creating/maintaining islands or rafts within impoundments 61 3
Providing artificial nest sites (for agricultural activities) 59 2
Providing artificial nest sites (for forest habitat management activities) 59 2
Snowshoeing/snow skiing/sledding 42 2
Snowmobiling 42 2
Harvesting wild mushrooms 26 1
* See Appendix for management activity codes and definitions.

in Habitat 15, Eastside (Interior) Grasslands. This asks the habitats. Thus, as should be expected, the potential
question, what array of HEs for wildlife are potentially influence of this kind of burn on wildlife species is different
influenced by low- to moderate-intensity burns, in eastside grasslands and forests, and a greater proportion
specifically in Eastside Grasslands? The result was a list of affected species in the forest habitat would be uniquely
of 73 HE categories. We next compared this with use of influenced.
low- to moderate-intensity burns in Wildlife Habitat 7,
Ponderosa Pine Forests and Woodlands, and this produced Discussion
a list of 78 HE categories. Stressors and Indicators. Why should the manager be
However, even though the number of HEs is similar concerned about evaluating influences of management
between these two habitats, the actual HE categories and activities on Habitat Elements and wildlife? Beyond the
the associated array of wildlife species may differ. We obvious reasons—that is, environmental and biological
tested this by listing wildlife species rather than HEs in assessments to meet legal and regulatory mandates—the
each query, saving those results, and comparing the results Management Activities Matrix provides a basis for
between the two habitats. It turns out that the HEs explicitly and repeatably describing the potential
potentially affected by low- to moderate-intensity burns influences of stressors on wildlife communities. The model
are virtually identical in these two habitats (72 of the of identifying such stressors (e.g., see the species influence
potentially affected HE categories are shared by these two diagram 11) can be central to identifying the most
habitats). However, of the 163 wildlife species in Eastside influential management activities for mitigation, and
Grasslands and 218 species in Ponderosa Pine Forests and prioritizing potential wildlife responses for monitoring.10,
16, 18
Woodlands that have HEs potentially influenced by this What needs empirical work is validating, refining, and
activity, only 99 of these species occur in common between quantifying the linkages between management activities
these two habitats. That is, 39% of the potentially fire- and habitat elements and associated wildlife populations.
affected wildlife species in the Grasslands habitat and 55% Although we have purposely avoided the problematic
of those in the Forest habitat occur uniquely in these concept of management indicator species, wildlife species
 504 Wildlife–Habitat Relationships in Oregon and Washington

most susceptible to particular stressors and management Obviously, activities vary in their effects, depending
activities might be good candidates for use as bioindicators on which HEs are involved. Some HEs will be directly
and “early warning” signals of impending changes to affected and others indirectly. Both direct and indirect
other aspects of their ecosystem.13 One example is stream effects can result from a single management activity, and
amphibians that indicate levels of aquatic ecosystem can influence wildlife populations through the HEs.
stress.20 However, cryptogams, plants, and invertebrates Although all of the above-mentioned factors will
often serve as more sensitive indicators,1, 12, 19 or, in many determine the ultimate impact of an activity, they are not
cases, it may be simpler to more directly monitor specific explicitly depicted in the Matrix because of the difficulties
biochemical responses to management activities. of addressing these complex factors in a simple, qualitative
matrix format. Managers need to consider the issues
Considerations and Caveats for described above and apply the findings of the Matrix to
Using the Matrix their own geographic area and project scope. Because the
The management activities listed in the Management influence of a management activity depends largely on
Activities Matrix are necessarily described in general local conditions and circumstances, variations in specific
ways, that is, at broad geographical scales. However, effects should be expected.
because the effects of management activities act as When evaluating effects of activities, managers might
“stressors” to native systems, they vary by degree of wish to consider the following:
perturbation based on duration, frequency, intensity 1. The influence of management activities on HEs can
(amplitude), and combination of activities. Furthermore, vary from those depicted in this simple Management
the specific influence of a management activity on Activities Matrix in several ways: some depicted
ecological processes and HEs will vary according to the effects may not occur, some may occur more saliently
scale, intensity, and the action of the effect (direct or than others, and there may be some effects not
indirect). Referring to the management activity, scale issues depicted in the Matrix;
include the level of spatial resolution (the pervasiveness 2. Effects can vary over space and time, including time
of the activity), overall geographic extent and context (over lags and off-site influences; and
how large an area does the activity occur, and in what 3. Not all effects are negative; identifying the positive
type of environment), and duration of the activity. ones could be most useful for some conservation
Specifically, spatial resolution refers to the geographic objectives such as habitat restoration.
extent over which the activities and associated effects
occur. For example, some activities affect entire Conclusions and Suggestions for Further
watersheds, whereas others affect only the stream Developments
environment. Identifying the level of spatial resolution of The Management Activities Matrix provides a rigorous,
the activity determines how finely the Management repeatable basis for considering the influence of land and
Activities Matrix can be applied. The Matrix should not resource management activities on wildlife habitats and
be used to predict effects at the scale of individual species. By querying the Matrix in conjunction with the
vegetation stands or point locations, but it can be used to other habitat- and species-based databases offered in this
predict overall, general effects averaged at the scale volume, the manager can be prompted to consider effects
provinces, habitats, and habitat structures. across a full range of environmental conditions and
Geographic extent and context influence an effect’s wildlife species groups. Certainly, some activities and
intensity. The geographic extent of agricultural wildlife species warrant far more detailed attention than
conversions in eastern Washington and Oregon, for can be provided here.
example, is quite widespread; agricultural activities in that Spatially, the Matrix might prove most useful when
region may have different effects on wildlife community applied to broad geographic areas such as National Forest
composition, structure, and function than will small Ranger Districts, or to general land use allocations. At finer
inclusions of the same activities in a matrix of native spatial scales, such as individual vegetation stands or point
grassland or forest. Likewise, agricultural activities locations, the Matrix might overestimate the number of
adjacent to a wetland will impact wildlife communities HEs and wildlife species affected by given management
differently than agricultural development in an upland activities.
context.2 Temporally, the Matrix does not specify how quickly
Time duration or temporal scale refers to the influence or or for how long an effect can be expected. Stressors to
persistence of effects over time, in terms of how long a ecosystems operate in different ways depending on their
management activity has persisted in an area and how initial intensity as well as duration. Also, effects of some
long wildlife communities and populations have had to management activities may be indirect or incur time lags.
respond. It also refers to the immediacy of an influence; The manager could query the Management Activities
some effects might be time-delayed. Two contrasting Matrix and determine likely HEs and species affected
examples are faunal relaxation (time-delayed loss of (positive and negative), and then sort those according to
species from isolated native habitats), and recolonization expected levels of intensity, duration, and time lag.
of restored native environments. Local conditions, site history, and the range of specific
activities greatly influence how any given management
 CHAPTER 20: MANAGEMENT ACTIVITIES 505

activity will affect habitats and wildlife. Thus, use of the Literature Cited
Management Activities Matrix may best be seen as helping 1. Anderson, E. W. 1986. Plant indicators of effective environment.
the manager to develop working hypotheses of effects Rangelands 8:70-73.
warranting expert review and, where needed, empirical 2. Boutin, C., and B. Jobin. 1998. Intensity of agricultural practices and
validation through local testing. At best, the Matrix can effects on adjacent habitats. Ecological Applications 8:544-557.
3. Chappell, C. B., R. C. Crawford, C. Barrett, J. Kagan, D. H. Johnson, M.
be used as a basis for devising and prioritizing adaptive O’Mealy, G. A. Green, H. L. Ferguson, W. D. Edge, E.L. Greda, and T.A.
management monitoring studies to better discern and O’Neil. 2001. Wildlife habitats: descriptions, status, trends, and system
refine the initial, crude estimates of effects. dynamics. Pages 22-114 in: D. H. Johnson and T. A. O’Neil, managing
The next level of development of the Management directors. Wildlife-habitat relationships in Oregon and Washington.
Activities Matrix could then entail refining the activity Oregon State University Press, Corvallis, OR.
categories for greater detail, and quantifying the specific 4. Costanza, R., F. H. Sklar, and M. L. White. 1990. Modeling coastal
landscape dynamics. BioScience 40:91-107.
influence of activities on HEs and wildlife species. Several
5. Dale,V. H. 1997. The relationship between land-use change and climate
tools may be useful for quantifying the influence of change. Ecological Applications 7:753-769.
management activities into the “causal web” of wildlife 6. Engle, D. M., D. J. Bernardo, T. D. Hunter, J. F. Stritzke, and T. G. Bidwell.
communities, including use of Bayesian belief networks8 1996. A decision support system for designing juniper control
and sequential Bayes statistics, 7 decision-aiding treatments. AI Applications 10:1-11.
programming,6 and dynamic, stochastic simulation 7. Gazey, W. J., and M. J. Staley. 1986. Population estimation from mark-
models.4 The influence diagram approach presented in recapture experiments using a sequential Bayes algorithm. Ecology
67:941-951.
Chapter 6 can provide an overall framework for such
8. Haas, T. C. 1991. A Bayesian belief network advisory system for aspen
further modeling exploits.17 In the end, the challenge to regeneration. Forest Science 37:627-654.
the manager is to consider the influence of management 9. Kryuchkov, V.V. 1993. Extreme anthropogenic loads and the northern
activities in the full web of ecological interactions within ecosystem condition. Ecological Applications 3:622-630.
the ecosystem. 10. Maltby, L. 1999. Studying stress: the importance of organism-level
responses. Ecological Applications 9:431-440.
Acknowledgments 11. Marcot, B. G., and M. Vander Heyden. 2001. Key ecological functions of
wildlife species. Pages 168-186 in: D. H. Johnson and T. A. O’Neil,
Many people contributed their expertise to the
managing directors. Wildlife-habitat relationships in Oregon and
development of the Management Links Matrix. We are Washington. Oregon State University Press, Corvallis, OR.
indebted to the following biologists and land managers: 12. McGoech, M. A. 1998. The selection, testing, and application of
terrestrial insects as bioindicators. Biological Review 73:181-201.
Database development: Susan Tank, Marla Trevithick, and
13. McLaren, M. A., I. D. Thompson, and J. A. Baker. 1998. Selection of
Thomas O’Neil. vertebrate wildlife indicators for monitoring sustainable forest
Classification, definition, and review of management management in Ontario. Forestry Chronicle 74:241-248.
14. O’Neil, T. A., K. A. Bettinger, M.Vander Heyden, B. G. Marcot,
activities: Susan Tank, Thomas O’Neil, David H. C. Barrett, T. K. Mellen, W. M.Vander Haegen, D. H. Johnson, P. J. Doran,
Johnson, Pat Chapman, Curt Leigh, Randy Carman, L. Wunder, and K.L. Boula. 2001. Structural conditions and habitat
Cheryl Friesen, Donavin Leckenby, and Doug Runde. elements of Oregon and Washington. Pages 115-139 in: D. H. Johnson
and T. A. O’Neil, managing directors. Wildlife-habitat relationships in
Literature Review: Susan Tank, Derek Stinson, and Kelly Oregon and Washington. Oregon State University Press, Corvallis, OR.
Bettinger. 15. Rose, C. L., B. G. Marcot, J. L. Ohmann, K. L. Waddell, B. Schreiber, T. K.
Mellen, and D. L. Lindley. 2001. Decaying wood in Pacific Northwest
Expert panelists: Susan Tank, Cheryl Broyles, Paul
forests: concepts and tools for habitat management. Pages 580-623 in:
Wagner, David H. Johnson, Kelly Bettinger, E. Charles D. H. Johnson and T. A. O’Neil, managing directors. Wildlife-habitat
Meslow, and Thomas O’Neil. relationships in Oregon and Washington. Oregon State University,
Corvallis, OR.
16. Schindler, D. W. 1987. Detecting ecosystem responses to
anthropogenic stress. Canadian Journal of Fisheries and Aquatic Science
44:6-25.
17. Schlapfer, F., and B. Schmid. 1999. Ecosystem effects of biodiversity: a
classification of hypotheses and exploration of empirical results.
Ecological Applications 9:893-912.
18. Sibly, R. M. 1999. Efficient experimental designs for studying stress and
population density in animal populations. Ecological Applications 9:496-
503.
19. Stolte, K., D. Mangis, R. Doty, and K. Tonnessen. 1993. Lichens as
bioindicators of air quality. U.S. Forest Service General Technical
Report RM-224. Rocky Mountain Forest and Range Experiment Station,
Fort Collins, CO.
20. Welsh, H. H., Jr, and L. M. Ollivier. 1998. Stream amphibians as
indicators of ecosystem stress: a case study from California’s redwoods.
Ecological Applications 8:1118-1132.
 506 Wildlife–Habitat Relationships in Oregon and Washington

Appendix Code Activity Definition

Land use and management activities depicted in 2G Burning wetlands to Periodic, low intensity burning of
the Management Activities Matrix. maintain successional wetlands that occurs in association
stages with rivers, lakes, and streams. The
extent, intensity, and timing are
Code Activity Definition either planned or controlled.
2H Restoration of Revegetation with native wetland
1 Fire Management wetlands species, and the maintenance of
1A Suppressing wildfire Actively extinguishing or water levels for the majority of the
preventing wildfires. year.
1B Low- to moderate Fires that are usually intentionally 2I Wetland management Describes a variety of methods by
-intensity burns lit (or natural fires that are allowed techniques which wetland ecosystem function
to burn) for a specific management is maintained.
objective. The extent, intensity, and 2J Flooding fields and Flooding that would, without
timing are either planned or wetlands human intervention, occur
controlled. Low-intensity burns normally in agricultural habitats
usually are repeated at regular time and other nonforest environments
intervals. due to site conditions and the
1C High-intensity burns Usually natural fires that are water table.
allowed to burn for a specific 2K Removing riparian Removal of trees and shrubs within
management objective.The extent, vegetation 30 m of a waterway.
intensity, and timing are either 2L Livestock grazing of The effects of primary grazers
planned or controlled. High- riparian areas (cattle, horses, and sheep) within
intensity burns are usually a one- 366 m of a waterway.
time event or occur very 2M Adding coarse woody Intentional addition of materials to
infrequently. debris and boulders to enhance aquatic habitat conditions
streams and rivers within streams and rivers.
2 Freshwater Wetland, Riparian, and Aquatic 2N Removing coarse Intentional removal of materials
Resource Management woody debris from from streams and rivers resulting
2A Creating and Activities include the construction streams and rivers in degradation of aquatic habitat
maintaining of main and retention dams that conditions.
impoundments create impoundments >4 ha. 2O Restoring/maintaining Restoring or maintaining beaver
2B Controlling water The effects of raising and lowering beaver populations populations will retain the primary
levels water levels within an function of beavers: to deliver
impoundment, assuming the high down wood to aquatic systems and
and low water marks are already produce small impoundments
well established, and considering (<0.8 ha).
only those effects that occur 2P Retaining riparian Activities associated with
within the variable zone. buffer strips maintaining trees and shrubs within
2C Creating/maintaining Naturally occurring islands that 30 m of a waterway.
islands or rafts result from high water levels 2Q Armoring banks for Enhancing bank stability within
within impoundments cutting off peninsulas, and human- erosion control streams and rivers (e.g., riprap).
made rafts created from a variety 2R Controlling sediment- Bank stabilization technique in
of materials. Both rafts and islands ation by revegetation streams and rivers.
are <0.8 ha. Also includes dredge of banks with grass-
spoil islands. sedge-forb mixtures
2D Draining wetlands, Effects associated with the draining 2S Controlling water Controlling point-source pollution
marshes, ponds, lakes of fully functional aquatic systems. pollution discharge into lakes, streams, rivers,
2E Increasing water Assumes, within the context of a or nearshore marine waters.
supply stream, wetland, or small lake (<4 2T Disposing/assimilating Controlling waste water effluent
ha), that flooding results in an wastewater discharge into lakes, streams, rivers,
increase in water supply that is or nearshore marine waters.
sustained for 2-3 months (or 2U Dredging Periodic dredging and deposition
more) over several years. of spoils by large ships or barges
2F Decreasing water Flow withdrawal occurs within the within large rivers.
supply context of a stream, wetland, or 2V Locating/constructing The construction of roads and
small lake (<4 ha) and these water stream crossings bridges across creeks and small
bodies normally contain open rivers; includes heavy equipment,
water for most of the year. blasting, and landscape alteration.
 CHAPTER 20: MANAGEMENT ACTIVITIES 507

Code Activity Definition Code Activity Definition

2W Controlling aquatic Activities, including herbicide 4F Maintaining grasses Leaving vegetation around desired
plants application and water drawdowns, and forbs within crops to provide habitat for
to reduce or remove emergent or orchards, Christmas wildlife.
submergent plants usually tree farms, etc.
associated with reservoirs or 4G Providing/maintaining Includes providing cover in the
impoundments. vegetation along field form of hedge rows, shelterbelts,
2X Channelization Creating passageways to direct the and ditch margins or other vegetated corridors.
flow of water. 4H Retaining crop residue Practicing harvest methods that
leave crop remains on the ground
3 Road Management Roads that are engineered and
over the winter.
maintained; the surfaces of
4I Implementing farm- Activities focused on the
which can be pavement,
land conservation restoration and maintenance of
gravel/rock/cinder, or dirt.
programs predominantly native vegetation
Does not include skid roads.
and erosion control measures on
3A Road and bridge The actual construction (or
lands formerly managed for
construction/ obliteration) of roads and bridges,
agricultural commodity production.
obliteration which includes heavy equipment,
4J Irrigating Routine application of water to
blasting, and landscape alteration.
row crops or pastureland.
3B Operational aspects Activities associated with the
4K Altering drainage Includes ditching and tiling on a
of road maintenance maintenance and use of roads and
recurrent basis on lands used in
and use bridges, which includes roadside
the production of agricultural
vegetation management (mowing,
commodities.
herbicides, ditch cleaning,
4L Decreasing water Pumping water out of streams,
revegetating roadsides, introducing
supply: flow rivers, and wetlands for irrigation,
exotic vegetation, removing hazard
withdrawal resulting in decreased water supply
snags), removing beaver dams that
in aquatic habitats.
cause road flooding, spreading oil
4M No-till or minimum- Crop production techniques that
for dust abatement, and runoff
till farming minimize soil disturbance.
management (e.g., culverts) to
4N Clean farming Intensive agriculture that uses all
reduce erosion, turbidity, and
available land surface, leaving no
contamination of waterways by
crop residues (often due to tilling
heavy metals.
and burning post-harvest).
3C Road closures Limiting road use (seasonal and
4O Strip intercropping Harvest technique to conserve soil
yearly closures with gates or some
by removing alternate rows of
other system, but road is still
vegetation.
maintained).
4P Conversion of native Replacing native forest or
3D Bridges (in general) Literature search resulted in
habitats shrubland/grassland habitats with
article(s) that described the
agriculture.
general effects of bridges.
4Q Control of verte- Use of repellents, including
3E Roads (in general) Literature search resulted in
brates considered to chemical, visual, and noise, also
article(s) that described the
be agricultural pests includes trapping to remove
general effects of roads.
animals causing damage to crops.
4 Agricultural Activities 4R Providing artificial Nest boxes placed along roads and
4A Applying fertilizers Periodic application of fertilizers to nesting sites near ponds to enhance passerine
agricultural habitats such as row and waterfowl productivity.
crops, orchards, nurseries, etc. 4S Agriculture (in Literature search resulted in
4B Applying pesticides Periodic application of pesticides general) article(s) that described the
to agricultural habitats such as row general effects of agriculture.
crops, orchards, nurseries, etc.
5 Shrubland and Grassland Management
4C Applying herbicides Periodic application of herbicides
5A Mechanical vegetation Physical removal of vegetation,
to agricultural habitats such as row
management including chaining.
crops, orchards, nurseries, etc.
5B Burning Fires that are intentionally lit (or
4D Applying fungicides Periodic application of fungicides
natural fires that are allowed to
to agricultural habitats such as row
burn) for a specific management
crops, orchards, nurseries, etc.
objective. The extent, intensity, and
4E Haying/mowing Vegetation removal on row crops
timing are either planned or
and pasturelands.
controlled.
5C Use of herbicides Periodic application of herbicides
to shrubland/grassland habitats.
 508 Wildlife–Habitat Relationships in Oregon and Washington

Code Activity Definition Code Activity Definition

5D Restoration Activities used to recreate or 8B Surface/strip mining Surface and strip mining are
enhance native grassland or and processing techniques that allow the
shrubland habitats. extraction of shallow ores and
5E Conversion of Conversion to grassland to provide coal. Processing associated with
shrubland to native forage for livestock. these techniques involves the
or non-native extraction of valuable materials
grassland from mixed ore and can include
5F Livestock grazing Allowing cattle, horses, or sheep to heap leach, vat leach, flotation, and
forage on open rangeland. other techniques. These
5G Shrubland manage- Literature search resulted in techniques involve the removal of
ment (in general) article(s) that described the overburden; ore processing; waste
general effects of shrubland rock disposal; tailings disposal and
management. embankment construction; water
5H Grassland manage- Literature search resulted in supply development, storage, and
ment (in general) article(s) that described the runoff management; and power
general effects of grassland supply development.
management. 8C Underground mining Underground mining is the
and processing extraction of mineralized zones by
6 Livestock Management (cattle, sheep, and horses)
underground methods. Processing
6A Livestock grazing Allowing cattle, horses, or sheep to
associated with these techniques
forage over wide areas.
involves the extraction of valuable
6B Conversion of shrub- Conversion to grassland to provide
materials from mixed ore and can
land to native or non- forage for livestock.
include heap leach, vat leach,
native grassland
flotation, and other techniques.
6C Creating or providing The presence of stockponds and
These techniques involve site
stockponds their influence on livestock impacts
preparation; ore processing; waste
to the environment (e.g., may
rock disposal; tailings disposal and
reduce impacts to naturally
embankment construction; and
occurring streams).
water supply development, storage,
6D Excluding livestock Removing or preventing livestock
and runoff management.
from riparian areas from access to streams and the
8D Maintaining access to Leaving openings allows for wildlife
resultant restoration of riparian
abandoned subsurface use of mines and tunnels.
habitats.
mines and tunnels
7 Fencing 8E Placer prospecting Searching for, and recovery of,
7A Fencing to control Includes fencing to (1) exclude and mining minerals from streamborne
or direct wildlife predators from other wildlife or deposits. Usually involves the use
access livestock, (2) exclude ungulates of water to aid recovery from
from orchards, hay stacks, and these deposits. These activities
seedlings, and (3) control wildlife include excavation of materials
access and movements along from these deposits and processing
roadways. with various equipment including
7B Fencing to protect Nonriparian, includes aspen and pans, sluice boxes, suction dredges,
or restore habitat special botanical areas. and highbankers. The primary
7C Fencing to exclude Preventing livestock access to impacts of these activities are the
livestock from riparian streams and the resultant redistribution of existing in-stream
areas restoration of riparian habitats. sediment, introduction of new
sediment from adjacent uplands,
8 Mining Activities negative impacts to in-stream
8A Site reclamation The purpose of reclamation is to invertebrates, fish, and fish
return the disturbed areas to a spawning areas, and alteration of
stabilized and productive condition the stream channel dynamics.
following mining and milling 8F Mineral exploration This activity reflects the
activities to protect long-term land, exploration of potential mineral
water, and air resources in the deposits and primarily involves
area. This most often involves significant amounts of road
modifying the final grade of gravel construction (often in a grid
and substrate materials, providing pattern), the drilling of test holes,
for soil stability, planting vegetative and associated land-clearing
cover, and addressing water flow activities by heavy equipment.
and quality aspects.
 CHAPTER 20: MANAGEMENT ACTIVITIES 509

Code Activity Definition Code Activity Definition

8G Sand, gravel Surface mining to extract building 9B6 Prescribed burning Fires that are intentionally lit for a
(aggregate), and peat and construction materials specific management objective such
mining including sand, gravel, and rock. as forest health or site preparation.
Activities are typified by rock and The extent, intensity, and timing
gravel quarries. Aggregate mining are either planned or controlled.
does not involve the chemical 9B7 Applying insecticides Periodic application of insecticides
processing of materials. Peat to prevent loss of tree rigor and
mining is included here as it mortality.
reflects similar methods for the 9B8 Forest management Literature search resulted in
extraction of subsurface materials. (in general) article(s) that described the
8H Mining (in general) Literature search resulted in general effects of forest
article(s) that described the management.
general effects of mining. 9C Site Preparation/Tree Establishment Activities
8I Mining activities Literature search resulted in 9C1 Applying herbicides Application of herbicides to reduce
involving blasting article(s) that described the competition to seedlings from
general effects of blasting. encroaching vegetation.
8J Oil and gas extraction Literature search resulted in 9C2 Fertilizing plantations Application of fertilizers to
article(s) that described oil and/or increase tree growth and site
gas extraction. productivity.
9C3 Removing slash Includes use of heavy equipment
9 Forest Managementa
and piling, burning, or hauling off of
9A Harvest Operation Activities
slash for chipping; all slash is
9A1 Clearcutting The harvesting of all standing trees
removed from site.
in a given area at the same time.
9C4 Planting/seeding Includes the use of vexar tubing,
9A2 Shelterwood cuts Harvesting in which trees on a site
shade cards, and plastic sheeting.
are removed in a series of cuts
9C5 Tilling prior to Mechanical preparation of the
over time to create an even-aged
planting ground to facilitate tree planting.
stand.
9D Habitat Management Activities
9A3 Seed tree cuts Harvesting in which a cut removes
9D1 Maintaining mature/ Includes maintaining forest
almost all the trees in an area, but
old growth corridors.
leaves a few scattered mature trees
9D2 Grazing livestock Allowing horses, sheep, or cattle to
of good genetic stock to produce
freely forage within forested
seed to regenerate the stand.
stands.
9A4 Group selection The selective removal of small
9D3 Retaining medium Leaving 28-48 cm dbh trees from
groups of trees in a system of
green trees prior stand.
uneven-age management.
9D4 Retaining large Leaving >51 cm dbh trees from
9A5 Selective harvest The selective removal of single
green trees prior stand.
across all tree sizes trees in a system of uneven-age
9D5 Retaining defective Deformities include cavities,
management.
trees broken tops, heart rot, conks,
9A6 Selective harvest of Includes varied silvicultural
multiple tops, etc.
specific tree sizes, prescriptions, including salvage
9D6 Creating/maintaining Many harvesting activities result in
conditions, or species harvests.
edges contrasting boundaries between
9B Silvicultural/Stand Improvement Activities
forest stands of various
9B1 Precommercial Removal of young trees to increase
successional stage or species
thinning the growth of remaining trees in a
composition.
stand. The cut trees are often left
9D7 Retaining mast trees Leaving hardwood species that
as slash on the forest floor.
produce soft or hard mast for
9B2 Commercial thinning Release cuttings to manipulate
wildlife use.
stocking densities for enhancement
9D8 Retaining forest Pertains to the creation of small
of dominant or codominant trees;
openings forest openings (gaps), or the
cut trees have commercial value
prevention of tree encroachment
and are removed from the stand.
into natural meadows.
9B3 Pruning The removal of lower limbs to
9D9 Retaining brush/ Leaving the unwanted vegetation
create higher quality wood and
slash piles left from a harvest operation in
enhance tree growth rates.
piles on the site for wildlife use.
9B4 Simplifying species Genetically selecting trees, favoring
9D10 Retaining/providing Leaving or providing dead wood
composition and/or one or few commercially desired
dead/down wood from harvest operations on the
structure species.
forest floor for wildlife use (instead
9B5 Type conversion Changing grasslands to tree farms,
of collecting or burning it).
or hardwood stands to conifer
stands.
 510 Wildlife–Habitat Relationships in Oregon and Washington

Code Activity Definition Code Activity Definition

9D11 Retaining/creating Allowing standing dead trees to 10B Harbor, marina, and Includes both the development and
snags remain in the stand after ferry terminal subsequent use of harbors,
harvesting, or topping/blasting live development marinas, and ferry terminals. This
green trees to create new snags. category reflects both fresh and
9D12 Retaining riparian Activities associated with saltwater environments. Includes
buffers maintaining trees and shrubs within commercial shipping, associated
30 m of a waterway. cargo handling, and ferry transport.
9D13 Providing artificial Employing nest boxes to enhance Recreational boat marinas and
nest sites productivity of selected forested associated infrastructure (e.g.,
species that use cavities. parking lots, floats, breakwaters,
9D14 Creating/maintaining Deliberately providing or retaining fueling stations). Commercial
corridors connective habitat to facilitate harbors and ferry terminals are
wildlife travel and use. typified by Elliott Bay, Port Angeles,
9E IncidentalActivities and Bellingham Bay,Washington;
9E1 Introducing exotic An example is elk forage mix. and Newport, Coos Bay, and
vegetation Portland, Oregon. Recreational
9E2 Creating water Digging pumper ponds or wildlife marinas are typified by Olympia
sources ponds. and Des Moines, Washington; and
9E3 Removing hazard Removal of trees deemed to be Astoria, Oregon. Impacts extend
trees hazardous to human safety from to include bilgewater and wakes
roadsides and campgrounds to from large ships.
comply with federal regulations. 10C Residential docks in Floating and fixed docks, piers, and
9E4 Building skid roads The creation of skid roads and marine and associated pilings in marine and
and landings landings opens up forest canopies, freshwaters freshwater environments. Physical
increasing the amount of edge and dimensions of docks tend to be
early successional habitats. about 2 m wide and 15-30 m long.
9E5 Forest vertebrate Activities to prevent animal Typical dock structures have
pest control damage or to remove offending associated pilings and deck
individuals. surfaces.
9F Special Forest Products 10D Toxic spills in fresh Spills or depositions of chemicals
9F1 Firewood cutting Removal of live trees or snags for and saltwater into freshwater and marine
the purpose of obtaining fuel. habitats. This is typified by, but not
9F2 Harvesting wild Commercial harvest of wild limited to, petroleum spills, railroad
mushrooms mushroom species. car incidents, semi-truck turnovers,
9F3 Bough collection The pruning of lower limbs for and marine Superfund sites (e.g.,
decorative purposes. Commencement Bay, Elliott Bay,
9F4 Special forest Literature search resulted in Washington). Chemicals are
products (in general) article(s) that described the primarily represented by
general effects of collecting special hydrocarbons, dioxins,
forest products. petrochemicals, fertilizers,
pesticides, and heavy metals. This
10 Marine Activities
category does not include spill and
10A Marine dredging Mechanically or hydrologically
deposition sites that are entirely
and filling removing bed materials (sand,
terrestrial-based.
gravel, mud) and moving them to a
10E Marine shoreline Placement of rock, wood, or
new location to provide increased
armoring concrete at the water’s edge to
depth for boat and ship navigation.
prevent shoreline erosion or bank
Filling is the placement of dredged
failure. Bulkheads are sometimes
material or upland materials in
placed in no-eroding areas.
marine aquatic areas. In Puget
10F Developing under- The active creation of underwater
Sound, fill materials are typically
water marine structures, normally involving
placed to create uplands for
structures placement of large concrete and
commercial purposes (e.g., marina,
rock substrates. Objective is to
port developments). Fill material
provide vertical relief to create
has been used to create dredge
habitat structures for various
spoil islands along the lower
marine fish and shellfish. These
Columbia River.
underwater reef structures could
be 15 m wide, 61 m long, and 3 m
tall. The structures are located
primarily in Puget Sound,
 CHAPTER 20: MANAGEMENT ACTIVITIES 511

Code Activity Definition Code Activity Definition

Washington. Note: oil exploration

12 Recreational Activities
and associated drilling platforms
12A Trail use and camping All activities associated with trails
are currently prohibited off the
and camping. Includes the use of
Oregon and Washington coastlines,
pack animals and mountain bikes.
and thus are not considered in this
12B Snowshoeing/snow Nonmotorized snow travel.
assessment.
skiing/sledding
10G Marine fisheries This activity reflects marine-based
12C Mountain/rock Disturbance caused to wildlife
harvest and processing of fish.
climbing from climbing of all kinds.
Primary effects are derived from
12D Motorized boating Disturbance by and effects of
trawlnet, purse-seine, and gillnet
motor boats on wildlife and
fishing techniques. Primary wildlife
aquatic habitats.
issues in Oregon and Washington
12E Nonmotorized Nonmotorized boat traffic
reflect the bycatch of marine
boating including rafts, canoes, sailboats,
seabirds (and marine mammals to a
and rowboats.
much lesser degree) in active or
12F Swimming The effects of humans in
lost fishing gear (e.g., “ghost nets”).
waterbodies.
10H Aquaculture Commercial production and
12G Off-road driving Includes the effects of all
harvest of fish (i.e., grown in net
motorized off-road vehicles,
pens) and shellfish (e.g., oysters,
including ATVs, four-wheel drive
geoducks, clams, and mussels). This
trucks, and dune buggies.
also includes impacts associated
12H Snowmobiling Motorized snow travel.
with recreational harvest of
12I Aircraft use Includes planes, helicopters, and
shellfish.
other motorized aircraft.
11 Urban Development 12J Recreational Includes ski areas, and other
11A Paving Creation of impervious surfaces, developments resorts.
(e.g., concrete or asphalt), and the 12K Fish stocking Providing fish for recreational use.
subsequent impacts associated
13 Right-of-way Management
with the loss of natural vegetation
13A Utility corridors Linear rights-of-way including
and substrates.
power lines, telephone lines, oil
11B Building houses and Converting natural habitats for
pipelines, etc.
businesses human occupation, including, for
example, single homes, apartments, Some of the Forest Management Activities were defined by D.
a

businesses, subdivisions, shopping Patton, 1992, in Wildlife Habitat Relationships in Forested

malls or industrial parks, and the Ecosystems, published by Timber Press, Portland, Oregon.
subsequent impacts associated
with the loss of natural vegetation.
11C Presence of domestic Refers to the disturbance and
animals impacts caused by dogs and cats.
11D Urban aquatic habitat Activities associated with the
management restoration and modification of this
habitat, including paving, ditching,
and channelization of urban
watercourses. Also includes the
development and maintenance of
ponds and lakes in an urban
context, for example, in golf
courses or parks.
11E Landscaping and The conversion of native
vegetation vegetation to exotic and
management ornamental plant species, including
lawns.
11F Water quality and Activities associated with the
stormwater prevention of water contamination
management from runoff.
11G Establishing and Providing undeveloped areas of
maintaining primarily natural vegetation within
greenways/greenbelts the urban matrix.
 21
An Overview of Models and Their Role in
Wildlife Management
Gary J. Roloff, George F.Wilhere,Timothy Quinn, & Steven Kohlmann

Introduction Wildlife Modeling Concepts,

Much of ecology is concerned with the search for repeated Terms, and Mechanics
patterns. When expressed in a structured, meaningful When formulating a model we: (1) identify the system of
representation of natural systems, these patterns form the interest, (2) state specific objectives, (3) hypothesize
foundation of modeling in wildlife management. Models relationships between system inputs and outputs, and (4)
are built to define problems, clarify ideas, organize construct a useful mechanism (e.g., an equation, a
concepts, communicate information, develop and test simulation, etc.) that represents these relationships. To
hypotheses, and to make predictions. Thus, models are illustrate the model building process, assume that we
crucial to the resource decision-making process.180, 183 wanted to construct a model for canopy closure. Canopy
Models should be viewed not as scientific laws that closure is defined as the percentage of ground area shaded
represent absolute truth, but rather, as hypotheses that by overhead foliage.47 It is a forest attribute associated with
offer a “purposeful representation.”182 The models we habitat quality for a number of wildlife species and it has
discuss here are used to characterize how animals respond been incorporated into some forest practices regulations.
to their abiotic (e.g., landform, soils) and biotic (vegetation, Unfortunately, canopy closure is notoriously difficult to
species interactions) environment. Animal response can measure precisely.27, 42, 198 Hence, a model that accurately
take the form of occurrence, distribution, abundance, relates canopy closure to some other forest attribute might
movements, productivity, or survival. The intent of this serve as a cost-efficient tool for assessing canopy closure.
chapter is to: Thus far in our modeling process, we have identified a
1. Introduce wildlife managers to common types of system that we wish to understand better and we have
resource management models. stated an objective—develop a tool for efficiently assessing
2. Illustrate the use of models in resource management canopy cover.
by offering simple examples. Models describe systems. A system is “any
3. Offer guidance in selecting an appropriate model for phenomenon, either structural or functional, having at
a specific use. least two separable components and some interaction
4. Provide references where managers can get more between these components . . . .” 74 For the purposes of
information. modeling, systems are decomposed into input variables,
This chapter is divided into four primary sections. The output variables, and the relationships among them.
first section presents modeling concepts, terminology, an Inputs and outputs are also known as independent and
overview of how a simple model works, and a discussion dependent variables, respectively. Typically,
of the role of models in resource management. The second “relationships” describe output variables as functions of
section discusses three general approaches to wildlife the input variables, but if feedback exists in the system,
modeling: habitat modeling, population modeling, and then outputs are a function of both inputs and outputs.
spatial population modeling. We provide examples for The complexity of relationships varies greatly from a single
each modeling approach and offer decision trees to assist mathematical equation to a multitude of equations,
wildlife managers in selecting the type of model(s) best inequalities, and logical expressions. Equations,
suited for their needs (Figures 1-3). Also in this section inequalities, and logical expressions contain variables and
we compare and contrast different model types in terms parameters. As their name implies, variables do not have
of their practical application to resource management a constant value. The simplest model has only two
issues. The third section of this chapter presents a case variables—input and output. More complex models may
study that uses a modeling approach for wildlife have multiple inputs or output variables, and may also
management. We discuss factors that should be considered have intermediate variables that represent steps in the
prior to initiating a modeling effort and demonstrate the translation of input to output. Parameters are quantities
use of the decision trees for selecting a particular model that mediate the relationship(s) among variables.
type. Finally, we surmise on the future of modeling in Parameter values are estimated from data and are often
wildlife management. treated as constants. In simulation models, however,

512
 CHAPTER 21: MODELING—INTRODUCTIONS 513

certain parameter values may vary according to a One challenge in developing meaningful models is
statistical distribution, but the distribution is described selecting the proper inputs. In the absence of empirical
by other parameters that are constants. data, inputs are identified from expert opinion, personal
For this example, assume that a forest inventory has experiences, or existing literature (which may be based
been completed that measured canopy closure. After on empirical data). If empirical data exist, there are many
summarizing our data, we infer that stand basal area is statistical techniques that can assist modelers in choosing
the independent (or input) variable most highly correlated the “best” inputs (where “best” can be defined as the set
with canopy closure, the dependent (or output) variable. of inputs that explains the most variability in the output).
Based on a priori knowledge of canopy closure, we make Consider our simple canopy closure model. The choice to
two modeling assumptions—a stand with nonzero basal use a single input variable (i.e., basal area) and a linear
area must have nonzero canopy closure, and a stand with relationship implies that we have identified the most
zero basal area must have zero canopy closure. This forces important variable affecting canopy closure. However, we
all mathematical equations through the zero-zero point know that other attributes can affect canopy closure such
of our canopy closure-basal area relationship. Upon as tree species and topography. So, did we create a good
further analysis of the data, we infer that the relationship model? The equivocal and correct answer is “it depends.”
between input and output variables is best described by Simple models are more general, easier to understand, and
the equation for a line: easier to communicate than more complex models, but
may lack realism when applied to specific cases. Including
Canopy Closure = K * Basal Area
more detail in our model (e.g., adding input variables to
The relationship has one parameter, K (although the account for tree species or topography) may make it more
intercept could be considered a parameter with value realistic but would also require more data and possibly
zero), and the value for K is derived from the data. Using more assumptions. The challenge for wildlife modelers is
other statistics, which are beyond the scope of this review, to evaluate the tradeoffs between realism, cost, and
we can evaluate the reliability of our model. The validity applicability. This evaluation depends on the complexity
of the model should also be tested by collecting new data of the system under study, availability of data, desired
on canopy closure and basal area and then comparing the outputs, and most importantly, the risk associated with
data with the model’s predictions. Validity of a model also making a wrong prediction. For example, a model that
depends on the “scope of inference”. For example, if data predicts canopy cover ±20% of the true canopy cover 80%
for our canopy closure-basal area relationship were of the time is probably useful to wildlife managers that
collected only from western hemlock (Tsuga heterophylla) need to predict the distribution of a common bird species
types in western Washington the scope of inference is in a strategic forest plan. Conversely, this type of model
limited to similar types. Therefore, the model would be may not be acceptable for a manager who may be required
invalid for Douglas-fir (Pseudotsuga menziesii) types in by law to provide >70% canopy cover around the nest of
eastern Washington. Finally, keep in mind that the an endangered species.
mathematical relationship we have developed is only a
hypothesis. Our model is the “best” relationship we could The Roles Models Play in
infer given our data. More data or more precise data might Wildlife Management
show that the relationship is not a straight line but actually We believe models are an important part of wildlife
a curvilinear function. management and thus, the question is not whether to
There is a great deal of literature on using model model but how to do it effectively.183 Starfield183 lists several
outputs and testing model validity.34, 71, 165, 180 Model validity misconceptions regarding the use of models and notes that
refers to a broad spectrum of performance standards and these often act as impediments to model use. Nonetheless,
criteria including credibility, realism, generality, precision, decisions that affect resources are continually made
breadth, and depth. 137 The common theme among despite a seemingly perpetual lack of data and
validation studies is that the utility of model output understanding. Models built for wildlife management will
depends on input data quality and realism of never perform to the standards of physicists or chemists
hypothesized relationship(s). However, the utility of because it is unlikely that ecological models can ever be
modeling is not limited to generating outputs. One of the tested conclusively.166, 180 We use models in wildlife
greatest benefits of a model is derived from the thought management to help define or reduce risk associated with
process required to build it. The modeling process requires complex decisions when data are scarce (see chapter 19).
us to explicitly state assumptions, helps us envision the The need for models in wildlife management has
system’s relationships, and increases our understanding increased in recent years for several reasons. First, models
of the critical components of systems.75 While nearly all can help quantify tradeoffs, including the risks, of
wildlife professionals know they are dealing with complex alternative management scenarios even as natural resource
systems, often the holistic thought processes are lost at issues grow increasingly complex. Second, models can
the decision-making level where attention is focused on a produce easily communicable results that transcend
“problem.” Building a model highlights aspects of systems disciplinary boundaries that are essential if all
that must be considered in making decisions.181 stakeholders are to be involved in decision-making. Third,
many models are specifically designed to help us predict
 514 Wildlife–Habitat Relationships in Oregon and Washington

the long-term consequences of our actions, a rare feature available on habitat modeling through 1986. Morrison et
of past natural resource management. Finally, models help al. 136, 137 combined habitat theory and the practical
provide a framework for formulating hypotheses and applications of research on animals and their habitats in a
research designs, which are essential parts of adaptive modeling context.
management. Wildlife modelers use habitat to model the occurrence,
abundance, distribution, or population performance of
Types of Wildlife Models animals. Habitat models can be categorized as theoretical
To help build understanding, we separated common and empirical.137 Theoretical habitat models are based on
wildlife models (hereafter models) into three classes: inputs and relationships hypothesized as important in
habitat models, population models, and spatial population describing the system. The modeler says “This is how I
models. This classification is based primarily on the think my system is.”74 This model building process may
characteristics of model inputs and outputs. Habitat be based on logic, experience, or on a model constructed
models typically have habitat quality and quantity as for some other system.75 In contrast, empirical models use
model outputs rather than population performance data collected specifically for constructing that model.
measures. Output from habitat models is an index of Following Morrison et al.,137 we developed a decision
population response to its environment. Habitat models tree to assist managers in identifying model(s) appropriate
can be designed to represent species’ occurrence, density, for their management issue (Figure 1). The decision branch
productivity, or survival. Population models, alternatively, points in Figure 1 are related to data availability and the
often use historic estimates of population performance to questions to be answered. There are undoubtedly
predict future population performance. Both inputs and exceptions to the logic presented in Figure 1. For example,
outputs of population models are measures of the habitat suitability index models have been applied to
demographic rates (e.g., fecundity, mortality, density, and multiple species situations171 and empirical data sets may
persistence). Spatial population modeling integrates be used to develop the more theoretical guild and life form
information about the geometry of habitats in space and models. Thus, the decision tree should not be viewed as a
time, how organisms interact to position themselves in rigid rule set but rather as a guide for reducing the number
their habitat and how they move. Like population of choices and for selecting the appropriate model.
modeling, the output of spatial population modeling is
often demographic. Spatial population modeling can also Theoretical Habitat Models
generate a variety of spatial outputs including Theoretical habitat models include modeling approaches
immigration and emigration rates and success, that use expert opinion, personal experience, qualitative
probabilities of population persistence, movement descriptions, and data that were collected extraneous to
patterns, and “source-sink” dynamics. the modeling effort to portray wildlife-habitat
relationships. They range from simple, qualitative
Habitat Modeling depictions of wildlife-habitat systems137: 312 to complex,
The underlying ecological concept for most habitat models multi-scale simulation models. 12 These models are
is niche theory. Elton61 is credited with defining the niche typically inexpensive to develop, easy to understand, and
as “the functional role and position of the organism in its easy to communicate (Table 1). Theoretical habitat models
community.” Many wildlife modelers attempt to quantify include the habitat evaluation procedures and habitat
important niche components and to predict a population’s effectiveness models; Bayesian and pattern recognition
response to changes in those components. This can be a models; habitat suitability index and habitat capability
daunting task, considering that niches for most species models; species-habitat matrixes; and coarse-filter/fine-
are extremely complex.93 A challenge for wildlife modelers filter and guild/life form models (Figure 1).
is to mathematically represent this complexity in a context
suitable for practical use and application. Habitat Evaluation Procedures and Habitat Effectiveness
Texts on habitat modeling date back to the 1960s8, 209 Models. One group of theoretical habitat models includes
although most work on modeling wildlife-habitat the habitat evaluation procedures 196 and habitat
relationships started in the 1970s 75, 130, 180 For general effectiveness models (Figure 1). The procedures are based
reviews of wildlife-habitat relationships modeling see on two primary components: 1) an index to habitat
Starfield and Bleloch,180 Verner et al.,204 and Morrison et condition, i.e., quality, and 2) a measure of habitat quantity.
al.136, 137 Starfield and Bleloch180 and Verner et al.204 wrote Habitat quality is determined from input data on
and edited some of the first comprehensive works relating vegetation structure, composition, and spatial
habitat modeling to wildlife management issues. Starfield arrangement of vegetation types (e.g., distance between
and Bleloch180 focused on the mechanics of applying suitable forage and cover). Habitat quantity is determined
models to wildlife management. They described several by multiplying the quality index by the corresponding
model types and supplemented their descriptions with amounts of that habitat type. The result is termed a
examples. The work in Verner et al.204 consists of 60 “habitat unit.”196 The assumption is that greater numbers
chapters (mostly case studies) that cover theory, of habitat units represent more favorable conditions (i.e.,
techniques, and the validation of habitat models. This also higher carrying capacity) for the wildlife species under
contains one of the most thorough literature reviews consideration.157 Although not intended to estimate animal
 CHAPTER 21: MODELING—INTRODUCTIONS 515

Figure 1. Decision tree for selecting a wildlife habitat

modeling approach.
that is discounted by human activity185, 213 (e.g., roads,
disturbance). For example, Suring et al.185 portrayed
numbers,6 both modeling procedures must be tested using
habitat potential for brown bears (Ursus arctos) on the
demographic data.159, 200 Once tested, these models can be
Kenai Peninsula using a habitat suitability index.
a useful tool for population viability planning.157
Subsequently, they developed a “human activities” sub-
Habitat evaluation procedures are appealing and have
model based on locations of urban areas, roads, hunting
been used extensively by regulatory agencies in the United
camps, and livestock grazing. The “human activities sub-
States to assess the adequacy of mitigation projects.196
model” was used to discount habitat potential. The
Habitat effectiveness models can be developed for low
modeling effort resulted in a single index of habitat
costs, apply to a range of similar systems, and readily
effectiveness for brown bear on the Kenai Peninsula.185
associated to real-world systems (Table 1). Both of these
model types have been criticized for their high levels of Habitat Suitability Index Models and Habitat Capability
subjectivity and lack of perceived validity (Table 1). Models. Another common theoretical habitat modeling
There are slight differences in how the habitat approach includes habitat suitability index models and
evaluation procedures and habitat effectiveness models habitat capability models. This approach produces single
are used.137 This difference depends on the use and patch indices of habitat quality (Figure 1). Developed by
interpretation of model outputs. In habitat evaluation the United States Fish and Wildlife Service, habitat
procedures, outputs are habitat units (the product of suitability modeling is part of the habitat evaluation
habitat quality index and the amount of that habitat type procedures.196 These models are typically single-species
available). In habitat effectiveness models outputs are focused and represent “within-patch” habitat conditions
indices that represent habitat conditions relative to some (Figure 1). Habitat suitability index models have also
biologically meaningful optimum213 (e.g., maximum use, received limited use for community assessments. 172
carrying capacity). Habitat evaluation procedures use Habitat suitability models generate a numerical value that
habitat units as a representation of carrying capacity indexes the capacity of a habitat patch to support a species.
whereas habitat effectiveness models score entire Inputs to habitat suitability models include measures of
evaluation areas with a single index.213 Also, habitat vegetation structure, composition, and spatial
effectiveness indices typically represent habitat potential arrangement. These inputs are mathematically combined
 516 Wildlife–Habitat Relationships in Oregon and Washington

Table 1. A coarse comparison of habitat-based wildlife modeling approaches.

Each model was ranked as unfavorable (U), moderately favorable (MF), or favorable (F),
according to the practical application of the modeling approach.
Rankings are subjective evaluations as to the historic use, performance, and perception of the models.

Model Evaluation Criteria

Applicability Model Structure Model Output

Appeal Common Cost of Cost of Training Level of Complexity Generality Face Communicability
data data model required subjectivity validity
acquisition develop-
ment
Theoretical models
Bayesian & pattern recognition
models F MF MF F MF U MF MF MF U
Habitat evaluation procedures F MF MF F MF U MF F U MF
Habitat capability modeling U MF MF F MF U MF F U F
Habitat effectiveness modeling MF MF MF F MF U MF F U F
Habitat suitability index models F MF MF F F U F F U F
Species-habitat matrixes F F F F F F F F MF F
Coarse- & fine-filter models U U U U U MF U MF MF MF
Guild and life form models MF MF MF F MF U MF F U MF

Empirical models
Community & ecosystem
simulation models F F U U U F U F MF U
Hierarchy models MF MF U MF MF MF MF MF MF MF
Community structure models MF F U MF U F U U MF U
Gap analysis models MF U U U MF MF MF F MF F
Biodiversity models U U U MF U F MF U F MF
Life history models U U MF MF F F MF F F MF
Correlation models NF MF MF MF F F MF U F F
Multivariate statistical models MF U U U U F U U F U
Habitat preference models MF U U U U F U U F MF
Optimal foraging models U U U U U F U U F U
a
Some criteria adapted from Marcot et al.124 where Appeal = How much the model framework has been used in the past and
practically applied; Common data = Data that are collected as part of routine inventories (e.g., rimber cruises, game check stations,
remote sensing); Cost of data acquisition = Cost to acquire the data for model development and use; Cost of model development = Cost
(monetary and time) of developing the model; Training required = Amount of training required to use and understand the model; Level
of subjectivity = Degree that subjective evaluations are used to develop the model relationships; Complexity = Complexity of the
modeling framework, synonymous with Marcot et al.’s “wholeness”; Generality = How well the model represents a broad range of
similar systems; Face validity = Perceived credibility of a modeling framework; Communicability = Ability to associate model output to
real world systems.

into an index that ranges from 0.0 to 1.0 representing to abundance,108, 147 capture success,63 abundance of nesting
totally unsuitable areas to optimum habitat, respectively.197 or denning structures, 156 animal locations, 107 or
In generating this index, a linear relationship between demographic performance.22
carrying capacity and the index value is assumed.197 The Habitat suitability index models are appealing as
linear relationship implies that a change in index score indicated by their historic use and common application,
will always correspond to the same proportionate change have low development costs, require minimal training for
in carrying capacity. implementation, are structured in an understandable
Habitat suitability models have been applied to manner, can typically be applied to a broad range of similar
numerous wildlife management situations,204 and their systems (but see Roloff and Kernohan),159 and are readily
performance has varied.189 Because of the relationship communicated to resource managers (Table 1). Drawbacks
between habitat suitability modeling and the habitat of using habitat suitability index models include a high
evaluation procedures, terminology for the two modeling level of subjectivity and lack of perceived validity (Table
approaches is sometimes used interchangeably in the 1).
literature. 111 The important difference between these Habitat capability models are closely related to habitat
approaches is reflected in model output. Habitat suitability suitability index models.137 In contrast to habitat suitability
models generate an index of habitat condition for a patch. index models that generate an index to habitat potential,
The habitat evaluation procedures, in contrast, generate output from habitat capability modeling has been used to
habitat units that are based on habitat quality and quantity represent population levels.135 Habitat capability models
(index score times the amount of the habitat available). typically have low development costs, can be applied to a
Both types of models can be tested by comparing outputs broad range of similar systems, and are readily
 CHAPTER 21: MODELING—INTRODUCTIONS 517

communicable to managers (Table 1). This model type has resource management (see below). Drawbacks of using
not been used extensively and has a high level of Bayesian and pattern recognition models include the
subjectivity and lack of perceived validity (Table 1). subjectivity often associated with assigning initial
probabilities, and difficulties in communicating the
Bayesian and Pattern Recognition Models. Bayesian and
outputs (Table 1).
pattern recognition models are another form of theoretical
habitat model that focuses on single species (Figure 1). Species-habitat Matrixes. Two classes of theoretical
The underlying framework of these models is Bayes’s habitat models address multiple wildlife species (Figure
Thereom, which is a statistical approach that integrates 1). Species habitat matrices (such as those outlined in this
both empirical data and subjective judgement.64 Simply, book) are tables that list vegetation types, seral stages of
Bayes’s Thereom states that the probability of an event ecological communities, or environmental conditions with
occurring is related to the degree of belief that the event which wildlife species associate. This group of models is
will occur. Knowledge and data influence this degree of designed in part to provide a rapid assessment
belief. As our knowledge regarding a system grows, our methodology for predicting the presence or relative
knowledge of system behavior improves. For example, abundance of multiple species within particular ecological
assume that we believe, a priori, that a grassland habitat communities.16, 60, 137 Species-habitat matrix information
patch becomes suitable for sparrows when vegetation may also be useful in crafting habitat evaluation
reaches a height of 2 m. The a priori probability for these procedures.
conditions, i.e., the event of sparrows occupying habitat The habitat rankings in species habitat matrices are
when it is grassland and vegetation is 2 m is 1.0 (or 100% often based on expert opinion or existing information.
probability of occupancy). We later observe that this Thus, species-habitat matrices can be built at relatively
sparrow only occurs in habitats meeting this description low costs (Table 1). Based on historical use and application,
80% of the time, thus, we may use this knowledge to adjust these models are one of the most favorable types (Table
our a priori probability down (perhaps to 0.8). This process 1), however, their lack of objectivity has caused some
is iterated repeatedly to refine a priori probabilities and to concern.16, 48, 124, 153 Also, since habitat rankings are by
create descriptions of new events. Iversen95 and Carlin32 community type, species-habitat matrices typically offer
provide thorough descriptions of Bayes’s Theorem and only within-habitat patch assessments and their validity
Williams et al.212 describe the adaptation of Bayesian theory decreases for species relying on spatial arrangements of
to wildlife habitat assessments. habitat patches.
In terms of wildlife habitat modeling, Bayesian theory Some of the more intensive species-habitat matrix work
most often occurs in the form of pattern recognition has been conducted as part of the California Wildlife
models. Pattern recognition models were initially Habitat Relationships program.1 Other models include
developed for the medical profession to assist in the Thomas’191 and Brown’s23 database for the Blue Mountains
diagnosis of disease.120 Pattern recognition models of Oregon, DeGraaf and Rudis’s49 work on New England
integrate Bayesian probabilities across habitat space. Three wildlife habitat, Marcot’s123 Wildlife Habitat Information
factors must be understood to use the pattern recognition Matrix Program for northwestern California, Verner and
approach.101 First, categories of habitat suitability into Boss’s202 model for the western Sierra Nevada, and this
which a patch can be classed must be identified. Second, book.
the habitat attributes that dictate habitat suitability must
Coarse-filter/Fine-filter Models and Guild and Life Form
be identified. Lastly, the set of probabilities that reflect the
Models. The other group of theoretical habitat models that
association between individual habitat attributes and each
address multiple wildlife species function at the
suitability class must be assigned. In the sparrow example
community-level, i.e., they either use ecological
above, we classified “grasslands” as the most suitable
communities as surrogates for population response or they
habitat type. We identified vegetation height as the
group organisms into guilds or life forms (Figure 1).
attribute that determined patch quality. Our a priori
Coarse-filter and fine-filter models are one approach for
probability was 1.0 if habitat was grassland and had
simultaneously assessing multiple species.137 Coarse-filter
vegetation 2 m high (i.e., we were certain that sparrows
models use the occurrence, abundance, and locations of
would occur if these conditions were provided). In pattern
ecological communities to predict animal responses.84, 85
recognition models, this process is repeated for multiple
When applying a coarse-filter model, considerable
habitat patches across space, thus, these models permit
attention must be directed at defining ecological
the incorporation of landscape configuration into the
communities. Haufler et al.84, 85 outline a process for
probability estimates. For a conceptual review of pattern
stratifying landscapes into “ecological land units” that are
recognition theory, see Flather and Hoekstra.64 Examples
based on similar bio-geo-climatic conditions, existing
of using pattern recognition for modeling habitats can be
vegetation structure, site potential, and historical
found for bighorn sheep,89 bald eagles,72 and pileated
disturbance regimes. These units are used to describe floral
woodpeckers.154
and faunal diversity for planning landscapes.84, 85 It is often
Bayesian and pattern recognition models can be
unclear, however, how to determine the amount and
developed for low costs, and they are easily updated or
spatial distribution of ecological land units that best
modified (Table 1). This model type is used less than other
function as a coarse-filter. Thus, fine-filter models (or
comparable model types but has been linked to adaptive
 518 Wildlife–Habitat Relationships in Oregon and Washington

species assessments) are used to check the adequacy of Empirical habitat models include life history models;
the coarse-filter.84, 85 A check is necessary because it is correlation and multivariate statistical models; community
reasonable to assume that some species will “slip through and ecosystem simulation models; hierarchical models;
the cracks” of the coarse-filter.137 Any of the previously community structure, gap analysis, and biodiversity
discussed single species habitat models that index species’ models; and habitat preference and foraging models
response (e.g., survival, fecundity) could be used as a fine- (Figure 1).
filter.
Life History Models. Empirical habitat models can be
Because of data requirements and complexity, use of
loosely categorized into single and multi-species
the coarse-filter and fine-filter approaches has been limited
approaches (Figure 1). A relatively simple, empirically
(Table 1). Recent advances in quantifying existing
derived single species approach is life history modeling.
vegetation from remotely sensed data have facilitated the
Life history models have been proposed for understanding
use of this approach. Development of large-scale models
how phenotypic traits evolved and how these traits can
that predict ecological site potentials and simulate
be used to predict habitat requirements.137 The underlying
historical disturbance regimes have also made the coarse-
premise of life history models is that the combination of
filter more user-friendly.
ecology and morphology provide consistent expressions
Guild or life form models are another multi-species,
of ecological and evolutionary adjustments between
community-based type of theoretical habitat model
phenotype and the environment.155 Here, “ecology” refers
(Figure 1). These models group wildlife species with
to the manner in which an organism relates to its
similar habitat requirements, morphology, or behaviors
environment and “morphology” refers to the physical
into guilds or life forms. For example, Inger and Colwell94
structure of an organism. Life history models are based
grouped amphibians and reptiles based on measured
on the premise that an organism’s morphology is an
associations to microhabitat characteristics. Because
expression of its environment, i.e., the physical structure
members of the groups share common characteristics, they
of an organism has evolved to optimize fitness in
are assumed to respond to changes in environmental
particular environments. For example, research has
conditions in a similar way. Guild and life form models
demonstrated relationships between organism body size
can be viewed as coarse-filter approaches and thus the
and the sizes of prey.154 The development of reliable life
concerns of individual species “slipping through the
history models requires fairly detailed data on animal
cracks of the coarse-filter” apply. The “coarseness” of a
behavior or morphology and associated habitat use
guild or life form model depends on the specificity of the
information.
animal grouping and the complexity of the environment
Life history models often require relatively little
being described. For example, Mannan et al.122 found that
training to use, have low subjectivity, and are perceived
guilds with >4 bird species in older forests in northeastern
as valid (Table 1). These models can be applied to a broad
Oregon had less predictive power than individual species
range of similar systems but only for species for which
models. The advantages of these types of models are that
data are collected (Table 1). Life history models require
they can be constructed from existing literature, they
specific data often not collected as part of routine
condense a large amount of information into tabular form,
inventories and their practical application may be limited
and they are appealing (Table 1). Drawbacks of using guild
(Table 1).
or life form models include the level of subjectivity and
low perceived validity (Table 1). Correlation Models and Multivariate Statistical Models.
Considerable research on the theory of guilds and life Another group of empirical habitat models involves the
forms has been conducted.15, 122, 174, 175 Verner203 provides a use of correlation and multivariate statistics (Figure 1).
clear understanding of a guild approach for the pine-oak These models typically portray relationships between
woodlands of California. Most often, guild models are environmental characteristics and measures of animal
used to evaluate the effects of changes in the environment abundance or distribution.137 Correlation models are most
on songbirds.25, 102, 160 Inger and Colwell94 and Hariston80 useful for demonstrating trends in wildlife habitat
applied the approach to amphibians and reptiles. The most relationships. For example, Steeger and Hitchcock184 used
familiar examples of the life form approach include a correlation analysis to demonstrate that red-breasted
Thomas191 and Brown23 for depicting habitat relationships nuthatch nests were positively associated with the density
in the western United States. of trees infected with Armillaria root disease. These types
of models are common in the habitat literature and can be
Empirical Habitat Models. Empirical habitat models
adapted to other models that require habitat quality
include approaches that are developed from data
information (e.g., habitat suitability index models, pattern
specifically collected for model building (Figure 1).
recognition models). Their ease of creation and
Empirical models can be purely descriptive, as derived
communication of results make this model type one of
from case studies, or statistical, as based on data
the more favorable empirically based approaches for
sampling.137 These models, in comparison to theoretical
practical application although their generality may be
habitat models, are often more difficult and expensive to
limited (Table 1).
apply and communicate but they lend greater scientific
credibility to the modeling process and outputs (Table 1).
 CHAPTER 21: MODELING—INTRODUCTIONS 519

Multivariate statistical models are also common in the Recently, classification and regression tree analysis and
literature. These models simultaneously relate multiple fuzzy logic modeling have received attention by wildlife
environmental attributes to animal abundance and modelers as useful multivariate approaches. Classification
distribution.137 They often take the form of “exploratory and regression tree analysis (also called CART) is used to
data techniques” for examining which input variables recursively partition data sets on the basis of a set of
(habitat characteristics) had the greatest effect on output independent variables.142 The technique is commonly used
variables (population response). For example, if songbird to develop classification trees for modeling and mapping
abundance and several habitat variables were collected species distributions,37, 90 biodiversity,141 and vegetation
at multiple sample points, multivariate models could be types (Mike McGrath, National Council for Air and Stream
used to identify which habitat characteristics appear to Improvement, Corvallis, Oregon, pers. comm.).
explain the most variation in bird abundance. Volumes of Classification and regression tree analysis tests several
literature exist on different multivariate techniques and independent variables and identifies which of those
how they can be used as predictive tools for resource variables most effectively partitions variation in the
managers. Sources for more generalized information on dependent variable. The modeling approach uses this
multivariate methods and their application to wildlife subset of independent variables to identify threshold
management include Capen,30 Dillon and Goldstein,54 and values that constitute splits in the decision tree.142 Each
Digby and Kempton.53 branch of the decision tree is subjected to this procedure
The complexity and breadth of multivariate wildlife- until certain stopping rules are encountered.142 Once the
habitat models are virtually limitless. Multivariate models decision tree is completely structured, the branches are
all use more than two input (or independent) variables in pruned to an optimal fit using cross-validation
various mathematical formulations to predict output (or procedures.142 The result of classification and regression
dependent) variables. One of the most common forms of tree analysis is a hierarchically organized decision tree.142
multivariate habitat analysis involves the use of multiple Fuzzy logic is relatively new to wildlife modeling but
regression. For example, Maurer 125 used multiple it has been used for decision-making in a multitude of
regression to develop predictive models of bird density applications, including chemistry, engineering, financial
using multiple environmental input variables. Similarly, forecasting, and mineral exploration. Fuzzy logic allows
Puttock et al.151 used an expanded form of regression one to express a degree of uncertainty around model
modeling, termed log-linear modeling, to estimate moose parameters and variables.39 The appeal of fuzzy logic to
density based on vegetation characteristics and spatial wildlife modelers is that answers to wildlife problems are
arrangements of habitats. Logistic regression and typically qualified. For example, we can seldom predict a
discriminate function models are also two forms of dependent wildlife variable with absolute certainty. More
multivariate models that are commonly used in wildlife often, we qualify our answers with fuzzy terms like
analyses. The output of these model types is categorical, “sometimes”, “probably”, or “most often”. Wildlife
most commonly probability of presence or absence. For modelers use fuzzy logic to portray these qualitative terms.
example, Nadeau et al. 139 used logistic modeling to Advantages of using fuzzy logic to model wildlife systems
determine the presence or absence of muskrat burrows include: 1) fewer values, rules, and decisions, 2) the use
along stream reaches based on vegetation characteristics of linguistic instead of numerical variables, 3) a direct
and stream morphology. Capen et al.31 used discriminate relationship between inputs and outputs without having
function analysis to categorize songbird habitats as “used” to understand all the relationships, 4) easy design and
or “unused” based on plot samples of vegetation prototyping; and 5) often a few rules encompass great
characteristics. Verner et al.204 provide some excellent complexity.132 Disadvantages of using fuzzy logic for
papers on the design and improvement of regression wildlife modeling are: 1) more simulation and fine-tuning
models. is required before models are operational, and 2) the
Principal components analysis is yet another common cultural bias that our profession has for mathematical
form of multivariate modeling approach. Principal precision may make these models less appealing. 132
components is used to identify important components of Because the very nature of fuzzy logic is imprecise, the
a multivariate data set. For example, Harestad and use of fuzzy logic models for modeling wildlife species
Keisker79 used principal components analysis to identify with regulatory impacts may be limited.
which tree attributes were important to primary cavity- Other, less common multivariate methods exist and a
nesting birds. Although principal components analysis complete discussion of these is beyond the scope of this
does not yield a predictive model per se, Maurer125 used chapter. Multivariate statistical models are appealing
the output of principal components analysis (namely the because they have a low level of subjectivity relative to
principal components) to generate a regression model. The other model types and they are perceived as valid (Table
primary difference between standard regression 1). Applicability of multivariate models is often limited
approaches (i.e., multiple and log-linear) and discriminate by data availability, cost of model development, and
function and principal components analyses is that the training that is required to interpret the model (Table 1).
former typically use a subset of input variables to develop These models are also complex, difficult to communicate,
the predictive equation whereas the latter use all of the and often apply to a narrow range of environmental
input variables to generate the model. conditions (Table 1). For more information on the use of
 520 Wildlife–Habitat Relationships in Oregon and Washington

multivariate models for wildlife habitat assessments see responses to changing environmental conditions, 2) the
Capen,30 Verner et al.,204 (Chapter 2) and Morrison et al.136 effects of catastrophic changes on populations, and 3)
(Chapter 7). overall ecosystem function.137 The simulation component
of these models permits the integration of randomness
Habitat Preference Models and Foraging Models.
into the analysis. Thus, community and ecosystem
Another category of empirical habitat models that
simulation models have been used to optimize systems
emphasizes single species includes two model types that
based on the re-occurrence of desired patterns from
use data on habitat use and availability (Figure 1). These
iterative model runs.21 The startup and maintenance costs,
models are termed habitat preference models and foraging
complexity, and difficulty in communicating the results
models (Figure 1). Habitat preference models rely on
reduce the utility of community and ecosystem simulation
statistical inference on habitat selection, i.e., the differential
models for practitioners. But once a process is established,
use of habitats as compared with their availability.137 These
these models offer powerful planning capabilities,
models are most frequently based on individual animal
particularly in combination with coarse-filter models.
selection as monitored via radio telemetry or mark-
Community and ecosystem simulation models are
recapture. The idea behind these models is that habitat
appealing, often use readily available data, reduce the
preference can be inferred from the amount of time (or
level of subjectivity by incorporating randomness, and
frequency of visits) an animal spends in a particular habitat
may apply to a broad range of conditions (Table 1). The
patch relative to the amount of the habitat available to
primary drawbacks of using these model types are the
that animal. For example, Quinn152 used radio-telemetry
costs associated with data collection, model development,
to show how coyotes in urban areas of Washington
and training (Table 1). Community and ecosystem
selected certain habitat types during routine movements.
simulation models are also complex and can be difficult
Foraging models are closely related to habitat
to communicate (Table 1).
preference models. These models predict animal welfare
or fitness based on the bioenergetics of forage intake. Hierarchy Models. The second group of multi-species
Optimal foraging theory suggests that when resources are empirical habitat models is called hierarchy models. These
not limiting, species should concentrate foraging on the models portray information at multiple spatial scales
best types of food.38 In other words, organisms chose (Figure 1). Although several of the previously discussed
habitat to optimize their energy budgets. Inputs to modeling approaches consider relationships across
foraging models include nutritional content of forages and multiple scales (e.g., coarse-filter and fine-filter models,
the organism’s energetic balance. Hobbs’s87 foraging community and ecosystem simulation models), hierarchy
model for mule deer simulates energy flow from forage models retain scale relationships and depict these
resources to the animal and the allocation of that energy relationships as a series ranking one above the other. The
to individual expenditures. The output generated by purest hierarchy models include the models developed
Hobbs’s87 model portrays the energy balance of mule deer for ecological land classification, 59, 128 however the
and he uses this information to predict deer condition and hierarchy concept is implicit in many of the previously
mortality. Foraging models have been developed for other discussed modeling approaches. For example, With and
species including caribou,17 black-tailed jackrabbits,40 Crist214 used hierarchical concepts to model grasshopper
insectivorous birds,126 and waterfowl.67 response to different habitat types and Roloff and
Both habitat preference and foraging models are Haufler157 used hierarchy theory in combination with the
analytical approaches that require extensive field data. habitat evaluation procedures to model population
These characteristics and their overall complexity often viability. Morrison et al.137 noted the relationship of
make them less appealing to practitioners (Table 1). hierarchy models to many of the community and
However, these model types can be some of the most ecosystem simulation models. Hierarchical models are
statistically robust and scientifically credible of the habitat- gaining favor throughout the natural resource sciences
based models (Table 1). Also, data from these types of because they offer a means of logically structuring
studies can be used to construct other, more simplistic complex ecosystems. The strength of hierarchical models
models (e.g., habitat effectiveness, habitat evaluation is in combination with other modeling approaches. By
procedures). building and presenting some of the more complex
models in a hierarchical fashion they often become more
Community and Ecosystem Simulation Models. We
appealing to practitioners.
divided empirical habitat models for multiple species into
three groups (Figure 1). One group of models can be used Community Structure, Gap Analysis, and Biodiversity
to project environmental conditions or evaluate the effects Models. The final category of empirical habitat models
of environmental stochasticity on wildlife habitat. These portrays multiple species habitat relationships at a
models are categorized as community and ecosystem particular spatial scale at one point in time (Figure 1). This
simulation models.143, 145, 146, 190 These models are based on category includes community structure models, gap
systems analysis and consist of inputs, outputs, and analysis models, and biodiversity models (Figure 1).
feedback loops that describe processes affecting that Community structure models describe species
system. Community and ecosystem simulation models distribution, abundance, or diversity as functions of
were designed to evaluate 1) population and system environmental complexity. 137 Like single species
 CHAPTER 21: MODELING—INTRODUCTIONS 521

multivariate statistical models, community structure expand their breadth.137 For further reading on the topic
models portray responses of wildlife communities or of biodiversity models, refer to Maurer 127 and
guilds based on empirical data. James96 used a community Rosenzweig.162 Biodiversity models have low subjectivity
structure model to display a three-dimensional ordination and thus are perceived as valid (Table 1). Drawbacks of
of a bird community in Arkansas. Each dimension these models include historically minimal use in land
represented a habitat gradient. For example, the first management decisions, costs, and their site-specificity
dimension, which accounted for 65% of the total variation (Table 1).
in bird distribution, ranged from areas with high ground Habitat models have a number of appealing attributes
cover and few trees and shrubs to areas with mature forests from which they derive their utility. First, theory, practice,
and low ground cover. 96 For another example of a and common sense tell us that habitat quality, quantity,
community structure model see James and Wamer.97 and the spatial distribution of habitats are important
Community structure models often use commonly determinants of population performance. Second, wildlife-
collected data and have low subjectivity (Table 1). These habitat relationship data (this book) and habitat data (e.g.,
models can be expensive, complex, site-specific, and vegetation inventories, remotely sensed data) are
difficult to communicate (Table 1). frequently the only type of information available to
Gap analysis models are another form of multi-species wildlife professionals. Third, inputs and outputs of habitat
habitat model. These models are based on geographic models are typically easy to understand and act upon.
distribution of many species, and can thus depict gaps in Finally, several habitat models assess management
species distributions or identify areas of relatively high impacts on a large number of species for which there often
species richness.173 The input data on vegetation are exists little more than life history or coarse-grained
derived from satellite imagery and wildlife species habitat distribution information. These multi-species models are
associations are assigned in a manner similar to the appealing because they provide information on numerous
species-habitat matrices discussed earlier. The United species across larger landscapes and are consistent with
States Fish and Wildlife Service initially established the contemporary perspectives on ecosystem management
gap analysis program at the state-level, however, use of and landscape-level planning.
the concept has spread 18 (e.g., into Mexico). First
generation gap models identified patches of habitat in Population Modeling
which wildlife species were known to occur and the Population modeling forms the basis of numerous harvest
locations of similar patches were used to infer species management plans world wide.119, 144, 170 A population is a
distributions.173 Second generation gap models use a group of individuals that is spatially, temporally or
refined vegetation classification scheme and integrate genetically separated from other groups.210 Population
minimum area requirements and landscape configuration models use demographic information such as fecundity,
into habitat assessments (Richard Minnis, Cooperative mortality, immigration, emigration, density, and age
Fish and Wildlife Research Unit, pers. comm.). structure as inputs to portray population dynamics.69
An abundance of published literature exists on the Population models can also be used to estimate which
implementation of gap analysis models; citations are parts of a population (e.g., juveniles, adult males, females)
provided at http://www.gap.uidaho.edi/gap. This site have the greatest effect on population growth.
also contains an overview of the gap analysis process, a The extensive history and theory of population
handbook for applying gap analysis, and a guide to the modeling is beyond the scope of this introductory chapter
status of gap analysis for each State in the United States. (see Andrewartha4 and Slobodkin178 for general reviews).
Other than startup costs of data acquisition and General texts on general wildlife biology and management
compilation, gap analysis models are one of the more often address population ecology and various modeling
appealing multi-species, coarse-filter modeling approaches.5, 19, 38 Dempster50 and Moss et al.138 discuss
approaches for portraying habitat relationships across basic demographic theory and environmental influences
large areas (Table 1) but see Morrison et al.137: 332 for a on population growth, with Dempster 50 providing
discussion on the limitations of the approach. examples from long-term data sets. Sutherland186 discusses
Biodiversity models are similar to gap analysis models; how certain population ecology aspects of vertebrates can
they map species richness based on vegetation attributes. be related to behavior. Fryxell and Lundberg69 address
Biodiversity models have most commonly appeared in the predator-prey modeling.
form of logarithmic series, Shannon-Wiener functions, Population models can be single- or multi-species and
Simpson’s diversity indices, Brillouin’s indices, and can integrate multiple demographic and environmental
evenness measures. 104 These approaches assess factors (Figure 2). Single-species models project population
community structure (e.g., richness, evenness, diversity) dynamics for individual species typically as an extension
based on the number of species in a sample and the of historic population trends 170 or in relation to
number of individuals of each species in a sample. environmental variables.144 Multiple species models
Although the measurement of species richness has represent interactions between two or more populations
dominated community ecology in the past, species of different species, most commonly interactions between
richness is only a partial expression of biodiversity and predator and prey. Population models and spatial
thus, there is a need to further develop these models to population models (below) can also be categorized as
 522 Wildlife–Habitat Relationships in Oregon and Washington

Figure 2. Decision tree for selecting a wildlife population

modeling approach.

deterministic and stochastic. Deterministic models assume different forms of trend models (logistic, Ricker, and
that parameter values are constant throughout the Beverton-Holt) on data from a Serengeti wildebeest
projections. For example, in a deterministic Leslie matrix (Connochaetes taurinus) population. Each of the three
(see below), demographic rates are treated as constants models require different assumptions on the feedback
for each age- or stage-class. 11, 58 Stochastic models between population size and population growth. Thus,
incorporate variability into the parameter estimates.86 Data each portrayed different population dynamics and
re-sampling techniques (e.g., Monte Carlo methods, demonstrated the importance of understanding a model’s
bootstrapping) are often used to incorporate this assumptions and structure to interpreting model output.144
variability. For example, in a stochastic Leslie matrix
Univariate Regression. To determine statistical
demographic rates by age- or stage-class are generated
significance of a population trend, a simple linear
multiple times by sampling from underlying data
regression applied to population time series (i.e., a series
distributions. Deterministic and stochastic methods exist
of estimates of population size taken at regular intervals)
for the three common approaches to modeling
is often appropriate. Transformation of population count
populations: trend analysis models, life-table or matrix
data (e.g., taking the natural logarithm of the population
models, and multiple species modeling.
measure) and various smoothing techniques (to help
Trend Analysis Models discern a trend from highly variable data) are available to
Trend analysis models are the simplest and often most estimate population change over time.168 These types of
useful form of population model. Trend analysis uses a models provide an estimate or prediction of population
dependent variable (e.g., number of animals harvested, size through time and can be used to infer if the population
number of waterfowl surveyed) to index population size. is stable (i.e., the trend line has a slope = 0), declining
Measures of dependent variables over time are used as (negative slope), or increasing (positive slope). For
inputs to trend models and population changes (trends) example, McCullough131 used univariate regression to
over time are estimated. Trend models portray average determine the relationship between adult population size
rates of population change, require few assumptions, and and recruitment of young for grizzly bears. Using the
can incorporate data that are readily available from magnitude of the regression coefficients and the regression
standard wildlife surveys. Pascual et al.144 evaluated three slope, McCullough131 argued that young grizzly bear
 CHAPTER 21: MODELING—INTRODUCTIONS 523

Table 2. A coarse comparison of population modeling approaches.

Each model was ranked as unfavorable (U), moderately favorable (MF), or favorable (F),
according to the practical application of the modeling approach.
Rankings are subjective evaluations as to the historic use, performance, and perception of the models.

Model Evaluation Criteria

Applicability Model Structure Model Output

Appeal Common Cost of Cost of Training Level of Complexity Generality Face Communicability
data data model required subjectivity validity
acquisition develop-
ment
Single species
Trend analysis
Univariate regression F F F F F MF F F F F
Multivariate techniques MF F F F F U U F MF U
Age/Stage based
Life tables or matrixes F MF MF F F F F F F F
Multiple species U MF U MF U U U U MF F
a
Some criteria adapted from Marcot et al. where Appeal = How much the model framework has been used in the past and
124

practically applied; Common data = Data that are collected as part of routine inventories (e.g., rimber cruises, game check stations,
remote sensing); Cost of data acquisition = Cost to acquire the data for model development and use; Cost of model development = Cost
(monetary and time) of developing the model; Training required = Amount of training required to use and understand the model; Level
of subjectivity = Degree that subjective evaluations are used to develop the model relationships; Complexity = Complexity of the
modeling framework, synonymous with Marcot et al.’s “wholeness”; Generality = How well the model represents a broad range of
similar systems; Face validity = Perceived credibility of a modeling framework; Communicability = Ability to associate model output to
real world systems.

recruitment into populations was highly dependent on contest competition) and intensities of intra-specific
adult population size. Univariate regression is one of the competition.
most practical approaches to population modeling (Table Multivariate techniques to population modeling often
2). use commonly collected data, can be developed for low
costs, and require minimal training to apply (Table 2).
Multivariate Techniques. To further account for variance
These models also can be applied to a broad range of
in population rates of change from trend data, it is possible
similar systems (Table 2). Drawbacks of these models
to examine the contribution of additional environmental
include their level of subjectivity and complexity (Table
variables that may regulate the population. Additional
2). Outputs can also be difficult to communicate (Table 2).
independent variables (e.g. weather, harvest, predation)
can be input to construct a multiple regression model that Age/Stage-Based Models
expresses population growth as a function of several If structure or composition of a population is of interest,
independent variables including multiple time lags. Time then age- or stage-based models are usually employed.
lags are a means of capturing the effects of independent These models contain elements that correspond to data
variables that appear after some period (lag) of time.164 collected on individuals that are extrapolated to the
For example, a summer drought might not immediately population using matrix algebra.86 Matrix elements can
influence adult mortality, but rather, the effects of the include survival of individuals, probability of transition
drought may be expressed 10 months later in the number from one stage to the next, or fecundity.201 Most life tables
of offspring produced by females. or matrix models focus on life cycle components with high
Caution should be exercised when projecting future potential impacts on population growth and how these
population size based on historic trends. This is akin to components interact.201
predicting the dependent variable outside the range of the Leslie 113 introduced matrix methods to wildlife
independent variable (time). Moreover, relationships biologists in the mid-1940s. Textbooks by Caswell35 and
between population growth rate and time are not cause Burgman et al.28 advanced matrix theory in biology and
and effect. That is, simple trend analysis does not indicate the tools are widely used for population projections on a
if populations are regulated in a predictable fashion, multitude of species. Life tables and matrix models were
perhaps by intrinsic (density-dependent) factors.14 One of developed from matrix theory and they represent age- or
the most commonly used hypotheses to detect density- stage-specific summaries of mortality or birth rates that
dependent population regulation is based on the logistic operate on populations. These tables have been presented
model.51, 163, 167 The logistic model often takes the form of in several formats (e.g., Leslie matrix) and often use a
the familiar S-shaped curve. The logistic model can specialized notation (reviewed in Krebs).104: 413 The original
incorporate a wide variety of density-dependent effects work of Leslie113 laid the foundation for age-based analysis,
that can imitate different forms (e.g., scramble competition, whereby individuals in a population are projected through
 524 Wildlife–Habitat Relationships in Oregon and Washington

different age classes. Lefkovitch112 recognized that Leslie’s killer whales, 201 semipalmated sandpipers, 86 spotted
work was a specialized version of a more general matrix. owls,29, 110 marbled murrelets,10 loggerhead sea turtles,43
He noted that Leslie’s113 matrix ideas could be used to and fur seals,58, 68 to name a few. These models are some
project the abundance of individuals in a particular life of the most widely used and practical of the population
stage at a certain time and he established the foundation modeling tools (Table 2).
for stage-based analysis. In stage-based analysis, the
Multiple Species Models. Often, a key question in wildlife
assumption is made that all individuals in a particular
population management relates to the role that predation
stage (e.g., body size or development classes) exhibit the
plays in creating prolonged prey suppression. 134 To
same mortality, fecundity, and growth schedules,
determine if predation is causing low prey density,
regardless of age. Stage-based models are useful when the
researchers must quantify the response of predators to
age of individual organisms cannot be exactly determined,
changing prey density.134 A diversity of models exist that
but when individuals can be classified according to their
describe population dynamics in communities with
developmental stage (e.g., subadults-adults, larvae-pupae-
interacting species.129 Two-species interaction models are
adults). For example, van Tienderen201 identified four life
created when the dynamics of two interacting populations
cycle stages (yearlings, juveniles, reproductive females,
(i.e., predator-prey, herbivore-plant, competition for a
and post-reproductive females) regulated by three
resource) are of interest.
probabilities (the probability of going to the next life stage,
Generally, the mathematics of multiple species
the probability of remaining in the current life stage, and
population models can be grouped into those that project
fecundity) in his matrix population projection of the
continuous population growth rates (i.e., use differential
demography of killer whales. Van Tienderen 201
equations) and those that project discrete population
demonstrated the importance of correlation among inputs
growth rates (i.e., use difference equations). Differential
on the population projection. These considerations become
models are used with wildlife species that exhibit
especially important when matrix models are used to
continuous reproduction, i.e., there is no “break” in the
project populations subjected to alternative management
potential for population growth. In contrast, difference-
activities, particularly if the management activity cascades
equation models (originally developed by entomologists)
through the life table because of interrelationships in the
have application to wildlife species that exhibit notable
demographic parameters.
generation breaks in the birth process (e.g., salmon,
Stage-based modeling can also be performed on
insects). Generally, difference equations tend to be less
habitat-structured populations. 20 Since animals that
stable than the corresponding differential equations
occupy habitats of different quality will survive and
because the time lapse between generations of growth in
reproduce at different rates (especially in the absence of
the difference models has the destabilizing effects
density dependent effects), it is logical to consider that a
associated with any time lag in an interactive system.129
more preferable approach to life table or matrix modeling
These models are not simply restricted to the growth
is to structure populations according to habitat quality.
equations for predator and prey, other environmental
Bowers 20 compared age-structured versus habitat-
influences can also be integrated (e.g., habitat quality).134
structured population projections using computer
Multi-species trend models are often based on the
simulation. He demonstrated that the demography of a
logistic growth model, where the population growth rate
habitat-structured population could change fitness by
of a species is not only a function of its own density, but
changing habitats. In age-structured populations fitness
also is influenced by the density of a second species. The
could only be changed through aging. Realistically, a
most well-known multi-species trend models are the
combination of age- or stage- and habitat-based
predator-prey models (more commonly called Lotka-
approaches probably provides the best model. Some
Volterra models) which estimate the demographics of two
modelers have taken this advice and integrated habitat
species in a predator and prey relationship. These models
and demographics into population viability analyses.2, 161
predict prey recruitment, prey mortality, predator
Age- and stage- models can employ key-factor analysis,
recruitment, and predator mortality as functions of each
which is often used to compare the effects of different
other.69 By using ratio-dependent interspecific effects (i.e.,
survival or fecundity rates among the different classes on
the ratio of the predator per prey) instead of a pure
population dynamics.199 In other words, by using key-
predator density effect the realism of the model is
factor analysis, age- and stage-models can be used to
increased.13 This approach is analogous to incorporating
determine the age or stage at which conservation actions
extrinsic covariates into the logistic model (see above). This
have the greatest effect on population size.44 This is often
generates an interesting conclusion, that carrying capacity
useful for evaluating the effects of density-dependence
is not a fixed entity, but depends on the covariates. In this
and environmental variation on population growth rates.24,
176, 199 case, the covariate is the ratio of predator to prey. Such
The key-factor analysis is data intensive in that it
covariates mediate and alter the strength of the density-
requires age- or stage-specific mortality and fecundity
dependent affects on carrying capacity. Examination of
data.14
covariate affects can greatly increase our understanding
Life table and matrix models have been used to project
of complex systems and enhance our ability to manage
populations for a wide array of wildlife species including
these systems successfully.103
 CHAPTER 21: MODELING—INTRODUCTIONS 525

Age- or stage-structured multi-species models are Space complicates population models in three
rarely constructed in wildlife management due to their fundamental ways: (1) non-uniform patterns of habitat
extensive data requirements (i.e., age- or stage-specific quality, (2) non-uniform patterns of animal density, and
demographic rates for predators and prey; Table 2), but (3) animal movements from one location to another.114
they occasionally find application in the field of These complications can be modeled with various degrees
entomology and pest management. Age-structured multi- of realism, and models can be classified accordingly—
species models can be implemented as linked Leslie- spatially implicit, spatially explicit, or spatially realistic
models.73 In these models, a realistic representation of the (Figure 3). The taxonomy of spatial models can be further
order of biological process (i.e., the sequence of events refined by classifying models according to their population
within the population) is of critical importance. structure, i.e., either as single population or
metapopulation models, and according to their
Spatial population modeling formulation—analytical, matrix, or individual-based
Widespread recognition of space as an important (Figure 3). Analytical models occur in the form of an
consideration for both conservation theory and equation or equations. Classic examples of analytical
management practice can be traced to the equilibrium models are the logistic and Lotka-Volterra population
theory of island biogeography. 56, 177 MacArthur and models. The appeal of analytical models stems from their
Wilson121 proposed that a dynamic equilibrium between mathematical elegance, basis in ecological principles, and
extinction and immigration could explain species richness tractability, either through exact solutions or numerical
on oceanic islands, and that species richness was a function methods. Most analytical models describe the dynamics
of an island’s area and distance from the mainland. An of idealized populations (e.g., closed-population, constant
analogy between oceanic islands and patches of habitat carrying capacity), hence their importance lies in
surrounded by inhospitable areas has been used to contributing to conservation theory which in turn forms
advance concepts in wildlife habitat management.52, 70, 188 the foundation of management practice. While the
Levins115, 116 is credited with establishing the contemporary generality of analytical models makes them less useful for
foundation for the study of spatially structured wildlife quantitative answers to specific management questions,
populations. His metapopulation concept pertains to a these models can provide qualitative insights.
collection of interacting populations that occur in a distinct The formulation of matrix models was previously
habitat patch separated from other patches by non-habitat. discussed in the “Population Modeling” section of this
The number of populations that occur within the chapter. In that section, we noted that the dimensions of
metapopulation (or the number of occupied habitat non-spatial matrix models were determined by the
patches) is determined by the rates of population number of population classes. The dimensions of spatial
extinction and patch colonization. There are obvious matrix models are determined by both the number of
similarities between island biogeography and population classes and types of habitat patches.
metapopulations, but for a number of reasons, the latter Individual-based models track all the individuals in a
has supplanted the former as a dominant paradigm in population; hence they can incorporate local interactions
conservation biology.78 Hanski and Gilpin,77 Hastings and between organisms and habitat.92, 98 Every individual in
Harrison, 83 and Hanski and Simberloff 78 provide the simulated population is represented numerically, and
explanations of metapopulation theory, its history, and data on every individual is “observed” and tabulated for
models. birth, death, reproduction, location, and movement. The
While the theoretical import of the metapopulation behavior and fate of individuals are stochastic. That is,
concept is undeniable, the practical application of the actions or events occur randomly according to probability
concept is limited, particularly with respect to large or distributions that are either empirically estimated from
mobile animals for which demographic isolation is less the real population or based on plausible assumptions.
likely. Rarely have managers faced the problem of Rules direct the reproductive, foraging, social, and
managing a “population of populations”, which was movement behaviors of individuals. The collective success
Levins’s 116 original definition for a metapopulation. or failure of all individuals determines the fate of the
Managers usually deal with a single population or a sub- population. In general, individual-based models are not
population. Members of that population will distribute mathematically sophisticated, but they are
themselves across space according to the spatial computationally complex. Hence, well-honed computer
distribution of habitats. This pattern of habitat across programming skills are indispensable for tackling the task
landscapes may have profound affects on population of model development. There are no standards for model
viability (i.e., the population’s survival). “Habitat structure, but modular, top-down, object-oriented
fragmentation” has become a well-known cause célèbre for programming is highly recommended.
conservation biologists, and many of their concerns are Figure 4 contains examples of different analytical,
well-justified.169 This is an important motivation behind matrix, and individual-based spatial population models.
many of the models used for investigating the relationship Figure 4(a) is an analytical spatially implicit model.109, 110
between the spatial pattern of habitats and population The equation represents the territory dynamics of a
dynamics across landscapes. population containing only females at demographic
equilibrium. Figure 4(b) is a spatially implicit model
 526 Wildlife–Habitat Relationships in Oregon and Washington

Figure 3. Decision tree for selecting a wildlife spatial

population modeling approach.
with environmental variability, describing dispersal, and
portraying dynamics within patches, together combine to
formulated as a matrix. The matrix represents movement yield an assortment of models that is overwhelming in its
among four habitat patches where F1i is the fecundity of variety.”
individuals in age class 1 and patch i; P1i is the probability
Spatially Explicit Models. Spatially explicit models are
of individuals in age class 1 surviving and staying in patch
those that “keep track of the exact positions of plants and
i; and d1ij is the probability of individuals in age class 1
animals”100 or “have a structure that specifies the location
surviving and dispersing to patch j from patch i (Figure
of each object of interest”.57 Hanski and Simberloff78 regard
4b). Figure 4(c) is the stylized landscape model developed
the key feature distinguishing spatially explicit models
by Lamberson et al.106 Each circle contains a territory
from spatially implicit models to be localized interactions,
cluster and arrows depict the dispersal paths between
i.e., the population dynamics within a patch are influenced
clusters. It is assumed that each cluster is identical and
only by the state of nearby patches in explicit models. A
contains the same number of territories. Movement within
wide variety of modeling approaches are considered
a cluster is governed by an equation similar to that
spatially explicit including reaction-diffusion models,
presented in Figure 4(a). Figure 4(d) is an example of a
coupled lattice models, cellular automata models, stepping
spatially realistic model. The input to the model is Landsat
stone models, and spatially realistic models (Figure 3).78
Thematic Mapper imagery classified into four forest cover
These approaches can be applied to single populations or
categories (left); a non-habitat class and three levels of
metapopulations.
habitat quality. A hexagonal grid was superimposed on
the image and each hexagon was assigned a territory Reaction-Diffusion Models and Coupled-Lattice
quality based on its habitat composition. This type of Models. Reaction-diffusion models are based on the
model was a two-sex, individual-based model.208 mathematics used to describe the kinetics of reacting and
The classification system for spatial population diffusing chemicals. 9, 114 In population modeling,
modeling we propose here should impart some order on “reaction” refers to within-patch population dynamics and
a rapidly evolving branch of population ecology, but it “diffusion” refers to the dispersal of individuals between
undoubtedly fails to convey the variety of spatial models patches. A reaction-diffusion model consists of a set of
that exist today. The problem we faced was well expressed partial differential equations (Figure 3) hence they are
by Kareiva 99 who wrote, “The different ways of continuous in both time and space. Each equation
representing space, keeping track of populations, dealing represents a patch’s population density and consists of
 CHAPTER 21: MODELING—INTRODUCTIONS 527

Figure 4. Examples of
spatial population
modeling approaches.

three terms: (1) a function describing within-patch of each cell in an automaton model is a discrete variable
population dynamics, (2) a function describing net (Figure 3). For example, Caswell and Etter36 describe a two-
exchange with other patches, and (3) a function describing species cellular automaton in which each cell could possess
net exchange with the region surrounding a patch.114 To one of four states: species S1 present, species S2 present,
date, the use of reaction-diffusion models has produced both species present, or neither species present. Discrete
mostly theoretical results (Table 3); practical applications states are ideal for many metapopulation analyses where
have dealt mainly with invasions by pests or the spread patches may be either occupied or unoccupied. The state
of disease.99 of each cell is determined by a set of rules that account for
Coupled lattice, cellular automata, and stepping stone the current state of the cell and the states of surrounding
models all arrange habitat patches, or local populations, cells.9, 45 The rules may be mathematical equations or
into a regular two-dimensional array of cells. Coupled logical operations or some combination of both. Cellular
lattice models are basically a discrete time, discrete space automata are difficult to solve analytically, so they are often
simplification of reaction-diffusion models;9 hence they implemented as computer simulations.36 We are unaware
are analytical models that take the form of difference of any applications of cellular automata models to address
equations. Each difference equation represents population real wildlife management issues and their practical
density within a particular cell. Cells within the lattice are application is limited (Table 3).
“coupled” through diffusion processes that represent
Stepping Stone Models. Because of their conceptual
animal dispersal. Like reaction-diffusion models, the
simplicity, stepping stone models have proven to be the
results from coupled lattice models have been theoretical
most practical approach for incorporating spatial structure
(Table 3). They are becoming well known for portraying
into population models. Stepping stones, or patches of
complex dynamics and chaotic behaviors that evoke
habitat, are simply arranged in a regular grid. Patch
intriguing speculations about real ecosystems.9
dependent demographics or local population dynamics
Reaction-diffusion models tend to have low appeal
are defined by a set of rules (Figure 3). Similar to cellular
because these models have, in general, been confined to
automata models, these rules can consist of mathematical
the realm of theory (Table 3). Costs and complexity of these
equations or logical operations or some combination of
models can also limit their practical application (Table 3).
both. Most stepping stone models simplify the space
The theoretical foundation and mathematical rigor
between patches by assuming it consists of entirely
associated with reaction-diffusion models reduce
uniform non-habitat. Stepping stone models played an
subjectivity, increase generality, and increase perceived
important role in analyses of federal management plans
validity
for spotted owl habitat.192 Like Doak,55 Thomas et al192 (and
Cellular Automata Models. A cellular automaton consists see Lamberson et al.106) modeled owl habitat as territory
of a regular array of interacting cells. Unlike a coupled clusters, but they added explicit spatial structure by
lattice model that describes the state of each cell by a arranging clusters in a rectangular array. Territories within
continuous variable (e.g., population density), the state clusters were all equally accessible, but movement from
 528 Wildlife–Habitat Relationships in Oregon and Washington

Table 3. A coarse comparison of spatial population modeling approaches. Spatial population models can be
applied to both individual populations or metapopulations. Each model was ranked as unfavorable (U),
moderately favorable (MF), or favorable (F), according to the practical application of the modeling approach.
Rankings are subjective evaluations as to the historic use, performance, and perception of the models.

Model Evaluation Criteria

Applicability Model Structure Model Output

Appeal Common Cost of Cost of Training Level of Complexity Generality Face Communicability
data data model required subjectivity validity
acquisition develop-
ment
Spatially explicit models
Reaction-diffusion models U U MF MF U F U F F MF
Coupled lattice models U U MF MF U F U F F MF
Cellular automata U U MF MF U MF MF F F MF
Stepping stone models F U MF MF MF U MF U MF F
Spatially realistic models MF U U U U U U U MF U
Spatially implicit models
Analytical models F MF MF F F F MF F F F
Matrix models F U MF F F F F MF F F
Individual-based models MF U MF MF MF U U U MF MF
a
Some criteria adapted from Marcot et al.124 where Appeal = How much the model framework has been used in the past and
practically applied; Common data = Data that are collected as part of routine inventories (e.g., rimber cruises, game check stations,
remote sensing); Cost of data acquisition = Cost to acquire the data for model development and use; Cost of model development = Cost
(monetary and time) of developing the model; Training required = Amount of training required to use and understand the model; Level
of subjectivity = Degree that subjective evaluations are used to develop the model relationships; Complexity = Complexity of the
modeling framework, synonymous with Marcot et al.’s “wholeness”; Generality = How well the model represents a broad range of
similar systems; Face validity = Perceived credibility of a modeling framework; Communicability = Ability to associate model output to
real world systems.

cluster to cluster was limited to adjacent clusters. Dispersal of powerful new technologies for the collection and
between clusters was more hazardous than dispersal analysis of spatial data, e.g., satellite imaging, global
within clusters because regions between clusters contained positioning systems, improved microelectronics for radio-
non-habitat. This model was an individual-based model. telemetry, and geographic information systems, have
A stepping stone model may also be implemented as a provided the wherewithal to conduct large-scale empirical
transition matrix.35 Spatial structure can be incorporated studies.
into a matrix model through elements that represent the The first published results obtained from applying a
probability of successful dispersal between patches.211 spatially realistic model to a wildlife population are
Demographic parameters would be patch-dependent, i.e., attributed to Fahrig and Paloheimo.62 They analyzed the
different vital rates can be assigned to patches according effects of habitat patch arrangement on the population
to habitat quality. Stepping stone models have high appeal density of cabbage butterflies (Pieris rapae) in Ontario.
as a compromise between the complexity of spatially Since then, interest in such models has continually
realistic models (see below) and the mathematical rigor increased. Spatially realistic models have been used to
of reaction-diffusion and cellular automata models (Table analyze the effects of habitat change on forest-interior birds
3). in Wisconsin;187 Bachman’s sparrow (Aimophila aestivalis)
in the southeastern United States;118, 150 an endangered
Spatially Realistic Models. The stylized landscapes of the
possum in Australia; 117 the endangered helmeted
aforementioned models have been eclipsed by the
honeyeater (Lichenostomus melanops cassidix) in Australia;2
empirically derived landscapes of spatially realistic
an endangered butterfly in Finland;205 the California
models. Spatially realistic models use habitat data from
gnatcatcher3 (Polioptila c. californica); the California spotted
actual landscapes that are tied to real-world spatial
owl105, 140 (S. o. occidentalis); and northern spotted owl.88, 91,
coordinates (Figure 3), and these habitat data can be 133, 208
manipulated to represent habitat conditions likely to result
Hanski76 developed an analytical spatially realistic
from management. Spatially realistic population models
metapopulation model. This model, which she called an
arose from a fortuitous convergence of theory and
incidence function, could accurately model the patch
technology. A critical mass of research in landscape
turnover rate and minimum patch sizes of several butterfly
ecology, albeit predominately theoretical, has been
species, and was described as a practical tool for
attained during the past 15 years.66, 81, 193, 195 This theory has
management problems.
led wildlife managers to recognize that the spatial
Hanski’s 76 spatially realistic models are typically
arrangement and quality of habitat can have a significant
stochastic and individual-based. For many applications,
impact on population viability.157 Concurrently, an array
 CHAPTER 21: MODELING—INTRODUCTIONS 529

spatially realistic individual-based models will require metapopulation concepts to model the effects of habitat
large computer memory and huge on-line data storage loss on a single population of a territorial species and he
because: (1) information on every living individual in the used this analytical model to examine management of the
population is maintained during a simulation, and (2) northern spotted owl.110 An analytical spatially implicit
geographic information system data, particularly population model was also used to explore the concept of
rasterized data, can occupy large amounts of hard-drive “source” and “sink” habitats by Pulliam,148 who claimed
space. High computer processing speeds are also desirable. that the model could be useful for natural resource
Data required by spatially explicit models fall into three management problems.149
separate categories: (1) habitat data, (2) demographic data, Lamberson et al.106 (and see Thomas et al.,192 Appendix
(3) and movement data. Habitat maps may be generated M) developed a spatially implicit model to analyze the
through remote sensing methods (e.g,. aerial photography effects of habitat fragmentation on the northern spotted
or satellite imaging), on-the-ground habitat surveys, owl. All suitable territories were equally accessible to
habitat-based modeling, or most likely some combination dispersing owls, but the probability of successfully finding
of both. The most important demographic data are a territory was a function of: (1) the proportion of
estimates of survivorship and fecundity. Ideally, territories unoccupied, and (2) a search efficiency
demographic data would also provide information on the parameter, which was assigned different values for
non-breeding portion of the population, such as the juveniles and adults. Matrix models can also incorporate
fraction of the population that is not breeding. Both implicit spatial structure.20 Unlike typical matrix models
demographic data and movement data should be collected that divide populations into age classes, Bowers20 divided
such that parameter estimates can be related to habitat a population into habitat-type classes, according to the
quality. habitat types in which individuals bred.
Generally, agreement exists that spatially explicit Spatially implicit models can possess a hierarchical
models have utility in land management decisions.57, 100, structure. Doak55 represented habitat as clusters of spotted
194
However, concerns have been expressed about model owl territories that imparted a two-level hierarchy to its
reliability.7, 41, 56, 165, 211 These concerns stem from two spatial structure. At the lower level, all territories within
fundamental problems with spatially explicit models. a cluster were equally accessible to owls in that cluster,
First, such models require data that are often difficult to but other clusters were much less accessible. At the higher
collect, in particular data relating demographics to habitat level, all clusters were equally accessible to all other
quality and data on movements (Table 3). These data clusters. In other words, the relative locations of clusters
requirements are expensive (Table 3). Therefore, such were ignored, and within clusters the relative locations of
models will be feasible only for species that can garner territories were ignored. His model was an individual-
substantial commitments to data collection. Second, based model—each year the location and age of each owl
because of the large spatial and long temporal scales to was known. Carroll and Lamberson 33 developed a
which the models are typically applied, there is no theoretical analytical model that also had a hierarchical
possibility of validating the model. In other words, spatial structure.
validation is impossible because true validation requires Among spatially implicit models, individual-based
measuring population dynamics and landscape patterns formulations scored the most “unfavorable” (Table 3). This
that are replicated and randomized. These concerns can can be explained, in part, by their relative newness.
be assuaged by intelligent model design, thorough model Practitioners have some exposure to analytical and matrix
evaluation, appropriate model application, and prudent formulations of population models, but individual-based
interpretation of model outputs.11, 26, 41, 183 models are still foreign. Therefore, individual-based
models score “unfavorable” for appeal, training required,
Spatially Implicit Models. Population models can
perceived validity, and communicability (Table 3).
incorporate aspects of space but not possess spatial
structure. For example, the classic logistic model of
population dynamics implicitly incorporates the amount Wildlife Modeling: A Case Study
of habitat available to a population through density Implementation of a modeling approach to address a
dependence. The logistic model, however, does not wildlife management issue involves an explicit statement
address the spatial distribution of habitats, and hence, it of the issue and a review of relevant information. The issue
is a spatially implicit model. Spatially implicit models can be characterized in terms of how output will be used,
assume that habitat is spatially structured, but simplify risks associated with producing imprecise or biased model
the structure by ignoring the geometry or relative locations output, and data availability or resources available to
of habitat (Figure 3). Typically, spatially implicit models collect data. In addition, scale is important and includes
assume that habitats are arranged as discrete patches and such things as the plan’s time frame, planning landscape
that each patch is equally connected to all other patches.78 extent, and species involved. More often that not, the “best
The original metapopulation model of Levins115, 116 was an model approach” is not selected because limited time and
analytical spatially implicit model. resources (i.e., data and knowledge of the system)
Hastings 82 developed a more detailed analytical constrain the scope and detail of the modeling effort.
metapopulation model that incorporated the local In this section, we demonstrate the process of selecting
population dynamics of each patch. Lande 109 used and building a model to address a real resource
 530 Wildlife–Habitat Relationships in Oregon and Washington

management issue. We frame the management issue, relationships between lynx and habitat existed for this
review the information pertinent to addressing the issue, area, the time frames were too short to initiate a large data
select a modeling approach using the decision trees collection effort. Thus, the project was single species
presented in figures 1-3, and discuss how the model output focused, supported by vegetation inventories and a
was used. geographic information system, and needed to be
completed in a short time frame. A theoretical habitat
Frame the Management Issue. The State of Washington
modeling approach was selected (Figure 1).
designated lynx as a threatened species in 1993. As part
of the listing, state agencies are responsible for establishing Select the Modeling Approach. Several single species
forest practice rules that protect lynx or its critical habitat. habitat models could have been used for this project
State agencies have jurisdiction over state-owned and (Figure 1). The rarity of lynx in the planning area and lack
large, privately owned lands. In lieu of forest practice rules of occurrence data precluded assigning probabilities of
for lynx, landowners within critical habitat areas and the occupancy by habitat patch. Thus, the landowners did not
state agencies agreed to develop and implement select Bayesian or pattern recognition models. Since the
conservation plans to protect lynx and their habitat. The amount of habitat was deemed important, they chose to
process was appealing to landowners because it provided index the contribution of habitat patches towards lynx
an opportunity to develop conservation strategies conservation using the habitat evaluation procedures
consistent with their landscape’s ecology and compatible (Figure 1). Part of the allure of the habitat evaluation
with their land management objectives and information procedures was the notion that smaller patches of high
systems. The conservation plans were subject to scientific quality habitat could function similar to larger patches of
review and approval by Washington State’s Forest marginal quality habitat.157 This tradeoff was attractive to
Practices Board. the landowner because it allowed for operational
The first step in developing the plan was for each flexibility. Another appealing characteristic of the habitat
landowner to state their goal and objectives. The goal was evaluation procedures was that they could be used to
to craft a scientifically credible lynx conservation plan that assess habitat at scales ranging from a single forest stand
could be integrated into the landowner’s long-term to a landscape composed of hundreds of stands. 157
resource management plan. For one landowner, objectives Spatially explicit output at the stand-level was appealing
to achieve this goal included: in that it permitted the evaluation of individual
• Create a lynx conservation plan that was compatible silvicultural prescriptions that linked to the operational
with the landowner’s resource information and harvest plans. Similarly, a spatially explicit, multi-stand
forest growth models. view of habitat was also important to address lynx
• Maintain or enhance lynx habitat condition over the movement requirements and the effects of harvest patterns
life of the plan. on within-home range habitat conditions. Finally, output
• Monitor the lynx population in compliance with the from the habitat evaluation procedures offered a currency
State’s recovery plan. that foresters, agency biologists, and the public could
• Maintain a reasonable flow of timber from the land. readily understand.
• Use existing inventory and mapping systems
How the Model Output Was Used. The final conservation
whenever possible.
plan consisted of a 5-year harvest schedule that outlined
Review the Information Pertinent to the Management the stands to be managed, silvicultural prescriptions, post-
Issue. Based on this, the landowners decided that harvest site preparation, and lynx mitigation measures
modeling was an appropriate tool to assist them in (Figure 5). The short-term effects of these activities on lynx
addressing the management issue. The next step in the habitat quality were assessed (Figure 5). In addition to
modeling process was to determine which model type (i.e., the spatially explicit harvest plan and lynx habitat
habitat-based, population-based, or spatial population assessment, a 25-year trend in lynx habitat condition was
modeling) was most suited to the situation and to decide estimated using a wildlife decision support system that
on a specific modeling approach (i.e., use the decision trees consisted of a tree growth and yield model, forest planning
in figures 1-3 to select a model). To make these decisions, model, and a spatial disaggregation tool.158 In this decision
information and data pertinent to the management issue support system, the lynx habitat model served several
were reviewed. Planning capabilities influenced the purposes. It provided the landowner a means to
landowner’s ability to generate a conservation plan. The quantitatively assess habitat in a repeatable, timely
landowner had a timber-based land classification scheme manner. The landowner used this information to evaluate
coarsely grouped by major plant association type, 46 different silvicultural prescriptions (e.g., tree spacing in
overstory tree species, forest structure, and vegetation thinning operations); to provide home-range-level
density. This scheme was catalogued in an extensive stand assessments that permitted an analysis of habitat
inventory database and mapped to a resolution of 5 acres configuration; to position human-made den sites, travel
(2 ha) in a geographic information system. The geographic corridors, and road closures; and to organize management
information system also contained transportation and activities in time and space.
hydrology map layers. Since the conservation plan needed The model and its output were easily communicable
to be completed in one year, and no empirical data on the to state biologists and regulators and the landowner’s
 CHAPTER 21: MODELING—INTRODUCTIONS 531

Figure 5. Components of a lynx conservation

plan that was developed using a wildlife
decision support system that included a tree
growth and yield model, forest planning
model, spatial disaggregation tool, and a
habitat evaluation procedures model for
lynx.

foresters and management. Considering the multiple Future Directions of Wildlife Modeling
stakeholders associated with this project (stage agencies, More and more, wildlife managers are being asked to
Forest Practices Board, landowner management) this apply their expertise toward holistic, often intangible,
characteristic of the model was critical. Also, because the management goals such as “sustainability,” “biodiversity,”
model was habitat-based, the model and its output were and “ecological integrity.” To be effective, wildlife
easily translated to on-the-ground management practices. managers cannot work within the confines of conventional
The model was run on map layers that corresponded to wildlife biology. Rather, managers must also be broadly
operational stand boundaries thereby permitting a trained and versed in statistics, ecology, law, sociology,
straightforward crosswalk to the forest plan. The habitat and economics. Managers are also expected to adapt to
evaluation procedures used in this project are also changing circumstances brought about by dynamic
conducive to adaptive improvements that may result from budgets, new information, or changes to regulations or
a more refined lynx habitat model, modifications in how public opinion. These new expectations will shape the
the landowner surveys and maps their forests, or from an future directions of wildlife modeling as a management
expanded view of habitat across multiple land tool.
ownerships. Modeling will play a key role in adaptive management.
Here, wildlife modeling provided a tool for addressing Adaptive management is a process that uses feedback for
regulatory issues on private lands management. A improving the management process. Models can fit neatly
defensible, meaningful, and practical plan was developed into this process in that they help inform management
and accepted by the Washington Forest Practices Board decisions. The adaptive management process was
in 1996. The landowners have integrated lynx habitat originally invented for fisheries based on control system
conservation into their resource planning process and they theory.179, 206, 207 In practice, a model of the fisheries system
continue to collect data and refine the relationships was developed and initial management decisions would
contained in the wildlife model. be based on information derived from the model.
 532 Wildlife–Habitat Relationships in Oregon and Washington

Management actions were viewed as perturbations to the Literature Cited

fishery and by measuring the response of the system to 1. Airola, D. A. 1988. A guide to the California wildlife habitat relationship
these perturbations data were acquired to improve the system. California Department of Fish and Game, Sacramento, CA.
model. The adaptive management process would use the 2. Akçakaya, R. H., M. A. McCarthy, and J. L. Pearce. 1995. Linking landscape
new data to revise the model and parameter estimates with data with population viability analysis: management options for the
helmeted honeyeater. Biological Conservation 73:169-176.
the intent of improving the information for management 3. ———, and J. L. Atwood. 1997. A habitat-based metapopulation model
decisions. This cycle continues indefinitely. of the California gnatcatcher. Conservation Biology 11:422-434.
Adaptive management is complex. The information 4. Andrewartha, H. G. 1971. Introduction to the study of animal
feedback mechanisms for wildlife systems may be direct populations. University of Chicago, Chicago, IL.
(as in the case of animal harvest), but more often indirect 5. Bailey, J. A. 1984. Principles of wildlife management. John Wiley & Sons,
(as is the case with habitat manipulation). For models New York, NY.
6. Bart, J., D. R. Petit, and G. Linscombe. 1984. Field evaluation of two
designed to predict indirect responses of wildlife, the link
models developed following the habitat evaluation procedures.
between management action and system response is subtle Transactions of the North American Wildlife and Natural Resources
and hence, inferences from the model may be weak. Also, Conference 49:489-499.
wildlife managers often deal with management issues that 7. ———. 1995. Acceptance criteria for using individual-based models to
range from decades to centuries in length. Thus, feedback make management decisions. Ecological Applications 5:411-420.
for model improvement may be delayed beyond the life 8. Bartlett, M. S. 1960. Stochastic population models in ecology and
of the management issue. To alleviate this time lag in epidemiology. Wiley & Sons, New York, NY.
9. Bascompte, J., and R. V. Solé. 1995. Rethinking complexity: modelling
receiving feedback, modelers have turned to iterative
spatiotemporal dynamics in ecology. Trends in Ecology and Evolution
modeling approaches that attempt to account for the 10:361-366.
different combinations of environmental and population 10. Beissinger, S. R. 1995. Population trends of the marbled murrelet
events. Iterative or simulation models tend to mimic projected from demographic analyses. Pages 385-393 in C. J. Ralph, G. L.
stochastic events and provide insight to the sensitivity and Hunt, Jr., M. G. Raphael, and J. F. Piatt, editors. Ecology and conservation
breadth of model output to the range of inputs. For of the marbled murrelet. United States Forest Service General
Technical Report PSW-GTR-152.
example, iterative modeling is commonly used in
11. ———, and M. I. Westphal. 1998. On the use of demographic models
extinction models65 to understand minimum population of population viability in endangered species management. Journal of
numbers required for population persistence under a Wildlife Management 62:821-841.
range of environmental stochasticities. Ruckelshaus et al.165 12. Benson, G. L., and W. F. Laudenslayer, Jr. 1986. DYNAST: Simulating
used iterative modeling to test the sensitivity of spatially wildlife responses to forest management strategies. Pages 351-356 in J.
explicit population models to parameter error. Bowers20 Verner, M. L. Morrison, and C. J. Ralph, editors. Wildlife 2000: modeling
used simulation modeling to evaluate the dynamics of age- habitat relationships of terrestrial vertebrates. University of Wisconsin,
Madison, WI.
and habitat-structured populations. These are just a few
13. Berryman, A. A. 1990. POPSYS Series 2 - two-species analysis. User
examples of the use of simulation modeling in an adaptive manual, Ecological Systems Analysis, Pullman, WA.
improvement mode. These simulations provided feedback 14. ———. 1994. Population dynamics: forecasting and diagnosis from
that facilitated the understanding of system dynamics and time-series. Pages 119-128 in S. R. Leather, A. D. Watt, N. Mills, and K. F.
model behaviors. A. Walters, editors. Individuals, populations and patterns in ecology.
Clearly modeling will remain an important part of Intercept, Andover, England.
wildlife management and undoubtedly a component of a 15. Block, W. M., L. A. Brennan, and R. J. Gutiérrez. 1986. The use of guilds
and guild-indicator species for assessing habitat suitability. Pages 109-
good decision-making process. As wildlife practitioners,
113 in J.Verner, M. L. Morrison, and C. J. Ralph, editors. Wildlife 2000:
we often become preoccupied with rejecting specific modeling habitat relationships of terrestrial vertebrates. University of
models or questioning the assumptions of a particular Wisconsin, Madison, WI.
modeling approach when we should be evaluating 16. ———, M. L. Morrison, J. Verner, and P. N. Manley. 1994. Assessing
whether or not a model allows us to make a more informed wildlife-habitat-relationships models: a case study with California oak
decision.165 This is not to say that models should be taken woodlands. Wildlife Society Bulletin 22:549-561.
at face value. It is critical for model users to understand 17. Boertje, R. D. 1985. An energy model for adult female caribou of the
Denali herd, Alaska. Journal of Range Management 38:468-472.
and communicate the limitations and meaning of a 18. Bojorquez-Tapia, L. A., I. Azuara, E. Ezcurra, and O. Flores-Villela. 1995.
model’s assumptions, relationships, and outputs relative Identifying conservation priorities in Mexico through geographic
to the “real world.” The complexity of wildlife information systems and modeling. Ecological Applications 5:215-231.
management dictates that no one model will suffice. 19. Bookout, T. A. 1994. Research and management techniques for wildlife
Rather, we must develop, apply, evaluate, and improve and habitats. The Wildlife Society, Bethesda, MD.
all models in our toolbox as we see fit, tailoring each to 20. Bowers, M. A. 1994. Dynamics of age- and habitat-structured
populations. Oikos 69:327-333.
meet the demands of the particular situation.
21. Boyce, S. G. 1985. Forestry decisions. United States Forest Service
General Technical Report SE-35.
22. Breininger, D. R., V. L. Larson, B. W. Duncan, and R. B. Smith. 1998.
Linking habitat suitability to demographic success in Florida scrub-jays.
Wildlife Society Bulletin 26:118-128.
23. Brown, E. R. 1985. Management of wildlife and fish habitats in forests
of western Oregon and Washington. Part I. United States Forest
Service, Portland, OR.
 CHAPTER 21: MODELING—INTRODUCTIONS 533

24. Brown, D., N. D. E. Alexander, R. W. Marrs, and S. Albon. 1993. 48. Dedon, M. F., S. A. Laymon, and R. H. Barrett. 1986. Evaluating models
Structured accounting of the variance of demographic change. Journal of wildlife-habitat relationships of birds in black oak and mixed-conifer
of Animal Ecology 62:490-502. habitats. Pages 115-119 in J.Verner, M. L. Morrison, and C. J. Ralph,
25. Brush, T., and E. W. Stiles. 1986. Using food abundance to predict editors. Wildlife 2000: modeling habitat relationships of terrestrial
habitat use by birds. Pages 57-63 in J. Verner, M. L. Morrison, and C. J. vertebrates. University of Wisconsin, Madison, WI.
Ralph, editors. Wildlife 2000: modeling habitat relationships of 49. DeGraaf, R. M., and D. D. Rudis. 1986. New England wildlife: habitat,
terrestrial vertebrates. University of Wisconsin, Madison, WI. natural history, and distribution. United States Forest Service General
26. Bunnell, F. L. 1989. Alchemy and uncertainty: what good are models? Technical Report NE-108.
United States Forest Service General Technical Report PNW-232. 50. Dempster, J. P. 1975. Animal population ecology. Academic Press, New
27. ———, and D. J. Vales. 1990. Comparison of methods for estimating York, NY.
forest overstory cover: differences among techniques. Canadian Journal 51. Dennis, B., and M. L. Taper. 1994. Density dependence in time series
of Forest Research 20:101-107. observations of natural populations: estimation and testing. Ecological
28. Burgman, M. A., S. Ferson, and R. H. Akçakaya. 1993. Risk assessment in Monographs 64:205-224.
conservation biology. Chapman and Hall, London, England. 52. Diamond, J. M. 1975. The island dilemma: lessons of modern
29. Burnham, K. P., D. R. Anderson, and G. C. White. 1996. Meta-analysis of biogeographical studies for the design of natural reserves. Biological
vital rates of the northern spotted owl. Studies in Avian Biology 17:92- Conservation 7:129-146.
101. 53. Digby, P. G. N., and R. A. Kempton. 1987. Multivariate analysis of
30. Capen, D. E. 1981. The use of multivariate statistics in studies of ecological communities. Chapman and Hall, London, England.
wildlife habitat. United States Forest Service General Technical Report 54. Dillon, W. R., and M. Goldstein. 1984. Multivariate analysis: methods
RM-87. and applications. John Wiley & Sons, New York, NY.
31. ———, J. W. Fenwick, D. B. Inkley, and A. C. Boynton. 1986. 55. Doak, D. 1989. Spotted owls and old growth logging in the Pacific
Multivariate models of songbird habitat in New England forests. Pages Northwest. Conservation Biology 3:389-396.
171-175 in J.Verner, M. L. Morrison, and C. J. Ralph, editors. Wildlife 56. Doak, D. F., and L. S. Mills. 1994. A useful role for theory in
2000: modeling habitat relationships of terrestrial vertebrates. conservation. Ecology 75:615-626.
University of Wisconsin, Madison, WI. 57. Dunning, J. B., D. J. Stewart, B. J. Danielson, B. R. Noon, T. L. Root, R. H.
32. Carlin, B. P. 1996. Bayes and empirical Bayes methods for data analysis. Lamberson, and E. E. Stevens. 1995. Spatially explicit population models:
Chapman & Hall, New York, NY. current forms and future uses. Ecological Applications 5:3-11.
33. Carroll, J. E., and R. H. Lamberson. 1993. The owl’s odyssey: a 58. Eberhardt, L. L. 1990. A fur seal population model based on age
continuous model for the dispersal of territorial species. SIAM Journal structure data. Canadian Journal of Fisheries and Aquatic Sciences
of Applied Mathematics 53:205-218. 47:122-127.
34. Caswell, H. 1976. The validation problem. Pages 296-308 in H. H. 59. Ecomap. 1993. National hierarchical framework of ecological units.
Shugart and R.V. O’Neill, editors. Systems ecology. Dowden, United States Forest Service, Washington, D.C.
Hutchinson and Ross, Stroudsburg, PA. 60. Edwards, T. C., Jr., E. T. Deshler, D. Foster, and G. G. Moisen. 1996.
35. ———. 1989. Matrix population models. Sinauer, Sunderland, MA. Adequacy of wildlife habitat relation models for estimating spatial
36. ———, and R.J. Etter. 1993. Ecological interactions in patchy distributions of terrestrial vertebrates. Conservation Biology 10:263-
environments: from patch occupancy models to cellular automata. 270.
Pages 93-109 in S. A. Levin, T. M. Powell, and J. H. Steele, editors. Patch 61. Elton, C. 1927. Animal ecology. Macmillan Company, New York, NY.
dynamics. Springer-Verlag, New York, NY. 62. Fahrig, L., and J. Paloheimo. 1988. Effect of spatial arrangement of
37. Caughley, G., J. Short, G. C. Grigg, and H. Nix. 1987. Kangaroos and habitat patches on local population size. Ecology 69:468-475.
climate: an analysis of distribution. Journal of Animal Ecology 56:751- 63. Fitzgerald, E. C., and C. H. Nilon. 1993. Testing the accuracy of an HSI
761. model in an urban county. Transactions of the North American Wildlife
38. ———, and A. R. E. Sinclair. 1994. Wildlife ecology and management. and Natural Resources Conference 58:117-123.
Blackwell Scientific Publications, Boston, MA. 64. Flather, C. H., and T. W. Hoekstra. 1985. Evaluating population-habitat
39. Chang, C.-L. 1997. Fuzzy-logic-based programming. World Scientific, models using ecological theory. Wildlife Society Bulletin 13:121-130.
Singapore, Singapore. 65. Foley, P. 1997. Extinction models for local populations. Pages 215-246
40. Clark, W. R., and G. S. Innis. 1982. Forage interactions and black-tailed in I. Hanski and M. E. Gilpin, editors. Metapopulation biology: ecology,
jack rabbit population dynamics: a simulation model. Journal of Wildlife genetics, and evolution. Academic Press, San Diego, CA.
Management 46:1018-1035. 66. Forman, R. T. T., and M. Godron. 1986. Landscape ecology. John Wiley
41. Conroy, M. J.,Y. Cohen, F. C. James, Y. G. Matsinos, and B. A. Maurer. and Sons, New York, NY.
1995. Parameter estimation, reliability, and model improvement for 67. Frederick, R. B., W. R. Clark, and E. E. Klass. 1987. Behavior, energetics,
spatially explicit models of animal populations. Ecological Applications and management of refuging waterfowl: a simulation model. Wildlife
5:17-19. Monographs 96:1-35.
42. Cook, J. G., T. W. Stutzman, C. W. Bowers, K. A. Brenner, and L. L. Irwin. 68. Frisman, E. Y., E. I. Skaletskaya, and A. E. Kuzin. 1982. A mathematical
1995. Spherical densiometers produce biased estimates of forest model of the population dynamics of a local northern fur-seal herd.
canopy cover. Wildlife Society Bulletin 23:711-717. Ecological Modelling 16:151-172.
43. Crouse, D. T., L. B. Crowder, and H. Caswell. 1987. A stage-based 69. Fryxell, J. M., and P. Lundberg. 1998. Individual behavior and
population model for loggerhead sea turtles and implications for community dynamics. Chapman & Hall, New York, NY.
conservation. Ecology 68:1412-1423. 70. Goeden, G. B. 1979. Biogeographic theory as a management tool.
44. Crowder, L. B., D. T. Krouse, S. S. Heppell, and T. H. Martin. 1994. Environmental Conservation 6:27-32.
Predicting the impact of turtle excluder devices on loggerhead sea 71. Grant, W. E. 1986. Systems analysis and simulation in wildlife and
turtle populations. Ecological Applications 4:437-445. fisheries sciences. John Wiley & Sons, New York, NY.
45. Czárán, T., and S. Bartha. 1992. Spatiotemporal dynamics models of 72. Grubb, T. G. 1988. Pattern recognition-a simple model for evaluating
plant populations and communities. Trends in Ecology and Evolution wildlife habitat. United States Forest Service Research Note RM-487.
7:38-42. 73. Gutierrez, A. P. 1996. Applied population ecology - a supply-demand
46. Daubenmire, R. 1952. Forest vegetation of northern Idaho and approach. John Wiley & Sons, New York, NY.
adjacent Washington, and its bearing on concepts of vegetation 74. Hall, C. A. S., and J. W. Day, Jr. 1977a. Systems and models: Terms and
classification. Ecological Monographs 22:301-330. basic principles. Pages 6-36 in C. A. S. Hall and J. W. Day, Jr., editors.
47. ———. 1959. A canopy-coverage method of vegetation analysis. Ecosystem modeling in theory and practice. John Wiley & Sons, New
Northwest Science 33:43-64. York, NY.
 534 Wildlife–Habitat Relationships in Oregon and Washington

75. ———, and J. W. Day, Jr. 1977b. Ecosystem modeling in theory and 99. Kareiva, P. 1990. Population dynamics in spatially complex
practice. John Wiley & Sons, New York, NY. environments: theory and data. Pages 53-68 in M. P. Hassell and R. M.
76. Hanski, I. 1994. A practical model of metapopulation dynamics. Journal May, editors. Population regulation and dynamics. Royal Society, London,
of Animal Ecology 63:151-162. England.
77. ———, and M. Gilpin. 1991. Metapopulation dynamics: brief history 100. Kareiva, P., and U. Wennergren. 1995. Connecting landscape patterns
and conceptual domain. Biological Journal of the Linnean Society 42:3- to ecosystem and population processes. Nature 373:299-302.
16. 101. Kling, C. L. 1980. Pattern recognition for habitat evaluation. Thesis,
78. ———, and D. Simberloff. 1997. The metapopulation approach, its Colorado State University, Fort Collins, CO.
history, conceptual domain, and application to conservation. Pages 5-26 102. Knopf, F. L., J. A. Sedgwick, and R. W. Cannon. 1988. Guild structure of
in I. Hanski and M. E. Gilpin, editors. Metapopulation biology: ecology, a riparian avifauna relative to seasonal cattle grazing. Journal of Wildlife
genetics, and evolution. Academic Press, San Diego, CA. Management 52:280-190.
79. Harestad, A. S., and D. G. Keisker. 1989. Nest tree use by primary 103. Kohlmann, S. G., and M. J. Hedrick. In Press. Density dependence in
cavity-nesting birds in south central British Columbia. Canadian Journal two Great Basin desert ungulates: implications for adaptive
of Zoology 67:1067-1073. management. Journal of Wildlife Management.
80. Hariston, N. G. 1981. An experimental test of a guild: salamander 104. Krebs, C. J. 1989. Ecological methodology. HarperCollins Publishers,
competition. Ecology 62:65-72. New York, NY.
81. Harris, L. D. 1984. The fragmented forest: island biogeography theory 105. Lahaye, W. S., R. J. Gutiérrez, and H. R. Akçakaya. 1994. Spotted owl
and the preservation of biological diversity. University of Chicago, metapopulation dynamics in southern California. Journal of Animal
Chicago, IL. Ecology 63:775-785.
82. Hastings, A. 1991. Structured models of metapopulation dynamics. 106. Lamberson, R. H., B. R. Noon, C.Voss, and K. S. McKelvey. 1994.
Biological Journal of the Linnean Society 42:57-71. Reserve design for territorial species: the effects of patch size and
83. ———, and S. Harrison. 1994. Metapopulation dynamics and genetics. spacing on the viability of the northern spotted owl. Conservation
Annual Review of Ecology and Systematics 25:167-188. Biology 8:185-195.
84. Haufler, J. B., C. A. Mehl, and G. J. Roloff. 1996. Using a coarse-filter 107. Lancia, R. A., S. D. Miller, D. A. Adams, and D. W. Hazel. 1982. Validating
approach with species assessment for ecosystem management. Wildlife habitat quality assessment: an example. Transactions of the North
Society Bulletin 24:200-208. American Wildlife and Natural Resources Conference 47:96-110.
85. ———, C. A. Mehl, and G. J. Roloff. 1998. Conserving biological 108. ———, and D. A. Adams. 1985. A test of habitat suitability index
diversity using a coarse-filter approach with a species assessment. models for five bird species. Proceedings of the Annual Conference of
Pages 107-125 in R. K. Baydack, H. Campa III, and J. B. Haufler, editors. the Southeastern Association of Fish and Wildlife Agencies 39:412-419.
Practical approaches to the conservation of biological diversity. Island 109. Lande, R. 1987. Extinction thresholds in demographic models of
Press, Washington, D.C. territorial populations. American Naturalist 130:624-635.
86. Hitchcock, C. L., and C. Gratt-Trevor. 1997. Diagnosing a shorebird 110. ———. 1988. Demographic models of the northern spotted owl
local population decline with a stage-structured population model. (Strix occidentalis caurina). Oecologia 75:601-607.
Ecology 78:522-534. 111. Laymon, S. A., and R. H. Barrett. 1986. Developing and testing habitat-
87. Hobbs, N. T. 1989. Linking energy balance to survival in mule deer: capability models: pitfalls and recommendations. Pages 87-91 in J.Verner,
development and test of a simulation model. Wildlife Monographs M. L. Morrison, and C. J. Ralph, editors. Wildlife 2000: modeling habitat
101:1-39. relationships of terrestrial vertebrates. University of Wisconsin,
88. Hof, J., and M. G. Raphael. 1997. Optimization of habitat placement: a Madison, WI.
case study of the northern spotted owl in the Olympic Peninsula. 112. Lefkovitch, L. P. 1965. The study of population growth in organisms
Ecological Applications 7:1160-1169. grouped by stages. Biometrics 21:1-18.
89. Holl, S. A. 1982. Evaluation of bighorn sheep habitat. Desert Bighorn 113. Leslie, P. H. 1945. On the use of matrices in certain population
Council Transactions, 47-49. mathematics. Biometrika 33:183-212.
90. Hollander, A. D., F. W. Davis, and D. M. Stoms. 1994. Hierarchical 114. Levin, S. A. 1976. Population dynamic models in heterogeneous
representations of species distributions using maps, images, and sighting environments. Annual Review of Ecology and Systematics. 7:287-310.
data. Pages 71-88 in R. I. Miller, editor. Mapping the diversity of nature. 115. Levins, R. 1969. Some demographic and genetic consequences of
Chapman and Hall, London, England. environmental heterogeneity for biological control. Bulletin of the
91. Holthausen, R. S., M. G. Raphael, K. S. McKelvey, E. D. Forsman, E. E. Entomological Society of America 15:237-240.
Starkey, and D. E. Seaman. 1995. The contribution of federal and 116. ———. 1970. Extinction. Pages 77-107 in M. Gerstenhaber, editor.
nonfederal habitat to persistence of the northern spotted owl on the Some mathematical problems in biology. American Mathematical
Olympic Peninsula, Washington: report of the reanalysis team. United Society, Providence, RI.
States Forest Service General Technical Report PNW-352. 117. Lindenmayer, D. B., and H. P. Possingham. 1996. Ranking conservation
92. Huston, M., D. DeAngelis, and W. Post. 1988. New computer models and timber management options for Leadbeater’s possum in
unify ecological theory. Bioscience 38:682-691. southeastern Australia using population viability analysis. Conservation
93. Hutchinson, G. E. 1957. Concluding remarks. Cold Spring Harbor Biology 10:235-251.
Symposium on Quantitative Biology 22:415-427. 118. Liu, J., J. B. Dunning, and H. R. Pulliam. 1995. Potential effects of a
94. Inger, R. F., and R. K. Colwell. 1977. Organization of contiguous forest management plan on Bachman’s sparrow (Aimophila aestivalis):
communities of amphibians and reptiles in Thailand. Ecological linking a spatially explicit model with GIS. Conservation Biology 9:62-
Monographs 47:229-253. 75.
95. Iversen, G. R. 1984. Bayesian statistical inference. Sage Publications, 119. Lubow, B. C., G. C. White, and D. R. Anderson. 1996. Evaluation of a
Beverly Hills, CA. linked sex harvest strategy for cervid populations. Journal of Wildlife
96. James, F. C. 1971. Ordinations of habitat relationships among breeding Management 60:787-796.
birds. Wilson Bulletin 83:215-236. 120. Lusted, L. B. 1968. Introduction to medical decision making. Charles
97. ———, and N. O. Wamer. 1982. Relationships between temperate C. Thomas, Springfield, IL.
forest bird communities and vegetation structure. Ecology 63:159-171. 121. MacArthur, R. H., and E. O. Wilson. 1967. The theory of island
98. Judson, O. P. 1994. The rise of the individual-based model in ecology. biogeography. Princeton University, Princeton, NJ.
Trends in Evolution and Ecology 9:9-14. 122. Mannan, R. W., M. L. Morrison, and E. C. Meslow. 1984. Comment: the
use of guilds in forest bird management. Wildlife Society Bulletin
12:426-430.
 CHAPTER 21: MODELING—INTRODUCTIONS 535

123. Marcot, B. G. 1979. California wildlife/habitat relationships program: 146. Poiani, K. A., and W. C. Johnson. 1993. A spatial simulation model of
north coast/Cascades zone.Volumes 1-4. United States Forest Service, hydrology and vegetation dynamics in semi-permanent prairie wetlands.
Eureka, CA. Ecological Applications 3:279-293.
124. ———, M. G. Raphael, and K. H. Berry. 1983. Monitoring wildlife 147. Prosser, D. J., and R. P. Brooks. 1998. A verified habitat suitability
habitat and validation of wildlife-habitat relationships models. index for the Louisiana waterthrush. Journal of Field Ornithology
Transactions of the North American Wildlife Conference 48:315-329. 69:288-298.
125. Maurer, B. A. 1986. Predicting habitat quality for grassland birds using 148. Pulliam, R. H. 1988. Sources, sinks, and population regulation.
density-habitat correlations. Journal of Wildlife Management 50:556- American Naturalist 132:652-661.
556. 149. ———, and B. J. Danielson. 1991. Sources, sinks, and habitat
126. ———. 1990. Extensions of optimal foraging theory for selection: a landscape perspective on population dynamics. American
insectivorous birds: implications for community structure. Studies in Naturalist 137:550-566.
Avian Biology 13:455-461. 150. Pulliam, R. H., J. B. Dunning, and J. Liu. 1992. Population dynamics in
127. ———. 1994. Geographical population analysis: tools for the analysis complex landscapes: a case study. Ecological Applications 2:165-177.
of biodiversity. Blackwell Scientific Publications, Oxford, England. 151. Puttock, G. D., P. Shakotko, and J. G. Rasaputra. 1996. An empirical
128. Maxwell, J. R., C. J. Edwards, M. E. Jensen, S. J. Paustian, H. Parrott, and habitat model for moose, Alces alces, in Algonquin Park, Ontario. Forest
D. M. Hill. 1995. A hierarchical framework of aquatic ecological units in Ecology and Management 81:169-178.
North America (Nearctic zone). United States Forest Service General 152. Quinn, T. 1997. Coyote (Canis latrans) habitat selection in urban areas
Technical Report NC-176. of western Washington via analysis of routine movements. Northwest
129. May, R. M. 1973. On relationships among various types of population Science 71:289-297.
models. American Midland Naturalist 107:46-57. 153. Raphael, M. G., and B. G. Marcot. 1986.Validation of a wildlife-habitat-
130. Maynard-Smith, J. 1974. Models in ecology. Cambridge University relationships model: vertebrates in a Douglas-fir sere. Pages 129-138 in
Press, Cambridge, England. J. Verner, M. L. Morrison, and C. J. Ralph, editors. Wildlife 2000: modeling
131. McCullough, D. R. 1981. Population dynamics of the Yellowstone habitat relationships of terrestrial vertebrates. University of Wisconsin,
grizzly bear. Pages 173-196 in C. W. Fowler and T. D. Smith, editors. Madison, WI.
Dynamics of large mammal populations. John Wiley & Sons, New York, 154. Renken, R. B. 1988. Habitat characteristics related to pileated
NY. woodpecker densities and territory size in Missouri. Dissertation,
132. McNeill, F. M., and E. Thro. 1994. Fuzzy logic: a practical approach. University of Missouri, Columbia, MO.
Academic Press, Cambridge, MA. 155. Ricklefs, R. E., and D. B. Miles. 1994. Ecological and evolutionary
133. McKelvey, K. 1992. A spatially explicit life-history simulator for the inferences from morphology: an ecological perspective. Pages 13-41 in
northern spotted owl. Appendix 4-P in Eugene District Resource P. C. Wainwright and S. M Reilly, editors. Ecological morphology.
Management Plan & EIS. United States Bureau of Land Management, University of Chicago, Chicago, IL.
Eugene, OR. 2 vols. 156. Robel, R. J., L. B. Fox, and K. E. Kemp. 1993. Relationship between
134. Messier, F. 1994. Ungulate population models with predation: a case habitat suitability index values and ground counts of beaver colonies in
study with the North American moose. Ecology 75:478-488. Kansas. Wildlife Society Bulletin 21:415-421.
135. Mills, T. R., M. A. Rumble, and L. D. Flake. 1996. Evaluation of a habitat 157. Roloff, G. J., and J. B. Haufler. 1997. Establishing population viability
capability model for nongame birds in the Black Hills, South Dakota. planning objectives based on habitat potentials. Wildlife Society Bulletin
United States Forest Service Research Paper RM-RO-323. 25:895-904.
136. Morrison, M. L., B. G. Marcot, and R. W. Mannan. 1992. Wildlife- 158. ———, B. Carroll, and S. Scharosch. 1999. A decision support
habitat relationships: concepts and applications. University of system for incorporating wildlife habitat quality into forest planning.
Wisconsin, Madison, WI. Western Journal of Applied Forestry 14:91-99.
137. ———, B. G. Marcot, and R. W. Mannan. 1998. Wildlife habitat 159. ———, and B. J. Kernohan. 1999. Evaluating the reliability of habitat
relationships: concepts and applications. Second edition. University of suitability index models. Wildlife Society Bulletin 27:973-985.
Wisconsin Press, Madison, WI. 160. Root, R. B. 1967. The niche exploitation pattern of the blue-gray
138. Moss, R., A. Watson, and J. Ollason. 1982. Animal population gnatcatcher. Ecological Monographs 37:317-350.
dynamics. J.W. Arrowsmith Limited, Bristol, England. 161. Root, K.V. 1998. Evaluating the effects of habitat quality, connectivity,
139. Nadeau, S., R. Décarie, D. Lambert, and M. St-Georges. 1995. and catastrophes on a threatened species. Ecological Applications
Nonlinear modeling of muskrat use of habitat. Journal of Wildlife 8:854-865.
Management 59:110-117. 162. Rosenzweig, M. L. 1995. Species diversity in space and time.
140. Noon, B. R., and K. McKelvey. 1992. Stability properties of the Cambridge University Press, New York, NY.
spotted owl metapopulation in southern California. United States 163. Rotella, J. J., J. T. Ratti, K. P. Reese, M. L. Taper, and B. Dennis. 1996.
Forest Service General Technical Report PSW-133. Long-term population analysis of gray partridge in eastern Washington.
141. O’Connor, R. J., M. T. Jones, D. White, C. T. Hunsaker, T. Loveland, B. Journal of Wildlife Management 60:817-825.
Jones, and E. Preston. 1996. Spatial partitioning of environmental 164. Rothery, P., I. Newton, L. Dale, and T. Wesolowski. 1997. Testing for
correlates of avian biodiversity in the conterminous United States. density dependence allowing for weather effects. Oecologia 112:518-
Biodiversity Letters 3:97-110. 523.
142. ———, and M. T. Jones. 1997. Using hierarchical models to index the 165. Ruckelshaus, M., C. Hartway, and P. Kareiva. 1997. Assessing the data
ecological health of the nation. Transactions of the North American and requirements of spatially explicit dispersal models. Conservation
Natural Resources Conference 62:501-508. Biology 11:1298-1306.
143. Pacala, S. W., C. D. Canham, and J. A. Silander, Jr. 1993. Forest models 166. Rykiel, E. J., Jr. 1996. Testing ecological models: the meaning of
defined by field measurements. I. The design of a northeastern forest validation. Ecological Modelling 90:229-244.
simulator. Canadian Journal of Forest Research 23:1980-1988. 167. Saitoh, T., N. C. Stenseth, and O. N. Bjornstadt. 1997. Density
144. Pascual, M. A., P. Kareiva, and R. Hilborn. 1997. The influence of model dependence in fluctuating gray-sided vole populations. Journal of Animal
structure on conclusions about the viability and harvesting of Serengeti Ecology 66:14-24.
wildebeest. Conservation Biology 11:966-976. 168. Sauer, J. R., and S. Droege. 1990. Survey designs and statistical
145. Pastor, J., and W. M. Post. 1985. Development of a linked forest methods for the simulation of avian population trends. United States
productivity-soil process model. Publication of Oak Ridge National Fish and Wildlife Service Biological Report 90(1).
Laboratory, Environmental Sciences Division, Number 2455, Oak Ridge,
TN.
 536 Wildlife–Habitat Relationships in Oregon and Washington

169. Saunders, D. A., R. J. Hobbs, and C. R. Margules. 1991. Biological 192. ———, E. D. Forsman, J. B. Lint, E. C. Meslow, B. R. Noon, and J.
consequences of ecosystem fragmentation: a review. Conservation Verner. 1990. A conservation strategy for the northern spotted owl.
Biology 5:18-32. Report of the Interagency Scientific Committee to address the
170. Schmutz, J. A., R. F. Rockwell, and M. R. Petersen. 1997. Relative effects conservation of the northern spotted owl. Appendix M. United States
of survival and reproduction on the population dynamics of emperor Forest Service, Portland, OR.
geese. Journal of Wildlife Management 61:191-201. 193. Turner, M. G. 1989. Landscape ecology: the effect of pattern on
171. Schroeder, R. L. 1986. Habitat suitability index models: wildlife process. Annual Review of Ecology and Systematics 20:171-197.
species richness in shelterbelts. United States Fish and Wildlife Service 194. ——— G. J. Arthaud, R. T. Engstrom, S. J. Hejl, J. Liu, S. Loeb, and K.
Biological Report 82(10.128). McKelvey. 1995. Usefulness of spatially explicit population models in
172. ———, and S. L. Haire. 1993. Guidelines for the development of land management. Ecological Applications 5:12-16.
community-level habitat evaluation models. United States Fish and 195. Urban, D. L., R. V. O’Neill, and H. H. Shugart. 1987. Landscape ecology.
Wildlife Service Biological Report 8, Washington, D.C. Bioscience 37:119-27.
173. Scott, J. M., F. Davis, B. Csuti, R. Noss, B. Butterfield, C. Groves, H 196. United States Fish and Wildlife Service. 1980. Habitat evalaution
Anderson, S. Caicco, F. D’erchia, T. C. Edwards, Jr., J. Ulliman, and R. G. procedures (HEP). United States Fish and Wildlife Service, Division of
Wright. 1993. Gap analysis: a geographic approach to protection of Ecological Services, ESM 102, Washington, D.C.
biological diversity. Wildlife Monographs 123:1-41. 197. ——— 1981. Standards for the development of habitat suitability
174. Severinghaus, W. D. 1981. Guild theory development as a mechanism index models. United States Fish and Wildlife Service Division of
for assessing environmental impacts. Environmental Management 5:187- Ecological Services, ESM 103, Washington, D.C.
190. 198. Vales, D. J., and F. L. Bunnell. 1988. Comparison of methods for
175. Short H. L., and K. P. Burnham. 1982. Technique for structuring estimating forest overstory cover. I. Observer effects. Canadian Journal
wildlife guilds to evaluate impacts on wildlife communities. United of Forest Research 18:606-609.
States Fish and Wildlife Service Special Science Report on Wildlife, 199.Varley, G. C., G. R. Gradwell, and M. P. Hassell. 1973. Insect population
Number 244, Washington, D.C. ecology—an analytical approach. Blackwell Press, Oxford, England.
176. Sibly, R. M., and R. H. Smith. 1998. Identifying key factors using lambda 200.Van Horne, B., and J. A. Wiens. 1991. Forest bird habitat suitability
contribution analysis. Journal of Animal Ecology 67:17-24. models and the development of general habitat models. United States
177. Simberloff, D. 1988. The contribution of population and community Fish and Wildlife Service Research Number 8, Washington, D.C.
biology to conservation science. Annual Review of Ecology and 201.Van Tienderen, P. H. 1995. Life cycle trade-offs in matrix population
Systematics 19:473-511. models. Ecology 76:2482-2489.
178. Slobodkin, L. B. 1980. Growth and regulation of animal populations. 202.Verner, J., and A. S. Boss. 1980. California wildlife and their habitats:
Dover Publishing, New York, NY. western Sierra Nevada. United States Forest Service General Technical
179. Smith, A. D. M., and C. J. Walters. 1981. Adaptive management of Report PSW-37.
stock-recruitment systems. Canadian Journal of Fisheries and Aquatic 203. ———. 1984. The guild concept applied to management of bird
Sciences 38:690-703. populations. Environmental Management 8:1-14.
180. Starfield, A. M., and A. L. Bleloch. 1986. Building models for 204. ———, M. L. Morrison, and C. J. Ralph. 1986. Wildlife 2000: modeling
conservation and wildlife management. Macmillan Publishing Company, habitat relationships of terrestrial vertebrates. University of Wisconsin,
New York, NY. Madison, WI.
181. ———, and A. L. Bleloch. 1991. Building models for conservation 205. Wahlberg, N., A. Moilanen, and I. Hanski. 1996. Predicting the
and wildlife management. Burgess International Group, Edina, MN. occurrence of endangered species in fragmented landscapes. Science
182. ———, K. A. Smith, and A. L. Bleloch. 1994. How to model it: 273:1536-1538.
Problem-solving for the computer age. Burgess International Group, 206. Walters, C. J., and R. Hilborn. 1976. Adaptive control of fishing
Edina, MN. systems. Journal of the Fisheries Research Board, Canada 33:145-159.
183. ———. 1997. A pragmatic approach to modeling for wildlife 207. ———, and R. Hilborn. 1978. Ecological optimization and adaptive
management. Journal of Wildlife Management 61:261-270. management. Annual Review of Ecology and Systematics 9:157-188.
184. Steeger, C., and C. L. Hitchcock. 1998. Influence of forest structure 208. Washington Department of Natural Resources. 1996. Draft
and diseases on nest-site selection by red-breasted nuthatches. Journal environmental impact statement: habitat conservation plan. Washington
of Wildlife Management 62:1349-1358. State Department of Natural Resources, Olympia, WA.
185. Suring, L. H., K. R. Barber, C. C. Schwartz, T. N. Bailey, W. C. Shuster, 209. Watt, K. E. F. 1966. Systems analysis in ecology. Academic Press, New
and M. D. Tetreau. 1998. Analysis of cumulative effects on brown bears York, NY.
on the Kenai Peninsula, southcentral Alaska. Ursus 10:107-117. 210. Wells, J. V., and M. E. Richmond. 1995. Populations, metapopulations,
186. Sutherland, W. J. 1996. From individual behaviour to population and species populations: what are they and who should care? Wildlife
ecology. Oxford University Press, New York, NY. Society Bulletin 23:458-462.
187. Temple, S. A., and J. R. Cary. 1988. Modeling dynamics of habitat- 211. Wennergren, U., M. Ruckelshaus, and P. Kareiva. 1995. The promise
interior bird populations in fragmented landscapes. Conservation and limitations of spatial models in conservation biology. Oikos 74:349-
Biology 2:340-347. 356.
188. Terborgh, J. 1974. Preservation of natural diversity: the problem of 212. Williams, G. L., K. R. Russell, and K. R. Seitz. 1977. Pattern recognition
extinction prone species. Bioscience 24:715-722. as a tool in the ecological analysis of habitat. Pages 521-531 in
189. Terrell, J. W., and J. Carpenter. 1997. Selected habitat suitability index Classification, inventory, and analysis of fish and wildlife habitat. United
model evaluations. United States Geological Survey Information and States Fish and Wildlife Service FWS/OBS-78/76.
Technical Report USGS/BRD/ITR-1997-0005. 213. Wisdom, M. J., L. R. Bright, C. G. Carey, W. W. Hines, R. J. Pedersen, D.
190. Tester, J. R., A. M. Starfield, and L. E. Frelich. 1997. Modeling for A. Smithey, J. W. Thomas, and G. W. Witmer. 1986. A model to evaluate
ecosystem management in Minnesota pine forests. Biological elk habitat in western Oregon. United States Forest Service, Pacific
Conservation 80:313-324. Northwest Region, Publication Number R6-F&WL-216-1986.
191. Thomas, J. W. 1979. Wildlife habitats in managed forests: the Blue 214. With, K. A., and T. O. Crist. 1996. Translating across scales: simulating
Mountains of Oregon and Washington. United States Forest Service species distributions as the aggregate response of individuals to
Agriculture Handbook Number 553. heterogeneity. Ecological Modeling 93:125-137.
 22
Five Case Studies in Wildlife Modeling
Timothy Quinn & David H. Johnson, editors

Introduction to the Case Studies

The five case studies that make up this chapter were
Case Study 1 presented at the Landscape Management of Pacific Northwest
A spatially realistic population model for Forest Workshop,* held in Olympia, Washington in February
informing forest management decisions 1998. Designed for land mangers and other natural
G.F. Wilhere, N.H. Schumaker, & S.P. Horton resource professionals, the purpose of the workshop was
Case Study 2 to help bridge the gap between the theory and practice of
A model to assess potential vertebrate habitat at conservation biology. In particular, as workshop
landscape scales: HABSCAPES organizers, we (T. Quinn and D.H. Johnson) were
M. Huff, T.K. Mellen, & R. Hagestedt interested in studies describing practical techniques and
tools that provide insights or solutions to complex, large-
Case Study 3 scale or multi-species conservation issues.
Cross-scale classification trees for assessing risks The five studies presented below represent an accurate
of forest practices to headwater stream range, and sampling, of the many projects presented at
amphibians the workshop. They are unique in the sense that each case
G.D. Sutherland & F.L. Bunnell study presents a methodology used in a real conservation
Case Study 4 setting, and one that also addresses common problems
Applying Gap analysis to county and regional with data quality and quantity. In other words, they used
land use planning data they had “at hand.” We present these case studies as
M.R. Stevenson examples of creative approaches rather than solutions to
Case Study 5 specific conservation issues. There are no perfect answers
A model to determine potential Northern spotted here, but there is abundant creative energy about
owl nesting areas important, and ever-improving, work by practitioners.
N.W. Darby & T. Young Case Study #1 by Wilhere et al. is an example of a
potentially powerful management tool to address a
fundamental management issue: predicting the effect of
landscape-level habitat changes to wildlife species or
populations. The single-species, spatially realistic
population model described here requires movement and
demographic data (see previous chapter), which typically
are available only for the best-studied species. To address
data limitations, Wilhere et al. introduce a process called
parameter tuning. Parameter tuning is a way of estimating
specific parameter values for which no specific data exist
by iteratively solving for those values during model
simulations. By treating known parameters as constants,
Wilhere et al. could search for appropriate values of the
unknown parameter by comparing the characteristics of
the simulated and real populations. Model output tended
*
The sponsors of the workshop were: Washington State Timber, to become more concordant with empirical data, as
Fish, and Wildlife Program; Wildlife-Habitat Relationships Project; increasingly realistic values are used to estimate the
Washington Dept. Of Fish and Wildlife; U.S. Fish and Wildlife unknown parameter. While this process does not lessen
Service; Washington and Oregon Chapters of The Wildlife the need for good data, it may provide managers with
Society; Organization of Fish and Wildlife Information Managers; better ways of using existing data.
and Environmental Systems Research Institute (ESRI).

537
 538 Wildlife–Habitat Relationships in Oregon and Washington

Case Study #2 by Huff et al. describes a process called conservation. Similar to HABSCAPES, the authors outline
HABSCAPES for determining distributions of suitable a process for assessing the distribution of potential
habitat for a species or collections of species (guilds), which northern spotted owl (NSO) nesting sites using vegetation
are grouped by resource use. The process uses vegetation cover type data (including vegetation structural data) and
cover type data (including vegetation structural autecological information about the NSO. The model is
characteristics) in conjunction with data on species-habitat built using routines commonly found in GIS software. This
associations, home range or territory size, and species life case study is not included here because we believe it has
history, to identify all patches of potential habitat. Users just the right recipe for spotted owls, but rather because it
can specify criteria, such as the size and composition of reminds us of the power of GIS as an integrator of
patches that comprise habitat to quickly delineate potential information from the fields such as forestry, biology, and
habitat at relatively large spatial scales. One unique feature economics.
of HABSCAPES, and an obvious place to issue caution, is
that it can streamline multi-species assessments by
guilding species. It is best thought of as a coarse filter Case Study 1
approach to habitat assessments. A Spatially Realistic Population Model
Case Study #3 by Sutherland and Bunnel explored the for Informing Forest Management
use of classification and regression tree analysis (CART) Decisions
for determining habitat relationships of tailed frogs at
George F.Wilhere, Nathan H. Schumaker, & Scott P. Horton
different spatial scales. They argue that many of the
spotted frog studies are contradictory because they were
done at different places, or they explained frog response Introduction
using independent variables representing different spatial Spatially realistic population models (SRPMs) address a
scales. Sutherland and Bunnel suggest that CART can be fundamental problem commonly confronted by wildlife
more appropriate than multiple or logistic regression at managers—predicting the effects of landscape-scale
teasing out relationships occurring at different spatial scale habitat management on an animal population. SRPMs
when only inventory data (relative abundance or typically consist of three submodels: (1) a habitat
occurrence) are available. While noting that it is heuristic submodel, (2) a movement submodel, and (3) a
in nature, they concluded that CART has four important demographic submodel. We describe the submodels and
strengths as demonstrated in their study: 1) it reveals data requirements for the typical SRPM. The most
pathways by which environmental variation may frustrating problem with SRPMs is the lack of data needed
determine species distribution, 2) it can use commonly to relate movement and demographic parameters to
available information comprised of numerical and habitat quality. We developed a SRPM to evaluate the
nominal data, 3) it can facilitate the use of separate data relative effects of different habitat management strategies
without making assumptions about how environmental on the spotted owl subpopulation of the Olympic
variables interact at all locations, and 4) it produces Peninsula. This case study documents some plausible
outputs that are easy to interpret and to convert into assumptions we made to circumvent the data problem,
hypotheses. and explains an approach called “parameter tuning” that
we used to generate parameter values.
Case Study #4: Stevenson describes a process of applying
data from Washington Gap and other sources to help Focus of Case Study
Spokane County plan for human population growth and The Washington State Department of Natural Resources
the protection of biodiversity. He outlines five steps to the (DNR) has developed a habitat conservation plan7 for
planning process: 1) identify the minimum amount of land 107,000 ha of state lands on the western Olympic Peninsula
for each species, 2) establish species richness thresholds that emphasizes the habitat needs of the Northern spotted
to located candidate reserves, 3) delineate reserves as an owl. The plan will alter landscape conditions across more
iterative process to account for local knowledge, than 4000 km2 of inter-mixed state, federal, and private
economics and politics, 4) delineate corridors connecting land, and thus influence the fate of the Peninsula’s spotted
reserves, and 5) evaluate success. While the steps reflect owl subpopulation. The critical question was how much
common approaches to conservation, this case study of an influence? Specifically, would the conservation plan
demonstrates how local knowledge and multiple data sets appreciably reduce the likelihood of survival and recovery
can be combined in useful ways during planning of spotted owls on the Peninsula? In addition, since
processes. Perhaps the most important thing about this Washington’s State Environmental Policy Act dictates that
case study is that someone despite the potential problems state agencies shall consider more than one alternative
with data and bureaucracy took the time to work with when planning large projects, DNR needed to answer
local jurisdictions that were struggling with managing questions about the relative effects of different habitat
human population growth. management strategies. During development of the plan,
Case Study #5: Darby and Young discuss how to build a spatially realistic population models (SRPMs) were
geographical information system (GIS) model to help emerging as a practical approach for answering these types
managers make more informed decisions about of questions,24, 29 so the DNR funded the development of a
 CHAPTER 22: MODELING—CASE HISTORIES 539

new SRPM that offered some advantages over existing The data requirements of a habitat submodel
models.6, 7, 31 We will refer to the model of Schumaker31 as correspond with its multiple dimensions. An intermediate
SOS for “Spotted Owl Simulator”. SOS is a pre-cursor to data product must be a habitat map showing the location
the model known as PATCH.32 of various vegetation classes or habitat categories. The
simplest map could consist of two categories: habitat and
An Introduction to Spatially Realistic nonhabitat.20, 29 Such maps may be generated through
Population Models remote sensing methods (e.g,. aerial photography or
Spatially realistic population models address a satellite thematic mapper imaging), through on-the-
fundamental problem commonly confronted by wildlife ground habitat surveys, or most likely some combination
managers—predicting the effects of landscape-scale of both. The habitat map is digitized using a geographic
habitat management on an animal population. SRPMs are information system and the habitat submodel translates
a type of spatially explicit population model.16 Spatially the digitized map into a simulated landscape. Forest
explicit models “keep track of the exact positions of plants growth models or information about future land use
and animals,”39 but most types of spatially explicit models patterns can be used to create maps of likely changes in
are based on simplistic, stylized landscapes (Roloff et al., habitat conditions over time.
this volume). In contrast, a SRPM uses empirically derived Ideally, a movement submodel would simulate realistic
habitat data that are referenced to real geographic movement behavior. That is, metrics of simulated
coordinates. These “realistic” landscapes can be movement, such as total distance moved, net distance
manipulated to represent conditions likely to result from moved, net direction moved, linear auto-correlation (i.e.,
future habitat management. SRPMs are usually the tendency to move in a straight line), and habitat
individual-based models. That is, location, birth, death, preference would be statistically representative of real
reproduction, and movement are “observed” and recorded movements. Certainly, habitat quality must influence
for every individual in the simulated population. The movement behavior, as must interactions with
behavior and fate of individuals are stochastic. This means conspecifics, but unfortunately, our understanding of such
that most actions or events occur randomly according to behavior is crude and the cost of improving our
probability distributions that are either estimated from understanding has been prohibitive. Consequently, a
empirical data or based on plausible assumptions. SRPMs common approach has been to simply intuit the cognitive
are typically comprised of three submodels: (1) a habitat mechanisms, such as “memory,” habitat “selectivity,” and
submodel, (2) a movement submodel, and (3) a intraspecific conflict “aversion” that reputedly manifest
demographic submodel. movement behaviors.25, 28
The habitat submodel creates an abstract representation If a habitat submodel uses a hexagonal grid to define
of a real landscape. SRPMs are usually limited to two landscape units, then movement paths consist of a
spatial dimensions (X and Y coordinates), a third sequence of steps from hexagon to hexagon. At each
dimension related to habitat quality, and a fourth hexagon, an individual can move in one of six directions
dimension is time, which implies dynamic habitat quality. or not move at all. Submodel parameters define how
The habitat submodel must interface with the “cognitive mechanisms” influence individual movement
demographic and movement submodels. That is, the decisions. Movement submodels in SRPMs usually allow
submodel should delineate landscape units that have a range of behaviors from totally random to completely
demographic significance, such as territories, or that may directed by habitat quality and the presence of
be relevant to movement behavior, such as habitat conspecifics. Because long-distance dispersal is considered
corridors. Also, a habitat submodel must provide metrics a risky behavior, some SRPMs incorporate dispersal
of habitat quality that can be used by habitat dependent related mortality into the movement submodel.
functions in the other submodels. A frustrating problem with SRPMs has been a lack of
A common approach to representing landscapes uses movement data.30 Movement data should be collected such
a hexagonal grid to define landscape units. 25, 28, 32 A that parameter estimates can be related to habitat quality.
hexagonal structure is often adopted for its geometric Mark-recapture studies can yield some information on
simplicity, and because, unlike a rectangular or triangular animal movements, but the most efficient means of
grid, the distances between a polygon center and centers studying movement is radio-telemetry. Understanding
of all adjacent polygons are equal. This simplifying how conspecific interactions influence movement
assumption—that the spatial distribution and connectivity behavior would require detailed knowledge of a local
of habitat can be adequately represented by a grid— population.
obviously imposes some limitations on realism of the The demographic submodel defines the population
habitat submodel. The most significant limitations are that structure and deals with changes in population size over
(1) the number of hexagons sets the carrying capacity of time. SRPMs can be two-sex or one-sex. The population is
the entire landscape, (2) the hexagon size determines the often broken into stage classes, such as juvenile, subadult,
maximum local territory density, and (3) and habitat and adult.25 The birth and death of individuals are usually
quality is usually characterized by one only variable: an modelled as stochastic processes with the probabilities of
index calculated for each hexagon. nesting success or producing a certain number of offspring
or death being functions of local habitat quality.20, 28 The
 540 Wildlife–Habitat Relationships in Oregon and Washington

crux of the demographic submodel is developing these habitat quality and no data to relate the movement of owls
quantitative relationships between demographic to habitat quality. The movement data available to us only
parameters and habitat quality. described the minimum, maximum, and mean distances
The most important data for the demographic (with standard error of the mean) for dispersing juveniles.
submodel are those needed for estimates of stage-specific
mortality and maternity parameters. These data are The Habitat Submodel
obtained through mark-recapture studies and SOS uses a hexagonal grid to represent the spatial
observations on the number of offspring produced per distribution of owl habitat. Six parameters govern the
breeding female (or breeding pair), respectively. Ideally, translation of rasterized habitat maps to a hexagonal
demographic data would also provide information on the landscape: hexagon size, “territory expansion,” a “territory
nonbreeding portion of the population, e.g., floaters. threshold,” and the relative habitat values of three land
Demographic data should be collected such that parameter cover categories. The hexagon size corresponds to the
estimates can be related to habitat quality. minimum home range area of an owl pair. We assumed
The basic output of SRPMs is the same as other that the smallest home ranges on the Peninsula would exist
population models: the number of individuals over time. in the high-quality habitat of Olympic National Park. The
SRPMs generate an output that other populations’ models density of owl pairs in the low-elevation old-growth
do not: the spatial distribution of individuals over time. forests of the Park was estimated to be 0.08/100 ha.34 This
One can compare the likely outcomes of different habitat density is equivalent to an exclusive home range of 1,250
management strategies, which could be useful for ha/pair, and we equated the size of the exclusive home
optimizing land management.18 An optimal strategy might range to the minimum home range. The minimum home
be one that maintains a desired population density for range is probably not equivalent to the exclusive home
the minimum cost. range, but theoretically, in a sparsely populated region
In general, SRPMs are not mathematically composed of high-quality habitat it should be nearly so.
sophisticated, but they are computationally complex. Well- The classified Landsat TM data38 had eight discrete
honed computer programming skills are indispensable for forest categories, but only three categories (old-growth,
tackling the task of model development. There are no large saw, and small saw) were assumed to have value as
standards for model structure, but modular, top-down, spotted owl habitat—an assumption based on Forsman
object-oriented programming is highly recommended. For et al.13 and Carey et al.4 A habitat utilization index (HUI)
most applications, a spatially realistic, individual-based was calculated for each forest category. It is defined by a
model will require large computer memory and huge on- ratio: (percent of owl relocations within a category)/
line data storage because: (1) information on every living (percent of home range area in that category).
individual in the population is maintained during a HUI describes habitat preferences. If HUI is greater than
simulation, and (2) GIS data, particularly rasterized data, 1.0, then owls expressed a preference for that forest
can occupy gigabytes of nonvolatile memory. High CPU category. The ratio of two HUI quantifies the relative
speeds are also very desirable. habitat value of two forest categories. HUIs were
calculated using radio-telemetry data collected from 20
An Example: The Spotted Owl Simulator owl home ranges on the western Olympic Peninsula (E.
The Spotted Owl Simulator6, 31 was developed to evaluate D. Forsman, U. S. Forest Service, unpublished data). The
the effects of different habitat management strategies on shortcomings of indices based on habitat use versus
the spotted owl subpopulation of the Olympic Peninsula. habitat availability were known to us,21, 37 but the data
With respect to the abundance and richness of data, the available to us precluded any other approach.
Olympic Peninsula was an excellent place to attempt The hexagon habitat scores were calculated with the
analyses using a SRPM. The Peninsula’s owl following equation:
subpopulation has been the subject of intensive study and
HS = HVOGAOG + HVLSALS + HVSSASS
the results of numerous investigations were available.4, 10,
11, 12, 23, 26, 33, 34
Demographic parameter estimates had been where HVX is the relative habitat value of either old-
published.3, 14 Habitat use versus habitat availability growth (OG), large-saw (LS), or small-saw (SS) forest, and
studies (E. D. Forsman, U.S. Forest Service, unpubl. data), AX is the proportion of a hexagon covered by that forest
and juvenile dispersal studies (E.D. Forsman, U. S. Forest category. Hexagons having scores above a threshold value
Service, pers. comm.) had been conducted. A gross are classified as potential owl territories. The threshold
estimate of population size had been issued in a draft value was determined by applying the same equation to
report.19 Data for habitat maps were also available. Landsat the 20 owl home ranges studied by Forsman, where AX
Thematic Mapper (TM) data for the western Peninsula, refers to the proportion of a home range. We found that
the actual planning area, had been classified into land setting the threshold value to the mean habitat score of
cover categories,38 and classified Landsat TM data also these 20 home ranges (which was less than the median
existed for the remainder of the Peninsula.27 score) yielded an acceptable spatial distribution of
Even with all this information, other essential potential territories when compared to the known
information was missing. There were no data that could locations of territorial owls on the Olympic Peninsula (E.
be used to relate mortality and maternity parameters to D. Forsman, U.S. Forest Service, unpubl. data; D.E.
 CHAPTER 22: MODELING—CASE HISTORIES 541

Seaman, National Biological Service, unpubl. data; S. P. The degree to which owl movements are guided by
Horton, DNR, unpubl. data). The minimum and first habitat quality is specified by the “Bias to Quality”
quartile scores each yielded a worse correspondence parameter. This parameter determines the frequency at
between predicted potential territories and known which owls move to the adjacent hexagon with the best
locations. habitat quality. If Bias to Quality equals 0.60, then owls
The maximum home range of an owl pair is determined move to the hexagon with the highest habitat score about
through an expansion parameter. This parameter 60 percent of their steps. If an owl does not consider habitat
represents two aspects of owl behavior. First, in areas of quality when moving, then the “Linearity” parameter
fragmented habitat, owls expand the size of their home specifies the probable direction of the next step. It
range, and second, the home ranges of neighboring owls determines the tendency to move in a forward direction.
may overlap.4 The expansion parameter represents the When Linearity equals zero, there is an equal probability
maximum amount of neighboring hexagons that may be of moving to any of the neighboring six hexagons. As
included in a home range. For 20 owl home ranges studied Linearity increases from zero, the simulated owls have a
on the western Olympic Peninsula (E. D. Forsman, U. S. greater tendency to move in a straight line. The values of
Forest Service, unpubl. data), the maximum home range Bias to Quality and Linearity were determined through
size was 11,248 ha, but the third quartile size was 6301 ha. parameter tuning (described below).
With a hexagon size of 1250, the maximum home range Little is known about the behavioral mechanisms that
that can be modeled is 8750 ha (a hexagon plus its six direct the movement of individual animals across
neighboring hexagons). The expansion parameter value landscapes. Hence, the strategy for developing the
was assigned its maximum value. The difference between movement submodel was to avoid parameters that are
the observed maximum home range and the maximum behavioral. This is contrary to the modeling strategy
size that can be modeled will cause the number of potential recommended by others5 who believe that parameters
territories to be underestimated. should be biologically meaningful. At least one parameter,
Hexagon “expansion” is a novel aspect of SOS. namely Bias to Quality, was needed to link habitat quality
Expansion does not actually increase the score or size of to movement. The other movement parameters—
hexagons. It is simply a mechanism that allows hexagons minimum and maximum number of steps and Linearity—
with low scores to function as territories by laying claim are basically statistical descriptors of the movement path.
to unallocated habitat present in adjacent hexagons. The Our current understanding is that males, females,
expansion of a sub-threshold hexagon continues until it juveniles, and adults probably exhibit different movement
reaches the potential territory threshold or until it reaches behaviors.15, 22 However, besides the minimum movement
the expansion limit. Only hexagons that are below the distance, all movement behavior was modeled the same
threshold expand, and hexagons with low scores require way, regardless of sex or stage class. A lack of data
more expansion to become potential territories. This motivated this simplification.
process does not affect the score of any hexagons. The No mortality occurs during movement. When juvenile
habitat submodel produces three types of hexagons: supra- owls are dispersing substantial mortality probably occurs,
threshold potential territory, sub-threshold potential and it presumably is a function of habitat quality and
territory, and not a potential territory. dispersal distance. But, having no data with which to
develop such a relationship, we elected to lump all death
The Movement Submodel into a single annual mortality.
The simulated landscape in SOS is a hexagonal grid. From
each location, owls can move in only six directions and T he Demographic Submodel
every move must be the same length, from hexagon center The Spotted Owl Simulator has a two-sex, three stage-class
to hexagon center. For hexagons of 1,250 ha, the “step” population structure. As with other spotted owl
size is about 3.80 km. Owls stop moving when a territory population models,3, 24 the stage classes are juvenile,
or mate is found, or a maximum number of steps have subadult, and adult. Estimates for demographic
been taken. parameters were available,14 but the crux was developing
The submodel requires specification of the minimum quantitative relationships between these demographic
number of steps that must be moved by dispersing parameters and habitat quality—a challenging task with
juveniles. Moving adults ignore the minimum movement no habitat data. The first step was to select the form of the
distance. This assumption was made because no data on relationships. A logistic function is intuitively appealing,
minimum movement distances exist for adults. The ratio but it requires five parameters to define completely. A
of total distance to net distance for owl dispersal equals logistic function may be approximated with a three-
about 2, 35 and the minimum net dispersal distance segment, piece-wise linear function that requires only four
observed for juveniles on the Peninsula is 8.67 km (E.D. parameters. We assumed that the least important domain
Forsman, U.S. Forest Service, unpubl. data). So, the of this function was the flat segment over lower quality
minimum number of steps equaled two times the net habitat. Hence, we eliminated this segment and adopted
dispersal distance divided by the mean step size, or about a two-segment, piece-wise linear function with only three
four hexagons. Likewise, the maximum net dispersal parameters. The shape of this function was based on the
distance observed for juveniles on the Peninsula is 58.25 plausible assumption that above some value of habitat
km, and so the maximum number of steps was set to 31.
 542 Wildlife–Habitat Relationships in Oregon and Washington

quality, survivorship (the complement of mortality) and Conclusions

maternity do not increase appreciably and can be Spatially realistic population models are an exciting new
considered constants. The same assumption was adopted technology in wildlife management, but the excitement is
by Holthausen et al.20 The value at which survivorship somewhat tempered by their extraordinary data
and maternity become constants is known as the requirements. Part of the excitement surrounding SRPMs
“saturation value” of habitat quality. undoubtedly arises from the array of powerful new
Adult and subadult stage classes had separate habitat technologies for the collection and analysis of spatial data:
dependent functions for the probability of death and the satellite thematic mapper imaging, global positioning
probability of nesting success. The values for the minimum systems, improved microelectronics for radio-telemetry,
and maximum parameters were determined through and geographic information systems. Yet, even with this
parameter tuning (described below). Juvenile survivorship technology, the extraordinary data requirements of most
was not a function of habitat quality because there is SRPMs are difficult to satisfy, and this has elicited some
considerable uncertainty about the value of this concerns about model reliability.1, 5, 8, 30, 39 Many of these
parameter.14 Instead, separate simulations were run using concerns can be assuaged by thorough explication of
different constant values of juvenile survivorship. These model assumptions, rigorous model evaluation,
runs represented a range of plausible scenarios from worst thoughtful interpretation of results, and cautious use of
case to best case, and were a form of sensitivity analysis the results for decision making. But, for those who choose
for juvenile survivorship. to develop a SRPM for analyzing their wildlife
management problems, grappling with insufficient data
Parameter Tuning remains a vexing challenge.
The values of some model parameters, such as fledgling Developing a SRPM will invariably require
sex ratio, came directly from published studies of spotted assumptions about the population or species autecology.
owls.13 The value of other model parameters, such as Assumptions will be necessary to circumvent insufficient
minimum number of steps for movement were calculated information or to make computer programming a practical
using available data. Unfortunately, SRPMs require types task, but each assumption or simplification made during
of information that are often difficult to acquire, such as model development can introduce poor judgement or bias.
the relationships between habitat quality and Nearly all other categories of models familiar to wildlife
demographic parameters. SRPMs also entail modeling biologists—habitat suitability indices, matrix models,
processes that are poorly understood, such as dispersal equations derived from statistical regression, etc.—have
behavioral. For these situations there were no data with well-established principles or methods for their
which to derive parameter values directly, but there were development, and much of their credibility as decision-
data that allowed us to iteratively search for values such making tools derives from the process of model
that the average characteristics of the simulated formulation that reduces subjective judgements. Likewise,
population approximated known characteristics of the real the credibility of a SRPM depends upon the handling of
population. This is parameter tuning. For example, to subjective judgements, i.e., assumptions, by the modeller.
determine values for the movement parameters, The most satisfactory approach to dealing with model
simulations were run on SOS with all other parameters assumptions is collecting the data needed to eliminate
assigned fixed values. Upon completion of a simulation, them. When this is impractical, each assumption should
statistical descriptors of the simulated dispersal paths were be explicitly stated and justified, and a thoughtful
compared to empirical data. Based on this comparison, consideration of its potential effects on model output
Bias to Quality and Linearity were adjusted, and the should be documented. The effects of each assumption
process was repeated until the mean net dispersal distance can be rigorously evaluated through sensitivity analyses.
was approximately equal to the mean observed on the Models which link habitat quality to population
Peninsula (24 km; E.D. Forsman, U.S. Forest Service, demographics are a relatively new tool. This may account
unpubl. data) and the ratio of mean net distance to mean for the scarcity of critical data needed by SRPMs. In fact,
total distance was approximately 2. To enable tuning of considering the needs of nongame species at large spatial
the dispersal parameters, mean net distance and mean scales is a relatively recent intellectual development in
total distance had to be model outputs. wildlife management. A critical mass of research in
To establish the relationship between demographic landscape ecology, albeit predominately conceptual or
parameters and habitat quality, a maximum value was theoretical, has been attained during the past fifteen
assumed using available information20 and the minimum years.2, 9, 17, 36 As is often the case, empirical research has
value was determined through parameter tuning. The not matched the pace of theoretical research, nor has it
minimum parameter value was iteratively adjusted until matched the pace of the technological developments that
the realized mean parameter value of the simulated have enabled SRPMs. Nevertheless, the outlook is
population was approximately those reported by Forsman promising. Awareness of important processes that take
et al.14 To enable tuning of the demographic parameters, place at the landscape scale has begun to direct field
adult and subadult mortality and mean maternity had to research in wildlife biology and data needed by SRPMs
be outputs of the model. will accumulate.
 CHAPTER 22: MODELING—CASE HISTORIES 543

Acknowledgments 18. Hof, J., and M.G. Raphael. 1997. Optimization of habitat placement: a
We gratefully acknowledge the reviewers of this paper, case study of the northern spotted owl in the Olympic Peninsula.
Ecological Applications 7:1160-1169.
Eric Forsman and John Dunning Jr., whose comments 19. Holthausen, R.S., M.G. Raphael, K.S. McKelvey, E.D. Forsman, E.E.
motivated major revisions. Timothy Quinn added Starkey, and D.E. Seaman. 1994. The contribution of federal and
considerable polish through his conscientious editing. We nonfederal habitat to persistence of the northern spotted owl on the
thank David Johnson for the opportunity to present our Olympic Peninsula, Washington. Draft Report of the Reanalysis Team,
work. U.S. Forest Service, Portland, OR.
20. ———, M.G. Raphael, K.S. McKelvey, E.D. Forsman, E.E. Starkey, and
Literature Cited D.E. Seaman. 1995. The contribution of federal and nonfederal habitat
1. Bart, J. 1995. Acceptance criteria for using individual-based models to to persistence of the northern spotted owl on the Olympic Peninsula,
make management decisions. Ecological Applications 5:411-420. Washington: report of the Reanalysis Team. U.S. Forest Service, General
Technical Report PNW-352.
2. Bissonette, J. A., editor. 1997. Wildlife and Landscape Ecology: effects of
patterns and scale. Springer-Verlag, New York, NY. 21. Hobbs., N.T., and T.A. Hanley. 1990. Habitat evaluation: do use/
3. Burnham, K.P., D.P. Anderson, and G.C. White. 1994. Estimation of vital availability data reflect carrying capacity? Journal of Wildlife
Management 54:515-522.
rates of the northern spotted owl. Appendix J, pp. 1-26 in Final
supplemental environmental impact statement on management of 22. Johnson, M.L., and M.S. Gaines. 1990. Evolution of dispersal:
habitat for late-successional and old-growth forest related species theoretical models and empirical tests using birds and mammals.
Annual Review of Ecology and Systematics 21:449-480.
within the range of the northern spotted owl. USDA Forest Service,
Portland, OR. 23. Lehmkuhl, J.F., and M.G. Raphael. 1993. Habitat pattern around
4. Carey, A.B., S.P. Horton, B.L. Biswell. 1992. Northern spotted owl: northern spotted owl locations on the Olympic Peninsula, Washington.
Journal of Wildlife Management 57:302-315.
influence of prey base and landscape character. Ecological Monographs
62:223-250. 24. McKelvey, K. 1992. A spatially explicit life-history simulator for the
5. Conroy, M.J., Y. Cohen, F.C. James,Y.G. Matsinos, and B.A. Maurer. 1995. northern spotted owl. Appendix 4-P in Eugene District Resource
Management Plan & EIS. U.S. Department of Interior, Bureau of Land
Parameter estimation, reliability, and model improvement for spatially
explicit models of animal populations. Ecological Applications 5:17-19. Management, Eugene, OR.
6. Department of Natural Resources. 1996. Draft Environmental Impact 25. ———, B.R. Noon, and R.H. Lamberson. 1993. Conservation planning
for species occupying fragmented landscapes: the case of the northern
Statement: Habitat Conservation Plan. Washington State Department
of Natural Resources, Olympia, WA. spotted owl. Pages 424-450 in P.M. Kareiva, J.G. Kingsolver, and R.B.
7. ———. 1997. Final Habitat Conservation Plan. Washington State Huey, editors. Biotic Interactions and Global Change. Sinauer
Associates Inc., Sunderland, MA.
Department of Natural Resources, Olympia, WA.
8. Doak, D.F., and L.S. Mills. 1994. A useful role for theory in conservation. 26. Mills, L.S., R.J. Fredrickson, and B.B. Moorhead. 1993. Characteristics
Ecology 75:615-626. of old-growth forests associated with northern spotted owls in the
Olympic National Park. Journal of Wildlife Management 57:315-321.
9. Forman, R.T.T., and M. Godron. 1986. Landscape Ecology. John Wiley
and Sons, New York, NY. 27. PMR. 1993. Washington State forest cover classification and
10. Forsman, E.D. 1990. Habitat use and home range characteristics of cumulative effects screening for wildlife and hydrology. Pacific Meridian
Resources, Portland, OR.
spotted owls on the Olympic Peninsula, Washington. pages numbers
unknown in Wildlife habitat relationships in western Washington and 28. Pulliam, R.H., J.B. Dunning, and J. Liu. 1992. Population dynamics in
Oregon. PNW Annual Report—Fiscal Year 1990. USDA Forest Service, complex landscapes: a case study. Ecological Applications 2:165-177.
29. Raphael, M.G., J.A. Young, K. McKelvey, B.M. Galleher, and K.C. Peeler.
Pacific Northwest Research Station, Wildlife Ecology Team, Olympia,
WA. 1994. A simulation analysis of population dynamics of the northern
11. ———. 1991. Habitat use and home range characteristics of spotted spotted owl in relation to forest management alternatives. Final
environmental impact statement on management of habitat for late-
owls on the Olympic Peninsula, Washington. pages 12-16 in Wildlife
habitat relationships in western Washington and Oregon. PNW Annual successional and old-growth forest related species within the range of
Report—Fiscal Year 1991. USDA Forest Service, Pacific Northwest the northern spotted owl. Vol. 2, Appendix J-3. U.S.D.A. Forest Service,
Portland, OR.
Research Station, Wildlife Ecology Team, Olympia, WA.
12. ———. 1992. Demographic characteristics of spotted owls on the 30. Ruckelshaus, M., C. Hartway, and P. Kareiva. 1997. Assessing the data
Olympic Peninsula, Washington, 1987-1992. pages 18-27 in Wildlife requirements of spatially explicit dispersal models. Conservation
Biology 11:1298-1306.
habitat relationships in western Washington and Oregon. PNW Annual
Report—Fiscal Year 1992. USDA Forest Service, Pacific Northwest 31. Schumaker, N.H. 1995. Habitat Connectivity and Spotted Owl
Research Station, Wildlife Ecology Team, Olympia, WA. Population Dynamics. Dissertation. University of Washington, Seattle,
WA.
13. ———, E. C. Meslow, and H. M. Wight. 1984. Distribution and biology
of the spotted owl in Oregon. Wildlife Monographs 87:1-64. 32. ———. 1998. A user’s guide to the PATCH model. U.S. EPA/600/R-
14. ———, S. G. Sovern, E. D. Seaman, K.J. Maurice, M. Taylor, and J.J. Zisa. 98/135, Environmental Protection Agency. Environental Research
Laboratory, Corvallis, OR.
1996. Demography of the northern spotted owl on the Olympic
Peninsula and east slope of the Cascade Range, Washington. In E.D. 33. Seaman, D.E., R.J. Fredrickson, D.B. Houston, B.B. Moorhead, and R.A.
Forsman, S. DeStefano, M.G. Raphael, and R.J. Gutierrez, editors. Hoffman. 1992. Northern spotted owl inventory, Olympic National
Park. Unpublished progress report.
Demography of the Northern Spotted Owl. Studies in Avian Biology,
vol. 17. Cooper Ornithological Society, Camarillo, CA. 34. ———, S. A. Gremel, S. L. Roberts, and D. W. Smith. 1996. Spotted
15. Greenwood, P.J., and P.H. Harvey. 1982. The natal and breeding Owl Inventory-Monitoring in Olympic National Park, Final Report.
National Biological Service and Olympic National Park, Port Angeles,
dispersal of birds. Annual Review of Ecology and Systematics. 13:1-21.
16. Hanski, I., and D. Simberloff. 1997. The metapopulation approach, its WA.
history, conceptual domain, and application to conservation. Pages 5-26 35. Thomas, J.W., E.D. Forsman, J.B. Lint, E.C. Meslow, B.R. Noon, and J.
Verner. 1990. A conservation strategy for the northern spotted owl.
in I. Hanski and M.E. Gilpin. Metapopulation Biology: Ecology, Genetics,
and Evolution. Academic Press, San Diego, CA. Report of the Interagency Scientific Committee to Address the
17. Harris, L.D. 1984. The Fragmented Forest: Island Biogeography Theory Conservation of the Northern Spotted Owl. Appendix M. U.S. Forest
Service, Portland, OR.
and the Preservation of Biological Diversity. University of Chicago
Press, Chicago, IL.
 544 Wildlife–Habitat Relationships in Oregon and Washington

36. Turner, M.G. 1989. Landscape ecology: the effect of pattern on information, for example, on structural conditions and
process. Annual Review of Ecology and Systematics 20:171-197. specific habitat features used by species, and spatial
37. Van Horne, B. 1983. Density as a misleading indicator of habitat relations of species including home range size class and
quality. Journal of Wildlife Management 47:893-901.
38. WDW. 1993. Experimental forest classification project. Unpublished
use at a landscape scale. These matrixes are provided for
report to the Washington Department of Natural Resources, species found in Washington and Oregon in the CD-ROM
Washington Department of Wildlife, Olympia, WA. included in this book. The distribution of each species also
39. Wennergren, U., M. Ruckelshaus, and P. Kareiva. 1995. The promise and needs to be identified for the analysis area.
limitations of spatial models in conservation biology. Oikos 74:349-356.
Step 2. Group species into guilds based on known
habitat use at the stand and landscape scales.
Case Study 2 HABSCAPES was designed to use a guild approach,
where all vertebrate species are grouped by similar uses
A Model to Assess Potential Vertebrate of resources.3 To assign a species to a single guild, we first
Habitat at Landscape Scales: divided all of the vertebrate species in our analysis area
HABSCAPES (Mt. Hood National Forest) into three groups: special and
Mark Huff,T. Kim Mellen & Rich Hagestedt unique habitat obligates (e.g., scree slopes or springs and
seeps), riparian habitat obligates, and terrestrial habitat
users. Only species belonging to the riparian habitat
Introduction obligate and terrestrial habitat groups were placed in
Our objective was to develop a model and analysis guilds; only the terrestrial guild is presented here. The
procedure targeted for land managers that (1) assesses the special and unique habitat obligate species were difficult
amount and distribution of “potential” habitat of all to place into groups and therefore did not fit well into our
terrestrial vertebrates and aquatic amphibians relative to guild approach. In addition to guilds, HABSCAPES can
large landscape patterns (e.g., 1 million acres) and (2) be used to assess potential habitat of individual species
provides options for developing maps and databases to using specific habitat and spatial attributes.
display and analyze “potential” habitat distribution Guilds for the terrestrial habitat group were based on
patterns. Our modeling approach, termed HABSCAPES, combinations of three attributes: home range size,
takes the user’s spatially explicit vegetation data (i.e., landscape-use categories, and structural habitat
vegetation maps linked to electronic coordinates), which conditions. Home range sizes were divided systematically
have been classified into mapped areas for wildlife habitat into three categories: small (<60 acres [24 ha]), medium
purposes, and links it to stand- and landscape-level (60-1,000 acres [24-400 ha]), and large (>1,000 acres [400
species-habitat relations and life history information. We ha]). These categories were selected based on observed
used the life history information as a surrogate for detailed gaps in home range sizes for species reported in the
demographic data that are largely unavailable for most literature and on the size of management units within the
vertebrate species in any given planning area. We planning area. Landscape-use categories were assigned
developed a series of computer programs and queries to to each species based on interpreting the literature on how
access this information, and to determine distributions of species use habitat within their home range and
suitable habitat for individual species and groups of professional judgment. Four categories were developed:
species (hereafter, guilds) with similar habitat needs patch (tend to use one homogeneous habitat patch),
relative to their spatial requirements (e.g., home range mosaic (aggregate patches of homogeneous habitat),
area). The results are spatially referenced in a database contrast (use of patches of two different structural stages
and stored for visual display and further analyses. in close proximity), and generalist (use a variety of
structural stages). The third attribute, structural habitat
Methods conditions, was derived from the habitat relations
Development and use of HABSCAPES follows the five information in the matrices, but simplified considerably
general steps described below. Two separate databases are into three categories: open, small tree (8-21 inches [20-53
needed to implement HABSCAPES: a vertebrate habitat cm] dbh), or large tree (> 21 inches [53 cm] dbh) conditions.
relations database (Step 1) and a spatially-referenced In developing HABSCAPES using Mt. Hood National
vegetation database with sufficient information to Forest data, we had 36 possible terrestrial guilds
characterize vegetation features into habitat conditions for (combinations of 3 home range, 4 landscape use, and 3
animal species (Step 3). HABSCAPES version 1.0 operates structural habitat categories); species were represented in
in a DOS environment for 486 MHz or higher, IBM 18 of the possible guilds.
compatible PCs. In May 2000, HABSCAPES version 2.0
was completed to run in an ARC/INFO platform. Step 3. Develop a habitat relations-based classification
of vegetation using composition and structure. The
Process vegetation database stores the spatially referenced
Step 1. Define vertebrate stand-level habitat information on which habitat relationship assessments are
relationships and distribution patterns. A series of made. At a minimum, these spatially explicit data
species-habitat and life history matrixes1, 7 have to be provided by the user should be organized into
developed for each species. These matrixes organize composition-classified communities (e.g., Pinus ponderosa
 CHAPTER 22: MODELING—CASE HISTORIES 545

series) and into successional or structural development neighborhood analysis that aggregates like pixels together
(e.g., late seral, open canopy) types. Vegetation must be based on the user’s rules (e.g., minimum patch size for
classified accurately. At large spatial scales these data are given set of objectives and analysis requirements and the
usually obtained from interpreted aerial photographs type and amount of adjoining habitat surrounding each
and/or satellite imagery and stored as raster-based (pixel) pixel).5 PATCH provides the user with a consistent and
maps. The attributes of the vegetation (e.g., canopy cover) repeatable method for delineating portions of a map as
are stored in a database with specific variables that are potential habitat. The output from the PATCH program is
spatially linked to vertebrate species distribution (e.g., a spatially-referenced database used to create individual
range maps). As we developed HABSCAPES, county “patch” maps for specific sets of conditions. In developing
boundaries were used to delineate distribution (i.e., HABSCAPES, for example, we created separate patch
potential occurrence) for vertebrates. To complete the maps for three structural states (open, small tree, and large
linkage to vegetation; county boundaries were integrated tree structural habitats) that were used as the basis to
into the classified vegetation database. determine suitable habitat for different guilds.
Step 4. Delineate groups of like vegetation pixels into Step 5. Evaluate suitability of patches for different guilds
patches. Interpretation of vertebrate habitat relations is using landscape-level characteristics and habitat
done usually at the patch or “patchiness” scale. To make relationships. Habitat suitability can be assessed for each
predictions of suitable habitat for different species and guild once patch maps are made. Suitability classes are
guilds, patches need to be created. Typically, patches are assigned using programs that evaluate patch size,
areas of relatively homogeneous conditions (e.g., tree neighborhood relationships of patches, and the amount
sizes), with fixed boundaries and delineated for a specific of area within fixed home ranges for each guild (or species)
purpose other than animal habitat relations. To provide that have similar patch-vegetation characteristics.
users of HABSCAPES with flexibility to delineate their Examples of the parameters we used for our guild-level
own patches for determining suitable habitat, we analysis are shown in Table 1.
developed a program called PATCH that aggregates The generalist guild is the easiest one to interpret for
habitat-related data stored at the pixel scale into patches. potential habitat: all vegetation patches are considered
PATCH “grows” patches through a pixel by pixel suitable. For the guilds with species confined to a single

Table 1. Parameters used to run HABSCAPES programs for each guild.

Blanks indicate parameter was not needed to assess habitat for the guild.

Guild Home Patch Structure Min. % Home Total Buffer % total

Code1 Range Configuration Stage Patch Range in Adjacent Width habitat of
Size Type Size Habitat Habitat2 each type3

TSPO 52 (Small) Patch Open 20 50

TSPST 52 (Small) Patch Small tree 20 50
TSPLT 52 (Small) Patch Large tree 20 50
TSMO 52 (Small) Mosaic Open 4.8 50 30
TSMST 52 (Small) Mosaic Small tree 4.8 50 30
TSGOS 52 (Small) Generalist Open/small tree 4.8 50 30
TSGSL 52 (Small) Generalist Small/large tree 4.8 50 30
TSGG 52 (Small) Generalist Generalist 4.8
TMPO 1000 (Med) Patch Open 500 50
TMMO 1000 (Med) Mosaic Open 20 50 700
TMMLT 500 (Med) Mosaic Large tree 20 50 350
TMGG 500 (Med) Generalist Generalist 4.8
TLMO 3000 (Large) Mosaic Open 40 50 2100
TLMLT 3000 (Large) Mosaic Large tree 40 50 2100
TLGG 3000 (Large) Generalist Generalist 4.8
TSC 52 (Small) Contrast Open/large tree 4.8 20 100 >=25
TMC 500 (Med) Contrast Open/large tree 10 20 200 >=25
TLC 3000 (Large) Contrast Open/large tree 20 20 400 >=25
1
Guild codes all begin with “T” to indicate terrestrial guilds. The remainder “first letter” code sequentially indicates home range size
(S, M, and L), patch configuration(P, M, G, and C) and structure stage (O, ST, and LT); contrast guild codes do not include a structure
stage.
2
Used for mosaic guild only to determine Adjacent-Neighborhood habitat. Acres are 70% of home range size.
3
Used for contrast guilds only; a type 1 and type 2 contrast habitat must each be at least 25% of total contrast to ensure adequate
contrasting environments on the landscape.
 546 Wildlife–Habitat Relationships in Oregon and Washington

patch (patch species), suitable habitat is assessed by minimum patch size criteria but does not contribute to
determining if a patch created by the PATCH program the suitability of another patch; and (7) unsuitable small
exceeds the designated minimum patch size for the given patch, does not meet the minimum patch size.
structure conditions. We developed specific programs to The CONTRAST program assesses the distribution of
assess potentially suitable habitat for groups of species contrasting habitats within a home range by merging two
that tend to aggregate multiple patches together to make different (structure stage) patch maps (e.g., mature forest
suitable habitat, and groups of species that tend to adjacent to open/early successional conditions classified
aggregate patches based on contrasting conditions; they together as a patch). Rules are defined by the user for the
are termed Mosaic and Contrast species, respectively. The width, amount, and proportions of contrast conditions
landscape-level habitat relationships for the different between the different patch types. The program produces
guilds of Mosaic and Contrast species are conceptual and four potential outcomes of patches that are suitable
based on the limited information available for the few contrasts or contribute to making a patch a suitable
species studied at landscape scales. Consequently, much contrast.
professional judgment was needed to build these types of
models.5 Examples
The SUIT program assesses habitat suitability for the Below we illustrate uses of HABSCAPES using the
Mosaic species (guilds) by centering a guild-defined, home programs SUIT, PATCH, and CONTRAST, and
range-sized circle on each pixel in each patch. For each demonstrate guild and single species approaches.
pixel, a series of menu-based questions are asked (i.e., set
of analysis rules) to determine the amount and distribution Bull Run Watershed Analysis
of patches within the circle.5 This process has seven HABSCAPES was used in the Bull Run Watershed
possible outcomes: (1) unsuitable patch, not the correct Analysis on the Mt. Hood National Forest to assess
structural stage; (2) suitable large patch, a single patch that potential change in amount of habitat for different
is larger than some specified portion of the guild home terrestrial guilds between historic and current conditions.
range size (e.g., 50%); (3) suitable aggregated patch, a Only guilds with species occurring in the watershed were
habitat patch which meets minimum patch size and which analyzed. The historic vegetation condition was based on
falls within a home range size area of which some specific 1940 county forest surveys. The survey provided stand
portion (e.g., 50%) consists of suitable habitat; (4) suitable structure information prior to most timber management
adjacent-neighborhood patch, a habitat patch which meets activities.
minimum patch size and is made suitable when patches All three HABSCAPES programs were used for the
on the edge of the home range extend beyond the home terrestrial guild assessment. See Table 1 for guild codes
range size are combined with patches within the home and the parameters used for each guild. The assessment
range size meet some specific portion of suitable habitat showed declines in habitat for TSPLT, TMMLT, and
(e.g., 70%); (5) unsuitable isolated-contributing patch, a TLMLT guilds (Table 2). All these guilds use large tree
patch which meets the minimum patch size criteria but is habitat. Consequently, the guild analysis was followed up
too isolated from other patches to be aggregated into with an analysis of two individual species of concern
suitable habitat and does contribute to making another within guilds experiencing declining habitat, northern
set of patches suitable; (6) unsuitable isolated patch, meets spotted owl and red tree vole. The other guild showing a

Table 2. Historic and Current Habitat Available for Terrestrial Guild

Guild Home Patch Structure Historic % Current % % Change from

Code Range Association Stage ofWatershed ofWatershed Historic Habitat

TSPO Small Patch Open 6 13 +117

TSPLT Small Patch Large tree 58 31 -46
TSMO Small Mosaic Open 6 13 +117
TSGOS Small Generalist Open/small tree 35 65 +86
TSGSL Small Generalist Small/large tree 59 83 +41
TSGG Small Generalist Generalist 95 98 +3
TMMO Medium Mosaic Open 5 9 +125
TMMLT Medium Mosaic Large tree 20 13 -35
TMGG Medium Generalist Generalist 95 98 +3
TLMO Large Mosaic Open 5 2 -60
TLMLT Large Mosaic Large tree 58 28 -52
TLGG Large Generalist Generalist 95 98 +3
TSC Small Contrast Open/large tree 2 7 +250
TMC Medium Contrast Open/large tree 4 9 +125
TLC Large Contrast Open/large tree 6 14 +133
 CHAPTER 22: MODELING—CASE HISTORIES 547

decline in habitat was associated with habitat of large open habitat.9 Specific analyses was done on the northern
areas (TMLO), however, this habitat was rare even spotted owl, a threaten species belonging to the TLMLT
historically. The guild analysis also revealed that habitat guild. Analysis attributes for locating suitable nesting and
increased substantially for the following guilds: TSPO, foraging habitat on Mt. Hood National Forest included
TSMO, TSGOS, TSGSL, TMMO, TSC, TMC, and TLC patches below 5,000 feet (1,610 m) elevation with >60%
(Table 2). canopy closure and trees >21 inch dbh or a mix of trees
>21 and 8-21 inches (13-53 cm) dbh. Patches with greater
North Willamette Late Successional Reserve than 40% canopy closure and a mix of trees 5-21 inches
(LSR) Assessment dbh were classified as suitable dispersal habitat. Other
The Record of Decision for Amendments to Forest Service analysis parameters specific to the 1990 guidelines to avoid
and Bureau of Land Management Planning Documents a “taking”11 were home range size of 2,995 acres (1198 ha),
within the Range of the Northern Spotted Owl (ROD) minimum patch size of 40 acres (16 ha), and minimum
(USDA and USDI, 1994) required the two agencies to acres of suitable habitat within a home range of 40% (1,182
develop LSR Assessments. The following two examples acres [474 ha]). The “Large Patch” polygons shown in Map
come from the North Willamette LSR Assessment.10 1 from Program SUIT represent the largest, least
fragmented blocks of suitable habitat. “Aggregated
Northern Spotted Owl Habitat. HABSCAPES was used
Patches” are more fragmented but still represent areas
to evaluate the current distribution, amount and quality
where at least 40% of the home range area is in suitable
of late-successional habitat within and outside LSRs. The
habitat. Other patch types do not meet this 40%
mosaic guild representing species with large home ranges
assumption. The majority of the “Large Patch” polygons
and associated with late-successional habitat (TLMLT) was
are within LSRs or wilderness areas. Additional “Large
used as a “first screening” of potential late-successional

Large patch
Aggregated patch
Adjacent neighborhood patch
Unsuitable patch
Critical habitat units
Late successional reserves
Wilderness
Map 1. North
Willamette LSRA
Spotted Owl Habitat
Quality
HABSCAPES
Assessment.
 548 Wildlife–Habitat Relationships in Oregon and Washington

Primary reproductive habitat

Primary habitat
Potential habitat
RTV areas of concern
Late successional reserves
Wilderness
Map 2. North Willamette LSRA Red Tree
Vole Habitat and Connectivity Areas of
Concern.

Patch” polygons occur in the northern part of the the ROD for the Northwest Forest Plan, the red tree vole
assessment area due to differences in the definition of was identified as a Survey and Manage Species requiring
suitable habitat. There are large amounts of “Aggregated surveys before initiating “ground disturbing”
and Adjacent-Neighborhood Patches” habitat outside management activities.8 Because of these circumstances,
these protected areas. Dispersal habitat was added to the a suitable habitat assessment was done for red tree voles
map after suitable habitat was run through the model. as part of the LSR assessment on Mt Hood National Forest.
The condition of spotted owl habitat in Critical Habitat The PATCH program in HABSCAPES was used to
Units (CHUs) was also a management concern. In the assess amount and distribution of red tree vole habitat.
western Oregon Cascade Range, overlap is low between Habitat descriptions and model parameters were based
LSRs and CHUs,8 and more site-specific analysis was on Huff et al.4 Red tree voles are most abundant in late-
needed for Mt. Hood National Forest. Output from successional forests; however, they do occur in younger
HABSCAPES was compared to assess the quality of conifer forest with a closed canopy,2 which provides
habitat between LSRs/wilderness areas and CHUs based dispersal habitat. Habitat was divided into primary
on distribution of patch types (Map 1). There were more reproductive, late-successional patches >100 acres (40 ha);
acres of “Large and Aggregated Patches” in CHUs than primary other than reproduction, late-successional patches
in LSRs/wilderness areas primarily because of more total of any size; and potential habitat, patches with closed-
acres in the CHUs, however, the proportion of habitat in canopy conifers and tree diameters 8-21 inches. Forests
the different patch categories was similar between CHUs above about 3,000 feet (970 m) elevation were considered
and LSRs/wilderness areas. The proportion of suitable unsuitable habitat.
habitat in large, contiguous patches was highest in areas Suitable red tree vole habitat identified using
where CHUs and LSRs/wildernesses areas overlap, HABSCAPES was limited primarily by elevation (Map 2).
indicating that areas with the least amount of fragmented High elevations create a barrier to the vole movement
habitat are covered by both designations. between the Salmon-Huckleberry Wilderness and the
Roaring River LSR in the north-central portion of the
Red Tree Vole. The red tree vole is a species of
assessment area. The HABSCAPES maps helped to
management concern because of its close association with
identify areas to sustain connectivity for the voles at lower
late-successional forests, it lives almost exclusively in the
elevations to the west and for four areas to the south
canopy of conifers, and its need for habitat connectivity
between LSRs.
in landscapes because of limited dispersal capability.8, 9 In
 CHAPTER 22: MODELING—CASE HISTORIES 549

Map 3. Large Home Range Contrast Guild,

Mt. Hood National Forest

Open contrast habitat

Large tree habitat
Open contributing habitat
Large tree contributing habitat

Large Home Range, Contrast Guild (TLC) Conclusions

The CONTRAST program in HABSCAPES was used to HABSCAPES is a “coarse filter” approach to assess
create a map of habitat for the TLC guild, large tree and potential habitat and to develop conservation strategies
open habitat within 400 m of each other.12 Species in this for many species simultaneously. HABSCAPES assists
guild include elk, great horned owl, and red-tailed hawk. biologists and resource managers with complex multi-
Minimum patch size was set at 20 acres (8 ha), percent scale assessments, such as watershed analyses required
habitat in the home range needed to be at least 20 percent, in the Northwest Forest Plan. Our approach of grouping
and each type of contrast habitat needed to contribute at species into similar habitat guilds, which considers
least 25 percent to total contrast habitat (based on species’ spatial requirements, and of providing suitability
professional judgement) to ensure adequate contrast indices of potential habitat gives biologists a tool to make
between seral stages. The results (Map 3) indicated where relatively quick habitat assessments for large numbers of
the best potential elk habitat could be found on the Mt. vertebrate species in a planning area.
Hood National Forest. Although not shown, separate HABSCAPES is versatile, as shown from our case
queries on winter and summer range attributes can done studies and assessments of individual species and guilds
and overlain together to assess the relations of seasonal that addressed many natural resource planning issues.
habitat patterns. Mapping the TLC guild also predicts HABSCAPES provides users with options to create
potential great horned owl habitat. This map could be patches based on specific objectives, and to change patch
compared with the map of suitable spotted owl habitat to shape, size, and composition for different objectives using
identify potential places where predation of great horned program PATCH. As PATCH aggregates like pixels into
owls on spotted owls6, 11 is probable. patches, it stores information about patches at the pixel
level, thus giving the user flexibility to assign a pixel to as
many patches as warranted. It is the user who defines the
 550 Wildlife–Habitat Relationships in Oregon and Washington

rule sets for the analysis programs PATCH, SUIT, AND 9. ———, and ———. 1994. Record of Decision for Amendments to
CONTRAST and for the queries to determine potential Forest Service and Bureau of Land Management planning Documents
habitat. Within the Range of the Northern Spotted Owl, and Standards and
Guidelines for Management of Habitats for Late Successional and Old
Tradeoffs have to be made to assess multi-species issues Growth Forest Related Species Within the Range of the Northern
at large landscape scales, as done with HABSCAPES. Spotted Owl. Forest Service, Bureau of Land Management. Portland,
Generalizations are made about species habitat relations OR. 3 vols.
and spatial requirements to fit groups of species into 10. ———, and ———. 1998. North Willamette LSR Assessment. Mt.
guilds. The amount of habitat for certain species Hood National Forest, Sandy, OR. Unpublished report.
unknowingly can be over- or under-estimated, especially 11. USDI Fish and Wildlife Service. 1992. Recovery plan for the northern
spotted owl—draft. Portland, OR.
for those species that are closely associated with specific
12. Wisdom, M.J., L.R. Bright, C.G. Carey, W.W. Hines, R.J.Pedersen, D.A.
habitat features that are not evaluated. Therefore, the guild Smithey, J.W. Thomas, and G.W. Witmer. 1986. A model to evaluate elk
approach adopted in HABSCAPES should not be used to habitat in western Oregon. Pub. No. R6-F&WL-216-1986, USDA Forest
evaluate population and species viability issues associated Service, Pacific NW Region, Portland, OR.
with habitat. Further, HABSCAPES projections of suitable
habitat are dependent on estimates of minimum home
range sizes, however many species have not been studied Case Study 3
sufficiently for different habitats and life history functions. Cross-scale Classification Trees for
Currently few tests have been done to validate the
HABSCAPES suitable habitat predictions; though, we did
Assessing Risks of Forest Practices to
examine how well spotted owl nest occurrence Headwater Stream Amphibians
corresponded with suitable habitat projects. We placed the Glenn D. Sutherland & Fred L. Bunnell
known spotted owl nest sites for Mount Hood National
Forest over habitat suitability types in Map 1. Sixty-five
Introduction
percent of the nests were concentrated in the “most”
Because organisms respond to environmental conditions
suitable types for spotted owls, while 22 and 18% were in
and processes operating at different scales, extrapolation
other suitable habitat patch types and dispersal habitat,
of specific research findings to guide general forest
respectively. We expected a higher percentage of nests in
practices can be misleading.3 We illustrate the use of
the most suitable habitat; we suspect it will be higher as
classification and regression tree (CART) methods to
the vegetation database is refined (e.g., narrower tree
reveal relationships occurring at different scales when only
diameter classes).
inventory data are available. Our primary objectives are
Literature Cited to: 1) identify the range of habitat attributes permitting
1. Brown, E.R., technical editor. 1985. Management of wildlife and fish the species presence; 2) reveal potential differences in
habitats in forests of western Oregon and Washington. Part 2— habitat relationships within the species range; and 3)
Appendices. Publ. R6-F&WL-192-1985. U.S. Department of Agriculture, identify sites where conservation measures would be most
Forest Service, Portland, OR. effective. We chose the tailed frog for five reasons: 1) it is
2. Carey, A.B. 1991. The biology of arboreal rodents in Douglas-fir designated “at risk” or “of special concern” in most of its
forests. In Biology and management of old-growth forests. M.H. Huff, range; 2) available data are contradictory, suggesting that
R.S. Holthausen, and K.B. Aubry (technical coordinators). PNW-GTR-
276. USDA Forest Service, Pacific Northwest Research Station,
broader scale phenomena are acting;2, 5, 11, 14 3) generalized
Portland, OR. conservation measures could have enormous impact on
3. Hale, W.G., and J.P. Margham. 1991. The HarperCollins dictionary of timber harvest; 4) similar approaches can be developed
biology. HarperCollins Publishers, New York, NY. for other species in headwater streams; and 5) given the
4. Huff, M.H., R.S. Holthausen, and K.B. Aubry. 1992. Habitat management number of studies on the species, it is worthwhile
for red tree voles in Douglas-fir forests. In: Biology and management of developing an approach that permits analysis of
old-growth forests. M.H. Huff, R.S. Holthausen, and K.B. Aubry
geographically scattered data.3
(technical coordinators). PNW-GTR-302, USDA Forest Service, Pacific
Northwest Research Station, Portland, OR.
5. Mellen, K., M. Huff, and R. Hagestedt. 1995. “HABSCAPES” Interpreting
Methods
landscape patterns: a vertebrate habitat relationships approach. Pages Bunnell et al.4 interpreted the disparate results of studies
135-144 in Thompson, J.E., compiler. Analysis in support of ecosystem evaluating impacts of forest practices on tailed frogs as a
management. Analysis workshop III, April 10-13, 1995, Fort Collins, function of different physical settings. Common
Colorado. USDA Forest Service, Ecosystem Manage. Cntr., Washington, approaches to describing environmental influences on
D.C. species response variables—multivariate habitat models
6. Miller, G.S. 1989. Dispersal of juvenile northern spotted owls in
western Oregon. M.S. thesis, Oregon State University, Corvallis, OR
(e.g. logistic or multiple regression)—can be misleading
7. Thomas, J.W., editor. 1979. Wildlife habitats in managed forests: the Blue if different factors limit a species in different portions of
Mountains of Oregon and Washington. Washington DC: U.S. its range, or if the interactions between variables are non-
Department of Agriculture, Forest Service. linear. Moreover, spatial autocorrelation within and
8. USDA Forest Service and USDI Bureau of Land Management. 1994. between environmental variables can alter or disguise the
Final supplemental environmental impact statement on management of apparent response.8
habitat for late successional and old growth forest related species
First, we developed empirical multivariate species-
within the range of the northern spotted owl. Forest Service, Bureau of
Land Management. Portland, OR. 3 vols.
habitat models from extensive inventory data, unstratified
 CHAPTER 22: MODELING—CASE HISTORIES 551

by management activities or ecological gradients, using risks to the species from management activities. Combined
CART analysis. CART models create hierarchical trees by with spatial analyses they can help to identify the
recursive partitioning of sets of numeric or categorical appropriate spatial scale for applying management
habitat predictor variables into mutually exclusive subsets, guidelines. The approach also provides probability
which are most homogeneous with respect to a response rankings that help to classify new sites where only some
variable (species occurrence or abundance).1 Each step predictor variables have been measured, and to identify
finds the variable most important in reducing remaining potential classifications for sites in areas not yet surveyed.
variation, dependent on all previous steps. The output tree Once such empirically-based habitat models are
diagram represents a nested set of ecological dependencies developed, they can be used to make predictions in two
among habitat factors, exposing how key environmental ways. Using the original data set, the likelihood of any
variables can act to constrain the ranges of other variables, rule in the tree applying in similar environmental
given the observed species response. That permits conditions elsewhere can be estimated from the proportion
inference about the consequences of management or of points falling into each node. This statistic should be
conservation actions in different environmental settings. used cautiously, because it will be imprecise when samples
Because broad-scale variability due to climatic, are not representative of the new environmental
geologic, and cumulative effects of disturbance were conditions.13 One also can use the original tree to examine
potential covariates of tailed frog responses, we then new data, and use the misclassification rate as an estimate
assessed potential spatial autocorrelation between of the goodness-of-fit of the tree to the new data. 6
variables and sites to help interpret model structure. We O’Connor et al.12 suggested that while tree-based models
used a multivariate approach based on Mantel tests, which tend to be robust to moderate sampling biases, prediction
quantifies the degree of concordance between two accuracy degrades significantly if key environmental
matrixes.9 We compared a matrix of abiotic and biotic site variables are missing or are undersampled.
descriptors using the CART node structure as a classifier
of ecological similarity between sites, with a matrix of Example
geographic distances obtained from UTM coordinates. We sought to determine where tailed frogs occurred and
For practitioners seeking empirical species-habitat which attributes of headwater stream habitats were
models, CART methods have advantages over logistic and necessary to classify, map, and evaluate potential
generalized linear methods.15 They are easier to interpret protection measures. We georeferenced 846 records of
when both continuous and categorical predictor variables tailed frog occurrence, abundance, and habitat from three
are used. They assume no specific multiplicative sources: an extensive survey across the range of the tailed
relationship between predictor variables, so resulting frog in British Columbia7 (L. Dupuis, unpubl. data), and
models are robust to both the shape of frequency two intensive studies of habitat association in replicated
distributions of predictor variables, and the presence of forest management treatments.14 Most records were of
outliers. They can be tested on other datasets, even when tadpoles, but adults were recorded when found. Although
some variable values are missing. In short, they are these sources had different objectives and methods of data
appropriate to the kinds of data commonly available. collection, they could be combined in CART analysis.
Details of data collection are found in the sources noted.
Data Requirements Access to sites was usually by road, restricting most
We used both occurrence and relative abundance data samples to watersheds with some harvesting history and
routinely collected in broad-scale inventory surveys, and encouraging opportunistic sampling. At each site,
density data summarized from site-specific and longer- occurrence of tailed frogs (tadpoles or adults) was
term research studies. For predictor variables, we chose determined. For 59.9% of the sites, estimates were also
site-specific habitat variables that could be associated with made of tadpole density (area constrained searches; n =
local abundance of tadpoles, including: substrate 134) or relative abundance (time constrained searches; n
composition, channel disturbance, stream width, water = 374). Up to 20 stream characteristics were recorded at a
temperature, and recovery of streambeds from site, but many sites (n = 263) recorded fewer than 10 of
disturbance.2, 14 We added summary variables of climate, these.
topography and forest structure and/or disturbance effects from To this basic dataset, we added broad-scale climate,
existing GIS databases, standard climate records, or maps. geological and topographic attributes of sub-basins
potentially relevant to disturbance regimes and stream
Analytical outputs productivity. We used GIS databases to estimate
Primary outputs of the CART approach are heuristic harvesting attributes of each sub-basin, including: mean
prediction rules that can be used to classify site suitability road density within 100m of streams inventoried in the
across a wide range of environmental conditions for a provincial stream database, percent area of recent harvest
given species. The rules provide guidance on key (since 1978) at different elevations (300m, 300-800m and
environmental factors (physical, climatic, habitat above 800m), and area harvested on slopes >50% and
structure) determining population status, and expose within 100m of streams. From the BC Ministry of Forests
ecological and environmental thresholds that can be used climate database, we extracted mean precipitation and air
in GIS software to delineate protection zones or possible temperature variables representing long-term climate
 552 Wildlife–Habitat Relationships in Oregon and Washington

averages for each biogeoclimatic subzone in our samples. at broad spatial scales (macro- and meso scales in Table 1)
We measured stream gradients, and determined parent accounted for >54% of the explained deviance in all
lithology from Geological Survey of Canada maps. We models. Micro-scale characteristics of streams contributed
eliminated redundant variables (correlations >0.75 with less to explaining occurrence and abundance patterns,
other variables), and samples with many missing predictor although they were associated with patterns of tadpole
variables. density in coastal sites (Table 1). Parent bedrock was the
To assess whether different environmental relationships dominant predictor of distribution and abundance on the
were acting in different portions of the tailed frog range, coast, and relative abundance in the Kootenays (Table 1;
we divided sites into regional subsets reflecting influences Figures 1 and 2). Variables reflecting extent and location
of coastal and inland climates (Coast Mountain Range and of harvests and roads near streams at the sub-basin scale
Kootenay Mountains of southeastern BC). We derived also predicted distribution and abundance in both regions
models for occurrence (Coast: n=180; Kootenays: n=113) (Table 1). Second, positive spatial autocorrelations between
and, on sites where frogs occurred, relative abundance geographic distances separating sites and their
(Coast n=31; Kootenays: n=27), and density (Coast: n=92). classification by CART was apparent in patterns of
Here we excluded sites outside these two regions. occurrence (Mantel tests; coast: P<0.001; Kootenays:
We first developed initial classification trees using P=.036), and in the relative abundance data for the coast
CART10 for each response variable and sample subset, by (P < 0.036). Coastal densities and Kootenay relative
recursively splitting each node until either the proportion densities showed no spatial autocorrelation (P>0.5).
of the deviance remaining for sample sites in the node Underlying geology governed occurrence, but variation
was less than 5% of the original deviance, or there were in stream characteristics related to tailed frog abundance
fewer than 5 sample sites left in the node. For these trees were independent of geographic location. Third, while
we used only sites with no missing data for all predictor there is overlap in habitat variables predicting occurrence
variables. We then “pruned back” each initial tree using a in the two regions (Table 1), the relative importance of
ten-fold cross-validation analysis.6 That process iteratively each variable differed between the two regions. In
removes random subsets of the original observations, and particular, variables linked to forest harvesting (areal
uses the remaining observations to refit the classification extent and location of recent harvest, and stream substrate)
tree, allowing the analyst to select the number of nodes were more important in the Kootenays than in coastal sites
that provides a parsimonious fit to the data. (Table 1; Figures 1 and 2). The % cover of fines and stream
Much can be inferred from CART analysis. We note temperature helped distinguish sites of high and low
five major points. First, environmental variables operating tadpole density on the coast. Differences in the importance

Table 1. Percentages of original deviance explained by the CART models for tailed frog occurrence (Figures 1
and 2) and abundance (see text) among the various classes of predictor variable. Scales defined by range of
detectable spatial autocorrelation in the original variables (macro = 80-120 km; meso = 25-60 km; micro = <25
km). Sample sizes indicate the number of sites used in each model.

Predictor class Spatial scale Coast Kootenays

Occurrence Relative Density Occurrence Relative
(n = 180) abundance (#/ m2) (n = 113) abundance
(#/ 20 min.) (n = 92) (#/ 20 min.)
(n = 31) (n = 27)

Climate1 macro - - 5.4 - -

Topography2 meso 9.5 - 11.1 3.5 9.9
Bedrock Lithology meso 21.1 - 22.5 2.3 24.5
Sub-basin Harvesting meso 9.6 7.5 2.1 18.8 9.9
and Road Density3
Stand Type4 micro - - - - -
Stream Substrate5 micro 6.9 - 37.0 13.8 10.6
Stream Temperature micro - - 9.1 6.0 -
Stream Width micro - 35.2 - 12.1 -
Fine Organic Debris micro 2.1 - - - -
Reach Slope (%) micro - - - - -
1
means for: annual precipitation, summer precipitation (in mm), annual temperature, temperature of warmest month (oC),
2
elevation (m), aspect (o), stream gradient (%)
3
% sub-basin area logged since 1978; % logged area within 100m of streams, road density within 100m of streams (km/ha)
4
age class of adjacent stand (0-10 yrs; 11-20 yrs; 21-60 yrs; 61-100 yrs, > 100 yrs), presence of riparian buffers (yes/no)
5
% cover of boulders, cobbles, and fines
 CHAPTER 22: MODELING—CASE HISTORIES 553

Figure 1. Classification and

Regression Tree (CART) model for
tailed frog occurrence in the
western coastal mountain zone of
British Columbia. Numbers inside
the rectangles (end nodes) are the
proportion of sampled sites (sample
size in brackets) with tailed frogs
present. The splitting predictor
variable and its threshold value is
shown for each branch of each node.
The length of each branch (growing
to the right) is proportional to the
percent of deviance explained by the
splitting variable at each node.

of macro-scale environmental variables relative to micro- presence or absence of buffers) had no apparent predictive
scale variables may reflect differences in regional capacity.
hydrology regimes or histories of forest management. The process can derive qualitative and quantitative
They could also reflect the restricted spatial distribution criteria for locating conservation measures and refining
of occurrence records in the Kootenays,7 which limits the site-specific forest practices for species at risk. Physical
ability to adequately separate components of variation. setting had a greater influence on occurrence and
Fourth, important dependencies between habitat variables abundance of larval frogs than did adjacent forest
were exposed by the occurrence trees (Figures. 1 and 2). practices. We offer two cautions when interpreting these
On the coast, sites with the highest probability of tailed models: 1) our source data for extent of recent logging
frog occurrence were associated with relatively may be at too coarse a scale to reliably detect an influence,
undeveloped watersheds in regions of highly competent and 2) our models are directly relevant only to the in-
bedrock (granodiorites or volcanic breccias) or, in regions stream life stage. Nonetheless, managers can use
of less competent bedrock types, with boulder-dominated classification trees to identify habitat protection options
streams, higher elevations, or steep stream gradients if a in several ways. For example, if the goal is maintenance
moderate amount of logging had occurred in the riparian of current habitat, then all sites meeting the environmental
zone. In the Kootenays, sites with the highest probability criteria with a >0 probability of occurrence become
of tailed frog occurrence were associated with watersheds candidates for protection measures. Our analyses suggest
having extensive harvesting history, in smaller boulder- that practices designed to conserve existing tailed frog
dominated streams, but, in less developed watersheds, tadpole populations (e.g., buffers around streams) will be
with sites having cobble-producing bedrock (argillite; most successful if: 1) they are situated in intrusive or
quartzite). Finally, variables directly reflecting forest metamorphic bedrock formations; 2) they target streams
management practices adjacent to sample sites (e.g. on intermediate gradients with significant boulder cover;
 554 Wildlife–Habitat Relationships in Oregon and Washington

Figure 2. CART model for tailed

frog occurrence records in the
eastern population of tailed frogs
in southeastern British Columbia.
Numbers as described in the
caption for Figure 1.

and 3) they are in watersheds with low or moderate levels There are four strengths to this approach in evaluating
of historical harvest. Conservation efforts will be less species-environment relationships. First, CART techniques
effective where weathering products from surrounding complement landscape-scale methods for predicting
geology produces fine-grained or rubbly stream presence and diversity of species (e.g., Gap analysis) by
sediments, at low elevation sites, and in sites with revealing pathways by which environmental variation
increased chances of dry periods during the summer. In may be determining species’ distribution. Second, CART
portions of the tailed frog range, entirely natural factors techniques make efficient use of minimal, low cost data
have much greater influence than human-induced sets involving combinations of nominal and numerical
changes. Such areas are not good candidates for expensive data. For relatively unstudied species, robust estimation
protective measures. More complex and quantitative rules techniques like CART can quickly explore existing habitat
can be derived from the trees (see Methods) to develop databases to yield provisional rules to guide management
stream classifications for species sensitivity, or compare options. Third, CART models permit data of several
predictions with other datasets. independent, small-scale studies to be “scaled up” into a
more spatially extensive analysis without making
Conclusions restrictive assumptions about how the same
Hierarchical classification trees can expose complex environmental variables interact in all locations. Finally,
species-environmental relationships operating at different CART models are easy to interpret, leading to hypotheses
scales, using field inventory samples overlaid with readily that may predict observed distribution and abundance,
available data describing climate, topography, and and ranking the likelihood of responses of species to
geology. Protection measures for tailed frogs must account combination of environmental variables. Our models can
for spatial variation in the sensitivity of sites to be effective. easily be tested and used in other areas where standard
More generally, for species in which the effects of forest inventory data is routinely collected (e.g., stream surveys).
practices act most strongly through altered gradients There are weaknesses in the approach. Most
affecting growth, fecundity and movements, a importantly, classification tree approaches are heuristic in
classification tree approach yields practical design criteria nature—processes inferred from them must be treated as
for management more quickly and with less effort than hypotheses until their predictions are confirmed by more
do standard linear models. detailed experimental studies. Secondly, interpretations
 CHAPTER 22: MODELING—CASE HISTORIES 555

of model outputs are constrained to the precision of the

datasets used as input. Finally, classification tree Case Study 4
approaches by themselves cannot infer effects of specific
Applying Gap Analysis to County and
habitat protection measures on future extirpation
probabilities, because of basic uncertainties in species’ life Regional Land Use Planning
history. Matthew R. Stevenson

Acknowledgments Introduction
We thank Linda Dupuis and Pierre Friele for the tailed
Biodiversity is the range of living organisms and the
frog survey database for British Columbia. Malcolm Gray
processes which created and sustain them. Human
(BC Ministry of Environment, Lands and Parks) provided
activities on the landscape in the form of resource
his Watershed Ranking database for coastal BC, while
extraction, agricultural conversion, and urban
Matthew Craig provided detailed GIS data for the
development have had a substantial impact on
Kootenay region. Arnold Moy assisted with analyses.
biodiversity. In fact, biological diversity is being lost to
Comments by Linda Dupuis, David Huggard and two
such a degree that the present rate of extinction rivals the
anonymous reviews improved the manuscript. Research
catastrophic loss of species at the end of the Paleozoic and
support provided by Forest Renewal BC, the BC.Ministry
Mesozoic eras. The rate at which species are becoming
of Environment, Lands and Parks, the BC Ministry of
extinct is 1,000 to 10,000 times higher than before human
Forests, and Wildlife Habitat Canada to FLB. This is
intervention.20 The Nature Conservancy estimates that 200
contribution M-10 of the Centre for Applied Conservation
plant species and 71 species and subspecies of vertebrates
Biology, UBC.
have gone extinct in North America (excluding Mexico)
Literature Cited since European settlement.9 The condition of ecosystems
1. Brieman, L., J.H. Friedman, R.A. Olshen, and C.J. Stone. 1984.
is similar: according to the Office of Technology
Classification and regression trees. Wadsworth International Group, Assessment, “twenty-three ecosystem types that once
Belmont, CA. covered about half the coterminous United States now
2. Bull, E.L., and B.E. Carter. 1996. Tailed frogs: distribution, ecology and cover about seven percent.”21
association with timber harvest in northeastern Oregon. USDA Forest The reasons for this ecological catastrophe are complex,
Service Research Paper PNW-RP-497. but to a large degree the responsibility can be attributed
3. Bunnell, F.L., and D.J. Huggard. 1999. Biodiversity across spatial and
to the way in which cities, counties and regions have
temporal scales: problems and opportunities. Forest Ecology and
Management 115: 113-126. chosen to locate, organize, and regulate land uses. Perhaps
4. ———, L.L. Kremsater, and R.W. Wells. 1997. Likely consequences of in partial recognition of this, the State of Washington
forest management on terrestrial, forest-dwelling vertebrates in adopted the Growth Management Act (RCW 36.70A) in
Oregon. Oregon Forest Resources Institute, Portland, OR. 1990, which includes in its requirement that jurisdictions
5. Bury, R.B. 1968.The distribution of Ascaphus truei in California. planning under the act “conserve fish and wildlife habitat”
Herpetologica 24: 39-46. and “protect the environment.” An additional legal
6. Clark, L.A., and D. Pregibon. 1992. Tree-based models. Pages 377-419 in
J.M. Chambers and T.J. Hastie, editors.Statistical Models in S.
imperative facing counties and regions is the Federal
Wadsworth & Brooks/Cole Computer Science Series, Pacific Grove, Endangered Species Act (ESA), which can trigger severe
CA. limitations upon local activities once a species (or a
7. Dupuis, L.D., and F.L. Bunnell. 1997. Status and distribution of the tailed population of a species) has been listed.17
frog in British Columbia. Unpublished report for Forest Renewal In addition to meeting these regulatory requirements,
British Columbia. cities, counties, and regions should be concerned with
8. Legendre, P. 1993. Spatial autocorrelation: trouble or new paradigm?
preserving biodiversity because only properly functioning
Ecology 74: 1659-1673.
9. ———, and M.-J. Fortin, 1989. Spatial pattern and ecological analysis.
ecosystems can provide the free ecosystem services
Vegetation 80: 107-138. civilization requires. However, one problem confronting
10. Mathsoft, Inc. 1998. SPLUS 4 Guide to Statistics. Mathsoft, Seattle, jurisdictions interested in protecting biodiversity is the
WA. lack of information describing the distribution of
11. Metter, D.E. 1964. A morphological and ecological comparison of two biologically important areas. In an attempt to provide this
populations of the tailed frog, Ascaphus truei Stejneger. Copeia 1964: information and improve upon previous fragmented
181-195.
approaches to single species preservation, the National
12. O’Connor, R.J., M.T. Jones, D. White, C. Hunsaker, T. Loveland, B. Jones,
and E. Preston. 1996. Spatial partitioning of environmental correlates of Gap Analysis Program (NGAP) was charged with
avian biodiversity in the conterminous United States. Biodiversity conducting the largest effort ever to map the distribution
Letters 3: 97-110. of biological resources in the United States.
13. Thompson, J.D., G. Weiblen, B.A. Thomson, S. Alfaro, and P. Legendre. Gap analysis, which uses land cover, wildlife habitat
1996. Untangling multiple factors in spatial distributions: lilies, gophers, relationship models, and other data to predict the
and rocks. Ecology 77: 1698-1715. distribution of terrestrial vertebrates, provides information
14.Wahbe,T.R. 1996.Tailed frogs (Ascaphus truei) in natural and managed
coastal temperate rainforests of Southwestern British Columbia,
which can be applied to landscapes and regions in order
Canada. Thesis, University of British Columbia, Vancouver, BC, Canada. to ensure that land management and land use plans more
15.Verbyla, D.M. 1987. Classification trees: a new discrimination tool. effectively incorporate the protection of biodiversity. This
Canadian Journal of Forest Research 17: 1150-1152. case study details one approach for applying these data
to land use planning.
 556 Wildlife–Habitat Relationships in Oregon and Washington

Methods instance, the violet-green swallow—might occur in 480

polygons where the Brewer’s blackbird has also been
Description of Process and Data Requirements predicted to occur. Selecting the polygons where both the
The process developed for applying Washington GAP data blackbird and the violet-green swallow occur reduces the
to county and regional land use planning involves five number of polygons to 480 needed to represent both
steps. species. Each additional species added to the set reduces
1. Finding the minimum amount of land required to the number of polygons in a similar fashion until finally
represent each species predicted to occur in the study the addition of one more species results in the selection of
area. no polygons because no occurrences overlap. This process
This first step is intended to ensure that every species is repeated until all species in each taxonomic group are
predicted to occur in the County is represented at least represented. The approach is similar (though less
once by any system of protected lands. This concept, sophisticated) to the representative set solutions discussed
commonly referred to as “representation,” does not by Pressey and others.2, 3, 11, 15, 19
necessarily mean that all life requisites of any given species 2. Establishing species richness thresholds to locate
can be accommodated within the representative land area, candidate areas for habitat reserves.
but does guarantee that at least some habitat for every Establishing species richness thresholds by taxonomic
species predicted to occur in the County is included.3, 11 group is necessary before areas of high species richness
Unfortunately, WAGAP did not determine home range can be identified as potential locations for reserves. This
size for each species, which is necessary to determine the process requires the creation of a decision rule for which
amount of habitat required to maintain minimum viable polygons should be selected or “turned on,” based upon
populations for all vertebrates. Consequently, the number of species predicted to occur in each.
representation means simply that at least one vegetation Determining a universally applicable richness threshold
polygon per species is included within the proposed is not as straightforward as is representation, since richness
network of reserves and corridors. is relative to the total number of vertebrates (by group)
Finding representation begins with clipping raw within the study area. For this analysis, threshold levels
WAGAP data to the study area. The WAGAP data are were arbitrarily set at 75 percent of possible richness for
organized into three discrete groups: land cover, which mammals and birds and 50 percent of possible richness
consists of 100 ha minimum mapping unit (MMU) land for reptiles and amphibians. A lower threshold for reptiles
cover polygons delineated using 1991 Landsat TM data; and amphibians was used because their populations tend
vertebrate distributions, which were derived using to be concentrated in discrete habitat patches such as
wildlife/habitat relationship models in conjunction with wetlands, rivers, and lakes, particularly for amphibians.
the land cover map; and stewardship, which rates the
management of large land holdings on a scale of 1 to 4 3. Delineating potential habitat reserves.
based upon the level of protection of biodiversity. I considered all polygons with relatively high species
First, using the clipped species distribution data, the richness for designation as a habitat reserve area. Factors
polygon or set of polygons with the highest species used in the selection of habitat reserves included polygon
richness (greatest number of species) for each given size, land cover, level of human development and intensity
taxonomic group is located. At the same time, species of human activity within the area, adjacent development,
which are not predicted to occur within this initial polygon relationship to Priority Habitats and Species (PHS)
or set of polygons are also identified. Next, these non- mapped designations, and proximity to other polygons
represented species within each group are ranked in satisfying the richness requirement. The Washington
descending order according to the number of occurrences Department of Fish and Wildlife (WDFW) established and
within the study area. Then, starting with the polygons maintains the PHS database, which was developed to help
for the non-represented species with the highest predicted guide management decisions regarding biologically
number of occurrences, polygons are eliminated by adding important areas. PHS differs from WAGAP in two
species to the representative set. important ways: first, it does not use wildlife-habitat
In other words, out of 180 total bird species in a given relationship models to predict species distributions; and
study area, 100 species may be present in the richest second, it does not encompass all terrestrial vertebrates
polygon, meaning 80 species are not represented. Of these within Washington.18 Since PHS information is limited to
80 non-represented species, the number of polygons in priority species (priority species include threatened,
which each occurs will be different. As an example, if the endangered, or sensitive species; vulnerable aggregations;
Brewer’s blackbird were not represented in the richest and species of recreational, commercial, and/or tribal
polygon but occurred most frequently among all non- importance) it should be used in conjunction with WAGAP
represented species, its distribution would form the initial data to corroborate areas indicated as important for
set of polygons for the second round of selection. If the establishing representation, corridors, and habitat
Brewer’s blackbird were to occur in 600 polygons, then reserves.
600 polygons would make up the largest set of polygons The polygons selected for potential habitat reserves
that would appear in the second round of selection. The were large polygons with high species richness, natural
next most representative species from that set—for land cover, low internal and adjacent human development,
 CHAPTER 22: MODELING—CASE HISTORIES 557

and high PHS polygon and point coverage overlap.6 Data on land use, zoning, growth area boundaries, and
Additionally, heron rookeries, waterfowl concentrations, critical areas are maintained by all counties and can be
and other vulnerable wildlife aggregations received used as an indication of how the landscape will likely
special consideration when potential reserves were change in the future and can also be used to detect areas
delineated.18 of potential conflict between planned or zoned
development and areas that may be important as habitat
4. Refining polygon sets to determine corridor
reserves or corridors. Digital orthophotos and stereo pairs
alignments.
were used to examine actual land cover conditions during
A considerable amount of land was selected during the
the process of refining boundaries for reserves and
process of identifying key habitat areas based on
corridors.
representation and richness of species, and the next step
was to refine this area into a comprehensive system of 5. Establishing an ecological and political context for
habitat reserves and wildlife corridors. Three kinds of data system evaluation.
were used to reduce and refine the area resulting from The reality of land management is political, so it is
steps 1-3: predicted species distributions from WAGAP useful to establish a context for decision making by
for all counties surrounding the study area county; PHS analyzing how policy decisions may affect stewardship
data from WDFW, and a variety of thematic data unique status within both ecologically and politically determined
to the County, such as large-scale land cover, topography, boundaries. I chose the statewide extent of the vegetation
wetlands and hydrology, land use, zoning, growth area zones passing through the study area (a county) as the
boundaries, critical areas, digital orthophotography, and ecological unit of analysis for the relative levels of
air photo stereo pairs. preservation of vegetation zones and vertebrates and I
The larger WAGAP data set for the counties used the county as the politically determined boundary.
surrounding the study area county were used during the The stewardship status of vegetation zones can serve as a
process of refining the selected set to ensure that reserves rough indicator of how well a county is protecting
and corridors connected to potential reserves in adjacent biodiversity relative to the entire extent of the vegetation
counties. The PHS database was used to ensure that zones that pass through the county. I used four
important habitats were retained as the larger WAGAP measurements to establish the political and ecological
polygons were refined into corridors. The corridors context. First, I determined the percentage of land within
themselves were delineated using the county-specific data the statewide extent of vegetation zones which is managed
in conjunction with the WAGAP and PHS data. primarily for the protection of biodiversity. Second, I
The most important county-specific data set was a determined the percentage of land within all vegetation
large-scale land cover map with a minimum mapping unit zones found within the study area. Third, I determined
(MMU) that is substantially smaller than the WAGAP the percentage of study area vegetation zones managed
MMU of 100 ha. The smaller MMU is necessary because primarily for the protection of biodiversity. Lastly, I
it lends greater accuracy to the process of delineating determined the new percentage of study area vegetation
potential corridor alignments, particularly through highly zones that would be managed primarily for the protection
developed or disturbed areas.9, 13 I created this map in the of biodiversity within any proposed system of corridors
summer of 1997 by interpreting and digitizing land use and reserves.
and land cover from digital orthophotos supplied by
WDFW. Unfortunately, this type of mapping was Process Output
expensive, time-consuming, and did not cover the entire Using the methods described above, this process will
county. Outside the mapped area, a parcels coverage and produce four primary products: an inventory of the
primary roads coverage were used as proxies for the predicted distributions of terrestrial vertebrates and their
location and intensity of development and human activity. level of protection within the study area and the state; an
A USGS digital elevation model (DEM) was used to inventory of vegetation zones and their level of protection
ensure that corridors follow realistic gradients (i.e. within the study area and the state; a set of potential
corridors should not run over cliffs) and that the complete habitat reserves and wildlife corridor alignments within
system “maintain(s) natural ecosystems and biodiversity the study area; and lastly, a “before and after” description
across the full extent of environmental gradients”.9 of the effect of any new system of corridors and reserves
Encompassing all environmental gradients will become upon the protection of vertebrates and vegetation zones.
increasingly important as global temperatures continue It is important to remember that WAGAP data was
to rise and ecological communities slowly shift northward designed for landscape-scale analyses; thus, it may be too
and upward to compensate.10 coarse to use directly for acquiring conservation lands.
Additionally, wetlands, rivers, and streams data were Additional field work, ground truthing, mapping, and
extremely important for delineating corridors in general surveying are critical before any specific land management
but especially for amphibians and birds. As demonstrated decisions are made.12 However, it is also important to note
by Klaus Richter in a study of 19 wetlands in King County, that the WAGAP data base is the only one of its kind in
Washington, wetlands are used disproportionately by existence, it is readily available (for a nominal fee) to any
birds and amphibians and are therefore critical habitat county or city that is interested in using it, and it can be
areas for these taxa.Richter, cited in 1 analyzed and manipulated on a PC within a realistic time
frame and budget.
 558 Wildlife–Habitat Relationships in Oregon and Washington

Example: MMU used by WAGAP. The IUGA does not cover the
Spokane County Comprehensive Plan entire county, so we used a parcels coverage and primary
The process described above was developed in Spokane roads coverage as indicators of human development and
County, Washington through an interdisciplinary studio activity in areas lacking higher resolution land cover
class in the Department of Urban Design and Planning at information. A parcels coverage can serve as an effective
the University of Washington in the fall of 1997. The initial proxy because smaller parcels typically indicate higher
motivation for conducting the analysis came from staff in densities and intensities of development, impervious
the WDFW Region 1 office, who were working with the surfaces, non-native vegetation, fences, and automobile
Spokane County Division of Long Range Planning to traffic. Conversely, larger parcels tend to have lower
locate “wildlife corridors” and “landscape linkages,” two densities, less impervious surface, and more native
categories within the County’s recently adopted Critical vegetation (with the exception of agricultural areas). Based
Areas Ordinance (CAO) which had been defined but not upon field visits, we determined that parcels smaller than
yet identified on the ground. In order to map these specific approximately 20 acres were too disturbed to be included
critical areas, the County was willing to serve as the pilot in potential corridor alignments.
project for the application of WAGAP data. I went on to Roads (particularly interstates and highways) were an
further develop this application in my Master’s thesis at equally important consideration when determining
the University of Washington.14 potential reserve locations and corridor alignments,
The first step in the analysis was to examine patterns because they may impact all of the taxonomic groups
of vertebrate richness by taxonomic group (in this case modeled by WAGAP (see for example transportation and
mammals, birds, reptiles, and amphibians). The second wildlife: reducing wildlife mortality and improving wildlife
step involved the establishment of representation for all passageways across transportation corridors16).
vertebrates (i.e., the minimum configuration of polygons The refined corridor system is based primarily upon
that captures all vertebrate taxa in at least one polygon). the existing riparian network and utilizes upland
Output from the richness analysis played a key role in the connections to complete and complement this foundation.
selection of WAGAP polygons considered as starting Riparian corridors were selected as the backbone for the
points for reserves. These three data sets were combined system in Spokane County because of their
with a variety of additional data sets to produce the disproportionately significant contribution to
potential habitat reserve locations and wildlife corridor biodiversity. 8 Wherever current land use permitted,
alignments. corridors were drawn at a minimum of 1/4-mile to 1/2-
The initial system of habitat reserves was determined mile wide on center in order to protect riparian vegetation
as described above in step three. In Spokane County, areas and provide realistic adjacent upland movement corridors
of habitat with high species richness, natural land cover, (particularly in forested areas) that would be wide enough
low internal and adjacent human development, and high to avoid the “edge” effects, which can be deleterious to
PHS polygon and point coverage overlap were located in some species.
the northeast, west, and southwest portions of the county.
These potential habitat reserve areas encompass such well- Results
known places as Mt. Spokane State Park, Turnbull If the refined reserve and corridor system were to be fully
National Wildlife Refuge, and portions of the Spokane implemented within Spokane County, the amount of
River. additional land managed primarily for the protection of
The most crucial piece of data used in refining the biodiversity would increase from 4% to 30%. As shown in
initially-selected reserves and corridors was the fine- the last column of Table 1, all vegetation zones within the
grained land-use land-cover map I created of the Interim County would gain at least 10% in protected area in
Urban Growth Area (IUGA) surrounding greater addition to currently protected land. These increases are
metropolitan Spokane. This map was created with a five important within the County but also address statewide
acre (2.5 ha) MMU, substantially smaller than the 100 ha conservation needs. As described by WAGAP in their final
report summary, “The vegetation zones with the highest

Table 1. Conservation status (acres, percent) of vegetation zones in

Spokane County under proposed system of corridors and reserves.

Vegetation zone Acres currently Percent of vegetation Acres in proposed Additional percent of
protected zone system vegetation zone within system

Ponderosa pine 21,917 1.9 165,821 26.5

Palouse 907 <0.01 36,404 10.6
Interior Doug fir 17,441 1.5 121,125 55.7
Three-tip sage 0 0 13,188 23.9
Grand fir 47 <0.01 2,025 97.3
Interior redcedar 0 0 49 89.2
 CHAPTER 22: MODELING—CASE HISTORIES 559

Table 2. Protected habitat as a percentage of total habitat for terrestrial vertebrates in Spokane County
before and after proposed system of reserves and corridors.

Mean (%) Median (%)

Taxonomic group Count Before After Before After

Mammals 66 5 41 4 39
Birds 161 6 37 6 38
Reptiles/Amphibians 17 5 44 6 43
All vertebrates 244 6 39 6 39

Table 3. Test results comparing the original and modified reserve and corridor system to field data specific to
Spokane County (results given in % of original database points or area [if polygons]).*

System PHS Heritage Herp Sensitive Heron Elk Public Input

Plants Rookeries

Original <1% omitted 31% omitted 33% omitted 30% omitted None omitted 3% omitted 3.5% omitted
Modified <1% omitted 7.8% omitted 7.7% omitted 7.5% omitted None omitted <1% omitted <1% omitted
* (Adapted from Ferguson, Robinette, and Stevenson 1999)

Conservation Priority Index (CPI) are steppe zones (which backbone for their entire conservation strategy. WDFW
include Palouse and Three-tip Sage).” Additionally, has used the data to evaluate parcels nominated for a
WAGAP describes the Ponderosa Pine and Oak zones as recently re-approved and taxpayer funded Conservation
“zones of high reptile and avian diversity” which have Futures Program, and has collaborated with Spokane
“moderately high CPIs.”7 The Ponderosa Pine zone and County to evaluate and further refine the proposed system
both of the steppe zones found in Spokane County would into a Final Open Space Plan.
be substantially more protected under the system designed As shown in Table 3, the WDFW evaluation of the
with this process. Furthermore, the majority of existing original system of reserves and corridors was made with
protected areas in Washington State are located at high field data, WDFW Heritage points, WDFW reptile and
elevations in the Cascade and Olympic mountains. The amphibian (herp) points, DNR sensitive plant species
system proposed for Spokane County would improve points, great blue heron rookery locations, elk habitat data,
statewide conservation not just by increasing the area of and public input. WDFW biologists detected relatively
protected land in relatively under-represented low high omission rates for Heritage points, herps, and
elevation vegetation zones, but also by connecting sensitive plant species, but fairly good coverage of all other
important habitats in an area of the state with very little evaluation criteria. By using these data to increase the land
biodiversity protection. area included in the system of reserves and corridors by
Table 2 shows current habitat protection in Spokane just 2%, omission rates were substantially reduced, as
County for all terrestrial vertebrates at a mean and median indicated in the second row of Table 3. According to
of 6% of predicted distributions. If the entire system of Ferguson et al,4 “this final version appears to have satisfied
reserves and corridors were implemented with no our goals—a biologically defensible open space plan with
substantial modifications, the median area of protected a reasonable amount of land being identified in just over
habitat for all species would increase to 39%, more than two months.”
six times the existing level of protection. This additional
protection occurs by increasing the amount of protected Conclusions
land to just 30% of the County’s total land area.
Based upon interviews I conducted with the Spokane Strengths and Weaknesses
County Division of Long Range Planning, WDFW, and The methodology described above has the capability of
the Inland Northwest Land Trust (INLT), the information informing the process by which land use plans are created
produced through this application of WAGAP is unique and land management policies are formulated. However,
and valuable to all of them. These results were finalized this process should be used with an understanding that it
in January 1998, and so far Spokane County is using the is only a tool for developing the solution required to
information to update its Critical Areas Ordinance and conserve biodiversity in the face of expanding human
evaluate a proposed system of extensive, high capacity population and human activity. Additionally, it is
roads encircling the greater Spokane area. The INLT has important to realize that although there are many strengths
used and will continue to use the results as the foundation to this approach, there are also shortcomings which must
for their Threads of Hope campaign, which serves as the be acknowledged and addressed accordingly.
 560 Wildlife–Habitat Relationships in Oregon and Washington

Strengths. Gap data predicts distributions for all Additional Research

vertebrates in the state, not just rare, threatened, or The Spokane County pilot project provided a tremendous
endangered species. Combining richness, representation, opportunity for the development of a methodology for
and connectivity ensures that every species is contained applying WAGAP to real problems. The process described
within any proposed system of protected or specially above can provide important information to decision
managed areas. Working at a variety of resolutions allows makers. However, there are several key areas of additional
land planners and managers to see patterns that would research which would enhance the quality, accuracy, and
otherwise not be visible, and integrating the results of this usability of the finished product. Regional coordination
process into land use planning can be an effective way of is indispensable because the effectiveness of any system
preventing additional populations of vertebrates and other of corridors and reserves will be severely constrained if
organisms from becoming threatened or endangered. similar biodiversity planning activities in adjacent
Identifying the most important areas of habitat can provide jurisdictions are not coordinated. Ground truthing should
some degree of certainty to the development and resource be conducted in parallel with the expanded mapping of
extraction industries. The successful adoption and land cover to ensure that what is being mapped actually
implementation of the results of this process can lead to a exists on the ground. Developing (or applying existing)
significant increase in the amount of land managed set coverage and optimization algorithms would allow
primarily for the protection of biodiversity. A number of for the evaluation of a variety of approaches to
management and policy-making bodies are necessarily representation, such as comparing the differences between
involved in the process and are therefore more likely to representing each taxonomic group and adding the results
find the outcome valuable. Finally, creating a connected versus representing all groups simultaneously. Land use
system of habitat areas maintains ecosystem services, modeling would be beneficial for determining which of
enhances the aesthetic qualities of the landscape, raises those areas within the proposed system are also the most
property values, and can provide recreational likely to come under intense development pressure and
opportunities to the burgeoning Washington population. should consequently be top priorities for protection. Lastly,
Weaknesses. The bulk of the process is based upon Gap public involvement is extremely important, because the
analysis, which being predictive in nature contains errors level of support for this approach will be related to the
of commission and omission, and overlaying these degree to which the people affected by it understand,
distribution maps to create composite richness maps can appreciate, and acknowledge its usefulness.
lead to additional error accumulation. 5 The satellite
Literature Cited
imagery used to conduct WAGAP was collected in 1991,
1. Azous, A.L. and R.R. Horner. 1997. Wetlands and urbanization:
and substantial changes may have occurred in the implications for the future. Final report of the Puget Sound Wetlands
landscape in the intervening eight years. Vertebrate and Stormwater Management Research Program, Washington State
distributions are not always the most accurate proxy for Department of Ecology, Olympia, WA.
the distributions of other taxa. The various data sets used 2. Church, R.L., D.M. Stoms, and F.W. Davis. 1996. Reserve selection as a
in this analysis were created at different scales for different maximal covering location problem. Biological Conservation 76:105-
purposes and do not always work well in conjunction with 112.
3. Csuti, B., J.D. Camm, B. Downs, M. Huso, R. Hamilton, M. Kershaw, A.R.
one another. Capturing the presence of any given taxon Kiester, S. Polasky, R.L. Pressey, K. Sahr, and P.H. Williams. 1996. A
does not guarantee long-term persistence (i.e., geographic comparison of reserve selection algorithms using data on terrestrial
units or polygons do not necessarily contain sustainable vertebrates in Oregon. Biological Conservation. 80:83-97.
populations). The coarse scale of the analysis necessitates 4. Ferguson, H.L., K. Robinette, and M.R. Stevenson. 1999. Searching for
a substantial amount of follow up fieldwork and the best science available: a method for identifying a corridor-open
additional fine-scale analysis, as detailed above. Lastly, space system for land use planning. Proceedings, Fourth International
Symposium on Urban Wildlife Conservation, Tucson, AZ.
classification of vegetation may be inadequate for certain
5. Flather, C. H., K.R. Wilson, D.J. Dean, and W.C. McComb. 1997.
taxa, and habitat relationship models may be weak for Identifying gaps in conservation networks: of indicators and uncertainty
certain taxa. in geographic-based analyses. Ecological Applications 7:531-542.
6. Forman, R.T.T. 1995. Land mosaics: the ecology of landscapes and
Relevance to the Landowner regions. Cambridge University Press, Cambridge, England.
This process is relevant to landowners in that it can 7. Grue, C.E., K.M. Cassidy, and K.M. Dvornich. 1998. GAP Bulletin
provide a considerably higher degree of certainty Number 6: Final report summary, Washington GAP analysis project.
concerning the use of their land. The current ad-hoc, U.S. Department of the Interior, U.S. Biological Service, Washington
D.C.
reactionary, species-by-species approach to protecting 8. Naiman, R.G., H. Decamps, and M. Pollock. 1993. The role of riparian
biodiversity is divisive and counter-productive. Land corridors in maintaining regional biodiversity. Ecological Applications
owners will probably be more interested in actively 3:209-212.
participating in the stewardship of their land when they 9. Noss, R.F., and A.Y. Cooperrider. 1994. Saving nature’s legacy: protecting
realize it is biologically significant on a regional scale. and restoring biodiversity. Island Press, Washington, D.C.
However, it is important that landowners be engaged and 10. Peters, R.L., and T.E. Lovejoy, eds. 1992. Global warming and biological
diversity. Yale University Press, New Haven, CT.
involved through a process relying on incentives for
11. Pressey, R.L., H.P. Possingham, and J.R. Day. 1997. Effectiveness of
cooperation, not government command regulations. alternative heuristic algorithms for identifying indicative minimum
requirements for conservation reserves. Biological Conservation
80:207-219.
 CHAPTER 22: MODELING—CASE HISTORIES 561

12. Scott, J.M., F. Davis, B. Csuti, R.F. Noss, B. Butterfield, C. Groves, H. over large landscapes could provide natural resource
Anderson, S. Caicco, F. D’Erchia, T.C. Edwards Jr., J. Ullman, and R.G. managers with greater flexibility in the design of forest
Wright. 1993. Gap analysis: a geographic approach to protection of management plans that promote the conservation of
biological diversity. Wildlife Monographs 123.
13. Stine, P.A. 1995. A multi-scale conservation assessment of plant
northern spotted owls and their habitat. Finally, the model
communities in southern California. Dissertation, University of can be adapted for other species of concern by identifying
California at Santa Barbara, Santa Barbara, CA. other vegetation attributes and spatial parameters that
14. Stevenson, M.R. 1998. Protecting biodiversity: applying gap analysis in describe habitat conditions conducive for that species’
Spokane County, Washington. Thesis. Department of Urban Design and needs.
Planning, University of Washington, Seattle, WA.
15. Stokland, J.N. 1997. Representativeness and efficiency of bird and Methods
insect conservation in Norwegian boreal forest reserves. Conservation
ArcView GIS 3.0a with the Spatial Analyst Extension (ESRI
Biology 11:101-111.
16. United States Department of Transportation, Federal Highway 1997) was used to process the model. The model consists
Administration, State of Florida Department of Transportation. 1996. of several iterative steps. In each step, specific features in
Transportation and wildlife: reducing wildlife mortality and improving a GIS map are selected for and modified to create a new
wildlife passageways across transportation corridors. Proceedings of GIS map, until the final GIS map depicting high potential
the Florida Department of Transportation/Federal Highway nesting areas is created. ArcView GIS commands used in
Administration Transportation-Related Wildlife Mortality Seminar.
each step are all found within the default menus, requiring
Federal Highway Administration, Washington, D.C.
17. United States Fish and Wildlife Service. 1988. Endangered Species Act
no special programming. This model process was
of 1973. As amended through the 100th Congress. United States developed using the Gifford Pinchot National Forest’s
Department of the Interior Washington, D.C. (GPNF) GIS vegetation map and interim vegetation table
18. Washington Department of Fish and Wildlife. 1996. Priority habitats (I-Veg). Both the GPNF GIS vegetation map and I-Veg table
and species list. Olympia, WA. are publicly available on the World Wide Web (http://
19. Williams, P., D. Gibbons, C. Margules, A. Rebelo, C. Humphries, and R. www.fs.fed.us/gpnf/gis/index.html).
Pressey. 1996. A comparison of richness hotspots, rarity hotspots, and
To begin the model process, a GIS vegetation map with
complementary areas for conserving diversity of British birds.
Conservation Biology 10:155-174.
forest stand attribute information detailed enough to allow
20. Wilson, E.O., and F.M. Peter, editors. 1988. Biodiversity. National the delineation of suitable northern spotted owl habitat is
Academy Press, Washington, D.C. required (Figure 1, Step A). The organization and detail
21. Winckler, S. 1992. Stopgap Measures. The Atlantic Monthly January: 74- of forest stand attribute information can vary widely
81. within and between federal, state and private agencies.
Thus, a biologist needs to review the GIS vegetation map
database and, using local knowledge and professional
Case Study 5
judgment, determine if it is adequate to confidently
A Model to Determine Potential identify the extent of stands containing suitable northern
Northern Spotted Owl Nesting Areas spotted owl habitat throughout the area of interest. It is
Neal Darby and TimYoung not important to have precise attribute information, only
that all GIS vegetation map features representing suitable
habitat be distinguishable from unsuitable habitat.
Introduction The GPNF GIS vegetation map and I-Veg table were
Private, State and Federal wildlife managers need used to establish a base GIS vegetation map (Figure 1, Step
planning tools that assist in determining the potential A). The I-Veg table is a database of forest stand attribute
occurrence of sensitive, threatened and endangered information (i.e., average tree diameter at breast height
species on their respective lands. These planning tools are (DBH), trees per acre, stand age, etc.). The I-Veg table is
particularly needed when wildlife managers lack the directly joined to the GIS vegetation map feature table by
physical or financial resources to conduct field surveys. matching fields (table columns) that contain identical
To meet this need a Geographical Information System information, such as a stand or polygon identifier number.
(GIS) model was developed to synthesize existing GIS Other GIS vegetation maps may not require a separate
vegetation maps and create a separate landscape level map table be joined, having the forest stand attribute
depicting areas with a high potential to contain nesting information in its feature table. Some GIS vegetation map
northern spotted owls. This model was designed for feature tables may have a field designating spotted owl
landscape level planning in areas where surveys for habitat. In fact, the I-Veg table contains a field titled
northern spotted owls may no longer be practicable. The “Spotted Owl Habitat.” Under the “Spotted Owl Habitat”
model uses desktop GIS technology and existing forest field, each polygon representing a forest stand in the GIS
inventory GIS data representing various northern spotted vegetation map is given one of four habitat descriptions:
owl habitat quality parameters (e.g., tree size, percent Unsuitable, Dispersal, Foraging or Nesting. Each habitat
canopy closure, tree species composition, etc.). Threshold description is based on forest stand attribute information
levels used to delineate high potential nesting areas were found under other fields in the I-Veg table for each
based on spatial analyses of habitat located around known respective polygon. Suitable northern spotted owl habitat,
northern spotted owl locations and nest sites as identified as defined here, consisted of stands designated as Foraging
in the literature. The ability to identify areas that have a and Nesting in the GPNF I-Veg table and included forest
high potential to support northern spotted owl nesting
 562 Wildlife–Habitat Relationships in Oregon and Washington

Figure 1. Stepwise process (A-F) to

create a map displaying potential
northern spotted owl nesting site
areas. Statements outside boxes
represent ArcView GIS Spatial
Analyst menu commands used to
generate each subsequent GIS map.
 CHAPTER 22: MODELING—CASE HISTORIES 563

assigning pixel values based on overlaying polygon

attribute information, pixel values in the Distance Grid
Map depict the actual distance of the pixel from selected
Unsuitable habitat. As a result, distance bands are created
from the edges of unsuitable habitat, consisting of
openings and early seral forests, into mid seral forests and
suitable habitat. This allows for the elimination of edge-
affected areas, defined by a specified edge width, to
highlight interior forest habitat on the landscape. Also,
by having distance bands radiating through unsuitable
habitat that consist of mid seral forest, a GIS edge is not
created between mid and late seral forests.
The display of forest interior patches is a better indicator
of forest fragmentation because it quantifies landscape
composition and configuration.9 The higher the amount
of fragmentation the less interior forest that is available.
Research on forest landscape patterns around known
spotted owl nest sites has indicated that the quantity of
forest interior habitat patches is important for continued
nest site use.3, 6, 7, 8, 10
The next step simplified the values of the Habitat Grid
Map and combined the Habitat and Distance Grid Maps
using the Map Query command (Figure 1, Step C1-2). The
Map Query command allows for selecting specific values
from one or more grid maps and combining them to create
a new grid map with pixel values of 0 and 1. The value of
1 represents selected values from the Habitat and Distance
Figure 2. Extent of spotted owl suitable habitat on the Gifford Grid maps and the value 0 represents the rest of the Habitat
Pinchot National Forest (solid gray pattern) and areas and Distance Grid maps’ extent. Using the Map Query
modeled as a high potential to support spotted owl nesting
command on the Habitat Grid Map, pixels valued as 2
sites (crosshatch).
(Foraging) and 3 (Nesting) were selected and queried
creating a Suitable Habitat Grid map (Figure 1, Step C1).
stands approximately 80-years-old and older, an average This created a grid map whose pixel value of 1 outlined
DBH >41 cm, one or more canopy layers, and an average northern spotted owl suitable habitat across the GPNF
canopy closure >40 percent. landscape (Figure 2).
Once the base GIS vegetation map displaying all To create a grid map displaying interior northern
spotted owl habitat attributes was established, it was then spotted owl suitable habitat, the Map Query command was
converted into a grid map using Spatial Analyst’s Convert used to combine selected values from the Habitat Grid
to Grid command (Figure 1, Step B). The Convert to Grid map and Distance Grid map. Pixel values 2 (Foraging)
command involved overlaying the vector (polygon) base and 3 (Nesting) from the Habitat Grid Map were selected
GIS vegetation map with a grid pattern. Each pixel in the and, from the Distance Grid Map, pixel values
grid is assigned a value (a whole number beginning with representing distances >100 meters from an edge were
1) representing the attribute it mostly overlays on the selected. After completing the query a new Interior
vector based map. Thus, the command created a Habitat Suitable Habitat Grid Map was created depicting only
Grid Map with pixel values of 1 for Dispersal; 2 for suitable habitat >100 meters from an edge. The 100 m forest
Foraging; 3 for Nesting; and, 4 for Unsuitable spotted owl edge buffer was based on work by Johnson.6 He noted
habitat. that spotted owl nest sites were predominately located
In addition to the Habitat Grid Map described above, >100 m from forest edges.
a grid map depicting interior forest habitat is needed Step D involves summarizing or, in this case, totaling
(Figure 1, StepB2). Interior forest habitat is defined here the number of pixels valued as 1 within a specified
as an area of suitable northern spotted owl habitat beyond distance from each individual pixel for both the Suitable
a specified edge distance from Unsuitable habitat. Creating Habitat Grid Map and Interior Suitable Habitat Grid Map
an interior forest habitat grid map involved selecting all (Figure 1, Step D). This involves using the Neighborhood
Unsuitable habitat that consisted of openings or early seral Statistics command in Spatial Analyst. The specified
forest stands on the base GIS Vegetation Map and then distance to evaluate on the GPNF was a 1.26 km radius
executing Spatial Analyst’s Find Distance command (Figure (500 ha). Research has noted the importance of the quantity
1, Step B2). As with the Convert to Grid command, the Find of northern spotted owl suitable habitat within an
Distance command overlays the vector based GIS approximate 500 ha area surrounding a nest site. The
Vegetation Map with a grid pattern. But, instead of quantity of suitable habitat within a 500 ha area appears
 564 Wildlife–Habitat Relationships in Oregon and Washington

to influence nest site location at landscape scales.5, 7, 8, 10 In 30 percent of the 500 ha area summarized in step D1. The
addition, this area receives concentrated use as a foraging threshold value of 146 ha used to create the Reclassified
site for the female and as a staging area for the owlets Interior Suitable Habitat Grid Map was derived from
after they leave the nest but prior to dispersal.4 Johnson6 and Ripple et al.10 Johnson6 stated that spotted
Two grid maps were created following steps D1 and owl nest sites in central Oregon were predominately
D2: a Summed Suitable Habitat Grid Map and a Summed greater than 100 m from the forest edge, indicating the
Interior Suitable Habitat Grid Map. The value for a pixel importance of interior forest for nest site location.
in each new grid map being the sum of the values for all Furthermore, Johnson6 found the majority of spotted owl
pixels surrounding it within a 1.26 km radius. responses to survey calls occurred in areas that contained
Each pixel value in the Summed Suitable Habitat and >146 ha of interior suitable habitat. Johnson6 defined
Interior Suitable Habitat Grid Maps was then converted interior forests as mature and old-growth stands buffered
to display the total area in hectares of suitable habitat and 100 m from an opening. In addition, Johnson6 noted that
interior suitable habitat that surrounds the respective pixel areas with <146 ha of interior suitable habitat contained
within the 1.26 km radius. To determine the area the majority of great horned owls. Carey et al.3 also noted
represented by the pixel value, the pixel value was an increased occurrence of great-horned owls in
multiplied by the area of the pixel. Back in steps B1 and fragmented landscapes as compared to larger intact,
B2 when the Convert to Grid and Find Distance commands forested landscapes.
were executed the user is prompted for pixel size. Thus, if For the final step, step F, the Reclassified Suitable and
a pixel size of 50 m was chosen: each pixel would represent Reclassified Interior Suitable Habitat Grid Maps were
an area of 2500 m2 or about 0.25 ha. If a summed pixel joined together using the Map Calculator command (Figure
value from the Summed Suitable Habitat Grid Map is 100 1, Step F). This step involved adding the weighted pixel
(100 pixels representing suitable habitat within 1.26 km), values of each overlaying pixel from each reclassified grid
multiply 100 by 0.25 ha to obtain the area (25 ha) of suitable map and dividing by the number of grids being added
habitat within 1.26 km of that pixel. together. This was a simple step since only two grids were
With this understanding of the pixel values in the combined. The grid with the most restricted area (a pixel
Summed Suitable Habitat and Summed Interior Suitable value of 2 for each map) would be the result (Figure 2).
Habitat Grid Maps, they can be reclassified using the Because only two grids were joined with weighted values
Reclassify command (Figure 1, Step E). The Reclassify of 1 and 2, the final Spotted Owl Nesting Potential Grid
command allows grouping of pixel values to represent Map exhibited pixel values of 1 and 2. Pixels with a value
ranges of specified values. The specified values are of 1 on the Nesting Potential Grid Map represented an
provided by the user and should, for purposes of area with a low potential to support a spotted owl nest
delineating high potential nesting areas, represent values site due to inadequate amounts of suitable habitat and
greater than the minimum area of the habitat parameters interior suitable habitat in a surrounding 500 ha area.
(e.g., suitable habitat and interior suitable habitat) shown Pixels with a value of 2 would represent an area with a
to support northern spotted owl nesting. Reclassified pixel high potential to support spotted owl nest sites due to
values need to be weighted when reclassified so that the sufficient quantities of suitable habitat and interior suitable
grid maps can be joined in the final step. For the analysis habitat within the surrounding 500 ha (Figure 2). The
on the GPNF, summed pixel values <150 ha in the Spotted Owl Nesting Potential Grid Map was converted
Summed Suitable Habitat Grid Map were reclassified to to a shapefile using the Convert to Shapefile command, so
a weighted pixel value of 1, while summed pixel values it could be overlain with other GIS vector map layers.
>150 ha were reclassified to a weighted pixel value of 2,
creating a Reclassified Suitable Habitat Grid Map. Examples
Likewise, summed pixel values in the Summed Interior An example of how the model’s Spotted Owl Nesting
Suitable Habitat Grid Map representing <146 ha and >146 Potential Grid Map could be used is as follows. A timber
ha were reclassified to weighted pixel values of 1 and 2, sale is proposed and the GIS vegetation map depicts the
respectively, creating a Reclassified Interior Suitable entire planning area with extensive coverage of suitable
Habitat Grid Map. habitat for the spotted owl, though it has been subjected
In the example for the GPNF, 150 ha and 146 ha were to past harvest activities. Spotted owl surveys have not
the threshold values that delineated low potential nesting been conducted and are not planned for the area.
areas from high potential nesting areas. The selective use Therefore, it is not known where spotted owls are nesting
of areas containing a higher proportion of mature and old- and it must be assumed that all suitable habitats are
growth forest (suitable habitat) by nesting spotted owls is occupied. Without further information, measures
supported in scientific literature. 2, 3, 6, 7, 8, 10 Bart and designed to minimize adverse impacts to spotted owls,
Forsman,2 Bart1 and Ripple et al.10 have further shown that such as timing restrictions on when logging could occur,
reproductive success may be depressed when the would be implemented across the entire planning area.
proportion of suitable habitat falls below 30 percent of a The impact of these restrictions could make the timber
1.26 km radius circle, as well as larger radius circles. The sale more difficult and costly to implement.
threshold value of 150 ha used to create the Reclassified However, after running the model for the planning area,
Suitable Habitat Grid Map for the GPNF approximates a wildlife manager could overlay the Spotted Owl Nesting
 CHAPTER 22: MODELING—CASE HISTORIES 565

Potential Grid Map with the proposed project map. After information to differentiate unsuitable and suitable
assessing the degree of overlap of High Potential Nesting spotted owl habitat. However, the model is flexible in that
Areas to proposed project sites and, accounting for additional GIS map layers can be incorporated to further
personal knowledge of habitat conditions and spotted owl define suitable habitat. For example, a GIS layer showing
use, the wildlife manager could suggest protective barred owl, a competitively advantaged species over the
measures for the spotted owl be applied specifically to spotted owl, distribution could be processed through each
those areas within High Potential Nesting Areas. No iterative step to develop a Reclassified Barred Owl Grid
protective measures to protect spotted owl nesting would Map with weighted pixel values. This layer could then be
be provided in Low Potential Nesting Areas. This process joined with the Reclassified Suitable Habitat and
would allow more flexibility in planning and conducting Reclassified Interior Suitable Habitat Grid Maps under
the timber sale while still considering the needs of spotted step F. Some iterative steps could be skipped to
owls. This is clearly shown by comparing the extent of accommodate map layer modifications. The final GIS map
the Spotted Owl Nesting Potential Grid Map to the GPNF can then be saved and used as an overlay for project
GIS vegetation map of spotted owl habitat types (Figure planning.
2). An analysis of the GPNF shows 107,030 ha, or It is recommended that any GIS map layer used in the
approximately 49 percent of the total area of suitable model should be reviewed based on local knowledge,
spotted owl habitat (218,070 ha) shown in the GPNF GIS personal judgment, and field reconnaissance. It is also
vegetation map was identified as having a high potential recommended that the model be run every time the GIS
to support spotted owl nesting (Figure 2). vegetation maps are updated.
For a second example, another timber sale is planned Threshold levels used in the High Potential Spotted
in an area known to have supported spotted owls in the Owl Nesting Area Model to differentiate low and high
past. After selecting the general area in which to conduct potential nesting areas were derived from published
the sale, the Spotted Owl Nesting Potential Grid Map scientific literature.2, 3, 6, 7, 8, 10These studies exhibited the
could be used as a planning and project design tool. One common occurrence of a high proportion of suitable
alternative would be to place units that fall outside of areas habitat and large suitable habitat patch sizes (interior
depicted as having a high potential to support spotted habitat) around spotted owl nest sites.
owl nesting. A second alternative could be to place units To determine if modeled results were realistic, known
in forest patches that constitute primarily edge, to maintain spotted owl activity center locations on the GPNF were
the highest quantity of interior suitable habitat that defines compared with the Spotted Owl Nesting Potential Grid
high potential spotted owl nesting areas. A third Map to see how well they correlated. A Chi-square test of
alternative would be to propose silvicultural prescriptions independence showed highly significant distributions
that would maintain the high quantity of suitable habitat. with modeled areas containing both >30 percent suitable
Such silvicultural prescriptions could include higher habitat (p <0.0001) and >146 ha of interior suitable habitat
canopy retention levels and protection of large trees and (p <0.01) within a 1.26 km radius circle. Still, more activity
snags. Thus, harvest units could be placed and designed centers were located in areas depicting low potential for
to minimize impacts to both suitable habitat and the spotted owl nesting. This was because since the
spotted owl. establishment of most activity centers, timber harvest had
As a final example, the Interior Suitable Habitat Grid reduced suitable habitat below these thresholds.
Map derived in Step C2 of the model (Figure 1, Step C2) Furthermore, scattered surveys for spotted owls on the
could be used to analyze effects of proposed projects on GPNF have not established any nest sites or activity
interior forests and the extent of forest fragmentation that centers within areas modeled as low potential nesting
may occur. Forest fragmentation is a common issue areas. Three new activity centers, including a nest site,
brought out during scientific and public review. By were all located within modeled high potential nesting
overlaying proposed project areas on the interior suitable areas. Another weakness involves the exclusion of
habitat grid map, the amount of interior suitable habitat contiguous suitable habitat stands along the periphery of
that would be lost or affected and the amount of edge high potential spotted owl nesting areas. This is because
created as a result of the proposed project could be easily of the nature of the Neighborhood Statistic command. As
assessed. neighborhood statistics are derived for each pixel, fewer
pixels valued as habitat are summed as it goes further from
Conclusions a habitat patch. Therefore, the edges of a large suitable
The High Potential Spotted Owl Nesting Area Model was habitat patch would be delineated as low potential spotted
designed for use on ArcView GIS software with the Spatial owl nesting areas due to insufficient quantities of suitable
Analyst extension. ArcView is commonly used by and interior suitable habitat on one side, even though they
numerous agencies to meet their GIS needs so there is no are the edges of a large contiguous, interior patch.
need to download a specific software or program and train Another similar situation involved the delineation of a
for its use. All commands to run the iterative steps in the circular high potential area that, when overlaid with the
model are located in ArcView’s default menus. suitable habitat map, mostly consist of unsuitable habitat
The only data requirement is a GIS vegetation map with suitable habitat located around the periphery. This
layer that contains sufficient forest stand attribute occurs in highly fragmented areas. Since the Neighborhood
 566 Wildlife–Habitat Relationships in Oregon and Washington

Statistics command summarizes each pixel regardless of

whether it is suitable or unsuitable, the high potential area
is situated over the landscape that maximizes the greatest
amount of suitable habitat in a 1.26 km radius.
These two situations can be overcome by selecting
suitable habitat polygons that intersect high potential
spotted owl nesting areas, but only when the polygons
do not cover more than approximately 100 ha. If polygons
cover extensive areas, then buffering the high potential
spotted owl nesting area designation by 0.25 mile appears
to incorporate the majority of stands that extend beyond
the boundary. It would be advised that biologists
determine the status of suitable habitat stands along the
periphery of modeled high potential spotted owl nesting
areas before accepting any decision of it being a low
potential spotted owl area.
Finally, this process need not be restricted to an analysis
for spotted owls. Other habitat parameters for other
species could be developed and modeled much like were
done here. An array of similarly modeled spatial
relationships of wildlife and their habitats could greatly
assist in ecosystem level planning and project design.

Literature Cited
1. Bart, J. 1995. Amount of suitable habitat and viability of northern
spotted owls. Conservation Biology 9:943-946.
2. Bart, J. and E. D. Forsman. 1992. Dependence of northern spotted owls
on old-growth forests in the western USA. Conservation Biology
62:95-100.
3. Carey, A. B., S. P. Horton, and B. L. Biswell. 1992. Northern Spotted
Owls: Influence of prey base and landscape character. Ecological
Monographs 62:223-250.
4. Forsman, E. D., E. C. Meslow, and H. M. Wight. 1984. Distribution and
biology of the spotted owl in Oregon. Wildlife Monographs 87:1-64
5. Hanson, E., D. Hays, L. Hicks, L. Young, and J. Buchanan. 1993. Spotted
owl habitat in Washington: A report to the Washington Forest
Practices Board. Final Report.
6. Johnson, D. H. 1993. Spotted owls, great horned owls, and forest
fragmentation in the central Oregon Cascades. M.S. Thesis. Oregon
State University, Corvallis, OR.
7. Lehmkuhl, J. F. and M. G. Raphael. 1993. Habitat pattern around
northern spotted owl locations on the Olympic Peninsula, Washington.
Journal of Wildlife Management 57:302-315.
8. Meyer, J. S., L. L. Irwin and M. S. Boyce. 1992. Influence of habitat
fragmentation on spotted owl site location, site occupancy, and
reproductive status in western Oregon. Progress Report.
9. McGarigal, K., and B. J. Marks. 1995. FRAGSTATS: Spatial pattern
analysis program for quantifying landscape structure. General Technical
Report PNW-GTR-351. USDA, Forest Service, Pacific Northwest
Research Station, Portland, OR.
10. Ripple, W. J., P. D. Lattin, K. T. Hershey, F. F. Wagner and E. C. Meslow.
1997. Landscape composition and pattern around northern spotted
owl nest sites in southwest Oregon. Journal of Wildlife Management
61:151-158.
 23
Integrating Wildlife Species Habitat Goals and
Quantitative Land Management Planning Processes
Pete Bettinger, Kevin Boston, John Sessions, & William C. McComb

Introduction
Land management planning is an iterative process. Goals spatial relationships of harvest units.6 As a result of
are developed and adjusted during planning processes, managing forest land within regulatory frameworks, forest
and desired outcomes are displayed in a management management planning now often requires multiple
plan. Wildlife habitat goals are becoming more common resource goals and the use of spatial rules for the selection
in management plans, and resource allocation models are of timber harvest units.17
used to attempt to resolve conflicts among management In this chapter we illustrate how to develop and
goals. For example, one goal may be to provide an communicate wildlife habitat goals and provide an
adequate supply of habitat for one or more selected example of how they can be included in land management
species, while another may be to provide a minimum planning processes. To accomplish this, we first briefly
amount of timber harvest volume. Quantitative land describe a generic, yet typical, planning process. Next we
management planning techniques can be used to represent describe several types of wildlife species habitat goals that
multiple resource goals, and to plan activities that are could be used in natural resource planning. We then define
compatible with resource sustainability. several common terms used in quantitative management
Quantitative planning approaches commonly have planning and illustrate their use with the species habitat
helped guide silviculture and the biological and economic goals. Several management plans are developed with these
management of forests. Using quantitative goals or goals in mind. Finally, we discuss four important areas of
objectives implies that desired conditions have been planning and their influences on management planning.
defined for a planning area. These goals and objectives
may change over time, as societal, regulatory, or Quantitative Management Planning
organizational goals change. For example, in the Pacific
Northwest we have seen a shift during the last two
Processes
decades from commodity production objectives to forest We undertake planning processes every day. We decide
structure objectives on public lands. Quantitative planning what we want to do, express some impression of what
methods can be applied to a wide range of landowner the desired goals should be, examine a few ways to achieve
goals (e.g., social, biological, economic) to provide the goals, pick an alternative, implement the alternative,
managers with management alternatives. and perhaps assess the results. Several examples come to
Planning should be viewed as a continuous and mind: driving to work (how do I get there? when should
incremental process that assists managers in making I leave? which route do I take?); shopping for groceries
decisions. The decisions faced by each organization could (what do I need to purchase? what do I want to purchase?
require short- or long-term plans, or strategic (forest-wide) what is my budget? what groceries do I already have at
or tactical (site-specific) plans. The potential limitations home?); and writing a report (when is it due? how long
to planning generally fall into four categories: should it be? what other tasks must I accomplish during
organizational, personnel, database, and technological. this same period?).
Despite the limitations, planning helps managers (1) make Organizations may undertake planning processes less
decisions, (2) establish credibility, (3) accomplish tasks frequently, yet the steps in planning are similar, and
efficiently, (4) achieve continuity in management, and (5) certainly more formal. Formal planning processes can take
provide a standard for comparison against other plans and many forms, but generally consist of the following tasks:
monitoring results. (1) define the scope of the planning problem, (2) articulate
The spatial arrangement of wildlife habitat and forest future goals, (3) isolate alternative solutions to the
management activities is important for a number of problem, (4) set criteria for evaluating the alternatives, (5)
reasons, such as adhering to regulatory rules, complying evaluate the alternatives, (6) choose an alternative for the
with organizational goals and policies, and maintaining plan, (7) monitor the plan, and (8) reevaluate alternatives
aesthetic conditions. Forest regulations, for instance, are after implementation.
placing increasingly restrictive limits on the size and

567
 568 Wildlife–Habitat Relationships in Oregon and Washington

The nature of planning problems ranges from those 2. Cooper’s hawk prefers 30- to 70-year-old even-aged
with relatively simplistic goals (e.g., habitat management conifer stands with a mean stand density of 367
for ruffed grouse in aspen forests), to problems that may trees/acre (907 trees/ha).
include many resources and goals over large areas (e.g., 3. Northern goshawk prefers •150-year-old conifer
national forest planning). During planning processes, the stands with a mean tree density of 195 trees/acre
goals, objectives, and constraints should become apparent, (482 trees/ha).
and may be based on an organization’s mission, goals, or 4. Red tree vole prefers old-growth forests that are •195
objectives. Goals include outcomes such as providing years old.
habitat for certain wildlife species. An objective may be to A planning team would define how much area with
maximize the amount of habitat produced, timber volume these characteristics is desired (e.g., 100 acres, 1000 acres,
produced, or profit, or to minimize costs to achieve habitat as much as possible) in order to use these goals as
goals. objectives or constraints in a quantitative planning process,
Constraints may include minimum harvest ages for although they still could simply be measured. A relaxation
timber stands or budget limitations. These parts of of the tree density requirements would permit increased
planning processes are very important and will be flexibility in the achievement of forest goals. For example,
discussed in more detail later in this chapter. The isolation three of the goals could be restated as:
of alternatives and the determination of criteria for 1. Sharp-shinned hawk prefers 25- to 50-year-old even-
evaluating alternatives often involve the use of decision aged conifer stands with a mean tree density of 405-
models. The objectives should be quantitative, providing 525 trees/acre (1000-1300 trees/ha).
criteria by which plans are judged and their 2. Cooper’s hawk prefers 30- to 70-year-old even-aged
implementation monitored. conifer stands with a mean stand density of 324-405
Much of the remainder of this chapter focuses on three trees/acre (800-1000 trees/ha).
types of wildlife goals that could be used in planning 3. Northern goshawk prefers •150-year-old conifer
problems. These goals are categorized as either spatial or stands with a mean tree density of 174-214 trees/acre
nonspatial. After discussing the goals in general terms, (430-530 trees/ha).
we define common terminology that could be used
throughout planning process discussions. This Spatial Goals: Minimum Patch Size
terminology is directed toward the professional involved Some forest planning goals use spatial characteristics of
with natural resource management. the landscape to determine their value to particular
wildlife species. One type of spatial goal might be to
Wildlife Species Habitat Goals require patches to be a minimum size before they
contribute positively toward the achievement of habitat.
Species habitat goals can be either qualitatively or
For example, the varied thrush, winter wren, and
quantitatively defined. Whereas qualitative goals may be
Hammond’s flycatcher may need intact stands of single
important for certain organizations, this chapter focuses
story open canopy forests •49.4 acres (•20 ha) in size.
on quantitative goals. Quantitative goals for wildlife
From a planning perspective, the definitions of single
species can be categorized in many ways: do the goals
story open canopy forest are important. While structural
require a measurement of snags per unit area; do the goals
attributes of forests (i.e., variation in tree diameter classes,
involve land classification (uplands vs. riparian areas); are
level of coarse woody debris, vertical complexity, etc.)
the goals expressed as linear or nonlinear equations? One
could be used to define these conditions, we will assume
categorization on which we concentrate is whether the
for discussion purposes that these are forested stands •90
goals require spatial information to measure their
years old. The minimum size requirements could then be
attainment. We describe three wildlife species habitat goals
met with individual stands that are •49.4 acres (•20 ha)
below; two of them require spatial information in their
and •90 years old. The goal could also be met with groups
computations. Nonspatial goals may consist of assembling
of contiguous stands, where the sum of the area is •49.4
a number of acres in certain age classes, or strata. Spatial
acres (•20 ha), and all stands have forests •90 years old.
goals may include configurations such as requiring
Finally, the planning team would need to determine, for
minimum patch sizes, or adjacent habitat types.
the analysis area, how many of these patches would be
Nonspatial Goals required (e.g., provide at least 3 distinct patches/mile2
These types of planning goals generally are based on [7.7/km2]), or how much area with these characteristics is
achieving the amount of some resource in a planning area. required (e.g., 100 acres, 1000 acres, as much as possible).
For example, goals could be developed with the criteria
Spatial Goals: Adjacency Requirements
that some amount of habitat, such as mature closed canopy
Goals for wildlife species may also be developed that
forest, will be achieved. Examples of nonspatial goals
indicate, for optimal benefit to a particular species, that
related to the type of nesting habitat required for four
one type of habitat should be placed next to another. For
wildlife species in the Pacific Northwest include:
example, the great gray owl prefers early seral stage forests
1. Sharp-shinned hawk prefers 25- to 50-year-old even-
(clearcuts) for foraging, yet these areas should be adjacent
aged conifer stands with a mean tree density of 477
to single story open canopy forests containing snags or
trees/acre (1,180 trees/ha).
 CHAPTER 23: INTEGRATING GOALS 569

large trees with broken tops. Alternatively, great gray owls Activities and Decision Variables. Activities include the
may show a preference for older forest stands adjacent to suite of actions that can be used to achieve the goals (e.g.,
permanent meadows. silvicultural activities), and are essentially the types of
Further clarification of this goal is important. For projects that can be implemented to achieve the goals.
example, do all early seral stage forests need to be adjacent Management activities could include a broad variety of
to single story open canopy forests, or just some portion silvicultural prescriptions. For example, we could thin a
of them? Conversely, the amount of single story open timber stand at age 30 years, and remove 25 ft2 /acre (5.7
canopy forests adjacent to early seral stands might be of m2/ha) of tree basal area, or, we could remove 30 ft2 /acre
interest. At some point a quantitative measure or threshold (6.9 m2/ha) of tree basal area, or, we could create 10 snags/
must be defined before the goal can be used in a planning acre (24.7 snags/ha), etc. Fertilization treatments, the
process. We will proceed with a rather simple assumption: creation of snags, leaving trees on a harvest site at the time
acceptable habitat for great gray owls requires •49.4 acres of regeneration harvest, and other silvicultural treatments
(•20 ha) of single story open canopy forest adjacent to •24.7 are all activities that can be combined to produce a single
acres (•10 ha) of early seral habitat. forest management prescription.
Decision variables represent the level or levels to which
Placing Wildlife Species Habitat Goals in a each activity is assigned (e.g., land units, roads). For
Management Planning Context example, an activity (thinning) may be applied to all, or
The process we suggest in the development of a part of, a decision variable (a timber stand), to open up
management plan consists of describing the planning the canopy and promote vertical structure complexity.
problem in words, then converting the descriptions to
Objective Function. An objective function is a
quantitative relationships. In the early stages of planning
mathematical statement of a goal, or goals, combining the
it may be beneficial for those involved in developing
criterion for measurement of goal achievement and the
management plans to identify the decision-maker. The
decision variables specified for the problem. Although
decision-maker is a person or group who has designated
many problems in business and engineering adopt a single
stewardship responsibilities, who chooses the policies or
goal (often a statement to maximize profit or minimize
actions that will be undertaken, and thereby accepts
cost), a broader definition is often useful in wildlife
responsibility for the choice of management plan.
management planning. The solution with the best
The applied economics and management science fields
objective function value, all else being equal, will be chosen
have defined several terms that are commonly associated
as the best solution. The objective function value serves
with quantitative management planning processes. We
as a decision guide for sorting through the possible feasible
place the goals we discussed above into context using
decision variable combinations to determine what
some of these terms. First we will restate the goals, and
combination is best according to the criteria used to
then develop goal criteria. Then we will determine the
evaluate the objective function.
activities and decision variables that could contribute to
Goals for wildlife management planning can be
goal achievement. Finally, we describe the objective
expressed using one of several forms: (1) maximize or
function and constraints of the resulting problems.
minimize 1 goal with the other goals expressed as
Goals. Goals are the outcomes of a planning process; they constraints, (2) minimize deviations from •1 goal with
are relevant to the planning problem and are expected to other goals expressed as constraints, or (3) a hybrid or
be achieved (e.g., increase habitat for marbled murrelets). combination of the previous 2 forms. In form 1, the goals
These outcomes are useful when they can be stated in represented as constraints must be met and the goal in the
quantitative terms, for example: (1) to provide the greatest objective function might be viewed as the residual goal
amount of varied thrush habitat, or (2) to provide the that is achieved to the extent possible given the goals
greatest amount of habitat for all species, each species represented as constraints. In form 2, the objective function
given equal value. is expressed as the sum of the deviations from targets for
•1 goal and the remaining goals are expressed as
Goal Criteria. Goal criteria measure goal achievement,
constraints. Using form 2 the decision-maker must
and help to determine whether or not the activities
develop targets for the goals and the optimization attempts
employed are, or will, move a landscape toward or away
to minimize the deviations between the targets and the
from goals. For example: (1) goal: to provide the greatest
goal achievement while satisfying any remaining goals
amount of varied thrush habitat, and (2) goal criterion:
represented as constraints. Form 2 is sometimes referred
the amount of land considered to be varied thrush habitat
to as goal programming. Goal programming can involve
in each planning period. Multiple goals could tend to
either linear or nonlinear programming models. In Form
compete against one another (e.g., sharp-shinned hawk
3 the objective function maximizes or minimizes 1 goal
and red tree vole). With multiple goals, it may be hard to
minus the sum of the deviations from targets of •1 goal.
determine whether a particular management plan
The remaining goals are represented as constraints.
achieves, in sum, more of the multiple goals than another
plan does, and if so, by how much. How the multiple goals Constraints. Anything that limits the achievement of a
and goal criteria are designed is a major factor in goal (i.e., a minimum amount of marbled murrelet habitat
evaluating which goal may be more influential in the must be maintained) is a constraint. Usually constraints
development of a management plan.
 570 Wildlife–Habitat Relationships in Oregon and Washington

arise from resource limitations (e.g., cannot schedule Table 1. Problem formulation for the nonspatial
activities to more than what you manage), decision-maker species habitat goals.
goals (e.g., need to produce X amount of timber volume/
year), or externally imposed policies or regulations (e.g.,
do not harvest within Y feet of an owl location). A Goals:
management plan is assumed to be feasible when all To provide the greatest amount of sharp-shinned hawk habitat
over time.
constraints are met. Of course, the level of the objective
To provide the greatest amount of Cooper’s hawk habitat over
function after meeting the constraints may not be
time.
acceptable, and new activities and a revision of the
To provide the greatest amount of northern goshawk habitat
constraints may be needed. over time.
Next we will define the goals, goal criteria, activities, To provide the greatest amount of red tree vole habitat over
decision variables, objective function, and constraints for time.
a planning problem that uses the species goals we have
described above. Then we will make some assumptions Goal criteria
about a planning exercise for a hypothetical forest. Finally, Amount of land considered to be sharp-shinned hawk habitat in
we illustrate several management plans that emphasize each planning period.
the achievement of the maximum amount of each of the Amount of land considered to be Cooper’s hawk habitat in each
three types of goals. planning period.
Amount of land considered to be northern goshawk habitat in
Nonspatial Goals each planning period.
Amount of land considered to be red tree vole habitat in each
The nonspatial goals we have defined for the four wildlife
planning period.
species in the Pacific Northwest are to provide the greatest
amount of habitat for each over the length of the planning Activities and decision variables
horizon (Table 1). The criteria to evaluate the goals are to Silvicultural activities include no harvest or regeneration harvest.
measure the amount of land considered to be habitat for Decision variables are individual land units allocated to
each species, in each planning period, and divide by the silvicultural activities.
total number of acres on the forest, to arrive at a percentage Objective function
of land in habitat for each species. The tree age and species Maximize the percent of land considered to be habitat for each
composition on these land units can be used to define species in each planning period.
whether habitat goals have actually been achieved for each
species. To simplify our example, we assume that the age maximize
of the stand is the most important factor, and assume that
the composition of trees within the stand meet the Where: i = land unit
requirements for each species. A more detailed analysis j = time period
could incorporate growth and yield estimates to determine k = wildlife species
the estimated number of trees/acre (ha), or the structural Ai = acres in land unit i
composition of stands (to determine whether they meet a Hijk = habitat for species k, on land unit i, during
criterion for old growth). planning period j (binary, 0 for no, 1 for yes).
The activities we will illustrate include just two options: The determination of habitat is a function of
regeneration or no harvest. Activities that affect the status average tree species age.
of decision variables, and either contribute positively or Constraints:
negatively to the condition of the decision variables (with Only 1 regeneration harvest can occur during the planning
respect to the goals) could include management actions horizon.
such as thinning, regeneration harvests, fertilization
treatments, and snag creation, although here we model
regeneration harvests only. The decision variables for
sharp-shinned hawk, Cooper’s hawk, northern goshawk, Where: Xij = binary (0 for no, 1 for yes) variable indicating a
and red tree vole are units of land allocated to various age regeneration harvest on land unit i during
classes. If the decision variables and the activities are planning period j
combined, we can track the amount of land, within a land Timber volume produced must be above a minimum volume.
unit, that is scheduled to receive a silvicultural treatment.
minimum harvest value
This implies that not all of a timber stand, for example,
could be harvested. We assume, however, that if a land
Where: Vij = timber volume on land unit i during planning
unit is scheduled for an activity, the activity will occur on
period j
the entire land unit (i.e., it is a binary decision). This
assumption makes choosing the size of the average timber
stand important.
The objective function for this problem is to maximize considered habitat for each species, while satisfying the
the amount of land, for all four species, that can be constraints (Table 1).
 CHAPTER 23: INTEGRATING GOALS 571

Spatial Goals: Table 2. Problem formulation for the minimum patch

Minimum Patch Size size species habitat goal.
The minimum patch size spatial goals for varied thrush,
winter wren, and Hammond’s flycatcher are to provide Goals:
the greatest amount of habitat over time (Table 2). The To provide the greatest amount of varied thrush habitat over
criteria we use to measure these goals consist of measuring time.
the percentage of land, in each planning period that meets
the habitat requirements. The activities and decision Goal criteria:
variables are similar to those described for nonspatial Amount of land considered to be varied thrush habitat in each
goals. The objective function is also similar to that of planning period.
nonspatial goals, yet only maximizes the percentage of Activities and decision variables:
land in patches •49.4 acres (•20 ha), and •90 years old. Silvicultural activities include no harvest or regeneration harvest.
The definition of habitat could, in this case, be considered Decision variables are individual land units allocated to
a constraint, because the aggregation of land units •90 silvicultural activities.
years old may be required before they are (in aggregate)
Objective function:
•49.4 acres (•20 ha) in size. The other constraints follow
Maximize the percent of land considered to be habitat for each
those of the nonspatial goals.
species in each planning period.
Spatial Goals:
maximize
Adjacency Requirements
The adjacent patch spatial goal for the great gray owl is to Where: i = land unit
provide the greatest amount of habitat over time (Table j = time period
3). The criteria we use to measure these goals consist of k = wildlife species
measuring the percentage of land, in each planning period Ai = acres in land unit i
that meets the habitat requirements. The activities and Hijk = habitat (binary, 0 for no, 1 for yes) for species k,
decision variables are similar to those described for on land unit i, during planning period j. The
nonspatial goals. The objective function is also similar to determination of habitat is a function of
that of nonspatial goals, yet only maximizes the percentage average tree species age and size of
of land in patches •49.4 acres (•20 ha), and •90 years old, contiguous habitat.
and adjacent to patches •24.7 acres (•10 ha) and •10 years Constraints:
old. The definition of habitat could, in this case, be Only 1 regeneration harvest can occur during the planning
considered a constraint, because to be eligible to be horizon.
considered, the aggregation of land units •90 years of age
may be required before they are (in aggregate) •49.4 acres
(•20 ha) in size, and similarly an aggregation of patches
may be required for early seral forested stands before the Where: Xij = binary (0 for no, 1 for yes) variable indicating a
goal can be achieved. In addition, we add one more level regeneration harvest on land unit i during
of complexity to this problem by limiting the size of planning period j
clearcuts to •120 acres (•48.6 ha). We add this last Timber volume produced must be above a minimum volume.
constraint to illustrate an increasing complexity in
planning goals. The other constraints follow those of the minimum harvest value
nonspatial goals. Where: Vij = timber volume on land unit i during planning
period j
Developing a Management Plan Habitat patches must be larger than a minimum size.
We will now develop a management plan with the wildlife Hijk = 1 if size of contiguous habitat of •90 year-old forest is
goals we have developed in the preceding sections. •49.4 acres (•20 ha), otherwise Hijk = 0. The
Described below are the assumptions we use regarding a determination of habitat is made using an area
hypothetical forest landscape, including a description of restriction model, similar to that described in Murray
the initial wildlife habitat classes, how habitat types (1999), which is a recursive function that evaluates all
transition after regeneration harvests, and how we make adjacent units y to unit i, and all adjacent units z to unit
decisions concerning the activities that are included in the y, and so on.
management plan.
units (polygons) defining the forest, with an average size
Case Study Landscape of 33.8 acres (13.7 ha). The forest is considered to be
We use a hypothetical forest (Figure 1) to illustrate the primarily a conifer-dominated forest, with hardwoods
implementation of the wildlife species habitat goals. This dominating in the riparian areas.
forest is 2,500 acres (1,012 ha), and has an initial age class Management Activities. As mentioned earlier, we
distribution illustrated in Figure 2. There are 74 analysis simplify the management choices for this discussion. The
 572 Wildlife–Habitat Relationships in Oregon and Washington

Table 3. Problem formulation for the adjacent patch Constraints:

species habitat goal. Only one regeneration harvest can occur during the planning
horizon.

Goals:
To provide the greatest amount of great gray owl habitat over
Where: Xij = binary (0 for no, 1 for yes) variable indicating a
time.
regeneration harvest on land unit i during
Goal criteria: planning period j
Amount of land considered to be great gray owl habitat in each Timber volume produced must be above a minimum volume.
planning period.
minimum harvest volume
Activities and decision variables:
Silvicultural activities include no harvest or regeneration harvest. Where: Vij = timber volume on land unit i during planning
Decision variables are individual land units allocated to period j
silvicultural activities. Habitat patches must be of certain sizes, and adjacent to each
Objective function: other.
Maximize the percent of land considered to be habitat for each Hijk = 1 if size of contiguous habitat of •90-year-old forest •49.4
species in each planning period. acres (•20 ha), is adjacent to a contiguous area of •10-
year-old forest that is •24.7 acres (•10 ha) (or vice
maximize versa), otherwise Hijk = 0. The determination of habitat
is made using two area restriction models, similar to
Where: i = land unit that described in Murray (1999), which are recursive
j = time period functions that evaluate all adjacent units y to unit i, and
k = wildlife species all adjacent units z to unit y, and so on.
Ai = acres in land unit i Regeneration harvests must be equal to or smaller than a
Hijk = habitat (binary, 0 for no, 1 for yes) for species k, maximum size. An area restriction model is used to control
on land unit i, during planning period j. The the size of regeneration harvests (Murray 1999).
determination of habitat is a function of maximum clearcut size
average tree species age and size of
contiguous habitat. Where: Xyj = binary (0 for no, 1 for yes) variable indicating a
regeneration harvest on land unit y during
planning period j
Ni = set of land units adjacent to unit i
Si = subset of regenerated land units containing all
units adjacent to the neighbors of land unit i,
and all land units adjacent to neighbors of
neighbors, and so on.

timing of the regeneration harvest will be the only Length of Planning Horizon. Planning horizons vary
silvicultural choice. The quantitative management from a few years to several decades, depending on the
planning technique will determine when to apply the objectives of the organization(s) guiding the planning
regeneration harvest. Of course, one choice is that no process. Here, we assume that the planning horizon will
harvest will occur during the planning horizon. In be 50 years, and that the analysis will be divided into ten
summary, we will allow the following types of 5-year planning periods.
management activities: (1) regeneration harvest in any
Activity Scheduling Considerations. Most forest resource
land unit with an age •40 years; and (2) grow only, no
organizations have developed or adapted mathematical
harvest activity.
programming techniques to enable them to produce
Vegetation Transition after Regeneration Harvest. In management plans. On public lands in the western United
order to emulate a dynamic forested landscape, we could States, land management planning has been increasingly
develop and use vegetation type transition probabilities, emphasizing both economic efficiency (or commodity
which we could then use to determine a new wildlife production) and wildlife goals,3 such as the goals we have
habitat type after harvest. The decision on vegetation type described that require the achievement of certain levels
transitions would be based on such factors as previous of wildlife habitat. Forest planning algorithms that
forest type, and distance from the stream system. We have optimize the spatial arrangement of forest resources to
chosen, however, to use very simple transition rules for meet a set of management goals vary from the more
this chapter’s examples. If regeneration harvests occur, traditional optimizations techniques, such as linear or
we will assume that the resulting forest will return to the mixed integer programming,10 to heuristic techniques such
vegetation type that existed prior to harvesting. This is, as binary search and simulated annealing.15 These types
of course, a simplification of forest succession, especially of problems are generally considered to be difficult to solve
if intensive stand establishment activities occur. with traditional mathematical programming techniques.
 CHAPTER 23: INTEGRATING GOALS 573

Table 4. A general heuristic procedure.

Step Process

1 Number all land units from 1 to n.

2 Number all activities j eligible for each land unit i from 1
to ki.
3 Set the initial value of the objective function to a large
negative number.
4 Choose a land unit randomly by choosing a random
number between 1 and n.
5 Choose an activity to assign to that land unit by choosing
a random number between 1 and ki.
6 Calculate the value of the objective function.
7 If this value is better than the last, continue to Step 10,
otherwise return to Step 4.
8 Update the solution by incorporating the activity from
Step 5.
9 Have we reached a stopping criterion? In other words,
has the objective function value stabilized, or have we run
out of time?
10 If we want to continue, return to Step 4, or else stop.
0 - 10 years old
For example, mixed integer and integer programming
11 - 30 years old
techniques have been used to produce management plans
31 - 89 years old with adjacency concerns, but these types of techniques
have severe limitations (directly related to problem size)
90+ years old when applied to large combinatorial problems.13
Riparian area The potential types of mathematical programming
techniques that are available to develop management
plans with spatial goals are therefore limited, because of
Figure 1. The hypothetical forest that is used in the planning
the nonlinear, and integer nature of the resulting planning
exercise.
problem. This should not, however, discourage the use of
complex habitat goals. Heuristic programming techniques
300
have been applied to traditional forest harvest scheduling
problems11 as well as to forest transportation problems,15,
16, 18, 20, 21
wildlife conservation and management,1, 4, 9 aquatic
250
system management,5 and biological diversity.12 Although
a heuristic technique does not guarantee that a global
200 optimum solution can be identified for a particular
problem, it can produce a feasible, efficient solution to a
RA = Riparian area
complex problem in a reasonable amount of time.
150
The nature of the planning goals will be a major factor
in determining how difficult it may be to perform planning
100 exercises with each technique. This chapter does not delve
into the differences between these techniques, but on the
implementation of the goals. We have chosen a heuristic
50
programming technique (tabu search) to use as the
demonstration tool to help develop a management plan.
0 The general structure of tabu search has been previously
RA 5 15 25 35 45 55 65 75 85 95 105 115 125 135 145 155 165 175 185
described.2, 5, 7, 8 A traditional optimization technique (e.g.,
AGE CLASS (years)
integer programming) can be used to develop
Figure 2. Age class distribution of the hypothetical forest management plans for the nonspatial goals, however the
use of traditional optimization techniques is limited for
more spatially complex planning problems. For
comparison, we developed a management plan for the
nonspatial goals using an integer programming technique
as well as tabu search, and found the tabu search produced
a result which was within 0.02% of the integer
programming result.
 574 Wildlife–Habitat Relationships in Oregon and Washington

Generally, heuristic techniques operate on a set of rules

(see Table 4). Rule-based procedures allow the
development of a solution by moving from 1 location in a
“solution space” to another; in our example, this happens
by a single change in the status of a land unit. This
procedure will lead to “good” solutions, but may become
stalled at local optima and may not be able to reach other
areas of the solution space, where even better solutions
might be found. These procedures can be modified to
allow the acceptance of changes (in solutions) that lead
temporarily to decreases in the objective function value.
Procedures like this allow movement out of local optima
of the solution space, and thus allow the search process to
explore other arrangements of activities across a
landscape, which may prove to be better overall. Tabu
search, simulated annealing, and Monte Carlo simulation19
all have more sophisticated rules to allow search processes
to do just that. Figure 3. Results from maximizing nonspatial wildlife species
habitat goals.
Management Plan Alternatives
The management plans that we will illustrate include the
following three scenarios:
1. Maximize the nonspatial goals for the sharp-shinned
hawk, Cooper’s hawk, northern goshawk, and red
tree vole, subject to the following: minimum harvest
age constraints (40 yr), timber volume goals (3
million board feet per 5-yr period [approximately
19,000 m3 per 5-yr period]), and only one
regeneration harvest per land unit during the
planning horizon. Both of the spatial goals will be
measured in a posterior (after-the-fact) manner on
the resulting “best” solution found for the nonspatial
goals.
2. Maximize the spatial goal requiring a minimum patch
size for the varied thrush, subject to the following:
minimum harvest age constraints (40 yr), timber
volume goals (3 million board feet per 5-yr period
[approximately 19,000 m3 per 5-yr period]), and only
one regeneration harvest per land unit during the
planning horizon. Both the nonspatial goals and the
spatial goal requiring adjacent patch types will be Figure 4. Timber volume produced for three planning
measured in a posterior (after-the-fact) manner on problems.
the resulting “best” solution found for this spatial
goal.
3. Maximize the spatial goal requiring adjacent patch
types for the great gray owl, subject to the following:
minimum harvest age constraints (40 yr), timber
volume goals (3 million board feet per 5-yr period
[approximately 19,000 m3 per 5-yr period]), and only
one regeneration harvest per land unit during the
planning horizon. In addition, clearcut sizes must be
•120 acres (48.6 ha). Both the nonspatial goals and
the spatial goals requiring minimum patch sizes will
be measured in a posterior (after-the-fact) manner on
the resulting “best” solution found for this spatial
goal. Figure 5 (right). Cooper’s hawk habitat (shaded area) in the
In addition to illustrating increasingly complex wildlife initial landscape condition (top), after period 5 (middle), and
goals, these three plans will serve to illustrate the trade- after period 10 (bottom), when nonspatial goals are
offs planners encounter when the emphasis of a plan is maximized.
CHAPTER 23: INTEGRATING GOALS 575

Figure 6. Results from maximizing the spatial goal requiring

minimum patch sizes for varied thrush habitat.

placed on goals that may not necessarily be

complementary, but perhaps competitive.

Results from Case Study

We developed feasible solutions to the three planning
problems described earlier in this chapter. The solutions
can be considered good feasible solutions to the problem,
but not necessarily optimal solutions, since heuristic
techniques do not guarantee optimal solutions will be
located during the search process.

Maximizing the Nonspatial Goals

When we used the heuristic technique to maximize the
nonspatial goals, we found that the percentage of land in
sharp-shinned hawk and Cooper’s hawk habitat was quite
high, averaging 40% and 46%, respectively and tended to
increase over the last half of the planning horizon (Figure
3). Northern goshawk and red tree vole habitat types
averaged 8% and 4% per period, respectively, with the
goshawk goal remaining constant and the red tree vole
goal increasing gradually over time. The percentage of
land in varied thrush habitat (spatial goal requiring
minimum patch sizes) remained constant over time, and
averaged 4.5% of the landscape, whereas the percentage
of land suitable for great gray owl habitat (spatial goal
requiring adjacent habitat types) was relatively high in
the first and last periods, yet was negligible in the
intermediate periods, as older forest patch size was limited
and little harvesting occurred (creating few young seral
forests. The average great gray owl goal was 4.5%. Timber
volume per 5-year planning period averaged 8.2 million
board feet (approximately 51,500 m3) (Figure 4), although
it ranged from about 3 million (approximately 18,900 m3)
to about 27.6 million board feet (approximately 173,900
m3) per period. The large harvest area in the first period
was beneficial to both the sharp-shinned and Cooper’s
hawk goals in the later periods. The increase in harvest
volume in period 10 had little effect on any of these goals,
576 Wildlife–Habitat Relationships in Oregon and Washington

since the stands harvested were greater than 70 and less

than 150 years old. The spatial arrangement of Cooper’s
hawk habitat over time is illustrated in Figure 5.

Maximizing the Spatial Goal Requiring

Minimum Patch Sizes
When we used the heuristic technique to maximize the
spatial goal requiring minimum patch sizes, we found that
the average percentage of land in sharp-shinned hawk and
Cooper’s hawk habitat decreased about 10%, and dropped
dramatically in period 10. Fewer stands were harvested
in period 1 of this scenario than in the previous scenario,
thus as the forest gets older (>70 yrs) these habitat goals
decline. Their level of goal achievement is illustrated in
Figure 6. The percentage of land in varied thrush habitat
(spatial goal requiring minimum patch sizes) gradually
increased over time, and was about 28% in period 10 of
this scenario. Therefore about 23% more land is in this
habitat type during period 10 than in the previous
scenario, because we attempted to maximize the amount
of varied thrush habitat here. The percentage of land in
great gray owl habitat (spatial goal requiring adjacent
habitat types) increased dramatically in period 10, as more
stands were clearcut, yet averaged about 10% per period.
Timber volume per 5-year planning period averaged 7.9
million board feet (approximately 49,500 m3) (Figure 4),
and ranged from 3 million (approximately 18,900 m3) to
45.9 million board feet (approximately 288,600 m3) per
period. Harvest volume is quite high in the last period
because some cutting here did not constrain the
achievement of varied thrush habitat, and because we did
not control harvest levels with a maximum harvest level
constraint. The spatial arrangement of varied thrush
habitat is illustrated in Figure 7.

Maximizing the Spatial Goal Requiring

Adjacent Patch Types
When we used the heuristic technique to maximize the
spatial goal requiring adjacent patch types, we found that
the percentage of land in sharp-shinned hawk and
Cooper ’s hawk habitat were lower than when the
nonspatial goals were maximized (23 and 32%,
respectively), with both generally declining over time
(Figure 8). Northern goshawk, and red tree vole habitat
types (the nonspatial goals) were approximately 50-70%
lower here than when these nonspatial goals were
maximized. The percentage of land in varied thrush
habitat (spatial goal requiring minimum patch sizes)
gradually increased over time, yet was about 1% lower
here than when we attempted to maximize this spatial
goal. The percentage of land in great gray owl habitat
averaged 27% over the planning horizon, with dramatic
increases in the last 3 planning periods. One reason for

Figure 7 (left). Varied thrush habitat (shaded area) in the

initial landscape condition (top), after period 5 (middle), and
after period 10 (bottom), when the spatial goal requiring
minimum patch sizes is maximized.
 CHAPTER 23: INTEGRATING GOALS 577

70

Sharp-shinned hawk
60 Cooper's hawk
Northern goshawk
Red tree vole
50 Varied thrush
Great gray owl

40

30

20

10

0
1 2 3 4 5 6 7 8 9 10
PLANNING PERIOD

Figure 8. Results from maximizing the spatial goal requiring

adjacent patch types for great gray owl habitat.

these patterns is that the planning model tries to develop

both old and young forests simultaneously, so it allows
some of the older stands to continue to grow, while the
younger ones are harvested between the ages of 40 and
50, in order to create as much early seral forest as possible.
Therefore, the amount of land in Cooper’s hawk habitat
types (stands between the ages of 30 and 70 yr) gradually
decreases over time. The spatial arrangement of great gray
owl habitat is illustrated in Figure 9. Timber volume per
5-year planning period averaged 9.5 million board feet
(approximately 60,000 m3) (Figure 4), and ranged from 3
million (approximately 18,900 m3) to 24.6 million board
feet (approximately 154,500 m3) per period. Harvest
volume is high in the last few periods because some
cutting is required to create early seral patches to
complement the older forest patches, and because we did
not control harvest levels with a maximum harvest level
constraint. The landscape that results at the end of the
planning horizon is much different than those that result
from maximizing the other goals (Figure 10).

Discussion
Formulating management plans with a variety of goals
(e.g., economic and wildlife species habitat) can be
challenging. We have illustrated a process in which
wildlife species habitat goals can be identified, and criteria
for their measurement developed. This type of process can
be used by planning teams to facilitate the incorporation
of these goals into planning. The complexity of the goal
criteria should not be a barrier to this process, as we have
shown that there are techniques to incorporate nonlinear

Figure 9 (right). Great gray owl habitat in the initial

landscape condition (a), after period 5 (b), and after period 10
(c), when the spatial goal requiring adjacent patch types is
maximized.
 578 Wildlife–Habitat Relationships in Oregon and Washington

0 - 10 years old 0 - 10 years old

11 - 30 years old 11 - 30 years old
31 - 89 years old 31 - 89 years old
90+ years old 90+ years old
Riparian area Riparian area

Fig. 10. Landscape condition in planning period 10 when

maximizing the nonspatial goals (a, top left), maximizing the
spatial goal requiring minimum patch sizes (b, lower left),
maximizing the spatial goal requiring adjacent patch types (c,
above).

integer type variables in a planning process. Spatial goals

require information about the locations of resources, and
can be represented by linear or nonlinear relationships.
For example, a nonlinear relationship could require one
type of habitat patch to be adjacent to another type of
habitat. A linear relationship could be a variable such as
distance from the ocean, which essentially can be assumed
to be a constant.
Whereas the development of goals and goal criteria are
important for planning processes, the specialists involved
in developing goal criteria, and the planners whose task
it is to implement the goals in a planning process,
encounter potential challenges in four focus areas; those
dealing with people, databases, technology, or
organizational commitment. People are the instrument by
0 - 10 years old which databases and technology are joined to help develop
management alternatives. The types of tools planners
11 - 30 years old possess will generally guide a process toward the use of a
31 - 89 years old particular set of tools and data structures. Databases are
constantly evolving, as are the computer and software
90+ years old technology we use to manipulate data. Organizations
should match the planning objectives (e.g., the types of
Riparian area
goals to be measured, how they will be measured, the
 CHAPTER 23: INTEGRATING GOALS 579

spatial extent of the planning area, etc.) with the databases Literature Cited
and the technology at their disposal (or which could be 1. Arthaud, G.J., and D. Rose. 1996. A methodology for estimating
acquired). In doing this, estimates of budget levels and production possibility frontiers for wildlife habitat and timber value at
cash flow needs, and other institutional limitations (e.g., the landscape level. Canadian Journal of Forest Research 26:2191-2200.
personnel), will become apparent. The reaction of the 2. Bettinger, P. and K. Boston. 1999. Intensifying a heuristic forest harvest
scheduling search procedure with 2-opt decision choices. Canadian
leaders of the organization to the planning needs will be Journal of Forest Research 29:1784-1792.
one indication of the commitment of the organization to 3. Bettinger, P., K.N. Johnson, and J. Sessions. 1996. Forest planning in an
the planning process. If •1 of these four essential focus Oregon case study: defining the problem and attempting to meet goals
areas (people, databases, technology, organizational with a spatial-analysis technique. Environmental Management 20:565-
commitment) is limiting, an organization may consider 577.
re-examining the expected results of the planning process. 4. Bettinger, P., J. Sessions, and K. Boston. 1997. Using tabu search to
schedule timber harvests subject to spatial wildlife goals for big game.
Ecological Modeling 94:111-123.
Conclusions
5. Bettinger, P., J. Sessions, and K.N. Johnson. 1998. Ensuring the
The central role of forest managers is good land compatibility of aquatic habitat and commodity production goals in
stewardship, which is achieved through decision making, eastern Oregon with a tabu search procedure. Forest Science 44:96-
or choosing among alternative courses of action. 112.
Quantitative planning techniques allow managers to 6. Daust, D.K., and J.D. Nelson. 1993. Spatial reduction factors for strata-
examine variations in goals, goal criteria, and activities, based harvest schedules. Forest Science 39:152-165.
and to develop alternative management plans. We are 7. Glover, F. 1989. Tabu search—Part I. ORSA Journal of Computing
1:190-206.
fortunate that advances in applied economics and 8. Glover, F. 1990. Tabu search—Part II. ORSA Journal of Computing 2:4-
management science have allowed the natural resources 32.
fields to use quantitative planning tools, and to say with 9. Haight, R.G., and L.E. Travis. 1997. Wildlife conservation planning using
some confidence that recommended land management stochastic optimization and importance sampling. Forest Science
plans are efficient and best meet the objectives of forest 43:129-139.
managers. Managers are then relied on to balance the 10. Hof, J., M. Bevers, L. Joyce, and B. Kent. 1994. An integer programming
approach for spatially and temporally optimizing wildlife populations.
divergent goals of an organization and appraise the
Forest Science 40:177-191.
reliability and meaning of information developed in plans 11. Hoganson, H.M., and D. Rose. 1984. A simulation approach for optimal
that are developed and designed to meet those goals. timber management scheduling. Forest Science 30:220-238.
Four major aspects of planning projects are people, 12. Kangas, J., and T. Pukkala. 1996. Operationalization of biological
databases, technology, and commitment by the diversity as a decision objective in tactical forest planning. Canadian
organization to the development of the plan. Assuming Journal of Forest Research 26:103-111.
databases, technology, and organizational commitment are 13. Lockwood, C., and T. Moore. 1993. Harvest scheduling with spatial
constraints: a simulated annealing approach. Canadian Journal of Forest
available at appropriate levels to facilitate a planning
Research 23:468-478.
process, the limiting factor (if there is one) is people. The 14. Murray, A.T. 1999. Spatial restrictions in harvest scheduling. Forest
communication process among individuals can be Science 45:45-52.
enhanced with a common understanding of the pieces of 15. Murray, A.T., and R.L. Church. 1995. Heuristic solution approaches to
a planning process, including the basic steps planners and operational forest planning problems. OR Spektrum [Operations
biologists must take to understand the goals: Research] 17:193-203.
1. develop quantitative description of species goal; 16. Nelson, J., and J.D. Brodie. 1990. Comparison of a random search
algorithm and mixed integer programming for solving area-based forest
2. establish criteria to measure the goals; plans. Canadian Journal of Forest Research 20:934-942.
3. select the activities that can be used to help achieve 17. O’Hara, A.J., B.A. Faaland, and B.B. Bare. 1989. Spatially constrained
the goals; timber harvest scheduling. Canadian Journal of Forest Research 19:715-
4. work with planners to develop mathematical 724.
statements of the goals, in the form of an objective 18. Pulkki, R. 1984. A spatial database—heuristic programming system for
function and constraints, to help guide the selection aiding decision-making in long-distance transport of wood. Acta
Forestalia Fennica 188.
of a combination of activities that will comprise a
19. Reeves, C.R. and J.E. Beasley. 1993. Introduction. Pages 1-19 in C.R.
management plan. Reeves, editor. Modern heuristic techniques for combinatorial
problems. John Wiley & Sons, Inc., New York, NY.
20. Weintraub, A., G. Jones, A. Magendzo, M. Meacham, and M. Kirby. 1994.
A heuristic system to solve mixed integer forest planning models.
Operations Research 42:1010-1024.
21. Weintraub, A., G. Jones, M. Meacham, A. Magendzo, A. Magendzo, and D.
Malchuk. 1995. Heuristic procedures for solving mixed-integer harvest
scheduling—transportation planning models. Canadian Journal of
Forest Research 25:1618-1626.
 24
Decaying Wood in Pacific Northwest Forests:
Concepts and Tools for Habitat Management
Cathy L. Rose, Bruce G. Marcot,T. Kim Mellen, Janet L. Ohmann,
Karen L.Waddell, Deborah L. Lindley, & Barry Schreiber

Introduction
Decaying wood has become a major conservation issue in Historical interest in decaying wood has centered
managed forest ecosystems.16, 64, 69a, 149, 201 Of particular mainly on impediments to reforestation and on economic
interest to wildlife scientists, foresters, and managers are consequences of decay to timber production. In the first
the roles of wood decay in the diversity and distribution half of this century, wood decay or “decadence” was
of native fauna, and ecosystem processes. Numerous viewed simply as an undesirable attribute of overmature
wildlife functions are attributed to decaying wood as a forests at increased risk of damage by fire, insects, or
source of food, nutrients, and cover for organisms at disease. Recognition of the diverse ecological benefits
numerous trophic levels.231, 232, 234, 346, 369 Principles of long- derived from decaying wood in aquatic and terrestrial
term productivity and sustainable forestry include ecosystems has grown appreciably over the past two
decaying wood as a key feature of productive and resilient decades.56, 198, 201, 234 Wood decay in forests of the Pacific
ecosystems. 10, 229, 291, 293, 386 In addition to a growing Northwest has recently become a topic of renewed interest
appreciation of the aesthetic, spiritual, and recreational at national and global scales, regarding the role of
values of forests, society increasingly recognizes ecosystem terrestrial carbon storage in the reduction of atmospheric
services of forests as resource “capital” with tangible CO2 (a greenhouse gas). In light of recent projections for
economic value to humansy, such as air and water quality, accelerated climatic warming during the next several
flood control, and climate modification.15, 262, 290 decades,255 regulatory targets for terrestrial carbon storage
The ecological importance of decaying wood is are being considered.16
especially evident in coniferous forests of the Pacific New research over the past three decades has
Northwest. In this region, the abundance of large decaying emphasized the significance of decaying wood to many
wood is a defining feature of forest ecosystems, and a key fish and wildlife species,48, 56, 230, 233, 234, 369 and to overall
factor in ecosystem diversity and productivity.127 Native ecosystem function.389 The importance of decaying wood
forests west of the Cascade Crest are highly productive to ecosystem biodiversity, productivity, and sustainability
and accumulate large amounts of live and dead wood—a is a keynote topic in two recent regional ecosystem
result of the temperate climate that favors tree growth.125, assessments in Oregon and Washington.114, 225 These, and
126, 395
Large accumulations of decaying wood provide other publications address both the specific roles of wood
wildlife habitat and influence basic ecosystem processes decay in ecosystem processes and functions, as well as
such as soil development and productivity, nutrient ecological functions of wildlife species associated with
immobilization and mineralization, and nitrogen wood decay.13, 68, 216, 250
fixation.85, 115, 218, 233 Forests east of the Cascade Crest are also Interactions among wildlife, other organisms, and
strongly influenced by accumulations of decaying wood decaying wood substrates are essential to ecosystem
that set the stage for ecosystem disturbances from fire, processes and functions. In the process of meeting their
insects, and disease.56, 137, 390 needs, animals accomplish ecosystem “work” with respect
Decaying wood has a pivotal role in both estuarine and to transformation of energy and cycling of nutrients in
coastal marine ecosystems, supporting complex trophic wood. For example, chipmunks and squirrels disperse
webs from benthos to higher vertebrates. More than half mycorrhizal fungi which play key roles in nutrient cycling
of the total organic carbon content of Washington’s off- for tree growth; birds, bats, and shrews consume insects
shore, midshelf sediment originates from coniferous that decompose wood or feed on invertebrates and
forests.139 Inputs of decaying wood are crucial to most microbes; beavers and woodpeckers create habitats by
aspects of stream processes, such as channel morphology, modifying physical structures; arthropods build and
hydrology, and nutrient cycling.233, 260, 262, 354 Although aerate soil by decomposing wood material. Relations
knowledge of these processes is incomplete, decaying between wood decay and wildlife have been examined in
wood in freshwater and marine food webs illustrates the several recent analyses.56, 225, 282, 283, 284, 286 Species-specific
potentially far-reaching implications of terrestrial wood associations have been identified for some tree species,
management. types of decay fungi, and different forms of wood, such

580
 CHAPTER 24: WOOD LEGACIES 581

as bark piles at the base of snags, hollow living trees, and Since the publication of Thomas et al.369 and Brown,48
broomed trees. Simplified classification schemes and new research has indicated that more snags and large
inventory procedures have been developed for decaying down wood are needed to provide for the needs of fish,
wood, particularly wood habitat structures relevant to wildlife, and other ecosystem functions than was
wildlife.56 previously recommended by forest management
Wildlife species associated with wood decay, and their guidelines in Washington and Oregon. For example, the
ecosystems are affected by management activities. density of cavity trees selected and used by cavity-nesters
Intensively managed forest plantations have replaced old- is higher than provided for in current management
growth throughout most of the commercial forest land guidelines.53, 102 Reductions in the wood content of
base in Oregon and Washington.41, 42 Intensive forest estuarine and marine environments, and consequences to
management regimes have substantially altered the assorted aquatic organisms have also been documented.139
abundance and composition (species, size, decay class) of Reductions in forest productivity linked to management
decaying wood in forest ecosystems in the Pacific practices such as stem removal, slash burning, and soil
Northwest. Managed forests, on average, have lower disturbance have also been recognized.10, 13, 44, 198, 199
amounts of large down wood and snags than do natural Critical ecosystem functions of wood, coupled with
forests.59, 81, 82, 114, 154, 225, 276, 344, 368 Furthermore, in forests east incomplete knowledge for management, make the topic
of the Cascade crest, fire suppression has altered stand of decaying wood a priority for future research and
dynamics and produced accumulations of fine fuels adaptive management.78, 149, 164 Effective approaches to
conducive to stand-replacement fires.5, 211, 302 Forest health managing decaying wood require that dead wood
problems and declining populations of some vertebrate components of wildlife habitats be viewed within the
and invertebrate wildlife species have coincided with context of the larger interacting ecosystem. To help
changes in forest structure. These changes have raised managers achieve the goal of effective management of
concerns about the future biodiversity, productivity, and decaying wood, this chapter seeks to provide a focus on
sustainability of the region’s forests, particularly in coastal the ecological context for wood decay and associated
and eastside forests.21, 170, 187, 222, 367 wildlife in forests of the Pacific Northwest. Emphasis on

Wood Legacies in Managed Forests

John Hayes
characteristics may never be produced in forests
Legacies are structures or components of ecosystems managed intensively on short- to moderate-length
that exist prior to a disturbance and are “inherited” rotations. Habitat quality for species that depend
by the post-disturbance community. Legacies can upon or are closely associated with these structures
provide important temporal connectivity within a can be seriously diminished with their loss from
stand, allowing organisms present in a pre- forest stands. The ecological importance of wood
disturbance community to persist in an area legacies combined with the difficulties of creating
following disturbance. In addition, legacy wood can replacement structures provide convincing reasons
provide structural elements and complexity in a to conserve legacy structures during management
stand that would otherwise require very long activities.
periods of time to develop. In managed forests, Managing wood legacies through time in
wood legacies, including large diameter trees, snags, managed forests is a multi-staged process. Existing
and down wood, are ecologically important structures that will serve as legacy structures in the
structures that play central roles in diverse post-disturbance environment should be identified
ecosystem processes and functions, such as prior to a disturbance event, such as logging. In
geomorphic processes, hydrology, nutrient cycling, some cases, it may be adequate to rely on the timber
and habitat for fish and wildlife. The ecological value sale administrator or loggers to identify appropriate
of wood legacies has begun to gain widespread structures and implement the management strategy
recognition only within the past two decades.122, 164 in the field. Since one intent of legacy structures is
As a result of a variety of operational, safety, and to provide various functions through time, it will
economic considerations, application of intensive often be valuable to either individually mark
forest management practices often results in removal important legacy structures, or to document their
of legacy structures from stands and minimal location and purpose so that future managers can
retention of future legacy structures. Growing take the structures into account. Of equal
replacement structures with similar characteristics importance, plans for recruitment of future legacy
(e.g., large diameter trees with large diameter structures should be prepared to ensure that legacy
branches, thick and deeply-furrowed bark, and structures will be available in future stands.
complex crown structure) requires decades or longer. Innovative silvicultural practices can be employed
Moreover, unless special provisions are made, large to create conditions favorable to development of
diameter trees, snags, and logs with these future legacy structures.
 582 Wildlife–Habitat Relationships in Oregon and Washington

concepts of long-term productivity in this chapter reflects

an underlying principle that habitat functions of decaying Ecosystem Definition
wood are inextricably linked to ecosystem processes. Ecosystems can be described at a variety of spatial
Careful attention to the whole ecosystem is a prerequisite scales and the spatial hierarchy of ecosystems is
to successful management of decaying wood for wildlife. helpful for understanding the ecological functions
This chapter provides a synthesis of knowledge on of wood decay. Therefore, we use the “macrosystem”
processes and functions of wood decay in forest definition of ecosystems,382 which recognizes the
productivity and wildlife habitat, and summarizes functional continuum of terrestrial, stream, estuarine
available information on the current regional status of and coastal ocean environments. Functional linkages
decaying wood. It then ofers managers a stepwise include both the physical and biological processes
assessment process to set goals and objectives, and select and the interactions between organisms and
silvicultural tools to manage wood decay for desired decaying wood that contribute to nutrient, energy,
results. Although the primary emphasis is on upland and material cycling this continuum.233 Smaller sub-
forests, the chapter also highlights wood decay structures units of ecological organization273 are recognized
and functions in forested wetlands, riparian forests, and within this macrosystem, such as the forest floor and
aquatic systems. The analysis addresses a wide range of riparian communities.
woody plant structures, with emphasis on large down
wood and snags that form ‘legacy’ structures linking
successive generations of stands. forests, biological activity is the primary determinant of
wood decay rates.164, 346, 404 In drier forest regions, fire is
often the primary agent of wood breakdown. In forest
ecosystems with infrequent fire, the principal mechanisms
Ecological Significance of of wood breakdown include leaching, fragmentation,
Decaying Wood transport, collapse, settling, seasoning, respiration, and
biological transformation.164 Decay processes in wood can
Wood Decay Processes be divided into two groups depending on whether the
Decay processes in wood form the basis for understanding mode of action is mechanical fragmentation, or chemical
ecological roles and relationships among wildlife and breakdown. In the following section, we describe
decaying woodand. Familiarity with these relationships biologically mediated chemical breakdown of wood, and
facilitates the translation of these concepts into successful that various physical and biological processes fragment
management strategies. Down wood, snags, and other wood structure.
persistent wood structures (such as stumps, root wads, Knowledge of wood structure and composition is
and coarse roots) occur in most forest ecosystems. In moist useful for understanding processes of wood decay. Wood
tissues of tree stems include the outer bark, cork cambium,
inner bark (phloem), vascular cambium, outer xylem
Terminology of Decaying Wood
(living sapwood), and the inner xylem (non-living
Decaying wood includes all portions of a tree, heartwood). Outer bark is a non-living barrier between
standing or down, that are dead and in the process the inner tree and harmful factors in the environment, such
of decay. This is a broad definition that seeks to as fire, insects, and diseases. The cork cambium
address wildlife uses56 and other ecological functions (phellogen) produces bark cells. The vascular cambium
of decaying wood. It includes the following wood produces both the phloem cells (principal food-conducting
structures: tissue) and xylem cells of the sapwood (the main water
• living trees with decay (e.g., heart-rot, root-rot, storage and conducting tissue) and heartwood. Wood
broken or decaying tops, large decaying properties are determined mainly by plant cell walls,
branches, broken branch stubs, bole wounds, consisting ofcellulose fibrils impregnated within a cement-
loose bark, stem cavities, and old trees with like matrix of lignin and other organic polymers in the
deep, fissured bark and thick, bulky branches) approximate proportions: cellulose (40-50%),
• living hollow trees (advanced heartwood decay) hemicellulose (20-35%), and lignin (15-35%).
• broomed trees (diseased branches)
• snags (dead trees or portions of trees that remain Wood Fragmentation
upright) Mechanical disruption of wood structure may be initiated
• down wood (decaying trees or portions of trees abiotically by natural events, such as heavy snows, rime
that have fallen to the forest floor; includes ice, avalanches, mass soil movement, floods, and
other commonly used terms, such as logs, windstorms that uproot or break trees.86, 230, 270, 301, 316 Thermal
coarse woody debris, and large woody debris) expansion and contraction of wood, and water in woody
• stumps and other tree parts (rootwads, bark tissues also promotes the physical disruption of wood
piles, coarse roots, and fine litter) structure. These disturbance agents affect wood decay
variously by: 1) directly or indirectly inducing mortality
of wood tissues, 2) altering the microclimate surrounding
the affected wood structure, 3) increasing the surface area
 CHAPTER 24: WOOD LEGACIES 583

of wood exposed to the environment, and 4) increasing

wood accessibility to microbes.
Fragmentation of decaying wood during the early Figure 2. Conk of
stages of decomposition is accomplished by wood-boring the fungus
insects such as beetles, carpenter ants, and termites.23, 89, 230 Fomes pini.
As fissures develop in wood, invertebrates from minute Photo:
mites to centipedes, millipedes, slugs and snails find Washington
suitable habitat, and contribute to further fragmentation. Department of
In the soil, an abundance of soil animals (including Natural
nematodes, earthworms, and microarthropods) contribute Resources.
to the fragmentation of decaying wood and increase the
surface area for microbial attack.186 Functions and
interactions between these soil fauna are varied and
complex.230
A variety of vertebrate species contribute to the wood and by the enzymatic action of microbes. Insects
fragmentation of decaying wood in both live and dead also serve as vectors for other decay organisms and
trees. Most notable are the primary cavity excavators, such increase wood surface area accessible to both microbial
as woodpeckers and nuthatches that break wood fibers and fungal attack.23, 47, 89, 118, 230 Fungi accomplish wood
in the process of cavity excavation216 (Figure 1). Other decay by enzymatically decomposing the cell wall. Wood
animal activities that fragment wood include limb pruning decay fungi can be separated into two main groups—the
or breakage, foraging, denning, and burrowing. As pathogenic heart-rot and root-rot fungi that colonize
decaying wood softens, tunneling and burrowing by other primarily the heartwood or roots of living trees (primary
vertebrates such as salamanders, shrew, shrew-moles, and decay), and the saprobic fungi that colonize decaying
voles continue the fragmentation process in rotten wood wood (secondary decay) (Figure 2).
and bark, and promote favorable conditions for further On live trees, vulnerable locations for primary decay
decomposition by invertebrates, fungi, and bacteria. include heartwood, branch nodes, roots, dead and broken
tops or branches, and wounds from fire, logging activities,
Principal Biological Agents of Wood Decay
windthrow, and lightning. Wounding is instrumental to
Biologically mediated chemical breakdown of wood can
primary decay because germinating fungal spores cannot
be grouped into two basic categories: primary decay that
penetrate intact bark or living sapwood, but must land
establishes on living trees, and secondary decay that
directly on exposed dead wood connected to the
commences after tree mortality caused by other factors.
heartwood. Generally, decay fungi located in the upper
Of the biological agents of wood decay, insects and fungi
stems of living trees are heart-rot fungi.220 Heart-rot fungi
are the principal players in coniferous forest ecosystems.
are not restricted to the heartwood, but usually infect
Wood-boring insects contribute to the early stages of wood
through wounds in the sapwood of living trees. Every tree
decomposition by consuming and chemically digesting
species is susceptible to at least one heart-rot fungus, but
few fungi can cause heart-rot in living trees.56, 175, 220
Root rot is caused by a group of primary decay fungi
that infect roots and spreads to other trees through root
contact. These root pathogens act by causing nutrient
Figure 1. The deficiency after disrupting root tissue, altering root
pileated morphology, parasitizing the root cambium, or
woodpecker, a functioning as heart-rotters of the roots and lower tree
primary bole.220, 322 By weakening the root structure, root rot may
excavator. Photo:
directly kill trees, or indirectly contribute to mortality from
David H.
windthrow and insects.
Johnson.
Processes of secondary decay are similar in both dead
trees and in dead portions of live trees. Heart-rot fungi
already present in a tree at the time of death usually
contribute little to further decay of the dead tree. Instead
they are replaced by other fungi termed saprobes, that are
better adapted to degrade dead wood. Sapwood rotting
fungi are quick to colonize a dead tree, particularly if tree
mortality occurred without depletion of nutrient and
carbohydrate stores. In general, once the sapwood is fully
decayed, the sapwood fungi are replaced by saprobes that
continue to degrade the sapwood and proceed into the
heartwood. Saprobes may also invade larger patches of
dead tissue on live trees, however.
 584 Wildlife–Habitat Relationships in Oregon and Washington

Libe plant roots contribute to wood decay by producing functional and structural properties of wood, and
enzymes and other exudates that accelerate cellulose eliminate the need to determine the date of tree mortality.
decomposition and catalyze nutrient release from organic The classes are based on a variety of tree characteristics,
matter, and by forming symbioses with mycorrhizal fungi, including the presence or absence of bark and fine twigs,
N-fixing bacteria and actinomycete fungi to aid lignin bole shape, wood texture, and extent of log contact with
decomposition.1 soil. The decay status of a tree when it falls to the forest
floor depends on its condition as a live tree or snag. For
Stages and Structures of Decaying Wood example, a live or recently dead tree that falls to the forest
Various woody tissues in a tree have inherent differences floor becomes a Class 1 log, however, a fallen snag may
in decay susceptibility. Decay normally begins at the create a Class 1 to a Class 4 log.
exterior of a tree and progresses inward, though heart rot A classification system for Douglas-fir snags describes
may degrade the interior of a tree with intact sapwood. a continuum of nine decay stages.234, 370 The nine stages of
The outer bark disappears mainly by fragmentation and snag decay correspond to the five decay classes for logs
sloughing from the top and sides of down trees. Despite a described above, with four additional stages: two earlier
high concentration of nitrogen, decay of outer bark is stages for live, but declining trees, and two later stages
inhibited by a high content of lignin and other anti- for progressive breakage and decay of the lower bole.230
microbial compounds.176, 203, 230 Inner bark rapidly decays This nine-stage system was modified and refined82, 263 into
due to its small volume and high nutritional quality (high a five-stage system by eliminating Stages 1 and 2, and
digestibility and nutrient content). The outer ring of living combining stages 7-9.
sapwood is usually highly resistant to decay despite its A simplified system of three structural classes each for
high nutrient content, and may remain so even when conifer logs and snags has been developed based on the
adjacent tissues are extensively decayed. Dead sapwood premise that fewer classes are sufficient to describe wildlife
decays rapidly, however. Compared to sapwood, use and easier to apply in the field (Figure 3). Visual,
heartwood decays slowly because of low nutrient quality chemical, and physical properties of conifer wood were
and digestibility to microbes (high lignin and low evaluated in relation to degree of decay. Wood density,
nitrogen).342 cellulose, and lignin content were found to be the best
All dead wood progresses through discreet stages of indicators of decay, whereas visual characteristics were
decay characterized by progressive changes in wood relatively poor indicators.182 Consequently, although the
attributes, decomposition rates, and decomposer species. three class system of visually classifying wood decay may
This chronology has been described in more detail for be satisfactory for assessing wood conditions important
down wood than for snags, however. At each stage, decay to many vertebrate wildlife species, the five-class system
is influenced by conditions in the physical environment, may better describe more subtle changes in wood
and by the structural and chemical composition of wood. properties (density, moisture) important to other
As decomposition advances, changes in wood properties organisms, and to ecosystem functions.
include decreased density and digestibility (higher ratio
of lignin to cellulose), and increased water content and Ecological Functions of Decaying Wood
nutrient concentrations (including nitrogen, phosphorus,
calcium, and magnesium in microbial tissue). Microbial Wildlife Habitat Roles
decomposers vary in their nutrient requirements and in Down wood, snags, and live trees with decay serve vital
their inherent ability to degrade the structural matrix of roles in meeting the life history needs of wildlife species
wood (cellulose and lignin). Altered nutrient and in Oregon and Washington. Literature describing habitat
structural attributes of wood with advancing decay thus relationships for wildlife and decaying wood in the Pacific
creates sequential niches for different decomposer Northwest are most comprehensive for vertebrate
species.220 Decay rates for dead wood are thus logarithmic foraging ecology and cavity-nesting relations. Knowledge
rather than linear, because the more readily-decomposed is more extensive for snags than for down wood, and for
fraction of wood is digested first, leaving the less digestible westside than for eastside ecosystems. Major regional
fraction,184 and because nitrogen availability to microbes ecosystem assessments recently completed in Oregon and
decreases over time. Washington have examined the roles of decaying wood
in habitat and in ecosystem biodiversity, productivity and
Classification Systems For Decaying Wood sustainability.114, 225, 302
To facilitate the description and quantification of habitat Recent significant advancements have defined wildlife
attributes provided by decaying wood, several systems species-specific relationships with particular
to classify decay conditions in conifers have been characteristics and components of decaying trees, both
developed. New studies are needed to adapt such systems standing and fallen,56, 95, 185, 284, 351, 373, 386, 402 and implications
to hardwoods, which typically do not follow the same for management.13, 68, 223, 226, 250, 327 Much of this recent
decay phases as conifers. A five-class system has been research has improved our understanding of poorly
commonly used to describe the extent of decay in down documented wildlife uses of microhabitats. Some of these
wood.28, 234, 339, 376 Decay classes are useful to ecological are represented in the Habitat Element matrixes on the
studies of wood decomposition because they describe CD-ROM accompanying this book, and in the DecAID
 CHAPTER 24: WOOD LEGACIES 585

Figure 3. Structural classes of snags and down wood, from

Bull et al.56

model.226 For example, research on amphibians previously are more readily decomposed by organisms creating
known to be generally associated with down wood, suitable conditions for cavity excavators.284, 323 Tree species
elucidated ways in which these species use distinctly may also be correlated with environmental conditions that
different features of down wood. 22, 59a For example, influence habitat suitability, such as microclimate, canopy
ensatinas occur primarily in bark piles at the base of snags, position, geographical location, etc.
redback salamanders are found most often under
Decaying Wood Habitat Structures. In a recent review,
moderately decayed logs, clouded salamanders prefer
decaying wood structures were grouped into five
habitat under bark on logs, and Oregon slender
categories representing key habitat elements for wildlife,
salamanders inhabit the interior of logs.22, 59a
including living trees with decay, hollow trees, trees with
Up-to-date and specific information on species’ habitat
brooms, dead trees (snags), and down woody material
associations and key ecological functions is available in
(logs).56 An overview of these habitat structures is
the matrixes on the CD-ROM accompanying this book and
provided below, with reference to the three-class systems
in the DecAID model.226 The remainder of this section
for classifying wood decay in snags and logs.56
presents a sampling of new findings about resource
partitioning among various wildlife communities in • Standing Trees. Living trees often have pockets of decay,
Pacific Northwest forests, with examples of current dieback, wounds, or broken tops and limbs that serve as
information available in the matrices and in the DecAID wildlife habitat. To be useful to most cavity excavators,
model. live trees usually must contain wood in a Class 2 stage of
decomposition.56, 238 For example, strong excavators, such
Tree Species. Few studies have demonstrated the causal
as Williamson’s sapsuckers, pileated woodpeckers, and
relationship between tree species per se, and the habitat
black-backed woodpeckers, select trees with a sound
functions of decaying wood. However, some habitat
exterior sapwood shell and decaying heartwood to
functions of wood are provided primarily by certain tree
excavate their nest cavities.238, 284, 307 Armillaria root rot and
species. Woodpeckers, sapsuckers, and nuthatches are
other saprophytic fungi produce softer exterior wood
highly specific in their selection of tree species for nesting
suitable for cavity construction byweaker excavators such
and roosting, and this selectivity is attributed to the
as red-breasted nuthatch and chickadee.216 Trees with
presence of decay fungi. 29, 56, 95, 351 Tree species also
decaying tops or large dead or deformed branches provide
influences other life requisites, such as prey production.
roosting and drumming habitat for birds, and nesting
For example, carpenter ants, a primary prey species of
platforms for murrelets, owls, and raptors (Figure 4).102,
pileated woodpeckers in northeastern Oregon, prefer 114, 153, 269
In addition, localized wood decay resulting from
down larch logs (Larix laricina).373 Decay characteristics,
tree wounding and scarring (particularly at the tree base)
size, and wood density may account for more variation in
provides substrate for ants and other insects consumed
wildlife use of wood than tree species. These parameters
by foraging birds.53
are likely to be highly correlated; that is, some tree species
 586 Wildlife–Habitat Relationships in Oregon and Washington

resistant sapwood remains structurally sound for

centuries. In the Interior Columbia Basin, grand fir and
western larch form the best hollow trees for wildlife uses.
Broomed trees caused by mistletoe, rust, or needlecast
fungi may remain alive for decades, and have attributes
distinct from decay patches in live trees. Abundant forage
is produced from mistletoe shoots and fruits. Regardless
of the extent of decay, broom infections provide various
habitat functions to wildlife depending on how and where
they form along the bole.371, 372 For example, mistletoe
brooms form platforms used for nesting, roosting, and
resting sites by owls, hawks, and song birds; roosting by
grouse; and resting cover by squirrels, porcupines, and
marten.56, 114 Little information is published on the use of
rust or needlecast brooms by wildlife, but it is likely they
Figure 4. Spotted owl roost. Photo: Deborah Lindley. serve similar functions as dwarf mistletoe. Documented
uses include rest sites for American marten and nest sites
for squirrels and great horned owls.50, 56, 256 Because the
dieback of infected tree parts may occur over a protracted
time frame, a single tree infected with these types of
pathogens may contain discrete patches of dead wood in
various stages of decay.51, 256
Recent studies have provided valuable insight on
wildlife uses of snags (dead trees).21, 56, 314, 402 Snags provide
essential habitat features for many wildlife species (Figure
6). The abundance of cavity-using species is directly
related to the presence or absence of suitable cavity trees.
Habitat suitability for cavity-users is influenced by the size
(diameter and height), abundance, density, distribution,
species, and decay characteristics of snags.307 In addition,
the structural condition of surrounding vegetation
determines foraging opportunities.402
Figure 5. Hollow snag of ponderosa pine. Photo: Deborah The Habitat Elements matrix on the CD-ROM with this
Lindley. book lists a total of 96 wildlife species associated with
snags in forest (93 species) or grassland /shrubland (47
species) environments. Most of these species use snags in
A hollow tree (live or dead) is formed when advanced
both environments. In forests, this includes 4 amphibian,
heartwood decay (Class 3) causes the heartwood cylinder
63 bird, and 26 mammal species. Additionally, 51 wildlife
to detach from the sapwood, leaving a hollow core
species are associated with tree cavities, 45 with dead parts
surrounded by a reinforcing band of sapwood. The
of live trees, 33 with remnant or legacy trees (which may
softened heartwood of trees colonized by heart-rot fungi
have dead parts), 28 with hollow living trees, 21 with bark
provides suitable conditions for excavating a nest chamber,
crevices, and 18 with trees having mistletoe or witch’s
and the living sapwood functions to maintain the tree’s
brooms. Habitat uses include nesting, roosting, preening,
structural integrity (Figure 5).56 The structural elements
foraging, perching, courtship, drumming, and hibernating
produced by advancing heartwood decay include hollow
(Figure 7).
cavities from small patches to larger cylinders, to entire
Of the 93 wildlife species associated with snags in forest
hollow trunks. These decay legacies and resulting habitat
environments, 21 are associated with hard snags (Stages
elements are described in detail by Bull et al.56
1 and 2), 20 with moderately decayed snags (Stage 3), and
Hollow trees larger than 20 inches (51 cm) in diameter
6 with soft snags (Stages 4-5) in the five-stage classification
at breast height (dbh) are the most valuable for denning,
system. According to the matrixes,188 most snag-using
shelter, roosting, and hunting by a wide range of animals.7,
52, 55, 56, 265, 282 wildlife species are associated with snags >14.2 inches (36
Hollow chambers are used as dens by black
cm) diameter at breast height (dbh), and about a third of
bears, as night roosts by woodpeckers, and as dens, shelter,
these species use snags >29.1 inches (74 cm) dbh.
roosts, and hunting sites by a variety of animals, including
This query of the Habitat Elements matrix illustrates
flying squirrels, wood rats, bats, American marten,
the breadth of updated information about wildlife and
northern flickers and Vaux’s swift.7, 51, 52, 55, 56, 265, 282 Hollow
snag habitat relations. Research results have expanded the
trees and down wood are formed from only a few tree
number and variety of decaying wood categories over
species that can maintain bole structural integrity as the
what was previously presented in Thomas366 and Brown.48
heartwood decays. 56 Western redcedar is especially
For example, the portion of the query (above) about cavity-
valuable in providing hollow trees because the decay-
 CHAPTER 24: WOOD LEGACIES 587

Figure 6. Figure 7.Brown creeper nest

Snags in sloughing bark. Photo:
provide Harry Hartwell.
essential
habitat for
wildlife.
Photo:
Deborah
Lindley.

nesters includes subsets of species associated with Lynx select dense patches of downed trees for denning.200
different types of cavities, such as bark crevices, hollow Jackstrawed piles of logs form a habitat matrix offering
trees, cavities in dead trees (snags), and cavities in living thermal cover, hiding cover, and hunting areas for species
trees. Further differences in matrix summaries of species’ such as marten, mink, cougar, lynx, fishers, and small
relations may result from changes in the taxonomic mammals305, 313 (Figure 8). Smaller logs benefit amphibians,
classification of species, updated species’ range and reptiles, and mammals that use wood as escape cover and
geographic distribution data, as well as data limitations shelter. Small mammals use logs extensively as runways234
for some species’ associations with specific habitat (Figure 9). California red-backed voles use Class 2-3 down
elements. It should be noted that the Habitat Elements logs for cover, and feed on fungi (especially truffles) and
matrix denotes wildlife species’ associations with the lichens growing in close association with down wood.378
various habitat elements, but does not distinguish species’ The orientation of down wood also influences wildlife
uses of habitat elements for life history needs (eg., nesting, habitat use.56 Logs oriented along slope contours may be
feeding, hibernating, etc.), nor does it indicate the degree useful travel lanes for wildlife, whereas logs oriented
of association or species’ versatility. However, species’ across contours impede travel.
habitat use information is available in the DecAID The moist environment beneath loose bark, bark piles
advisory model.226 and in termite channels of logs with advanced decay
Stumps provide a variety of wildlife habitats. Stumps provides a protected area for foraging by salamanders.22
with sloughing bark (Class 2) provide sites for bat roosts,387 The cool, moist environment of rotten wood may be
and foraging sites for flickers, and downy, hairy and required for some species of salamanders to survive heat
pileated woodpeckers. In openings, tall stumps with stress during summer. Decaying wood also provides
advanced decay (Class 3) provide nest sites for flickers, habitat for invertebrates on which salamanders and other
and subsequently for blue birds and other secondary foraging vertebrates feed (e.g., collembolans, isopods,
cavity-nesters associated with openings. Squirrels and millipedes, mites, earthworms, ants, beetles, flies, spiders
chipmunks also use stumps as lookouts and platforms for and snails).230 The folding-door spider constructs a silk
cone-shredding. The matrixes on the CD-ROM with this tube within the cracks and crevices of wood with advanced
book do not include stumps as a habitat element. decay.
• Down Woody Material (logs). Down wood affords a Generalizations regarding decay classes of habitat
diversity of habitat functions for wildlife, including structures must be made cautiously. Habitat functions may
foraging sites, hiding and thermal cover, denning, nesting, encompass several decay classes or be specific to particular
travel corridors, and vantage points for predator patterns and classes of decay.56, 233
avoidance.56, 64, 230 Larger down wood (diameter and length) Habitat structures in upper layers of the forest floor
generally has more potential uses as wildlife habitat. Large (soil, litter, duff) result from processes involving organic
diameter logs, especially hollow ones are used by material (litter, decaying roots, vertebrate and invertebrate
vertebrates for hiding and denning structures.214, 230 Bears carrion, and fecal matter) and a diverse community of
forage for invertebrates in logs during summer and fall. organisms, including bacteria, fungi, algae, protozoa,
Fishers use large logs to a limited degree as den sites.297 nematodes, arthropods, earthworms, amphibians, reptiles,
 588 Wildlife–Habitat Relationships in Oregon and Washington

in streamside forests.233, 261, 262, 376 The matrices on the CD-

ROM with this book list 27 wildlife species associated
specifically with down wood in riparian areas.
The role of down wood in salmon habitat has received
much attention over the past two decades. Large wood is
a key component of salmonid habitat both as a structural
element and as cover and refugia from high flows. Large
wood serves key functions in channel morphology, as well
as sediment and water routing.33, 38, 262 The importance of
wood to salmon habitat varies from headwater to stream
mouth.235 As stream order increases and gradient decreases
in third- to fifth-order streams, down wood is a dominant
channel-forming feature. Larger wood deflects water and
increases hydraulic diversity, producing a range of pool
Figure 8. Jackstrawed logs offer hunting areas and den sites
conditions that serve as habitats for juvenile salmonids in
for lynx. Photo: Gary Koehler.
summer.37 Diverse channel margins are a primary aspect
of rearing habitat. Flow obstructions created by large wood
provide foraging areas for young salmonid fry that are
not yet able to swim in fast currents,40 and provide refugia
to juvenile salmonids at high flow.257 In higher order
streams, flow deflections created by large wood trap
sediments and nutrients, and enhance the quality of
gravels for spawning. Down wood is less of a channel-
forming feature along large rivers, but defines meander
cutoffs and provides cover and increased invertebrate
productivity for juvenile salmonids.392

Long-Term Productivity
Long-term productivity is technically founded on the
concept of net primary productivity—the conversion of
solar energy to plant biomass, expressed as grams of
organic matter per unit area per year. At any point in time,
ecosystem productivity may vary depending on the
successional stage of vegetation, disturbances, and
variation in climate and other environmental factors. In
terms of present-day forest management objectives, long-
term ecosystem productivity typically refers to the
Figure 9. Townsend’s chipmunk, Photo: D. Anderson, production, in perpetuity, of diverse resources, including
National Museum. timber, fish, wildlife, and other ecosystem services, such
as air and water quality. Processes that sustain the long-
and small mammals167, 185, 207, 230, 249, 376 The complex trophic term productivity of ecosystems have become the
web supported by nutrient and moisture conditions within centerpiece of new directives in ecosystem management
the litter and duff layers transforms plant material into a and sustainable forestry.78, 229, 291, 320 Given the key role of
variety of degradation products, thereby storing and decaying wood in long-term productivity of forest
releasing nutrients within the ecosystem. ecosystems in the Pacific Northwest,122, 169, 261, 302 the topic
According to the Habitat Elements matrix on the CD- should remain of keen interest to scientists and managers
ROM with this book, in forest environments, 86 vertebrate during the coming decade. 149 Below, we highlight
wildlife species are associated with down wood, 38 with functions of decaying wood directly linked to long-term
litter (undecomposed fine wood), and 14 with duff productivity, including influences on the frequency and
(decomposed litter and other vegetation matter severity of disturbances such as fire, disease, and insect
underlying the litter layer). Of these species, 58 are outbreaks.5, 6, 133, 137
associated exclusively with down wood, 10 exclusively
with litter, and none exclusively with duff. Wildlife uses Nutrient Cycling and Soil Fertility. Decaying wood has
include reproduction, hibernation, feeding, resting, been likened to a savings account for nutrients and organic
sunning, drumming, preening, dusting, lookout and matter,376 and has also been described as a short-term sink,
travel. All decay stages of down wood are used by wildlife. but a long-term source of nutrients in forest ecosystems.164
Decaying wood forms many habitat structures in The total amount and distribution of nutrients in different
riparian forests. Accumulations of large wood on stream woody tissues varies from region to region and among
banks provide habitat for small mammals and birds that forest types. Coarse wood contains 0.3-4.4% of the total
feed on stream biota,103, 104 and provide structural diversity nitrogen and 4.2-10.6% of the total phosphorus in westside
 CHAPTER 24: WOOD LEGACIES 589

forests. Nutrient cycling via foliage and fine litter has been that improves soil development.164, 242 An estimated 9-68%
well-described.1, 161, 388, 408 Substantial amounts of nitrogen of the forest floor in Douglas-fir/western hemlock (Tsuga
are returned to the soil from coarse wood inputs, yet even heterophylla) clearcuts in the Pacific Northwest is derived
where annual rates of wood input are high, 4 to 15 times from decaying wood. 213 On the H. J. Andrews
more nitrogen is returned to the forest floor from foliage Experimental Forest of western Oregon, 20-30% of the soil
than from large wood.164 This is a consequence of the higher volume consists of decaying wood dispersed throughout
nutrient concentrations and shorter turnover times of leaf a matrix of litter and duff.294 Because wood is a relatively
litter compared to wood.164, 300 The relative contribution of inert substance, it may help to stabilize pools of organic
large wood to the total nutrient pool in an ecosystem matter in forests by slowing soil processes and buffering
depends to a large extent, on the size of other organic pools against rapid changes in soil chemistry. Humus and soil
in the system. organic matter are critical to site productivity on dry sites,
The proportion of tree nutrition derived from large and may limit site productivity on excessively wet and
wood during normal stand development is unknown. The cold sites.169, 377, 381
low nutrient content in wood, small mass of tree boles Litter decomposition processes in the upper soil are the
relative to foliar litterfall, and slow rates of wood decay major locus of nutrient cycling within forests of the Pacific
suggest that large wood plays a minor role in forest Northwest.1 Soil organic matter and wood can abiotically
nutrition.18, 159, 162 After large scale disturbance such as fire (chemically) retain large amounts of nitrogen and other
and blowdown, however, the large nutrient pool stored elements in forest ecosystems. A significant fraction of the
in woody structures of trees (bole, branches, twigs, roots) nitrogen is believed to be in soil organic matter pools.58, 189,
219, 266, 355, 385
becomes available to the regrowing forest. Large down Down wood and soil organic matter thus, may
wood may thus be an ample source of nutrients regulate productivity in some forest ecosytems by limiting
throughout secondary succession.281 nutrient availability. 106 Numerous studies have
The slow rate of nutrient release from decomposing demonstrated that losses in soil productivity often are
wood may serve to synchronize nutrient release with closely linked to losses in soil organic matter.298
nutritional demands in forests, and also to minimize
Moisture Retention. Water stored in large decomposing
nutrient losses via leaching to the ground water.8, 34, 83, 174,
341 wood accelerates microbial decay rates by stabilizing
In addition to nitrogen bound chemically within wood,
temperature and preventing dessication during the
down wood reduces nutrient losses from ecosystems by
summer.11, 160, 376 Moist conditions within the wood favor
intercepting nutrients in litterfall and throughfall.
decay by attracting burrowing and tunneling mammals
Favorable temperature and moisture conditions also
and invertebrates that improve aeration of wood, and by
makes large decaying wood sites of significant nitrogen
providing colonization substrate and moisture for
inputs via N-fixation.
mycorrhizae and other fungi.88, 168, 206 Moist “nurse logs”
Chronosequence studies indicate that large decaying
also provide excellent sites for seedling establishment and
wood accumulates nitrogen, calcium, and magnesium
production of sporocarps. These processes increase
over time, but loses phosphorus and potassium.76, 117, 146, 164,
205, 340 retention and cycling of nutrients within ecosystems and
The time frame for nutrient immobilization and
contribute to higher biodiversity and biomass production.
release (mineralization), as well as the specific processes
responsible for changes in nutrient content of down wood Mycorrhizae. Mycorrhiza, meaning “fungus-root”, is a
have not been fully elucidated. Mineralization of nitrogen symbiotic association of fungi with plant roots. The fungus
has been associated with a critical carbon-to-nitrogen ratio improves nutrient and water availability to the host in
of approximately 100. That is, nitrogen and other elements exchange for energy derived from plant sugars.
stored in microbial tissues are gradually released to plants Mycorrhizae are necessary for the survival of numerous
as ratios decline below 100. 105, 340 Despite growing tree families, including pine, hemlock, spruce, true fir,
recognition of the roles of decaying wood in soil Douglas-fir, larch, oak, and alder. Mycorrhizal associations
productivity, details of many processes are poorly are a source of nutrients to promote wood decay. By the
understood.125, 131, 164, 376 Harmon et al.164 describe patterns time a log reaches more advanced stages of decomposition
in the accumulation and release of major nutrients in (Class 3) fungal colonization leads to the accumulation of
decaying wood through time. nutrients in hyphae, rhizomorphs and sporocarps,23, 87
Recent studies indicate that wood may release nutrients especially for ectomycorrhizal fungi, where >90% of the
more rapidly than previously thought through a variety fungal activity is associated with organic material.119, 168, 310
of decay mechanisms mediated by means other than Ectomycorrhizal fungi decrease the ratio of carbon to
microbial decomposers, i.e. fungal sporocarps, nitrogen in decomposing wood, and mediate nutrient
mycorrhizae and roots, leaching, fragmentation, and availability to plants while improving nutrient retention
insects.107, 158, 159, 162, 339, 405 Harmon et al.162 found that during by forest ecosystems.13, 206, 251
early stages of decomposition, fungal sporocarps
Mass Wasting and Surface Erosion. In the Pacific
concentrated nitrogen, potassium and phosphorus from
Northwest, combined effects of steep slopes, high rainfall,
down wood. Annual dieback of the sporocarps returned
a history of tectonic uplift, and rapid weathering of weak
nutrients to the available soil nutrient pool.
rocks make unstable slopes a dominant erosion process.
Soil is the foundation of the forest ecosystem.68, 348 Large
Large wood helps to anchor snowpacks, limit the extent
wood is a major source of humus and soil organic matter
 590 Wildlife–Habitat Relationships in Oregon and Washington

of snow avalanches, and may even stabilize debris flows, Key Ecological Functions of Wildlife Species
depending on the depth of the unstable area.125, 356, 358 The Associated With Decaying Wood
energy derived from falling or flowing water is the driving As previously described, decaying wood provides habitat
force behind erosion processes in Pacific Northwest elements for many wildlife species. Beyond this, the
forests. By covering soil surfaces and dissipating energy associated wildlife species play diverse ecological roles
in flowing and splashing water, logs and other forms of in their ecosystems, which in turn can influence other
coarse wood significantly reduce erosion.357 Large trees species. These roles are termed “key ecological functions”
lying along contours reduce erosion by forming a barrier or KEFs, which are categorized and denoted for each
to creeping and raveling soils, especially on steep terrain. species in the Key Ecological Functions matrix on the CD-
Material deposited on the upslope side of fallen logs ROM with this book.224
absorbs moisture and creates favorable substrates for Detailed descriptions of the patterns of KEFs of wildlife
plants that stabilize soil and reduce runoff.230 species associated with wood decay have been summed
Stand Regeneration and Ecosystem Succession. from the Key Ecological Functions matrix.223 An associated
Decomposing wood serves as a superior seed bed for some “functional web” summarizing the ecological roles of
plants because of accumulated nutrients and water, wildlife associated with down wood included 86 wildlife
accelerated soil development, reduced erosion, and lower species in Washington and Oregon and traced various
competition from mosses and herbs.160, 376 In the Pacific functional categories.
Northwest, decaying wood influences forest succession Similarly, a functional web can be described here for
by serving as nursery sites for shade-tolerant species such the 96 vertebrate species associated with snags in forest
as western hemlock, the climax species in moist Douglas- or grassland/shrubland environments, as denoted by the
fir habitat.80, 123, 160, 163, 244 Wood that covers the forest floor Habitat Elements matrix on the CD-ROM with this book.
also modifies plant establishment by inhibiting plant For example, a query of the KEF matrix for these species
growth, and by altering physical, microclimatic, and shows that 40% of snag-associated species are primary
biological properties of the underlying soil. For example, consumers, 95% are secondary consumers, and 8% are
elevated levels of nitrogen fixation in Ceanothus velutinus carrion feeders. The remaining >5% are tertiary
and red alder35, 88 have been reported under old logs. consumers, cannibalistic feeders, coprophages (feed on
fecal matter), or feed on human refuse. The primary
Streams and Riparian Forests. Long-term productivity consumers include spermivores or seed-eaters (63% of all
in streams and riparian areas is closely linked to nutrient primary consumers associated with snags), frugivores or
inputs, to attributes of channel morphology, and to flow fruit-eaters (50%), sap feeders (18%). The others include
dynamics created by decaying wood.144, 233, 360 Small wood (each <15%) fungivores or fungus-eaters, foliovores or leaf-
contributes to nutrient dynamics within streams and eaters, grazers, and feeders on flowers or buds, aquatic
provides substrates to support biological activity by plants, bark or cambium, and roots. Secondary consumers
microorganisms, as well as invertebrates and other aquatic associated with snags include insectivores (90% of all
organisms.145, 262 Much of the organic matter processed by secondary consumers associated with down wood),
the aquatic community originates in riparian forests and vertebrate predators (36%), ovivores or egg-eaters (13%),
is stored as logs.90, 259 Down wood also helps to retain and piscivores or fish-eaters (8%). Percentages may sum
nutrients in streams, by trapping carcasses of dead salmon to >100% because some species have multiple roles.
and increasing carcass availability to terrestrial Various symbiotic relations can be described for the 96
scavengers.72 snag-associated species. Sixteen species are primary cavity
Large wood is the principal factor determining the excavators and 35 are secondary cavity users; 8 are
productivity of aquatic habitats in low- and mid-order primary burrow excavators and 11 are secondary burrow
forested streams.262 Large wood stabilizes small streams users; 5 are primary terrestrial runway excavators and 6
by dissipating energy, protecting streambanks, regulating are secondary runway users. Nine snag-associated species
the distribution and temporal stability of fast-water create nesting or denning structures and 8 use created
erosional areas and slow-water depositional sites, shaping structures. Sixteen species might influence vertebrate
channel morphology by routing sediment and water, and population dynamics and 22 might influence invertebrate
by providing substrate for biological activity. 361 The population dynamics. Snag-associated species also
influence of large wood on energy dissipation in streams contribute to dispersal of other organisms including seeds
influences virtually all aspects of ecological processes in and fruits (21 snag-associated wildlife species perform this
aquatic environments, and is responsible for much of the function), invertebrates (8 species), plants (8 species), fungi
habitat diversity in stream and riparian ecosystems.262, 376 (2 species), and lichens (1 species). Six snag-associated
The stair-step gradients produced by wood in small stream species can improve soil structure and aeration through
basins supports higher productivity and greater habitat digging, 2 species fragment standing wood, and 2 species
diversity than that found in even-gradient streams lacking fragment down wood. One snag-associated species creates
wood structure.32, 40, 142, 173, 254, 357 snags, and at least 1 can alter vegetation structure and
succession through herbivory.
The ecological roles of wildlife associated with various
decaying wood structures have both many similarities and
 CHAPTER 24: WOOD LEGACIES 591

salient differences. For example, both snag- and down Wood Input Rates
wood-associated wildlife more or less equally participate Primary Productivity. Regionally, ecosystem
in dispersal of seeds and fruits (although the particular accumulations of decaying wood are closely related to
species they disperse may differ); however, snag- gradients in net primary productivity as a result of
associated wildlife play a greater role in dispersal of moisture and temperature limitations. Consequently, dead
invertebrates and plants, and down wood-associated wood abundance in the Pacific Northwest declines along
wildlife play a greater role in dispersal of fungi and lichens. an easterly gradient from the highly productive Douglas-
Down wood-associated species might contribute more to fir zone120 to the high desert. A summary of existing studies
improving soil structure and aeration through digging, in Washington and Oregon164 showed greater input rates
and to fragmenting wood. of decaying wood biomass in mature and old-growth
This is one example of the far greater differentiating Douglas-fir and Sitka spruce/western hemlock forests
power afforded by a well-constructed set of matrixes than (0.20 - 12.14 Mg/ac/yr) than in higher elevation Pacific
was previously available in Thomas366 and Brown.48 That silver fir (0.12 Mg/ac/yr). No data were available for
is, a combined query of the Habitat Elements matrix and eastside forests; however, lower accumulations of dead
the Key Ecological Functions matrix can produce a list of wood in juniper forests are closely tied to low biomass
species with unique combinations of habitat element productivity, as well as fire. Detailed discussions of
associations and particular ecological functions. For regional patterns in forest productivity have been
example, other decaying wood elements, such as hollow provided elsewhere.138, 148, 208, 274, 315, 396, 398, 399
live trees, cavities in dead trees, dead parts of live trees,
Successional Development. Pulses in tree mortality occur
bark crevices and bark piles, support different arrays of
normally in stand regeneration and development. Self
wildlife species; their ecological roles (KEFs) may also
thinning from competition-induced suppression, for
differ from those discussed here.
example, reduces stand density during the course of even-
It should be recognized, too, that invertebrates play
aged stand development in western Oregon and
critical major roles in wood decay ecology in both standing
Washington. 277, 343 Suppressed trees are also more
and down wood.327 The full array of ecological roles of
susceptible to insects and diseases. Suppressed trees are
invertebrates associated with wood decay and related soil
typically of small diameter and often remain standing until
formation processes has been little studied, but is likely
blown down by wind. Compared to young regenerating
to be substantial. Included in 11 functional invertebrate
stands, natural mortality in old-growth stands usually
groups described by FEMAT,114 based on ecological roles,
affects a smaller number of trees, usually older and larger
were coarse wood chewers, litter and soil dwellers,
trees with slower inherent rates of decay and longer
understory and forest gap herbivores, canopy herbivores,
residence times compared to small trees. Natural aging
epizootic forest species, pollinators, riparian herbivores,
processes may directly cause tree mortality by impairing
and riparian predators. A 1 square meter of undisturbed
tree physiological and hydraulic functioning, or indirectly
forest soil is estimated to contain from 200 to 250 species
by predisposing trees to other agents of mortality, such as
of arthropods, including up to 75 species of fungivorous
insects, diseases, and windthrow. 194, 317, 318 393, 394, 400
mites reaching densities of 200,000 individuals.249
Consequently, older trees with a higher ratio of heartwood
to sapwood are more susceptible to heart-rot fungi, root-
Current Regional Patterns of rotting fungi, sapwood fungi, and insects such as bark
Decaying Wood beetles.79, 108, 237, 400
The input rates and average piece size of dead wood
Factors Influencing Regional Abundance generally increase with stand age,164 although the amount
of Decaying Wood of decaying wood can follow a U-shaped pattern if young
Quantities and characteristics of decaying wood in an forests inherit large amounts of decaying wood and live
ecosystem represent a balance between additions through trees from preceding stands.346 Age-related inputs and
tree death, breakage, transport, and losses through accumulation of wood throughout stands succession in
processes of decomposition and fire consumption, all of the Pacific Northwest have been summarized.71, 164, 277, 346
which can be influenced by forest management Information is lacking to describe natural patterns of
activities.164 Management effects on the abundance of decaying wood inputs and accumulation during stand
decaying wood can be accurately assessed only within the development in forests east of the Cascade Range.
context of underlying natural patterns of wood dynamics. Fire. Prior to European settlement, natural disturbance
The following section describes predominant influences regimes dominated by fire had a major influence on input
on accumulations of decaying wood in forests of the Pacific rates and accumulations of dead wood in forests of the
Northwest, including inputs (primary productivity and Pacific Northwest,2, 5, 390 but other factors such as wind,
mortality), and outputs (decay and combustion), and pathogens, and insects also influenced forest development
interactions with various disturbances of natural and to varying degrees. Fire effects on ecosystem inputs of
human origin. wood are varied and complex, depending on specific
ecosystem attributes of soil, vegetation characteristics, and
historic disturbance patterns.1 Fire plays a major role in
 592 Wildlife–Habitat Relationships in Oregon and Washington

the dynamics of decaying wood by altering the rates and in flooded soils can inhibit decay rates of wood in close
composition (size, species) of dead wood inputs, as well contact with the soil, such as soil humus, small diameter
as combustions losses. The historic frequency and severity wood, and large wood in advanced stages of decay. Poor
of fire events varies widely across the region. East of the aeration inhibits nitrogen mineralization and induces
Cascade Range, pre-settlement forests were subject to denitrification, reducing nitrogen supply to wood
frequent low-intensity fires that killed only a small decomposers. 179 Prolonged flooding that eliminates
percentage of living trees, but consumed much of the beneficial populations of soil decomposer organisms thus,
decaying wood (85%), due to low moisture contents of can also inhibit decay rates of wood.353
fuel wood during late summer.5, 24, 95 In moist forests west Wind is one of the most common disturbances to
of the Cascade crest, by comparison, fire disturbances were forests, with damage ranging from single trees to many
infrequent, but of high intensity and magnitude, hectares. In historic times, wind was the major disturbance
producing high inputs of wood from tree mortality.2, 345, 346 factor controlling inputs of decaying wood to forests west
For example, Agee and Huff6 reported a 10-fold increase of the Cascade crest in Washington and Oregon, and a
in snags and a 150% increase in down wood in old-growth primary influence in maintaining soil productivity.46, 399
hemlock-Douglas-fir forests after wildfire. Wind produces pulsed inputs of decaying wood, often
selecting for larger trees, though smaller trees in dense
Insects and Pathogens. Insects and pathogens deserve
stands may also be highly vulnerable to wind, especially
special mention because they have had a major influence
along natural topographic breaks, and on the windward
on historic inputs of dead wood in the Pacific Northwest.
side of harvest edges. Windthrow frequently also triggers
Although these disturbance agents are natural
outbreaks of bark beetles. Windthrow risk is difficult to
components of the ecosystems, their prevalence and
predict, as it is is influenced by a complex interactions at
degree of influence on ecosystem structure and function
scales ranging from individual trees to stands, to the
may be greatly modified by human activities. Insects and
surrounding landscape. Excellent discussions of
pathogens play a key role in maintaining diverse and
windthrow processes in ecosystems have been presented
productive forests by creating habitat and stimulating
elsewhere.270, 335, 349 A recent study by Sinton et al.337 is the
nutrient cycling. Catastrophic insect and pathogen
first to evaluate the landscape context of forests as well as
infestations, though, may be symptomatic of imbalances
environmental factors contributing to historic windthrow
causing reduced ecosystem productivity.
patterns.
Insects and pathogens of Pacific Northwest forests are
The Pacific Northwest is a region of active volcanic
too numerous to describe in entirety here, but detailed
influence, most recently impacted by the explosion of Mt.
information is available elsewhere.133, 220, 322 Insects can be
St. Helens.128, 129 Volcanic events variably contribute to large
separated into general groups of defoliators, sap-suckers,
input, redistribution, and consumption of dead wood to
bark beetles, wood borers, cone and seed insects, and
ecosystems depending on proximity to the blast zone,
excavators of seasoned wood. Insects may be primary
pyroclastic flows, and debris flows. Thus, effects of
agents of tree mortality or incidental agents that take
volcanic activity on the subsequent abundance and decay
advantage of trees weakened by other stresses. Bark beetle
rates of wood may be highly variable and difficult to
species represent both groups and may inhabit trees that
predict. Mudflows from volcanic events can completely
are healthy, dying, or recently dead. Of the numerous
cover down wood and create a sanitized soil surface with
varieties of tree pathogens, those that cause brooming in
reduced populations of decomposers.101 However, plant
conifers induce substantial inputs of decaying wood in
succession after volcanic events may include many N-
Pacific Northwest forests. These include the dwarf
fixing species that serve to increase N availability and
mistletoes (Arceuthobium spp.), rust fungi (Chrysomyxa spp.
accelerate wood decomposition.
or Melampsorella spp.), or the needle cast fungus
Landslides are a natural but infrequent occurrence
(Elytroderma deformans). Dwarf mistletoes are perennial
(estimated interval of 500-1500 years) on steeper slopes
parasitic plants that derive water and nutrition from their
within the region.31, 190, 361 Altered slope hydrology and soil
hosts, progressively killing branches, trees, and sometimes
stability due to activities such as road building and
entire stands of vulnerable species.171 Broom rusts are
logging, however, can increase the intensity, magnitude,
obligately parasitic pathogenic fungi that form witches’
and frequency of such events.262 Landslides in the upper
brooms on their coniferous hosts.407 Needle cast fungus
basins of steep mountainous terrain form debris flows that
causes early needle drop in pines, and can form witches’
transport most of the sediment and wood in first- and
brooms similar in appearance to dwarf mistletoe.77
second-order (i.e. low-order) channels.359 Landslides and
Other Natural Disturbances. Flooding has diverse effects debris flows also contribute most of the sediment and
on ecosystems processes, thus specific effects on net coarse material into larger third to fifth-order channels
accumulation of decaying wood may be difficult to (i.e. mid-order). These geomorphic events increase and
discern. Streamside forests are often perturbed and even redistribute large amounts of wood in Northwest
destroyed by flooding. Periodic flooding produces pulsed ecosystems.
inputs of wood to forest ecosystems, but chronic flooding
may eventually cause conversion to non-timber vegetation
by impairing site productivity for trees. Reduced aeration
 CHAPTER 24: WOOD LEGACIES 593

Wetlands Perpetuate Wood Legacies Wood structures contribute to wetland ecological

Richard Bigley function. Water storage in wetlands serves to reduce
flooding at lower elevations, and to moderate
Forested wetlands in the Pacific Northwest often seasonal variability in streamflow. Hydrological
contain a disproportionally high concentration of functions and surface runoff thus are major concerns
snags and down wood compared to most upland in managing forests in and around wetlands. In
forests. Inundation promotes the accumulation of forested wetlands where ground vegetation can be
down wood by limiting decay and creating snags. sparse, down wood is often a dominant influence
Decaying wood contributes to many wetland in water routing and channel morphology.
functions and exerts considerable influence on water Down wood significantly reduces erosion by
quality and flow routing in the forest. dissipating flow energy and trapping sediments.
Environmental conditions for wood decay in Stable wood structures in wetlands create effective
wetlands. Flooding often produces anaerobic soil barriers to downslope transport of unconsolidated
conditions that curtail the decomposition of wood sediments, and facilitate vegetation establishment.
and other organic matter for many years. Buried Major fluvial erosion and deposition processes are
wood thousands of years old has been recovered influenced by overland flow resulting from heavy
from deep peat and other anaerobic environments. rainfall events and the limited infiltration capacity
Anaerobic conditions in wetlands inhibit wood of some forest soils in the Pacific Northwest.
decomposition rates by favoring facultative and As with upland sites, down wood in wetlands
obligate anaerobes that are less efficient dcomposers provides sites for plant germination and
than areobic bacteria. Many fungal decomposers establishment, protects vegetation from grazing, and
become dormant when water-saturated, and serves as travel corridors for wildlife.
continue to grow only when oxygen is restored. Wetland management and wood legacies. Forest
Clues as to the frequency, duration, and management in and around wetlands is primarily
seasonality of wetland conditions can be inferred concerned with soil disturbance and windthrow.
from soil features. Short periods of flooding may not Selective harvest and salvage of valuable western
be sufficient to inhibit wood decomposition in red cedar (Thuja plicata) has been the main impetus
upland forests. Soils in areas without prolonged for management activities in forested wetlands of
flooding usually have evenly-distributed red and the Pacific Northwest. Management objectives to
yellow colors characteristic of oxidized iron. In retain snags and down wood in forested wetlands
contrast, forests with prolonged flooding often have require more detailed consideration of the different
deep organic surface horizons and underlying ecological functions of fast-growing hardwoods like
mineral soils lacking oxidized iron. Fluctuating popular (Populus tremulodes) and more decay-
water tables can usually be identified by the degree resistant but slower-growing species like western
of “mottling” (red and yellow flecks) resulting from red cedar.
variable oxygen levels (and hence iron oxidation)
in soils.

Wood Output Rates Cascades. Mesoscale atmospheric phenomena (e.g.

Aside from wood removal in commercial harvesting, orographic effects) that introduce finer-scale climate
decay and combustion processes are the primary factors patterns also produce more variability in wood decay rates
influencing the ouputs of decaying wood over time. Wood within ecosystems. Microclimatic variability at the scale
decay rates have been estimated for a number of forest of stands and individual trees is also affected by forest
types, but are highly sensitive to wood structure and type, successional stage, aspect, slope, elevation, soil
chemistry as described earlier, and to climatic conditions, drainage, shading, and other factors.131, 141, 146, 234 Wood size,
especially temperature, moisture, and aeration.18, 105, 113, 146, shape, and placement affect decomposition rates by
164, 300, 340
Decay rates of wood are most rapid under changing microclimates and wood exposure to
conditions of warm temperature and good aeration that decomposers.56, 115, 205, 218, 341
limit the metabolic rates of decomposer organisms. In addition to stimulating inputs of dead wood, fire
Adequate moisture also is required to soften wood eliminates dead wood via processes of decay and
structure.376 combustion. Decay rates are influenced by climatic and
Seasonal patterns of temperature and moisture in nutritional conditions for decay organisms in the post-fire
decaying wood have been reviewed in detail.164, 230 In the environment.193, 245, 379 Higher rates of wood decay in the
western Cascades, decay is limited more by temperature warmer temperatures of forest openings created by fire,
during the winter and by moisture in the summer. may be counterbalanced by moisture limitation. Similarly,
Aeration may limit decay where poor drainage results in changes in nutrient availablility after fire, may either
saturated soil and a perched water table. The brief warm stimulate or inhibit decay. Cool-burning fires consume less
season limits decay processes more in the eastern wood and produce a pulse of nutrients to decomposers.
 594 Wildlife–Habitat Relationships in Oregon and Washington

Conversely, high intensity fires consume more wood and Hence, wood accumulation and decay processes may
may cause nutrient losses via volatilization and leaching, exhibit considerable variation from one ecosystem to
or by depressing populations of beneficial microbial and another.
invertebrate decomposer organisms in soil. 2, 5, 43 In
Anthropogenic Pollution. Anthropogenic pollution can
addition, charring and case-hardening of wood surfaces
affect stand microclimates, soil chemistry, populations of
by fire can retard decay processes.218
decay organisms, and a variety of basic ecosystem
Effects of Human Disturbances on Wood processes, thus it is not surprising that pollutants alter
Dynamics processes of wood decay.1, 280, 329 Ecological effects on forest
Timber Management. Intensive forest management for ecosystems are highly specific to the source, type, and
timber objectives has been shown to simplify stand and magnitude of the pollutants. Of the diverse pollutants
landscape structure.154 Harvest and burn cycles of 60-100 affecting forest ecosystems, nitrogen pollution is the most
years in the western Cascades have been estimated to be widespread and has been thoroughly studied.166, 219, 264, 288,
329
equivalent to a 4- to 5 -fold increase in fire frequency Nitrogen pollution is of concern because biomass
compared to natural conditions, in terms stand structure.368 production and storage in many forest ecosystems is
Only two rotations of managed Douglas-fir have been tightly regulated by nitrogen availability. One of the most
estimated to reduce the abundance of dead wood by 90%, comprehensive studies of pollution impacts to date
compared to levels in natural old-growth systems. In documented inhibited decomposition rates of organic
addition to removing wood, management activities also material at a site receiving chronic inputs of nitrogen and
influence wood decay rates. For example, warmer sulfur.166 Elevated nitrogen loading has been documented
microclimates after harvesting stimulate decay. in coastal watersheds and estuarine environments;380
Fertilization conducted to improve wood production in consequently, wood dynamics may be altered in these
commercial forest lands312 can accelerate decomposition environments, as well.
rates of down wood by supplying decomposer organisms Fire Suppression. In the eastern Cascades and through
with limiting nutrients, particularly nitrogen. Conversely, much of the intermountain area, extensive forest insect
intensive practices that deplete soil nitrogen, such as short- and disease problems have resulted from decades of fire
rotation, whole tree harvesting198 may inhibit rates of wood suppression in combination with selective harvesting of
decomposition over the longer term. Models estimate that pines.177, 194, 236, 401, 403 An analysis of landscape dynamics in
between 50 to 100 years are required to replace nitrogen the Interior Columbia River Basin302, 379 revealed that fire
losses after timber harvest.1 suppression resulted in a decreased abundance of large-
Carbon Storage and Climate Change. Douglas-fir forests diameter trees, and caused fuel accumulations that
in low elevation temperate rain forests have the capacity predisposed forests to stand-replacement fires. As
to store more carbon in the form of biomass and dead mentioned previously, more intense fires not only
wood than any forest ecosystem on earth. 197, 258 consume more wood, but can inhibit wood decay by
Consequently, these forests provide another benefit of reducing nitrogen availability (and other elements)
global significance—the ability to sequester carbon from through volatilization and leaching, especially for wood
the atmosphere, and potentially to counteract global in close association with the soil.245 Wood decay in post-
warming.16 Intensive forestry practices that released large fire regenerating forests also may be exacerbated by a
quantities of carbon stored as biomass and dead wood in decline in symbiotic nitrogen-fixing plant species in stands
Northwest forests during the 1980s have been implicated subject to prolonged fire suppression.169
as a contributing factor in the rising atmospheric
concentrations of carbon dioxide. 19, 161 The relative Summary of Regional Patterns of Abundance
contribution of carbon storage in terrestrial forests to for Snags and Down Wood
climatic warming remains equivocal, however.16 The Regional variation in dead wood abundance reflects strong
potential for warmer climates to accelerate decomposition underlying gradients in physical environment,
of decaying wood in terrestrial systems remains an issue disturbance, and biological processes that affect
of concern.294 community composition and structure, forest dynamics,
Furthermore, ecosystem accumulation of total wood and rates of dead wood input and output, as reviewed
(live and dead) has been postulated to increase in response earlier in this chapter. These factors interact in complex
to temporary stimulation of biomass productivity by ways in influencing abundance of dead wood on a given
higher temperatures, carbon dioxide, and nitrogen site. Basic information about regional ecological patterns
availability. 19, 308, 309 The increased rates of wood of dead wood can provide context for management
accumulation may level off, however, as nitrogen decisions at a variety of scales, and for analyzing forest
availability to forests stabilizes.100 As a result of global policy at regional and national levels. A wealth of
warming, complex linkages between temperature, carbon information on snags and down wood has recently become
dioxide, plant growth, decomposition, and nutrient available from grids of field plots established by regional
cycling could alter the size and relative distributions of forest inventories across Washington and Oregon. This
carbon and nutrient pools in forest soils and vegetation. section summarizes findings of a recent analysis of these
data by Ohmann and Waddell.276
 CHAPTER 24: WOOD LEGACIES 595

Ohmann and Waddell276 analyzed data on dead wood down wood in montane mixed-conifer forests were
collected on 16,867 field plots in 9 forested wildlife habitat significantly different from almost all other wildlife habitat
types.188 The plots represent about 49 million ac (20 million types.
ha) of forest within Washington and Oregon, about 54 Much of the regional variation in dead wood
percent of which is publicly owned, 24 percent is owned abundance, expressed as differences among the wildlife
by timber industry, and 22 percent is owned by habitat types, probably can be attributed to strong
nonindustrial private landowners.296 The field plots were gradients in net primary productivity. For example, the
measured from 1984-1997 as part of inventories conducted large amount of dead wood in westside conifer-hardwood
by the BLM (Natural Resource Inventory, NRI); the USDA forests probably can be explained by high rates of input
Forest Service, National Forest System (Current Vegetation within these forests, which are the most productive of the
Survey, CVS); and the USDA Forest Service, Pacific wildlife habitat types.120 In contrast, the large amount of
Northwest Research Station (Forest Inventory and dead wood in montane mixed-conifer forest also may be
Analysis, FIA). Dead wood data were unavailable for explained by slow rates of decomposition in the cold
national and state parks, and down wood data were temperatures at high elevations. The high density of snags
unavailable for nonfederal lands in Oregon and western in the subalpine parkland and montane mixed-conifer
Washington. Snags on plots were sampled on fixed- and types may be a function of high mortality rates and low
variable-radius circular plots, and down wood was fall rates in these habitats.
sampled on line transects. See Ohmann and Waddell276
for information about inventory design, field methods, Successional Patterns of Dead Wood
and data summary. Snag density generally increased with stand age. Within
Plots in upland forest were classified into 1 of 9 wildlife wildlife habitat types, total snag density always was
habitat types based on potential natural vegetation and lowest in the early successional stage and usually was
ecoregion.75 Each plot was also classified into 1 of the highest in the late stage, although no differences were
following 3 successional stages, which are groupings of detected among stages of subalpine parkland, ponderosa
the structural stages based on current vegetation pine, and western juniper (Table 1). Large snag abundance
structure188: early (tree stocking [sensu217] <10 percent, or increased with successional development in all of the
tree stocking >=10 percent and quadratic mean diameter wildlife habitat types except western juniper woodland,
[QMD] 1.0-9.8 in [2.5-24.9 cm]), middle (tree stocking >=10 where no trends were evident (Table 1).
percent and QMD 9.8-19.6 in [25.0-49.9 cm]), late (tree The volume of total and large down wood also
stocking >=10 percent and QMD >=19.7 in [50.0 cm]). The generally increased with forest development, but
QMD is the diameter of the tree of average cross-sectional successional patterns differed somewhat among the
area at breast height (4.5 ft; 1.37 m) on the plot. wildlife habitat types (Table 2, 3). Late successional stages
contained the largest concentrations of both total and large
Differences in Dead Wood Among Wildlife Habitat down wood in 7 of the 9 wildlife habitat types (Table 2, 3).
Types In the westside wildlife habitats and in montane mixed-
The abundance of snags and down wood varied conifer forest, the volume of total and large down wood
substantially across the region. The greatest differences in the late stage usually was significantly different from
in dead wood abundance were among the wildlife habitat the early and mid stages, but early and mid stages usually
types, although differences among successional stages were not significantly different from one another (Table
within wildlife habitat types also were significant in many 2, 3). Large down wood volumes differed significantly
cases. Total snag densities were greatest at higher between the early and middle successional stages in all of
elevations: 15.1/ac (37.2/ha) in montane mixed-conifer the eastside wildlife habitat types except western juniper
forest and 14.6/ac (36.0/ha) in subalpine parks (Table 1). woodland (Table 3).
Snags were least dense in the drier wildlife habitat types No wildlife habitats exhibited a U-shaped pattern in
on the eastside: 0.3/ac (0.8/ha) in western juniper snag abundance, which can occur if large amounts of wood
woodland and 2.0/ac (5.0/ha) in eastside ponderosa pine is inherited from a preceding stand.346 Down wood also
(Table 1). Large snags were most abundant in montane most often increased with succession, but this pattern was
mixed-conifer forest (3.8/ac; 9.6/ha) and in westside less consistent than for snags, and some wildlife habitat
conifer-hardwood forest (2.2/ac; 5.5/ha), and least types did exhibit a U-shaped pattern. The lack of a U-
abundant in western juniper woodland (0.1/ac; 0.2/ha) shaped successional pattern for snags is not surprising.
and ponderosa pine (0.4/ac; 1.0/ha) (Table 1). The Snags have much shorter lag times in the forest than down
volumes of both total and large down wood were greatest wood. Natural processes of fragmentation and
in westside conifer-hardwood forest and lowest in western decomposition begin much sooner, and they disappear
juniper woodland (Table 2, 3). Total down wood volume as recognizable structures much faster.164 In addition, much
among wildlife habitat types ranged from 105.6 to 2,619.7 of the dead wood in westside forests is input directly as
ft3/ ac (7.4 to 183.3 m3/ha), and large wood from 64.3 to down wood rather than snags.164 Snags also are much more
1,883.7 ft3/ac (4.5 to 131.8 m3/ha). Differences in total dead likely than down wood to be damaged or intentionally
wood generally were more pronounced among wildlife removed by humans through the course of forest
habitat types on the west side of the Cascades than among management and harvest activities. In a previous analysis
the eastside types, and the amounts of total snags and of regional plot data, Hansen et al.154 found that large snags
 596 Wildlife–Habitat Relationships in Oregon and Washington

Table 1. Weighted mean (standard error) density of snags by wildlife habitat type, successional stage, and
snag size, Oregon and Washington.1

Successional Stage and Snag Size

Early Middle Late All stages
Wildlife habitat type Total Large Total Large Total Large Total Large

Mean (SE) trees per acre

Westside conifer-hardwood 2.1 a 0.8 a 7.2 b 2.4 b 12.7 c 5.8 c 5.8 2.2
(0.1) (0.0) (0.2) (0.1) (0.4) (0.2) (0.1) (0.1)
Westside white oak-Douglas-fir 2.5 a 0.6 a 4.6 a 1.1 a 6.9 b 2.4 b 4.1 1.0
(0.4) (0.2) (0.4) (0.1) (1.0) (0.4) (0.3) (0.1)
SW OR mixed conifer-hardwood 3.8 a 1.0 a 6.9 b 2.0 b 8.5 b 3.8 c 6.2 2.1
(0.4) (0.1) (0.4) (0.1) (0.4) (0.2) (0.2) (0.1)
Montane mixed-conifer 7.2 a 1.2 a 20.0 b 4.2 b 16.3 c 8.8 c 15.1 3.9
(0.6) (0.1) (0.5) (0.2) (0.6) (0.3) (0.4) (0.1)
Subalpine parkland 14.1 0.7 a 15.1 2.3 b NA NA 14.6 1.5
(3.0) (0.2) (1.7) (0.4) (1.7) (0.2)
Eastside mixed-conifer 6.0 a 0.8 a 8.7 b 1.7 b 8.4 3.2 c 7.9 1.5
(0.4) (0.1) (0.2) (0.0) (0.2) (0.2) (0.2) (0.0)
Lodgepole pine 6.7 a 0.3 a 11.2 b 0.9 b NA NA 8.0 0.5
(0.6) (0.0) (1.1) (0.1) (0.5) (0.0)
Ponderosa pine (eastside) 2.0 0.4 2.0 1.1 2.1 0.6 2.0 0.4
(0.2) (0.0) (0.2) (0.0) (0.3) (0.1) (0.1) (0.0)
Western juniper 0.3 0.1 0.6 0.2 0.3 0.1 0.3 0.1
(0.1) (0.0) (0.2) (0.1) (0.2) (0.1) (0.1) (0.0)
All wildlife habitat types 3.7 a 0.8 a 8.6 b 2.1 b 11.5 c 5.4 c 7.0 2.0
(0.1) (0.0) (0.1) (0.0) (0.2) (0.1) (0.1) (0.0)

1
NA: Not applicable — sample size <10 plots.
Significantly different means (alpha=0.05) within rows (among successional stages for a given snag size-class) are indicated by
different letter footnotes.
“Total” includes snags >= 10.0 in DBH, decay classes 1-5, and >=6.6 ft tall
“Large” includes snags >=19.7 in DBH, decay classes 1-5, and >=6.6 ft tall.
Table modified from Ohmann and Waddell (in press).

were 3-5 times more abundant in stands that had never vs. 323.1 ft 3/ac [22.6 m 3/ha] inside wilderness) and
been clearcut than in stands that had been clearcut at least montane mixed-conifer (1,061.9 ft3/ac [ 74.3 m3/ha] vs.
once. These factors taken together suggest that snag levels 502.9 ft3/ac [35.2 m3/ha]).
would more closely track recent disturbance and forest The wilderness stratification of the plots was intended
succession, while down wood amounts would be more to separate plots with different likelihoods of having been
strongly influenced by the long-term history and disturbed by timber harvest and management. However,
productivity of the site. comparisons of dead wood within and outside of
wilderness areas must be interpreted with caution. Plots
Dead Wood in Wilderness Areas in wilderness are strongly biased towards higher
Over all wildlife habitat types, large snags were more than elevations and lower productivities, which may account
twice as dense in wilderness areas than outside wilderness for much of the higher amounts of down wood outside
(Figure 10 a). The strongest differences were for westside wilderness. In addition, even though wilderness areas are
conifer-hardwood forest (2.06/ac [5.1/ha] outside off-limits to future timber harvesting, they have been
wilderness vs. 6.15/ac [15.2/ha] within wilderness), affected by other human activities to some degree (e.g.,
eastside mixed-conifer forest (1.30/ac [3.2/ha] vs. 3.84/ roads, recreation, exotic species introduction, fire
ac [9.5/ha]), and lodgepole pine (0.3/ac [0.8/ha] vs. 1.1/ suppression). Furthermore, many plots outside wilderness
ac [2.7/ha]). In contrast, large down wood was more areas sample old growth and younger natural forest.
abundant outside wilderness than within wilderness in Nevertheless, if snags are more strongly influenced by
all of the wildlife habitat types except eastside ponderosa timber management activities than down wood, then
pine (Figure10 b), although the differences usually were wilderness areas would be more likely to contain greater
not significant. The most pronounced differences in down amounts of snags than areas outside wilderness, as the
wood volume were in southwest Oregon mixed conifer- data show. In fact, OSHA standards historically have
hardwood (918.9 ft3 /ac [64.3 m3/ha] outside wilderness
 Table 2. Weighted mean (standard error) volume, percent cover, and density of total down wood by
wildlife habitat type and successional stage, Oregon and Washington.1

Successional Stage
Wildlife habitat type Early Middle Late All stages Early Middle Late All stages Early Middle Late All stages

Mean (SE) cubic feet per acre Mean (SE) percent cover Mean (SE) pieces per acre

Westside conifer- 2,169 a 2,396 a 3,234 b 2,619 4.4 a 4.5 a 5.5 b 4.8 132.1 a 102.9 a 101.5 b 109.7
hardwood (109) (89) (120) (61) (0.2) (0.1) (0.2) (0.1) (5.6) (2.8) (2.8) (2.1)
Westside white oak- 772 632 1,017 733 1.8 1.5 2.2 1.7 70.5 41.1 51.9 48.9
Douglas-fir (167) (87) (176) (73) (0.3) (0.1) (0.3) (0.1) (11.1) (4.1) (7.5) (3.8)
SW OR mixed conifer- 1,213 a 983 a 1,695 b 1,227 2.5 a 2.2 a 3.2 b 2.5 64.3 a 59.7 a 69.7 b 63.5
hardwood (123) (66) (114) (54) (0.2) (0.1) (0.2) (0.1) (5.9) (3.2) (3.8) (2.3)
Montane mixed-conifer 1,459 a 1,608 a 2,837 b 1,769 3.8 a 4.3 b 5.1 c 4.3 102.8 a 101.3 b 90.0 c 99.8
(91) (47) (151) (46) (0.2) (0.1) (0.2) (0.1) (4.1) (2.6) (3.8) (1.9)
Subalpine parkland 502 789 NA 629 1.6 2.1 NA 1.8 45.8 52.2 NA 47.8
(104) (149) (90) (0.3) (0.4) (0.2) (7.0) (7.9) (5.1)
Eastside mixed-conifer 672 a 780 b 840 b 753 2.0 2.2 a 1.9 b 2.1 63.9 57.8 a 41.3 b 58.5
(31) (19) (70) (16) (0.1) (0.0) (0.1) (0.0) (2.0) (1.1) (2.7) (0.9)
Lodgepole pine 714 792 NA 734 2.9 2.8 NA 2.9 81.6 65.2 NA 77.3
(36) (60) (31) (0.1) (0.2) (0.1) (3.4) (5.0) (2.8)
Ponderosa pine (eastside) 304 a 406 b 282 362 0.8 a 1.0 b 0.7 a 0.9 28.4 a 31.9 b 18.1 a 29.7
(19) (20) (43) (14) (0.0) (0.0) (0.1) (0.0) (1.6) (1.2) (2.8) (0.9)
Western juniper 107 104 107 106 0.2 0.3 0.3 0.2 6.6 9.3 10.9 7.8
CHAPTER 24: WOOD LEGACIES 597

(44) (27) (67) (27) (0.1) (0.1) (0.2) (0.0) (1.7) (2.5) (6.6) (1.4)
c
All wildlife habitat types 1,000 a 1,109 b 2,306 1,255 2.6 a 2.7 b 4.1 c 2.9 74.0 a 67.7 b 77.4 c 71.0
(27) (19) (64) (17) (0.1) (0.0) (0.1) (0.0) (1.4) (0.8) (1.6) (0.7)

1
NA: Not applicable — sample size <10 plots.
Total down wood includes pieces >=4.9 in large end diameter, decay classes 1-4, and >=6.6 ft long.
Significantly different means (alpha = 0.05) among successional stages are indicated by different letter footnotes.
Table modified from Ohmann and Waddell (in press).
 Table 3. Weighted mean (standard error) volume, percent cover, and density of large down wood
by wildlife habitat type and successional stage, Oregon and Washington.1

Successional Stage
Wildlife habitat type Early Middle Late All stages Early Middle Late All stages Early Middle Late All stages

Mean (SE) cubic feet per acre Mean (SE) percent cover Mean (SE) pieces per acre
598

Westside conifer- 1,408 a 1,709 a 2,459 b 1,883 1.9 a 2.2 a 3.1 b 2.4 23.9 a 19.7 a 24.1 b 22.3
hardwood (97) (81) (114) (57) (0.1) (0.1) (0.1) (0.1) (1.6) (0.9) (1.1) (0.7)
Westside white oak- 442 373 609 432 0.6 0.6 0.9 0.6 9.0 5.9 6.5 6.6
Douglas-fir (147) (79) (154) (64) (0.2) (0.1) (0.2) (0.1) (2.4) (1.1) (1.8) (0.9)
SW OR mixed conifer- 766 a 609 a 1,197 b 803 0.9 a 0.9 a 1.6 b 1.1 7.9 a 7.4 a 12.0 b 8.9
hardwood (107) (57) (106) (49) (0.1) (0.1) (0.1) (0.1) (1.2) (0.8) (1.2) (0.6)
Montane mixed-conifer 699 a 660 a 2,071 b 906 1.0 a 1.0 a 2.9 b 1.3 10.3 a 8.7 a 23.4 b 11.6
(80) (34) (137) (40) (0.1) (0.0) (0.2) (0.1) (0.9) (0.4) (1.5) (0.4)
Subalpine parkland 104 319 NA 207 0.2 0.5 NA 0.4 2.6 4.4 NA 3.4
(47) (91) (54) (0.1) (0.1) (0.1) (1.2) (1.4) (0.9)
Eastside mixed-conifer 247 a 329 b 513 c 317 0.4 a 0.5 b 0.7 c 0.5 3.6 a 4.2 b 6.4 c 4.2
(26) (14) (60) (13) (0.0) (0.0) (0.1) (0.0) (0.2) (0.2) (0.7) (0.1)
Lodgepole pine 80 a 209 b NA 116 0.2 a 0.3 b NA 0.2 1.7 a 3.2 b NA 2.1
(13) (40) (14) (0.0) (0.1) (0.0) (0.3) (0.7) (0.3)
Ponderosa pine (eastside) 147 a 221 b 160 191 0.2 a 0.3 b 0.3 0.3 2.3 a 2.9 b 2.5 2.7
(16) (17) (34) (11) (0.0) (0.0) (0.1) (0.0) (0.2) (0.2) (0.6) (0.2)
Western juniper 71 50 89 64 0.1 0.1 0.2 0.1 0.4 0.8 4.5 0.7
(40) (23) (66) (24) (0.0) (0.0) (0.2) (0.0) (0.2) (0.4) (3.3) (0.2)
c
All wildlife habitat types 496 a 579 b 1,695 720 0.7 a 0.8 b 2.2 c 1.0 7.1 a 6.9 b 17.5 c 8.5
(23) (16) (59) (14) (0.0) (0.0) (0.1) (0.0) (0.3) (0.2) (0.6) (0.2)
Wildlife–Habitat Relationships in Oregon and Washington

1
NA: Not applicable — sample size <10 plots.
Large down wood includes pieces >=19.7 in large end diameter, decay classes 1-4, and >=6.6 ft long.
Significantly different means (alpha = 0.05) among successional stages are indicated by different letter footnotes.
Table modified from Ohmann and Waddell (in press).
 CHAPTER 24: WOOD LEGACIES 599

Figure 10. Abundance of dead wood by habitat type and Comparisons with Other Studies
reserved status, Oregon and Washington. (a) Weighted mean Very few estimates of dead wood abundance at broad
density of snags >=19.7 in DBH, decay classes 1-5, and geographic scales are available for comparison with results
>=6.6 ft tall. (b) Weighted mean volume of down wood from Ohmann and Waddell.276 Direct comparisons with
>=19.7 in diameter at the large end, decay classes 1-4, and published studies are extremely difficult to make because
>=6.6 ft long. Error bars indicate one standard error of the of differences in geographic location; the vegetation types,
mean. WCNH=westside conifer—hardwood, WODF=white stand ages, and disturbance histories sampled; sampling
oak—Douglas-fir, SWOMCH=southwest Oregon mixed design; definitions (e.g., dead wood sizes and decay
conifer-hardwood, MMC=montane mixed-conifer, classes); and units of measure (numbers of trees, volume,
PARK=subalpine parkland, EMC=eastside mixed-conifer, density, cover, or linear meters). Other regional studies of
LP=lodgepole pine, EPPWO=eastside ponderosa pine—white
dead wood in Washington and Oregon have been
oak, JUN=western juniper, ALL=all habitats. There were <10
restricted either to federal or to nonfederal lands, which
plots in reserved western juniper.
usually represent very different ecological conditions.274
A study by Ohmann et al.275 was limited to snags on
nonfederal lands, because data were unavailable for dead
required the removal of most snags from harvest units for
wood on federal lands and down wood on nonfederal
worker safety. Therefore, fewer snags were expected in
lands at that time. The study by Spies et al.346 was confined
managed stands outside wilderness. If snags are cut and
to natural Douglas-fir forests >40 years old on federal
left on site, this would contribute to the larger amount of
lands on the westside. Published information for eastside
down wood we observed outside wilderness areas. High
forests is not available,111 or consists of summaries of a
snag densities in the high elevation habitats (subalpine
few local studies.56 Scientists for the Interior Columbia
parkland and montane mixed conifer) also result from
River Basin Ecosystem Management Project relied on
inaccessibility for timber and firewood cutting.
expert opinion and local studies to estimate current and
Distribution of Dead Wood Abundance historical amounts of decaying wood.202 Harmon et al.164
Estimates of wood abundance in Ohmann and Wadell276 did not include any studies from eastern Washington or
are regional averages within wildlife habitats. The eastern Oregon.
standard errors of these estimates were fairly low because Estimates of down wood volume presented by
of the very large sample sizes for most of the wildlife Ohmann and Waddell276 are somewhat lower than other
habitat types and successional stages. In reality, the plot- published numbers, but this is expected for several
level amounts of dead wood within the wildlife habitat reasons: the minimum diameters are larger than in many
types were extremely variable. This variability reflects the other studies; the data describe both managed and natural
high spatial and temporal variability in the many forests of all ages, not just older natural forests originating
interacting environmental and disturbance factors that after fire; down wood of decay class 5 is excluded; the
influence dead wood on a site. All of the wildlife habitat values represent means across many stands, including
types examined had similar patterns: distributions were stands where no dead wood was observed, and maximum
non-normally distributed and strongly skewed to the values are not presented. Estimates of percent cover of
right. A large proportion of the plots contained no snags down wood also may be lower than in other studies that
or down wood, and a very small proportion of the plots used plot sampling or total tallies, as percent cover
contained extremely large accumulations of dead wood. calculated from line intersect sampling has been shown
Mean values for these skewed distributions must be to underestimate true values.27
interpreted with caution. The distribution of snags for the Although estimates of mean total down wood volume
conifer alliance of westside conifer-hardwood forest in successional stages of westside conifer-hardwood forest
(Figure 11) illustrate this pattern. In this wildlife habitat ranged from 2,169.2 to 3,233.8 ft3/ac (151.8 to 226.3 m3/
type, 39 percent of the area sampled had no snags. ha) (Table 2), the maximum value on a plot was 30,625.9
ft3/ac (2,142.9 m3/ha). This compares to a range of 4,416
 600 Wildlife–Habitat Relationships in Oregon and Washington

It is clear that wilderness and parks generally provide

substantial standing dead wood habitat, but this provides
for wildlife associated more with montane through alpine
environments than with lower elevation environments.
This would favor such species as black-backed and
northern three-toed woodpeckers. The apparent dearth
of large snags in Ponderosa pine may mean lower
suitability for the 54 wildlife species associated with large
snags (20+ in or 51+ cm dbh) in that wildlife habitat.
Intensive forest management activities that have
decreased the density of large snags in early forest
successional stages (sapling/pole and small tree stages)
may have had adverse impacts on the 61 associated
wildlife species (Figure 12). Similarly, the lesser amount
Figure 11. Density of snags >=10.0 in DBH, decay classes of large down wood in early forest successional stages may
1-5, and >=6.6 ft tall across plots in the conifer alliance of not provide as well for the 24 associated wildlife species.
westside conifer-hardwood forest, Oregon and Washington, Such results suggest the continuing need for specific
displayed as a percent of the sampled area. management guidelines to provide large standing and
down dead wood in all successional stages.
to 20,306 ft3/ac (309 to 1,421 m3/ha) in various studies in Ecological Processes
westside Douglas-fir-western hemlock summarized by Natural patterns of wood distribution and legacy retention
Harmon et al.,164 and to 2,115 to 4,473 ft3/ac (148 to 313 are characterized by high variability through space and
m3/ha) reported by Spies et al.346 The large snag densities time. The lack of data on historical conditions of snag and
in westside conifer-hardwood forest (Table 1) were down wood abundance, combined with highly altered
substantially less than those reported by Spies et al.;346 the disturbance regimes, limits conclusions that may be drawn
estimate of 0.9 large snags/ac (2.1 large snags/ha) in early regarding temporal changes in decaying wood across the
stages probably represent stands younger than the 40 yr region.164, 178 The interpretation of inventory data will be
minimum sampled by Spies et al.;346 the estimate for improved in future years only as more detailed
middle-successional stages of 2.4/ac (6.0/ha) compares background information on stand conditions and
to 10.9/ac (27/ha) in their young stands; and the estimate disturbance histories becomes available. Until then, it may
for late stages of 5.70/ac (14.3/ha) compares to their be more productive for managers to assess future trends
mature 6.48/ac (16/ha) and old growth 10.9/ac (27/ha) in dead wood abundance and composition using new
classes. models that evaluate effects of specific management
Estimates of dead wood presented by Ohmann and practices on wood habitat functions, such as DecAID.223,
Waddell276 are not directly comparable to those reported 226
To assist with the assessment process, the following
in most wildlife studies.226 Wildlife studies typically section examines evidence regarding impacts of past
describe dead wood around nest sites, where dead wood silvicultural practices on wood dynamics in the Pacific
abundance may be substantially higher than in Northwest.
surrounding stands because many wildlife species select During the past century, management practices in the
nest sites within clumps of snags.226 Limited evidence Douglas-fir region have favored even-aged stands,
suggests that dead wood is most often distributed although there was some selective cutting prior to the
randomly within stands, but sometimes is clumped,82, 215 1950s (see reviews94, 365) Clear-cut harvesting on short
(Marcot et al. in prep., unpublished data). Twenty-five rotations was the dominant silvicultural system,
percent of stands sampled in the Oregon Coast Range were depending on site productivity, market preferences, and
found to contain patches of 5-10 trees that died accessibility. To facilitate site preparation, reserved trees
simultaneously.82 were usually absent and standing snags were felled and
slash was burned. Management priorities emphasized a
Management Considerations shift in stand composition from species mixtures to
monocultures of the most valuable and fast-growing
Management Ramifications of Snag and species, such as Douglas-fir. The economic rotation age
Down Wood Abundance varied from 40 years to over 100 years.59 In addition to
fire suppression, forest management east of the Cascade
Wildlife Species crest emphasized even-aged silviculture via clear-cutting,
By querying the CD-ROM matrixes, some generalizations and uneven-aged silviculture using selective removal of
can be drawn about the implications to wildlife of the overstory dominants.59, 137 Piling and burning of slash was
current status of decaying wood as detailed above. common in even-aged management regimes. These
Unfortunately, such inventory data are not available for silvicultural practices clearly altered the abundance and
historic conditions, so little can be quantified about recruitment of large down wood and snags in managed
potential changes in wildlife communities.
 CHAPTER 24: WOOD LEGACIES 601

forests of the Pacific Northwest,1, 2, 5, 59, 94, 112, 122, 130, 154, 198, 228, 230, sediment input to streams, and increase nutrient losses in
261, 262, 275, 276, 279, 302, 330, 344, 346, 357, 376
including: litter.164, 358, 359, 360, 361 Some data support a linkage between
1. Lower abundance of large diameter snags and down intensive management (especially depletion of decaying
wood “legacies” in managed forests (and streams); wood) and reduced forest biomass productivity,
e.g. lack of the U-shaped pattern; higher particularly on less productive sites. Lower productivity
accumulation of smaller-diameter fuels in eastside is attributed to nutrient losses from managed forests,
forests. reduced nutrient availability in older stands, and
2. Reduced recruitment and retention of large trees to decreased nutrient storage, particularly in the soil.272, 383, 384
provide future “legacies” Depletion of soil organic matter has been cited as a
3. Shorter mean residence time for down wood (i.e. primary factor contributing to declining forest
faster decomposition as a function of reduced log productivity and biodiversity in the Pacific Northwest and
diameter). elsewhere.17, 137, 198, 199, 228, 292, 293, 298, 299
4. Altered species composition of forests (westside: more One study of a western hemlock stand in Oregon
Douglas-fir, less western red cedar; eastside: less determined that after 150 years of intensive utilization and
pine, more true fir species). short rotations (Forcyte-10 model), reduced timber yields
were linked to depleted soil organic matter and lower
Depletion of Large Wood. The loss of large wood
nutrient availability.319 Such imbalances frequently reduce
structures has numerous potential impacts on ecological
tree growth in early stages, and impair ecosystem
functions of forests, although available information is
resilience to disturbance in later stages.9, 291, 292, 293 In contrast,
inadequate for a definitive assessment. The lack of large
Curtis et al.94 review evidence that forest management does
logs on steep slopes can decrease water percolation into
not degrade site productivity in moist temperate forests.
soil, impair slope stability, accelerate soil erosion and
Management effects on site productivity are likely to be
highly variable, depending on site-specific attributes of
climate, soil, and vegetation.
Small Wood Accumulation. In eastside forests subject to
fire suppression, large accumulations of small wood fuels
have been shown to improve site productivity due to
increased soil organic matter and nutrient retention. These
nutritional benefits to tree growth may be short-lived,
however, as the forests are predisposed to high intensity
fires that consume wood and accelerate nutrient losses.137,
170

Riparian Forests. Human alteration of riparian and

aquatic ecosystems in the Pacific Northwest over the past
century has proceeded with limited understanding of the
ecosystems and the effects of human disturbances. The
few studies that have examined streams as ecological
systems (see review in Naiman et al.262) highlight the
spatial and temporal complexity of riparian ecosystems,
and the importance of connectivity to diverse ecosystem
functioning. A balanced ecosystem is one in which
biodiversity, productivity, biogeochemical cycles are in
balance with the geological and climatic conditions of the
region.191, 192 The delivery of woody material to stream
channels has been identified as a key process determining
the ecological functioning of watersheds in the Pacific
Northwest coastal ecoregion.33, 40, 260, 262 This statement may
be extended to most areas west of the Cascade crest in
Oregon and Washington. Far-reaching effects of the
absence of large wood structures in streams include: 1)
simplification of channel morphology, 2) increased bank
erosion, 3) increased sediment export and decreased
nutrient retention, 4) loss of habitats associated with
diversity in cover, hydrologic patterns, and sediment
Figure 12. Management practices of the past have simplified retention.33, 144, 262 In coastal environments and estuaries,
forest structure. Note the lack of snags in this regenerating the loss of large wood may disrupt trophic webs and alter
forest on the right side of this hillslope near Satsop, coastal sediment dynamics.233
Washington. Photo: Deborah Lindley.
 602 Wildlife–Habitat Relationships in Oregon and Washington

Other Management Effects. In forests throughout the • Setting a goal of 40% of habitat capability for primary
region, multiple stand entries and stump residues have excavators, mainly woodpeckers,369 is likely to be
spread diseases, particularly root pathogens. Root diseases insufficient for maintaining viable populations.
initiate changes in stand composition, structure, and • Numbers and sizes (dbh) of snags used and selected
growth that alter nutrient and biomass dynamics of by secondary cavity-nesters often exceed those of
ecosystems.137, 220, 237 Root diseases also predispose trees to primary cavity excavators.
windthrow. Injury to residual trees during partial cutting • Clumping of snags and down wood may be a natural
contributes to the spread of stem decays that influence pattern, and clumps may be selected by some
input rates of dead wood and provide wood structures species, so that providing only even distributions
for wildlife habitat. Silvicultural practices that produce may be insufficient to meet all species needs.
multi-layered stands of susceptible species may also affect • Other forms of decaying wood, including hollow
ecosystem processes by favoring the spread of mistletoe. trees, natural tree cavities, peeling bark, and dead
In general, it is important to recognize that stresses parts of live trees, as well as fungi and mistletoe
imposed by management have the potential to alter the associated with wood decay, all provide resources for
abundance and dynamics of decaying wood by altering wildlife, and should be considered along with snags
ecosystem energy and nutrient cycles, stand and down wood in management guidelines.
microclimates, and forest susceptibility to disturbance. • The ecological roles played by wildlife associated with
decaying wood extend well beyond those structures
Lessons Learned During the Last Fifteen Years per se, and can be significant factors influencing
What new developments have ensued since publication community diversity and ecosystem processes.
of Thomas366 and Brown 48 in our applied knowledge of We have also learned that managing forests with decay
wildlife-wood decay relations? Much basic research has processes should be done as part of a broader management
been conducted on terrestrial vertebrates associated with approach to stand development, with attention paid to
snags, although much remains to be learned of wildlife retaining legacies of large trees and decaying wood from
associated with the other forms of decaying wood. Major original or prior stands. Further lessons have been learned
research initiatives such as the Demonstration of in the area of technical and operational developments;
Ecosystem Management Options (DEMO) study by USDA some of these are discussed below.
Forest Service is helping to quantify relations of wildlife
presence to down wood. Nothing can replace field zoology Habitat Assessment
and ecology in providing such basic understanding of Regional summaries of the current abundance of snags
wildlife relations with decaying wood. and down wood presented by Ohmann and Waddell276
Several models have been introduced that help track have several potential applications to forest management,
the demography of snags,60, 183, 221, 286, 306 and several other planning, and policy. One use is in broad-scale assessments
unpublished models provide further evidence. More of wildlife habitat. Managers and planners can compare
recently, limited capabilities for handling snag dynamics the inventory information on current dead wood resources
have been incorporated into the Forest Service’s Forest to characteristics selected and used by wildlife and other
Vegetation Simulator (FVS) model. Some of these models organisms in developing management guidelines for
track numbers of snags over time by dbh and decay class, federal lands, or in evaluating forest practice regulations
and variously incorporate creation, decay, and fall rates or incentive programs for state and private lands. Such
of snags. The only available down wood dynamics model guidelines currently are based on very limited scientific
to date is the Coarse Wood Dynamics Model for the data.
western Cascade Mountains of Oregon and Washington.247 Comparisons of inventory estimates to those reported
No model, however, had been advanced to replace the in most wildlife studies are complicated, however, by the
“biological potential” models263, 369 until the DecAID model fact that inventory estimates represent average conditions
was developed for this book188 and USDA Forest Service within a wildlife habitat type at the regional level rather
(see next section). Several major lessons have been learned than around nest sites (see earlier discussion), and few
in the period 1979-1999 that have tested critical wildlife studies report both values (nest site and stand
assumptions of these earlier management advisory average) for comparison.226 Furthermore, although the
models: analysis of inventory data presents data on dead wood
• Calculations of numbers of snags required by abundance, management actions at the local level may best
woodpeckers based on assessing their “biological be focused on the ecological processes that lead to
potential” (that is, summing numbers of snags used development of these forest structures rather than on the
per pair, accounting for unused snags, and abundance of structures themselves. Management
extrapolating snag numbers based on population decisions also may require information on the spatial
density) is a flawed technique. Empirical studies are distribution (landscape pattern) of dead wood, which
suggesting that snag numbers in areas used and cannot be estimated from sample-based inventories.
selected by some wildlife species are far higher than Information on regional patterns of dead wood is
those calculated by this technique.226 currently being incorporated into the DecAID model248
depicting the range of natural conditions. This information
 CHAPTER 24: WOOD LEGACIES 603

is intended to help guide managers in considering dead of a given size and decay class may be the identified
wood and processes of decomposition in forest objective. The functional objectives should be determined
management. The regional inventory database contains to be within the productive capability of the ecosystem.
information on occurrence of pathogens such as stem Productive capability may be evaluated by examining
decays and root diseases that contribute dead wood. In information on primary productivity for the ecoregion and
addition, the data contain new information about the forest type (described earlier) and if available, data
range of variability in dead wood—both historically and summaries on the historical range of natural conditions
in the current landscape. The range of variability in dead of dead wood status.
wood abundance among plots in the region can help guide Wood status may include various descriptors of the
management decisions regarding the desired distribution amount (volume or weight), size class, decay class, and
of dead wood within a large landscape or watershed being perhaps species of wood structures. Current inventory
managed, but not spatial variation within stands. methods for quantifying the status of dead wood,
Caution must be exercised in using the regional plot including plot design and sampling techniques, have been
data to describe the historical range of conditions in dead reviewed. 56, 71, 164, 165, 276, 285 Field inventories should be
wood. The regional plot data sample only current conducted at the scale at which management guidelines
conditions and lack data on site history, as discussed are intended to be developed and implemented, such as
earlier. Even if plots in “natural” forest could be identified, among watersheds within basins.
current levels of dead wood have been altered to an
Incorporate Wood Dynamics Into Planning. The
unknown degree by fire suppression and other human
extended residence time for dead wood legacies
influences. On the eastside in particular, current levels of
necessitates that snag and log dynamics models be
dead wood may be elevated above historical conditions
incorporated into the process of long-term management
due to fire suppression and increased mortality, and may
planning. New and existing wood dynamics models are
be depleted below historical levels in local areas burned
reviewed below. It is important to remember that such
by intense fire or subjected to repeated salvage and
models typically operate at the stand level, whereas
firewood cutting. Plot data from natural forests on the
management guidelines for snags and down wood are
westside, where fire return intervals are longer,5 may
directed at broader scales. Thus, model output should be
provide a reasonable approximation of historical
averaged across stands within watersheds, or
conditions.
management guidelines should address stand- (meaning
Template for a Stepwise Assessment Process successional stage) specific conditions.
The following template presents a stepwise process for Monitor Management Effects on Wood Dynamics. Due
habitat assessment and management planning to achieve to the general lack of information guiding management
objectives for down wood and snags. These of decaying wood, wood dynamics and associated wildlife
recommendations are based on existing knowledge of response is a priority topic for monitoring and adaptive
wood dynamics and decay processes in ecosystems, and ecosystem management.149 Approaches for monitoring
insights developed by the inventory analysis of Ohmann natural resources and for modifying management
and Waddell. 276 This process also can be applied in activities must be carefully planned and evaluated, with
managing for other elements of forms of decaying wood consideration of appropriate landscape scale.
or wood legacies.
How To Set Quantitative Objectives Using DecAID
Set Quantitative Objectives. To develop effective
How should the manager determine appropriate
management plans for down wood and snags, it is
objectives for managing decaying wood? This chapter is
imperative to evaluate: 1) land ownership and desired
not intended to prescribe standards for wood decay
biodiversity, in terms of wildlife species to be supported,
management. Instead, we offer the following set of
2) specific land allocation, 3) site capability and history,
questions that the land manager could use to develop
and 4) management scale. Consideration of these factors
overall objectives and specific guidelines for wood decay
assists in identifying a range of target values as guideposts
management.
for management activities.
What is the land ownership and overall expectation for
Evaluate Current and Historical Status of Decaying
biodiversity conservation? It may be socially acceptable
Wood. Information on the current and historical status of
to have different levels of expectation for conservation and
dead wood is helpful in determining how the existing
maintenance of biological diversity, including decaying
patterns in abundance, composition, and distribution of
wood and associated wildlife, on different land
down wood and snags relate to objectives identified for
ownerships and land use allocations. Land ownerships
the area of interest. If detailed data on the current and
are subject to different sets of laws and regulations
historical range of natural conditions is lacking (which is
regarding biodiversity and habitat conservation. For
likely), it may be preferable to substitute functional target
example, commercial forestry landowners may be
values for specific wildlife species. For example, to provide
regulated by the appropriate state forest practices act, as
maximum habitat elements for specific cavity-nesting
well as any habitat conservation plans, but are not subject
species, a designated quantity and distribution of snags
to the more stringent wildlife habitat regulations found
 604 Wildlife–Habitat Relationships in Oregon and Washington

in the National Forest Management Act that regulate ecosystem functions associated with particular amounts
national forests. Consequently, commercial tree farms and and sizes of snags and down wood and other decaying
commercial forestry lands may call for conservation of wood, in specific wildlife habitats, over time, and at several
decaying wood and associated wildlife at moderate statistical levels. Using such tools, the manager may
statistical levels presented in the cumulative species curves consider alternative scenarios for retention, recruitment,
of the DecAID advisory model, whereas some Federal and restoration of decaying wood for wildlife and
lands such as national forests and national parks may ecosystem function. Uncertainty in wildlife response,
strive for the high statistical levels. expected levels of decaying wood on site, likelihoods of
fire, insect, and disease conditions, and future
What is the specific land use allocation? For a given land
disturbances, may factor into specific management
ownership, specific land use allocations may guide
guidelines, as well as into monitoring programs to help
objectives for the abundance of decaying wood. For
evaluate efficacy of the guidelines and objectives.
example, on commercial forestry lands, intensive timber-
use zones on upland slopes may provide for lower levels How To Evaluate The Current And Historical
of decaying wood than in riparian buffer zones, and on Status of Decaying Wood
some Federal lands, old-forest conservation areas (such To determine a quantitative objective for dead wood, it is
as Managed Late-Successional Reserves) may warrant necessary to evaluate the functional target for wood
even higher levels. relative to the historic range of natural consditions and
What is the site capable of producing overall and given current status of dead wood in the area of interest. A new
its particular site history? For a given site, such as a modeling tool named DecAID is available to assist with
particular forest stand, the manager may wish to this task. DecAID (as in “decayed” or “decay aid”) is a
determine what the site is capable of producing in terms new Decayed Wood Advisory Model being developed to
of sizes and densities of decaying wood structures such address some of the recent lessons learned.226, 247 DecAID
as snags, hollow live trees, and down wood. Then, is based on a thorough review of literature, available
considering current stand conditions and recent site research and inventory data, and expert judgement. It
history, the manager can craft reasonable management broadens the paradigm for wildlife species and habitat
direction to provide for appropriate levels of decaying assessment by considering the key ecological functions of
wood as integrated into other site management activities wildlife (see below) as well as the ecosystem context of
such as tree pruning, precommercial thinning, final tree wood decay in terms of secondary effects on forest
harvests, etc. productivity, fire, pest insects, and diseases.
DecAID has four components:
What is the appropriate spatial scale for managing
decaying wood? The tools discussed in this chapter (1) Species-habitat associations, specifically, the use and
pertain to helping prescribe or predict effects of the selection by terrestrial wildlife species of snag dbh, snag
abundance of decaying wood at the scale watersheds or density (number per unit area), down wood diameter, and
larger, rather than for particular sites such as individual down wood percent cover (percent of forest floor covered
forest stands. However, this does not mean that attending by down wood or wood litter). This information is based
to individual stands is unimportant. Studies suggest that on a synthesis of empirical data, and interpretations made
wood habitat structures function best for wildlife when by the authors and experts.226 DecAID presents cumulative
they are broadly distributed as well as occurring in locally- species curves for combinations of wildlife habitats and
dense clumps, such as with scattered snag or down wood habitat structures, showing the size and density of snags
patches. The manager might use some of the concepts, (or diameter and percent cover of down wood)
suggestions, and tools in this chapter at stand levels but corresponding with each species as reported in field
would need to validate them at broader geographic scales. studies. Where data permit, these curves are shown for
Even broader geographic areas such as subbasins might low (30% tolerance interval), moderate (mean), and high
need to be evaluated when considering or projecting (80% tolerance interval) statistical levels. Some studies
natural and human-caused disturbance regimes. pertain to snag (or down wood) densities averaged across
Disturbances can radically change local conditions. study stands, whereas others pertain to locally high
Borrowing from landscape ecology, one rule of thumb that densities in snag (or down wood) clumps centered on
the manager might follow is to evaluate, as analysis units, nest/breeding, den/roost, or foraging sites.
a land area large enough to fully encompass the occurrence (2) Key ecological functions of wildlife species associated
and full extent of the infrequent (e.g., 50-year) disturbance with decaying wood, and full lists of wildlife species
event such as a major wildfire or harvest schedule rotation associated with decaying wood. This information is based on
period. Planning should be targeted for average levels of the species functions database of the matrices on the CD-
decaying wood across the analysis unit realizing that ROM with additional information from the literature and
individual sites within the unit might be subject to greater other databases.223, 224
uncertainty of specific conditions at various times.
The manager can use tools such as the DecAID model (3) Range of natural conditions. This information is an
and matrixes on the CD-ROM to perform a risk analysis analysis of current and recent past inventory data on snags
of the likelihood of providing for wildlife species and and fallen trees.276 The inventory data are taken from
 CHAPTER 24: WOOD LEGACIES 605

several sources, including the Forest Inventory and DecAID also ties into the CD-ROM matrix on wildlife
Analysis of USDA Forest Service, Pacific Northwest species and their key ecological functions. The 30 bird and
Research Station; the Current Vegetation Survey of USDA 10 mammal species associated with snags in Upland
Forest Service, National Forest System; and the Natural Aspen Forest collectively perform some 60 categories of
Resource Inventory of USDI Bureau of Land Management. key ecological functions. These include primary
Collectively, these inventory data cover all national forests consumers or herbivores (16 species), secondary
and BLM lands, and all other non-reserved lands. Use of consumers or primary carnivores (38 species), and tertiary
inventory data is limited for eastside, fire-dominated consumers or secondary carnivores (3 species). Other
systems. ecological functions performed by this set of species
include potential control of insect populations (10 species)
(4) Ecosystem processes and productivity. This
or vertebrate populations (7 species); dispersal of fungi,
information is based on a synthesis of publications and
lichens, insects and other invertebrates, seeds, fruits, or
research identifying how wood decay provides for
vascular plants (11 species); creating sapwells (2 species),
ecosystem productivity.
creating nesting structures (4 species), excavating cavities
The manager will be able to use DecAID for advice on
(11 species), digging burrows (3 species), and creating
the following topics by first specifying wildlife habitat,
terrestrial runways (2 species) potentially used by other
structural stage, and statistical (confidence) level: 1)
species, some of which are also associated with snags in
wildlife species associated with particular sizes and
Upland Aspen Forest. Other ecological functions of snag-
densities of snags and down wood, or, conversely, the sizes
associated wildlife in Upland Aspen Forest are potential
and densities required to meet specified wildlife
improvement of soil structure and aeration through
management objectives, at three levels of confidence; 2)
digging and tunneling (2 species), physically fragmenting
the array of key ecological functions of wildlife associated
standing or down wood (2 species), creating snags (1
with decaying wood; 3) the recent-historic and current
species), and potentially altering vegetation structure or
range of natural conditions of snags and fallen trees; 4)
succession through herbivory (1 species). Using the CD-
advice on fire risk assessment and mitigation; 5) advice
ROM, the manager also can determine the specific
on the roles of insects and diseases associated with various
ecological roles of just those species expected to be
amounts of decaying wood; 6) and the influence of the
provided at given sizes and amounts of snags and down
abundance of decaying wood on ecosystem processes and
wood, and trade-offs with other functions. Overall,
productivity.
through such lists, DecAID provides the manager with
As an example of using Component 1 and 2 of DecAID,
insights into the array of ecological roles performed by
a manager might be interested in the role of snags in the
wildlife species associated with decaying wood structures
Upland Aspen Forest wildlife habitat type. A query of the
in specific wildlife habitats. The model also indicates how
CD-ROM portion of DecAID reveals that 30 bird and 10
those functions might respond to changing attributes of
mammal species are coded as being associated with snags
decaying wood. Thus, the model offers an indication of
in this forest type. DecAID has compiled all available
how different management practices affect the
empirical data on aspen-related species, which totals 9
relative “functionality” of ecosystems.
birds (8 primary and 1 secondary cavity-nester) for snag
density data, and 23 bird (16 primary and 6 secondary How To Incorporate Wood Dynamics Models Into
cavity-nesters, the primaries including 4 sapsucker Planning
hybrids) and 1 mammal species for snag dbh data. For Once an appropriate objective is determined with
Upland Aspen Forest, the cumulative species curves quantitative targets, the next step is to integrate wood
suggest that providing for all nesting or breeding species dynamics into the planning process. This is a critical step
as found in field studies entails locally dense clumps of because of the extended time frames required to replace
snags averaging 32.4 snags per acre larger than 33 in (80 large wood structures. Dynamics models predict the
snags per ha) with at least some snags 20.3 in (52 cm) or general rate of fall of snags and decay of snag and logs on
greater in dbh. a site to determine the amount of decaying wood occurring
If the manager had a known or expected condition through time (e.g. a rotation). These models can also be
different from this, the expected associated species could used to estimate the number of green replacement trees
also be determined. An example is an Upland Aspen Forest to leave on a site and when to convert them to snags or
stand with locally dense snag clumps averaging 20.3 snags logs. Mortality estimates from growth and yield models
per ac (60 snags per ha) of 15.7 in (40 cm) dbh. In this case, (Forest Vegetation Simulator, Organon, DFSIM, etc.) can
the snag density corresponds to providing for 6 of the 9 be input into all the models reviewed here to track snag
studied bird species, and the snag dbh corresponds to and log recruitment in forest stands. The following are
providing for all of the 21 species (100%) at low statistical three dynamics models available for Washington and
levels, 14 species (67%) at average, and 6 species (35%) at Oregon.
high statistical levels; DecAID also lists the specific species
in each case. The manager can use the information on Snag Recruitment Simulator (SRS)—Marcot. 221 SRS is
statistical levels as a risk analysis and also as a means of based on the snag life-table approach.263 The model was
setting objectives, where the high statistical level might built to generalize snag decay and falling rates, and can
correspond with highly conservatory objectives such as be parameterized with any such rates. Data limitations,
for Late-Successional Reserves.
 606 Wildlife–Habitat Relationships in Oregon and Washington

however, necessitated that the westside version of the required to meet these new objectives, particularly the goal
model (i.e., Oregon and Washington west of the Cascades to produce and retain legacies of decaying wood.122, 228, 292
crest) predict fall and decay of Douglas-fir snags based Management goals for dead wood, thus, are inextricably
on data from the Oregon Coast Range,82 and the eastside embedded within the broader goals for diverse forest
version predict fall rates of ponderosa pine snags from resources. To accomodate multiple objectives, Franklin122
the Blue Mountains of Oregon and decay rates from the has suggested that new management should do the
Coast Range Douglas-fir data. SRS consists of a series of following:
compiled Lotus spreadsheets and runs in DOS on a PC. It 1. Emphasize adding new silvicultural tools to the old.
is the easiest of the three models to use. Several people 2. Address stand and landscape-level objectives.
have parameterized SRS for specific geographic areas. A 3. Maintain stands that are structurally and
similar model based on a Leslie Matrix was later compositionally diverse (practices will differ from
published.253 stand to stand, depending on forest type, condition,
environment and specific objectives).
Snag Dynamics Projection Model (SDPM)—McComb
New silvicultural practices are a high priority in
and Ohmann.240 SDPM uses logistic regression analysis
regenerating stands where one or more harvest cycles have
to predict the probability of a snag falling over a 10-year
simplified stand structure and composition and removed
period. A straight rate is used for the probability of a snag
wood legacies. In young stands, Franklin122 recommends
changing from hard to soft over the same period. Forest
that management should:
Inventory and Analysis (FIA) remeasurement data from
1. Aggressively create stands of mixed composition to
western Washington was used to develop the model.
maintain habitat for a broad array of species (and to
SDPM models Douglas-fir, western hemlock and western
achieve diversity in quality and timing of nutrient
redcedar snags. SDPM incorporates site factors such as
inputs to streams).
climate, slope, and aspect which affect snag fall and decay
2. Delay the process of early canopy closure (wide
rates. The model is an executable DOS-based program
spacings, pre-commercial thinning etc.).
written in C language.
3. Provide for adequate amounts and a continuous
Coarse Wood Dynamics Model (CWDM)—Mellen and supply of large wood, including snags and down
Ager, 246 Mellen and Ager, 247 Mellen et al. 248 CWDM logs, for maintaining structural divesity in forests
analyzes the dynamics of both snags and down logs in and streams and maintaining all other ecosystem
forests on the westside of the Cascade Mountains. The processes associated with wood.
model assesses snag fall, height loss and decay, and log The basic theme of these revisions of intensive forestry
decay for Douglas-fir and western hemlock. Snag fall and practices is to retain the higher levels of complexity found
height loss rates are from FIA remeasurement data from in natural forests, and in so doing, to protect processes
western Oregon and Washington. Decay rates are based and structures that retain future options for ecosystem
on work in the western Oregon Cascades.140 A single- management. Effective management of decaying wood
exponential equation164 was used to model the dynamics must do more than simply provide for inputs of dead trees.
of snags and logs. CWDM is available in both DOS and Rather, management should strive to provide for diversity
Windows versions. An east-side version of CWDM will of tree species and size classes, in various stages of decay
be developed as data on decay rates of snags and logs in and in different locations and orientations within the stand
eastern Washington and Oregon become available. and landscape.122, 233 Examples of new practices include
retention of large woody structures at the time of harvest,
Management Tools and Opportunities long rotations, and creation of snags and logs from green
trees. An overview of different silvicultural practices to
Silvicultural Methods regain (i.e. retain and create) wood legacies in managed
Traditional forestry practices in the Pacific Northwest94 stands is provided below. Detailed discussions of
have been based on the simplification and homogenization alternative silvicultural practices are available. 94, 132
of forests to achieve economical wood fiber production.122, Through the integrated application of silvicultural
228, 289
As a result, the amount of old-growth forests has practices described below, it becomes possible to conserve
declined over 50% in the last 60 years,41 and remaining biodiversity in managed forests by retaining legacy
stands are highly fragmented.347 Such practices have been structures and accelerating the development of structural
at the center of intense controversy over the past three complexity.61, 69a
decades.271, 272 Concerns center on the loss of biological
“legacies” resulting from natural disturbances—surviving Structural Retention. Structural retention refers to any
organisms and wood structures that contribute to resilient practice that retains significant structural elements from
ecosystems with structural, compositional, and functional a harvested stand for incorporation into a new stand. The
diversity.291 focus of this practice has been on harvesting in old and
New management paradigms, originally coined “New mature stands. Objectives in structural retention include
Forestry,”121 have shifted the emphasis from maximized maintenance of organisms and processes on harvested
fiber production over the short-term to diverse forest areas, structural enrichment of the regenerating forest, and
values and maintenance of long-term forest productivity enhanced connectivity across managed landscapes. To a
(see also 201). Alternative forest management systems are limited degree, structural retention mimics the effects of
 CHAPTER 24: WOOD LEGACIES 607

variable disturbance patterns to create legacy structures. in soil, stem, and canopy habitats of old-growth forests
Retention of wood structures from a harvested stand for do not disperse well, and thus, do not readily recolonize
carryover into a new stand offers an infinite array of clear-cut areas.207, 326 The same concept holds for many
alternatives to clearcutting and other regeneration harvest mycorrhizae-forming fungal species.293 Added benefits of
systems.132, 364 Variables in structural retention include the green tree retention include moderated microclimates of
type, size, number of structures, and spatial distribution the cutover area, which may increase seedling survival,
of the retention. Large, old, and decadent trees, standing reduce additional losses of biodiversity on stressed sites,293
dead trees (snags), and down wood are examples of and facilitate movement of organisms through cutover
structures selected for retention (Figure 13). patches of the landscape. Green trees retained across
Retention of snags and large logs is a particularly harvest cycles can also be used to grow very large trees
effective practice in maintaining large wood when for either ecologic or economic goals. This may be an
harvesting stands that already contain significant wood especially valuable practice in providing large wood to
legacies—such as young and mature stands of natural riparian forests subject to harvest: green tree retention can
origin and old-growth forests. Practices including the be implemented to favor species such as Douglas-fir and
removal or burning of unmerchantable trees and down redcedar that produce larger and more persistent wood.
wood in forests and near stream channels should be Other benefits of green tree retention include reduced
minimized or eliminated. Retention of snags provides hazards of landslides via maintenance of root strength and
numerous habitat benefits.154, 239, 402 However, safety and reduced potential for rain-on-snow flood events.
liability issues associated with snag retention have posed Green tree retention offers many benefits to wildlife.
an operational barrier to management objectives for For example, the higher structural diversity in young
structural retention. Two approaches useful in reducing stands that contain legacy trees from previous stands
hazards associated with snags are: 1) to cluster snags in provides much improved habitat values to late
patches rather than wide dispersal, and 2) to create snags successional species such as the northern spotted owl, as
from green trees after cutting.122 well as other vertebrates that use late-successional stands
Many questions remain regarding the most desirable for some elements of their life history.69, 122, 314 Such stands
density and spatial distribution of snags and down wood may provide wildlife habitat as early as age 70-80 years
for wildlife.94 New information suggests that snags and rather than 200-300 years, the approximate time interval
down wood may follow a naturally “clumped” required for old-growth conditions to develop after
distribution. Therefore, managers can take opportunistic secondary succession. Green tree retention on a harvest
advantage of site-specific occurrences of snags and down cycle of 120 years has been proposed as a method to
wood without having to match a particular spatial provide habitat for late successinal species in only 40-50
distribution pattern of clumps. This offers managers broad years.278 Different scenarios for green tree retention have
flexibility to provide varying local densities of snags and been offered.122, 278 To meet needs of all species likely to
down wood across the ground, within and among stands. occur in area, it is critical to identify tree species for
Managers must also consider the temporal dimension to retention. Updates on wildlife and vegetation response to
decaying wood, to ensure that sufficient sufficient snag
and down wood densities are provided through time.
Franklin et al.132 review information on the type, amount,
and distribution of structural retention for various Figure 13.
objectives. Additional examples of wildlife benefits from Retention of
structural retention have been described.20, 21, 63, 64, 69, 152, 154, old-growth
210, 314, 326
“legacy”snags
in a commercial
Live (Green) Tree Retention. Retention of living trees on thinning unit
cutover areas is one form of structural retention that can in western
provide for future recruitment of snags and down wood Washington.
(created artificially or through natural processes).122 Other Photo: Deborah
terms commonly used for this method include partial Lindley.
cutting or partial retention to distinguish it from selection
cutting. Green tree retention involves reserving a
significant percentage (10-40%) of living trees, including
dominants through the next rotation. The density,
composition, condition, size classes, and spatial
distribution of the retained trees varies according to
management objectives, stand and site conditions, and
other constraints. The objective is to maintain a more
structurally diverse stand than could be achieved through
even-aged management.
Green trees function as a refugium of biodiversity in
forests. For example, many species of invertebrate fauna
 608 Wildlife–Habitat Relationships in Oregon and Washington

green tree retention are recently available for the DEMO the rotation cycle. Rather, retention of snags at harvest,
program in Oregon and Washington, and other combined with long rotations would be necessary to
silvicultural trials.20, 30, 152, 155, 156, 210, 268, 311, 406 provide for present and future recruitment of such snags.
Long rotations are unlikely to provide forests with
Variable Retention Harvest Systems. Franklin et al.132
structural, functional, and compositional features
recommended the term “variable retention harvest
comparable to late-successional reserves.12, 132, 156
systems” to refer to harvesting practices that allow a
continuous spectrum of removal and retention in mature Thinning. Thinning in plantations can be used to decrease
and older stands, depending on objectives. The the time required to develop larger trees and multiple-
development and maintenance of structurally complex canopy forests for species associated with late-successional
managed forests is the primary rationale for retaining forests, while producing ecomonic benefits from thinnings
structural elements of the harvested stand. Retention of and shorter rotations.94, 364 It may not produce desired
various decaying wood structures through variable amounts of down wood or heavy-limbed tree crowns.
retention harvest provide many benefits to many wildlife However, benefits of accelerating the rate of stand
species and functions,20, 26, 64, 70, 73, 156, 326 as well as other forest development must be weighed against the detrimental
resource commodities.12 effects of logging disturbances on species, particularly
More recently, there has been increasing interest in those with limited dispersal capabilities. 343 Variable
aggregated or “patch” retention—the maintenance of density thinning is a variant of the traditional unifrom
small forest patches, instead of dispersion of retained spacing that shows promise for accelerating structural and
structures.64 These patches can provide refugia, while also compositional diversity.66 In general, thinning is beneficial
providing microclimatic gradients for more sensitive to the development of more structural diversity in young
species and functions.151 By providing for the maintenance stands, and to a variety of wildlife species, particularly in
of refugia, tree and patch retention may benefit species more complex and patchy stand structures, with legacy
limited by slow dispersal rates, rather than by particular structures.26, 70, 73, 150, 155
habitat structures.13, 239, 343 Franklin et al.132 review major
issues in developing harvest prescriptions based on the Restoration Techniques
variable retention harvest concept, including the type, Restoration with respect to decaying wood involves
amount, and distribution of structural retention for various silvicultural manipulations to develop and create wood
objectives. Franklin 122 described several alternative legacies in stands lacking suitable existing structures. In
management approaches for maintaining wood addition to retention of existing snags and down wood at
production and complex forest structure suggested by fire the time of harvest, and young stand management to favor
history research. In a multi-aged management strategy, structural complexity (retain green trees), intentional
selective cutting practices can be used to sustain complex methods have been developed and tested to create snags,
stand structure and composition for long periods. It may down wood, cavities and other habitat niches. 96, 132
be beneficial that the management system mimic the Techniques include girdling, injection with herbicide,
natural disturbance regime to the extent possible, so that topping, explosives, fungal innoculation, and use of
the site can accomodate future natural disturbances as they pheromones to attract beetles.25, 54, 57, 84, 212, 284 Various
interact with managed stands. techniques for creating artifical cavities have been tested,
including creation of cavities by den routing,63, 65 or by
Long Rotations. Long rotations involve the use of rotation cutting a hole with a chainsaw and covering with a
ages that are significantly longer than that defined by the faceplate.62
culmination of mean annual increment.91, 92, 93, 94 Long Many of these techniques have produced favorable
rotations provide for the recruitment of larger and more habitat for wildlife. However, characteristics of artificially
complex wood structures by allowing tree to grow to a created snags may not always be comparable to natural
larger size and by eliminating logging disturbances that snags. Additional research and monitoring is needed to
damage or remove wood legacies. When coupled with a better evaluate the attributes and habitat uses of snags
series of silvicultural treatments, long rotations can and down wood created by artificial methods. Costs for
produce complex managed forests, increase commodity creating artificial snags and down wood are important
yields, and address cumulative issues where too much of management considerations, thus it is desirable to retain
the landscape is in a recently-harvested condition. Long existing legacy structures, as well as trees for future snag
rotations also may be applied to patches of trees at a scale recruitment. In addition, it may be both biologically and
smaller than individal stands. As an added benefit, operationally beneficial to create and preserve snags
extended rotations can reduce the need for permanent within patches of uncut trees, rather than to distribute
transportation systems.196 them uniformly across stands.
Long rotations may not provide adequate protection Few long-term data are available to assess the
for all structural elements, processes, and organisms, effectiveness of different methods of snag creation. Results
particularly those most sensitive to disturbance, or with of a recent study328 monitoring natural and artificially
more rapid turnover rates. For example, large-diameter, created snags on the Siuslaw National Forest in the Oregon
moderately-decayed snags removed at the time of Coast Range provide useful guidance to managers. The
harvesting (via clearcutting) may not be replaced within study tracked longevity and wildife use in 150 green leave
 CHAPTER 24: WOOD LEGACIES 609

trees, 91 natural (Class 1) snags, 27 intentionally-topped

Douglas-fir trees, and 23 hardwood leave trees (big-leaf
maple and alder) and snags in variable retention harvest
units from 1987 to 1998 (Figure 14).
The study identified several interesting facts about
topped leave trees and residual snags:
1. Topped leave trees (blasted or cut) are far more
windfirm than natural snags or green leave trees
(Figures 15, 16). Greater windfirmness of topped
trees (0.7% rate of windthrow) compared to natural
snags (11%) or green leave trees (17%) is attributed to
both lower rates of root failure and stem breakage.
Windthrow of green leave trees can be substantial,
particularly in wind-prone areas. Numerous cavities
and other defects in the lower bole indicate that bole
Figure 14. Stand treated to restore wildlife snags. Treatments
breakage could increase in the future. Thus, long- included topping with and without retention of live branch
term cavity habitat requires periodic topping, rather whorls. Photo: Barry Schreiber.
than a single entry, especially for small-diameter
trees that decay more rapidly.
Topped trees present fewer conflicts with harvest and
2. Live leave trees experience high rates of windthrow
silvicultural activities than snags, and if trees are topped
and breakage due to increased exposure in clearcuts.
prior to harvest, some of the tops may be salvaged to offset
Thus, live green trees may not provide for snag
the cost of topping. In planning for the desired density of
recruitment for an extended period post-harvest.
leave trees, managers need to consider the vulnerability
3. Degradation and loss of natural snags is high due to
of green leave trees to wind and fire, and degree of
root throw, but more significantly to bole breakage.
competition to living topped trees by the regenerating
Breakage is most frequent near cavities, areas of
forest. Thinning may be desirable to extend the lifespan
advanced decay, and structural deformities. The
of green trees or live-topped trees. Tree defects will affect
longevity of Class 1 snags is limited. Continued
the quality and longevity of leave trees as snag habitat.
recruitment of new snags requires snags to be created
For example, bole crooks are weak points that can
periodically from green trees during the course of
determine where stem breakage will occur, hence this
stand development.
feature can be selected to provide future snag habitat of
4. Trees topped above two branch whorls survive and
desired height. Trees with butt rot, hollow stems, large
develop new tops. Continued diameter growth in
bole sweeps, forks at the lower bole, or leaning trees are
these trees provide higher values as wildlife snags.
less suitable for retention, as they are less stable, and
Large crooks formed in these trees also provide
succumb more rapidly to wind and gravity. Live-topping
platform nest sites and create future breaking points
of green trees has been found to have great potential to
to form a tall snag. The greater longevity of these
provide long-term cavity habitat in managed stands. This
live-topped trees should reduce the need to cause
possibility should be more thoroughly investigated.328
intentional mortality in leave trees in the future.
Additional considerations for active management of
5. Methods for topping trees by either blasting or
decaying wood are available in a recent review.69a
chainsawing produce similar results for both snags
and live-topped trees. Blasting provides a more Management of Wood in Streams. In riparian areas, the
natural look, but the chainsaw method allows for development and maintenance of large trees is required
directional felling if the salvage of tops is planned. to provide inputs of large wood to streams and rivers33, 143
6. Natural and created snags show high levels of use by The forest adjacent to channels is particularly critical in
cavity-nesting birds. Topped trees rapidly develop small streams, since wood inputs from other sources may
cavities throughout the bole. Live-topped trees be negligible.241 Natural stable wood in streams should be
develop cavities ten years after topping, with cavities undisturbed 376 and supplementary wood recruited
forming first near the upper bole. The creation of live through management of riparian forests to the maximum
trees with cavity habitat is highly desirable, as it extent possible. The delivery and routing of wood to
allows cavity habitat to be maintained over longer streams is important to the vitality of watersheds and their
periods. component drainages in the Pacific Northwest.262 Integrity
7. Big-leaf maple has relatively high survival and of the lateral, longitudinal and vertical components, as
provides a high density of cavity sites. The rapid well as the temporal and spatial characteristics of wood
diameter growth of big-leaf maple allows a tree size cycling within basins must be maintained. Thus, managers
suitable for cavity production to be developed within must pay attention to geomorphic considerations and
thirty years. connectivity between parts of the larger system when
planning for inputs of large wood in riparian areas.
 610 Wildlife–Habitat Relationships in Oregon and Washington

adjacent harvest unit. Where harvesting is allowed, group

selection or single tree selection is preferred to clear-
cutting.3 Thinning should strive to leave trees dispersed
in irregular patches.

Summary of Management Recommendations

The information presented in this chapter emphasizes
several properties of decaying wood in forest ecosystems:
(1) each structure formed by decaying wood helps support
a different functional web in the ecosystem; (2) no one
decaying wood structure supports all functions equally;
and (3) all decaying wood habitats together support the
widest array of ecological functions and associated wildlife
species. The CD-ROM with this book in combination with
the DecAid model provides managers with a powerful
Figure 15. Snag created by blasting. Photo: Barry Schreiber. tool that makes it possible to assess the degree of “full
functionality” of ecosystems as supported by the various
decaying wood structures, and which functions are
strengthened, diminished, or lost through alternative
silvicultural management practices.
Lessons for managers are:
Figure 16. Snag created by 1. Examine forestry practices for how they influence the
topping with a saw. Photo: distribution and abundance of down wood and snags
Barry Schreiber. in relation to forest landscape patterns. In situations
where forest management objectives extend beyond wood
production to broader biological and human values,
intensive forestry practices by themselves may
inadequately maintain or restore biodiversity, especially
in early and late successional forest development phases.
Species, processes, and values associated with older stages
of stand development (transition and and shifting gap
stages)277 are likely impaired or absent from intensively
managed stands.343 Species and processes associated with
the early establishment phase also have shorter duration
than may occur naturally. This does not mean that
intensive forest management practices are incompatible
with multiple forest objectives at a landscape scale, but
Aquatic functions dependent upon connectivity rather that species and processes associated with early and
include: 1) requirements for habitat access during discrete late stages of forest development should be assessed over
life history stages of fish and wildlife, 304, 321 2) large areas such as landscapes, subregions, and regions.291,
343
synchronization of emergence and migration of aquatic Management for certain species must consider habitat
organisms to stream temperature,303, 362 3) regulation of requirements at different spatial and temporal scales.181 It
nutrient and material exchanges between forests and may then be possible to modify silvicultural practices at
streams,295, 361 4) and maintenance of hydraulic regimes the stand scale to meet multiple objectives at landscape
within boundaries of evolutionary adaptation for specific and larger scales. The landscape perspective also is
organisms.350 For example, in coastal Oregon, debris flows pertinent to managing riparian systems,143 where the role
from tributaries provide most of the large wood inputs.39 of wood decay in riparian environments varies according
However, in southeastern Alaska, floodplain forests to the type and geography of the associated water body.
adjacent to the stream are the primary source of wood to 2. Emphasize retention of wood legacies, and secondarily
the stream system.233 promote restoration where legacies are deficient to meet
New silvicultural methods being developed and tested stated objectives. The decline of species associated with
in riparian forests143 include re-establishing conifer species, late-successional forest structures, as well as the prolonged
retaining snags, down logs, and green trees, and avoiding time needed to produce wood legacies, suggests that it is
salvage. Underburning may be used to re-establish both ecologically and economically advantageous to retain
conifers, if necessary, to reduce competition with legacy structures across harvest cycles wherever possible,
understory shrubs. Thinning and underplanting may also rather than attempt to restore structures that have been
be used to create snags and provide large wood. In heavily depleted. This is especially obvious for slow-growing tree
degraded riparian areas, forest complexity can be species and very large wood structures. Retention of old-
improved by retaining large, and broken trees in the
 CHAPTER 24: WOOD LEGACIES 611

growth structural legacies has been identified as critical some discretion in determining the hazard area around a
to conservation of biodiversity between large reserves and danger tree, by “...allowing work to commence within two
conservation areas.222, 267 tree lengths of a marked danger tree, provided that the
employer demonstrates that a shorter distance will not
3. Use an adaptive management approach to assess
create a hazard for an employee.”(OSHA Logging
management options where possible. Our ability to
Preamble, Section V). Determining a safe working distance
sustain forest ecosystem values while producing
requires a case-by-case “...evaluation of various factors
commodities is uncertain. Given the imperfect state of our
such as, but not limited to, the size of the danger tree,
knowledge regarding management effects on biodiversity
how secure it is, its condition, the slope of the work area,
and long-term productivity of forests, prudence calls for
and the presence of other employees in the area.” The
an adaptive management approach that spreads
employer is responsible for marking hazard trees and
environmental risk across a range of management
making this determination. Washington State guidelines
strategies. Management must seek a blend of practices to
for reserve tree selection provide definitions of hazard
meet biological and social objectives.343 Guidance on
areas and examples of operational techniques that are
approaches to adaptive management is available in several
compatible with safe work practices.401a
publications.45, 110, 157, 338, 391
Oregon (Chapter 437, Oregon Administrative Rules)
4. Address management objectives and data needs to and Washington (Chapter 296-54, Washington
protect functions of dead wood in basic processes of Administrative Code) state safety regulations require
forest ecosystems, not solely to fish and wildlife habitat. employers to have a site-specific safety plan before logging
At the forest policy level, broad-scale assessments of down activities begin. The operator can include strategies for
wood are needed to address Criteria and Indicators for safe retention of reserve trees in the safety plan. A safety
the Conservation and Sustainable Management of consultation program, completely separate from OSHA’s
Temperate and Boreal Forests, developed through the inspection program, is available through state safety
Montreal Process. Although dead wood was not agencies. The operator may request free, confidential, on-
considered in the first national-level assessment, dead site assistance with safety planning, without risk of
wood abundance will be addressed in the first assessment citations or penalties. State safety plan requirements and
of forest sustainability to be conducted by any state in the OSHA consultation services are two key mechanisms to
U.S., by the Oregon Department of Forestry.36 resolve operational safety issues associated with structural
retention and restoration. Reserve tree requirements are
Operational Considerations best implemented through descriptive criteria in contract
Management for decaying wood will require new and or application conditions, rather than marking individual
innovative operational approaches. Structural retention trees and snags. Because the employer is responsible for
and restoration pose many new challenges to forest identifying hazards and preventing worker exposure, pre-
operations, both in current and future harvest activities. marked trees could reduce the operator’s flexibility to deal
To be successful, any plan for provision of wood decay with unforeseen hazards which arise in the course of
elements should address operational aspects of logging activities. If a tree marked by a timber sale
implementation. This includes scoping operational administrator or forest practices officer turned out to be a
approaches for plan implementation, and identifying danger tree, confusion over safety compliance authority
strategies for resolving potential conflicts ahead of time. also might result.
Two common sources of operational difficulties include Potential safety conflicts can be addressed in advance
safety concerns and associated administrative costs. by requiring a pre-work conference with the operator and
a state safety consultant to review the operator’s site-
Worker safety specific strategies for meeting reserve tree objectives in
From 1980-1989, the average annual fatality rate for compliance with safety regulations. At this time, the types,
workers in the logging industry was more than 23 times locations, and distributions of leave trees can be specified,
that for all U.S. workers. More than half of these fatalities considering the logging system to be used, and site-specific
occurred when workers were struck by falling or flying topography, unit layout and decay conditions. Once
objects or were caught in or between objects; most of these landings and tail holds are identified, reserve trees may
fatalities involved trees, logs, snags or limbs.264a An be mapped; as an example, clumps of reserve trees may
analysis of claims data from 1990-1997, ranked workers be left between cable roads. For the variety of situations
in the logging industry as having the highest risk of that may be encountered during operations, it is important
traumatic head and brain injuries in Washington state.401b to devise alternative strategies. This is a good way for
In recognition of the need for tree retention in harvests, managers to improve operational knowledge and expedite
OSHA revised the federal Logging Standard (29 CFR administrative compliance time.
1910.266) in 1995, to clarify its intent that danger trees may Concerns frequently arise where high public use creates
be avoided, rather than being removed or felled.72a A a risk of third party liability. Considerations include the
danger tree is any standing tree (live or dead) that poses a proximity of reserve trees to roads, trails, campgrounds,
hazard to workers, from unstable conditions such as ski areas, and other recreation areas and public access
deterioration, damage, or lean. The revised rule allows points. Methods for addressing these concerns include
 612 Wildlife–Habitat Relationships in Oregon and Washington

signage and clear delineation of potential hazard areas, Literature Cited

fencing and other barriers to discourage public access, 1. Aber, J. D., and J. M. Melillo. 1991. Terrestrial ecosystems. Saunders
snag height reduction and use of setbacks to minimize College Publishing, of Holt, Rinehart and Winston, Orlando, FL.
exposure. 1b. Acker, S. A., E. K. Zenner, and W.H. Emmingham. 1998. Structure and
yield of two-aged stands on the Willamette National Forest, Oregon:
Costs and Forest Management Complexity implications for green tree retention. Canadian Journal of Forest
Research 28:749-758.
Legacy retention generally results in higher logging costs
2. Agee, J. K. 1981. Fire effects on Pacific Northwest forests: flora, fuels,
than clearcutting and may require alternative logging and fauna. Pages 54-66 in Northwest Fire Council Proceedings,
technology. Compared to clearcutting, logging costs have Northwest Fire Council, Portland, OR.
been estimated to be significantly greater for dispersed 3. ———. 1988. Successional dynamics of forest riparian zones. Pages 31-
retention, but only slightly higher for aggregated 43 in K. J. Raedecke, editor. Streamside management: riparian wildlife
retention. 201,406a Aggregated retention has also been and forestry interactions. Institute of Forest Resources, Contribution
recommended to minimize interference with aerial Number 59. University of Washington, Seattle, WA.
4. ———. 1993. Fire ecology of the Pacific Northwest forests.
application of pesticides, fertilizers, and herbicides.201
Washington, D.C. Island Press.
Other concern regarding tree retention are effects on 5. ———. 1994. Fire and weather disturbances in terrestrial ecosystems
stocking density and genetic composition of the of the eastern Cascades. U.S. Forest Service, Pacific Northwest
regenerating forest. Compared to live trees, the retention Research Station, Portland, Oregon. General Technical Report PNW-
of snags, decadent trees, and down wood within a harvest GTR-320.
unit have little influence on future stand genetics or 6. ———, and M. H. Huff. 1987. Fuel succession in a western hemlock /
stocking density. Retention of live trees and decaying Douglas-fir forest. Canadian Journal of Forest Research 17:697-704.
7. Akenson, J. J., and M. G. Henjum. 1994. Black bear den site selection in
wood may reduce wood yields due to volume in wood
the Starkey study area. Natural Resource News 4(2):1-2. Blue
structures permanently retained on a site, or reduced Mountains Natural Resource Institute, La Grande, OR.
growth in the regenerating stand due to shading.1b, 311, 8. Allen, R. B., P. W. Clinton, and M. R. Davis. 1997. Cation storage and
311a,401c, 406
Conversely, decaying wood may increase future availability along a Nothofagus forest development sequence in New
wood yields by improving seedling survival and growth, Zealand. Canadian Journal of Forest Research 27:323-330.
and site productivity.164 9. Amaranthus, M. P., and D. A. Perry. 1987. Effect of soil transfer on
extomycorrhiza formation and the survival and growth of conifer
Long Range Planning Considerations seedlings on old, nonreforested clear-cuts. Canadian Journal of Forest
Long-range plans should provide criteria for tree selection Research 17:944-950.
10. ———, R. J. Molina, and J. M. Trappe. 1989 a. Long-term productivity
and distribution that are flexible enough to account for and the living soil. Pages 36-52 in: D. Perry, D., R. Meurisse, B. Thomas, R.
ongoing tree decay processes, and changes in harvest Miller, and J. Boyle, editors. Maintaining the long-term productivity of
plans. For instance, a time lag between unit layout and Pacific Northwest ecosystems. Timber Press. Portland, OR.
harvest may result in changes in reserve tree decay classes. 11. ———, D. S. Parrish, and D. A. Perry. 1989 b. Decaying logs as
It is also important to consider how reserve trees will fit moisture reservoirs after drought and wildfire. Pages 191-194 in: E. B.
into plans for other forest practices after harvest, such as Alexander, editor. Proceedings of Watershed ’89: A Conference On The
Stewardship Of Soil, Air, And Water Resources. U.S. Forest Service,
thinning operations, fertilization, vegetation management,
Juneau, AK.
and fire control. This is particularly important in uneven- 12. ———, J. F. Weigand, and R. Abbott. 1998. Managing high-elevation
age management schemes where multiple entries are forests to produce American matsutake (Tricholoma magnivelare), high-
planned. When developing conceptual long range plans quality timber, and non-timber forest products. Western Journal of
and site-specific plans, it is thus important to consider Applied Forestry 13(4): 1-9.
whether a hazard would exist by the time future activities 13. ———, J. M. Trappe, L. Bednar, and D. Arthur. 1994. Hypogeous fungal
take place. production in mature Douglas-fir forest fragments and surrounding
plantations and its relation to coarse woody debris and animal
New Approaches mycophagy. Canadian Journal of Forest Research 24:2157-2165.
14.Anderson, N. H.1982. A survey of aquatic insects associated with
Few studies have analyzed operational challenges to
wood debris in New Zealand streams. Mauri Ora 10:21-23.
managing for decaying wood. However, a variety of forest 15. Aplet, G. H., N. Johnson, J. T. Olson, and V. A. Sample, editors. 1993.
managers are currently testing various methods to retain Defining sustainable forestry. Island Press, Washington, D.C.
and restore decaying wood in various forms as snags, 16. Apps, M. J. and D.T. Price. 1996. Forest ecosystems, forest
decadent trees, and down wood. Recent publications offer management and the global carbon cycle. Springer, New York, NY.
useful discussions of concepts and silvicultural 17. Arnolds, E. 1991. Decline of ectomycorrhizal fungi in Europe.
approaches to managing for structurally complex forest Agriculture, Ecosystems & Environment 35:209-244.
18. Arthur, M. A. and T. J. Fahey. 1990. Mass and nutrient content of
stands, including operational considerations; they also decaying boles in an Engelmann spruce—subalpine fir forest, Rocky
discuss recent lessons from experimental forestry Mountain National Park, Colorado. Canadian Journal of Forest
operations.31a, 69a, 201, 288a, 328, 406a Research 20:730-737.
19. Asner, G. P., T. R. Seastedt, and A. R. Townsend. 1999.The decoupling of
terrestrial carbon and nitrogen cycles. BioScience 47(4):226-234.
20. Aubry, K. B., Amaranthus, M. P., Halpern, C. B., White, J. D., Woodard, B.
L., Peterson, C. E., Lagoudakis, C. A., and A. J. Horton. 1999. Evaluating
the effects of varying levels and patterns of green-tree retention:
experimental design of the DEMO study. Northwest Science 73: 12-26.
 CHAPTER 24: WOOD LEGACIES 613

21. ———, J. G. Hallett, S. D. West, M. A. O’Connell, and D. A. Manuwal. 38. ———, and R. E. Bilby. 1998. Organic matter and trophic dynamics.
1997. Wildlife use of managed forests: a landscape perspective,Volume Pages 373-398 in R. J. Naiman, and R. E. Bilby, editors. River ecology and
1. Timber Fish and Wildife Program, Report No. TFW-WL4-98-001. management. Springer, New York, NY.
22. ———, L. L. Jones, and P. A. Hall. 1988. Use of woody debris by 39. ———, T. P. Quinn, G. H. Reeves, and S. V. Gregory. 1992. Best
plethodontid salamanders in Douglas-fir forest in Washington. Pages management practices, cumulative effects, and long-term trends in fish
32-37 in R. C. Szaro, K. E. Severson, and D. R. Patton, technical abundance in Pacific Northwest river systems. Pages 189-232 in R.J.
coordinators. Management of amphibians, reptiles, and small mammals Naiman, editor. Watershed management balancing sustainability and
in North America. Proceedings of a symposium, July 19-21 1988 in environmental change. Springer Verlag, New York, NY.
Flagstaff, AZ. U.S. Forest Service, Rocky Mountain Forest and Range 40. ———, R. E. Bilby, M. D. Bryant, C. A. Dolloff, G. B. Grette. R. A. House,
Experiment Station, Fort Collins, CO. General Technical Report RM- M. L. Murphy, K. V. Koski, and J. R. Sedell. 1987. Large woody debris in
GTR-166. forested streams in the Pacific Northwest: past, present, and future.
23. Ausmus, B. S., G. J. Dodson, and D. G. Todd. 1975. Microbial - Pages 143-190 in E. O. Salon and T. W. Cundy, editors. Streamside
invertebrate interactions: The mechanism of wood decomposition. management: forestry and fishery interactions. Contribution 57,
Ecological Society of America Bulletin 56(2):42. Institute of Forest Resources, University of Washington, Seattle, WA.
24. Avery, C.C., F.R. Larson, and G. H. Schubert. 1976. Fifty year records of 41. Bolsinger, C. L. and K. L. Waddell. 1993. Area of old-growth forests in
virgin stand development in southwestern ponderosa pine. U.S. Forest California, Oregon, and Washington. U.S. Forest Service, Pacific
Service, Rocky Mountain Forest And Range Experiment Station, Fort Northwest Research Station, Resource Bulletin PNW-RB-197. Portland,
Collins, Colorado. General Technical Report RM-GTR-22. OR.
25. Baker, F. A., Daniels, S. E., and C. A. Parks. 1996. Inoculating trees with 42. ———, N. McKay, D. R. Gedney, and C. Alerich. 1997. Washington’s
wood decay fungi with rifle and shotgun. Western Journal of Applied public and private forests. U.S. Forest Service, Pacific Northwest
Forestry 11(1): 13-15. Research Station, Resource Bulletin PNW-RB-218. Portland, OR.
26. Bailey, J. D., and J. C. Tappeiner. 1998. Effects of thinning on structural 43. Borchers, J. G. and D. A. Perry. 1990. Effects of prescribed fire on soil
development in 40- to 100-year-old Douglas-fir stands in western organisms. Pages 143-175 in J. D. Walstad, S. R. Radosevich, and D.V.
Oregon. Forest Ecology and Management 108: 99-113. Sandberg, editors. Natural and prescribed fire in Pacific Northwest
27. Bate, L. J. , E.O. Garton, and M. J. Wisdom. 1999. Estimating snag and forests. Oregon State University Press, Corvallis, OR.
large tree densities and distributions on a landscape for wildlife 44. ———, D. A. Perry, P. Sollins. et al. 1992. The influence of soil texture
management.U.S. Forest Service, Pacific Northwest Research Station, and aggregation on carbon and nitrogen dynamics in southwest
Portland, Oregon. General Technical Report PNW-GTR-425. Oregon forests and clear-cuts. Canadian Journal of Forest Research
28. Bartels, R., J. D. Dell, R. L. Knight, and G. Schaefer. 1985. Dead and 22:298-305.
down woody material. Pages 172-186 in E.R. Brown, editor. 45. Bormann, B. T., P. G. Cunningham, M. H. Brookes, V. W. Manning, and M.
Management of wildlife and fish habitats in forests of western Oregon W. Collopy. 1994. Adaptive ecosystem management in the Pacific
and Washington. U.S. Forest Service, Pacific Northwest Region. Northwest. U.S. Forest Service, Pacific Northwest Research Station,
29. Bednarz, J. C., D. M. Juliano, M. J. Huss, A. A. Beall, and D. E.Varland. General Technical Report PNW-GTR-341.
1998. Interactions between fungi and cavity nesters: applications to 46. ———, H. Spaltenstein, M. H. McClellan, F.C. Ugolini, K. Cromack, Jr.,
managing wildlife and forests in western Washington: 1997 annual S. M. Nay. 1995. Rapid soil development after windthrow disturbance in
report. Unpublished report. Arkansas State University. pristine forests. Journal of Ecology 83:1-20.
30. Beese, W. J., and A. B. Bryant. 1999. Effect of alternative silvicultural 47. Breznak, J. A. 1982. Intestinal microbiota of termites and other
systems on vegetation and bird communities in coastal montane xylophagous insects. Annual Review of Microbiology 36:323-343.
forests of British Columbia, Canada. Forest Ecology and Management 48. Brown, R. E. 1985. Management of wildlife and fish habitats in forests
115: 231-242. of western Oregon and Washington. U.S. Forest Service, Pacific
31. Benda, L. E. and T. Dunne. 1987. Sediment routing by debris flows. Northwest Region.
Pages 213-223 in R. L. Beschta, T. Blinn, G. E. Grant, G. G. Ice, and F. J. 49. Brown, T. K. 1996. Snags and wildlife tree preservation and
Swanson, editors. Erosion and sedimentation in the Pacific Rim. enhancement. Pages 71-75 in P. Bradford, T. Manning, and B. l’Anson.
International Association of Hydrological Sciences, Publication 165. Editors. Wildlife tree/stand-level biodiversity workshop proceedings.
Oxfordshire, England. October 17-18, 1995, Victoria, BC, Canada.
31a. Berg, D. R. 1995. Forest harvest-setting design evaluation 50. Bull, E. L. 1996. Owl nests in brooms caused by dwarf mistletoes and
incorporating logger’s preference. Ph.D. Dissertation. University of Elytroderma disease. U.S. Forest Service, Pacific Northwest Research
Washington, Seattle. Station, Portland, OR. Unpublished report.
32. Beschta, R. L. and W. S. Platts. 1986. Morphological features of small 51. ———. 1995. Progress report on American marten home range and
streams: significance and function. Water Resources Bulletin 22:369- habitat use. U.S. Forest Service, Pacific Northwest Research Station,
379. Portland, OR. Unpublished report.
33. Bilby, R. E. and P. A. Bisson. 1998. Function and distribution of large 52. ———, and C.T. Collins. 1993.Vaux’s swift (Chaetura vauci). In A.
woody debris. Pages 324-346 in R. J. Naiman, and R. E. Bilby, editors. Poole, and E. Gill, editors, The Birds Of North America. The Academy of
River ecology and management. Springer, New York, NY. Natural Sciences, Report No 77., Philadelphia, PA.
34. Binkley, D. and T. C. Brown. 1993. Management impacts on water 53. ———. and R. S. Holthausen. 1993. Habitat use and management of
quality of forests and rangelands. U.S. Forest Service, Rocky Mountain pileated woodpeckers in northeastern Oregon. Journal of Wildlife
Research Station, General Technical Report RM-GTR-239. Fort Collins, Management 57:335-345.
CO. 54. ———. and A. D. Partridge. 1986. Methods of killing trees for use by
35. ———, K. Cromack, Jr., and R. L. Fredricksen. 1982. Nitrogen cavity nesters. Wildlife Society Bulletin 14:142-146.
accretion and availability in some snowbrush ecosystems. Forest 55. ———, R. S. Holthausen, and M.G. Henjum. 1992. Roost trees used
Science 28(4):720-724. by pileated woodpeckers in northeastern Oregon. Journal of Wildlife
36. Birch, K. 1999. First approximation report for sustainable forest Management. 56:786-793.
management in Oregon. Oregon Department of Forestry. Salem, OR. 56. ———, C. G. Parks, and T. R. Torgerson. 1997. Trees and logs
37. Bisson, P. A., J. L. Nielsen, R. A. Palmason, and L. E. Grove. 1982. A important to wildlife in the interior Columbia River Basin. U.S. Forest
system of naming habitat types in small streams, with examples of Service, Pacific Northwest Research Station, General Technical Report
habitat utilization by salmonids during low streamflow. Pages 62-73 in PNW-GTR-391.
N. B. Armantrout, editor. Acquisition and utilization of aquatic habitat 57. ———, A. D. Partridge, and W. G. Williams. 1981. Creating snags with
inventory information. Western Division, American Fisheries Society, explosives. U.S. Forest Service, Pacific Northwest Research Station,
Portland, Oregon. The Hague Publishing, Billings, MT. Portland, OR. Research Note PNW-RN-393.
 614 Wildlife–Habitat Relationships in Oregon and Washington

58. Burge, W. D. and F. E. Broadbent. 1961. Fixation of ammonia by 75. Chappell, C., R. Crawford, J. Kagan, and P. J. Doran. 1997. A vegetation,
organic soils. Soil Science Society of America Proceedings 25:199-204. land use, and habitat classification system for the terrestrial and aquatic
59. Burns, R. M., compiler. 1983. Silvicultural systems for the major forest systems of Oregon and Washington. Wildlife habitats and species
types of the United States. U.S. Forest Service, Agriculture Handbook associations in Oregon and Washington—progress report #3,
No. 445. Washington, D.C. Washington Department of Fish And Wildlife, Olympia.
59a Bury, R. B., and P. S. Corn. 1988. Douglas fir forests in Oregon and 76. Chen, H. 1989. Studies on tree mortality and log decomposition of
Washington Cascades; relation of the herpetofauna to stand age and main species in conifer-deciduous mixed forest on Changbai Mountain
moisture. Pages 11-122 in R. C. Szaro, K. E. Severson, and D. R. Patton, Reserve. Thesis. Academia Sinica, Shenyang, People’s Republic of China.
technical coordinators. Management of amphibians, reptiles, and small 77. Childs, T. W., K. R. Shea, J. L. Stewart. 1971. Elytroderma disease of
mammals in North America. Proceedings of a symposium, July 19-21 ponderosa pine. U.S. Forest Service, Forest Insects And Disease
1988 in Flagstaff, AZ. U. S. Forest Service, Rocky Mountain Forest and Program, Leaflet No.42.
Range Experiment Station, Fort Collins, CO. General Technical Report 78. Christensen, N. L. et al. 1996. The report of the Ecological Society of
RM-GTR-166. America commitee on the scientific basis for ecosystem management.
60. Camp, A. E. (submitted). Demographics of dead trees on the eastern Ecological Applications 6:665-691.
slope of the Washington Cascades from the Cascades crest to the 79. Christiansen, E., R. H. Waring, and A. A. Berryman. 1987. Resistance of
Columbia Plateau. Northwest Science. conifers to bark beetle attack: searching for general relationships.
61. Carey, A. B. and R. O. Curtis. 1996. Conservation of biodiversity: a Forest Ecology and Management 22:89-106.
useful paradigm for forest ecosystem management. Wildlife Society 80. Christy, E. J. and R. N. Mack. 1984. Variation in demography of juvenile
Bulletin 24(4):610-620. Tsuga heterophylla across the substratum mosaic. Journal of Ecology
62. ———. and J. D. Gill. 1983. Direct habitat improvements: some 72:75-91.
recent advances. In J. W. Davis, G. A. Goodwin, and R. A. Ockerfells, 81. Cline, S. P. and C.A. Phillips. 1983. Coarse woody debris and debris-
technical coordinators. Snag habitat management: proceedings of a dependent wildlife in logged and natural riparian zone forests: a
symposium. U. S. Forest Service, Washington, D.C. Technical Report RM- western Oregon example. Pages 33-39 in J. W. Davis, G. A. Goodwin,
99. and R. A. Ockenfels, technical coordinators. Snag Habitat Management:
63. ———. 1995. Sciurids in Pacific Northwest managed and old-growth Proceedings of the Symposium. U.S. Forest Service, Rocky Mountain
forests. Ecological Applications 5(3):648-661. Research Station, General Technical Report, RM-GTR-99.
64. ———. and M. L. Johnson. 1995. Small mammals in managed, 82. ———, A. B. Berg, and H. M. Wight. 1980. Snag characteristics and
naturally young, and old-growth forests. Ecological Applications 5(2): dynamics in Douglas-fir forests, Western Oregon. Journal of Wildlife
336-352. Management 44(4): 773-786.
65. ———, and H. R. Sanderson. 1981. Routing to accelerate tree-cavity 83. Clinton, P. W., R. B. Allen, R. H. Newman, and M. R. Davis. 1997.
formation. Wildife Society Bulletin 9:14-21. Nitrogen availability and forest development in mountain beech. Soil
66. ———, D. R. Thysell, and A. Brodie. 1999. The forest ecosystem study: Biology and Biochemistry: in press.
background, rationale, implementation, baseline conditions, and 84. Conner, R. N., J. G. Dickson, and J. H. Williamson. 1983. Potential
silvicultural assessment. U.S. Forest Service, Pacific Northwest woodpecker nest trees through artificial innoculation with heart rots.
Research Station, Portland, OR. General Technical Report PNW-GTR- In J. W. Davis, G. A. Goodwin, and R. A. Ockerfells, technical
457. coordinators. Snag habitat management: proceedings of a symposium.
67. ———, C. Elliott, B. R. Lippke, J. Sessions, C. J. Chambers, C. D. Oliver, U.S. Forest Service, Washington, D.C. Technical Report RM-99.
J. F. Franklin, and M. J. Raphael. 1996. A pragmatic ecological approach to 85. Cornaby, B. W., and J. B. Waide. 1973. Nitrogen fixation in decaying
small-landscape management. Washington Forest Landscape chestnut logs. Plant and Soil 39(2):445-448.
Management Project—report No.2. Washington Forest Landscape 86. Coutts, M. P. and J. Grace. 1995. Wind and trees. Cambridge
Management Project Report. No. 2, Washington Department of Natural University Press, New York, NY.
Resources, Olympia, WA. 87. Cromack, K., Jr., R. L. Todd, and C. D. Monk. 1975. Patterns of
68. ———, D. R. Thysell, L. J.Villa, T. M. Wilson, S. M. Wilson, J. M. Trappe, W. Basidiomycete nutrient accumulation in conifer and deciduous forest
Colgan III, E. R. Ingham, and M. Holmes. 1996. Foundations of litter. Soil Biology And Biochemistry 7(4-5): 265-268.
biodiversity in managed Douglas-fir forests. Pages 68-82 in D. L. Pearson 88. ———, C. C. Delwiche, and D. H. McNabb. 1979. Prospects and
and C.V. Klimas, editors. The role of restoration in ecosystem problems of nitrogen management using symbiotic nitrogen fixers.
management. Society For Ecological Restoration, Madison, WI. Pages 210-223 in J. C. Gordon, C. T. Wheeler, and D. A. Perry, editors.
69. ———, T. M. Wilson, C. C. Maguire, B. Biswell. 1997. Dens of Symbiotic nitrogen fixation in the management of temperate forests.
northern flying squirrels in the Pacific Northwest. Journal of Wildife Forest Research Lab, Corvallis, OR.
Management 61(3): 684-699. 89. Crossley, D. A. Jr. 1976. The roles of terrestrial saprophagous
70. ———. 2000. Effects of new forest management strategies on arthropods in forest soils: current status of concepts. Pages 49-56 in W.
squirrel populations. Ecological Applications: 10(1):248-257. J. Mattson, editor. The role of arthropods in forest ecosystems. Springer
71. Caza, C. L. 1993. Woody debris in the forests of British Columbia: a Verlag, New York, NY.
review of the literature and current research. Ministry Of Forests, 90. Cummins, K. W., J. R. Sedell, F. J. Swanson, G. W. Minshall, S. G. Fisher, C.
British Columbia, Canada. E. Cushing, R. C. Petersen, and R. L.Vannote. 1982. Organic matter
72. Cederholm, C. J. and N. P. Peterson. 1985. The retention of coho budgets for stream ecosystems: problems in their evaluation. Pages
salmon (Oncorhynchus kisutch) carcasses by organic debris in small 299-353 in G.W. Minshall and J. R. Barnes, editors. Stream ecology:
streams. Canadian Journal of Fisheries and Aquatic Sciences 42: 1222- application and testing of general ecological theory. Plenum, New York,
1225. NY.
72a. CFR 1995. Logging Standard (29 CFR 1910.266) Code of Federal 91. Curtis, R. O. 1995. Extended rotations and culmination age of coast
Regulations. Washington, DC.:US Government Printing Office, Office of Douglas-fir: Old studies speak to current issues. U.S. Forest Service,
the Federal Register. Pacific Northwest Research Station, Portland, Oregon. Research Paper
73. Chambers, C. L., W. C. McComb, and J. C. Tappeiner, II. 1999. Breeding PNW-RP-485.
bird responses to three silvicultural treatments in the Oregon Coast 92. ———. 1997. The role of extended rotations. Pages 165-170 in K. A.
Range. Ecological Applications 9(1): 171-185. Kohm, and J. F. Franklin, editors. Creating a forestry for the 21st
74. Chapman, D.W. and K. P. McLeod. 1987. Development of criteria for century: the science of ecosystem management. Island Press,
fine sediment in the Northern Rockies Ecoregion. U.S. Environmental Washington, D.C.
Protection Agency, Water Division, Report 910/9-87-162, Seattle, WA. 93. ———, and D. D. Marshall. 1993. Douglas-fir rotations: time for
reappraisal? Western Journal of Applied Forestry 8(3): 81-85.
 CHAPTER 24: WOOD LEGACIES 615

94. ——— D. S. DeBell, C. A. Harrington, D. P. Lavender, J. B. St. Clair, J. C. 114. Federal Ecosystem Management Analysis Team (FEMAT). 1993.
Tappeiner, and J. D. Walstad. et al. 1998. Silviculture for multiple Chapter III, Option Development and Description, and Chapter IV,
objectives in the Douglas-fir region. U.S. Forest Service, Pacific Terrestrial Forest Ecosystem Assessment. Forest Ecosystem
Northwest Research Station, Portland, Oregon. General Technical Management: An Ecological, Economic, and Social Assessment. U.S.
Report PNW-GTR-435. Government Printing Office.
95. Daily, G. C. 1993. Heartwood decay and vertical distribution of red- 115. Fogel, R. and K. Cromack, Jr. 1977. Effect of habitat and substrate
naped sapsucker nest cavities. Wilson Bulletin 105(4):674-679. quality on Douglas-fir litter decomposition in western Oregon.
96. DeBell, D. S., R. O. Curtis, C. A. Harrington, and J. C. Tappeiner. 1997. Canadian Journal of Botany 55(12):1632-1640.
Shaping stand development through silvicultural practices. Pages 141- 116. Fogel, R., M. Ogawa, and J. M. Trappe. 1973. Terrestrial decomposition:
150 in K.A. Kohm, and J.F. Franklin, editors. Creating a forestry for the a synopsis. U.S. International Biological Program, Coniferous Forest
21st century: the science of ecosystem management. Island Press, Biome Internal Report 135. University of Washington, Seattle, WA.
Washington, D.C. 117. Foster, F. R. and G. E. Lang. 1982. Decomposition of red spruce and
97. Deyrup, M. A. 1975. The insect community of dead and dying Douglas- balsam fir boles in the White Mountains of New Hampshire. Canadian
fir. 1: The Hymenoptera. Ecosystem Analysis Studies Bulletin 6, Journal of Forest Research 12:617-626.
Coniferous Forest Biome. University of Washington, Seattle, WA. 118. Francke-Grosmann, H. 1967. Ectosymbiosis in wood inhabiting
98. ———. 1976. The insect community of dead and dying Douglas-fir : insects. Pages 142-205 in S. M. Henry, editor. Symbiosis.Volume 2.
Diptera, Coleoptera, and Neuroptera. Dissertation, University of Academic Press, New York, NY.
Washington, Seattle, WA. 119. Frankland, J. C., J. N. Hedger, and M. J. Swift, editors. 1982.
99. ———. 1981. Deadwood decomposers. Natural History 90:84-91. Decomposer basidiomycetes, their biology and ecology. Cambridge
100. Diaz, S., J. P. Grime, J. Harris, and E. McPherson. 1993. Evidence of a University Press, New York, NY.
feedback mechanism limiting plant response to elevated CO2. Nature 120. Franklin, J.F. 1988. Pacific Northwest forests. Pages 104-130 in M. G.
364:616-617. Barbour, and W. D. Billings, editors. North American terrestrial
101. Dickson, B. A., and R. L. Crocker. 1953. A chronosequence of vegetation. Cambridge University Press, New York, NY.
vegetation and soils near Mt. Shasta, California. II. The development of 121. ———. 1989. Toward a new forestry. American Forests 95 (11/12):
the forest floors and the carbon and nitrogen profiles of the soils. 37-44.
Journal of Soil Science 4:142-154. 122. ———. 1992. The scientific basis for new perspectives in forests and
102. Dixon, R. D. 1995. Ecology of white-headed woodpeckers in the streams. Pages 25-72 in R. J. Naiman, editor. Watershed management
central Oregon Cascades. Thesis, University of Idaho, Moscow, ID. balancing sustainability and environmental change. Springer Verlag, New
103. Doyle, A.T. 1985. Small mammal micro- and macrohabitat selection York, NY.
in streamside ecosystems. Dissertation, Oregon State University, 123. ———, and C.T. Dyrness. 1988. Natural vegetation of Oregon and
Corvallis, OR. Washington. Oregon State University Press, Corvallis, OR.
104. ———. 1990. Use of riparian and upland habitat by small mammals. 124. ———, and C. Maser. 1988. Looking ahead: some options for public
Journal of Mammalogy 71:14-23. lands. Pages 113-122 in C. Maser, R. F. Tarrant, J. M. Trappe, and J. F.
105. Edmonds, R. L. 1987. Decomposition rates and nutrient dynamics in Franklin, editors. U.S. Forest Service, Pacific Northwest Research
small-diameter woody litter in four forest ecosystems in Washington, Station, General Technical Report PNW-GTR-229, Portland, OR.
U.S.A. Canadian Journal of Forest Research 17:499-509. 125. ———, and T. A. Spies. 1991. Composition, function, and structure of
106. ———. 1991. Organic matter decomposition in western United old-growth Douglas-fir forests. Pages 71-82 in L. F. Ruggiero, K.B. Aubry,
States. Pages 118-128 in A.E. Harvey and L. F. Neuenschwander, editors. A.B. Carey, and M.H. Huff, technical coordinators. Wildlife and
Management and productivity of western-montane forest soils. U.S. vegetation of unmanaged Douglas-fir Forests. U.S. Forest Service,
Forest Service, Intermountain Research Station, Ogden, UT. Pacific Northwest Research Station, Portland, OR, General Technical
107. ———, and A. Eglitis. 1989. The role of Douglas-fir beetle and wood Report, PNW-GTR-285.
borers in the decomposition and nutrient release from Douglas-fir logs. 126. ———, and T. A. Spies. 1991b. Ecological definitions of old-growth
Canadian Journal of Forest Research 19:327-337. Douglas-fir forests. Pages 61-70 in L. F. Ruggiero. K. B. Aubry, A. B. Carey,
108. Entry, J.A., N. E. Martin, and K. Cromack, Jr. 1986. Light and nutrient and M. H. Huff, technical coordinators. Wildlife and vegetation of
limitation in Pinus monticola: seedling susceptibility to Armillaria unmanaged Douglas-fir forests. U.S. Forest Service, Pacific Northwest
infection. Forest Ecology and Management 17:189-198. Research Station, General Technical Report, PNW-GTR-285, Portland,
109. Everest, F. H., R. L. Beschta, J. C. Srivener, K. V. Koski, J. R. Sedell, and C. OR.
J. Cederholm. 1987. Fine sediment and salmon production: a paradox. 127. ———, and R. H. Waring. 1980. Distinctive features of the
Pages 98-142 in E. O. Salo, and T. W. Cundy, editors. Streamside Nortwestern coniferous forest: Development, structure, and function
management: forestry and fishery interactions, Institute of Forest Pages 59-85 in R. H. Waring, editor. Forests: fresh perspectives from
Resources, Contribution 57. University of Washington, Seattle, WA. ecosystem analysis. Proceedings of the 40th Biology Colloquium
110. Everett, R., C. Oliver, J. Saveland, P. Hessburg, N. Diaz, and L. (1979), Oregon State University. Press, Corvallis, OR.
Irwin.1993. Adaptive ecosystem management. Pages 351-364 in M. 128. ———, P. M. Frenzen, and F. J. Swanson. 1988. Re-creation of
Jensen and P. Bourgeron, editors. Ecosystem management: principles ecosystems at Mount St. Helens: contrasts in artificial and natural
and application. U. S. Forest Service, Northern Region. Missoula, MT. approaches. Pages 1-37 in J. Cairns, Jr., editor. Rehabilitating damaged
111. ———, L. Lehmkuhl, R. Schellhaas, P. Ohlson, D. Keenum, H. ecosystems.Vol. 2. CRC Press, Boca Raton, FL.
Riesterer, and D. Spurbeck. In press. Snag dynamics in a 129. ———, J. A. Mac Mahon, F. J. Swanson, and J. R. Sedell. 1985.
chronosequence of 26 wildfires on the east slope of the Cascade Ecosystem responses to the eruption of Mt. St. Helens. National
Range in Washington. International Journal of Wildland Fire. Geographic Research. 2: 198-216.
112. ———,compiler. 1994.Volume IV: restoration of stressed sites, and 130. ———, D. A. Perry, T. A. Schowalter, M. E. Harmon, A. McKee, and T. A.
processes. U.S. Forest Service, Pacific Northwest Research Station, Spies. 1989. Importance of ecological diversity to long-term site
Portland, Oregon. General Technical Report PNW-GTR-30. productivity. Pages 82-97 in D. A. Perry, R. Meurisse, B. Thomas, R. Miller,
113. Fahey, T. J., J. W. Hughes, M. Pu, and M. A. Arthur. 1988. Root J. Boyle, J. Means, C. R. Perry, and R. F. Powers, editors. Maintaining the
decomposition and nutrient flux following whole-tree harvest of productivity of Pacific Northwest forest ecosystems. Timber Press,
northern hardwood forest. Forest Science 34(3): 744-768. Portland, OR.
 616 Wildlife–Habitat Relationships in Oregon and Washington

131. ———, K. Cromack, Jr., W. Denison, A. McKee, C. Maser, J. R.Sedell, F. 152. ———, S. A. Evans, C. R. Nelson, D. McKenzie, D. Liquori, D. E. Hibbs,
J. Swanson, and G. Juday. 1981. Ecological characteristics of old-growth and M. G. Halaj. 1999. Response of forest vegetation to varying levels
forests. U.S. Forest Service, Pacific Northwest Research Station, and patterns of green-tree retention: an overview of a long-term
Portland, OR. General Technical Report, PNW-GTR-118. experiment. Northwest Science 73: 27-43.
132. ———, D. R. Berg, D. A. Thornburgh, and J. C. Tappeiner. 1997. 153. Hamer, T. E., and S. K. Nelson. 1995. Characteristics of marbled
Alternative silvicultural approaches to timber harvesting: variable murrelet nest trees and nesting stands. Pages 69-87 in C. J. Ralph, G. L.
retention harvest systems. 1997. Pages 111-139 in K.A. Kohm, and J. F. Hunt Jr., M. G. Raphael, and J. F. Piatt, editors. Ecology and conservation
Franklin, editors. Creating a forestry for the 21st century: the science of the marbled murrelet. U.S. Forest Service, Pacific Southwest Forest
of ecosystem management. Island Press, Washington, D.C. and Range Experiment Station, Albany, CA. General Technical Report
133. Furniss, R. L. and V. M. Carolin. 1977. Western Forest Insects. U.S. PSW-GTR-152.
Forest Service Publication No. 1339. 154. Hansen, A. J., Spies, T. A., Swanson, F. J., and J. L. Ohmann. 1991.
134. Gartner, B. L. 1995. Plant stems: physiology and functional Conserving biodiversity in managed forests. BioScience 41:382-392.
morphology. Academic Press, New York, NY. 155. ———, W. C. McComb, R.Vega, M. G. Raphael, and M. Hunter. 1995.
135. Gashwiler, J. S. 1959. Small mammal study in west-central Oregon. Bird habitat relationships in natural and managed forests in the west
Journal of Mammalogy 40(1): 128-139. Cacades of Oregon. Ecological Applications 5(3):555-569.
136. ———. 1970. Plant and mammal changes on a clear-cut in west 156. ———, S. L. Garman, J. F. Weigand, D. L. Urban, W. C. McComb, and
central Oregon. Ecology 51(6):1018-1026. M. G. Raphael. 1995. Alternative silvicultural regimes in the Pacific
137. Gast, W., D. Scatt, C. Schmitt, D. Clemens, S. Howes, C. G. Johnson, Jr., Northwest: simulations of ecological and economic effects. Ecologcial
R. Mason, F. Mofr, and R. A. Clapp. 1991. Blue Mountains Forest Health Applications 5(3):535-554.
Report: New Perspectives In Forest Health. U.S. Forest Service, Pacific 157. ———, S. L. Garman, B. Marks, and D. L. Urban. 1993. An approach
Northwest Region, Malheur, Umatilla, and Wallowa-Whitman National for managing vertebrate diversity across multiple-use landscapes.
Forest. Ecological Applications 3: 481-496.
138. Gholz, H. L. 1982. Environmental limits on aboveground net primary 158. Harmon, M.E. 1992. Long-term experiments on log decomposition
production, leaf area, and biomass in vegetation zones of the Pacific at the H. J. Andrews Experimental Forest. U.S. Forest Service, Pacific
Northwest. Ecology 63(2):469-481. Northwest Research Station, General Technical Report, PNW-GTR-
139. Gonor, J. J., J. R. Sedell, and P. A. Benner. 1988. What we know about 280.
large trees in estuaries, in the sea, and on coastal beaches. Pages 83-112 159. ———, and H. Chen. 1991. Coarse woody debris dynamics in two
in: Maser, C., R. F. Tarrant, J.M. Trappe, and J. F. Franklin, technical editors. old-growth ecosystems: comparing a deciduous forest in China and a
1988. From the forest to the sea: a story of fallen trees. U.S. Forest conifer forest in Oregon. BioScience 41:604-610.
Service, Pacific Northwest Research Station, Portland, OR. General 160. ———, and J. F. Franklin. 1989. Tree seedlings on logs in Picea-Tsuga
Technical Report PNW-GTR-229. forests of Washington and Oregon. Ecology 70(1):48-59.
140. Graham, R.L. 1982. Biomass dynamics of dead Douglas-fir and 161. ——— W. K. Ferell, and J. F. Franklin. 1990. Effects on carbon storage
western hemlock boles in mid-elevation forests of the Cascade Range. of conversion of old-growth forests to young forests. Science 247:699-
Ph.D. Dissertation. Oregon State University, Corvallis, OR. 702.
141. ———, and K. C. Cromack, Jr. 1982. Mass, nutrient, content, and 162. ———, J. Sexton, B. A. Caldwell, and S. E. Carpenter. 1994. Fungal
decay rate of dead boles in rain forests of Olympic National Park. sporocarp mediated losses of Ca, Fe, K, Mg, N, P, and Zn from conifer
Canadian Journal of Forest Research 12:511-521. logs in the early stages of decomposition. Canadian Journal of Forestry
142. Grant, G. E., F. J. Swanson, and M. G. Wolman. 1990. Pattern and Research 24: 1883-1893.
origin of stepped-bed morphology in high-gradient streams, western 163. ———. 1989. Effects of bark fragmentation on plant succession on
Cascades, Oregon. Geological Society of America Bulletin 102:340-352. conifer logs in Picea-Tsuga forests of Olympic National Park,
143. Gregory, S.V. 1997. Riparian Management In The 21st Century. Pages Washington. American Midland Naturalist 121:112-124.
69-86 in K. A. Kohm, and J. F. Franklin, editors, Creating a forestry for 164. ———, J. F. Franklin, F. J. Swanson, P. Sollins, S. V. Gregory, J. D. Lattin,
the 21st century. The science of ecosystem management. Island Press, N. H. Anderson, S. P. Cline, N. G. Aumen, J. R. Sedell, G. W. Lienkaemper,
Washington, D.C. K. Cromack, Jr., and K. W. Cummins. 1986. Ecology of coarse woody
144. ———, and P. A. Bisson. 1997. Degradation and loss of anadromous debris in temperate ecosystems. Advances in Ecological Research
salmonid habitat in the Pacific Northwest. Pages 277-214 in D. J. 15:133-302.
Stouder, P. A. Bisson, and R. J. Naiman, editors. Pacific salmon & their 165. Harrod, R. J., W. L. Gaines, W. E. Hartl, and A. Camp. 1998. Estimating
ecosystems. Chapman and Hall, New York, NY. historical snag density in dry forests east of the Cascade Range. U.S.
145. ———, F. J. Swanson, W.A. McKee, and K. W. Cummins. 1991. An Forest Service, Pacific Northwest Research Station, Portland, Oregon,
ecosystem perspective of riparian zones. BioScience 41:540-551. General Technical Report PNW-GTR-428.
146. Grier, C. C. 1978.A Tsuga heterophylla-Picea sitchensis ecosystem of 166. Hartmann, P., M. Scheitler, and R. Fischer. 1989. Chapter 2-C in E.D.
coastal Oregon: decomposition and nutrient balances of fallen logs. Schulze, O. L. Lange, and R. Oren, editors. Forest decline and pollution.
Canadian Journal of Forest Research 8:198-206. Springer-Verlag, New York, NY.
147. ———, and R. S. Logan. 1978. Old-growth Pseudotsuga menziesii 167. Harvey, A.E. 1995. Soil and the forest floor: what it is, how it works,
communities of a western Oregon watershed: biomass distribution and and how to treat it. Nat. Res. News (5)1. Blue Mountains National
production budgets. Ecological Monographs 47:373-400. Research Institute, La Grande, OR. U.S. Deptartment of Agriculture,
148. ———, K. M. Lee, and N. M. Nadkarni et al. 1989. Productivity of Forest Service, Pacific Northwest Research Station.
forests of the United States and its relation to soil and site factors and 168. ———, M. J. Larsen, and M. F. Jurgensen. 1976. Distribution of
management practices: a review. U.S. Forest Service, Pacific Northwest ectomycorrhizae in a mature Douglas-fir/larch forest soil in western
Research Station, Portland, OR. General Technical Report PNW-GTR- Montana. Forest Science 22:393-398.
222. 169. ———, M. F. Jurgensen, R. T. Graham. 1987. The role of woody
149. Hagan, J. M. and S. L. Grove. 1999. Coarse woody debris. Journal of residues in soils of a symposium. Washington State University, Pullman,
Forestry 97(1):6-11. WA.
150. Hagar, J. C., W. C. McComb, and W. H. Emmingham. 1996. Bird 170. ———, J. M. Geist, G. I. McDonald, M. F. Jurgensen, P. H. Cochran, D.
communities in commercially thinned and unthinned Douglas-fir stands Zabowski, and R. T. Meurisse. 1994. Biotic and abiotic processes in
of western Washington. Wildlife Society Bulletin 24(2):353-366. eastside ecosystems: the effects of management on soil properties,
151. Halpern, C. B., and T. A. Spies. 1995. Plant species diversity in natural processes, and productivity. U.S. Forest Service, Pacific Northwest
and managed forests of the Pacific Northwest. Ecological Applications Research Station, Portland, OR. General Technical Report PNW-GTR-
5(4):913-934. 323.
 CHAPTER 24: WOOD LEGACIES 617

171. Hawksworth, F. G. and D. Wiens. 1996. Dwarf mistletoes: biology, resource management. Ecological Applications 1:66-84.
pathology and systematics. U.S. Forest Service, Agricultural Handbook 192. ———, K. D. Fausch, P. L. Angermeier, P. R.Yant, and I. J. Schlosser.
No. 709, Washington, D.C. 1986. Assessing biological integrity in running waters: a method and its
172. Haynes, R. W., R. J. Alig, and E. Moore. 1994. Alternative simulations of rationale. Illinois Natural History Survey, Special Publication 5,
forestry scenarios involving carbon sequestration options: investigation Champaign, IL.
of impacts on regional and national timber markets. U.S. Forest Service, 193. Kauffman, J. B. 1990. Ecological relationships of vegetation and fire in
Pacific Northwest Research Station, General Technical Report PNW- Pacific Northwest Forests. Pages 39-54 in J. D. Walstad, S. R. Radosevich,
GTR-335, Portland, OR. and D.V. Sandberg, editors. Natural and prescribed fire in Pacific
173. Heede, B. H. 1972. Influences of a forest on the hydraulic geometry Northwest forests. Oregon State University Press, Corvallis, OR.
of two mountain streams. Water Resources Bulletin 8:523-529. 194. Kauffman, M. R. 1996. To live fast or not: growth, vigor, and longevity
174. Hedin, L. O., J. J. Armesto, and A. H. Johnson. 1995. Patterns of of old-growth ponderosa pine and lodgepole pine trees. Tree
nutrient loss from unpolluted temperate old-growth forests: evaluation Physiology 16:139-144.
of biogeochemical theory. Ecology 76:493-509. 195. Kauffman, J. B. and R. E. Martin. 1985. A preliminary investigation on
175. Hepting, G. H. 1971. Diseases of forest and shade trees of the the feasibility of preharvest prescribed burning for shrub control. Pages
United States. USDA Forest Service Agricultural Handbook 386. 89-114 in Proceedings of The 6th Forest Vegetation Management
176. Hergert, H. L. and E. F. Kurth. 1952. The chemical nature of cork Conference, Redding, CA.
from Douglas-fir bark. Tappi 35: 59-66. 196. Keenan, R. J., and J. P. Kimmins. 1993. The ecological effects of clear-
177. Hessburg, P. F., R. G. Mitchell, and G. M. Filip. 1994. Historical and cutting. Environmental Review 1:121-144.
current roles of insects and pathogens in eastern Oregon and forested 197. Kellomaki, S., and T. Karjalainen. 1995. Sequestration of carbon in
landscapes. U.S. Forest Service, Pacific Northwest rearch Station, the Finnnish boreal forest ecosystem managed for timber production.
Portland OR. General Technical Report PNW-GTR-327. Pages 59-68 in M. J. Apps and D.T. Price, editors. Forest ecosystems,
178. ———, B. G. Smith, and R. Brion Salter. 1999. Detecting change in forest management and the global carbon cycle. Springer, New York,
forest spatial patterns from reference conditions. Ecological NY.
Applications 9(4):1232-1252. 198. Kimmins, J. P. 1997. Forest ecology: A foundation for sustainable
179. Hixson, S. E., R. F. Walker, and C. M. Skau. 1990. Soil denitrification management, 2nd edition. Prentice Hall, NJ.
rates in four subalpine plant communities of the Sierra Nevada. Journal 199. ———. 1997. Balancing act. Environmental issues in forestry,
of Environmental Quality 19:717-620. second edition. UBC Press, Vancouver, B.C., Canada.
180. Holsten, E. H., R. A. Werner, and T. H. Laurent. 1980. Insects and 200. Koehler, G. M., and K. B. Aubrey. 1994. Pages 74-98 in L. F. Ruggiero,
diseases of Alaskan forests. USDA Forest Service, Alaska Region report K. B. Aubrey, S. W. Buskirk, L. J. Lyon, and W. J. Zielinski, editors. The
No. 75. scientific basis for conserving forest carnivores: American marten,
181. Holthausen, R. S., M. G. Raphael, K. S. McKelvey, E. D. Forsman, E. E. fisher, lynx, and wolverine. U.S. Forest Service, Rocky Mountain Forest
Starkey, and D. Erran Seaman. 1996. The contribution of federal and and Range Experiment Station, Fort Collins, CO. General Technical
non-federal habitat to persistence of the Northern Spotted Owl on Report RM-GTR-254.
the Olympic Peninsula, Washington: report of the reanalysis team. U.S. 201. Kohm, K. A., J. F. Franklin. 1997. Creating a forestry for the 21st
Forest Service, Pacific Northwest Research Station, Portland, OR. century: The science of ecosystem management. Island Press,
General Technical Report PNW-GTR-352. Washington D.C.
182. Hope, S. M. 1987. Classification of decayed Abies Amabalis logs. 202. Korol, J. J., M. A. Hemstrom, W. J. Hann, and R. A. Gravenmier. In
Canadian Journal of Forest Research 17:559-564. press. Snags and down wood in the Interior Columbia Basin Ecosystem
183. Huggard, D. J. 1999. Static life-table analysis of fall rates of subalpine Management Project. In P. J. Shea and W. Laudenslayer, editors.
fir snags. Ecological Applications 9(3):1009-1016. Proceedings of a conference on the ecology and management of dead
184. Hulme, M. A. and J. K. Shields. 1970. Biological control of decay fungi wood in western forests, Reno, Nevada. USDA Forest Service, Pacific
in wood by competition for non-structural carbohydrates. Nature Southwest Research Station General Technical Report.
227:300-301. 203. Kurth, E. F. 1948. The chemical analysis of western woods. Part I.
184a. Hunter, M. 1995. Residual trees as biological legacies. Management Paper Trade. Journal 126:56-58.
Communique no. 2. Corvallis, Oregon. Cascade Center for Ecosystem 204. ———, and F. L. Chan. 1953. Extraction of tannin and
Management. dihydroquercetin from Douglas-fir bark. Journal of American Leather
185. Ingham, E. R. 1995. Organisms in the soil: the functions of bacteria, Chemical Association 48:20-32.
fungi, protozoa, nematodes, and arthropods. National Research News 205. Lambert, R. L., G. E. Lang, and W. A. Reiners. 1980. Loss of mass and
(5)1. Blue Mountains Nat. Res. Institute, La Grande, OR. U.S. Forest chemical change in decaying boles of a subalpine fir forest. Ecology
Service, Pacific Northwest Research Station. 61:1460-1473.
186. ———, and A. R. Moldenke. 1995. Microflora and microfauna on 206. Larsen, M. J., M. F. Jurgensen, and A. E. Harvey. 1978. N2 fixation
stems and trunks: diversity, food webs, and effects on plants. Pages 241- associated with wood decayed by some common fungi in western
256 in B. L. Gartner editor. Plant stems: physiology and functional Montana. Canadian Journal of Forest Research 8:341-345.
morphology. Academic Press, New York, NY. 207. Lattin, J.D. 1990. Arthropod diversity in Northwest old-growth
187. Johnson, C. G., Jr., R. R. Clausnitzer, P. J. Mehringer, and C. D. Oliver. forests. Wings 15 (2):7-10.
1994. Biotic and abiotic processes of eastside ecosystems: the effects of 208. Law, B. E. and R. H. Waring. 1994. Combining remote sensing and
management on plant and community ecology, and on stand and climatic data to estimate net primary production. Ecological
landscape vegetation dynamics. U.S. Forest Service, Pacific Northwest Applications 4:717-728.
Research Station, Portland, OR. General Technical Report PNW-GTR- 209. Lee, K. N. 1999. Appraising adaptive management. Conservation
322. Ecology 3(2):1-19.
188. Johnson, D. H., and T. A. O’Neil. 2001. Wildlife-habitat relationships in 210. Lehmkuhl, J. F., S. D. West, C. C. Chambers, W. C. McComb, D. A.
Oregon and Washington. Oregon State University Press, Corvallis, OR. Manuwal, K. B. Aubry, J. L. Erickson, R. A. Gitzen, and M. Leu. 1999. An
189. Johnson, D.W. 1992. Nitrogen retention in forest soils. Journal of experiment for assessing vertebrate response to varying levels and
Environmental Quality 21:1-12. patterns of green tree retention. Northwest Science 73: 45-55.
190. Jones, J. A. and G. E. Grant. 1996. Peak flow responses to clear- 211. Lehmkuhl, J. F., P. F. Hessburg, R. L. Everett, M. H. Huff, and R. D.
cutting and roads in small and large basins, western Cascades, Oregon.
Water Resources Research 32: 959-974.
191. Karr, J. R. 1991. Biological integrity: a long-neglected aspect of water
 618 Wildlife–Habitat Relationships in Oregon and Washington

Ottmar. 1994. Historical and current forest landscapes of eastern 229. ———. 1994. Sustainable forestry, philosophy, science, and
Oregon and Washington. Part 1: vegetation pattern and insect and economics. St. Lucie Press. Delray Beach, FL.
disease hazards. U.S. Forest Service, Pacific Northwest rearch Station, 230. ———, and J. M. Trappe, technical editors. 1984. The seen and
Portland, OR. General Technical Report PNW-GTR-328. unseen world of the fallen tree.U.S. Forest Service, Pacific Northwest
212. Lewis, J. C. 1998. Creating snags and wildlife trees in commercial Research Station, Portland, OR. General Technical Report, PNW-GTR-
forest landscapes. Western Journal of Applied Forestry 13(3):97-101. 164.
213. Little, S. N. and J. L. Ohmann. 1988. Estimating nitrogen lost from 231. ———, J. M. Trappe, and R. A. Nussbaum. 1978. Fungal-small mammal
forest floor during prescribed fires un Douglas-fir/ western hemlock interrelationships with emphasis on Oregon coniferous forests. Ecology
clearcuts. Forest Science 34:152-164. 59(6):799-809.
214. Lofroth, E. C. 1993. Scale dependent analyses of habitat selection by 232. ———, ———, and D. Ure. 1978. Implications of small mammal
marten in the sub-boreal spruce biogeoclimatic zone, British Columbia. mycophagy to the management of western coniferous forests. Pages
M. S. Thesis, Simon Fraser University, Burnaby, B. C., Canada 78-88 in Transactions of the 43rd North American Wildlife and Natural
215. Lutes, D. C. 1999. A comparison of methods for the quantification of Resources Conference.
coarse woody debris and identification of its spatial scale; a study from 233. ———, R. F. Tarrant, J. M. Trappe, and J. F. Franklin. 1988. From The
the Tenderfoot Creek Experimental Forest, Montana. M. S. thesis. Forest To The Sea: A Story Of Fallen Trees. USDA Forest Service,
University of Montana, Missoula, MT. Pacific Northwest Research Station, Portland, OR. General Technical
216. Machmer, M. M., and C. Steeger. 1995. The ecological roles of wildlife Report PNW-GTR-229.
tree users in forest ecosystems. Land Management Handbook No. 35. 234. ———, R. G. Anderson, K. Cromack, Jr., J. T. Williams, and R. E.
Ministry of Forests, Victoria, B.C., Canada. Martin. 1979. Pages 78-95 in J. W. Thomas, editor. Dead and down
217. Maclean, C. D. 1979. Relative density: the key to stocking assessment woody material. Wildlife Habitats in Managed Forests; the Blue
in regional analysis—a Forest Survey viewpoint. U.S. Forest Service, Mountains of Oregon and Washington. USDA Forest Service,
Pacific Northwest rearch Station, Portland, OR. General Technical Washington, D.C.
Report PNW-78. 235. Mason, D. T. and J. Koon. 1985. Habitat values of woody debris
218. MacMillan, P., J. Means, G. M. Hawk, K. Cromack, Jr., and R. Fogel. accumulations of the lower Stehekin River, with notes on disturbances
1977. Log decomposition in an old-growth Douglas-fir forest. of alluvial gravels. Final report to the National Park Service, Contract
Northwest Science. Assoc. Program and Abstract. at 50th annual mtg. CX-9000-3-8066. Fairhaven College, Western Washington University,
Pullman, WA, Washington State University Press, WA. Bellingham, WA.
219. Magill, A. H., J. D. Aber, J. J. Hendricks, R. D. Bowden, J. M. Melillo, and P. 236. Mason, R. R., B. E. Wickman, R. C. Beckwith, and H. G. Paul. 1992.
A. Steudler. 1997. Biochemical response of forest ecosystems to Thinning and nitrogen fertilization in a grand fir stand infested with
simulated chronic nitrogen deposition. Ecological Applications western spruce budworm. Part I. Insect Response. Forest Science 38:
7(2):402-415. 235-251.
220. Manion, P. D. 1981. Tree Disease Concepts, second edition. Prentice- 237. Matson, P. A. and R. H. Waring. 1984. Effects of nutrient and light
Hall. Englewood Cliffs, NJ. limitation on mountain hemlock: susceptibility to laminated root rot.
221. Marcot, B. G. 1992. Snag Recruitment Simulator, Rel. 3.1 [computer Ecology 65:1517-1524.
program]. USDA Forest Service, Pacific Northwest Region, Portland, 238. McClelland, B. R., S. S. Frissell, S. S., W. C. Fischer, and C. H. Halvorsen.
OR. 1979. Habitat management for hole-nesting birds in forests of western
222. ———. 1997. Biodiversity of old forests of the west: A lesson from larch and Douglas-fir. Journal of Forestry 77: 480-483.
our elders. Pages 87 - 105 in K. A. Kohm, and J. F. Franklin, editors, 239. McComb, W. C., J. Tappeiner, L. Kellogg, C. Chambers, R. Johnson.
Creating a forestry for the 21st Century. The science of ecosystem 1994. Stand management alternatives for multiple resources: integrated
management. Island Press, Washington, D.C. management experiments. Pages 71-86 in M. H. Huff, L. K. Norris, J. B.
223. ———. In press. An ecological functional basis for managing Nyberg, and N. L. Wilkin, coordinators. Expanding horizons of
decaying wood for wildlife. In: P. J. Shea and W. Laudenslayer, editors. ecosystem management. U.S. Forest Service, Pacific Northwest
Proceedings of a Conference on The Ecology and Management of Dead Research Station, Portland, Oregon. General Technical Report PNW-
Wood in Western Forests, Reno, Nevada. USDA Forest Service, Pacific GTR-336.
Southwest Research Station General Technical Report. 240. ———, and J. L. Ohmann. 1996. Snag Dynamics Projection Model
224. ———, and M. Vander Heyden. 2001. Key ecological functions of (SDPM), [computer program], USDA Forest Service, Pacific Northwest
wildlife species. Pages 168-186 in: D. Johnson and T. O’Neil, managing Research Station, Corvallis, OR.
directors. Wildlife-habitat relationships in Oregon and Washington. 241. McDade, M. H., F. J. Swanson, W. A. McKee, and J. F. Franklin. 1990.
Oregon State University Press, Corvallis OR. Source distances for coarse woody debris entering small streams in
225. ———, M. A. Castellano, J. A. Christy, L. K. Croft, J. F. Lehmkuhl, R. H. western Oregon and Washington. Canadian Journal of Forest Research
Naney, R. E. Rosentreter, R. E. Sandquist, and E. Zieroth. 1997. Terrestrial 20:326-329.
ecology assessment. Pages 1498-1713 in Quigley, T. M, and S. J. 242. McFee, W. W. and E. L. Stone. 1966. The persistence of decaying wood
Arbelbide, technical editors. An assessment of ecosystem components in the humus layers of northern forests. Soil Science Society American
in the interior Columbian basin and portions of the Klamath and Great Proceedings. 30:513-516.
Basins. U.S. Forest Service, Pacific Northwest rearch Station, Portland, 243. McIntosh, B. A., J. R. Sedell, J. E. Smith, R. C. Wissmar, S. E. Clarke, G. H.
OR. General Technical Report PNW-GTR-405. Reeves, and L. A. Brown. 1994. Management history of eastside
226. ———, K. Mellen, S. A. Livingston, and C. Ogden. In press. The ecosystems: changes in fish habitat over 50 years, 1935 to 1992. U.S.
DecAID advisory model: wildlife component. In: P. J. Shea and W. Forest Service, Pacific Northwest rearch Station, Portland, OR. General
Laudenslayer editors. Proceedings of a conference on The Ecology and Technical Report PNW-GTR-321.
Management of Dead Wood in Western Forests, Reno, Nevada. USDA 244. McKee, A., G. Laroi, and J. F. Franklin. 1982. Structure, composition,
Forest Service, Pacific Southwest Research Station General Technical and reproduction behavior of terrace forests, South Fork Hoh River,
Report. Olympic National Park. Pages 22-29 in E. E. Starkey, J. F. Franklin, and J.
227. Marra, J. L. and R. L. Edmonds. 1994. Coarse woody debris and forest W. Matthews, editors. Ecological research in national parks of the
floor respiration in an old-growth coniferous forest on the Olympic Pacific Northwest. Forest Research Laboratory, Oregon State
Peninsula, Washington. Canadian Journal of Forest Research 24:1811- University, Corvallis, OR.
1817. 245. McNabb, D. H. and K. Cromack, Jr. 1990. Effects of prescribed fire on
228. Maser, C. 1988. The redesigned forest. R&E Miles, San Pedro, CA.
 CHAPTER 24: WOOD LEGACIES 619

nutrients and soil productivity. Pages 125-142 in J. D. Walstad, S. R. D.Taylor, and F. F. Wagner. 1985. Snags (wildlife trees). Pages 129-169 in
Radosevich, and D.V. Sandberg, editors. Natural and prescribed fire in E. R. Brown, technical editor. Management Of Wildlife And Fish Habitats
Pacific Northwest forests. Oregon State University Press, Corvallis, In Forests Of Western Oregon and Washington. USDA Forest Service,
OR. Pacific NW Region, Portland, OR. Publication No. R6-F&WL-192-1985.
246. Mellen, T. K., and A.. Ager. 1998. Coarse Wood Dynamics Model 264. Nihlgard, B. 1985. The ammonium hypothesis—an additional
(CWDM), [computer model], USDA Forest Service, Pacific Northwest explanation to the forest dieback in Europe. Ambio 14:2-8.
Region, Portland, OR. 264a. NIOSH. 1995. National Institute for Occupational Safety and
247. ———, and A. Ager. In press. A coarse wood dynamics model for Health. Preventing injuries and deaths of loggers. US Dept. of Health
the western Cascades. In P. J. Shea and W. Laudenslayer, editors). The and Human Services, Public Health Service, Centers For Disease
Ecology and Management of Dead Wood in Western Forests. Reno, Control and Prevention, National Institute for Occupational Safety and
NV. Health, Fatality Assessment and Control Evaluation (FACE) Report No.
248. ———, B. G. Marcot, J. L. Ohmann, K. L. Waddell, E. A. Willhite, B. B. 95-101.
Hostetler, S. A. Livingson, and C. Ogden. In press. DecAID: A decaying 265. Noble, W. O., E. C. Meslow, M. D. Pope. 1990. Denning habits of black
wood advisory model for Oregon and Washington. In: P. J. Shea and W. bears in the central Coast Range of Oregon. Oregon State University,
Laudenslayer, editors. Proceedings of a conference on The Ecology and Department of Fisheries and Wildlife, Corvallis, OR.
Management of Dead Wood in Western Forests, Reno, Nevada. USDA 266. Nommik, H. and K. Vahtras. 1982. Retention and fixation of
Forest Service, Pacific Southwest Research Station General Technical ammonium and ammonia in soils. Pages 123-171 in F. J. Stevenson,
Report. editor. Nitrogen In Agricultural Soils. American Society of Agronomy.
249. Moldenke, A. R. 1990. One hundred twenty thousand little legs. Madison, WI.
Wings 15(2):11-14. 267. North, M. and J. F. Franklin. 1990. Post-disturbance legacies that
250. ———, and J. D. Lattin. 1990. Density and diversity of soil enhance biological diversity in a Pacific Northwest old-growth forest.
arthropods as “biological probes” of complex soil phenomena. The Northwest Environmental Journal 6:427-429.
Northwest Environmental Journal 6:409-410. 268. ———, J. Chen, G. Smith, L. Krakowiak, and J. F. Franklin. 1996. Initial
251. Molina, R. and J. M. Trappe. 1982. Patterns of ectomycorrhizal host response of understory plant diversity and overstory tree diameter
specificity and potential among the Pacific Northwest conifers and growth to a green tree retention harvest. Northwest Science 70(1): 24-
fungi. Forest Science 28:423-458. 35.
252. Moore, K. M. S. and S. V. Gregory. Response of young-of-the-year 269. ———. 1993. Stand structure and truffle abundance associated with
cutthroat trout to manipulation of habitat structure in a small stream. northern spotted owl habitat. Ph.D. thesis, University of Washington,
Transactions of the American Fisheries Society 117:162-170. Seattle, WA.
253. Morrison, M. L., and M. G. Raphael. 1993. Modeling the dynamics of 270. Nowacki, G. J. and M. G. Kramer. 1998. The effects of wind
snags. Ecological Applications 3(2):322-330. disturbance on temperate rainforest structure and dynamics of
254. Moseley, M. P. 1981. The influence of organic debris on channel southeast Alaska. USDA Forest Service, Pacific Northwest Research
morphology and bedload transport in a New Zealand forest stream. Station, Portland, OR. General Technical Report, PNW-GTR-42.
Earth Surface Processes and Landforms 6:571-579. 271. NRC (National Research Council). 1990. Forestry Research: A
255. Mote, P. 1999. Impacts of climate variability and change. National Mandate For Change. National Academy Press, Washington, D.C.
Oceanic And Atmospheric Administration, Office of Global Programs, 272. ———. 1998. Changing conditions of the forest. Pages 57-71 in
University of Washington, JISAO/SMA Climate Impacts Group Forested landscapes in perspective. Prospects and opportunities for
Contribution #715, Seattle, WA. sustainable management of America’s nonfederal forests. Committee
256. Mowrey, R. A., and J. C. Zasada. 1984. Den tree use and movements on prospects and opportunities for sustainable management of
of northern flying squirrels in interior Alaska and implications for forest America’s nonfederal forests. National Academy Press, Washington, D.
managers. Pages 351-356 in W. R. Meehan, T. A. Hanley, and A. Thomas, C.
editors. Fish and wildlife relationships in old-growth forests: 273. Odum, E. P. 1971. Fundamentals of ecology. W. B. Saunders Co.,
proceedings of a symposium: 1982 April 12-14 in Juneau, Alaska. Philadelphia, PA.
American Institute of Fishery Research Biologists. 274. Ohmann, J. L., and T. A. Spies. 1998. Regional gradient analysis and
257. Murphy, M. L., K. V. Koski, J. Heifetz, S. W. Johnson, D. Kirchofer, and J. F. spatial pattern of woody plant communities of Oregon forests.
Thedinga. 1985. Role of large organic debris as winter habitat for Ecological Monographs 68:151-182.
juvenile salmonids in Alaska streams. Proceedings, W. Association of 275. ———, W. C. McComb, and A. A. Zumrawi. 1994. Snag abundance
Fish and Wildlife Agencies 1984:251-262. for cavity-nesting birds on nonfederal lands in Oregon and Washington.
258. Nabuurs, G. J. and G. M. J. Mohren. 1993. Carbon fixation through Wildlife Society Bulletin 22:607-620.
forestation activities. A study of the carbon sequestration potential of 276. ———, and K. L. Waddell. In press. Regional patterns of dead wood
selected forest types, commissioned by the FACE Foundation. IBN in forested habitats of Oregon and Washington. In: P. J. Shea and W.
Natural resource report 93/4. FACE/Institute for Forestry and Nature Laudenslayer, editors. Proceedings of a conference on The Ecology and
Research (IBN-DLO), Arnhem, Wageningen, The Netherlands. Management of Dead Wood in Western Forests, Reno, Nevada. USDA
259. Naiman, R. J. and J. R. Sedell. 1979. Benthic organic matter as a Forest Service, Pacific Southwest Research Station General Technical
function of stream order in Oregon. Archiv für Hydrobiologie 87:404- Report.
422. 277. Oliver, C. D., and B. C. Larson. 1996. Forest stand dynamics. John
260. ———, K. L. Fetherston, S. J. McKay, and J. Chen. 1998. Riparian Wiley and Sons, Inc.
forests. Pages 289-323 in R. J. Naiman and R. E. Bilby, editors. River 278. ———, D. R. Berg, D. R. Larsen, and K. L. O’Hara. 1992. Integrating
ecology and management. Springer, New York, NY. management tools, ecological knowledge, and silviculture. Pages 361-
261. ———, and R. E. Bilby. 1998. River ecology and management. 382 in R.J. Naiman, editor. Watershed Management. Balancing
Lessons from the Pacific coastal region. Springer, New York, NY. sustainability and environmental change. Springer Verlag, NY.
262. ———, T. J. Beechie, L. E. Benda, D. R. Berg, P. A. Bisson, L. H. 279. ———, L. L. Irwin, W. H. Knapp. 1994. Eastside forest management:
MacDonald, M. D. O’Connor, P. L. Olson, and E. A. Steel. 1992. historical overview, extent of their applications, and their effects on
Fundamental elements of ecologically healthy watersheds in the Pacific sustainability of ecosystems. U.S. Forest Service, Pacific Northwest
Northwest coastal ecoregion. Pages 127-18 in R. J. Naiman, editor. Research Station, Portland, OR. General Technical Report PNW-GTR-
Watershed Management: Balancing Sustainability and Environmental 324.
Change. Springer-Verlag, New York, NY. 280. Olson, R. K., D. Binkley, and M. Bohm. 1992. The Responses of
263. Neitro, W. A.,V. W. Binkley, S. P. Cline, R. W. Mannan, B. G. Marcot,
 620 Wildlife–Habitat Relationships in Oregon and Washington

Western Forests to Air Pollution. Ecological Studies Vol. 97. Springer- impacts on long-term productvity. Pages 127-145 in W. J. Dyck and C. A.
Verlag, New York, NY. Mees, editors. Impact of intensive harvesting on forest site productivity.
281. O’Neill, R.V., W.F. Harris, B. S. Ausmus, and D. E. Reichle. 1975. A Proceedings IEA/BE A3 Workshop. South Island, New Zealand, March
theoretical basis for ecosystem analysis with particular reference to 1989. Forest Research Institute, Rotorua, New Zealand, FRO Bulletin
element cycling. Pages 28-40 in F. G. Howell, J. B. Gentry, and M. H. No. 159.
Smith editors. Mineral Cycling in Southeastern Ecosystems. ERDA 300. Prescott, C. E., J. P. Corbin, and D. Parkinson. 1989. Input,
Symposium Series (Conf-740513). accumulation, and residence times of carbon, nitrogen, and phosphorus
282. Parks, C. G. 1996. Bear Trees - An Eastern Oregon Landscape Legacy. in four Rocky Mountain coniferous forests. Canadian Journal of Forest
USDA Forest Service, Pacific Northwest Research Station, Portland, Research 19:489-498.
OR. 301. Putz, F. E., P. D. Coley, K. Lu, A. Montalvo, and A. Aiello. 1983.
283. ———, and D. C. Shaw. 1996. Death and decay: a vital part of living Uprooting and snapping of trees: structural determinants and
canopies. Northwest Science. 70:46-53. ecological consequences. Canadian Journal of Forest Research
284. ———, E. Bull, G. M. Filip, and R. L. Gilbertson. 1996. Wood decay 13:1011-1020.
fungi associated with woodpecker nest cavities in living western larch. 302. Quigley, T. M., R. W. Haynes, and R. T. Graham, technical editors. 1996.
Plant Disease. 80:959. Integrated scientific assessment for ecosystem management in the
285. ———, ———, and T. R. Torgersen. 1997. Field guide to the interior Columbia basin and portions of the Klamath and Great Basins.
identification of snags and logs in the interior Columbia River basin. U.S. Forest Service. Pacific Northwest Research Station, Portland, OR.
U.S. Forest Service, Pacific Northwest rearch Station, Portland, OR. General Technical Report PNW-GTR-382.
General Technical Report PNW-GTR-390. 303. Quinn, T. P. and R. F. Tallman. 1987. Seasonal environmental
286. ———, D. A. Conklin, L. Bednar, and H. Maffei. 1999. Woodpecker predictability in riverine fishes. Environmental Biology of Fishes 18: 155-
use and fall rates of snags created by killing ponderosa pine infected 159.
with dwarf mistletoe. U.S. Forest Service, Pacific Northwest rearch 304. Raedecke, K. J. 1988. Streamside management: Riparian wildlife and
Station, Portland, OR. General Technical Report PNW-GTR-515. forestry interactions. Institute of Forest Resources, Contribution 59,
287. Parmeter, J. R. 1978. Forest stand dynamics and ecological factors in Univ. of Washington, Seattle, WA.
relation to dwarf mistletoe spread, impact, and control. Pages 16-31 in: 305. Raphael, M. G. and L. C. Jones. 1997. Characteristics of resting and
Proceedings of the Symposium on Dwarf Mistletoe Control Through denning sites of American marten in central Oregon and western
Forest Management. Pacific Southwest Research Station. PSW-GTR-31. Washington. Pages 146-165 in G. Proulx, H. N. Bryant, and P. M.
Berkeley, CA. Woodard, editors. Martes: taxonomy, ecology, techniques and
288. Paulus, W. and A. Bresinsky. 1989. Soil fungi and other management. Provincial Museum of Alberta, Edmonton, Alberta,
microorganisms. Chapter 2-A in E.-D. Schulze et al. editors. Forest Canada.
decline and pollution. Springer-Verlag, New York, NY. 306. ———, and M. L. Morrison. 1987. Decay and dynamics of snags in
288a. Pecore. M. 1992. Menominee sustained yield management: a the Sierra Nevada, California. Forest Science 33(3):774-783.
successful land ethic in practice. Journal of Forestry 90(7):12-16. 307. ———, and M. White. 1984. Use of snags by cavity-nesting birds in
289. Perlin, J. 1989. A forest journey: the role of wood in the the Sierra Nevada. Wildlife Monograph 86: 1-66.
development of civilization. Harvard University Press, Cambridge, MA. 308. Rastetter, E. B., R. B. McKane, G. R. Shaver, and J. M. Melillo. 1992.
290. Perry, D. A. 1994. Forest Ecosystems. John Hopkins University Press, Changes in C storage by terrestrial ecosystems: How C-N interactions
Baltimore and London. restrict responses to CO2 and temperature. Water, Air, And Soil
291. ———, and M. P. Amaranthus. 1997. Disturbance, recovery, and Pollution. 64:327-344.
stability. Pages 31-67 in K. A. Kohm, and J. F. Franklin, editors. Creating a 309. ———, G. I. Agren, and G. R. Shaver. 1997. Responses of N-limited
forestry for the 21st century. Island Press, Washington, D.C. ecosystems to increased CO2: A balanced-nutrition, coupled-element-
292. ———, and J. Maghembe. 1989. Ecosystem concepts and current cycles model. Ecological Applications 7(2): 444-460.
trends in forest management: time for reappraisal. Forest Ecology and 310. Rayner, A. D. M. and L. Boddy. 1988. Fungal decomposition of wood,
Management 26:123-140. its biology and ecology. John Wiley & Sons, New York, NY.
293. ———, M. P. Amaranthus, J. G. Borchers, S. L. Borchers, and R. E. 311. Rose, C. R. and P. S. Muir. 1997. Green-tree retention: consequences
Brainerd. 1989. Bootstrapping in ecosystems. Bioscience 39:230-236. form timber production in forests of the western Cascades, Oregon
294. ———, J. G. Borchers, D. P. Turner, S. V. Gregory, C. R. Perry, R. K. Ecological Applications 7(1): 209-217.
Dixon, S. C. Hart, B. Kauffman, R. P. Neilson, and P. Sollins. 1991. 311a. ———. 1994. Relationships of green-tree retention following
Biological feedbacks to climate change: terrestrial ecosystems as sinks timber harvest to forest growth and species composition in the
and sources of carbon and nitrogen. The Northwest Environmental western Cascade Mountains. M.S. Thesis. Oregon State University,
Journal 7:203-232. Corvallis.
295. Peterjohn, W. T., and D. L. Correll. 1984. Nutrient dynamics in an 312. Regional Forest Nutrition Research Project and Stand Management
agricultural watershed: observations on the role of a riparian forest. Cooperative (RFNRP/SMC) 1989. Planning for 2001: Research in Forest
Ecology 65:1466-1475. Nutrition, Silviculture, and Wood Quality in the 21st Century.
296. Powell, D. S., J. L. Faulkner, D. R. Darr, Z. Zhu, and D. W. MacCleery,. Proceedings of a Joint Review And Planning Meeting at Pack Forest.
1993. Forest resources of the United States, 1992. General Technical November, 1989.
Report RM-234. U.S. Department of Agriculture, Forest Service. 313. Ruggiero, L. F., and K. B. Aubry. 1994. The scientific basis for
297. Powell, R. A., and W. J. Zielinski. 1994. Fisher. Pages 38-74 in L. F. conserving forest carnivores. U.S. Forest Service, Rocky Mountain
Ruggiero and K. B. Aubrey, technical editors. The scientific basis for Forest and Range Experiment Station, Fort Collins, CO. General
conserving forest carnivores. U.S. Forest Service, Rocky Mountain Technical Report RM-GTR-254.
Forest and Range Experiment Station. Fort Collins, CO. General 314. ———, ———, A. B. Cary, and M. H. Huff. 1991. Wildlife and
Technical Report, RM-GTR-254. vegetation of unmanaged Douglas-fir forests. USDA Forest Service,
298. Powers, R. F. 1990. Do timber management practices degrade long- Pacific Northwest Research Station, Portland, OR. General Technical
term site productivity?: what we know and what we need to know. Report PNW-GTR-285.
Pages 87-106 in Proceedings of the Eleventh Annual Forest Vegetation 315. Runyon, J., R. H. Waring, S. N. Goward, and J. M. Welles. 1994.
Management Conference, November 8-9, 1989. Environmental limits on net primary production and light-use efficiency
299. ———. 1990. A soils research approach to evaluating management across the Oregon transect. Ecological Applications 4(2): 226-237.
316. Ruth, R. H., and R. A.Yoder. 1953. Reducing wind damage in the
 CHAPTER 24: WOOD LEGACIES 621

forests of the Oregon Coast Range. Tree. U.S. Forest Service, Pacific management studies. BC Ministry of Forests, Land Management
Northwest Research Station, Portland, OR. Research Paper, PNW-RP-7. Handbook No. 2.Victoria, BC, Canada.
317. Ryan, M. G., D. Binkley, and J. H. Fownes. 1997. Age-related decline in 339. Sollins, P. 1982. Input and decay of coarse woody debris in
forest productivity: pattern and process. Advances in Ecological coniferous stands in western Oregon and Washington. Canadian
Research 27:213-262. Journal of Forest Research 12: 18-28.
318. ———, and B. J. Yoder. 1997. Hydraulic limits to tree height and tree 340. ———, S. P.Cline, T.Verhoeven, D. Sachs, and G. Spycher. 1987.
growth. BioScience 47(4): 235-242. Patterns of log decay in old-growth Douglas-fir forests. Canadian
319. Sachs, D. L., and P. Sollins. 1986. Potential effects of management Journal of Forest Research 17(12):1585-1595.
practices on nitrogen nutrition and long-term productivity of western 341. ———, C. C. Grier, F. M. McCorrison, K. Cromack, Jr., R. Fogel, and R.
hemlock stands. Forest Ecology and Management 17:25-36. L. Fredricksen. 1980. The internal element cycles of an old-growth
320. Society of American Foresters (SAF). 1993. Task Force report on Douglas-fir forest in western Oregon. Ecological Monographs
sustaining long-term forest health and productivity. Society of 50(3):261-285.
American Foresters, Bethesda, MD. 342. Sperry, J. S. 1995. Limitations on water stem transport and their
321. Salo, E. O., and T.W. Cundy, editors. 1987. Streamside management: consequences. Pages 105-125 in B. L. Gartner editor. Plant stems:
Forestry and fishery interactions. Institute of Forest Resources, Physiology and functional morphology. Academic Press, New York, NY.
Contribution 57. University of Washington, Seattle, WA. 343. Spies, T. A. 1997. Forest stand structure, composition, and function.
322 Scharpf, R. F. 1993. Diseases of Pacific coast conifers. U.S. Forest Pages 11-30 in K. A. Kohm, and J. F. Franklin, editors. Creating a forestry
Service, Pacific Southwest Research Station, Washington, D.C. for the 21st century. Island Press, Washington, D.C.
Agriculture Handbook No 521. 344. ———, and S. P. Cline. 1988. Coarse woody debris in forests and
323. Scheffer, T. C. and E. B. Cowling. 1966. Natural resistance of wood to plantations of coastal Oregon. Pages 5-24 in C. Maser, R. F. Tarrant, J. M.
microbial deterioration. Annual Reviews of Phytopathology. 4:147-170. Trappe, and J. F. Franklin, editors. From the forest to the sea: A story of
324. Schowalter, T. D. 1989. Canopy arthropod community structure and fallen trees. U.S. Forest Service, Pacific Northwest Research Station,
herbivory in old-growth and regenerating forests in western Oregon. Portrland, OR. General Technical Report PNW-GTR-229.
Canadian Journal of Forest Research 19:318-322. 345. ———, and J. F. Franklin. 1991. The structure of natural young,
325. ———. 1990. Invertebrate diversity in old-growth versus mature, and old-growth Douglas-fir forests in Oregon and Washington.
regenerating forest canopies. The Northwest Environmental Journal Pages 111-121 in L. F. Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff,
6:403-404. technical coordinators. Wildlife and vegetation of unmanaged Douglas-
326. ———. 1995. Canopy arthropod communities in relation to forest fir forests. U.S. Forest Service, Pacific Northwest Research Station,
age and alternative harvest practices in western Oregon. Forest Portland, OR. General Technical Report PNW-GTR-285.
Ecology and Management 78:115-125. 346. ———, ———, and T. B. Thomas. 1988. Coarse woody debris in
327. ———, E. Hansen, R. Molina, and Y. Zhang. 1997. Integrating the Douglas-fir forests of western Oregon and Washington. Ecology 69(1):
ecological roles of phytophagous insects, plant pathogens, and 689-702.
mycorrhizae in managed forests. Pages 171-189 in: K. A. Kohm and J. F. 347. ———, W. J. Ripple, and G. A. Bradshaw. 1994. Dynamics and pattern
Franklin, editors. Creating a forestry for the 21st century: the science of a managed coniferous forest landscape in Oregon. Ecological
of ecosystem management. Island Press, Washington, D.C. Applications 4(3): 555-568.
328. Schreiber, B. 1998. Long-term monitoring of wildlife leave trees in 348. Starr, L., editor. 1995. Soil: the foundation of the ecosystem. National
clearcut harvest units and the Siuslaw National Forest, Northwest Research News (5)1. Blue Mountains National Research Institute, La
Oregon, 1987-98. Report to the Siuslaw National Forest. Grande, OR. U.S. Department of Agriculture, Forestry Service, Pacific
329. Schulze, E. D., O. L. Lange, and R. Oren. 1989. Forest decline and Northwest Research Station.
pollution. Springer-Verlag, New York, NY. 349. Stathers, R. J., T. P. Rollerson, and S. J. Mitchell. 1994. Windthrow
330. Sedell, J. R., and F. J. Swanson. 1984. Ecological characteristics of handbook for British Columbia forests. Ministry of Forests. Research
streams in old-growth forests of the Pacific Northwest. Pages 9-16 in Progress Working Paper 9401, Victoria, BC, Canada.
W. R. Meehan, T. R. Merrell, and T. A. Hanley, editors. Fish And Wildlife 350. Statzner, B., J.A. Gore, and V.H. Resh. 1988. Hydraulic stream ecology:
Relationships In Old-growth Forests. Transaction of the Annual observed patterns and potential applications. Journal of the North
Management of American Institute Of Fisheries Biologists, Juneau, AK. American Benthological Society 7: 307-360.
331. ———, G. H. Reeves, and P. A. Bisson. 1997. Habitat policy for 351. Steeger, C. and C. L. Hitchcock. 1998. Influence of forest structure
salmon in the Pacific Northwest. Pages 375-388 in D. J. Stouder, P. A. and disease on nest-site selection by red-breasted nuthatches. Journal
Bisson, and R. J. Naiman, editors. Pacific salmon & their ecosystems. of Wildlife Management 62(4):1349-1358.
Chapman and Hall, New York, NY. 352. Stevens, V. 1997. The ecological role of coarse woody debris. An
332. Sharp, R. F. and J. W. Milbank. 1973. Nitrogen fixation in deteriorating overview of the ecological importance of CWD in BC Forests. Ministry
wood. Experientia 29:895-896. of Forests Research Program. BC, Canada.
333. Shaw, D. C. and J. Bible. 1996. An overview of forest canopy 353. Stone, E. C., and R. B.Vasey. 1968. Preservation of coast redwood on
ecosystem functions with reference to urban and riparian systems. alluvial flats. Science 159:157-161.
Northwest Science 70:1-6. 354. Stouder, D. J., P. A. Bisson, and R. J. Naiman, editors. 1997. Pacific
334. Shearer, C. A. and S. B. von Bodman. 1983. Patterns of occurrence of salmon and their ecosystems. Chapman and Hall, New York, NY.
ascomycetes associated with decomposing twigs in a Midwestern 355. Strickland, T. C., P. Sollins, N. Rudd, and D. S. Schimel. 1992. Rapid
stream. Mycologia 75:518-530. stabilization and mobilization of 15N in forest and range soils. Soil
335. Shaetzl, R. J., D. L. Johnson, S. F. Burns, and T. W. Small. 1989. Tree Biology And Biochemistry 9:849-855.
uprooting: a review of terminology, process, and environmental 356. Swanson, F. J. 1980. Geomorphology and ecosystems. Pages 159-170
implications cations. Canadian Journal of Forest Research 19:1-11. in R. H. Waring, editor. Forests: Fresh perspectives from ecosystem
336. Silvester, W. B., P. Sollins, T. Verhoeven, and S. P. Cline. 1982. Nitrogen analysis. Proceedings of the 40th Annual Biology Colloquium. Oregon
fixation and acetylene reduction in decaying conifer boles: effects of State University Press, Corvallis, OR.
incubation time, aeration, and moisture content. Canadian Journal of 357. ———, and G. W. Lienkaemper. 1978. Physical consequences of large
Forest Research 12:646-652. organic debris in Pacific Northwest streams. U.S. Forest Service, Pacific
337. Sinton, D. S., J. A. Jones, J. L. Ohmann, and F. J. Swanson. 2000. Northwest Research Station, Portland, OR. General Technical Report,
Windthrow, disturbance, forest composition, and structure in the Bull PNW-GTR-69.
Run basin, Oregon. Ecology, 81(9):2539-2556. 358. Swanston, D. N. and F. Swanson. 1976. Timber harvesting, mass
338. Sit, V., and B. Taylor. Editors. 1998. Statistical methods for adaptive
 622 Wildlife–Habitat Relationships in Oregon and Washington

erosion, and steepland, forest geomorphology in the Pacific Northwest. forests and streams. Pages 171-190 in R. H. Waring, editor. Forests :
Pages 199-221 in D. R. Coates, editor. Geomorphology And Engineering. fresh perspectives from ecosystem analysis. Proceedings of the 40th
Hutchinson Ross, Stroudsburg, Pennsylvania. annual biology colloquium. Oregon State University Press, Corvallis,
359. Swanson, F. J., R. L. Graham, and G. E. Grant. 1985. Some effects of OR.
slope movements on river channels. Pages 273-278 in International 377. Ugolini, F. C. and D. H. Mann. 1979. Biopedological orgin of peatlands
Symposium on Erosion, Debris Flow and Disaster Prevention, Tsukuba, in southeastern Alaska. Nature 281:366-368.
Japan. 378. Ure, D. C., and C. Maser. 1982. Mycophagy of red-backed voles in
360. ———,G. W. Lienkaemper, and J. R. Sedell. 1976. History, physical Oregon and Washington. Canadian Journal of Zoology 60:3307-3315.
effects, and management and implications of large organic debris in 379. U.S. Forest Service. 1996. Status of the Interior Columbia Basin.
western Oregon streams. U.S. Forest Service, Pacific Northwest Summary of scientific findings. U.S. Forest Service. Pacific Northwest
Research Station, Portland, OR. General Technical Report, PNW-GTR- Research Station, Portland, Oregon. General Technical Report PNW-
56. GTR-385.
361. ———, S. V. Gregory, J.R. Sedell, and A.G. Campbell. 1982. Land- 380.Valiela, I., G. Collins, J. Kremer, K. Lajtha, M. Geist, B. Seely, J. Brawley,
water interactions: the riparian zone. Pages 267-291 in R. L. Edmonds, and C. H. Sham. 1997. Nitrogen loading from coastal watersheds to
editor. Analysis of coniferous forest ecosystems in the western United receiving estuaries: new method and application. Ecological Applications
States. US/ISB Synthesis Series. Hutchinson Ross Publ. Co., Stroudsberg, 7(2): 358-380.
PA. 381.Van Cleve, K. C., T. Dyrness, L. A. Viereck, J. Fox, F. S. Chapin, III, and W.
362. Sweeney, B. W. and R. L. Vannote. 1978. Size variation and the C. Oechel. 1983. Taiga ecosystems in interior Alaska. Bioscience 33: 39-
distribution of hemimetabolous aquatic insects: two thermal 44.
equilibrium hypotheses. Science 200:44-446. 382.Vannote, R. L., G. W. Minshall, K. W. Cummins, J. R. Sedell, and C. E.
363. Swezy, D. M., and K. K. Agee. 1991. Prescribed-fire effects on fine- Cushing. 1980. The river continuum concept. Canadian Journal of
root and tree mortality in old-growth ponderosa pine. Canadian Fisheries and Aquatic Science 37: 130-137.
Journal of Forest Research 21:626-634. 383.Vitousek, P. M. and P. A. Matson. 1984. Mechanism of nitrogen
364. Tappeiner, J. C., D. Huffman, D. Marshall, T. A. Spies, and J. D. Bailey. retention in forest ecosystems: a field experiment. Science 225: 51-52.
1997. Density, ages, and growth rates in old-growth and young-growth 384.Vitousek, P. M. and W. A. Reiners. 1975. Ecosystem retention: a
forests in coastal Oregon. Canadian Journal of Forest Research 27: 638- hypothesis. Bioscience 5:376-381.
648. 385.Vitousek, P. M. and P. A. Matson. 1985. Disturbance, nitrogen
365. Tesch, S. D. 1995. The Pacific northwest region. Pages 499-558 in J. W. availability, and nitrogen losses in an intensively managed loblolly pine
Barrett, editor. Regional silviculture of the United States.Third edition., plantation. Ecology 66:1360-1376.
John Wiley and Sons, New York, NY. 386.Vogt, K. A., J. C. Gordon, J. P. Wargo, D. J. Vogt, H. Asbjornsen, P. A.
366. Thomas, J. W. editor. 1979. Wildlife habitats in managed forests. The Palmiotto, H. J. Clark, J. L. O’Hara, T. Patel-Weynard, B. Larson, D.
Blue Mountains of Oregon and Washington. U.S. Forest Service. Tortoriello, J. Perez, A. Marsh, M. Corbett, K. Kaneda, F. Meyerson, and D.
Agriculture Handbook No. 553. Smith. 1997. Ecosystems: Balancing science and management. Springer-
367. Thomas, T. L., et al. 1993. Viability assessments and management Verlag, Inc., New York, NY.
considerations for species associated with late successional and old- 387.Vonhof, M. J., and R. M. R. Barclay. 1997. Use of tree stumps as roosts
growth forests of the Pacific Northwest. The report of the scientific by the western long-eared bat. Journal of Wildlife Management 61(3):
analysis team. U. S. Forest Service. 674-684.
368. ———, and J. K. Agee. 1986. Prescribed fire effects on mixed conifer 388. Vogt, K. A., C. C. Grier, and D. J.Vogt. 1986. Production, turnover, and
forest structure at Crater Lake, Oregon. Canadian Journal of Forest nutrient dynamics of above- and belowground detritus of world
Research 16:1082-1087. forests. Advances in Ecological research 15: 303-375.
369. Thomas, J. W., R. G. Anderson, C. Maser, E. L. Bull. 1979. Snags: Pages 389. Walker, B. H. 1992. Biodiversity and ecological redundancy.
60-77 in J. W. Thomas, editor. Wildlife habitats in managed forests: the Conservation Biology 6:18-20.
Blue Mountains of Oregon and Washington. USDA Forest Service, 390. Walstad, J. D., S. R. Radosevich, and D.V. Sandberg. 1990. Natural and
Agric. Handbook. 553. Washington, D.C. prescribed fire in Pacific Northwest forests. Oregon State University
370. ———., R. J. Miller, H. Black, J. E. Rodiek, and C. Maser. 1976. Press, Corvallis, OR.
Guidelines for maintaining and enhancing wildlife habitat in forest 391. Walters, C. J. 1986. Adaptive management of renewable resources.
management in the Blue Mountains of Oregon and Washington. Pages McGraw-Hill, New York, NY.
452-476 in Transactions of 41st North American Wildlife and National 392. Ward, G. W., K. W. Cummins, R. W. Speaker, A. K. Ward, S. V. Gregory,
Research Conference. Wildlife Management Institute, Washington, D.C. and T. L. Dudley. 1982. Habitat and food resources for invertebrate
371. Tinnin, R. O., and D. M. Knutson. 1982. Witches’ broom formation in communities in South Fork Hoh River, Olympic National Park. Pages 9-
conifers infected by Arceuthobium spp: an example of parasitic impact 14 in E. E. Starkey, J. F. Franklin, and J. W. Matthews, editors. Ecological
upon community dynamics. American Midland Botanist. 107:351-359. research in national parks of the Pacific Northwest. Forest Research
372. ———, and ———. 1985. How to identify brooms in Douglas-fir Laboratory, Oregon State University, Corvallis, OR.
caused by dwarf mistletoe. U.S. Forest Service, Pacific Northwest 393. Waring, R. H. 1987. Characteristics of trees predisposed to die.
rearch Station, Portland, OR. Research Note, PNW-RN-426. Bioscience 37:569-574.
373. Torgersen, T. R. and E. L. Bull. 1995. Down logs as habitat for forest 394. ———. 1983. Estimating forest growth efficincy in relation to
dwelling ants—the primary prey of pileated woodpeckers in canopy leaf area. Advances in Ecological Research 13:327-354.
northeastern Oregon. Northwest Science 69:294-303. 395. ———, and J. F. Franklin. 1979. Evergreen coniferous forests of the
374.Townsend, A. R., M. T. Sykes, M. J. Apps, I. Fung, S. Kellomaki, P. J. Pacific Northwest. Science 204:1380-1386.
Martinkainen, E. B. Rastetter, B. J. Stocks, W. J. A. Volney, S. C. Zoltai. 1996. 396. ———, J. J. Landsberg, and M. Williams. 1998. Net primary
WG1 Summary: Natural and anthropogenically-induced variations in production of forests: a constant fraction of gross primary production?
terrestrial carbon balance. Chapter 9 in M. J. Apps, and D. T. Price, Tree Physiology 18(2).
editors. Forest ecosystems: Forest management and the global Carbon 397. ———, and G. B. Pitman. 1985. Modifying lodgepole pine stands to
cycle. NATO ASI Series 1: Global Environmental Change,Vol. 40. change susceptibility to mountain pine beetle attack. Ecology 66:889-
Springer, New York, NY. 897.
375.Trappe, J. M. and C. Maser. 1976. Germination of spores of Glomus 398. ———, and W. E. Winner. 1996. Constraints on terrestrial primary
macrocarpus (Endogonaceae) after passage through a rodent digestive
tract. Mycologia 68:433-436.
376. Triska, F. J., and K. Cromack, Jr.1980. The role of wood debris in
 CHAPTER 24: WOOD LEGACIES 623

production in temperate forests along the Pacific Coast of North

America and South America. Pages 89-102 in R. G. Lawford, P. Alaback,
and E. R. Fuentes, editors. High latitude rain forests and associated
ecosystems of the west coast of the Americas: climate, hydrology,
ecology, and conservation. Springer Verlag, New York, NY.
399. ———, and S. W. Running. 1998. Forest ecosystems: Analysis at
multiple scales, 2nd edition. Academic Press.
400. ———, K. Cromack, Jr., P. A. Matson, R. D. Boone, and S. G. Stafford.
1987. Responses to pathogen-induced disturbance: decomposition,
nutrient availability, and tree vigor. Forestry 60:219-227.
401. ———, T. Savage, K. Cromack, Jr., and C. Rose. 1992. Thinning and
nitrogen fertilization in a grand fir stand infested with western spruce
budworm. Part IV. An ecosystem management perspective. Forest
Science 38:275-286.
401a. Washington Department of Labor and Industries. 1992. Guidelines
For Selecting Reserve Trees. Report# P 417-092-000, Olympia.
401b. ———. 1999. Work-related traumatic Head and Brain Injuries in
Washington State 1990-1997. Report # 57-1-1999. State and Health
Assessment and Research for Prevention Program. Wasington State
Dept. of Labor& Industries.
401c. Weigand, J.F. and A. L. Burditt. 1992. Economic implications for
management of structural retention on harvest units at the Blue River
Ranger District, Willamette National Forest, Oregon. USDA PNW
Research Station, Research Note PNW-RN-510.
402. Weikel, J. M., and J. P. Hayes. 1999. The foraging ecology of cavity-
nesting birds in young forests of the notthern coast range of Oregon.
The Condor 101: 58-66.
403. Wickman, B. E., R. R. Mason, and H. G. Paul. 1992. Thinning and
nitrogen fertilization in a grand fir stand infested with western spruce
budworm. Part II:Tree growth response. Forest Science 38:252-264.
404. Wright, P. J. 1998. The effects of fire regime on coarse woody debris
in the west central Cascades, Oregon. Master Thesis. Oregon State
University, Corvallis, OR.
405. Yavitt, J. B. and T. J. Fahey. 1985. Chemical composition of interstitial
water in decaying lodgepole pine bole wood. Canadian Journal of
Forest Research 15:1149-1153
406. Zenner, E. K., S. A. Acker, and W. H. Emmingham. 1998. Growth
reduction in harvest-age, coniferous forests with residual trees in the
western central Cascade Range of Oregon. Forest Ecology and
Management 102:75-88.
407. Ziller, W. G. 1974. The tree rusts of western Canada. Canadian
Forestry Service Publ. 1329. Department of the Environment.Victoria,
BC, Canada.
408. Zinke, P. J. A. Stangenberger, and W. Colwell. 1979. The fertility of the
forest. California Agriculture. 33(5): 10-11.
 25
Single Species, Multiple Species,
or Ecosystem Management:
A Perspective on Approaches to Wildlife Conservation
Hal Salwasser

The conservation of wildlife and of biological diversity at conditions be distributed across landscapes? And where
large has taken various approaches in the U.S.4 Sometimes, do we turn for answers? We usually turn to our knowledge
the focus is on the provision of life requisites for a single of what certain plant or animal species need for population
species of plant or animal, such as spotted owls, elk, or viability or levels of abundance that allow for sustainable
grizzly bears. Sometimes it is on the provision of habitats uses of surplus individuals. The upshot, then, is that it
for a suite of species, i.e., a guild or biological community, seems to make little difference what approach we start
such as cavity-dependent or wetland-associated animals. with, species or ecosystems; we eventually end up
And sometimes the focus is on ecosystems, i.e., integrated addressing both.
systems of land, water, and biota in contiguous areas, e.g., A more germane question for wildlife or biodiversity
watersheds, landscapes, or regions. Given this variety of conservation is this: how do we best take advantage of all
approaches, it is logical to pose a simple question: when the tools available for perpetuating desired conditions of
do you take a single species, multiple species, or ecosystem select species, biological diversity, and ecosystems? To gain
approach? perspective on possible answers to this question, we need
an historical understanding of what has been used.
The Filter Concept First, consider the possibility that the coarse filter—fine
One answer to this question has been offered through the filter idea is really just a subset of a much richer, longer-
so-called coarse filter—fine filter concept. In this approach standing concept, that of multiple overlapping and nested
(also known as macro—micro filter) the needs of most filters (to continue the metaphor). Consider also that the
species are assumed to be met through the coarse filter so-called filters do not even belong to the same dimension,
provision of mosaics of habitats, i.e., multi-species or i.e., that they all deal with habitats. We can see this through
ecosystem approaches. The needs of certain species, a brief exposition of how the system of biodiversity
usually endangered or game species, are then addressed conservation has unfolded in the United States during the
through a fine filter of special habitat components and past 250 years.
species-specific actions. This “filter concept” is often Think of each of the major factors affecting conservation
attributed to Hunter,3 who provided a good articulation as a kind of filter or overlay. The first overlay was
of the concept. But in practical use, the filter concept is ordinances and laws designed to protect certain species
richer than the coarse-fine dichotomy. And it has been from excessive harvest, some as early as the 1600s in the
around for a very long time in wildlife conservation. first colonies and others as late as the early 1900s in the
The debate over single species, multiple species or west. The next overlay was reservation of large tracts of
ecosystem approaches is essentially a false debate. In land in public parks, forests, and refuges, beginning locally
reality, except for zoos, botanical gardens, and gene pool in the 1700s in the east and extensively in the late 1800s in
banks, there is no such thing as pure single species the west. The next overlay was restrictions on
approach to wildlife conservation. This could even be said environmental toxics, poisons, and traps used to reduce
of multi-species approaches. Thus, the above question of pests and predators, mostly not occurring until the mid-
when you use one or the other is not really a useful to late-1900s. In most of these cases, only crude models or
question to pursue; we need to use all of them. Even when analytical tools were used to accomplish the intended
we try to address the needs of individual species or groups conservation purposes, and sometimes no tools at all. To
of species in actual habitats or landscapes, it is ultimately the extent that these three filters or overlays were applied,
ecosystems that are conserved, restored or managed. This rather significant positive effects were accomplished for
has the result that what start out as single species or multi- wildlife conservation.5, 6, 8, 13 We might think of such actions
species approaches, in the end evolve toward ecosystem as mega or meta filters (again to continue the filter
approaches. Conversely, when we set out to conserve, metaphor). In fact, by the present time, most populations
restore, or manage ecosystems, one of the first questions of native species have been protected from excessive
that arises is this: how much of what kinds of habitat harvest or unregulated poisoning for at least several
conditions are needed, and how should those habitat decades. And about one-third of the land area of the U.S.

624
 CHAPTER 25: ECOSYSTEM PERSPECTIVE 625

is protected to some degree in public stewardship, Where Do Models Fit?

managed in ways that are conducive to the maintenance So it might be useful to step back a bit from our focus on
of significant aspects of biological diversity. But these are analytical models such as WHR matrixes (Wildlife Habitat
only the direct actions that impinge upon wildlife Relationships tables), HSI (Habitat Suitability Indices),
populations and availability of habitats. PATREC (Pattern Recognition), PVA (Population Viability
Early in the twentieth century, improvements in Analysis) and AI (Artificial Intelligence) and think about
agricultural production and the advent of machines how each of these potential tools might serve us in the
powered by internal combustion engines provided future. First, though, what will the future in which these
another overlay of conservation, allowing the nation to tools must operate be like? We know from ecology and
meet its food and fiber needs without continued paleoecology that the future is going to be dynamic,1 and
conversion of forests and grasslands to agriculture.2, 5 This thus we will have limited ability to predict with any
had a substantial positive, indirect effect on wildlife precision. So we will need tools that help us deal with
conservation, as habitat conversion is often the most change and surprises. The future is also going to have
significant and permanent negative impact to populations. many more human beings seeking greater access to the
Then, several decades ago, another set of mega filters was land, water, air, and habitat resources upon which other
put into place that has had profound but often little species also depend for livelihood.9, 15 So we need tools
recognized beneficial effects on biological diversity: that will help us mitigate or reduce potential human
namely, the nation’s clean air and clean water statutes. impacts on those resources, essentially to optimize
Think of these indirect factors as another set of overlays potentially complementary land and resources uses. It also
on the coarse and fine filters of habitat. Think also what appears that information will be more readily available
the environment and habitats for many species might be to people in the future through emerging communications
like without agricultural efficiency, clean air, and clean technologies. So we will need tools that tap the information
water. sources, such as geographic information systems, remotely
Late in the conservation game, so to speak, we saw the sensed databases, and internet data, and tools that will
emergence of new policy overlays: endangered species help people interpret or translate that information into
legislation (1960s in principle and ESA of 1973 in particular meaningful understanding of what is going on in nature
(P.L. 93-205, 87 Stat. 884, as amended); environmental and what is most likely to deliver desired outcomes, i.e.,
policy require-ments (NEPA in 1969; P.L. 91-190, 83 Stat. predictive tools that work with reasonable simulations of
852); and forest and rangeland conservation laws (in the future.10 Going back to the human dimension, a realistic
particular, FLPMA in 1976; P.L. 94-579, 90 Stat. 2743, as outcome of the growing population is that less space and
amended, and NFMA in 1976; P.L. 94-588, 90 Stat. 2949, as resources will be left for wild things to occupy or use. Thus
amended). These laws essentially refined conservation we will need tools that help us choose wisely where the
tools and approaches, generally requiring more greatest returns for investment in those habitats might
quantitative assessments of populations and habitats, exist. These are essentially decision analysis or risk
leading to the emergence of books and journal articles on analysis tools.
wildlife-habitat-relationships tools.12 They also increased It might also be useful to look at the environment that
accountability for achieving desired results for certain the first hundred years of conservation has bequeathed
species. Eventually, highly quantitative, analytical tools to us, and to realize that 70%, maybe 80% (maybe even
came into being as a result of these laws, to assess such more) of what there was to accomplish (or what was
things as population viability, habitat capability, biological possible given the starting point of the late 1800s) has
diversity and ecosystem integrity.7, 11, 12, 14 The models and already been done through the land systems and
analytical procedures, such as those presented in this book environmental and conservation laws and policies now
and its predecessors, are among the tools we can now use set in place by federal, state, county and tribal
to determine both the coarse and fine filters. They have a governments. What is left to us with our models, concepts,
dual edge, however. On the one hand, they can give us and analyses, then, is fine-tuning around the edges of these
systematic ways to address species and ecosystem land conservation systems and various environmental
conservation matter. On the other, they can create the laws. This is not to denigrate new tools and their potential
illusion that we can, in fact, be comprehensive, accurate, utility. Nor is it to say that significant accomplishments
or precise in how we set out to achieve desired end results. are a thing of the past. It is, rather, to put our new tools
The reality is that we can never be very comprehensive, into their proper perspective, to recognize them for what
accurate, and precise, because nature is more complex than they bring to the already mature conservation business. If
we can account for in models, and it is never going to be nothing else, these tools will be needed to help us retain
fully understandable or predictable. or restore the health and productive capacity of lands and
environments, so they can accommodate the many more
people who will occupy the future without the significant
losses in biological diversity that would occur were we to
not use these tools.
We must also acknowledge that just as some seemingly
unrelated forces or events of the past had as much to do
 626 Wildlife–Habitat Relationships in Oregon and Washington

with conservation of biological diversity as the direct humans fit into the scheme. So we return to the
actions taken on its behalf, or perhaps even more, the same fundamental conservation challenge. We have only one
might also be true in the future. I remind the reader of Earth to work with, and it is finite. It has many different
one historic example in support of this point: the kinds of places, i.e., ecosystems, each with differing
agricultural revolution of the early twentieth century in capabilities to sustain desired conditions of environments
the U.S. that stemmed the tide of increasing conversion of and human well-being. All ecosystems are constantly
wildlands to farms. No conservation law, land reserve changing, though many are still well within geo-climatic
system, or WHR model affected this outcome, yet the ranges of variability. But Earth and all its ecosystems must
positive result for wild plants and animals has been contend with more people, well beyond anything these
enormous. ecosystems have ever experienced. Regardless of where
Given this context, and given that we will use single- we enter the stream of species conservation, we will need
species, multi-species, and ecosystem approaches to to address individual species, groups of species, and
conservation, let us consider the relative strengths and ecosystems in an integrated manner. Furthermore, because
weaknesses of these approaches. (The following borrows people are so numerous, and because they and their
heavily from Johnson).4 artifacts are so widely distributed in nature, we will have
to integrate species and ecosystem concerns with
Single-species Approaches to Conservation increasing human needs and impacts. And we will have
to operate in the knowledge that events, forces, and trends
Strengths beyond our control or even our understanding may exert
• Ability to selectively focus on species most valued for more influence on our desired outcomes for biological
some particular purpose, i.e., endangered, game, diversity than anything we purposefully set out to do.
commercially valuable Thus I close this perspective with a review of the
• Efficient proxy for protecting habitats or ecosystems strengths and weaknesses of integrative, ecosystem
• More easily understood by public, can be based on approaches to conservation as they are now often
charismatic species proposed as key parts of sustainable development and
Weaknesses adaptive management schemes:
• Species taxonomy might not be clear
Integrative, Ecosystem Conservation, i.e.,
• We don’t know much about most species, and nothing
at all about many Sustainable Development
• Monitoring can be difficult and costly; easier to
Strengths
measure habitats, for example • Plans can have higher likelihood of durable
• Prime habitat for one species is not necessarily prime implementation because human needs and
for others institutional factors have been incorporated with
• No species is a perfect surrogate for another or for an species and ecosystem conservation; yields broader
entire ecosystem social ownership in the approach
• Can link biological reasons for conservation with
Ecosystem Approaches to Conservation
social and economic forces to increase political
Strengths support for conservation
• Ability to conserve many species and genetic diversity • Can put biological, economic, social, and political
through adequate amount and distribution of factors into a more explicit policy and decision
different ecosystem types (need good ecosystem making forum
classifications)
Weaknesses
• Ability to conserve diversity and ecological processes
• Wildlife and biodiversity can be devalued compared
essential to long-term productivity and resilience,
to other social and economic goals
hence the foundations for species persistence and
• Experimental, still not as well developed as historical
adaptation
species and ecosystem conservation strategies
• Most cost-effective way to conserve multiple species
• Social, economic and biological data may not all be
• Effective for species conservation when little or
available or in consistent quality at appropriate
nothing is known about many species
scales, and some factors may change at different
Weaknesses rates, e.g., when an economy changes faster than
• There is no ecosystem classification scheme with the ecosystem conditions
national or international consistency of species The challenge in the future for those who wish to conserve
classifications wildlife or biological diversity in our increasingly human-
• Can miss rare, poorly distributed, or potentially dominated ecosystems is threefold. First, it is to use the
endangered species strengths of each approach, i.e., single species, multiple
As noted earlier, species and ecosystem conservation species, and ecosystem, to compensate for the weaknesses
can no longer be done without full consideration of how of the other approaches. Second, it is to adopt adaptive
 CHAPTER 25: ECOSYSTEM PERSPECTIVE 627

management as the overarching framework to allow us

to learn from our actions and respond to both systematic
change and surprise events. And third, it is never to fool
others or ourselves by believing that we can figure natural
systems out precisely or comprehensively. We will simply
do the best we can with the data and tools at hand, and
learn as we go.

Literature Cited
1. Botkin, D.B. 1990. Discordant harmonies: a new ecology for the 21st
century. Oxford University Press. New York, NY.
2. Fedkiw, J. 1989. The evolving use and management of the nation’s
forests grasslands, croplands and related resources. USDA Forest
Service General Technical Report RM-175, Fort Collins, CO.
3. Hunter, M.L., Jr. 1990. Wildlife, forests, and forestry. Prentice-Hall, Inc.,
Englewood Cliffs, NJ.
4. Johnson, N.C. 1995. Biodiversity in the balance: approaches to setting
geographic conservation priorities. Biodiversity Support Program,
World Wildlife Fund, Washington, D.C.
5. MacCleery, D.W. 1996. American forests: a history of resiliency and
recovery. Forest History Society, Durham, NC.
6. Matthiessen, P. 1959. Wildlife in America. The Viking Press, New York,
NY.
7. McCullough, D.R., and R.H. Barrett, editors 1992. Wildlife 2001:
Populations. Elsevier Science Publishers, Ltd., London, England.
8. Rieger, J.F. 1986. American sportsmen and the origins of conservation.
Oklahoma University Press, Norman, OK. Revised edition: Oregon
State University Press, Corvallis, OR, 2000.
9. RNRF. 1999. Congress on human population growth: impacts on the
sustainability of renewable natural resources. Renewable Natural
Resources Journal, Special Report.Volume 16, Number 4, Washington,
D.C.
10. Salwasser, H. 1993. Perspectives on modeling sustainable forest
ecosystems. Pages 176-181 in D.C. LeMaster and R.A. Sedjo, editors.
Modeling sustainable forest ecosystems. Forest Policy Center, American
Forests,Washington, D.C.
11. Soule, M.E. (editor) 1987.Viable populations for conservation.
Cambridge University Press, Cambridge, England.
12. Thomas, J.W. (editor) 1979. Wildlife habitats in managed forests: the
Blue Mountains of Oregon and Washington. USDA Forest Service,
Agricultural Handbook No. 553.Washington, D.C.
13. Trefethen, J.B. 1975. An American crusade for wildlife. Winchester
Press and the Boone and Crockett Club, New York, NY.
14.Verner, J., M.L. Morrison, and C.J. Ralphs (editors) 1986. Wildlife 2000.
University of Wisconsin Press, Madison, WI.
15. Wilson, E.O. (editor) 1988. Biodiversity. National Academy Press,
Washington, D.C.
 26
Pacific Salmon and Wildlife—Ecological Contexts,
Relationships, and Implications for Management
C. Jeff Cederholm, David H. Johnson, Robert E. Bilby, Lawrence G. Dominguez,
Ann M. Garrett,William H. Graeber, Eva L. Greda, Matt D. Kunze,
Bruce G. Marcot, John F. Palmisano, Rob W. Plotnikoff, William G. Pearcy,
Charles A. Simenstad, & Patrick C.Trotter

Introduction
The landscapes of Washington and Oregon at first glance spans an area of freshwater and ocean habitat in excess of
appear to have some disconnect between the terrestrial 4 million km2. The essence of the salmon is that they link
and ocean environments. Abundant rivers and streams together what humans generally consider distant, diverse,
flow from the interior to the coastal zones, actively and separate ecosystems, and relatively long time spans.
connecting the freshwater, estuarine, and ocean systems Scientific knowledge of salmon in Washington and Oregon
(Photograph 1). Within these environments there are was preceded by a rich legacy of aboriginal culture, which
countless abiotic and biotic processes which form a highly wove them into everyday life.98, 379, 431 (Photograph 2)
integrated ecosystem. Key inhabitants include wild Salmon were an important food staple and a basis of many
anadromous Pacific salmon (Oncorhynchus spp.) legends of the native people of these states, particularly
(anadromous fishes are those that spend much of their those that lived along rivers and marine areas. Salmon
lives feeding in the ocean and migrate to freshwater to were consumed by natives in large quantities, for example,
breed), 605 common vertebrate wildlife species, and Craig and Hacker109, cited in 379 calculate that pre-contact
numerous species of macroinvertebrates and other fishes. catches of salmon in the Columbia basin alone ranged
Complex relationships have evolved within and between between 4.5 and 5.6 million fish annually. Most of the
anadromous salmon and other inhabitants that may be salmon caught at that time were consumed within their
important for maintaining this ecosystem. Former highly
exploitive fisheries and poor land uses, an over-reliance
on salmon hatcheries, and a change in ocean environment
have contributed to many salmon stock declines in these
states.367, 397, 32, 153 It has been suggested that future salmon
conservation will need to take an ecosystem approach if
wild stocks are to survive.485 The purpose of this paper is
to identify known relationships between wild salmon and
wildlife, to discuss the ecological context of these
relationships, and to suggest new ways of managing the
salmon resource with an ecosystem perspective.
We define wild salmon as indigenous species that are
the progeny of streambed spawners. This definition is used
to distinguish wild salmon from hatchery (artificially)
propagated salmon. The genus Oncorhynchus includes
both salmon and trout; however, for our purpose, we
collectively refer to them simply as salmon. Wildlife are
divided into two main categories, indigenous
macroinvertebrates (aquatic and terrestrial) and
vertebrates (amphibians, reptiles, fishes, birds, and
mammals) found in Washington and Oregon.
It is important to recognize that the ecosystem of
Washington and Oregon salmon can be hemispheric in
scale. It reaches from local inland watersheds, where
spawning occurs, all the way to ocean feeding grounds
north of the Aleutian Islands of Alaska, west to the Asian Photograph 1. Hoh River on the western Olympic Peninsula.
side of the Pacific Ocean, and back again. This ecosystem (Photo by Jeff Cederholm)

628
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 629

Photograph 2. Indians
fishing for salmon at Celilo
Falls on the lower Columbia
River. (Photo courtesy of
Archive of the Spokesman
Review, Spokane,
Washington)

respective river drainage and some were traded with

distant tribes. Columbia River tribal records indicate that
salmon were transported long distances inland, including
trade routes over the Continental Divide.
The Wishram and Wasco (tribes along the lower
Columbia River near Celilo Falls) seem to have been the
focal point in the most extensive trade network in the
plateau—one that reached to the mouth of the Columbia
and out onto the plains east of the Rockies. They traded
dried fish (salmon) for bison hides and other
commodities that originated on the plains.”177, cited in 379
Some of the earliest Euro-Americans to view Pacific
salmon traveled to the Northwest with the Lewis and
Clark expedition in 1805. Near the confluence of the
Columbia and Snake rivers, they observed salmon in
unimaginable abundance, as William Clark reported: “The
number of dead Salmon on the Shores & floating in the
river is incrediable to say....”252 cited in 119
European settlement and commercial development of
Washington and Oregon brought significant habitat
problems for the salmon; resulting in many physical,
chemical, hydrological, and biological modifications to the
environment (Photograph 3). Varied effects on salmon Photograph 3. Urbanization and habitat loss in the Puyallup
habitat are often interrelated in complex ways, and the River estuary and floodplain, Tacoma, Washington. (Photo
effects of various activities and ecosystem modifications courtesy of Washington Department of Natural resources
can be cumulative.513 Some of the more harmful habitat Photo and Mapping Section)
losses caused by humans have been: river channel clearing
and channelization, log driving and splash damming,
extensive land clearing, major water diversions, livestock
grazing, mining runoff pollution, logging road-associated
erosion and removal of the old growth forest, filling and
diking of wetlands and estuaries, hydroelectric dam
development, urban runoff, water and sediment
contamination with toxicants, and recently recognized
 630 Wildlife–Habitat Relationships in Oregon and Washington

Photograph 4. Salmon catch at the

Seattle wharf. (Photo courtesy of
Washington State Historical
Society, Tacoma, Washington.
Negative Number 1994.123.121)

human-induced oligotrophication of waterways.16, 59, 101, 174, impacts, they have not been enough to prevent the
208, 245, 296, 365, 453, 485, 543
widespread decline of wild salmon stocks in these states.
Fishery exploitation of Columbia River salmon by Euro- Recent publications have chronicled the low abundance
Americans became a major factor after the middle to late of wild salmon stocks along the Pacific Coast in the lower
1800s (Photograph 4). To ensure primary access to the forty-eight states.53, 181, 367, 387, 542 In 1991, the American
salmon, commercial fisheries were strategically located Fisheries Society367 published a list of 214 naturally
downstream of popular Indian fishing grounds. The spawning stocks of salmon, steelhead, and cutthroat from
principle means used to catch the salmon were gillnets, California, Oregon, Idaho, and Washington, including: 101
traps, seines, and fish wheels.121, 274, 458 It was reported that stocks at high risk of extinction, 58 stocks at moderate risk
“...on a single spring day in 1913 the Seufert brothers’ of extinction, 54 stocks of special concern, and one stock
wheel no. 5 turned a record catch of 70,000 pounds.”103 classified as threatened under the Federal Endangered
After the 1870s and up to the early 1900s, the Columbia Species Act (ESA) of 1973. In spite of past salmon habitat
River salmon fishery grew from 1 to 40 canneries.368 cited in degradation and over-fishing, however, some stocks
365, 478
(Photograph 5) Fish wheels were prohibited on the remain healthy. 221 Since 1990, The National Marine
Columbia River after 1935.365 Commercial landings of Fisheries Service (NMFS) has received a number of
Columbia River salmon and steelhead peaked between petitions to list Pacific salmon stocks as threatened or
1880 and 1930, and then went into a long-term decline endangered under ESA, and the first salmon stock of this
through to present times.365 Depletion of the prime spring area to be listed as endangered was the Snake River
and summer chinook probably started earlier than this sockeye, in November 1991.366a
time frame, however, as the fishery shifted to the less
desirable coho and fall chinook.275 One estimate of annual General Salmon Life History
pre-Euro-American salmon and steelhead run size for the There are seven species of Pacific salmon and trout of the
Columbia River ranges between 8.2 and 16.3 million fish.379 genus Oncorhynchus in Washington and Oregon, and they
Early attempts to increase salmon catches using salmon include: chum (O. keta), pink (O. gorbuscha), sockeye (O.
hatcheries began as early as the 1870s, when concerns nerka), chinook (O. tshawytscha), and coho salmon (O.
about over fishing led the Oregon and Washington Fish kisutch); and rainbow (called steelhead when anadromous)
Propagating Company to construct a salmon-breeding (O. mykiss) and coastal cutthroat trout (O. clarki clarki).
station on the Clackamas River.365 By the 1960s, with the Some of these species, including the sockeye salmon
advent of the Oregon Moist Pellet medicated food, hatchery (kokanee) and rainbow and cutthroat trouts, have both
salmon production increased dramatically. Total annual anadromous and nonanadromous forms.
Columbia-Snake River system hatchery production A typical anadromous salmon life history has five main
(Washington, Idaho, and Oregon) reached 216 million stages: (1) spawning and egg incubation, (2) freshwater
smolts in 1989.390 By the middle 1990s there were well over rearing, (3) seaward migration, (4) ocean rearing, and (5)
100 state, federal, tribal and private salmon hatcheries in return migration and deposition of marine-derived
Washington and Oregon. The history of artificial nutrients into the freshwater ecosystem (Figure 1). Each
propagation reveals a recurring cycle of technological species has slightly different temporal phases of the
optimism followed by pessimism. anadromous life history. The chum, pink, sockeye,
While many attempts have been made at remedying chinook, and coho salmon all die after spawning just once,
the threats of habitat loss, over-fishing, and hatchery a life history strategy known as semelparity.322 This life
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 631

Photograph 5. Canned salmon at the

Apex fish company (1913). (Photo
courtesy of Washington State Historical
Society. Negative number 27683)

Figure 1. Generalized
resident forms anadromous and
nonanadromous (resident)
Pacific salmon life histories,
showing freshwater, estuary,
and ocean components (the
original diagram was from
Nicolas and Hankin372 and later
modified by Spence et al.485

anadromous forms
 632 Wildlife–Habitat Relationships in Oregon and Washington

strategy has evolved because of the need to have a greater and intra-specific competition.9 Declining streamflow
portion of the energy obtained from ocean feeding devoted conditions also may cause some fish (e.g., chinook, coho)
to gamete production and juvenile survival. Consequently, to emigrate to estuaries,196, 522 where they continue to rear.
survival after spawning no longer offered an advantage Where species overlap in fresh water, a number of
to these species.322 The iteroparous, repeat spawning temporal and behavioral differences facilitate
strategy, typical of the rainbow and cutthroat, probably coexistence.96, 282
occurred in the headwater reaches of larger rivers, where Upon the first rains and high waters of fall, coastal
nonanadromous populations could be maintained year species (juvenile coho, steelhead, and cutthroat) make a
around. These fish generally were smaller in size, less directed migration to seasonally alternate rearing habitats.
fecund, and had sparser distribution and lower abundance Juvenile coho and cutthroat exhibit major immigrations
than the anadromous forms. However, by retaining into side-channel swamps80 and riverine ponds,94, 398, 400, 162
iteroparity, calamitous losses of young due to floods or located along river flood plains. Juvenile coho, steelhead,
drought, could be compensated for in subsequent and cutthroat are also known to immigrate into small
breeding seasons.322 “runoff” tributaries (valley-wall tributaries) of rivers.94, 271
Chum, pink, sockeye, chinook, and coho salmon all Presumably these immigrations are to avoid high flows
spawn sometime from August through February, and and turbidity of main rivers, as well as to take advantage
cutthroat spawn between December and July. Pacific of good feeding conditions.398, 399 In contrast, interior
salmon are able to clean gravels by purging them of fine (Idaho) juvenile chinook and trout are known to move
sand and silt particles during redd (spawning nest) out of tributaries and into main rivers to over-winter,
excavation, but subsequent sediment transport processes probably to avoid winter ice conditions in the tributaries.57
and fine bedload flux tend to return this environment to Upon completing their freshwater stage, juvenile
the pre-spawning conditions.414 After approximately 2-4 salmon of all anadromous forms undergo a physiological
months of incubation, salmon fry swim up through the change called smoltification that includes osmoregulatory
gravel and emerge into the stream. Emerging fry can vary adjustments which prepare them to enter saltwater. For
widely in size at emergence, ranging from 20+ mm example, chum and pink salmon are nearly smolts upon
nonanadromous cutthroat to 35-40 mm chinook. Upon emergence from the gravel, going directly to estuaries and
emergence, fry actively feed on a variety of aquatic insects, the ocean;259, 440 while chinook197 and coho442 may either go
and for those that freshwater rear for extended periods of directly to sea the first summer of their life, or remain in
time (e.g., coho), the proportion of terrestrial food items freshwater for a whole year before smolting. Sockeye may
in the diet may increase to over 30%.338 Larger-sized rear in freshwater for one or two years before smolting,77
juvenile salmon such as older aged rainbow and cutthroat and steelhead283 and cutthroat160, 162 may not smolt for two
prey on a mixed diet of aquatic and terrestrial or three years or more.
macroinvertebrates, and may supplement their diet with Once in the estuary or ocean, most salmon prefer to
occasional salmon eggs or fry.498, 521 Sockeye fry are known feed on such prey as euphausiids, squid, herring,
to feed on cladocerens, copepods, and gammarid sandlance, rockfish, and anchovy.83b, 197 While in the ocean
amphipods in lakes.337 most salmon species migrate long distances to feeding
After a summer of rearing in fresh water, juvenile coho grounds along the North Pacific coast.135, 136, 178, 194, 196 In
average approximately 50 to 90 mm in length, and may contrast, anadromous cutthroat may only range several
weigh 2 to 5 g each.87,128 Summer low-flow is a crucial time kilometers from their natal stream without overwintering
in the life of juvenile salmon that extended rear in in the ocean.395
freshwater. During this period the volume of aquatic During their anadromous life history salmon make
habitat shrinks to a minimum, which can intensify inter- important ecological contributions (as prey) to various

Table 1. Key sources of life history information for the seven salmon species of Washington and Oregon.

Species Reference

chum Bjornn and Reiser,56 Everest et al.,140 Koski,259 Salo,440 Wydoski and Whitney573
pink Bjornn and Reiser,56 Everest et al.,140 Heard,198 Wydoski and Whitney573
sockeye Bjornn and Reiser,56 Burgner,77 Everest et al.,140 Foerster,146 Wydoski and Whitney573
chinook Bjornn,56 Bjornn,57 Everest et al.,140 Healey,197 Wydoski and Whitney573
coho Bjornn and Reiser,56 Bustard and Narver,80 Everest et al.,140 Peterson and Reid,401 Salo and Bayliff,439 Sandercock,442
Tagart,510 Wydoski and Whitney,573
steelhead Allee,9 Bjornn and Reiser,56 Cederholm and Scarlett,94 Everest et al.,140 Stolz and Schnell,498 Winter,560 Wydoski and
Whitney573
cutthroat Cederholm and Scarlett,94 Everest et al.,140 Fuss,160 Garrett,162 Glova,166 Hall et al.,181 Johnston,238 Pearcy et al.,395
Trotter,521 Wydoski and Whitney573
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 633

Figure 2. Some food web beneficiaries of Pacific salmon

nutrient in freshwater, estuary, and ocean environments.

predators in the Pacific Northwest ecosystem, regardless Water Quality and Quantity: temperature, dissolved
of whether a particular individual salmon completes all oxygen, dissolved nutrients, dissolved and particulate
life history stages or not (Figure 2). It is not uncommon organic matter, hydrography.
for overall salmon survival rates to average 0.1% from egg
Cover Factors: interstitial spaces (space between gravels),
to spawning adult. See Table 1 for more life history
undercut banks, woody debris, water surface disturbance.
information on the seven salmon species of Washington
and Oregon. Biological Factors: food availability, salmon carcass
nutrient inputs, competition, predation, disease, parasites,
Freshwater and Terrestrial Habitat and functioning riparian conditions.
The abundance of fish in a stream is greatly affected by
Relationships of Salmon the stream’s capacity to produce food. Many of the factors
influencing stream productivity change predictably with
Freshwater Habitat
changes in stream size, a pattern termed the river
Freshwater habitat of salmon includes all the physical,
continuum.529 Productivity is influenced by nutrient
chemical and biological elements within the aquatic
availability, input of organic matter from external sources,
environment. Geology, climate, topography, disturbance
and the capacity for the channel to store and process
history, nutrients from returning salmon, and
organic matter and light. Differences in these factors can
characteristics of the riparian vegetation typically govern
be very large and lead to a high degree of variability in
the characteristics and the distribution of habitat types in
production of fish, including salmon and trout
a watershed.33, 38, 332, 417 Components of freshwater habitat
populations.223 Some of the highest freshwater production
include:
values have been reported for trout in New Zealand
Physical Characteristics: channel width and depth, spring-fed streams, 54.7 g/m2/y;10 however, production
substrate composition, pool and riffle frequency, pool values from the Pacific Northwest are generally low when
types, and channel roughness. compared with other regions of the world, often below
1.0 g/m2/y and very rarely over 5.0 g/m2/y.55
 634 Wildlife–Habitat Relationships in Oregon and Washington

Riparian Habitat an increased susceptibility to disease,363, 386 have all been

In this section, we offer only a summary of riparian documented. Some species of amphibians and aquatic
aspects, as a chapter elsewhere in this book covers this macroinvertebrates also are thermally intolerant and
topic more fully. elevated water temperatures may have detrimental
Many of the functional and structural attributes of impacts on their populations.78, 137, 327
stream habitat are created and maintained through Riparian vegetation increases streambank stability and
interaction with riparian vegetation. Riparian areas resistance to erosion. Roots from woody and herbaceous
constitute the interface between aquatic and terrestrial vegetation bind soil particles together, helping to maintain
ecosystems,171, 506 performing a number of vital functions bank integrity during erosive high-streamflow events.485,
506
that affect the quality of salmon habitats as well as Riparian vegetation also facilitates bank-building
providing habitat for a large variety of terrestrial plants during high flow events by slowing stream velocities,
and animals. Riparian areas influence streams and which in turn helps to filter sediments and debris from
consequently salmon habitat in a variety of ways,485 suspension. This combing action helps to stabilize and
including: rebuild streambanks, allowing the existing channel to
narrow and deepen, and increases the effectiveness of
Shade which dampens seasonal and diel fluctuations in riparian vegetation in providing bank stability and
stream temperature and controls primary and secondary shade.132 During over-bank flows, water is slowed and fine
production. silts are deposited in the flood plain, increasing future
Streambank stabilization provides erosion resistant roots productivity of the riparian zone.485
that bind soil particles together, thus facilitating bank Forested riparian areas generate much of the organic
building during high flow events by slowing the stream matter that provides the energy source for the trophic
velocities. systems of small streams. In one study of forested
headwater channels in Oregon, Sedell et al.452 determined
Sediment control regulates sediment flow from upland that over 90% of the in-channel organic matter was
areas by acting as a filter, or storing sediments in the provided from the surrounding terrestrial environment.
primary floodplain. A 10-m wide stream in western Washington received over
Litter input contributes a significant amount of organic 75% of its annual organic matter supply from terrestrial
matter to streams, which acts as an important food sources.46 Even though this source of organic matter
resource for aquatic communities. decreases relative to autochthonous (biomass produced
from within the stream) organic matter in larger channels,
Large woody debris (LWD) provides important structure it remains vital to stream productivity.
to the stream channel for energy dissipation, fish habitat, Large woody debris (LWD) has been shown to be a
and salmon carcass retention. critical structural component in Pacific Northwest streams,
Nutrients input: riparian zones mediate the flow of forming pools, waterfalls, and overhead cover; and it also
nutrients to the stream and are, therefore, important regulates the transport of sediment, gravel and organic
regulators of stream production. Some riparian species matter, for fish and other aquatic biota.49, 50 (Photograph
such as red alder (Alnus rubra) also fix atmospheric 6) In forested watersheds LWD provides the most common
nitrogen and therefore augment N availability to the obstruction, often forming pools in various types of fluvial
ecosystem. channels.332 Without this material, pool abundance and
size is decreased,45 reducing habitat complexity and
Microclimate: streamside soils and vegetation can have a potentially reducing the diversity of the fish community.
significant effect on moderating the climate within riparian In addition to numerous habitat and morphological
zones. functions,50 wood (organic debris) helps retain salmon
Streamside vegetation moderates water temperature. carcasses in streams for many carnivorous wildlife
This relationship is influenced by elevation, air species91 and general biological activity.90
temperature, stream width, water depth, and aspect.41 Riparian areas play a key role in determining the
Removal of riparian vegetation has been associated with concentration of nutrients in stream water.42, 520 The
increased maximum water temperatures, and diurnal presence of even a narrow riparian buffer can profoundly
fluctuations in water temperature during summer; and influence stream water chemistry. Uptake and storage of
decreased winter water temperatures.41 Small, low- various elements carried by groundwater can be
elevation streams are the most susceptible to summer considerable, even where input rates have been
water temperature increases caused by canopy removal.504 substantially altered as a result of upslope land uses.284
The biological consequences of elevated water Riparian vegetation composition can influence nitrogen
temperature on aquatic communities are complex. There input to streams. Early successional vegetation in riparian
is little information indicating direct mortality of fishes as areas in the Pacific Northwest is often dominated by red
a result of temperature changes related to riparian canopy alder, a nitrogen (N) fixing species. As a result, the N
removal;41 however, reductions in growth rate,51, 572 changes content of litter beneath these riparian stands is 1.5 to 3-
in life history,215 changes in competitive interactions fold higher than sites where conifer species are the
between species,418 reductions in fecundity of adults,40 and dominant component of the over story.128a The result is
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 635

Photograph 6. Low gradient stream

showing large woody debris formed
habitat in Monroe Creek,
Washington. (Photo by Jeff
Cederholm)

higher N levels in the riparian soils58 and increased assemblages found in riparian areas compared to uplands.
delivery of N to the stream channel. Higher N levels in Wildlife also congregate seasonally in riparian areas where
stream water may elevate primary production and salmon spawn, to take advantage of an abundant food
decomposition (heterotrophy) in the channel and increase supply of live fish205 and carcass flesh.91 The high value of
food availability for the invertebrate community. Increased riparian habitats to wildlife has been recognized by
invertebrate production may elevate food availability for naturalists,380 and considered a bridge between upland
stream-dwelling fishes, amphibians, and other insect habitats and the aquatic environment. The combination
feeders such as bats and flycatchers. of shape, moisture, deposition soils, and disturbance
The riparian area may act as either a source or sink of regime unique to riparian areas contributes to their
organic matter and sediment during flood flows. The exceptional productivity in terms of plant growth, plant
manner in which the stream and riparian area interact at diversity, and structural complexity of the vegetation.237,
270, 329
these times depends upon the morphology and vegetation Wildlife dependency and diversity peak at this
of the riparian zone and the intensity of the discharge terrestrial/aquatic boundary. Brown69 reports that 359 of
event. 34 The structure and abundance of riparian 414 (87%) species of wildlife in western Washington and
vegetation plays a key role in moderating the movement western Oregon use riparian areas and wetlands during
of materials between the riparian area and the stream.361 some season or part of their life cycle. In their detailed
Vegetation in the riparian zone has been shown to be the examination of wildlife and habitats for all of Washington
single most important structural element for the retention and Oregon, Johnson and O’Neil236 reported that 393 of
of fluvially transported organic matter during high flow 456 (86%) of the common terrestrial and freshwater
events.484 Similarly, riparian vegetation promotes the wildlife species have seasonal use of riparian areas,
storage of sediment,216 that may provide germination sites wetlands, and streams. Of these 393 species, 110 were
for some species of riparian plants.370 The variations in found to be closely associated (e.g., obligates) with eastside
retentive capacity of different riparian areas for organic and westside riparian habitat types.
matter leads to large differences in the organic content of The close association may very well have evolved from
riparian soils, ranging from nearly all inorganic material the direct or indirect exploitation of the rich vegetative
in some locations to very high concentrations of organic habitat provided by riparian areas.255 Quantitative studies
matter in stream-adjacent swamps and wetlands.71 This conducted during the past several decades have supported
variation in substrate further contributes to riparian observations and have identified biological and physical
vegetation heterogeneity.2 attributes of riparian habitats which enhance their value
The area in which water exchange between the channel to wildlife. Brinson et al.64 and Oakley et al.381, cited in 380
and the underlying riparian soils occurs is termed the summarize these important biological and physical
hyporheic zone.491 The extent of the hyporheic zone varies features of riparian areas:
as a function of site topography and soil characteristics. • presence of surface water.
In riparian areas of low relief and porous soils, the • increased humidity, high rates of transpiration, and
hyporheic zone may extend as far as 3 km from the edge greater air movement.
of the channel.491 Riparian vegetation can influence the • complexity of biological and physical habitats.
amount of water stored in the hyporheic zones and soil • maximum edge effects with adjacent upland forests,
chemistry. which is beneficial for some species.
The attractiveness of riparian areas to wildlife likely • food supply.
reflects three main attributes: the presence of water, local • thermal cover.
microclimate condition, and more diverse plant
 636 Wildlife–Habitat Relationships in Oregon and Washington

According to O’Connell et al.380 stream type has a direct Alder stands begin to die-out and provide LWD to
influence on the riparian habitat and its associated wildlife channels after about 60 years.175 Shade tolerant conifers,
communities. In the smaller headwater streams the like western red cedar (Thuja plicata) and western hemlock
impacts of the upstream riparian vegetation on the streams (Tsuga heterophylla), colonize the site and begin to provide
is greater than downstream where flow volume increases, needles and other litter to the channel. Some stands of
flooding is more widespread, and the impact of riparian alder can persist and will repeat themselves several times
vegetation on the stream is less. Brinson et al.64 suggest (Slaney, pers. comm.). Woody debris amounts and average
that middle order perennial streams and associated piece size increase for 100 or more years following conifer
riparian areas have the greatest wildlife use. Periodic occupation of a site.44, 506 The morphology of the channel
flooding can enhance the availability of food for wildlife and the routing of sediment and organic matter evolve
by creating new feeding areas.64 Flooding can also make slowly as the riparian community changes, ultimately
riparian habitat unsuitable for other species. Species creating channels which are highly complex structurally
abundance of riparian mammal communities has been and support a macroinvertebrate community dominated
related to the timing of recent hydrologic events; by shredders.11
impoverished mammal populations have been attributed Forest practices and other land uses have accelerated
to recent flooding whereas more abundant populations the rate of occurrence of some types of disturbance. The
have been observed in areas not subject to recent acceleration in disturbance has led to the establishment
flooding.64 of early successional communities in the majority of
riparian areas on commercial forest land in the Pacific
Habitat-forming Processes Northwest,60, 86, 154 and has been shown to alter natural
Disturbance plays a major role in maintaining community hydrology.207 Practices such as splash damming of rivers
diversity and productivity in many ecosystems,105, 420 and to float logs to market,550 and removing all trees to the
is a key factor in creating and maintaining diverse stream channel’s edge,43 modify the riparian successional process.
habitat in the Pacific Northwest.54 These disturbances Timber harvest or roads constructed on unstable slopes
range in severity from minor events, such as seasonal or road drainage systems that were improperly
changes in flow, to less frequent high intensity events such maintained, dramatically increase the incidence of
as wildfire, debris torrents, and major floods. Riparian and landslides.93, 421, 456 Many hillslope failures enter stream
channel conditions evolve as impacted areas recover from channels and may move considerable distances
disturbance. 211 The result is a diverse set of riparian downstream, removing streamside vegetation and soil. On
community types and stream habitat conditions that vary the positive side, however, localized landslides also can
over both time and space.2 input massive amounts of spawnable sized gravels and
Disturbance contributes to both diversity of aquatic LWD into stream channels, where they may benefit salmon
fauna and productivity of these communities when populations.456 These disturbance events have affected a
considered at a watershed level.173 Aquatic communities large proportion of the riparian areas bordering streams
associated with early-successional riparian areas typically in the region over the last century, and have played a key
exhibit low diversity, but high productivity for certain role in determining channel form and habitat conditions.507
species. Removal of the channel shading canopy brings A study of fire history in Mt. Rainier National Park201
about dramatic increases in light and algal productivity; reported that alluvial terraces and valley bottoms were
however, input of terrestrial litter decreases. 46, 172 often forested with old stands, and that every major river
Invertebrates that feed on algal material (e.g., grazers) valley contained a stream-side old-growth corridor. These
typically dominate communities at recently disturbed observations support an inference that the moist
sites.137 These invertebrates form a major component of environment of riparian areas inhibits fire and reduces
the diet of some salmon and trout338 and can contribute to the fire return interval for riparian forests. The persistence
increased fish productivity following disturbance.46, 52, 352 of live trees in riparian forests may also provide a local
The increased productivity is typically observed during seed source that facilitates a more rapid development of a
summer, and often does not extend into winter months multi-layered, conifer-dominated forest.405
when the availability of shelter from high flows for
juvenile salmon becomes important.94, 162, 401 Beaver
After forest canopy closure, primary productivity in Beavers have long co-existed with salmon in the Pacific
streams decreases. The type of litter delivered to these Northwest, and have had a important ecological
systems and the physical characteristics of the channel relationship with salmon populations. The beaver created
differ from those at sites bordered by mature vegetation. and maintained a series of beneficial aquatic conditions
Hardwood trees, especially red alder, often dominate the in many headwater streams, wetland, and riparian
canopy at these sites. Litter from red alder trees systems, which serves as juvenile salmon rearing habitat.
decomposes much more rapidly than conifer litter, in part Beavers have multiple effects on water bodies and riparian
due to the higher N content.451 The high N content of the ecosystems that include altering hydrology, channel
litter improves its nutritional value for shredding morphology, biochemical pathways, and stream
macroinvertebrates but the high rate of decomposition productivity.385 Beaver ponds were of special importance
causes it to be scarce at some times of the year. in more arid regions, but also had important roles in
coastal systems.365
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 637

Beavers were once extremely abundant in the Pacific and this increases the water table, enhances summer flows,
Northwest, but as far back as 1778 trapping expeditions adds cold water during summer, and causes more even
into western North America began depleting their stream flows throughout the year. During winter, beaver
numbers. Between 1834 to 1837, pelts from 405,472 beavers ponds in cold environments prevent anchor ice from
from the area that would become southwest Washington forming and prevent super-cooling of the water. By storing
and Oregon were shipped to Europe. Past excessive spring and summer storm run-off, beaver ponds help to
trapping, and subsequent unregulated land- and water- reduce downstream flooding and the damage from rapid
use activities, significantly reduced abundance of beaver increases in stream flows.385
and beaver ponds. Additionally, excessive livestock Beavers also help shape riparian habitat. Beaver ponds
grazing in riparian areas has degraded habitat conditions increase the surface area of water several hundred times
for beaver.385 Severe declines of beaver in Washington and and thereby enhance the overall riparian habitat
Oregon have fundamentally altered important natural development.385 They also enhance vegetation growth by
aquatic ecosystem processes such as nutrient cycling, flood increasing the amount of groundwater for use by riparian
plain development, and stream hydrology. plants and wetland areas. The presence of beaver can have
Beaver dams can obstruct channels and redirect channel both positive and negative influence on salmon habitat,
flow and the flooding of streambanks and side channels. but on the whole, their presence is considered of great
By ponding water, beaver dams create enhanced rearing benefit to both water quality and salmon, particularly
and over-wintering habitat that protect juvenile salmon juvenile coho salmon and cutthroat trout, and to many
during high flow conditions.365 Studies in Oregon coastal other species of wildlife and invertebrates.
streams have suggested that where the amount of
spawning is adequate, the winter survival of juvenile coho, Estuary Habitat
which can be swept downstream in high winter flows, is By definition,413 an estuary is a region where salt water of
limited by the presence of adequate slow-water habitat.374 the ocean is measurably diluted by freshwater runoff from
Beaver dams are often found associated with riverine the land within a constricted body of water. In the Pacific
ponds called “wall-base channels”401 along main river Northwest, river flow plays a strong role in the structure
flood plains, and these habitats are used heavily by and dynamics of estuarine circulation.515 Estuaries of
juvenile coho salmon94, 400 and cutthroat trout94, 162 during lowland watersheds along the Washington and Oregon
the winter. Though their dams can occasionally block coasts tend to exhibit high peak flows associated with
upstream migration by adult and juvenile salmon, studies winter storms, but often extremely low flows associated
of trout movement indicate that fish can pass over beaver with the dry summers. This can frequently cause dramatic
dams during all seasons.385 Beaver dams may temporarily differences in the available estuarine habitat between
keep salmon adults in the lower parts of spawning winter and spring-summer periods, limiting summer
streams, where flows are greater and pools are deeper. rearing. In some southern Oregon and northern California
Then when dam breaching flows occur, free passage to estuaries river flow can decrease to the point that bars
upstream areas is allowed. form across the estuaries’ entrances, restricting juvenile
Beaver foraging can cause a loss of woody riparian salmon ocean emigration to extreme high (spring) tides.
vegetation and an increase of fine sediments, but it also Perhaps the most fundamental concept in
increases the input of large woody debris to streams, and understanding the estuarine ecology of juvenile salmon
beaver droppings may enrich pond productivity. Bank is that the salmon do not respond to singular habitats per
dens and channels can increase erosion potential, but se, but rather interact with a mosaic of habitats in response
because ponds fill with sediment to become wetlands over to changing migratory mandates, tidal cycles and
time, this helps to retard upstream erosion and retain freshwater runoff events (Figure 3). River flow and tide,
sediments that otherwise could adversely alter physiological change, prey and predator distributions, and
downstream areas. In a Wyoming study of an area that likely metapopulation genetic structure as well, all affect
had 10.5 beaver dams per km, each dam was found to the rate of fish movement through the estuary. But the
retain 5,350 m3 of sediment. In another Wyoming study, opportunity for juvenile salmon to exploit preferred
sediment loads were reduced by 90% after flowing 8 km habitats is just as likely dependent on the arrangement of
through an area with well developed riparian habitat and key landscape features such as tidal-freshwater and
beaver dams. brackish rearing zones, low-velocity refugia, migratory
Beaver ponds provide a sink for nutrients from corridors and foraging patches. Although this is a
tributary streams and create conditions that promote relatively new topic of research, with few definitive
anaerobic decomposition and de-nitrification. These experiments and tests, there is some emerging evidence
processes can cause nutrient enrichment and increased that the edge of marsh vegetation in dendritic tidal channel
primary and secondary production downstream from the and slough systems may relate directly to juvenile salmon
pond, while increasing nutrient retention time and production.473
enhanced invertebrate production in the pond.365 These Large watersheds with significant snow accumulations
factors help increase salmon growth and survival, and also and extended melting periods can create prolonged spring
helps improve water quality. Beaver ponds can cause freshets.308 Spring and winter freshets, and winter “rain-
increased storage of water in the banks and flood plains, on-snow” events associated with rapid snowmelt, produce
 638 Wildlife–Habitat Relationships in Oregon and Washington

Figure 3. Movements
and migrations of
juvenile Pacific
salmon across tidal-
freshwater delta-
estuarine landscapes.
(Contributed by C.
Simenstad,
unpublished
diagram)

flooding in tidal flood plains and estuaries that influences degree is a function of scale, as juvenile salmon can
short-term and long-term productivity of juvenile salmon respond to habitat features (e.g., LWD or tidal channels)
and their ecosystems.360, 566 Although flood plain and that are elemental to the broader habitats. Estuaries
estuarine wetland flooding increases flows in the main generally posses eight habitat components: (1) subtidal
distributary channels, likely diminishing the ability of distributaries; (2) mud- and sand-flats; (3) gravel-cobble
juvenile salmon to occupy them, considerable side-channel beaches; (4) low elevation emergent marshes; (5) high
and other flood plain wetlands (i.e., ponds, relict side- elevation emergent marshes; (6) forested and shrub
channels) are inundated and become available for refuge swamps; (7) eelgrass; and (8) kelp. Salmon communities
and rearing. This flooding recruits organic detritus and have been shown to utilize many of these habitat
dissolved nutrients from these peripheral wetlands and components. Juvenile coho (fry, fingerling) are often found
imports them to the estuary. While trapped in estuarine rearing during winter and early spring in the tidal flood
wetlands or circulation features such as estuarine turbidity plains of many large rivers such as the Chehalis River.323,
maxima,472 these materials contribute to primary and 470, 471, 472
These fish are either staging for migration through
secondary production by supporting food web pathways the estuary or are moving back into freshwater for
to juvenile salmon. extended rearing. Work in British Columbia,438, 522, 523 and
The structure of the watershed and estuary, with the Alaska 512 show that certain sub-populations have a
seasonal variability in river flow, shapes estuarine minimal juvenile freshwater rearing phase of their life
circulation, and strongly influences juvenile salmon history (“ocean-type”), and spend extended periods of
residence time, habitat use and production. Except where time either feeding in estuaries or in the ocean. Such
the river has been extensively diked and channeled, the subyearling migrant coho, may constitute significant
flood plain in the freshwater-tidal region is characterized portions (up to 50%) of the returning adult spawners.523
by extreme habitat complexity, abrupt changes in water Natural disturbance regimes are responsible for
velocity and low-velocity off-channel habitats. As the creating and maintaining habitat complexes important to
“estuarine gateway,” the tidal-freshwater mixing zone can juvenile salmon. Erosive flooding, channel
be exceedingly important to juvenile salmon469 because it: reconfiguration, and changes imposed by LWD all
(1) provides habitat for overwintering chinook, coho and promote increased habitat complexity and heterogeneity.
steelhead forced downstream during high river flows; (2) In the absence of disturbance, early successional habitats
contains complex low-velocity refugia such as off-channel such as mudflats and low elevation estuarine marshes
sloughs and LWD; (3) allows migrating juveniles to adapt (e.g., Carex lyngbyei sedge) would not persist or would be
physiologically as they encounter brackish waters of the relatively rare. Yet, these habitats can be some of the most
upper estuary; (4) drifting insects are trapped and productive and beneficial of salmon habitats and play
concentrated due to flow reversals, providing opportune unique roles for some salmon species. A number of
feeding conditions;523 and (5) is the first region of estuarine important studies have described the association with
settling of suspended sediments and detritus, which can specific migratory, rearing and residency times of salmon
fuel soft-sediment habitat formation and detritus-based species in estuaries.104, 192, 196, 197, 227, 247, 256, 273, 460, 466, 467, 565
food webs exploited by salmon.
Estuaries are composed of both discrete and highly
integrated habitat complexes and their associated plant
and animal communities. Categorizing habitats to a large
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 639

Ocean Habitat North Pacific Ocean vary among species and sizes of fish,
Upwelling along the coast of the Pacific Northwest often with season and year, and with location and proximity to
results in high primary and secondary productivity, the coast. Fishes, squids, amphipods, copepods, and
resulting in large standing stocks of fishes, seabirds, and pteropods are primary prey.178, 268, 396
marine mammals. The coastal upwelling domain extends
from British Columbia to Baja California, and is located Ecological Relationships of Salmon
inshore of the equatorial flowing California Current.
Coastal upwelling is driven by prevailing northwesterly
Macroinvertebrates
winds during the spring and summer. Those winds result Freshwater Macroinvertebrates and Salmon
in offshore displacement of near-shore surface waters and Freshwater ecosystems are inhabited by a large variety of
vertical advection of deep, cool and often nutrient-rich macroinvertebrates that play an integral part in the
waters into the euphotic zone along the coast and into salmon’s life history. They include insects, crustaceans, and
estuaries. Rich blooms of phytoplankton are observed other forms of macroinvertebrates (larger than 595 microns
along the coast following episodic upwelling events.286, 501, in their later instars or mature forms). Many species in
514
Upwelling varies seasonally and over longer annual their aquatic phase have been described from the
and semiannual cycles, with intensity generally increasing hyporheic zone, or zone below the surface of the stream
southward to northern California. Upwelling is most bottom.492 Given the variety of physical habitat across the
intense in regions of capes such as Cape Blanco in southern region’s landscape, there is an opportunity for freshwater
Oregon. There was a strong correlation between the invertebrate species to form diverse and specialized
intensity of coastal upwelling and the smolt-to-adult communities.
survival of hatchery coho salmon from the Oregon Freshwater macroinvertebrates play a significant role
Production Index region south of the Columbia River from in energy pathways of aquatic ecosystems. The
1960 to 1981.373 During the late 1970s, however, there was consumption of algae, detritus, and bacteria is the basis
a major change in ocean climate in the North Pacific Ocean, for transfer of this energy. A few invertebrate species are
called a regime shift or the Pacific Decadal Oscillation, known to actively derive their food base from higher life
which was correlated with a sharp decline in the survival forms (e.g., small fish). The food source used by an
of Oregon coho salmon between smolt release years 1975 invertebrate defines what function it performs in this food
and 1976.32, 153, 397 After this regime shift the production of web.
Oregon coho salmon has usually been low. The
relationship between coastal upwelling and coho survival Structure and Function in Macroinvertebrate
is no longer significant. The reason for this changed Communities
relationship is unclear, but is related to weak coastal The type and location of food in the aquatic environment
upwelling, warm sea temperatures, high sea levels, and consumed by invertebrates determines their functional
frequent El Niño events.394 Upwelling has probably not designation. Headwater streams or heavily canopied
been effective in injecting nutrient-laden water into the streams are dominated by leaf litter input, allochthonous
euphotic zone because of the deep lens of overlying warm, (biomass produced from outside the stream) material,
nutrient-depleted water along the coast. 195, 432 The which has been linked to significant shredder activity.111,
persistence of warm, unproductive ocean conditions is a 536
Shredders comprise a group of aquatic insects that
major reason for the decline of many stocks of anadromous utilize coarse particulate organic matter, such as leaf litter,
fishes along the west coast, and for the very large with a significant dependence on the associated microbial
variability in survival and reproduction of marine birds. biomass.529 Portions of a drainage where the riparian
Although the mechanisms that have resulted in poor canopy opens can result in substantial autochthonous
ocean survival of salmon are speculative, one hypothesis input (periphyton growth), and are consumed by scrapers
is that weak upwelling results in low growth and poor like the mayfly family Heptageniidae.315 Lower in a
survival of zooplankton and forage organisms, and drainage the channel can accumulate large deposits of
impacts juvenile salmon during their critical first summer detritus. Invertebrates distributed here are mainly
in the ocean. This lack of forage and the narrow band of collector-gathers and may constitute the bulk of juvenile
cool waters along the coast during weak upwelling years salmon diets.196 The distribution of dominant food sources
concentrates juvenile salmon near the coast where they throughout a drainage are influenced by a continuum of
are more vulnerable to predation by seabirds, marine physical changes as one travels from the steep headwater
mammals and fishes.145 During warm years predators from streams to the relatively low gradient flood plains.529
southern waters, e.g., Pacific and jack mackerel, invade Invertebrate community structure in a stream or pond
coastal waters and may either compete with or prey upon reflects physical characteristics of the living space.
juvenile salmon.394, 397 Numbers of species in a stream ecosystem are usually
The principal prey of juvenile salmon off the coast of greater in physically diverse habitats. Structural attributes
Oregon and Washington during the spring and summer like species richness change along a disturbance gradient.
are fishes and crustaceans.65b, 395 Salmon in the open ocean Two investigations found that species richness was
forage opportunistically on a diverse assemblage of consistently higher in streams with intermediate
pelagic organisms. The diets of maturing salmon in the disturbance of substrate.116, 519 The effect of disturbance and
 640 Wildlife–Habitat Relationships in Oregon and Washington

physical change over a continuum results in species Significance of Macroinvertebrate Life Cycles
replacement and sometimes adjustments of the functional There are two life strategies characteristic of freshwater
characteristics in the community.325 macroinvertebrate species.558 The simpler hemimetabolous
strategy inherent in stonefly and mayfly species contain
Physical and Chemical Influences on an egg, multiple nymph, and adult stages. A few of the
Macroinvertebrate Distribution stonefly species are long-lived (more than a year) in the
Factors that control distribution and abundance of aquatic nymphal form. Large-bodied stoneflies found in
macroinvertebrates are substrate, current velocity, streams indicate adequate flow in channels that are key
temperature, predators, and food resources.223 Substrate to survival of early salmon life stages and to some of the
heterogeneity often promotes greater species richness.326, invertebrate fauna they will eventually consume.
328
Interstitial spaces in stream gravels can serve as refuge The second life strategy contains representatives of the
from predators and physical disturbance, and entrap holometabolous invertebrates. Midges, blackflies, and
detritus. Water temperature in the interstitial caddisflies have egg, larva, pupa, and adult life stages.
microenvironment can be relatively constant and cooler These types are mostly short-lived having one or many
than the overlying surface water.557 generations per year in a population. Aquatic
Early life stages of the salmon can be affected by environments that are seasonally stressed by high
substrate quality. Factors that favor survival of salmon temperatures, low dissolved oxygen, or drought are
eggs and fry (low levels of fine sands and silts) are primarily colonized by holometabolous invertebrates.
coincident with requirements of aquatic invertebrates that These stressors increase the mortality of early life stages
have a narrow tolerance range to environmental in salmon, but encourage dominance of holometabolous
fluctuations. Protection from natural physical disturbance species in the aquatic invertebrate community.
is important for early life stages of salmon and mobile
aquatic invertebrates. Stable stream bottoms during Aquatic Macroinvertebrates as a Food Source for
periods of flood or freshet reduce predation on dislodged Salmon
animals. In some instances, salmon redd construction is a Aquatic ecosystems are frequently inhabited by both
natural disturbance that reduces invertebrate density in hemimetabolous and holometabolous macroinvertebrates.
localized areas of a stream.324 This disturbance also opened The hemimetabolous species richness is greater in mid-
niche space for other functional groups of aquatic insects, to upper-drainage streams and play a larger role in the
like blackflies, who feed on suspended particles and diet of juvenile chinook197 and coho salmon.442 Although
recolonized quickly along with stonefly nymphs and holometabolous invertebrates are dominant in lower-
midge larvae.324 Other invertebrates that enter the drift drainages, species in the family Chironomidae are present
behaviorally or unintentionally from substrate disturbance in all habitats and are a significant food source to salmon
are potential prey items for feeding salmon. Mayfly and in early freshwater stages.100, 399, 440
stonefly density and richness can be reduced by physical All species of salmon fry consume some life stages of
alterations to the stream corridor. These changes may have dipterans, primarily Chironomidae, during the freshwater
significant implications to the salmon food base. life phase.178 Stonefly and mayfly nymphs are consumed
Invertebrate drift is either voluntary, a behavioral by pink, chum, and chinook salmon fry. Coho fry are
activity, or coincides with catastrophic stream conditions, suspension and surface feeders whose diet is
especially during floods. Taxonomic groups prominent in predominately terrestrial insects. Ecologically important
behavioral drift are amphipods, Ephemeroptera freshwater invertebrates in coho natal habitat are emerging
(mayflies), Plecoptera (stoneflies), Tricoptera (caddisflies), and flying insects such as mayflies, stoneflies, and midges
and Simuliidae (blackfly larvae). Later stages of the nymph (Chironomidae). The rapid migration of chinook fry to
and larval forms are most active in the diel (24-hour cycle) the river estuary introduces terrestrial homopterans (leaf
drift.547 Behavioral drift occurs with a diel periodicity, hoppers and aphids) into their diet. Additional details of
typically at two peaks in a 24-hour time frame. Most prey items during the freshwater cycle have been
invertebrates that enter the drift are night-active, with described.95, 155, 178, 294, 338, 399, 449, 461, 466 The influence of riparian
photoperiods as the major cue. Fewer invertebrates are vegetation along streams and estuaries appears to be an
day-active and begin drifting by cues through change in important factor in determining abundance and type of
water temperature. terrestrial insects on which salmon forage.
Drifting invertebrates are a food source for certain
species of fish that forage in the stream water column. Salmon as a Food Source for Aquatic
Rader415 determined that the mayfly genus, Baetis, whose Macroinvertebrates
drift propensity was high, was a significant food source Freshwater macroinvertebrates such as caddisflies,
to juvenile and adult salmon. Other studies indicated that stoneflies, and midges are involved in processing the
food preference of juvenile fish was related to its microbially conditioned salmon carcass. Bilby et al.47
abundance and location within the stream channel. observed a significant contribution of nitrogen from
Juvenile coho salmon diet varied seasonally depending spawning salmon to the collector-gatherer invertebrate
on the type and abundance of invertebrates, salmon fry, community. (Photograph 7) Increases in aquatic
or salmon eggs in the benthos or drift.260 invertebrate density from the introduction of salmon
carcasses564 stimulated feeding by early life stages of select
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 641

Photograph 7. Aquatic insects feeding on a

salmon carcass. (Photo by Jason Walter and
Brian Fransen)

salmon species.48 Other stages of the salmon life history food-web that knits the water and land together. Wildlife
contribute to the invertebrate food base. Nicola376 observed species have likely had a very long, and probably co-
the stonefly nymph, Alloperla, scavenging dead pink and evolutionary, relationship with salmon in the Pacific
chum salmon embryos and alevins. Also, Elliott and Northwest. In their Natural History of Washington Territory
Bartoo 131 found the midge, Polypedilum (Diptera), and Oregon, Suckley and Cooper502 wrote of the California
associated with dead pink salmon embryos and alevins. condor:
Freshwater invertebrate shredder abundance increases The Californian vulture visits the Columbia river in
in the presence of salmon carcasses.564 Non-salmon- fall, when its shores are lined with great numbers of
bearing streams support a limited abundance of shredders dead salmon, on which this and the other vultures,
mediated through input of leaf matter. This organic food besides crows, ravens, and many quadrupeds, feast for a
base must first be conditioned by the microbial community couple of months.
to increase palatability to shredders. Cool water
temperatures characteristic of coastal streams slow the The “Five Mile Rapids” prehistoric archaeological site
microbial decay of the leaf litter food source resulting in along the banks of the Columbia River, five miles east of
limitations in distribution and abundance of the shredder the Dalles, Oregon, yielded bones from at least 63
community. The appearance of salmon and the additional individual California condors, plus remains of turkey
influx of biomass to streams appears to be a controlling vultures, cormorants, bald eagles, and gulls.110 Carbon-14
factor for shredder species. However, the role of shredders dating placed materials at this site from 10,000 to 7,500
in the presence of salmon carcasses continues to be years before present.321, 474 Miller321 suggested that these
investigated. Bilby et al.47 found no significant concen- birds were attracted to the site by the presence of abundant
trations of carbon contributed from decaying carcasses in living and dead salmon and human refuse resulting from
the shredder community. Undigestable animal tissue fishing.
consumed by shredders was excreted as fine particulate The life cycle stages of salmon (i.e., eggs, fry, smolts,
organic matter (FPOM). Nutritive food value for shredders adults, and carcasses) all provide direct or indirect foraging
may have been derived primarily from the microbial opportunities for terrestrial, freshwater, and marine
community on decaying carcasses. Aquatic insects of the wildlife (Photograph 8). While sometimes abundant and
collector-gatherers group typically benefit secondarily somewhat dependable from year to year, the availability
from the activity of shredders.316, 47 of salmon to wildlife is largely seasonal in nature. The high
The relationship between invertebrates and salmon can seasonal variability in a particular food resource is
be complex. Functions of invertebrates have not yet been reflected in the opportunistic foraging of many wildlife
fully defined, but we know they are essential to salmon consumers—however, “opportunistic” is not a synonym
survival. Invertebrates complete a loop beginning as for biological unimportance. Thus, one could hypothesize
recipients of food from adult salmon carcasses that, in turn, that while many wildlife species could develop important
fuel the growth and survival of early stages in the salmon’s food-web relationships with salmon, few wildlife species
life cycle. would likely be able to form an ecological “dependance”
on salmon. Only those species which are highly mobile,
Vertebrate Wildlife or are able to capture, consume, and store (in body tissues)
substantial quantities of salmon biomass in a short period
Vertebrate Wildlife and Salmon of time would be likely to develop a strong ecological
Anadromous salmon provide a rich, seasonal food dependance on salmon. It is more probable that the
resource that directly affects the ecology of both aquatic majority of wildlife which directly consume salmon will
and terrestrial consumers, and indirectly affects the entire
 642 Wildlife–Habitat Relationships in Oregon and Washington

Photograph 8. Juvenile glaucous-winged

gull eating a chum salmon (O. keta)
carcass at Kennedy Creek, Washington.
(Photo by Jeff Cederholm)

have flexible foraging strategies, utilizing salmon when anadromous fish stocks could have major effects on the
available, and alternate food sources during other times population biology (i.e., age class, longevity, dispersal
of the year. ability) of many species of wildlife, and thus, on the overall
Indirect relationships develop when a food resource is health and functioning of natural communities over the
providing foraging opportunities to a secondary majority of the region.
consumer. An example in our case is reflected by peregrine Research on predator-prey interactions in which
falcons which eat gulls that feed on salmon carcasses. As anadromous fish are the prey has strongly emphasized
salmon are a concentrated resource, this will serve to the effects of predation on the fish populations.225, 331, 430, 559,
570, 571, 576
concentrate otherwise dispersed wildlife species (e.g., Many existing studies describe predatory species
bears). In this scenario, there may well be competition, as competitors of human harvesters and attempt to control
parasitism, or other aggressive interactions between or the rate of predation to maximize human consumption.
among wildlife species. Some of these interactions, e.g., We hope to reverse this perspective by focusing on the
bald eagles disturbing common mergansers, serve to important interplay between salmon and wildlife
benefit salmon by reducing predation. The magnitude of populations. In the following sections, we discuss the
the salmon-wildlife interaction warrants special relationships between salmon and their vertebrate
examination and calls attention to the pervasive consumers, and the salmon’s role in enhancing ecological
occurrence of these important ecological functions and functions involving wildlife in terrestrial, freshwater, and
linkages across the region. The loss or severe depletion of marine systems.

Table 2. Relationship between Pacific salmon and 605 species of wildlife in Washington and Oregon.

Relationship
Salmon life stage Strong, Consistent Recurrent Indirect Rare Unknown None

Incubation—eggs 2 10 1 10
and alevin(23)
Freshwater Rearing— 4 31 4 10
fry, fingerling, parr (49)
Saltwater—smolts, 6 36 5 19
immature adults,
adults (63)
Spawning (16) 5 10 0 1
Carcasses (82) 5 28 22 38
(9) (58) (25) (64) (60) (408)

There were 137 species with a positive relationship with salmon (i.e., combined total for species with Strong, Consistent, Recurrent,
Indirect, and Rare relationships). The total number of individual wildlife species for columns and rows are shown in parenthesis; the
number of species shown in the rows and columns may not equate to the numbers shown as totals as 19 species had more than one
type of relationship with salmon, and 73 species are associated with salmon at more than one life stage.
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 643

Wildlife Species with a Relationship to Salmon between an aerial insectivore and salmon was supported
Johnson et al.233 examined the relationships between the by the presence of two or more of the following
Pacific salmon and 605 species of terrestrial and marine characteristics of the insectivore: (1) riparian obligate or
mammals, birds, reptiles, and amphibians currently or associate, (2) feeds below or near the canopy layer of
historically common to Washington and Oregon. They riparian trees, (3) known or perceived to feed on midges,
found a positive relationship between salmon and 137 blackflies, caddisflies, stoneflies, or other aquatic insects
species of wildlife, the relationship was “unknown” for that benefit from salmon-derived nutrients, and/or (4)
60 species, and a determination of “no relationship” was feeds near the water surface. While this category includes
made for 408 species (Table 2). Where a relationship general aspects of salmon nutrient cycling in stream/river
existed, they identified both the type(s) of relationship and systems, we are not including or examining the role of
the stage(s) of the salmon life cycle to which it applied. Of carcass-derived nutrient cycling on lentic system riparian
the 137 species with a relationship to salmon, 9 species and wetlands vegetation, and subsequent links to wildlife.
were categorized as having a Strong, Consistent
4) Rare Relationship. Salmon play a very minor role in
relationship (Appendix I), 58 as Recurrent (Appendix II),
the diet of these species, often amounting to less than 1
25 as Indirect (Appendix III), and 64 as Rare (Appendix
percent of the diet. Typically, salmon are consumed only
IV). (This tally totals more than 137 because 19 species
on rare occasions, during a shortage of the usual food and
had more than one type of relationship with salmon.)
may be especially evident during El Niño events. As
Of the 137 wildlife species, 88 were characterized as
salmon are often present in large quantities, they may be
having a routine relationship (combination of species with
consumed on rare occasions by species that normally do
Strong, Consistent, Recurrent; and Indirect) with salmon. Of
not consume them. Examples: red-tailed hawks are known
these 88 species, there were 25 mammals (8 of these were
to consume salmon carcasses in times of distress;
marine mammals), 60 birds, 2 amphibians, and 1 reptile.
trumpeter swans are primarily vegetarians, but on rare
The relationship categories are briefly described as
occasions will consume eggs, parr, as well as salmon
follows:
carcass tissue.
1) Strong, Consistent Relationship. Salmon play (or
5) Unknown Relationship. A relationship between this
historically played) an important role in this species
species and salmon may exist, but there is not enough
distribution, viability, abundance, and/or population
information to determine the scope or scale of the
status. The ecology of this wildlife species is supported
relationship at this time. Example: while it is logical to
by salmon, especially at particular life stages or during
speculate that riparian feeding bats may feed on salmon-
specific seasons. Timing of reproductive activities, and
derived insects, aspects of seasonality of both bats and
daily or seasonal movements often reflect salmon life
salmon carcasses are relevant, as is the nocturnal flight
stages. Relationship with salmon is direct (e.g., feeds on
behavior of the insects. Do bats and salmon carcasses
salmon, or salmon eggs) and routine. The relationship may
coincide seasonally, and if so, are salmon-derived insects
be regional or localized to one or more watersheds.
actually available to feeding bats? At this time, the
Examples: a significant portion of the diet of killer whales
evidence for this relationship is inconclusive and remains
is adult salmon (Saltwater stage); common mergansers may
to be examined.
congregate to feed on salmon fry (Freshwater Rearing stage)
when they are available. 6) No Relationship. There is no recognized or apparent
relationship between salmon and this species.
2) Recurrent Relationship. The relationship between
As part of the same study, Johnson et al.233 reported 60
salmon and this species is characterized as routine, albeit
species as having an “unknown” relationship with salmon
occasional, and often tends to be in localized areas (thus
(Appendix V), suggesting that the diets of these species
affecting only a small portion of this species population).
in Oregon and Washington, were not understood well
While the species may benefit from this relationship, it is
enough to characterize their relationship with salmon.
generally not considered to affect the distribution,
Additional observations on the diets of these species will
abundance, viability, or population status of this species.
help determine whether the relationships of these species
The percent of salmon in the diet of these wildlife species
with salmon is routine, a rare and unusual event, or
may vary from 5% to over 50%, depending on the location
whether a relationship exists at all. Johnson et al.233
and time of year. Example: turkey vultures routinely feed
identified 408 species as having “no relationship” to
on salmon carcasses, but feed on many other items as well.
salmon.
3) Indirect Relationship. Salmon play an important,
routine, but indirect link to this species. The relationship Wildlife Response to Salmon Congregations
could be viewed as one of a secondary consumer of The numerical response of predators to salmon
salmon; for example, salmon support other wildlife that congregations is often substantial, sometimes
are prey of this species. This includes aspects such as spectacularly so. The ability of wildlife species to
salmon carcasses that support insect populations that are concentrate at salmon sites is more than opportunistic
a food item for this species. Example: American dippers foraging, it provides for interaction of important ecological
feed on aquatic insects that are affected by salmon-derived processes. Anadromous fishes (including their eggs) are
nutrients. The hypothesis of an indirect relationship a major source of high-energy food that allows for
 644 Wildlife–Habitat Relationships in Oregon and Washington

Table 3. Wildlife species that have been observed or successful reproduction and enhanced survival of adults
are perceived to aggregate at salmon congregations and juveniles of many wildlife species, and support for
in Oregon and Washington. long-distance migrant birds. Wildlife movements to
salmon congregations can be seasonal (e.g., bald eagles
American crow along the Skagit River in Washington), or depending on
American white pelican the situation (e.g., hatchery fish released during an El Niño
Arctic tern high food-stress seabird breeding season) can occur within
Bald eagle a matter of hours. Perhaps as noteworthy, but much harder
Bank swallow to detect, is that some wildlife species that have been
Barn swallow reported to group at salmon sites in other areas (e.g., black
Barrow’s Goldeneye bears in southeast Alaska) do not appear to be doing so
Black beara with any regularity in Washington and Oregon. This may
Black-billed magpie well be reflecting the depressed nature of some salmon
Brandt’s cormorant stocks rather than the inherent behavior of the wildlife
Brown pelican species. Of the 88 species with a link to salmon,233 43 species
California gull (37 birds, 6 mammals) concentrate or form loose
California sea lion
aggregations at salmon sites (Table 3). Some reasons why
Caspian tern
other species do not congregate at salmon sites are: strong
Clark’s grebe
territoriality (e.g., great blue heron), foraging strategies
Cliff swallow
Common goldeneye
which require above-water structures or perches (e.g.,
Common merganser belted kingfisher), and limited movement capabilities (e.g.,
Common murre shrews).
Common raven That wildlife concentrate at salmon areas is well
Common tern established for some species, and for others we may be
Double-crested cormorant witnessing signs of the impacts of salmon declines. We
Elegant tern offer the following examples relevant for Oregon and
Forester’s tern Washington.
Glaucous gull
Bald Eagle. Suckley and Cooper502 state:
Glaucous-winged gull
Grizzly beara This noble looking bird is exceedingly abundant in
Harbor seal Oregon and Washington Territories, and in certain
Herring gull localities, especially during the salmon season, may be
Killer whale found in great numbers.
Northwestern crow
Northern rough-winged swallow The North Fork of the Nooksack River (coastal
Northern (Steller) sea lion Washington) currently hosts one of the largest and most
Red-breasted merganser visible concentrations of wintering bald eagles in the lower
Rhinoceros auklet forty-eight states. Peak concentrations (100 or more eagles)
Ring-billed gull occur along the Nooksack263 and Skagit218 rivers with
Thayer’s gull December- and January-spawning chum salmon.
Tree swallow
Tufted puffin Caspian Tern. The first breeding record of Caspian terns
Turkey vulture along the Oregon/Washington coast was a colony of 50
Violet-green swallow pairs in Grays Harbor, Washington in 1957.6 This, and
Western grebe other nesting colonies (mid-1950s through early 1990s)
Western gull along the Washington coast in Grays Harbor, Willapa Bay,
and near the mouth of the Columbia River, have been
a
now questionable; there may not be enough salmon in Oregon
abandoned or destroyed by human actions.429, 430, 479 A
and Washington for bears to gather at a site.
colony of Caspian terns originally settled on Rice Island,
a dredge material disposal island in the lower Columbia
River, in 1987. In 1997, an estimated 14,000 terns used this
island for nesting and/or roosting during the 80-100 day
(April-July) breeding season.429, 430 This represents the
largest known colony of Caspian terns in North America,
and possibly the world. In 1997, the terns appeared to be
largely dependent on juvenile salmon (roughly 75% of the
diet), consuming an estimated 14.5 million smolts, the
majority being hatchery fish.429, 430 Tern nesting success was
very low (roughly 5%) in 1997; predation on adult terns
by bald eagles, and gull predation on tern eggs and chicks
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 645

(caused by eagle and researcher disturbance) were the pers. comm.) and Oregon528 have been found to move to
primary causes.429, 430 and congregate at higher elevations in the fall to feed on
huckleberries (i.e., forming “traditional use areas”). Thus
Common Murre. The common murre is a seabird that
one could reasonably conclude that if salmon were to be
nests in large colonies along the Oregon coast; colonies in
found in substantial and predictable numbers, bears in
Washington have undergone significant declines in the
Oregon and Washington, like those studied by Chi in
last decade. It is only an occasional consumer of salmon;
Alaska, would also establish traditional use areas around
the vast majority of its diet is other small marine fishes. A
salmon. Black bears in western Washington typically den
severe El Niño event occurred in 1983 during the seabird
by 1 November and emerge around 1 April, thus salmon
nesting season along the eastern Pacific coast, with the
runs occurring during the winter will not be available to
majority of common murres (and other seabirds species)
bears. Recent bear studies in western Washington have
either not attempting to nest or abandoning their nests
included the Humptulips, Wishkah, Wynoochee, and
once initiated. 27, 210 Adult survival was also greatly
Quinault Rivers, and while these hold low levels of
reduced.169 Oregon Aqua-Foods, Inc. had released a total
hatchery-based salmon, bears do not congregate along
of 2 million or more salmon smolts into the Yaquina
them (G. Kohler, pers. comm.). D.H. Johnson (unpubl.
Estuary at roughly 2.5 day intervals between June and
data) summarized 1990-1998 hatchery return data of adult
August since 1977.28 Murres were more numerous at the
salmon for the Humptulips river system in western
mouth of Yaquina Estuary for the first two days post-
Washington. These fish return to the hatchery facility as
release in July of 1983 than in July of 1982 (a non-El Niño
early as late-September (most begin around mid-October),
year), as they were drawn in to feed on the released
and as typical with most, were done spawning by mid-
salmon. First day post-release averages of murres were
December. While there are additional fish in this system,
3,710 in 1983 and 3,053 in 1982. In August of 1983 however,
an average number of 217 (range 95-320) chinook, 6,496
murre numbers were significantly less than in 1982
(range 177-10,195) coho, and 165 (range 51-339) steelhead
(average 1983=106; 1982=1,860), as murres had begun
returned annually to the hatchery. The substantial majority
moving north earlier to feed on other food resources.27 In
of these fish species return to spawn after 1 November
summary, although not as a primary food resource, murres
(the average date of bear denning) and are not available
will make use of salmon resources during food-stress
to bears; an average of 73 chinook, 1,264 coho, and 0 (zero)
conditions.
returning steelhead were available to black bears. Here,
Black Bear. Contrary to popular image, Washington and “available” means simply present in the river system, and
Oregon black bears only rarely congregate at salmon sites. not located at spawning redds. In summary, bears have a
Poelker and Hartwell406 reported on three diet studies of strong relationship to salmon where they have access to
black bears in western Washington for the time periods of them, but it appears that in substantial measure, current
1952-54 and 1968, and found that fish represented 5.0% of salmon populations do not represent a predictable food
the diet. In their treatise on land mammals, Verts and supply to bears in Washington and Oregon.
Carraway535 describe black bears in Oregon as being
largely herbivorous, and do not mention salmon as part Review of Wildlife Relationships by Salmon
of their diets. Cederholm et al.91 found black bears on Life Stages
Washington’s Olympic Peninsula to heavily consume For the 137 species with a relationship to salmon, Johnson
salmon carcasses. In northern California, Kellyhouse244 et al.233 identified the salmon life stage(s) involved for each
found evidence of salmon in 10% of black bear fecal species. In this study, the five general life history stages of
samples analyzed from spawning areas. The California salmon were identified as: (a) Incubation (egg and alevin);
Department of Fish and Game84 also reported black bears (b) Freshwater Rearing (fry, fingerling, and parr); (c)
taking advantage of anadromous fish runs. The strong link Saltwater (smolt, subadult, adult); and (d) Spawner; and
between black bears and salmon was demonstrated in the (e) Carcass. The number of wildlife species associated with
Anas Creek drainage of southeastern Alaska (D. Chi, pers. each (in parenthesis) were: Incubation (23); Freshwater
comm.). This effort studied the behavior and activity Rearing (49); Saltwater (63); Spawning (16), and Carcass (82);
patterns of black bears (n=40 individuals) which had this tally of wildlife species totals more than 137 because
established movement patterns according to salmon 73 species are associated with salmon at more than one
migrations. Bears arrived in the lower reaches of the creek life stage (Appendixes I, II, III). See Appendix VI for a list
in June to begin feeding on spawning salmon, and stayed of published and unpublished observations of wildlife
through August and early September to feed on salmon predators and scavengers on salmon.
carcasses. Thirteen of the bears were radio-marked and
Incubation Stage (egg and alevin). Twenty-three wildlife
their movements indicated that they were moving in from
species are linked to salmon at this stage. Twenty-two
at least eight miles (12.9 km) away. Other bears were
wildlife species are direct consumers of “drift eggs” (eggs
assumed to be coming in from further away. The general
not buried in redds) or alevin (2 amphibians, 1 reptile, 19
lack of salmon in the published accounts of black bear
birds, and 1 mammal); and 1 bird (bald eagle) is an indirect
diets across Washington and Oregon exception: see91 is
consumer of eggs/alevin, feeding on the waterfowl that
somewhat counter to the observed salmon use in adjacent
consume eggs and alevin.
regions. Radio-marked bears in Washington (G. Kohler,
 646 Wildlife–Habitat Relationships in Oregon and Washington

Photograph 9. Garter
snake eating a
salmon smolt,
unknown Olympic
Peninsula stream,
Washington. (Photo
by Jim Rozell,
deceased)

Freshwater Rearing (fry, fingerling, and parr). Forty-nine Continued documentation of wildlife species—salmon
wildlife species are linked to rearing salmon, including 2 interactions, especially of the 60 species having an
amphibians, 5 reptiles, 34 birds, and 4 mammals. Forty- “unknown” relationship, will provide vital information
five of these species are direct consumers of salmon and 4 for ongoing developments in ecologically-based salmon
species (bald eagle, gyrfalcon, peregrine falcon, and snowy spawner escapement research and prescriptions for
owl) are indirect consumers, feeding on terns, waterfowl, riparian management practices. As Key Ecological
gulls, and other animals that eat rearing salmon Functions (KEFs) are identified through such research,292
(Photograph 9). tools for informed decisions will be made available to fish
and land managers operating under an ecosystem context.
Saltwater (smolt, subadult, adult). Sixty-three wildlife
species are consumers of salmon at this stage (51 birds Key Ecological Functions (KEFs) Provided to
and 12 marine mammals). Fifty-eight of these species are
the Ecosystems through Salmon—Wildlife
direct consumers of salmon and 5 species are indirect
consumers. This list is somewhat expansive due to the Interactions
geography being included, that is, the estuarine and all In striving to manage for healthy and sustainable
marine water habitats. ecosystems, we simultaneously are striving to provide for
the full range of ecological functions that these systems
Spawner. Sixteen species of wildlife are consumers of provide. Key ecological functions (KEFs) refers to the main
spawning salmon (6 birds and 10 mammals). This list is ecological roles of a species (or group of species) that
relatively small, as few wildlife species are physically influence diversity, productivity, or sustainability of
capable of capturing and handling live, adult fish. The ecosystems.334 A given KEF can be provided by a single
gray wolf and grizzly bear are on this list, but both have species or shared by many species, and a given species
undergone significant range contractions and declines in can have several KEFs. Main categories of KEFs include
their abundance (i.e., both are extirpated from Oregon and trophic relations; herbivory; nutrient cycling; interspecies
significantly reduced in Washington). relations; disease; pathogen and parasite relations; soil
Carcass. Carcasses are linked to the largest group of relations; wood relations; water relations; and vegetation
wildlife consumers of any salmon life cycle stage, with 82 structure and composition relations. Building upon work
species (1 reptile, 50 birds, and 31 mammals) being by Marcot et al., 291 Marcot and Vander Heyden 292
consumers of carcasses and/or carcass-derived insects. characterized the key ecological functions for each of the
Body sizes of these animals range from shrews to grizzly 605 common wildlife (i.e., terrestrial and marine birds,
bears. Seventy-two species of wildlife (1 reptile, 38 birds, mammals, reptiles, and amphibians) species in Oregon
and 31 mammals) are direct consumers of carcasses; 22 and Washington. Several questions can be thus posed:
species (14 birds and 8 mammals) are consumers of • In what way does providing for salmon also provide
carcass-derived insects; and 10 species (2 birds and 8 for a wider array of ecological functions of wildlife
mammals) are consumers of both carcasses and carcass- species associated with salmon?
derived insects. • What are those functions?
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 647

• How do different kinds of salmon-wildlife relations, 9 species inhabit freshwater and marine habitats, but some
and different salmon life stages, provide for an array of them also occur across the range of inland forest,
of ecological functions? woodland, shrubland, and grassland habitats. It is of
This somewhat innovative analysis describes the interest that from 1-7 of these 9 species can be found in
functional links among fish and wildlife species across each of the 32 habitats. In this way, salmon provide for a
aquatic and terrestrial communities. To conduct this set of wildlife species that occur well beyond just salmon-
analysis, we queried the database matrixes on salmon- inhabited aquatic systems.
wildlife relations, key ecological functions of wildlife In addition, the full set of key ecological functions
species, and habitats used by wildlife.236 The general performed by these 9 species also extends beyond the
conclusion is that salmon provide a causal mechanism for aquatic system. Each of these species provides a set of
movement behaviors and a nutrient source for a variety ecological functions to the various array of habitats that
of wildlife species which in turn perform a surprisingly they occur within. Figure 5 depicts the collective range of
broad array of ecological functions292 across a wide span ecological functions that these 9 wildlife species provide
of habitats. For this analysis, one can think of the array of to the number of habitats that they occupy. The functions
ecological functions performed by these wildlife species range from various trophic, organismal, and wood and
as a “functional web.” It focuses on salmon in their various soil relations. Some functions are more widespread (occur
life stages, and extends well beyond the aquatic realm to in more habitats) than are other functions. Examples of
influence the diversity, productivity, and ultimately some widespread functions are: potential control of
sustainability of habitats and ecosystems throughout vertebrate populations (through predation), carrion
Washington and Oregon. feeding, piscivory (fish-feeding), invertebrate feeding
(including insectivory), omnivory, transportation or
Wildlife with Strong Consistent Links to Salmon. The 9
dispersal of seeds and animals, creation of terrestrial
species of wildlife with strong consistent links to salmon
runways used by other species, and secondary use of
(bald eagle, American black bear, Caspian tern, common
burrows created by other species.
merganser, grizzly bear, harlequin duck, killer whale,
osprey, and river otter) comprise a functional group of All Wildlife with Links to Salmon. What of the full set
“salmon-eaters” with close affinities to salmon. There are of species showing either strong consistent, recurrent,
32 primary wildlife-habitats across Washington and and/or indirect links to salmon? (Some species have more
Oregon236; Figure 4 summarizes the occurrence of these 9 than one type of relation because they use more than one
wildlife species by habitat. Not surprisingly, most of these salmon life stage). Table 4 lists key ecological functions of

Figure 4. Occurrence by number

of vertebrate wildlife species in the
32 wildlife habitats in Washington
and Oregon, as used by the 9
wildlife species with a strong
consistent relationship to salmon.
 648 Wildlife–Habitat Relationships in Oregon and Washington

Figure 5 . The array of key

ecological functions performed
by the 9 vertebrate wildlife
species with a strong consistent
relationship to salmon, across
the 32 wildlife habitats in
Washington and Oregon.

wildlife more or less unique to each type of salmon- Managing the Functional Web. So what is the manager
wildlife link. Each of the three types of relations provides to do with this information? For one, be aware that salmon
for some unique set of ecological functions. For example, can be viewed as the center of a broad “functional web”
wildlife species indirectly linked to salmon can provide of wildlife and their ecological roles. Such roles extend
the following ecological functions: fungivory (fungus- well past the salmon populations and aquatic habitats
eating); tertiary consumption or secondary predation; prey themselves, and likely influence the structure and
source; regulate insect populations through predation; processes of the communities and ecosystems in which
serve as interspecific host for avian nest parasites; and they reside, thus a “keystone” species.559
create primary small ground burrows. These functions are Second, one can use the information presented here and
either not performed, performed by far fewer wildlife in the species data matrixes to list the collective set of
species, or by the wildlife with occasional or strong habitat elements and conditions used by wildlife species
consistent links to salmon. associated with salmon. For example, one can link the list
What this means is that different degrees of salmon- of wildlife associated with salmon life stages likely to be
wildlife relations provide for some unique kinds of wildlife found in low order headwater streams, and determine the
ecological functions. Only the full set of all wildlife-salmon set of habitat elements used by this set of wildlife species,
link relations can provide for all collective functions. Thus, by habitat type, and then establish habitat-specific
to manage the full set of all ecological functions, one management guidelines to provide for such habitat
should not focus solely on those few wildlife species with elements over time. Maintaining such habitat elements
strong consistent links to salmon, but on all types of links. and conditions would help maintain the full salmon-
wildlife functional web.
How Salmon Life Cycle Stages Provide for Ecological
Third, one can begin to predict—or, at least pose
Functions. In a similar way, most of the five life cycle
tentative management hypotheses about—which
stages of salmon provide for a unique set of wildlife
ecological functions may be in jeopardy if the wildlife that
species and their ecological functions (Table 5). For
performs such functions are not maintained. That is, one
example, wildlife associated with the incubation stage of
can now determine which wildlife species may be
salmon include secondary cavity users and primary
influenced by altering salmon populations and habitats
excavators of small ground burrows; these two ecological
that imperil specific salmon life stages, and the set of
functions are not provided, or only poorly provided, by
ecological functions associated with such wildlife species.
wildlife species associated with any of the other salmon
In some cases, other wildlife species not associated with
life cycle stages. Thus, to manage for the full set of
salmon may also perform some ecological function, but
ecological functions, one should focus on providing all
never in exactly the same manner and in the same set of
life cycle stages of salmon.
habitats and habitat elements.
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 649

Table 4. Importance of types of salmon-wildlife Table 5. Importance of salmon life stages to key
relations to key ecological functions. Listed are ecological functions. Listed are functions unique to
functions unique to each relationship category. each life stage category

1. Strong Consistent Relationships are important for: 1. Incubation Stage is important for:
trophic relations: organismal relations:
primary consumption: secondary cavity use
spermivory primary excavation of small ground burrows
grazing
2. Freshwater Rearing Stage is important for:
frugivory
(no specific function is mostly supported by this stage)
root feeding
organismal relations: 3. Saltwater Stage is important for:
controlling vertebrate populations organismal relations:
dispersing seeds, fruits, inverts, vasc plants creating aerial structures used by other spp.
creating feeding opportunities for other species creating aquatic structures used by other spp.
primary cavity excavation in trees and snags
primary creation of large ground burrows 4. Spawning Stage is important for:
primary creation and secondary use of ground trophic relations:
runways primary consumption:
wood relations: spermivory
fragmenting standing and down wood grazing
killing standing trees (creating snags) frugivory
root feeding
2. Occasional Relationships are important for: bark/cambium/bole feeding
trophic relations: organismal relations:
pirating of food controlling vertebrate populations
organismal relations: creating feeding opportunities for other species
secondary use of aerial and aquatic structures primary cavity excavation
created by other spp. primary excavation of large ground burrows
disease relations: primary creation of ground runways
carrier of domestic animal disease secondary use of ground runways
soil relations: wood relations:
improving soil structure and aeration by digging and fragmenting standing and down wood
burrowing killing standing trees (creating snags)
3. Indirect Relationships are important for: 5. Carcass Stage is important for:
trophic relations: trophic relations:
primary consumption: primary consumption:
fungivory fungivory
tertiary consumption organismal relations:
prey relations: controlling insect populations
providing prey for predators serving as interspecific host for avian nest parasite
organismal relations:
controlling insect populations
serving as interspecific host for avian nest parasite
primary creation of small ground burrows
 650 Wildlife–Habitat Relationships in Oregon and Washington

Salmon Fisheries enhance some fisheries, notably the rise of the coho salmon
Herein we provide only a very brief overview of fisheries runs in the 1960s, and chum salmon runs in Puget Sound.
as related to wildlife. A more thorough review of the In general, however, artificial propagation has failed to
magnitude and characteristics of Northwest Pacific Coast rebuild the runs to former levels,394 and in some instances
salmon fisheries and habitat issues are provided in a likely contributed to the further decline of wild stocks.209
number of recommended readings.103, 275, 365, 387, 390, 499 Admittedly, managing salmon fisheries is a challenge,
Washington and Oregon salmon fisheries include and past management approaches were generally
commercial, recreational, and treaty harvests that occur commodity/extraction-based; however, this approach has
in the states’ rivers, inland lakes, inland marine waters, significantly contributed to the decline of wild stocks.544
coastal embayments, and at sea. Fishing is an important Fishing activities that contribute to the problem include:
source of mortality, both for immature fish in the ocean indirect mortality due to catch and release of undersized
and mature fish on their return to freshwater to spawn. fish (bycatch), out-of-state domestic and foreign
Understanding past fishery effects is important because interception, conflicts among user groups, and the mixed-
man’s increasing efficiency as a predator augments the stock fisheries.
mortality rates that salmon encounter in natural situations. A major dilemma that fishery resource agencies find
Because of the presence of humans, salmon are themselves in is how to harvest hatchery salmon
significantly less available to a wide variety of natural selectively, while still meeting spawning escapement goals
consumers in both the freshwater, terrestrial, and marine of wild stocks of salmon. Hatchery-produced salmon co-
environments, and thus these ecosystems are suffering. mingle with wild salmon in ocean waters, and as a result,
According to the NRC,365 salmon habitat mortality factors a mixed-stock fishery is created. If harvests are allowed
caused by human activities and natural factors together in such mixed-stock fisheries, then wild and hatchery fish
usually exceed fishing mortality. Thus, although factors will be caught at rates that only hatchery fish can sustain.
other than fishing have a major effect on the production Wild fish cannot withstand the high hatchery exploitation
of adult fish, fishing is still the easiest salmon mortality rate because they are exposed to a full range of natural
factor to control.365 and human-caused selection pressures and mortalities.
The abundance of virtually all salmon species initially Hatchery fish are sheltered from mortality factors that
harvested in the Columbia River, Puget Sound, and coastal normally occur during incubation and freshwater rearing.
Washington and Oregon, have never recovered to their At smolt migration, many more progeny are still alive per
former levels, except for Puget Sound chum salmon hatchery female than per wild female. This condition
(Figures 6 and 7). Increased hatchery production did enables hatchery stocks to withstand higher rates of

Chinook Coho

Figure 6. Total commercial catch,

in millions of pounds, of Lower
Columbia River salmon from 1870s
to 1990s.390, 379, 387b, 540

Sockeye Chum

Steelhead Total
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 651

Chinook Coho

Figure 7. Estimated and

reported total catch, in
numbers, of Puget Sound
salmon from 1890s to
1990s.390, 57a, 538b

Chum Pink

Sockeye Total

harvest than wild stocks; however, even hatchery stocks According to the Pacific Fisheries Management Council
eventually succumb to the high exploitation rates through (PFMC)388 the state of Washington has established annual
changes to smaller adult size,517 or different time of escapement goals for coho, chum, and chinook salmon
return.190 Therefore, high fishing exploitation rates are and steelhead, and include wild and hatchery fish. Some
associated with fishery resource management agency escapement goals exist for pink and sockeye salmon that
policies, and agency policies also determine hatchery are mostly for wild fish. No determination has been made
policies and practices. The current demand by certain of the spawning escapement needs of sea-run cutthroat
interests to protect wild stocks can be in direct conflict trout. In Oregon, escapement goals have been established
with the agency mandate to enhance or supplement primarily for chinook and coho salmon, and include wild
current stocks with hatchery-produced fish to ensure and hatchery fish. No determination of the spawning
sustainable harvest for the major user groups. This conflict escapement needs has been made for wild steelhead, chum
will have an important bearing on future management of salmon, pink salmon, or sea-run cutthroat. Escapement
salmon fisheries and hatchery practices. Live capture goals may have been established for some of these stocks
selective fishing, including live release of wild unmarked in more recent years. Spawning escapement goals have
fish and retention of marked hatchery fish, is potentially been established for 98 wild salmon stocks and 15 wild
an option.279 stocks of steelhead in Washington.106, 390, 538a Analysis of
annual spawning escapement data by Konkel and
Spawning Escapement Goals McIntyre,257 collected for naturally spawning salmon
Spawning escapement goals (the number of spawners populations in the Pacific Northwest between 1969 and
required to perpetuate the population254) are set by fishery 1984, suggests that escapements are down for coho and
managers solely to determine the portion of the estimated chum; but up for chinook, sockeye, and pink salmon. The
returning adult population that can be harvested. No critical factor that salmon harvest managers need to face
spawning escapement goals currently provide an is how to reduce the annual salmon harvest, and achieve
identified portion of the escaping fish for wildlife or stock-by-stock ecosystem-based spawning escapement
ecological functions (Photograph 10).
 652 Wildlife–Habitat Relationships in Oregon and Washington

Photograph 10. Chum

salmon carcasses in
Kennedy Creek,
Washington. (Photo by
Jeff Cederholm)

goals. With the exception of carcass supplementation approaches do not account for impacts associated with
programs, there has not been a concerted effort to manage land use that drive down stock productivity (reduce
salmon populations for the benefits they provide to the survivals) and reduced habitat capacity, and decreased
recovery of listed wildlife species (e.g., grizzly bears) or genetic diversity, nor do they incorporate any
to the broader ecological systems. Salmon spawning consideration of temporal variability in environmental
escapement goals should not only replace a stock of conditions. Nonetheless, these approaches do represent a
salmon with sufficient numbers of recruits, but also meet shift from MSY to more ecologically based stock
the needs of the broader aquatic and terrestrial ecosystems management objectives.
that depend on salmon for nutrients and carbon influx.47,
48, 92, 319
Salmon harvest managers could take a lesson from Understanding Salmon Relationships
the worldwide conventions for herring harvest managers,
where ecosystem function has been explicitly recognized Salmon as a Key Linkage in Biodiversity And
through “...a precautionary, conservative approach to Productivity
fisheries management.” 541a Ecological processes have been so altered by human
There is increased interest in the Pacific Northwest in activities, especially in the more densely populated
developing harvest management strategies that address regions, that natural resource and environmental
nutrient delivery to freshwater ecosystems.266, 544 However, management will need to expand from current site- and
there is relatively little information on which to base case-specific methods, to landscape- and ecosystem-scale
escapement targets that will meet this objective. Ideally, approaches. The struggle to develop the tools required
estimates of the number of spawners needed to fulfill this for these scales of management has only just begun.269, 305,
function would be determined by experimentally altering 365, 460
The documents: From the Forest to the Sea,295 Forest
escapement levels for each stock and evaluating the impact Ecosystem Management: An Ecological, Economic, and Social
on system productivity. However, conducting such an Assessment,151 Pacific Salmon and Their Ecosystems,499 An
experiment on hundreds or thousands of stocks over a Ecosystem Approach to Salmonid Conservation,485 Fish Habitat
sufficient length of time is a daunting prospect and would Rehabilitation Procedures,477 and River Ecology and
not provide usable results for many years. Several other Management—Lessons From Pacific Coastal Ecoregion362 are
approaches to determining appropriate escapement levels recommended readings that describe the understanding
are currently being investigated. One option being of natural systems and processes and take a holistic
considered attempts to determine the amount of food approach to rehabilitation and restoration of watersheds
required to support a population of rearing fish that fully and ecosystems.
utilizes the habitat available in a stream. Escapement levels Anadromous salmon play an important role in
would be established which would ensure sufficient maintaining an ecosystem’s productivity. The seasonal
nutrients and organic matter are returned to the stream to migrations of millions of salmon between Pacific rim
produce this level of food. Another alternative is to streams and the subarctic Pacific Ocean appear to increase
develop a relationship between spawner density and the overall terrestrial productivity. Key processes discussed
proportion of marine-derived nutrients in the tissues of here are the transport of materials, energy and nutrients
juvenile fish. This type of relationship may enable a between marine, aquatic, and terrestrial ecosystems, with
“saturation level” for marine nutrients to be established emphasis on salmon as a transport vector. From a broad
and escapement goals set accordingly. However, these ecosystem management perspective, the status of salmon
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 653

metapopulations is a powerful indicator of human and nutrients carried in the biomass of the salmon runs is
adaptation to boreal biomes.348, 419, 464 Sibatani 464 has input to the trophic system through multiple pathways
suggested that salmon are the “canary in the mineshaft”; including direct consumption, excretion, decomposition,
the mineshaft in this instance being entire subarctic ocean and primary production. Direct consumption may occur
basin ecosystems. Cederholm et al.92 review and discuss in the form of predation, parasitism, or scavenging on the
the mechanisms of salmon nutrient transport and the live spawner, carcass, egg or fry cycle life stages. Carcass
significance to terrestrial and freshwater ecosystems. The decomposition and the particulate and dissolved organic
following discussion suggests a need to understand and matter released by spawning fish (e.g., eggs and milt,
apply information on the exchange of materials, energy, excrement) delivers nutrients to primary producers.
and nutrients between the aquatic and terrestrial Potential nutrient or energy pathways and factors
ecosystems of the northern Pacific Basin. Such an influencing biomass cycling of spawning salmon is
application would occur at multiple geographic and graphically depicted in Figure 8.
temporal scales, examining both healthy and depressed Freshwater ecosystem productivity depends upon
salmon populations under varying conditions. Effectively nutrient inputs and retention. Larkin and Slaney266 and
applied, managers would be able to define and achieve Munn et. al.339 discuss nutrient cycling and the nutrient
long-term ecosystem management success not just for spiraling concept, whereby nutrients unidirectionally
salmon, but for numerous other fish and wildlife resources spiraling downstream can influence aquatic system
and the overall health of the environment. functions for considerable distances. Larkin and Slaney266
and Munn et. al.339 also discuss the importance of instream
Biomass of Salmon Runs habitat complexity (wood debris complexes) for increasing
as an Energy Source productivity by increasing salmon carcass retention; citing
Organic matter that supports the trophic system of fresh Cederholm and Peterson90 and Cederholm et. al.91 The
water ecosystems is provided from both autochthonous discussion that follows emphasizes the necessity of
and allochthonous sources. Common types of retention mechanisms and how the physical and biological
autochthonous sources are: algae, mosses, vascular plants, complexity of the aquatic, riparian, and wetland zones
and phytoplankton. All of these factors are found in enhances this function.
freshwater, and generate organic matter through the Sportsmen and naturalists have long recognized the
process of photosynthesis. Common types of importance of salmon runs to the natural economy of
allochthonous input include leaves, needles, wood and streams, as this quotation by Haig-Brown180 reveals:
insects from the terrestrial environment and dissolved The death of a salmon is a strange and wonderful thing,
organic matter carried in groundwater that enters the a great gesture of abundance. Yet the dying salmon are
water body. Salmon provide an important source of not wasted. A whole natural economy is built on their
allochthonous organic matter for Pacific Northwest fresh bodies. Bald eagles wait in the trees, bears hunt in the
water ecosystems.47, 239, 248 Salmon spawning runs transport shallows and along the banks, mink and marten and
organic matter and nutrients from the northern Pacific coons come nightly to the feast. All through the winter
Ocean to their natal spawning grounds. The organic matter mallards and mergansers feed in the eddies, and in

Figure 8. Factors of
stream complexity and
marine-derived
nutrient pathways that
influence biological
productivity.92
 654 Wildlife–Habitat Relationships in Oregon and Washington

freshet time, the herring gulls come in to plunge down phosphorus total loads to the Elwha River watershed
on the swifter water and pick up the rotting drift. under existing conditions.226 These preliminary estimates
Caddis larvae and other carnivorous insects crawl over as to the levels of nutrient loadings to the water column
the carcasses that are caught in the bottoms of the pools and sediments, while crude, should be sufficient cause to
or against the rocks in the eddies. The stream builds its trigger some rethinking of what constitutes healthy
fertility on this death and readies itself to support a new baseline water and sediment quality for Puget Sound and
generation of salmon. Pacific Coast streams and estuaries.
The scientific community has also recognized the Even low spawning densities can provide significant
contribution of nutrients and organic matter from contributions of nutrients to the system. Juvenile coho,
spawning salmon for some time. Juday et al.241 estimated steelhead and cutthroat in a small stream in western
that sockeye salmon transported in excess of 2 million kg Washington obtained from 25% to 40% of their N and C
of organic matter and 5000 kg of phosphorus to the Karluk from dead coho salmon that spawned in the stream.47
River system in Alaska in an average year. This recognition Aquatic insects also contained high levels of marine-
resulted in sockeye lake fertilization programs in British derived N and C, and the foliage of plants growing along
Columbia and Alaska, to replenish lost nutrients caused the streams also contained nitrogen of marine origin.47 The
by fish harvest.496, 497 Over the last 10 years stable isotope direct feeding of salmon fry on salmon carcasses has been
analysis has enabled direct measurement of marine- known for a long time in the Amur River of Asia,377
derived nutrients in stream47, 239, 248 and lake249 ecosystems. however, the growth benefits for juvenile salmon has only
These studies have firmly established the need to consider recently been documented in North America.47
the importance of salmon biomass as a flow of energy and Macroinvertebrate communities in streams receiving
nutrients into the freshwater and estuarine food webs of salmon runs can change in response to spawning activity
the Pacific Northwest. and nutrient enrichment. In a Snoqualmie River tributary
Nutrient levels in freshwater and estuarine systems can and in Kennedy Creek, Washington, Minakawa324 found
be substantially enriched by the organic inputs of salmon the presence of salmon carcasses and eggs produced a two-
spawning runs. The majority of material transported to fold or greater increase in total insect densities and biomass
freshwater by some species of anadromous salmon is of compared to control reaches. Piorkowski404 found insect
marine origin. Mathisen et al.298 demonstrated that over taxa richness and diversity to increase in response to
95% of the body mass of some salmon species is produced nutrient enrichment from salmon carcasses in southeast
during ocean residence; the remainder represents mass Alaska, and suggested that insect colonization of carcasses
accumulated in freshwater prior to migration to the sea. facilitated decomposition and subsequent nutrient release.
The species of salmon that spawn at high densities (e.g., Bilby et al.47 found all functional feeding groups except
chum, pink, sockeye) significantly alter nutrient loadings insect shredders to be enriched with marine origin isotopes
and budgets in the freshwater systems where they spawn. of nitrogen and carbon in western Washington streams
In Kamchatka, Krokhin262 reported that 35-40% of the after coho salmon spawning. Some aquatic invertebrates
yearly total phosphorus input to a lake was transported such as stoneflies (Alloperla) 375 and Dipteran flies
by spawning sockeye salmon, as was much of the nitrogen (Chironomidae)131 will scavenge for dead salmon eggs and
input to the system. Similar results have been reported alevins within the gravel. Limnephilid caddisfly larvae
for the Iliamna Lake system in Alaska,122, 249, 297, 298 and for are attracted to recently expired salmon and have been
the Paratunka River basin, Kamchatka.261, 262 Nutrients observed feeding directly on fish flesh.324, 404
from spawning pink and chum salmon have been shown Terrestrial insects including fly maggots (Diptera) have
to enrich not only the freshwater and estuarine habitats also been observed feeding heavily on salmon carcasses
where they spawn, but also the estuarine habitats in streams in the Queen Charlotte Islands of British
downstream.62, 503 Munn et. al.339 consider changes in Columbia,422 but generally little work has been done to
nutrient loading and cycling and ecosystem productivity systematically document these activities. Maggot larvae
that could result from restoration of historic salmonid have commonly been observed consuming beached
populations to the Elwha River system in Washington salmon carcasses during the warmer months of the
state. The study indicates a potential 65-fold increase in spawning season along the spawning reaches of several
nitrogen and phosphorus loadings from salmon returns. Washington streams. Dead chum salmon along Kennedy
They concluded that restoration of the Elwha River system Creek in South Puget Sound often have their heads filled
salmon runs would have a profound effect on the primary with maggots (Cederholm, pers. obs.) (Photograph 11).
and secondary production in the system. Annual nutrient Chinook carcasses along Puget Sound Basin rivers can be
inputs of 29.8 mt of nitrogen and 3.5 mt of phosphorus reduced to skeletal remains by maggots within a two week
are projected for a 980 mt biomass of restored salmon runs period; fall freshets frequently have been observed to wash
based on Munn et al.339 Average sizes of salmon by species the carcasses and masses of larvae back into the stream
are from Ricker,426 as quoted in Larkin and Slaney.266 where they are then available as food for juvenile salmon
Proportions of body composition for nitrogen and and other organisms (W.Graeber, pers. obs.). Hornets have
phosphorus are from Larkin and Slaney.266 These nutrient also been observed to feed on carcass remains during
input levels would be approximately 40% and 13%, warm fall weather periods in the same areas; they are
respectively, of the 74.5 mt of nitrogen and 27.3 mt of especially attracted to exposed fresh flesh or blood
(W.Graeber, pers. obs.).
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 655

Photograph 11. Fly

maggots eating a
chum salmon carcass
at Kennedy Creek,
Washington. (Photo
by Jeff Cederholm)

Quantitative measurements of salmon carcass Pacific Salmon Provide

consumption in the terrestrial environment has focused Key Ecological Functions
on their utilization by high profile species like bald eagles The key ecological functions that spawning salmon play
along the Skagit River, Washington; 490 and grizzly bears within certain freshwater ecosystems may be illustrated
along the Columbia River.205 Cederholm et al.91 recorded with a well-documented case study from McDonald Creek
43 taxa of mammals and birds present on small Olympic in Glacier National Park, Montana. This stream is a
Peninsula streams at a time when coho salmon carcasses principal spawning tributary for the Flathead Lake—
were present, and found that 51% of those taxa had fed Flathead River ecosystem. The triggering event in the
on carcasses. Skagen et al.475 in their study of human series of changes that cascaded through this ecosystem
disturbance on an avian scavenging guild, observed was caused by the introduction of an exotic species, the
significant bird scavenging of chum and coho salmon opossum shrimp (Mysis relicta), from 1968 and 1975.486 The
carcasses along the North Fork of the Nooksack River, shrimp were added to the lake as a food source for kokanee
Washington. The primary bird scavengers were eagles, salmon, but behavioral patterns made them unavailable
crows, and glaucous-winged gulls. Information from the for consumption. Opossum shrimp are voracious
expert panel process and the published literature on predators of zooplankton, the principal food of the
observations of wildlife predation and scavenging on kokanee. The shrimp decimated the zooplankton in the
salmon is presented in Appendix VI. lake, and by the late 1980s the lake and McDonald Creek
Cederholm et al.91 also reported that black bears, spawning kokanee population had collapsed. These fish
raccoons, and river otters increase food availability for served as an important food source for various birds and
terrestrial species incapable of removing carcasses from mammals that had fed upon them in the spawning
the stream. The larger animals rarely completely tributaries. Among the most prominent predators and
consumed the carcasses they removed from the stream, scavengers utilizing this resource were bald eagles that
and were often followed by an array of other smaller birds gathered by the hundreds during the kokanee spawning
and animals who fed on the “leavings.” A similar period. In 1981, spawning kokanee in excess of 100,000
interaction occurred at McDonald Creek, Glacier National attracted 639 eagles, the densest eagle concentration south
Park, Montana, where kokanee salmon captured by of Canada. Beginning in 1987 eagle numbers declined
grizzly bears were incompletely consumed, leaving along with the kokanee, reaching a low of just 25 birds in
remains for birds and small mammals.486 1989. It is feared that loss of the kokanee spawning run
As the above studies indicate, spawning salmon could lead to higher eagle mortality during migration or
provide a source of carbon, nitrogen and phosphorus during winter, unless the birds can find alternate food
essential to maintaining the production of salmon resources, a prospect that is not likely in that ecosystem.486
juveniles and other trophic levels of the stream. A number of other bird and mammal species that used
Accumulating evidence suggests that spawning salmon the McDonald Creek kokanee also have been displaced.486
populations are an important link to the adjacent riparian Gulls, mergansers and mallards commonly fed on kokanee
and terrestrial communities, and indeed, fortifies the role carcasses, while Barrows and common goldeneyes and
of salmon as a keystone species, wherein the integrity and dippers fed on loose eggs. Mammals that fed on spawning
persistence of the entire community is contingent upon kokanee or carcasses along McDonald Creek, including
the population’s actions and abundance.559 grizzly bears, coyotes, mink, and river otters, are now less
common along the creek.
 656 Wildlife–Habitat Relationships in Oregon and Washington

Estuaries, where rivers and streams meet tidal influence birds, fish) that carry nutrients across ecosystem
and enter the ocean, act as traps for many of the nutrients boundaries.61, 349, 379, 463, 464 Therefore the role and importance
washed from watersheds. Some species of Pacific salmon of salmon in the freshwater and terrestrial ecosystems can
typically spawn near saltwater, even beginning within the be recognized within the context of broader nutrient
upper reaches of estuaries. Spawning very often occurs at cycling, and spawning migrations of salmon represent an
high densities. The effect of salmon carcasses on the obvious example of this process. Other means of moving
nutrient dynamics and trophic productivity of estuarine nutrients upstream, such as the emergence of the adult
systems is just beginning to be examined, Kline et al.248 stages of insects and meteorological vectors, are
reported that approximately 30,000 pink salmon spawned considered to be relatively insignificant compared to
within 1.2 km of the estuary of Sashin Creek in southeast anadromous fish.278 The nutrient-subsidy contributed by
Alaska. In southwestern Washington, the 5 km of Kennedy salmon in the North Pacific could potentially serve as a
Creek accessible to anadromous fishes has supported as model for a more general, and global, aspect of nutrient
many as 80,000 spawning chum salmon (WDFW unpub. circulation. Such studies may be supplemented by an
data). Using the size and body composition information assessment of the extent to which migratory birds, which
previously cited (under the Elwha River discussion), we travel long distances between boreal/subboreal zones and
estimate that this peak escapement to Kennedy Creek temperate and/or tropical regions, contribute to similar
delivered approximately 398 mt of salmon flesh containing effects.
12 mt of nitrogen and 1.4 mt of phosphorus to 0.075 km2 Using the discipline known as Resource Physics,
of stream channel area (5 km with an average channel Tsuchida524 has explained how the hydrologic cycle and
width of 15 m). The nutrient loading per unit of channel the convection of air are able to keep the earth in a low-
area would be 160 mt/km2 nitrogen and 18.7 mt/km2 entropy state.350 The hydrologic cycle is in turn driven by
phosphorus. Many of these salmon become carcasses solar thermal energy and the earth’s gravity. Subsequently,
within the stream system or their dissolved nutrients may inorganic salts (especially nitrates and phosphates), which
be carried to the estuary during freshets. Therefore, a are essential to formation and activities of animals, plants,
nutrient link may function between adult salmon carcasses and microorganisms, are eventually washed
and juvenile salmon rearing in the estuary. For example, downstream.525 Ultimately, these salts are dissolved in
Fujiwara and Highsmith159 found elevated stable isotope river water and transported to the ocean, where they attain
ratios of nitrogen in Ulva sp., an estuarine macroalga, the highest and most uniform concentration below the
following the decomposition of salmon carcasses in depth of 1,000 m, largely free from biological consumption
Seldovia Bay, Alaska. Ulva spp. are a major food source in the absence of photosynthesis. However, due in part to
for harpacticoid copepods, which in turn are a preferred ocean currents (local upwelling), these nutrients
prey of juvenile chum salmon fry in the estuary. Thus the eventually find their way back to the surface waters. This
contribution of nutrients and organic matter from salmon occurs in northern temperate or subpolar oceans by the
carcasses may be a substantial source in some systems, effective vertical mixing of seawater due to the
and may be a key factor in promoting estuarine approximation of water temperature between deep and
productivity. The importance of estuaries as nursery zones shallow water, primarily during colder seasons. Finally,
for anadromous salmon along the Pacific Northwest coast uptake of these salts by marine plants near the surface,
is well documented;194, 196, 289, 355, 397, 454 the role carcasses play where photosynthesis is possible during the warmer
in maintaining productivity of these systems may be seasons, allows a means through which other animals are
critical in supporting the health of salmon populations.159 able to derive and transport the nutrients inland. For
The role salmon populations play as a key vector in example, Tsuchida525 speculates that in coastal regions,
the recycling of energy and nutrients inland from the some birds will carry nutrients back to the land after
North Pacific Ocean to aquatic and the terrestrial deriving them from the consumption of marine organisms.
ecosystems is now gaining recognition as a critical Bird excrement is a fertilizer rich in inorganic matter,
component of ecosystem function.348, 350, 463, 464 River and lake especially phosphates, as evidenced by the material
fertilization with inorganic nutrients has been undertaken deposits (Peruvian bird guano) on tropical sea islands.
with ecosystem restoration in mind in some British Sibatani463, 464 points out that another, arguably more
Columbia systems; 15, 16, 476, 496 however, artificially significant way that nutrients are transported back onto
supplementing inorganic nutrients may not fully mitigate the land is by anadromous fish swimming up, spawning,
for the loss of the multiple pathway flow of energy and and dying in the many rivers of the Asian and North
materials provided by naturally spawning salmon. American continents. Murota et al.351 discussed how
migratory fish move ocean nutrients inland and benefit
Salmon as Vectors in the Siberian forest and its various wildlife inhabitants
Broader Nutrient Cycling (Figure 9).
The flux of nutrients is essential for the continuity and In the Edo era, some people in Japan started to notice
stability of any living system,412 and nutrients provide a that forests along seashores or rivers attracted fish
link between aquatic and terrestrial ecosystems.61, 317 towards them. It was considered that a forest could give
Biological vectors through which materials and energy are benefits to fish in the forms of shadow as shelter,
transported include migration of animals (i.e., mammals, nutrients, and so on. This consideration remained in
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 657

Figure 9. Anadromous fish and the

Siberian forest.351

the minds of people living near waterfronts or forests With an Ecosystem

after the Meiji Restoration (1868). When the first forest
act was introduced at the beginning of the twentieth
Perspective in Mind—
century, it contained the article ordering the Where Do We Go from Here?
conservation of uo-tsuki-rin, which literally meant The need for an ecosystem approach to salmon
“fish-attaching forest.’”This article is still valid in the management has inevitably grown in Washington and
present-day forest act of Japan.348 Oregon.305, 365, 499 Terrestrial ecologists have recognized the
The theories of Sibatani, 463, 464 Murota, 348, 349 and influence human uses (and thereby disturbances) have had
Tsuchida524, 525 cause one to reflect on the once bountiful on terrestrial and aquatic ecosystems of large scales.151, 295,
545
salmon runs of the Columbia River. Before Europeans Some investigators have explored the significance of
settled the Pacific Northwest, salmon and steelhead had the salmon as a key ecological process vector on the
access to over 20,000 km of main river and tributaries in broadest scales of energy and nutrient transport.348, 464, 559
the Columbia River basin.379 The annual Columbia River The use of salmon as an indicator of ecosystem health, or
salmon and steelhead run size was estimated to range “the canary” is complicated by the fact that this canary is
between 8.2 and 16.3 million fish.379 Using an average a food resource in high demand by humans.464
weight of 6.75 kg per salmon, spawning populations could The magnitude of the role of salmon populations as
have potentially contributed between 55,350 and 110,025 keystone vectors in energy and nutrient cycling inland
mt of nutrient annually. This amounts to potential average from the Northern Pacific to freshwater and terrestrial
annual contributions of salmon nutrient in the ecosystems is now gaining recognition as a critical
anadromous area of the Columbia River on the order of component to an overall understanding of ecosystem
2.76 to 5.50 mt/km. But what of the native people of the functions.92, 348, 349, 464 Application of this knowledge to
Columbia River? Many traveled long distances to partake understand the cumulative impacts of human land use
in the catch and consumption of salmon, and in doing so practices and fisheries exploitation on ecosystem functions
participated in the further cycling of nutrients over this is only just beginning. New tools are necessary to make
vast watershed, and beyond. Some tribes of the upper management actions toward regaining lost productivity
Columbia were known to cross the Continental Divide to and biodiversity objectives.305, 365, 499
trade dried salmon for buffalo hides,379 thus providing an Greater understanding of the hydrologic cycle has
additional mechanism for transfer of marine-derived helped us to improve land and water uses to better adapt
nutrients to the inland land mass. Salmon, wildlife, and to our environment. Better understanding of geologic
humans, therefore, may be the most prominent carriers processes including the role of hydrogeology in shaping
of ocean nutrients to inland ecosystems. the landscape and controlling the rates at which sediments,
and small and large organic debris cycle through
 658 Wildlife–Habitat Relationships in Oregon and Washington

watersheds is also leading to changes in views on land 3. How have humans altered the nutrient budget?
and water uses.295, 434 Understanding nutrient cycling 4. What adaptive management actions might be
processes, pathways and the effects of nutrient loading warranted and feasible to push the terrestrial and
has helped in managing water and sediment quality aquatic systems toward the identified goals?
problems. 245, 435, 538 In much the same manner, an 5. Are there some measures to employ in the interim
understanding of nutrient spiraling and cycling in until stocks of salmon can be restored?
streams369 and the keystone role of anadromous salmon266, 6. What are the desired future conditions?
339
will be valuable, if not essential, to understand how we Early European settlement of the eastern North Pacific
have affected coastal ecosystems and the processes that Rim territories provides many accounts of heavy
support them. Such an understanding will lead us to better extractions of forest resources.70, 110a, 276, 295, 348, 365, 464, 554
identification of those management action options we may Logging of the forests had the obvious effect of short-
take to achieve desired future conditions. circuiting the prior cycles that supplied woody debris to
Maser et al.295 have added substantially to the literature freshwater, estuarine, and marine ecosystems. In recent
on downstream and seaward movement of materials, decades, mechanized logging equipment combined with
energy and nutrient transport processes to and through highly efficient (“clean”) logging practices and slash
aquatic systems. Their report compiled and presented a burning to prevent wildfires and accelerate re-growth of
wealth of information on the inputs, fates, and effects of planted conifers has further resulted in very little debris
forest debris, particularly LWD, in freshwater, estuarine, left on the site or entering streams. The store of vast
and marine ecosystems. In essence, physical and chemical quantities of nutrients in the form of the decayed woody
processes were well described, now the biological is debris and trapped detritus that serves as substrate for
considered. Adding estimates of the upstream flow of long-term nitrogen fixation and retention no longer exists.
energy and nutrients via salmon to existing watershed Large woody debris may continue to enter stream
processes literature, will provide a more complete picture corridors, but not necessarily in the amount, size, and
of the large scale and long-term energy and nutrient cycles quality that it did in the past, thereby decreasing potential
for entire watersheds. An understanding of the overall to provide stream structure and organic matter to food-
cycles and levels of productivity at this scale will provide webs.299, 416
the context for interpretation of local level trends in The long-term loss of the function of LWD as a primary
production and materials transport, utilization and component of the floodplain waterways, resulting from
storage. It has been well established that aquatic systems land and channel clearing, may well be more significant
have metabolisms that function based upon physical than the loss of the wood material itself. The resistance of
processes, rates of loadings of materials, energy, and the abundant woody material slowed the flows of water,
nutrients, rates of primary and secondary production, and sediment, and smaller debris; resulting in very complex
resulting changes in standing stocks of fishes. valley floor stream-ways composed of multiple highly
As a keystone species to the productivity and sinuous channels that were generally well connected to
biodiversity of the ecosystems of the North Pacific basin, off-channel wetland systems by sloughs and high water
anadromous salmon closely link the management issues channels. Put this liquidation of natural resource capital
of the forests, the flood plains and lowlands, the estuaries into the context of long-term climate cycles189 and Maser
and nearshore areas, and the ocean domains as a et al.’s295 long-term geologic and successional cycles, and
continuum. Materials, energy and nutrient budget one can begin to formulate management goals for
analyses on an appropriate time scale will be necessary to ecological processes.
estimate the potential effects that past land use practices A combination of development activities have diverted
have had on production and discern which ones persist. water from, shortened, straightened, cleared, dammed,
Budgets for the North Pacific Basin can theoretically be diked, drained, filled and polluted the habitats of salmon.
calculated in a similar manner to that already used on Early in settlement, logging and splash damming, land
smaller systems. Then we can begin to use resulting clearing for agriculture, and channel clearing for
information to provide a context for management navigation appear to have had the most pronounced
decisions in various disciplines and forums that will affect effects. The development and consumptive utilization of
materials, energy, and nutrient flows and stocks at various natural resources was highly dependent upon water-borne
scales. To address the disconnects apparent in the current transportation and patterns of impacts are reflective of
state of terrestrial and aquatic systems will require some navigation-centered commerce. Continuing development
estimate of many factors, including: for agriculture, industry, and urban growth has resulted
1. What is the status of the nutrient capital and rates of in further losses through conversions to other uses.
transport within the domains and the basin as a Impacts have become more pervasive throughout the
whole (nutrient budget)? landscape as transportation infrastructure and vehicle
2. What is the range of the nutrient and materials capital capabilities have increased. Releases of persistent toxins
and rates of transport (how does the current budget has contaminated coastal sediments. The available
relate to the known ranges of standing stocks and freshwater and estuarine salmon habitats also continue
rates of metabolism and transport)? to be degraded by ongoing land uses.174, 245, 295, 365, 454, 526, 554
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 659

Desired Future Condition Acknowledgments

The inability of the various interest groups to resolve We thank the Washington Departments of Fish and
conflicting and agreed-to goals or conditions has been Wildlife (WDFW) and Natural Resources (WDNR) for
identified as the fatal gap in salmon management.54, 365 supporting this project. We appreciate greatly the
Describing the desired future condition of the terrestrial thoughtful reviews given by Peter Bisson of the United
and aquatic ecosystems and identifying and defining States Forest Service; Jim Lichatowich, Private Consultant;
human actions can influence movement of ecosystems Hal Michael, Jr., of the WDFW; Tom Mumford of WDNR;
toward those conditions. The institutional changes Pat Slaney of the British Columbia Watershed Restoration
suggested by Lichatowich 277 will be necessary to Program; and Mary F. Willson of the Great Lakes Program
implement the long-term management approach required. of The Nature Conservancy. We also thank Luis Prado of
Some of those institutions are known and have been in WDNR and Darrell Pruett of the WDFW for graphic art
use during other eras where human land and resource work on some of the figures in this report, and Billie
uses appeared to have been indefinitely sustainable.464, 554 Wyckoff of WDNR for her help.
The essence of the Pacific salmon is that it links together
what humans generally consider distant, diverse, and Literature Cited
separate ecosystems and long time spans. 1. Adams, C. 1998. Determining the effects of marine derived nutrients
In spite of the high potential commodity value of the on growth of juvenile coho salmon (Oncorhynchus kisutch) using stable
isotopes. The 1998 Annual General Meeting of the North Pacific
harvest and our knowledge that the Pacific salmon is a
International Chapter of American Fisheries Society Meeting, held
key driver for the biodiversity and productivity of the March 18-20, 1998 at Alderbrook, WA.
northern Pacific basin, we have not developed the 2. Agee, J. K. 1988. Successional dynamics of forest riparian zones. Pages
strategies for effective long-term management of the 31-43 in: K. J. Raedeke, editor. Streamside Management: Riparian Wildlife
resource’s health. The widespread use of salmon and Forestry Interactions. Contrib. No. 59, Institute of Forest
hatcheries has also significantly reduced the amount of Resources, University of Washington, Seattle, WA.
salmon carcass nutrients available for the aquatic food 3. Ainley, D. G. C. S. Strong T. M. Penniman, and R. J. Boekelheide. 1990. The
feeding ecology of Farallon seabirds. Pages 51-127 in: D. A. Ainley and R.
webs. Modern human culture has not fully adapted to the J. Boekelheide, editors. Seabirds of the Farallon Islands, Ecology,
environment of the northern Pacific Basin. The Dynamics, and Structure of an Upwelling-System Community. Stanford
environment of our region is showing the signs of stress University Press, Stanford, CA.
all around us which indicates the failure of our past and 4. ———, and D. W. Anderson. 1981. Feeding ecology of marine
current approaches. If humans are to thrive at present cormorants in southwestern North America. Condor. 83:120-131.
population levels within portions of the basin, we will 4a. ———, and G.A. Sanger. 1979. Trophic relations of seabirds in the
northeastern Pacific Ocean and Bering Sea. Pages 95-122 in J.C.
need to look more at multiple scales of ecosystem
Bartonek and D.N. Nettleship, editors. Conservation of marine birds of
management and at integrating that management to northern North America. U.S. Department of Interior, Fish and Wildlife
sustain productivity over very long time frames. Service, Wildlife Research Report No. 11.
The challenge, then, for the people of Washington and 5. Akande, M. 1972.The foods of feral mink (Mustela vison) in Scotland.
Oregon and the whole North Pacific Rim is to recognize Journal of Zoology 167:475-479.
that the character and the health of the northern Pacific 6. Alcorn, G.D. 1958. Nesting of the Caspian Tern in Gray’s Harbor,
Basin sub-ecosystems depend upon the material and Washington. Murrelet 39:19-20.
7. Alexander, G. R. 1979. Predators of fish in cold water streams. Pages
energy flow processes that link them. The flows of energy
153-170 in: Predator-Prey Systems in Fisheries Management. Sport
and of organic and inorganic materials among the various Fishing Institute, Washington, D.C.
aquatic and upland ecosystems determines the 8. Alexander, G. 1977. Food of vertebrate predators on trout waters in
productivity of each component and of the whole. In short, north-central lower Michigan. Michigan Academician 10:181-195.
the whole is greater than the sum of the parts, symbiosis 9. Allee, B. A. 1982. The role of interspecific competition in the
on a grand scale. Understanding the biological processes distribution of salmonids in streams. Pages 111-122 in: E. L. Brannon
and the impacts of our management practices upon them and E. O. Salo, editors. Proceedings of the Salmon and Trout Migratory
Behavior Symposium. 1981, First International Symposium. School of
will be necessary for our long-term (measured in Fisheries, University of Washington, Seattle, WA.
generations) success in the region. To do that we will need 10. Allen, K. R. 1951. The Horokiwi Stream: a study of a trout population.
to look beyond the plants, animals, and the habitats of a New Zealand Dept. Fish. Bull. 10:1-238.
given smaller-scale ecosystem to the processes that link 11. Anderson, N. H. 1992. Influence of disturbance on insect communities
them together, perhaps into the large-scale ecosystem of of Pacific Northwest streams. Hydrobiologia 248:79-92.
the anadromous salmon. The flow of sediments, woody 12. Anderson, M. E. 1977. Aspects of the ecology of two sympatric species
of Thamnophis and heavy metal accumulation within the species. M.S.
debris, detritus, and nutrients through a watershed
Thesis, University of Montana, Missoula, MT.
determines the character and productivity of the entire 13. Antonelis, G. A. Jr. and M. A. Perez. 1984. Estimated annual food
watershed, estuary, the near shore zone and even the consumption by northern fur seals in the California current. CalCOFI
domains of the northern Pacific Ocean. The flow of energy Rep. 25:135-145.
and nutrients back upstream via the Pacific salmon and 14. Antonelli, A. L., R. A. Nussbaum, and S. D. Smith. 1972. Comparative
the ability of the watershed to retain them, in large food habits of four species of stream-dwelling vertebrates
measure determines the productivity of the entire (Dicamptodon ensatus, D. copei, Cottus tenuis, Salmo gairdneri). Northwest
Science 46:277-289.
watershed.
15. Ashley, K., L. C. Thompson, D. C. Lasenby, L. McEachern, K. E.
Smokorowski, and D. Sebastian. 1997. Restoration of an interior lake
ecosystem: the Kootenay Lake fertilization experiment. Vol. 32: 295-323.
 660 Wildlife–Habitat Relationships in Oregon and Washington

16. ———, and P. A. Slaney. 1997. Accelerating recovery of stream, river 38. Benda, L., T. J. Beechie, R. C. Wissmar, and A. Johnson. 1992.
and pond productivity by low-level nutrient replacement. Pages 13-1 to Morphology and evolution of salmonid habitats in a recently
13-24 in: P. A. Slaney and D. Zaldokas, editors. Fish Habitat deglaciated river basin, Washington State, USA. Canadian Journal of
Rehabilitation Procedures. Watershed Restoration Program, Ministry of Fisheries and Aquatic Sciences 49:1246-1256.
Environment, Lands and Parks, Vancouver, BC, Canada. 39. Bent, A. C. 1921. Life histories of North American Gulls and Terns.
17. Ayles, G. B., J. G. I. Lark, J. Barica, and H. Kling. 1976. Seasonal mortality United States National Museum. Bulletin No.1131:65-71.
of Rainbow trout (Salmo gairdneri) planted in small eutrophic lakes of 39a. ———. 1926. Life histories of North American marsh birds. U.S.
central Canada. Journal of the Fisheries Research Board of Canada National Museum Bulletin No. 135.
33:647-655. 40. Berman, C., and T. P. Quinn. 1991. Behavioural thermoregulation and
18. Baird, P. H. 1990. Influence of abiotic factors and prey distribution on homing by spring chinook salmon, Oncorhynchus tshawytscha Walbaum,
diet and reproductive success of three seabird species in Alaska (USA). in the Yakima River. Journal of Fish. Biology 39:301-312-
Ornis Scandinavica . 21:224-235. 41. Beschta, R. L., R. E. Bilby, G. W. Brown, L. B. Holtby, and T. D. Hofstra.
19. Baird, R. 1991a. W. The Risso’s dolphin in British Columbia. Victoria 1987. Stream temperature and aquatic habitat: fisheries and forestry
Naturalist (Victoria, BC, Canada) 47:6-7. interactions. Pages 191-232 in: E. O. Salo and T. W. Cundy, editors.
20. ———. 1991b. Optimal foraging and intraspecific competition in the Streamside Management: Forestry and Fishery Interactions. Institute of
tufted puffin. Condor 93:503-515. Forest Resources Contribution Number 57. University of Washington,
21. Baker, R. C., F. Wilke, and C. H. Baltzo. 1970 The Northern Fur Seal. Seattle, WA.
United States Dept. of the Interior. U. S. Fish and Wildlife Service. 42. Bilby, R. E. 1988. Interactions between terrestrial and aquatic systems.
Circular No. 336. Washington D.C. Pages 13-29 in: K. J. Raedeke, editor. Streamside Management: Riparian
22. Balcomb, K. C., J. R. Boran, and S. L. Heimlich. 1982. Killer whales in Wildlife and Forestry Interactions. Contrib. No. 59, Inst. For. Res.,
Greater Puget Sound. Pages 681-685 in: International Whaling University of Washington, Seattle, WA.
Commission, Report of the Commission 32. 43. ———. 1984. Post-logging removal of woody debris affects stream
23. ———, J. R. Boran, J. R. Osborne, and N. J. Haenel. 1980. Observations channel stability. Journal of Forestry 82: 609-613.
of killer whales (Orcinus orca) in Greater Puget Sound, State of 44. ———, and J. W. Ward. 1989. Changes in characteristics and function
Washington. INTIS PB80-224728, U. S. Dept. of Comm., Springfield, VA. of woody debris with increasing size of streams in western Washington.
24. Baltz, D. M., and G. V. Morejohn. 1977. Food habits and niche overlap Transactions of American Fisheries Society 118:368-378.
of seabirds wintering on Monterey Bay, California. Auk 94:526-543. 45. ———, and ———. 1991. Characteristics and function of large
25. Banci,V. 1987. Ecology and behavior of wolverine in Yukon. M.S. Thesis. woody debris in streams draining old-growth, clear-cut, and second-
Simon Fraser University. 178 pp. growth forests in southwestern Washington. Canadian Journal of
26. Banfield, A. W. F. 1974. The mammals of Canada. University of Toronto Fisheries and Aquatic Sciences 48:2499-2508.
Press, Toronto, Canada. 438 pp. 46. ———, and P. A. Bisson. 1992. Allochthonous versus autochthonous
26a. Barr, J.F. 1973. Feeding biology of the common loon (Gavia immer) in organic matter contributions to the trophic support of fish populations
oligotrophic lakes of the Canadian shield. Ph.D. dissertation, University in clear-cut and old-growth forested streams. Canadian Journal of
of Guelph, Ontario, 192pp. Fisheries and Aquatic Sciences 49:540-551.
27. Bayer, R. D. 1986. Seabirds near an Oregon estuarine salmon hatchery 47. ———, B.R. Fransen and P.A. Bisson. 1996. Incorporation of nitrogen
in 1982 and during the 1983 El Niño. Fish Bulletin 84:279-286. and carbon from spawning coho salmon into the trophic system of
28. ———. 1986a. Nearshore flights of seabirds past Yaquina Estuary, small streams: evidence from stable isotopes. Canadian Journal of
Oregon, during the 1982 and 1983 summers. Western Birds 16:169- Fisheries and Aquatic Sciences 53:164-173.
173. 48. ———, ———, ———, and J. W. Walter. 1998. Response of juvenile
29. ———. 1986b. Breeding success of seabirds along the mid-Oregon coho salmon (Oncorhynchus kisutch) and steelhead (Oncorhynchus
coast concurrent with the 1983 El Niño. Murrelet 67:23-26. mykiss) to the addition of salmon carcasses to two streams in
29a. ———. 1989. The cormorant/fisherman conflict in Tillamook Southwestern Washington, U.S.A. Canadian Journal of Fisheries and
County, Oregon. Studies in Oregon Ornithology, No. 6. Gahmken Press, Aquatic Sciences 55:1909-1918.
Newport, OR. 101 pp. 49. ———, and P. A. Bisson. 1998. Function and distribution of large
30. Beach, U. S. 1937. The destruction of trout by fish ducks. Transactions woody debris, pages 324-346 In: R. J. Naiman and R. E. Bilby, editors.
of the American Fisheries Society 66:338-342. River Ecology and Management: Lessons From The Pacific Coastal
31. Beach, R., A. C. Geiger, S. J. Jeffries, S. D. Treacy and B. L. Troutman. Ecoregion. Springer-Verlag. New York City, NY.
1985. Marine mammals and their interactions with fisheries of the 50. ———, R.E. Bilby, M.D. Bryant, C.A. Dolloff, G.B. Grette, R.A. House,
Columbia River and adjacent waters, 1980-1982. Third Annual Report. M.L. Murphy, K V. Koski, and J.R. Sedell. 1987. Large woody debris in
Washington Department of Wildlife, Wildlife Management Division. forested streams in the Pacific Northwest: past, present, and future,
Olympia, WA. pages 143-190 in: E.O. Salo and T.W. Cundy, editors. Streamside
32. Beamish, R. J., and R. Boullion. 1993. Pacific salmon production trends Management: Forestry and Fishery Interactions. Contribution Number
in relation to climate. Canadian Journal of Fisheries and Aquatic 57, Institute of Forest Resources, University of Washington, Seattle,
Sciences 50:1002-1016. WA.
33. Beechie, T. J., and T. H. Sibley. 1990. Evaluation of the TFW stream 51. ——— and G. E. Davis. 1976. Production of juvenile chinook salmon,
classification system: stratification of physical habitat area and Oncorhynchus tshawytscha, in a heated model stream. NOAA Fisheries
distribution. Timber/Fish/Wildlife Final Report TFW-16B-89-006, Bulletin 74:763-774.
Washington Department of Natural Resources, Olympia, WA. 52. ———, and J. R. Sedell. 1984. Salmonid populations in streams in
34. Bell, D. T., and S. K. Sipp. 1975. The litter stratum in the streamside clearcut vs. old-growth forests of western Washington. Pages 121-130
forest ecosystem. Oikos 26: 391-397. in: W. R. Meehan, T. R. Merrell, Jr., T. A. Hanley, editors. Proceedings of a
35. Ben-David, M., R.W. Flynn, and D.M. Schell. 1997. Annual and seasonal Symposium: Fish and Wildlife Relationships in Old-growth Forests. April,
changes in diets of martens: evidence from stable isotope analysis. 1982, Juneau, AK.
Oecologia 111:280-291. 53. ———, T. Quinn, G. H. Reeves, and S.V. Gregory. 1992. Best
36. Ben-David, M. T. 1997. Timing of reproduction in wild mink: the management practices, cumulative effects, and long-term trends in fish
influence of spawning Pacific salmon. Canadian Journal of Zoology abundance in Pacific Northwest river systems. Pages 223-265 in: R. J.
75:376-382. Naiman, editor. Watershed management, Balancing Sustainability and
37. ———, A. Hanley, D. R. Klein, and D. M. Schell. 1997. Seasonal changes Environmental Change. Springer-Verlag. New York, NY.
in diets of coastal and riverine mink: the role of spawning Pacific 54. ——— G. H. Reeves, R. E. Bilby, and R. J. Naiman. 1997. Watershed
salmon. Canadian Journal of Zoology 75:803-811. management and Pacific salmon: Desired future conditions. Pages 447-
474 in: Pacific Salmon and Their Ecosystems: Status and Future Options.
Chapman and Hall, New York, NY.
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 661

55. ——— and R. E. Bilby. 1998. Organic matter and trophic dynamics. In: 72. Browne, P., R. L. DeLong, H. R. Huber, and J. L. Laake. 1997. Pinniped
R. J. Naiman and R. E. Bilby, editors. River Ecology and Management: predation on endangered salmonids in Washington and Oregon: harbor
Lessons From the Pacific Coastal Ecoregion. Springer-Verlag. New York, seal food habits on the Columbia River. Pages 109-118 in: P. S. Hill and
NY. D. P. DeMaster, editors. Marine Mammal Protection Act and Endangered
56. Bjornn, T. C., and D. W. Reiser. 1991. Habitat requirements of Species Act Implementation Program 1996. AFSC Processed Report
salmonids in streams. Pages 83-138 in: W. R. Meehan, editor. Influences 97-10.
of Forest and Rangeland Management on Salmonid Fishes and Their 73. Burcham, J. S. 1904. Notes on the habits of the water ousel (Cinclus
Habitats. American Fisheries Society Special Publication No.19. mexicanus). Condor. 6:50.
57. ———. 1971. Trout and salmon movements in two Idaho streams as 74. Burger, A. E, R. P. Wilson, D. Garnier, and M. P. T. Wilson. 1993. Diving
related to temperature, food, stream flow, cover, and population depths, diet, and underwater foraging of Rhinoceros Aucklets in British
density. Transactions of American Fisheries Society 100:423-438. Columbia. Canadian Journal of Zoology 71:2528-2540.
57a. Bledsoe, L. J. et al. The Puget Sound runs of salmon: An examination 75. Burgess, S. A. and J. R. Bider. 1980. Effects of stream habitat
of the changes in run size since 1896. Pages 50-61 in: C. D. Levings et improvements on invertebrates, trout populations, and mink activity.
al., editors. Proceedings of the National Workshop on Effects of Habitat Journal of .Wildlife Management 44:871-880.
Alteration on salmonid stocks. Canadian Special Publication Fisheries 76. Burger, A. E. 1993 (1994). Mortality of seabirds assessed from
and Aquatic Science. No. 105. beached-bird surveys in southern British Columbia. Canadian Field-
58. Bollen, W. B., and K. C. Lu. 1968. Nitrogen transformations in soils Naturalist 107:164-176.
beneath red alder and conifers. Pages 141-148 in: J. M. Trappe, J. F. 77. Burgner, R. L. 1991.The life history of sockeye salmon (Oncorhynchus
Franklin, R. F. Tarrant, and G. M. Hansen, editors. Biology of Alder. USDA nerka). Pages 1-118 in: C. Groot and L. Margolis, editors. Pacific Salmon
Forest Service. Portland, OR. Life Histories. University of British Columbia Press,Vancouver, BC,
59. Booth, D. B., and C. R. Jackson. 1997. Urbanization of aquatic systems: Canada.
Degradation thresholds, stormwater detection, and the limits of 78. Bury, R. B., and P. S. Corn. 1988. Responses of aquatic and streamside
mitigation. Journal of American Water Resources Association 33:1077- amphibians to timber harvest: A review. Pages 165-188 in: K. J. Raedeke,
1090. editor. Streamside Management: Riparian Wildlife and Forestry
60. Booth. D. E. 1991. Estimating prelogging old-growth in the Pacific Interactions. Contribution No. 59, Institute of Forest Resources,
Northwest. J. For. 89:25-29. University of Washington, Seattle, WA.
61. Bormann, F. H., and G. E. Likens. 1967. Nutrient cycling. Science 79. Busby, P. J., T. C. Wainwright, G. J. Bryant, L. J. Lierheimer, R. S. Waples, F.
155:424-429. W. Waknitz, and I. V. Lagomarsino. 1996. Status review of west coast
62. Brickell, D. C., and J. J. Goering. 1970. Chemical effects of salmon steelhead from Washington, Idaho, Oregon, and California U.S. Dept. of
decomposition on aquatic ecosystems. Pages 125-138 in: R. S. Murphy, Commerce. NOAA Tech. Memo. NMFS-NWFSC-27:261 pp.
editor. Proceedings of the Symposium on Water Pollution Control in 80. Bustard, D. R., and D. W. Narver. 1975. Aspects of the winter ecology
Cold Climates. U. S. Government Printing Office, Washington, DC. of juvenile coho salmon (Oncorhynchus kisutch) and steelhead trout
63. Briggs, D. T. and C. W. Davis. 1972. A study of predation by sea lions on (Salmo gairdneri). Canadian Journal of Fisheries and Aquatic Sciences. 32:
salmon in Montery Bay. California Fish Game 58:37-43. 667-80.
64. Brinson, M. M., B. L. Swift, R. C. Plantico, and J. S. Barclay. 1981. Riparian 81. Butler, R. L. 1991. Trout as predator. Pages 65-72 in: J. Stolz and J.
ecosystems: Their ecology and status. USDI Fish and Wildlife Service, Schnell, editors. Trout, the Wildlife Series. Stackpole Books. Harrisburg,
Biological Services Program. Kearneysville, WV. PA.
65. Brittell, J. D. Sweeney and S. T. Knick. 1979. Washington bobcats: diet, 82. Butler, R. W. 1973. Trumpeter swans eating salmon.Vancouver Nat.
population dynamics and movement. National Wildlife Federation Hist. Soc. Discovery (New Ser.). 2:120.
Scientific and Technical Service No. 6: 107-110. 83a. Brodeur, R. D. 1989. Neutonic feeding by juvenile salmonids in
65a. Brodeur, R. D. 1989. Neutonic feeding by juvenile salmonids in coastal waters of the Northeast Pacific. Canadian Journal of Zoology
coastal waters of the Northwest Pacific. Canadian Journal of Zoology 67:1995-2007.
67:1995-2007. 83b. ———, and W. G. Pearcy. 1990. Trophic relations of juvenile Pacific
65b. ———, and W. G. Pearcy. 1990. Trophic relations of juvenile Pacific salmon off the Oregon and Washington coasts. Fisheries Bulletin
salmon off the Oregon and Washington coasts. Fisheries Bulletin 88:617-630.
88:617-630. 84. California Department of Fish and Game. 1993. Final environmental
66. Brooks, A. 1928. Does the marbled murrelet nest inland? Murrelet document regarding bear hunting. Resource Agency, California
9:68. Department of Fish and Game, Sacramento, CA.
67. Brown, R. F., S. Riemer, and S. J. Jeffries. 1995. Food of pinnipeds 85. Campbell, R. W. and N. K. Dawe. 1990. The birds of British Columbia,
collected during the Columbia River area salmon gillnet observation Volume 1, Nonpasserines: Loons through waterfowl. Royal British
program, 1991-1994. Oregon Dept. of Fish and Wildlife, Wildlife Columbia Museum, Victoria, BC, Canada.
Diversity Program, Technical Report #95-6-01 (Available From Oregon 85a. ———, ———, I. McTaggart-Cowan, J.M. Cooper, G.W. Kaiser, and
Dept. of Fish and Wildlife, 2501 SW First, Portland, OR 97201). M.C.E. McNall. 1990. The birds of British Columbia, volume III:
67a. Brown, H.A., R.B. Bury, D.M. Darda, L.V. Diller, C.R. Peterson, and Passerines, flycatchers through vireos. Royal British Columbia Museum,
R.M. Storm. 1995a. Reptiles of Washington and Oregon. Seattle Victoria, BC, Canada.
Audubon Society, Seattle, WA. 176pp. 86. Carlson, A. 1991. Characterization of riparian management zones and
68. Brown, R. F., and B. R. Mate. 1983. Abundance, movements, and feeding upland management areas with respect to wildlife habitat.. Washington
habits of harbor seals, Phoca vitulina, at Netarts and Tillamook Bays, Timber/Fish/Wildlife Rep. T/F/W-WLI-91-001. Washington Department
Oregon. U. S. Fish and Wildlife Service Fish. Bull. 81(2):291-301. of Natural Resources, Olympia, WA.
69. Brown, E. R. 1985. Riparian zones and freshwater wetlands. 87. Carman, R. E., C. J. Cederholm, and E. O. Salo. 1984. A baseline
Management of wildlife and fish habitats in forests of western Oregon inventory of juvenile salmonid populations and habitats in streams in
and Washington, Part I—Chapter Narratives. USDA, Forest Service. Capitol Forest, Washington 1981-1982. Progress Report for Sampling
70. Brown, B. 1982. Mountain in the clouds: A search for wild salmon. During 1981 and 1982, University of Washington, Fisheries Research
Simon and Schuster, New York, NY. Institute. Report No. FRI-UW-8416, Seattle, WA.
71. Brown, S., M. M. Brinson, and A. E. Lugo. 1979. Structure and function 88. Carter, H. R. and S. G. Sealy. 1984. Marbled murrelet mortality due to
of riparian wetlands. Pages 17-31 In: R.R. Johnson and J. F. McCormick, gill-net fishing in Barkley Sound, British Columbia. Pages 212-220 in: D.
editors. Strategies for protection and management of floodplain N. Nettleship, G. A. Sanger, and P. F. Springer, editors. Marine Birds: Their
wetlands and other riparian ecosystems. USDA For. Ser. Gen. Tech. Rep. Feeding Ecology and Commercial Fisheries Relationships. Canadian
WO-12. Wildlife Service.
 662 Wildlife–Habitat Relationships in Oregon and Washington

89. Case, D. J., and D. R McCullough. 1987. White-tailed deer forage on 109. Craig, J. A., and R. L. Hacker. 1940. The history and development of
alewife. Journal of Mammalogy. 68:195-197. the fisheries of the Columbia River. Bulletin of the Bureau of Fisheries,
90. Cederholm, C. J., and N. P. Peterson. 1985. The retention of coho Vol. XLIX.
salmon (Oncorhynchus kisutch) carcasses by organic debris in small 110. Cressman, L.S. 1960. Cultural sequences at the Dalles, Oregon. A
streams. Canadian Journal of Fisheries and Aquatic Sciences 42:1222- contribution to Pacific Northwest prehistory. Transactions of the
1225. American Philosophical Society,Vol. 50, Part 10.
91. ———, D. B. Houston, D. L. Cole, and W. J. Scarlett. 1989. Fate of 110a. Crutchfield, J. A., and G. Pontecorvo. 1969. The Pacific salmon
coho salmon (Oncorhynchus kisutch) carcasses in spawning streams. fisheries: A study of irrational conservation. The John Hopkins Press,
Canadian Journal of Fisheries and Aquatic Sciences 46:1347-1355. Baltimore, MD.
92. ———, Kunze, M. D., Murota, T., and A. Sibatani. 1999. Pacific salmon 111. Cummins, K. W., M. A. Wilzbach, D. M. Gates, J. B. Perry, and W. B.
carcasses: Essential contributions of nutrients and energy for aquatic Taliaferro. 1973. Shredders and riparian vegetation. Bioscience 39:24-30.
and terrestrial ecosystems. Fisheries 24 (10):6-15. 112. Dahm, C. C. 1981. Pathways and mechanisms for removal of
93. ———, and E. O. Salo. 1979. The effects of logging road landslide dissolved organic carbon from leaf leachate in streams. Canadian
siltation on the salmon and trout spawning gravels of Stequaleho Creek Journal of Fisheries and Aquatic Science 38:68-76.
and the Clearwater River Basin, Jefferson County, Washington. 113. Day, J. W., Jr., C. A. S. Hall, W. M. Kemp, and A. Yáñez-Arancibia. 1989.
University of Washington, Fisheries Research Institute. Report No. FRI- Estuarine Ecology. John Wiley & Sons, New York, NY.
UW-7915. Seattle, WA. 114. Dalquest, W. W. 1948. Mammals of Washington. University of Kansas
94. ———, and W. J. Scarlett. 1982. Seasonal immigrations of juvenile Publication Volume 2, Museum of Natural History, University of Kansas,
salmonids into four small tributaries of the Clearwater River, Lawrence, KS.
Washington, 1977-198. Pages 98-110 in: E. L. Brannon and E. O. Salo, 115. Dawson, W. L. 1909. The Birds of Washington. A Complete, Scientific
editors. Proceedings of the Salmon and Trout Migratory Behavior and Popular Account of the 372 Species of Birds Found in the State.
Symposium, June 3-5, 1981. University of Washington, School of Seattle, WA.
Fisheries, Seattle,WA. 116. Death, R. G., and M. J. Winterbourn. 1995. Diversity patterns in
95. Chapman, D. W., and T. C. Bjornn. 1969. Distribution of salmonids in stream benthic invertebrate communities: the influence of habitat
streams, with special reference to food and feeding. Pages 153-176 in: T. stability. Ecology 76:1446-1460.
G. Northcote, editor. Symposium On Salmon and Trout In Streams, H. 117. DeGange, A. R. 1996. The marbled murrelet: a conservation
R. MacMillan Lectures In Fisheries. February 22-24, 1968, the University assessment. General Technical Report PNW-GTR-388. U.S.
of British Columbia. Vancouver, BC, Canada. Department of Agriculture, Forest Service, Pacific Northwest Research
96. ———. 1962. Aggressive behavior in juvenile coho salmon as a cause Station, Portland, OR.
of emigration. Canadian Journal of Fisheries and Aquatic Sciences 118. DeShazo, J. J. 1980. Sea-run cutthroat trout management in
19:1047-1080. Washington - An overview. Fisheries Management Division, Washington
97. Chi, D. K. 1999. The effects of salmon availability, social dynamics, and State Game Dept., Olympia, WA.
people on Black Bear (Ursus americanus) fishing behavior on an Alaskan 119. DeVoto, B. 1981. The journals of Lewis and Clark. Houghton Mifflin
salmon stream. Ph.D. Dissertation. Utah State University, Logan, UT. Company. Boston, MA.
98. Clark, E. E. 1984. Indian legends of the Pacific Northwest. University 120. Dolloff, C. A. 1993. Predation by river otters (Lutra canadensis) on
of California Press. Berkeley, CA. 225 pp. juvenile coho salmon (Oncorhynchus kisutch) and Dolly Varden
99. Clemens, W. A. and G.V. Wilby. 1933. Food of the fur seal off the coast (Salvelinus malma) in southeast Alaska. Canadian Journal of Fisheries
of British Columbia. Journal of Mammalogy 14:43-46. and Aquatic Science 50:312-315.
100. Coffman, W. P., and L. C. Ferrington, Jr. 1996. Chironomidae. Pages 121. Donaldson, I. J., and F. K. Cramer. 1971. Fishwheels of the Columbia.
635-754 In: R. W. Merritt and K. W. Cummins, editors. Kendall/Hunt Binfords and Mort, Publishers. Portland, OR.
Publishing. Dubuque, IA. 122. Donaldson, J. R. 1967. The phosphorus budget of Iliamna Lake, Alaska
101. Commencement Bay Natural Resource Trustees. 1997. as related to the cyclic abundance of sockeye salmon. Ph.D.
Commencement Bay Natural Resource Damage Assessment Dissertation, University of Washington, School of Fisheries, Seattle, WA.
Restoration Plan and Final Programmatic Environmental Impact 123. Drummond, H. 1983. Aquatic foraging in garter snakes: a comparison
Statement. Prepared by the U. S. Fish and Wildlife Service and the of specialists and generalists. Behaviour 86:1-30.
National Oceanic and Atmospheric Administration for the 124. Dunstone, N. 1993. The mink. T & A. D. Poyser Ltd, London.
Commencement Bay Natural Resource Trustees and Cooperating 125. Dzinbal, A., and R. L. Jarvis. 1982. Coastal feeding ecology of
Agencies. Olympia and Seattle, WA. harlequin ducks in Prince William Sound, Alaska, during summer. Pages
102. Conaway, C. H. 1952. Life history of the water shrew (Sorex palustris 6-10 in: D. N. Nettleship, G. A. Sanger and P. F. Springer, editors. Marine
navigator).The American Midland Naturalist 48:219-248. Birds: Feeding Ecology and Commercial Fisheries Relationships.
103. Cone, J., and S. Ridlington. 1996. The Northwest salmon crisis—a Proceedings of the Pacific Seabird Group Symposium, Seattle, WA.
documentary history. Oregon State University Press, Corvallis, OR. 126. Eagle, T. C., and J. S. Whitman. 1987. Mink. Pages 614-624 in: M.
104. Congleton, J. L., S. K. Davis, and S. R. Foley. 1982. Distribution, Novak, J. A. Baker, M. E. Obbard and B. Malloch, editors. Wild Furbearer
abundance and outmigration timing of chum and chinook salmon fry in Management and Conservation in North America. Ministry of Natural
the Skagit salt marsh. Pages 153-163 In: E. L. Brannon and E. O. Salo, Resources, Ontario, Canada.
editors. Proceedings of the Salmon and Trout Migratory Behavior 127. Eastman, D. E. 1996. Response of freshwater fish communities to
Symposium, June 3-5, 1981. University of Washington, School of spawning sockeye salmon (Oncorhynchus nerka). M. S.Thesis, University
Fisheries, Seattle, WA. of Washington, School of Fisheries, Seattle, WA.
105. Connell, J. H. 1978. Diversity in tropical rain forests and coral reefs. 128. Edie, B. G. 1975. A census of the juvenile salmonids of the
Science 199:1302-1310. Clearwater River Basin, Jefferson County, Washington, in relation to
106. Cooper, R., and T. H. Johnson. 1992. Trends in steelhead abundance in logging. M. S. Thesis. University of Washington, School of Fisheries,
Washington and along the Pacific coast of North America. Report #92- Seattle, WA.
90. Fisheries Management Division. Washington Department of Wildlife, 128a. Edmonds, R. L. 1980. Litter decomposition and nutrient release in
Olympia, WA. Douglas-fir, red alder, western hemlock, and Pacific silver fir ecosystems
107. Cottam, C. 1939. Food habits of North American diving ducks. U.S. in western Washington. Canadian Journal of Forestry Res. 10:327-337.
Departmen of Agriculture Technical Bulletin No. 643. 129. Ehinger, C. E. 1930. Some studies of the American Dipper or Water
108. ———, and Uhler F. M. 1937. Birds in relation to fishes. Wildlife Ouzel. Condor 47:487-498.
Research and Management Leaflet BS-83. 130. Eipper, A. W. 1956. Differences in the vulnerability of the prey of
nesting kingfishers. Journal of Wildlife Management 20:177-183.
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 663

131. Elliott, S.T., and R. Bartoo. 1981. Relation of larval Polypedilum 154. Franklin, J. F. 1992. Scientific basis for new perspectives in forests and
(Diptera: Chironomidae) to pink salmon eggs and alevins in an Alaska streams. Pages 25-72 in: R. J. Naiman, editor. Watershed Management:
stream. Prog. Fish-Culturist 43: 220-221. Balancing Sustainability and Environmental Change. Springer-Verlag,
132. Elmore, W. 1992. Riparian responses to grazing practices. Pages 442- New York, NY.
457 in: R. B. Naiman, editor. Watershed Management: Balancing 155. Friesen, W. 1990. Winter dietary studies of juvenile coho salmon
Sustainability and Environmental Change. Springer-Verlag. New York, (Oncorhynchus kisutch) utilizing two enhanced wall-base channels along
NY. the Clearwater River in Jefferson County, Washington. Master of
133. Elphick, C.S., and T.L. Tibbitts. 1998. Greater Yellowlegs (Tringa Environmental Studies, The Evergreen State College, Olympia, WA.
melaoleuca). The Birds of North America, No. 355. A. Poole and F. Gill, 156. Fraser, J. M. 1972. Recovery of planted brook trout, splake, and
editors. The Academy of Natural Sciences, Philadelphia, PA, and The rainbow trout from selected Ontario lakes. Journal of Fisheries
American Ornithologists Union, Washington D.C. Research Board Can. 29:129-142.
134. Elson, P. F. 1962. Predator-prey relationship between fish-eating birds 157. ———. 1974. An attempt to train hatchery reared brook trout to
and Atlantic salmon. Fisheries Research Board Can. Bulletin No 133. avoid predation by the common loon. Transactions of the American
135. Emmett, R. L. 1997. Estuarine survival of salmonids: The importance Fisheries Society 103:815-818.
of interspecific and intraspecific predation and competition. Pages 147- 158. French, J. M. and J. R. Koplin. 1977. Distribution, abundance, and
158 in: R.L. Emmett and M.H. Schiewe, editors. Estuarine and Ocean breeding status of ospreys in northwestern California. Trans. North
Survival of Northeastern Pacific Salmon. NOAA Technical American Osprey Research Conference U.S. National Park Service
Memorandum NMFS-NWFSC-29. Transactions Proceedings Series No. 2: 223-240.
136. ——— and M. H. Schiewe. 1997. Estuarine and ocean survival of 159. Fujiwara, M., and R. C. Highsmith. 1997. Harpacticoid copepods:
Northeastern Pacific salmon: Proceedings of the workshop. U.S. Dept. Potential trophic link between inbound adult salmon and outbound
Commerce., NOAA Technical Memo. NMFS-NWFSC-29. juvenile salmon. Marine Ecology Progress Series 158:205-216.
137. Erman, D. C., J. D. Newbold, and K. B. Roby. 1977. Evaluation of 160. Fuss, H. J. 1982. Age, growth, and instream movement of Olympic
streamside bufferstrips for protecting aquatic organisms. California Peninsula coastal cutthroat trout (Salmo clarki clarki). M. S.Thesis.
Water Resources Center, Contribution Number 165, University of University of Washington, School of Fisheries, Seattle, WA.
California, Davis, CA. 161. Gabrielson, I. N., and S. G. Jewett. 1940. Birds of Oregon. Oregon
138. Erickson, J. 1988. Competition for fish. Headlight Herald. June 4th(A- State College, Corvallis, OR.
4). 162. Garrett, A. M. 1998. Interstream movements of coastal cutthroat
139. Eriksson, M. O. G. D. Blomqvist M. Hake and O. C. Johansson. 1990. trout (Oncorhynchus clarki clarki) in the Clearwater River, Jefferson
Parental feeding in the Red-throated Diver Gavia stellata. Ibis.132:1-13. County, Washington. Master of Environmental Studies. The Evergreen
140. Everest, F. H., N. B. Armantrout, S. M. Keller, W. D. Parante, J. R. Sedell, State College, Olympia, WA.
T. E. Nickelson, J. M. Johnston, and G. N. Haugen. 1985. Salmonids. Pages 163. Gaston, A. J and S. B. C. Dechesne. 1996. Rhinoceros Auklet,
199-226 in: R. Brown, editor. Management of Wildlife and Fish Habitats Cerorhinca monocerata. In:A. Poole and F. Gill, editors.The Birds of
in Forests of Western Oregon and Washington. Volume 1. USDA, Forest North America No. 212. The Academy of Natural Sciences, Philadelphia,
Service. Pacific Northwest Region. PA, and The American Ornithologist’s Union, Washington, D.C.
141. Everitt, R. D., P. J. Gearin, J. S. Skidmore, and R. L. DeLong. 1981. Prey 164. Gearin, P. J., S. R. Melin, R. L. DeLong, H. Kajimura, and M. A. Johnson.
items of harbor seals and California sea lions in Puget Sound, 1994. Harbor porpoise interactions with a chinook salmon set-net
Washington. Murrelet 62:83-86. fishery in Washington state. Pages 427-438 in: W. F. Perrin, G. P.
142. Farley, L. C. 1980. The Behavioral-Ecology of Swans Wintering in Donovan, and J. Barlow, editors. Gillnets and Cetaceans: Incorporating
Southeast Alaska. MS Thesis. Idaho State University. the Proceedings of the Symposium and Workshop on the Mortality of
143. Felleman, F. L. J. R. Heimlich-Boran and R. W. Osborne. 1991. The Cetaceans in Passive Fishing Nets and Traps. La Jolla, CA. October
feeding ecology of killer whales (Orcinus orca) in the Pacific Northwest. 1990. Whaling Commission Special Issue No.15.
Pages 113-147 in: K. Pryor, and K. S. Norris, editors. Dolphin Societies, 165. ———, R. Pfeifer, S. J. Jeffries, R. L. DeLong, and M. A. Johnson. 1988.
Discoveries and Puzzles. University of California Press, Berkeley, CA. Results of the 1986-1987 California sea lion—steelhead trout
144. Fiscus, C. H. and G. A. Baines. 1966. Food and feeding behavior of predation control program at the Hiram M. Chittenden Locks. U. S.
Steller and California sea lions. Journal of Mammalogy 47:195-200. Department of Commerce. NWAFC Processed Report 88-30.
145. Fisher, J. P., and W. G. Pearcy. 1988. Growth of juvenile coho salmon 166. Glova, G. J. 1986. Interactions for food and space between
(Oncorhynchus kisutch) in the ocean off Oregon and Washington, USA, in experimental populations of juvenile coho salmon (Oncorhynchus
years of differing coastal upwelling. Canadian Journal of Fisheries and kisutch) and coastal cutthroat trout (Salmo clarki) in a laboratory
Aquatic Sciences 45:1036-1044. stream. Hydrobiologia 132:155-168.
146. Foerster, R. E. 1968.The sockeye salmon (Oncorhynchus nerka). 167. Gould,V. E. 1934. A monograph of the belted kingfisher Megaceryle
Fisheries Research Board Can. Bulletin No. 162. Ottawa, Canada. alcyon (Linnaeus).
147. Fitch, H. S. 1984. Thamnophis couchii. Catalogue of American 168. Graf, W. 1949. Observations on the salamander Dicamptodon .
Amphibians and Reptiles. 351.1-351.3. Copeia 1:79-80.
148. ———. 1941. The feeding habits of California garter snakes. 169. Graybill, M.R., and J. Hodder. 1985. Effects of the 1982-1983 El Niño
California Fish and Game 27:2-32. on reproduction of six species of seabirds in Oregon. Pages 205-210 in:
149. Fontaine, P. M. M. O. Hammill C. Barrette and M. C. Kingsley. 1994. W. S. Wooster and D. L. Fluharty, editors. El Niño North. Washington
Summer diet of the harbor porpoise (Phocoena phocoena) in the Sea Grant Program, University of Washington, Seattle, WA.
estuary and the Northern Gulf of St. Lawrence. Canadian Journal of 170. ———. 1981. Haul out patterns and diet of harbor seals, Phoca
Fisheries and Aquatic Science 5:172-178. vitulina, in Coos County, Oregon. M.S.Thesis, University of Oregon,
150. Forbes, L. C. and K. Simpson. 1982. Behavioral studies of great blue Eugene, OR.
herons at Pender Harbor and Sechelt, British Columbia, in 1980. 171. Gregory, S. V., F. J. Swanson, and W. A. McKee. 1991. An ecosystem
Unpublished Report., Canadian Wildlife Service, Delta, B.C, Canada. perspective of riparian zones. BioScience 40:540-551.
151. Forest Ecosystem Management Assessment Team (FEMAT). 1993. 172. ———. 1980. Effects of light, nutrients, and grazing on periphyton
Forest Ecosystem Management: An Ecological, Economic, and Social communities in streams. Ph.D. Dissertation. Oregon State University,
Assessment. USDA, Forest Service. Corvallis, OR.
152. Frame, G. W. 1974. Black bear predation on salmon at Olsen Creek, 173. ———, G. A. Lamberti, D. C. Erman, K. V. Koski, M. L. Murphy, and J. R.
Alaska. Zeitung der Tierpsychol. 35:23-38. Sedell. 1987. Influences of forest practices on aquatic production. Pages
153. Francis, R. C., and S. R. Hare. 1994. Decadal-scale regime shifts in the 233-255 In: E. O. Salo and T. Cundy , editors. Streamside Management—
large marine ecosystems of the Northeast Pacific: A case for historical Forestry and Fishery Interactions. College of Forest Resources,
science. Fish. Oceanogr. 3:279-291. University of Washington. Contribution No. 57. Seattle, WA.
 664 Wildlife–Habitat Relationships in Oregon and Washington

174. ———, and P. A. Bisson. 1997. Degradation and loss of anadromous 196. Healey, M. C. 1982. Juvenile Pacific salmon in estuaries: The life
salmonid habitat in the Pacific Northwest. Pages 277-314 in: D. J. support system Pages 315-341 in:V. S. Kennedy, editor. Estuarine
Stouder, P. A. Bisson, and R. J. Naiman, editors. Pacific Salmon and their Comparisons. Academic Press. New York, NY.
ecosystems. Chapman and Hall Publishers, New York, NY. 197. ———. 1991. Life history of chinook salmon (Oncorhynchus
175. Grette, G. 1985. The role of large organic debris in juvenile salmonid tshawytscha). Pages 311-393 in: C. Groot and L. Margolis, editors. Pacific
rearing habitat in small streams. M.S. Thesis, University of Washington, Salmon Life Histories, University of British Columbia Press.Vancouver,
School of Fisheries. Seattle, WA. BC, Canada.
176. Grinnell, J. J. Dixon and J. M. Linsdale. 1937. Fur-bearing mammals of 198. Heard,W. R. 1991. Life history of pink salmon (Oncorhynchus
California. Their natural history, systematic status, and relations to man. gorbuscha). Pages 121 to 230 in: C. Groot and L. Margolis, editors.
University of California Press, Berkeley, CA. Pacific Salmon Life Histories, University of British Columbia Press.
177. Griswold, G. 1953. Aboriginal patterns of trade between the Vancouver, B.C.
Columbia Basin and the northern plains. M. A. Thesis, Montana State 199. Heimlich-Boran, J. R. 1986. Fishery correlations with the occurrence
University. Missoula, MT. of killer whales in Greater Puget Sound. Pages 113-131 in: B. C.
178. Groot, C., and L. Margolis. 1991. Pacific Salmon Life Histories, Kirkevold, and J. S. Lockard, editors. Behavioral biology of killer whales.
University of British Columbia Press, Vancouver, BC, Canada. Alan R. Liss, Inc., New York, NY.
179. Gross, M. R., R. M. Coleman, and R. M. McDowell. 1988. Aquatic 200. ———. 1988. Behavioral ecology of killer whales (Orcinus orca) in
productivity and the evolution of diadromous fish migration. Science the Pacific Northwest. Canadian Journal of Zoology 66:565-578.
239:1291-1293. 201. Hemstrom, M. A., and J. F. Franklin. 1982. Fire and other disturbances
180. Haig-Brown, R. 1946. A river never sleeps. Crown Publishers. New of the forests in Mount Rainier National Park. Quaternary Research
York, NY. 18:32-51.
181. Hall, J. D., P. A. Bisson, and R. E. Gresswell. 1997. Sea-run cutthroat 202. Henny, C. J. and M. R. Bethers. 1971. Population ecology of the great
trout-biology, management , and future considerations. Proceedings of a blue heron with special reference to western Oregon. Canadian Field-
Symposium, Reedsport, Oregon. October 12-14, 1995. Published by the Naturalist 85:205-209.
Oregon Chapter of the American Fisheries Society. Corvallis, OR. 203. Herder, M. J. 1983. Pinniped fishery interactions in the Klamath River
182. ———. 1986. Notes on the distribution and feeding behavior of system, July 1979 to October 1980. Southwest Fish. Centennial
killer whales in Prince William Sound, Alaska. Pages 69-83 in: B. C. Administrative Report LJ-83-12C. (Available From Southwest Fisheries
Kirkevold, and J. S. Lockard, editors. Behavioral biology of killer whales. Science Center, National Marine Fisheries Services, NOAA, P.O. Box
Alan R. Liss, Inc., New York. NY. 271, La Jolla, CA 92038).
183. Hamilton, A. N. and W. R. Archibald. 1985. Grizzly bear habitat in the 204. Hewson, R. 1995. Use of salmonid carcasses by vertebrate
Kimsquit River Valley, coastal British Columbia: evaluation. Pages 50-57 scavengers. Journal of Zoology 235:53-65.
in: G. P. Contreras, and K. E. Evans, editors. Proceedings—Grizzly Bear 205. Hildebrand, G. V., S. D. Farley, C. T. Robbins, T. A. Hanley, K. Titus, and
Habitat Symposium, Missoula, Montana, April 30 - May 2, 1985. C. Servheen. 1996. Use of stable isotopes to determine diets of living
Intermountain Research Station, Ogden, UT. and extinct bears. Canadian Journal of Zoology 74:2080-2088.
184. ———, and F. L. Bunnell. 1987. Foraging strategies of coastal grizzly 207. Hicks, B.J., R. L. Beschta, and R. D. Harr. 1991b. Long-term changes in
bears in the Kimsquit River Valley, British Columbia. International streamflow following logging in western Oregon and associated
Conference on Bear Research Management 7:187-197. fisheries implications. Water Resources Bulletin 27:217-226.
185. Hampton, P. D. 1981. The wintering and nesting behavior of the 208. ———, J. D. Hall, P.A. Bisson, and J. R. Sedell. 1991a. Response of
trumpeter swan. M.S. Thesis. University of Montana. salmonids to habitat changes. Pages 483-518 in: W.R. Meehan, editor.
186. Hansen, W. R. 1980. Western aquatic garter snakes in central Influences of Forest and Rangeland Management On Salmonid Fishes
California: an ecological and evolutionary perspective. M.S. Thesis, and Their Habitats. American Fisheries Society Special Publication 19.
California State University, Fresno, CA. Bethesda, MD.
187. Hansen, A. J., E. L. Boeker, J. I. Hodges, and D. R. Cline. 1984. Bald 209. Hilborn, R. 1992. Hatcheries and the future of salmon in the
eagles of the Chilkat Valley, Alaska: ecology, behavior, and management. Northwest. Fisheries 17:5-8.
National Audubon Society. New York, NY. 210. Hodder, J., and M.R. Graybill. 1985. Reproduction and survival of
188. Hanson, L. C. 1993. The foraging ecology of harbor seals, Phoca seabirds in Oregon during the 1982-1983 El Niño. Condor 87:535-541.
vitulina, and California sea lions, Zalophus californianus, at the mouth of 211. Hogan, D.L., and J. W. Schwab. 1991. Stream channel response to
the Russian River, California. M.A. Thesis, Sonoma State University, landslides in the Queen Charlotte Islands, B.C.: changes affecting pink
Rohnert Park, CA. and chum salmon habitat. Pages 222-236 in: B. White and I. Gutherie,
189. Hare, S. R., N. J. Mantua and R. C. Francis. 1999. Inverse production editors. Proceedings Of the 15th Northeast Pacific Pink and Chum
regions: Alaska and West Coast Pacific salmon. Fisheries 24: 6-14. Workshop. Canada Department of Fisheries and Oceans.Vancouver,
190. Hartman, W., and R. Raleigh. 1964. Tributary homing of sockeye BC, Canada.
salmon at Brooks and Karluk lakes, Alaska. Canadian Journal of 212. Hoffman, T. and T. Hall. 1988. Tillamook trip report regarding smolt/
Fisheries and Aquatic Science 21:485-504. cormorant problem. U. S. Dept. of Agriculture, Animal Damage Control
191. Harvey, J. T. and M. J. Weise. 1997. Impacts of California sea lions and Program Interoffice Memorandum About Their Cormorants Collection
Pacific harbor seals on salmonids in Monterey Bay, California. MLML on 27 April 1988.
Technical Publication No.97-03. Moss Landing Marine Laboratories. 213. Holland, D. C. 1985a. Clemmys marmorata (western pond turtle).
192. Hasegawa, S., T. Hirano, T. Ogasawara, M. Iwata, T. Akiyama, and S. Arai. Feeding. Herpetol. Review 16:112-113.
1987. Osmoregulatory ability of chum salmon, Oncorhynchus keta, 214. ———. 1985b. An ecological and quantitative study of the Western
reared in fresh water for prolonged periods. Fish Physiol. Biochem. Pond Turtle (Clemmys marmorata) in San Luis Obispo County, California.
4:101-110. M.S. Thesis, California State University, Fresno, CA.
193. Hatler, D. F. 1976. The coastal mink on Vancouver Island, British 215. Holtby, L. B. 1988. Effects of logging on stream temperatures in
Columbia. Ph. D. Dissertation, University British Columbia,Vancouver, Carnation Creek, British Columbia, and associated impacts on the
BC, Canada. coho salmon (Oncorhynchus kisutch). Canadian Journal of Fisheries and
194. Hayman, R. A., E. M. Beamer, and R. E. McClure. 1996. Fiscal Year 1995 Aquatic Sciences 45:502-515.
Skagit River chinook restoration research. Final project performance 216. Horner, R. R., and B. W. Mar. 1982. Guide for water quality impact
report, National Marine Fisheries Service, Contract# 3311 for FY 1995. assessment of highway operations and maintenance. Report to
Skagit System Cooperative. La Conner, WA. Washington Dept. of Transp. FHWA WA-RD-39.14. Dept. of Civil
195. Hayward, T. L. 1993. Preliminary observations of the 1991-1992 El Engineering, University of Washington, Seattle, WA.
Niño in the California Current. California Cooperative Oceanic 216a. Hughes. J. 1983. On osprey habitat and productivity: a tale of two
Fisheries Investigations Reports 34: 21-29. habitats. Pages 269-273 in D.M. Bird, editor. Biology and management of
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 665

bald eagles and ospreys. Proceedings of 1st International Symposium in the ‘Early Stuart’ stock spawning streams, 1050 km from the ocean.
on Bald Eagles and Ospreys, Montreal, 28-29 October 1981. Harpell Fisheries Project Report No. RD55, Fisheries Branch, Ministry of
Press, Ste Anne de Bellevue, Quebec. Environment, Lands, and Parks, Province of British Columbia, Canada.
217. Hunt, W. G., B. S. Johnson, and R. E. Jackman. 1992. Carrying capacity 240. Jones, R. E. 1981. Food habits of smaller marine mammals from
for bald eagles wintering along a northwestern river. Journal of Raptor northern California. Proceedings of the California Academy of Sciences
Research 26:49-60. 42:409-433.
218. ———, J. M. Jenkings, R. E. Jackman, C. G. Thelander, and A. T. 241. Juday, C., W. H. Rich, G. I. Kemmerer, and A. Mean. 1932. Limnological
Gerstell. 1992. Foraging ecology of bald eagles on a regulated river. studies of Karluk Lake, Alaska 1926-1930. Bulletin of the U. S. Bureau of
Journal of Raptor Research 26:243-256. Fisheries 47:407-436.
219. Hunt, W. A. 1993. Jasper National Park harlequin duck research 242. Jurek, R. M. 1974. Special wildlife investigations.: American River
project: 1992 pilot projects - interim results. Jasper Warden Service Green Heron study, 1974. California Department of Fish and Game.
Biological Report Series, No. 1, Heritage Resource Conservation, Parks 243. Kajimura, H. 1983. Food of the Pacific White-sided Dolphin,
Canada, Jasper, Alberta, Canada. Lagenorhynchus obliquidens, Dall’s Porpoise, Phocoenoides dalli, and
220. Hunt, W. G., B. S. Johnson, and R. E. Jackman. 1992. Carrying capacity Northern Fur Seal, Callorhinus ursinus, off California and Washington
for bald eagles wintering along a northwestern river. Journal of Raptor with appendices of size and food of Dall’s porpoise from Alaskan
Research 26(2):49-60. waters. U.S. National Marine Fisheries Service Technical Memorandum;
221. Huntington, C. W., W. Nelhsen, and J. Bowers. 1996. A survey of F/NWC-2. June 1980. From Monthly Catalog of U.S. Government
healthy native stocks of anadromous salmonids in the Pacific Publications 83-6731.
Northwest and California. Fisheries 21(3). 244. Kellyhouse, D. G. 1975. Habitat utilization by black bears in northern
222. Huntsman, S. G. 1941. Cyclical abundance of birds versus salmon. California. International Conference on Bear Research and
Journal of the Fisheries Research Board of Canada 5:227-235. Management 4:221-227.
223. Hynes, H. B. N. 1970. The ecology of running waters. University of 245. Kennish, M. J. 1997. Practical handbook of estuarine and marine
Liverpool Press. Liverpool, England. pollution. (CRC Press, Marine Science Series). CRC Press. Boca Raton,
224. Imler, R. H. and H. R. Sarber. 1947. Harbor seals and sea lions in FL.
Alaska. U. S. Fish. Wildlife Services Special Sci. Report No.28. 246. Kingery, H. E. 1996. American Dipper. The Birds of North America,
225. Science Team. 1998. Pinniped and seabird predation: implications for No. 229 in: A. Poole and F. Gill, editors. The Academy of Natural
recovery of threatened stocks of salmonids in Oregon under the Sciences, Philadelphia, PA, and The American Ornithologist’s Union,
Oregon Plan for Salmon and Watersheds. Technical Report 1998-2 to Washington, D.C.
the Oregon Plan for Salmon and Watersheds. Governor’s Natural 247. Kjelson, M. A., P. F. Raquel, and F. W. Fisher. 1982. Life history of fall-
Resources Office, Salem, OR. run juvenile chinook salmon, Oncorhynchus tshawytscha, in the
226. Inkpen, E. L., and S. S. Embrey. 1998. Nutrient transport in the major Sacramento-San Joaquin estuary, California. Pages 393-411 in:V. S.
rivers and streams of the Puget Sound Basin, Washington. U. S. Kennedy, editor. Estuarine Comparisons. Academic Press, New York,
Department of the Interior-U. S. l Survey, National Water-Quality NY.
Assessment Program. 248. Kline, T. C., Jr., J. J. Goering, O. A. Mathisen, P. H. Poe, and P. L. Parker.
227. Iwata, M., T. Hirano, and S. Hasegawa. 1982. Behavior and plasma 1990. Recycling of elements transported upstream by runs of Pacific
sodium regulation of chum salmon fry during transition into seawater. salmon: I. ð15N and ð13C evidence in Sashin Creek, southeastern
Aquaculture 28:133-142. Alaska. Canadian Journal of Fisheries and Aquatic Science 47: 136-144.
228. Jameson, R. J. and K. W. Kenyon. 1977. Prey of sea lions in the Rogue 249. ———, ———, ———, ———, ———, and R. S. Scalan. 1993.
River, Oregon. Journal of Mammalogy 58: 672. Recycling of elements transported upstream by runs of Pacific salmon:
229. Jeffries, S. 1985. Marine mammals of the Columbia River Estuary. II. ð15N and ð13C evidence in the Kvichak River watershed, Bristol Bay,
Washingotn. State Department of Game; 1984. From U.S. Government southwestern Alaska. Canadian Journal of Fisheries and Aquatic
Reports 85(2):85. Available From NTIS As PB85-107050/GAR. Sciences 50: 2350-2365.
230. Jewett, S. G., W. P. Taylor, W. T. Shaw, and J. W. Aldrich. 1953. Birds of 250. Knick, S. T. S. J. Sweeney J. R. Alldredge and J. D. Brittell. 1984. Autumn
Washington State. University of Washington Press, Seattle, WA. and winter food habits of Bobcats in Washington State. Great Basin
231. Johnson, W. E. and A. D. Hasler. 1954. Rainbow trout production in Naturalist 44:70-74.
dystrophic lakes. Journal of Wildlife Management 18:113-134. 251. Knight, R. L., P. J. Randolph, G. T. Allen, L. S.Young, and R. J. Wigen.
232. Johnson, J. H. and A. A. Wolman. 1984. The Humpback Whale, 1990. Diets of nesting bald eagles, Haliaeetus leucocephalus, in western
Megaptera novaeangliae. U. S. National Marine Fisheries Service Marine Washington. Canadian Field-Naturalist 104:545-551.
Fisheries Review 6:30-37. 252. ———, and D. P. Anderson. 1990. Effects of supplemental feeding on
233. Johnson, D. H., M. M. Hoover, E. L. Greda, and C. J. Cederholm. (In an avian scavenging guild. Wildlife Society Bulletin 18:388-394.
prep.). Relationships between Pacific salmon and 605 species of birds, 253. Knight, S. K and R. L. Knight. 1983. Aspects of food finding by
mammals, reptiles, and amphibians in Oregon and Washington. wintering bald eagles. Biology and management of bald eagles and
234. Johnson, O. W., M. H. Ruckelshaus, W. S. Grant, F. W. Waknitz, A. M. ospreys. Pages 28-29 in: D. M. Bird, editor. Proceedings of First
Garrett, G. J. Bryant, K. Neely, and J. J. Hard. 1999. Status review of International Symposium on bald eagles and ospreys Montreal.
coastal cutthroat trout in Washington, Oregon, and California. U.S. October, 1981. Harpell Press, Ste Anne De Bellevue, Quebec, Canada.
Dept. Commerce, NOAA Tech. Memo. NMFS-NWFSC-37. 254. Knudsen, E. E. 1999. Managing Pacific salmon escapements: the gaps
235. Johnson, R. R., and S. W. Carothers. 1982. Riparian habitats and between theory and reality. Pages 237-272 in: E. E. Knudsen, C. S.
recreation: Interelationships and impacts in the Southwest and Rocky Steward, D. D. MacDonald, J. E. Williams, and D. W. Reiser, editors.
Mountain region. Eisenhower Consortium for West. Environmental For. Sustainable Fisheries Management: Pacific Salmon. C. R. C. Lewis
Res. Bulletin 12: 1-31. Publishers, Boca Raton, FL.
236. Johnson, D.H., and T.A. O’Neil. 2001. Wildlife—habitat relationships 255. Knutson, K. L., and V. L. Naef. 1997. Management recommendations
in Oregon and Washington. Oregon State University Press, Corvallis, for Washington’s Priority habitats. Washington Department of Fish and
OR. Wildlife. Olympia, WA.
238. Johnston, J. M. 1982. Life histories of anadromous cutthroat with 256. Kojima, H., M. Iwata, and T. Kurokawa. 1993. Development and
emphasis on migratory behavior. Pages 123-127 in: E. L. Brannon and E. temporal decrease in seawater adaptability during early growth in
O. Salo, editors. Proceedings of the Salmon and Trout Behavior chum salmon, Oncorhynchus keta.Aquaculture118:141-150.
Symposium. School of Fisheries, University of Washington, Seattle, WA. 257. Konkel, G. W., and J. D. McIntyre. 1987. Trends in spawning
239. Johnston, N. T., J. S. MacDonald, K. J. Hall, and P. J. Tschaplinski. 1997. A populations of Pacific anadromous salmonids. Technical Report 9,
preliminary study of the role of sockeye salmon (Oncorhynchus nerka) USDI—Fish and Wildlife Service.
carcasses as carbon and nitrogen sources for benthic insects and fishes
 666 Wildlife–Habitat Relationships in Oregon and Washington

258. Koplin, J. R., D. S. MacCarter, D. P. Garber, and D. L. MacCarter. 1977. editors. Pacific Salmon and Their Ecosystems. Chapman and Hall
Food resources and fledgling productivity of California and Montana Publishers, New York, NY.
ospreys. Pages 205-214 in: Trans. North American Osprey Research 278. Likens, G. E., and F. H. Bormann. 1974. Linkages between terrestrial
Conference U.S. National Park Service Transactions of Proceedings and aquatic ecosystems. BioScience 24: 447-456.
Series No. 2. 279. Link, R. M., and K. K. English. 1999. Long-term, sustainable monitoring
259. Koski, K V. 1975. The survival and fitness of two stocks of chum of Pacific salmon populations using fish wheels to integrate harvesting,
salmon (Oncorhynchus keta) from egg deposition to emergence in a management, and research. Pages 667-674 in: E. E. Knudsen, C. S.
controlled-stream environment at Big Beef Creek. Ph.D. Dissertation, Steward, D. D. MacDonald, J. E. Williams, and D. W. Reiser, editors.
University of Washington, School of Fisheries. Seattle, WA. Sustainable Fisheries Management: Pacific Salmon. C. R. C. Lewis
260. ———, and D. Kirchofer. 1984. A stream ecosystem in an old- Publishers, Boca Raton, FL.
growth forest in Southeastern Alaska. Part IV: Food of juvenile coho 280. Lind, G. S. 1976. Production, nest site selection, and food habits of
salmon in relation to abundance of drift and benthos. Pages 81-88 In: W. ospreys on Deschutes National Forest, Oregon. M.S. Thesis, Oregon
R. Meehan, T. R. Merrell, Jr., and J. W. Matthews, editors. Proceedings of a State University, Corvallis, OR.
Symposium on Fish and Wildlife Relationships in Old-Growth Forests, 281. Lingle, G. R. 1977. Food habits and sexing-aging criteria of the white
Held April, 12-15, 1982. Juneau, AK. pelican at Chase Lake National Wildlife Refuge, North Dakota. M.S.
261. Krokhin, E. M. 1967. Effect of the size of sockeye migration on the Thesis, Michigan Technological University.
phosphate regime of spawning lakes. Izvestia, Pacific Scientific Institute 282. Lister, D. B., and H. S. Genoe. 1970. Stream habitat utilization by
of Fisheries and Oceanography 64: 353-364 (in Russian). cohabiting under-yearlings of chinook (Oncorhynchus tshawytscha) and
262. ———. 1975. Transport of nutrients by salmon migrating from the coho (Oncorhynchus kisutch) salmon in the Big Qualicum River, British
sea into lakes. Pages 153-156 in: A. D. Hasler, editor. Coupling of Land Columbia. Journal of Fisheries Research Board of Canada 27:1215-
and Water Systems. Springer-Verlag. New York, NY. 1224.
263. La Tourrette, J. 1992. Washington Wildlife Viewing Guide. Falcon 283. Loch, J. J., S. A. Leider, M. W. Chilcote, R. Cooper, and T. H. Johnson.
Publishing, Inc. Helena, MT. 1988. Differences in yield, emigration-timing, size, and age structure of
263a. Lagler, K.F. 1943. Food habits and economic relations of the turtles juvenile steelhead from two small western Washington streams.
of Michigan with special reference to fish management. The American California Fish and Game 74:106-118.
Midland Naturalist 29(2):257-312. 284. Lowrance, R., R. Todd, J. Fail Jr., O. Hendriksen Jr., R. Leonard, and L.
264. ———, and J. C. Salyer II. 1945. Influence of availability on the Asmussen. 1984. Riparian forests as nutrient filters in agricultural
feeding habits of the common garter snake. Copeia. 1945:100-107. watersheds. BioScience 34:374-377.
265. Lampman, B. H. 1947. A note on the predaceous habits of water 285. Loegering, J. P. 1997. Abundance, habitat association, and foraging
shrews . 1947. Journal of Mammalogy 28:181. ecology of American dippers and other riparian-associated wildlife in
266. Larkin, G. A., and P. A. Slaney. 1997. Implications of trends in marine- the Oregon Coast Range. Ph.D. Thesis, Oregon State University,
derived nutrient influx to south coastal British Columbia salmonid Corvallis, OR.
production. Fisheries 22 (11):16-24. 286. Landry, M. R., J. R. Postel, W. K. Peterson, and J. Newman. 1989.
267. Leatherwood, S. and R. R. Reeves. 1983. The Sierra Club handbook of Broad-scale distributional patterns of hydrographic variables on the
whales and dolphins. Tien Wah Press, Singapore, Malaysia. Washington/Oregon shelf. Pages 1-40 In: M. R. Landry and B. M. Hickey,
268. LeBrasseur, R. J. 1966. Stomach contents of salmon and steelhead editors. Coastal Oceanography of Washington and Oregon. Elsevier
trout in the northeastern Pacific Ocean. Journal of Fisheries Research Oceanography Series 47, Amsterdam, Netherlands.
Board of Canada 23:85-100. 287. MacCarter, D. L. 1972. Food habits of ospreys at Flathead Lake,
269. Lee, K. N. 1993. Compass and gyroscope. Integrating science and Montana. M. S. Thesis. California State University. Humboldt, Arcata, CA.
politics for the environment. Island Press. Washington, D.C. 288. Macdonald, J. S. C. D. Levings C. D. McAllister U. H. M. Fagerlund and
270. Lee, L. C., T. A. Muir, and R. R. Johnson. 1987. Riparian ecosystems as J. R. McBride. 1988. A Field Experiment to Test the Importance of
essential habitat for raptors in the American West. Pages 15-26 in: Estuaries for Chinook Salmon (Oncorhynchus tshawytscha) Survival:
Proceedings of the Western Raptor Management Symposium and Short-Term Results. Canadian Journal Fisheries and Aquatic Sciences
Workshop. National Wildlife Fed. Washington, D.C. 45:1366-1377.
271. Leider, S. A., M. W. Chilcote, and J. J. Loch. 1986. Movement and 289. ———, I. K. Birtwell, and G. M. Kruzynski. 1987. Food and habitat
survival of presmolt steelhead in a tributary and the main stem of a utilization by juvenile salmonids in the Campbell River estuary.
Washington river. North American Journal of Fisheries Management Canadian Journal of Fisheries and Aquatic Sciences 44:1233-1246.
6:526-531. 290. Mace, P. M. 1983. Bird Predation on Juvenile Salmonids in the Big
272. Leonard, W. P. H. A. Brown L. L. C. Jones K. R. McAllister and R. M. Qualicum Estuary, Vancouver Island. Can. Technical Report, Fisheries
Storm. 1993. Amphibians of Washington and Oregon. Seattle Audubon Aquatic Science 1176:1-77.
Society, Seattle, WA. 291. Marcot, B. G., M. A. Castellano, J. A. Christy, L. K. Croft, J. F. Lehmkuhl,
273. Levy, D. A., and T. G. Northcote. 1982. Juvenile salmon residency in a R. . Naney, R. E. Rosentreter, R.E. Sandquist, and E. Zieroth. 1997.
marsh area of the Fraser River estuary. Canadian Journal of Fisheries Terrestrial ecology assessment. Pages 1497-1713 in: T. M. Quigley, S. J.
and Aquatic Sciences 39: 270-276. Arbelbide, and S. F. McCool, editors. An assessment of ecosystem
274. Lichatowich, J. A., and L. E. Mobrand. 1996. Chinook salmon components in the interior Columbia Basin and portions of the
(Oncorhynchus tshawytscha) in the Columbia River:The components of Klamath and Great Basins, USDA Forest Service General Technical
decline. In: Applied Ecosystem Analysis—Background—Mobrand Report PNW-GTR-405. Pacific Northwest Research Station, Portland,
Biometrics, Inc., Prepared for US Department of Energy Bonniville OR.
Power Administration, Environmental Fish and Wildlife. Portland, OR. 292. ———, and M. Vander Heyden. 2001. Key ecological functions of
275. ———. 1999. Salmon without rivers—A history of the Pacific wildlife species. Pages 168-186 in: D. H. Johnson and T. A. O’Neil,
salmon crisis. Island Press, Covelo, CA. managing directors. Wildlife—habitat relationships in Oregon and
276. ———. 1998. Habitat alteration and changes in habitat of coho Washington. Oregon State University Press, Corvallis, OR.
(Oncorhynchus kisutch) and chinook (O. tshawytscha) in Oregon’s 293. Marquiss, M. and K. Duncan. 1993. Variation in the abundance of red-
coastal streams. Pages 92-99 in: C. C. Levings, L. B. Holtby, and M. A. breasted Mergansers Mergus serrator on a Scottish river related to
Henderson, editors. Proceedings of the National Workshop on Effects season, year, river hydrography, salmon density and spring culling. Ibis.
of Habitat Alteration on Salmonid Stocks. Canadian Special Publication 135:33-41.
Fisheries and Aquatic Science 105. 294. Martin, D. J. 1985. Production of cutthroat trout (Salmo clarki) in
277. ———. 1997. Evaluating salmon management institutions: The relation to riparian vegetation in Bear Creek, Washington. Ph. D.
importance of performance measures, temporal scales, and production Dissertation. University of Washington, School of Fisheries, Seattle, WA.
cycles. Pages 69 to 90 in: D. J. Stouder, P. A. Bisson, and R. J. Naiman,
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 667

295. Maser, C., R. F. Tarrant, J. M. Trappe, and J. F. Franklin. 1988. From the 319. ———. 1998. Pacific salmon spawner escapement goals for the
forest to the sea: A story of fallen trees. USDA- Forest Service, and Skagit River watershed as determined by nutrient cycling
USDI- Bureau of Land Management. General Technical Report PNW- considerations. Northwest Science 72: 239-248.
GTR-229. 320. Miegs, R. C. and C. A. Rieck. 1958. Mergansers and trout in
296. ———, and J. R. Sedell. 1994. From the forest to the sea—the Washington. Pages 306-318 in: Proceedings of 47th Annual Conference
ecology of wood in streams, rivers, estuaries, and oceans. St. Lucie West Assoc. State Game Fish Commission.
Press. Delray Beach, FL. 297. Mathisen, O. A. 1972. Biogenic enrichment 321. Miller, L. 1957. Bird remains from an Oregon Indian midden. Condor
of sockeye salmon lakes and stock productivity. Verh. Int.Ver. Limnol. 59:59-63.
18:1089-1095. 322. Miller, R. J., and E. L. Brannon. 1982. The origin and development of
298. Mathisen, O. A., P. L. Parker, J. J. Goering, T. C. Kline, P. H. Poe, and R. S. life history patterns in Pacific salmon. Pages 296 to 309 in: E. L. Brannon,
Scalan. 1988. Recycling of marine elements transported into freshwater and E. O. Salo, editors. Proceedings of the Salmon and Trout Migratory
systems by anadromous salmon.Verh. Int. Ver. Limnol. 23:2249-2258. Behavior Symposium. June 3-5, 1981, First International Symposium,
299. McHenry, M. L., E. Shott, R. H. Conrad, and G. B. Grette. 1998. School of Fisheries, University of Washington, Seattle, WA.
Changes in the quantity and characteristics of large woody debris in 323. Miller, J. A., and C. A. Simenstad. 1997. A comparative assessment of a
streams of the Olympic Peninsula, Washington, U. S. A. (1982-1993). natural and created estuarine slough as rearing habitat for juvenile
Canadian Journal of Fisheries and Aquatic Science 55:1395-1407. chinook and coho salmon. Estuaries 20:792-806.
300. Matkowski, S. M. D. 1989. Differential susceptibility of tree species of 324. Minakawa, N. 1997. The dynamics of aquatic insect communities
stocked trout to bird predation. North American Journal of Fisheries associated with salmon spawning. Ph. D. Dissertation, University of
Management 9:184-187. Washington, School of Fisheries. Seattle, WA.
301. ———. 1984. Angler harvest and other causes of mortality of 325. Minshall, G. W., R. C. Petersen, K. W. Cummins, T. L. Bott, J. R. Sedell,
stocked salmonids in Duck Mountain Provincial Park, Maniotba. M.S. C. E. Cushing, and R. L. Vannote. 1983. Interbiome comparison of
Thesis. University of Manitoba, Winnipeg, Canada. stream ecosystem dynamics. Ecological Monographs 53:1-25.
302. Matthews, D. R. 1983. Feeding ecology of the common murre, Uria 326. ———, and J. N. Minshall. 1977. Microdistribution of benthic
aalge, off the Oregon coast. M. S. Thesis, University of Oregon, Eugene, invertebrates in Rocky Mountain (U.S.A.) Stream. Hydrobiologia
OR. 55:231-249.
303. Mattson, D. J. B. M. Blanchard and R. R. Knight. 1991. Food habits of 327. ———. 1968. Community dynamics of the benthic fauna in a
Yellowstone Grizzly Bears, 1977-1987. Canadian Journal of Zoology woodland springbrook. Hydrobiologia 32:305-337.
69:1619-1629. 328. ———. 1984. Aquatic insect-substratum relationships. Pages 358-
304. McClelland, B. R. 1973. Autumn concentrations of bald eagles in 400 in: V. H. Resh and D. M. Rosenberg, editors. The ecology of aquatic
Glacier National Park. Condor 75:121-123. insects. Praeger Publishers. New York, NY.
305. McMurray, G. R., and R. J. Bailey. 1998. Change in Pacific Northwest 329. Mitsch, W. J., and J. G. Gosselink. 1986. Wetlands. Van Nostrand
Coastal Ecosystems: Science for Solutions, Pacific Northwest Coastal Reinhold, New York, NY.
Ecosystems Regional Study (PNCERS). U. S. Department of Commerce. 330. Mizue, K. K. Yoshida and A. Takemura. 1966. On the ecology of the
NOAA Coastal Ocean Program Decision Analysis Series No. 11. Dall’s porpoise in the Bering Sea and the North Pacific Ocean. Fac.
306. McNeil, W. J., J. R. Gowan, and R. Severson. 1991. Offshore release of Fish., Nagasaki University, Bulletin 21:1-21.
salmon smolts. American Fisheries Society Symposium, 10:548-553. 331. Modde, T., A. F. Wasowicz, and D. K. Hepworth. 1996. Cormorant and
307. Meffe, G. K. 1992. Techno-Arrogance and halfway technologies: grebe predation on rainbow trout stocked in a southern Utah
Salmon hatcheries on the Pacific coast of North America. reservoir. North American Journal of Fisheries Management 16:388-
Conservation Biology 6(3):350-354. 394.
308. Melone, A. M. 1985. Flood producing mechanisms in coastal British 332. Montgomery, D. R., and J. M. Buffington. 1993. Channel classification,
Columbia. Canadian Water Research Journal 10:46-64. prediction of channel response, and assessment of channel condition.
309. Melquist, W. E., and A. E. Dronkert. River Otter. 1987. Wild Furbearer Draft report to the Sediment, Hydrology, and Mass Wasting Committee
Management and Conservation in North America. Pages 626-641 in: M. of the Washington State Timber/Fish/Wildlife Agreement. Department
J. A. Novak, M. E. Obbard, and B. Malloch, editors. Ministry of Natural of Geological Sciences and Quarternary Research Center, University of
Resources, Ontario, Canada. Washington. Seattle, WA.
310. ———, J. S. Whitman, and M. G. Hornocker. 1981. Resource 333. Morejohn, G. V. J. T. Harvey and L. T. Krasnow. 1978. The importance
partitioning and coexistence of sympatric mink and river otter of Loligo opalescens in the food web of marine vertebrates in Monterey
populations. Proceedings of the Worldwide Furbearer Conference Bay, California. California Department of Fish and Game Bulletin No.
1:187-220. 169: 67-98.
311. ———, and M. G. Hornocker. 1983. Ecology of river otters in west 334. Morrison, M.L., B.G. Marcot, and R.W. Mannan. 1998. Wildlife-Habitat
central Idaho. Wildlife Monographs No. 83:1-60. Relationships. Second edition. University of Wisconsin Press, Madison,
312. ———, and D. R. Johnson. 1984. Additional comments on the WI.
migration of northern Idaho and eastern Washington ospreys. Journal 335. Mossman, A. S. 1959. Selective predation of Glaucous-winged gulls
of Field Ornithol. 55:483-485. upon adult red salmon. Ecology 39:482-486.
313. Mendall, H. L. 1944. Food of hawks and owls in Maine. Journal of 335a. ———. 1958. Selective predation of glaucous-winged gulls upon
Wildlife Management. 8:198-208. adult red salmon. Ecology 39(3):482-486.
314. ———. 1939. Food habits of the herring gull in relation to 336. Moyle, P. 1966. Feeding behavior of the Glaucous-winged gull on an
freshwater game fishes in Maine. Wilson Bulletin 41(223-226). Alaskan salmon stream. Wilson Bulletin. 78:175-190.
315. Merritt, R. W., and K. W. Cummins. 1996. An introduction to the 337. Muir, W. D., and R. L. Emmett. 1988. The food habits of migrating
aquatic insects of North America. Kendall/Hunt Publishing Company. salmonid smolts passing Bonneville Dam in the Columbia River, 1984.
Dubuque, IA. Re. Riv. Res. Man. 2:1-10.
316. ———, ———, and T. M. Burton. 1984. The role of aquatic insects in 338. Mundie, J. H. 1969. Ecological implications of the diet of juvenile
the processing and cycling of nutrients. Pages 134-163 In:V. H. Resh and coho in streams. Pages 135-152 in: T. G. Northcote, editor. Symposium
D. M. Rosenberg, editors. The Ecology of Aquatic Insects. Praeger On Salmon and Trout In Streams, H. R. MacMillan Lectures in Fisheries.
Publishers. New York, NY. Held February 22-24, 1968, at the University of British Columbia,
317. Meyer, J. l., and eight co-authors. 1988. Elemental dynamics in Vancouver, BC, Canada.
streams. Journal of North American Benthological Society 7:410-432. 339. Munn, M. D., R. W. Black, A. L. Haggland, M. A. Hummling, and R. L.
318. Michael, J. H., Jr. 1995. Enhancement effects of spawning pink salmon Huffman. 1999. An assessment of stream habitat and nutrients in the
on stream rearing juvenile coho salmon: Managing one resource to Elwha River Basin: Implications for restoration. U.S. Geological Survey,
benefit another. Northwest Science 69:228-233.
 668 Wildlife–Habitat Relationships in Oregon and Washington

Water-Resources Investigations Report 98-4223. Prepared in North American Bioregion. Ecotrust/Interain Pacific and Island Press.
cooperation with the Lower Elwha Tribe and National Park Service. 361. ———., T. J. Beechie, L. E. Benda, P. A. Bisson, L. H. MacDonald, M. D.
340. Munro, J. 1941. Studies of waterfowl in British Columbia. Greater O’Connor, C. Oliver, P. Olson, and E. A. Steel. 1992. Fundamental
scaup duck, lesser scaup. Canadian Journal Research Section D elements of ecologically healthy watersheds in the Pacific Northwest
Zoological Sci.19:113-138. Coastal Ecoregion. Pages 127-188 in: R. J. Naiman, editor. Watershed
341. Munro, J. A. 1932. Food of the American Merganser (Mergus Management—Balancing Sustainability and Environmental Change.
merganser americanus) in British Columbia. Canadian Field-Naturalist Springer-Verlog. New York, NY.
46:166-168. 362. ———, and R. E. Bilby. 1998. River ecology and management:
341a. ———. 1939. Food of ducks and coots at Swan Lake British Lessons from the Pacific Coastal Ecoregion. Springer-Verlag, New York,
Columbia. Canadian Journal of Research 17:178-186. NY.
342. ———, and W. A. Clemens. 1939. The food and feeding habits of the 363. Nakatani, R. E. 1969. Effects of heated discharge on anadromous fish.
Red-breasted Merganser in British Columbia. Journal of Wildlife In: P. A. Krenkel and F. L. Parker, editors. Biological aspects of thermal
Management 3:46-53. pollution.Vanderbilt University Press. Nashville, TN.
343. ———. 1945. Observations of the loon in the Cariboo Parklands, 364. National Marine Fisheries Service (NMFS). 1997. Investigation of
British Columbia. Auk 62:38-49. scientific information on the impacts of California Sea Lions and Pacific
344. ———. 1923. A preliminary report on the relation of various ducks Harbor Seals on salmonids and on the coastal ecosystems of
and gulls to the propagation of sockeye salmon at Henderson lake, Washington, Oregon, and California. U.S. Dep. Commer., NOAA Tech.
Vancouver Island, BC. Canadian Field Naturalist 37:107-116. Memo. NMFS-NWFSC-28.
345. ———. 1938. Studies of waterfowl in British Columbia: Barrow’s 365. National Research Council (NRC). 1996. Upstream - Salmon and
goldeneye, American goldeneye. Transactions of the Royal Canadian society in the Pacific Northwest. National Academy Press. Washington,
Institute 22:259-318. D.C.
346. ———. 1938a. The northern bald eagle in British Columbia. Wilson 366. National Marine Fisheries Service (NMFS). 1998. Essential Fish
Bulletin 50:38-35. Habitat Advisory Report to the Pacific Fishery Management Council
347. Murie, O. J. 1959. Fauna of the Alutian Islands and Alaska Peninsula. for Pacific salmon species of California, Oregon, and Washington. Draft
North American Fauna. report prepared by the Pacific State Marine Fisheries Commission.
348. Murota, T. 1998a. Material cycle and sustainable economy. Pages 120- Gladstone, OR.
138 In: D. Bell, L. Fawcett, R. Keil, and P. Penz, editors. Political Ecology. 366a. ———. 1991. Final rule for endangered status of Snake River
Routledge, London & New York, NY. sockeye salmon. November 20, 1991. Federal Register Volume 56:
349. ———. 1998b. Nutrient shadow cast by anadromous fishes: 58619.
Perspectives in comparison with marine fishery and guano occurance. 367. Nehlsen, W., J. E. Williams, and J. A. Lichatowich. 1991. Pacific salmon
Paper presented at the Western Division American Fisheries Society at the crossroads: stocks at risk from California, Oregon, Idaho, and
Symposium, held in Anchorage, Alaska (Sept.-Oct., 1998). Washington. Fisheries 16 (2):4-21.
350. ———. 1987. The environmental economics of the water planet 368. Netboy, A. 1980. The Columbia River salmon and steelhead trout:
earth. In: Environmental Economics—The Analysis of a Major Interface. Their fight for survival. University of Washington Press, Seattle, WA.
Pages 185-199 in: G. Pilet and T. Murota, editors. R. Leimgruber, Geneva, 369. Newbold, J. D., J. W. Elwood, R.V. O’Neil, and W.V. Winkle. 1981.
Switzerland. Measuring nutrient spiralling in streams. Canadian Journal of Fisheries
351. ———, and Faculty of Environmental Studies. 1994. Material cycle and Aquatic Sciences 38:860-863.
and sustainable economy: A thermodynamical approach to political 370. Newton, M., B.A. El Hassen, and J. Zavitkovski. 1968. Role of red
ecology. A paper read at The Global Political Ecology Conference alder in western Oregon forest succession. Pages 73-84 in: J. M. Trappe,
(Harold Innis Centenary Celebration). Held March 3-6, 1994.York J. F. Franklin, R. F. Tarrant, and G. M. Hansen, editors. Biology of Alder.
University, Toronto, Canada. USDA Forest Service. Portland, OR.
352. Murphy, M. L., and J. D. Hall. 1981. Varied effects of clear-cut logging 371. Nichol, L. M. and D. M. Shackleton. 1996. Seasonal movements and
on predators and their habitat in small streams of the Cascade foraging behavior of northern resident killer whales (Orcinus orca) in
Mountains, Oregon. Canadian Journal of Fisheries and Aquatic Sciences relation to the inshore distribution of salmon (Oncorhynchus spp.) in
38:137-145. British Columbia. Canadian Journal of Zoology 74:983-991.
353. Myers, G. L. and J. J. Peterka. 1976. Survival and growth of Rainbow 372. Nicholas, J. W., and D. G. Hankin. 1988. Chinook salmon populations
trout (Salmo gairdneri) in four prairie lakes, North Dakota. Journal of in Oregon coastal river basins: Description of life histories and
the Fisheries Research Board of Canada 33:1192-1195. assessment of recent trends in run strengths. Oregon Department of
354. Myers, K. W. 1980. An investigation of the utilization of four study Fish and Wildlife Fisheries Div. Info. Report, No. 88-1.
areas in Yaquina Bay, Oregon, by hatchery and wild juvenile salmonids. 373. Nickelson, T. E. 1986. Influences of upwelling, ocean temperatures,
M. S. Thesis. Oregon State University, Corvallis, OR. and smolt abundance on marine survival of coho salmon
355. ———, and H. F. Horton. 1982. Temporal use of an Oregon estuary (Oncorhynchus kisutch) in the Oregon Production Area. Canadian
by hatchery and wild juvenile salmon. Pages 377-392 in:V. S. Kennedy, Journal of Fisheries and Aquatic Sciences 43:527-535.
editor. Estuarine comparisons. Academic Press, New York, NY. 374. Nickelson, T., J. Rodgers, S. Johnson, and M. Solazzi. 1992. Seasonal
356. Myers, J. M., R. G. Kope, G. J. Bryant, D. Teel, L. J. Lierheimer, T. C. changes in habitat use by juvenile coho salmon (Oncorhynchus kisutch) in
Wainwright, W. S. Grant, F. W. Waknitz, K. Neeley, S. T. Lindley, and R. S. Oregon Coastal Streams. Canadian Journal of Fisheries and Aquatic
Waples. 1998. Status review of chinook salmon from Washington, Idaho, Sciences 49:783-789.
Oregon, and California. U.S. Dept. of Commerce, NOAA Tech. Memo. 375. Nicola, S. J. 1966. The relationship of Alloperla nymphs to incubating
NMFS-NWFSC-35. pink (Oncorhynchus gorbuscha) and chum (Oncorhynchus keta) salmon
357. Myers, J. P. 1980. Territoriality and flocking by Buff-breasted eggs, alevins, and pre-emergent fry in a southeastern Alaska stream. M.
Sandpipers: variations in non-breeding dispersion. Condor 82:241-250. S. Thesis. University of Washington, School of Fisheries, Seattle, WA.
358. Nagorsen, D. W. Morrison K. F. and Forsberg J. E. 1989. Winter diet of 376. ——— 1968. Scavenging by Alloperla (Plecoptera: Chloroperlidae)
Vancouver Island marten (Martes americana). Canadian Journal of nymphs on dead pink (Oncorhynchus gorbuscha) and chum
Zoology 67:1394-1400. (Oncorhynchus keta) salmon embryos. Canadian Journal of Zoology
359. ———, Campbell W. R. and Giannico G. R. 1991. Winter Food 46:787-796.
Habits of Marten, Martes americana, on the Queen Charlotte Islands. 377. Nikol’skii, G. V. 1952. The type of dynamics of stocks and the
Canadian Field-Naturalist 105:55-59. character of spawning of the chum (Oncorhynchus keta,Walb.) and the
360. Naiman, R. J., and E. C. Anderson. 1997. Streams and rivers: their pink salmon (Oncorhynchus gorbuscha,Walb.) in the Amur River. Doklady
physical and biological variability. Pages 131-148 in: P. Schoomaker, B. Akademii Nauk. SSSR, 86(4): 873-875.
von Hagen, and E. Wolf, editors. The Rain Forests of Home: Profile of a
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 669

378. Norris, K. S., and J. H. Prescott. 1961. Observations on Pacific 396. ———, ———, J. M. Shenker, W. W. Smoker, and Y. Endo. 1988. Food
cetaceans of California and Mexican waters. University of California. habits of Pacific salmon and steelhead trout, midwater trawl catches
Publ. Zoo. 63:291-403. and oceanographic conditions in the Gulf of Alaska, 1980-1985. Bull.
378a North, M. R. 1994. Yellow-billed loon, Gavia adamsii. The Birds of Ocean Res. Instit., Univ. of Tokyo 26(2):29-78.
North America No.121. In: A. Poole and F. Gill, editors. The Academy of 397. ———. 1992. Ocean ecology of north Pacific salmonids. Washington
Natural Sciences, Philadelphia, PA, and The American Ornithologist’s Sea Grant Program, distributed by University of Washington Press.
Union, Washington, D.C. Seattle, WA.
379.Northwest Power Planning Council (NPPC). 1986. Compilation of 398. Peterson, N. P. 1980. The role of spring ponds in the winter ecology
information on salmon and steelhead losses in the Columbia River and natural production of coho salmon (Oncorhynchus kisutch) on the
basin. Appendix D of the 1987 Columbia River Basin Fish and Wildlife Olympic Peninsula, Washington. M. S.Thesis, University of Washington,
Program. Portland, OR. College of Fisheries. Seattle, WA.
380. O’Connell, M. A., J. G. Hallett, and S. D. West. 1993. Wildlife use of 399. ———. 1982a. Population characteristics of juvenile coho salmon
riparian habitats: A Literature review. Washington Department of (Oncorhynchus kisutch) overwintering in riverine ponds. Canadian
Natural Resources, Timber/Fish/Wildlife, Report TFW-WL1-93-001. Journal of Fisheries and Aquatic Science 39:1303-1307.
381. Oakley, A. L., J. A. Collins, L. B. Everson, D. A. Heller, J. C. Howerton, 400. ———. 1982b. Immigration of juvenile coho salmon (Oncorhynchus
and R. E. Vincent. 1985. Riparian zones and freshwater wetlands. Pages kisutch) into riverine ponds. Canadian Journal of Fisheries and Aquatic
58-76 in: R. E. Brown, editor. Management of Wildlife and Fish Habitats Science 39:1308-1310.
in Forests of Western Oregon and Washington. U. S. For. Serv. Portland, 401. ———, and L. M. Reid. 1984. Wall-base channels: their evolution,
OR. distribution, and use by juvenile coho salmon in the Clearwater River,
382. Obermayer, K. E. Hodgson A. and M. F. Willson. 1999. American Washington. Pages 215-226 in: J. M. Walton and D. B. Houston, editors.
Dipper, Cinclus mexicanus, foraging on Pacific salmon, Oncorhynchus spp., Proceedings of the Olympic Wild Fish Conference, March 23-25, 1983,
eggs. Canadian Field-Naturalist 113:288-290. Port Angeles, WA.
383. Ofelt, C. H. 1975. Food habits of nesting bald eagles in southeast 402. Phinney, L. A., and P. Bucknell. 1975. A catalog of Washington streams
Alaska. Condor 77:337-338. and salmon utilization. Washington Department of Fisheries, Olympia,
384. Olesiuk, P. F. 1993. Annual prey consumption by harbor seal (Phoca WA.
vitulina) in the Strait of Georgia, British Columbia. Fish. Bull. 91:491-515. 403. Pike, G. C. 1950. Stomach contents of whales caught of the coast of
384a. ———, M.A. Bigg, G.M. Ellis, S. J. Crockford, and R.J. Wigen. 1990. An British Columbia. Progress Report of the Pacific Coast Stations.
assessment of the feeding habits of harbour seals (Phoca vitulina) in the Fisheries Research Board Can. 83:27-28.
Strait of Georgia, British Columbia, based on scat analysis. Canadian 404. Piorkowski, R. J. 1995. Ecological effects of spawning salmon several
Technical Report of Fisheries and Aquatic Sciences. No. 1730. south-central Alaskan streams. Ph. D., University of Alaska, Fairbanks.
Department of Fisheries and Oceans. Biological Sciences Branch. Pacific 405. Poage, N. J., and T. A. Spies. 1996. A tale of two unmanaged riparian
Biological Station. Nanaimo, British Columbia. forests. COPE Report 9(1):6-9.
385. Olson, R. and W. Hubert. 1994. Beaver: Water resources and Riparian 406. Poelker, R.J., and H.D. Hartwell. 1973. Black bear of Washington.
Habitat Manager. University of Wyoming. Laramie, WY. Washington State Game Department. Biological Bulletin No. 14.
386. Ordal, E., and R. E. Pacha. 1963. The effects of temperatures on Olympia, WA.
disease in fish. in Water temperature: Influences, effects and control. 407. Pitcher, K.W. 1981. Prey of the Steller sea lion, Eumetopias jubatus, in
Proceedings of the twelfth Pacific Northwest symposium on water the Gulf of Alaska. U. S. Fish and Wildlife Service Fishery Bulletin
pollution research. U.S. Public Health Service, Pacific Northwest Water 79:467-472.
Laboratory. Corvallis, OR. 410. ———. 1977. Population Productivity and Food Habits of Harbor
387. Oregon Coastal Salmon Restoration Initiative (OCSRI). 1997. Seals in the Prince William Sound, Copper River Delta Area, Alaska.
OCSRI Conservation Plan. Draft Revision February 24, 1997. Salem, Report to U.S. Marine Mammal Commission for Contract MM5AC01.
OR. 411. ———, and D. G. Calkins. 1981. Reproductive biology of Steller sea
388. Pacific Fishery Management Council (PFMC). 1998. Review of 1997 lions in the Gulf of Alaska. Journal of Mammalogy 62:599-605.
ocean salmon fisheries. A Report of the Pacific Fishery Management 412. Pomery, L. R. 1970. The strategy of mineral cycling. Annual Review
Council pursuant to National Oceanic and Atmospheric Ecol. Syst. 1:171-190.
Administration, Award No. NA87FC0008. Portland, OR. 413. Pritchard, D. W. 1967. What is an estuary: Physical viewpoint. Pages
388a. ———. 1991. Review of 1990 ocean salmon fisheries. Portland, OR 3-5 in: G. H. Lauff, editor. Estuaries. American Association of the
388b. ———. 1992. Review of 1990 ocean salmon fisheries. 2130 SW Advancement of Science Publication No. 83, Washington, D.C.
Fifth Avenue, Suite 224, Portland, OR. 414. Quinn, T. P., and N. P. Peterson. 1994. The effects of forestry practices
389. Palmer. 1962. Handbook of North American Birds 1. Yale University on fish populations: Incubation Environment of Chum Salmon
Press. New Haven, CT. (Oncorhynchus keta) in Kennedy Creek - Part A. Persistence of Egg
390. Palmisano, J. F., R.H. Ellis and V. W. Kaczynski. 1993. The impact of Pocket Architecture in Chum Salmon Redds. Washington Department
environmental and management factors on Washington’s wild of Natural Resources, Olympia, WA. Timber/Fish/Wildlife Report. TFW-
anadromous salmon and trout. Washington Forest Protection FA-94-001.
Association and State of Washington Department of Natural 415. Rader, R. B. 1997. A functional classification of the drift: traits that
Resources. Olympia, WA. influence invertebrate availability to salmonids. Canadian Journal
391. Parker, M. S. 1994. Feeding ecology of stream-dwelling Pacific giant Fisheries Aquatic Sciences 54:1211-1234.
salamander larvae (Dicamptodon tenebrosus). Copeia 3:705-718. 416. Ralph, S. C., G. C. Poole, L. L. Conquest, and R. J. Naiman. 1994.
392. ———. 1991. Relationship between cover availability and larval Stream channel morphology and woody debris in logged and unlogged
Pacific giant salamander density. Journal of Herpetology 25:355-357. basins of western Washington. Canadian Journal of Fisheries and
393. ——— 1993. Predation by Pacific Giant Salamander larvae on Aquatic Sciences 51:37-51.
juvenile steelhead trout. Northwestern Naturalist 74:77-81. 417. Reeves, G. H., P. A. Bisson, J. M. Dambacher. 1998. Fish Communities.
394. Pearcy, W. G. 1997. What have we learned in the last decade? What Pages 200-234 in: R. J. Naiman and R. E. Bilby, editors. River Ecology and
are research priorities? Pages 271-277 in: R. L. Emmett and M. H. Management: Lessons From the Pacific Coastal Ecoregion. Springer-
Schiewe, editors. Estuarine and Ocean Survival of Northeastern Pacific Verlag. New York, NY.
Salmon, NOAA Technical Memorandum NMFS-NWFSC-29. 418. ———. F. H. Everest, and J.D. Hall. 1987. Interactions between the
395. ———, R. D. Brodeur, and J. P. Fisher. 1990. Distribution and ecology redside shiner (Richardsonius balteatus) and the steelhead trout (Salmo
of juvenile cutthroat trout (Oncorhynchus clarki clarki) and steelhead gairdneri) in western Oregon: the influence of water temperature.
(Oncorhynchus mykiss) in the ocean off Oregon and Washington. Canadian Journal Fisheries and Aquatic Science 44:1602-1613.
Fisheries Bulletin 88:697-711.
 670 Wildlife–Habitat Relationships in Oregon and Washington

419. Regier, H. A. 1997. Old traditions that led to abuses of salmon and 441. Salyer, J. C. and K. F. Lagler. 1940. The food and habits of the
their ecosystems. Pages 17-28 In: D. J. Stouder, P. A. Bisson, and R. J. American merganser during winter in Michigan, considered in relation
Naiman, editors. Pacific Salmon and Their Ecosystems: Status and to fish management. Journal of Wildlife Management 4:186-219.
Future Options. Chapman & Hall. New York, NY. 442. Sandercock, F. K. 1991. Life history of coho salmon (Oncorhynchus
420. Reice, S. R. 1994. Nonequilibrium determinants of biological kisutch). Pages 397 to 445 in: C. Groot and L. Margolis, editors. Pacific
community structure. Amer. Sci. 82:424-435. Salmon Life Histories, University of British Columbia Press.Vancouver,
421. Reid, L. M. 1981. Sediment production from gravel-surfaced forest BC, Canada.
roads, Clearwater Basin, Washington. M. S. Thesis, University of 442a. Sanger, G. A. 1983. Diets and Food Web Relationships of Seabirds in
Washington Department of Geological Sciences. Seattle, WA. the Gulf of Alaska and Adjacent Marine Regions. Pages 631-771 in: Final
422. Reimchen, T. E. 1994. Further studies of predator and scavenger use Report Outer Continental Shelf Environmental Assessment Program,
of chum salmon in stream and estuarine habitats at Bag Harbour, Gwaii US Fish and Wildlife Service Denver Wildlife Research Center
Haanas. Island Ecological Research, Queen Charlotte City, B.C., Migratory Bird Project.
Prepared for Canadian Parks Service. 443. Schlorff, R. W. 1978. Predatory ecology of the Great Egret at
423. Riemer, S. D. and R. F. Brown. 1997. Prey of pinnipeds at selected Humboldt Bay, California. Pages 347-353 in: A. J. Sprunt, J. C. Ogden, and
sites in Oregon identified by scat (fecal) analysis, 1983-1996. Oregon S. Winkler, editors. Wading Birds. National Audubon Society Research
Department of Fish and Wildlife, Wildlife Diversity Program, Technical Report No. 7.
Report No. 97-6-02. 444. Scheffer,V. B. and J. W. Slipp. 1944. The harbor seal in Washington
424. Reinhart, D. P. and D. J. Mattson. 1989. Bear use of cutthroat trout State. American Naturalist 32:374-416.
spawning streams in Yellowstone National Park. International 444a. ———, and ———. 1948. The whales and dolphins of Washington
Conference of Bear Research and Management 8:343-350. State with a key to the cetaceans of the west coast of North America.
425. Richey, J. E., M. A. Perkins, and C. R. Goldman. 1975. Effects of American Midland Naturalist 39:257-337.
kokanee salmon (Oncorhynchus nerka) decomposition on the ecology of 445. ———. 1950. The food of the Alaska Fur Seal. United States Dept. of
a subalpine stream. Journal of Fisheries Research Board of Canada the Interior, Fish and Wildlife Service. Wildlife Leaflet No. 329.
32:817-820. Washington D.C.
426. Ricker, W. 1980. Causes of the decrease in age and size of chinook 446. Scheffer, T. H. and C. C. Sperry. 1931. Food habits of the Pacific
salmon (Oncorhynchus tshawytscha). Canadian Technical Report Aquatic Harbor Seal, Phoca richardii. Journal of Mammalogy 12:214-226.
Science 944. 447. Schwartz, J. E. II and G. E. Mitchell. 1945. The Roosevelt elk on the
427. Robertson, I. 1973. Predation by fish-eating birds on stocks of Pacific Olympic Peninsula, Washington. Journal of Wildlife Management 9:295-
herring, Clupea pallasi, in the Gulf Islands of British Columbia. 319.
Unpublished Report, Pacific Biol. Stn., Nanaimo, BC, Canada. 448. Scott, J. M. 1973. Resource allocation in four synoptic species of
428. ——— 1974. The food of nesting Double-crested and Pelagic marine diving birds. Ph.D. Dissertation, Oregon State University,
Cormorants at Mandarte Island, British Columbia, with notes on Corvallis, OR.
feeding ecology. Condor 76:346-348. 449. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada.
429. Roby, D. D., D. P. Craig, K. Collis, and S. L. Adamany. 1998. Avian Fisheries Research Board Can., Bulletin 184.
predation on juvenile salmonids in the lower Columbia River—1997 450. Schuldt, and Hershey. 1995. Effect of salmon carcass decomposition
Annual Report. Oregon Coop. Wildl. Res. Unit, Oregon State University, on Lake Superior tributary streams. Journal of North American
Corvallis, OR. Benthological Society 14 (2) 259-268.
430. ———, ———, ———, and ———. 1998. (Unpublished) Avian 451. Sedell, J. R., F. J. Triska, and N. S. Triska. 1975. The processing of conifer
predators on juvenile salmonids in the lower Columbia River. Annual and hardwood leaves in two coniferous forest streams. 1. Weight loss
Report to Bonneville Power Administration and U. S. Army Corps of and associated invertebrates.Verh. Int. Ver. Limnol. 19:1617-1627.
Engineers. 452. ———, R. J. Naiman, K. W. Cummins, G. W. Minshall, and R. L.
431. Roche, J., and M. McHutchison. 1998. First fish/first people: Salmon Vannote. 1979. Transport of particulate organic material in streams as a
tales of the North Pacific rim. University of Washington Press. Seattle, function of physical processes. International.Verh. Int.Ver. Limnol.
WA. 20:1366-1375.
432. Roemmich, D., and J. McGowan. 1995. Climatic warming and the 453. ———, and K. J. Luchessa. 1981. Using the historical record as an
decline of zooplankton in the California Current. Science 267:1324- aid to salmonid habitat enhancement. Pages 210 to 223 in: N. B.
1326. Armantrout, editor. Aquisition and Utilization of Aquatic Habitat
433. Roffe, T. J. and B. R. Mate. 1984. Abundances and feeding habits of Information. Proceedings of a Symposium held 28-30 October, 1981.
pinnipeds in the Rogue River. Oregon. Journal of Wildlife Management Portland, OR.
48:1262-1274. 454. Seliskar, D. M., and J. L. Gallagher. 1983. The ecology of tidal marshes
434. Rosgen, D. 1996. Applied river morphology. Wildland Hydrology. of the Pacific Northwest coast: a community profile. USDI—Fish and
Pagosa Springs, CO. Wildlife Service, Division of Biological Services. FWS/OBS-82/32.
435. Ruttner, F. 1971. Fundamentals of limnology. English translation of Washington, D.C.
Third Edition Copyright University of Toronto Press. Toronto, Canada. 455. Senn, H. G. 1958. Merganser depredation. Unpublished Report to
436. Rowlett, R. A. 1980. Observarions of marine birds and mammals in Fisheries Management Division, State of Washington Department of
the northern Chesapeake Bight. U. S. Fish Wildl. Serv., Biol. Serv. Prog., Game, Olympia, WA.
FWS/OBS-80/04. 456. Serdar, C. F. 1999. Description, analysis, and impacts of the Grouse
437. Ruggerone, G. T. 1986. Consumption of migrating juvenile salmonids Creek Landslide, Jefferson County, Washington, 1997-98. Master of
by gulls foraging below a Columbia River dam. Transactions of the Environmental Studies, The Evergreen State College. Olympia, WA.
American Fisheries Society 115:736-742. 457. Servheen, C. W. 1975. Ecology of the wintering Bald Eagle on the
438. Ryall, R., and C. D. Levings. 1987. Juvenile salmon utilization of Skagit River, Washington. M.S.Thesis. University of Washington, Seattle,
rejuvenated tidal channels in the Squamish estuary, British Columbia. WA.
Can. Manuscript Rep. of Fisheries Aquatic Science 1904. 458. Seufert, F. 1980. Wheels of fortune. Oregon Historical Society.
439. Salo, E. O., and W. H. Bayliff. 1958. Artificial and natural production of 459. Shea, D. S. 1973. White-tailed deer eating salmon. Murrelet 54:23.
silver salmon, Oncorhynchus kisutch, at Minter Creek, Washington. 459a. Shetter, D.S., and G.R. Alexander. 1970. Result of predator
State of Washington Department of Fisheries. Res. Bulletin No. 4. reduction on brook trout and brown trout in 4.2 milse (6.76 km) of
440. ———. 1991. Life history of chum salmon. Pages 231-307 in: C. the North Branch of the Au Sable River. Transactions of the American
Groot and L. Margolis, editors. Pacific Salmon Life Histories, University Fisheries Society 2:312-319.
of British Columbia Press. Vancouver, BC, Canada. 460. Shreffler, D. K., and R. M. Thom. 1993. Restoration of Urban Estuaries:
New approaches for site location and design. Prepared for the
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 671

Washington State Department of Natural Resources by Battelle/ 477. ———, and D. Zaldokas. 1997. Fish habitat rehabilitation
Marine Sciences Laboratory. Sequim, WA. procedures. Watershed Restoration Technical Circular No. 9.
461. Shreffler, D. A., C. A. Simenstad, and R. M. Thom. 1992. Temporary Watershed Restoration Program, Ministry of Environment, Lands and
residence of juvenile salmon in a restored estuarine wetland. Canadian Parks, Vancouver, BC, Canada.
Journal of Fisheries and Aquatic Science 47:2079-2084. 478. Smith, C. L. 1979. Salmon fishers of the Columbia. Oregon State
462. Shuldt, J. A., and A. E. Hershey. 1995. Effect of salmon carcass University Press, Corvallis, OR.
decomposition on Lake Superior tributary streams. Journal of Wildlife 479. Smith, M. R., P. W. Mattocks, Jr., M. K. Cassidy. 1997. Breeding birds of
Management 14:1-9. Washington State. In: K. M. Cassidy, C. E. Grue, M. R. Smith, and K. M.
463. Sibatani, A. 1992. Naze sake ha kawa wo sozyousuruka (Why do Dvornich, editors. Vol. 4. Washington State GAP Analysis - Final Report.
salmon go up rivers). In: Tyuuou Kouron/Chuo Koron (Tokyo) 1992(4): Seattle Audubon Society, Publications in Zoology, No. 1. Seattle, WA.
286-295 (In Japanese). 480. Smith, J. L. and D. R. Mudd. 1978. Food of the Caspian Tern in Grays
464. ———. 1996. (English translation by: R. Davis). Why do salmon Harbor, Washington. Murrelet 59:105-106.
ascend rivers? Selected Papers on Entropy Studies 3:3-11. 481. Smith, M. W. 1968. Fertilization and predator control to increase
465. Simenstad, C. A., W. J. Kinney, S. S. Parker, E. O. Salo, J. R. Cordell, and growth rate and yield of trout in a natural lake. Journal of Fisheries
H. Buechner. 1980. Prey community structures and trophic ecology of Research Board of Canada 25:2011-2036.
outmigrating juvenile chum and pink salmon in Hood Canal, 482. Spalding, D. J. 1964. Comparative feeding habits of fur seal, sea lion,
Washington: A synthesis of three years’ studies, 1977-1979. Final Rep. and harbor seal on the British Columbia coast. Fisheries Research
to Washington Department of Fisheries. Fisheries Research Institute, Board of Canada Bull. No. 146.
University of Washington. FRI-UW-8026. Seattle, WA. 483. Spanier, E. 1980. The use of distress calls to repel night herons
466. Simenstad, C. A., K. L. Fresh, and E. O. Salo. 1982. The role of Puget (Nycticorax nycticorax) from fish ponds. Journal of Appl. Ecol. 17:287-294.
Sound and Washington coastal estuaries in the life history of Pacific 484. Speaker, R.W., K. J. Luchessa, J. F. Franklin, and S.V. Gregory. 1988. The
salmon: An unappreciated function. Pages 343-364 In:V. S. Kennedy, use of plastic strips to measure leaf retention by riparian vegetation in
editor. Estuarine Comparisons. Academic Press. New York, NY. a coastal Oregon stream. American Midland Naturalist 120:22-31.
467. ———, and E. O. Salo. 1982. Foraging success as a determinant of 485. Spence, B. C., G. A. Lomnicky, R. M. Hughes and R. P. Novitzki. 1996.
estuarine and nearshore carrying capacity of juvenile chum salmon An ecosystem approach to salmonid conservation. Report prepared by
(Oncorhynchus keta) in Hood Canal,Washington. Pages 21-37 in: B. R. ManTech. TR-4501-96-6057. Corvallis, OR.
Melteff and R. A. Neve, editors. Proceedings of the North Pacific 486. Spencer, C. N., B. R. McClelland, and J. A. Stanford. 1989. Shrimp
Aquaculture Symposium, 18-27 August 1980. Anchorage, Alaska and stocking, salmon collapse and eagle displacement: cascading interactions
Newport, Oregon. Alaska Sea Grant Rep. 82-2. University of Alaska, in the food web of a large aquatic ecosystem. BioScience 41:14-21.
Fairbanks, AK. 487. Stalmaster, Mark V. 1980. Salmon carrion as a winter food source for
468. ———, B. S. Miller, C. F. Nyblade, K. Thornburgh, and L. J. Bledsoe. red-tailed hawks. Murrelet 61(1):43-44.
1979. Food web relationships of northen Puget Sound and the Strait of 488. Stalmaster, M. V. J. R. Newman and A. J. Hansen. 1979. Population
Juan de Fuca. MESA Puget Sound Project, Eviron. Res. Labs. DOC/EPA dynamics of wintering bald eagles on the Nooksack River, Washington.
Interagency Energy /Environment R&D Program Rep., EPA-600/7-79- Northwest Science 53:126-131.
259. 489. Stalmaster, M. V. 1976. Winter ecology and effects of human activity
469. ———, M. Dethier, C. Levings, and D. Hay. 1997b. The Land-Margin on bald eagles in the Nooksack River Valley, Washington. M.S. Thesis.
Interface of Coastal Temperate Rain Forest Ecosytems: Shaping the Western Washington State College, Bellingham, WA.
Nature of Coastal Interactions. Pages 149-187 in: P. Schoonmaker, B. 490. ———, and J. A. Gessaman. 1984. Ecological energetics and foraging
von Hagen, and E. Wolf, editors. The Rain Forests of Home: Profile of a behavior of overwintering bald eagles. Ecological Monographs 54:407-
North American Bioregion. Ecotrust/Interain Pacific and Island Press. 428.
470. ———, J. R. Cordell, W. G. Hood, B. E. Feist, and R. M. Thom. 1997a. 491. Stanford, J. A., and J. V. Ward. 1988. The hyporheic habitat of river
Ecological status of a created estuarine slough in the Chehalis River ecosystems. Nature 335:64-66.
estuary: Assessment of created and natural estuarine sloughs, January- 492. Stanford, J. A., and A. R. Gaufin. 1974. Hyporheic communities of two
December 1995., Fisheries Research Institute, School of Fisheries, FRI- Montana rivers. Science 185:700-702.
UW-9621, University of Washington, Seattle, WA. 493. Steeger, C., H. Esselink, and R. C. Ydenberg. 1992. Comparative
471. ———, J. R. Cordell, J. A. Miller, W. G. Hood, and R. M. Thom. 1993. feeding ecology and reproductive performance of Ospreys in different
Ecological status of a created estuarine slough in the Chehalis River habitats of southeastern British Columbia. Canadian Journal of Zoology
estuary: Assessment of created and natural estuarine sloughs, January- 70:470-475.
December 1992. FRI-UW-9305. Fish. Res. Inst., University of 494. Stenson, G. B., G. A. Badgero and H. D. Fisher. 1984. Food habits of
Washington, Seattle, WA. the River Otter Lutra canadensis in the marine environment of British
472. ———, ———, W. G. Hood, J. A. Miller and R. M. Thom. 1992. Columbia. Canadian Journal of Zoology 62:88-91.
Ecological status of a created estuarine slough in the Chehalis River 495. Stewart, B. S. and S. Leatherwood. 1985. Minke whale. Balaenoptera
estuary: Report of monitoring in created and natural estuarine sloughs, acutorostrata Lacepede, 1804. Pages 91-136 in: S. H. Ridgway, and R.
January-December 1991. Fisheries Research Institute, University of Harrison, editors. Handbook of Marine Mammals. Volume 3. The
Washington, FRI-UW-9206. Seattle, WA. Sirenians and Baleen Whales. Academic Press, London & Orlando, FL.
473. ———, W. G. Hood, R. M. Thom, D. A. Levy and D. L. Bottom. In 496. Stockner, J. G., and E. A. MacIsaac. 1996. British Columbia lake
press. Landscape structure and scale constraints on restoring estuarine enrichment program: two decades of habitat enhancement for sockeye
wetlands for Pacific Coast juvenile fishes. In: M. P. Weinstein and D. A. salmon. Regulated Rivers: Research & Management 12: 547-561.
Kreeger, editors. Concepts and Controversies in Tidal Marsh Ecology, 497. ———, and K. R. S. Shortreed. 1978. Enhancement of autotrophic
Kluwer Academic Publ., Dordrecht, Netherlands. production by nutrient addition in a coastal rainforest stream on
474. Simons, D. D. 1983. Interactions between California Condors and Vancouver Island. Journal of Fisheries Res. Bd. Can. 35:28-34.
humans in prehistoric far western North America. Pages 470-494 in: 498. Stolz, J., and J. Schnell. 1991. Trout—The Wildlife Series. Stackpole
S.R. Wilbur and J. A. Jackson, editors.Vulture Biology and Management. Books. Cameron and Kelker Streets, Harrisberg, PA.
University of California Press, Berkeley, CA. 499. Stouder, D. J., P. A. Bisson, and R. J. Naiman, editors. 1997. Pacific
475. Skagen, S. K. R. L. Knight and G. H. Orians. 1991. Human disturbance salmon and their ecosystems: Status and future options. I. T. P. Chapman
of an avian scavenging guild. Ecological Applications 1:215-225. and Hall International Thomson Publishing. New York, NY.
476. Slaney, P. A., B. O. Rublee, C. J. Perrin, and H. Goldberg. 1994. Debris 500. Stroud, R. K. C. H. Fiscus and H. Kajimura. 1980. Food of the Pacific
structure placements and whole-river fertilization for salmonids in a White-sided Dolphin, Lagenorhynchus obliquidens, Dall’s Porpoise,
large regulated stream in British Columbia. Bulletin of Marine Science Phocoenoides dalli and Northern Fur Seal, Callorhinus ursinus, off
55:1160-1180.
 672 Wildlife–Habitat Relationships in Oregon and Washington

California and Washington. U. S. Fish and Wildlife Service Fisheries 517. Todd, I., and P. Larkin. 1971. Gillnet selectivity on sockeye
Bulletin 78(4):951-959. (Oncorhynchus nerka) and pink salmon (O. gorbuscha) of the Skeena
501. Strub, P. T., C. James, A. C. Thomas, and M. R. Abbott. 1990. Seasonal River system, British Columbia. Journal of Fisheries Research Board
and nonseasonal variability of satelite-derived surface pigment Can. 28:821-842.
concentration in the California Current. J. Geophy. Res. 95(C7):11503- 518. Toweill, D. E. 1974. Winter food habits of River Otters in western
11530. Oregon. Journal of Wildlife Management 38:107-111.
502. Suckley, G. and J. G. Cooper. 1860. The natural history of Washington 519. Townsend, C. R., and M. R. Scarsbrook. 1997. The intermediate
Territory and Oregon. Bailliere Brothers, New York, NY. disturbance hypothesis, refugia, and biodiversity in steams. Limnol.
503. Sugai, S. F., and D. C. Burrell. 1984. Transport of dissolved organic Oceanogr. 42(5):938-949.
carbon, nutrients, and trace metals from the Wilson and Blossom rivers 520. Triska, F. J., J. H. Duff, and R. J. Avanzino. 1990. Influence of exchange
to Smeaton Bay, southeast Alaska. Canadian Journal of Fisheries and flow between channel and hyporheic zone on nitrate production in a
Aquatic Sciences 41:180-190. small mountain stream. Canadian Journal of Fisheries and Aquatic
504. Sullivan, K., J., Tooley, K. Doughty, J. E. Caldwell, and P. Knudsen. 1990. Sciences 47:2099-2111.
Evaluation of prediction models and characterization of stream 521. Trotter, P. C. 1987. Cutthroat—Native Trout of the West. Colorado
temperature regimes in Washington. Washington Department of Associated University Press. Boulder, CO.
Natural Resources.Timber/Fish/Wildlife Rep. No. TFW-WQ3-90-006. 522. Tschaplinski, P. J. 1982. Aspects of the population biology of
Olympia, WA. estuarine-reared and stream-reared juvenile coho salmon in Carnation
505. Sumner, F. H. 1962. Migration and growth of coastal cutthroat trout Creek: A summary of current research. Pages 289-307 in: G. F. Hartman,
in Tillamook County, Oregon. Transactions of American Fisheries editor. Proceedings of the Carnation Creek Workshop: A Ten-year
Society 91:77-83. Review. Malaspina College, Nanaimo, BC, Canada.
506. Swanson, F. J., S. V. Gregory, J. R. Sedell, and A.G. Campbell. 1982. 523. ———. 1987. The use of estuaries as rearing habitats by juvenile
Land-water interactions: The riparian zone. Pages 267-291 In: R. L. coho salmon. Pages 123-141 in: T. W. Chamberlin, editor. Proceedings of
Edmonds, editor. Analysis of coniferous forest ecosystems in the the Workshop: Applying 15 Years of Carnation Creek Results. Pacific
western United States. U.S. Int. Biolog. Prog. Synthesis Series 14, Biol. Station. Nanaimo, BC, Canada.
Hutchinson Ross. Stroudsburg, PA. 524. Tsuchida, A. 1994. Resource physics and the limitations of nuclear
507. ———, L. E. Benda, S. H. Duncan, G. E. Grant, W. F. Megahan, L. M. fusion power generation. Selected Papers On Entropy Studies,Vol. 1.
Reid, and R. R. Ziemer. 1987. Mass failures and other processes of 525. ———. 1996. The importance of the “Cycle of Matter:” A discussion
sediment production in Pacific Northwest forest landscapes. Pages 9-38 of nutrient cycling in basins, land, and human society from the
in: E. O. Salo and T. W. Cundy, editors. Streamside Management: Forestry viewpoint of entropy, pages 150-170. In Toyohashi, Proceedings of Clean
and Fishery Interactions. College of Forest Resources, University of Sea ‘96. International Workshop and Symposium On Environmental
Washington. Institute of Forest Resources, Contribution No. 57. Restoration for Enclosed Seas.
Seattle, WA. 526. United States Army Corps of Engineers (USACOE), U. S.
508. Sweeney, S. J. 1978. Diet, reproduction and population structure of Environmental Protection Agency, U. S. Fish and Wildlife Service, and
the bobcat (Lynx rufus fasciatus) in western Washington. M. S.Thesis. National Marine Fisheries Service. 1993. Commencement Bay
University of Washington, WA. Cumulative Impact Study. U. S. Army Corps of Engineers, Seattle
508a. Sweonson, J.E. 1978. Factors affecting status and reproduction of District. Seattle, WA.
ospreys in Yellowstone National Park. Journal of Wildlife Management 527. Van Daele, L. J. and H. A. Van Daele. 1982. Factors affecting the
43(3):595-601. productivity of ospreys nesting in west-central Idaho. Condor 84:292-
509. Sydeman, W. J. K. A. Hobson P. Pyle and E. B. McLaren. 1997. Trophic 299.
relationships among seabirds in central California: combined stable 528.Vander Heyden, M. 1997. Female black bear habitat selection and
isotope and conventional dietary approach. Condor 99:327-336. home range ecology in the central Cascades of Oregon. M.S. Thesis,
510. Tagart, J. V. 1976. The survival from egg deposition to emergence of Oregon State University, Corvallis, OR.
coho salmon in the Clearwater River, Jefferson County, Washington. 529.Vannote, R. L., G. W. Minshall, K. W. Cummins, J. R. Sedell, and C. E.
M.S. Thesis, University of Washington School of Fisheries, Seattle, WA. Cushing. 1980. The river continuum concept. Canadian Journal of
510a.Tanner,W.W. 1949. Food of the wandering garter snake, Thamnophis Fisheries and Aquatic Sciences 37:130-137.
elegans vagrans (Baird & Girard) in Utah. Herpetologica 5:85-86. 530. Vermeer, K. and K. Devito. 1986. Size, caloric content, and association
511. Taverner, P. A. 1934. Birds of Canada. Canada Dept. Mines, Natl. Mus. of prey fishes in meals of nestling Rhinoceros Auklets. Murrelet 67:1-9.
Bull. No. 72 (Biol. Ser. 19). Ottawa, Canada. 531. ———, and S. J. Westrheim. 1984. Fish changes in diets of nestling
512. Thedinga, J. F., and K. V. Koski. 1984. A stream ecosystem in an old- rhinoceros auklets and their implications. Pages 96-105 in: D. N.
growth forest in southeast Alaska. Part VI: The production of coho Nettleship,, G. A. Sanger, and P. F. Springer, editors. Marine Birds: Their
salmon, Oncorhynchus kisutch, smolts and adults from Porcupine Creek. Feeding Ecology and Commercial Fisheries Relationships. Minister of
Pages 99-108. in: W. R. Meehan, T. R. Merrel, Jr., and T. A. Hanley, editors. Supply and Services, Canada.
Proceedings from a Symposium on Fish and Wildlife Relationhships in 532. ———. 1979. Nesting requirements, food and breeding distribution
Old-growth Forests. Am. Inst. Fish. Res. Biol., Juneau, AK. of rhinoceros auklets, Cerorhinca monocerata, and tufted puffins, Lunda
513. Thom, R. M., and A. B. Borde. 1997. Human intervention in Pacific cirrhata.Ardea 67:101-110.
Northwest coastal ecosystems. Pages 5-37 in: G. R. McMurray and R. J. 533. ———. 1982. Comparison of the diet of the glaucous-winged gull
Bailey, editors. Change in Pacific Northwest Coastal Ecosystems. on the east and west coasts of Vancouver Island. Murrelet 63:80-85.
Science for Solutions, Pacific Northwest Coastal Ecosystems Regional 533a. ———. 1992. The diet of birds as a tool for monitoring the
Study (PNCERS). U.S. Department of Commerce. NOAA Coastal biological environment. Pages 41-50 in K.Vermeer, R.W. Butler, and K.H.
Ocean Program Decision Analysis Series No. 11. Morgan, editors. The ecology, status, and conservation of marine and
514. Thomas, A. C., and P. T. Strub. 1989. Large scale patterns of shoreline birds on the west coast of Vancouver Island. Canadian
phytoplankton pigment distribution during the spring transition along Wildlife Service Occasional Papers, No. 75.
the west coast of North America. J. Geophy. Res. 94(C12):18-117. 534. ———, and K. H. Morgan. 1992. Marine bird populations and habitat
515. Thomson, R. E. 1981. Oceanography of the British Columbia Coast. use in a fjord on the west coast of Vancouver Island. Canadian Wildlife
Can. Spec. Publ. Fisheries and Aquatic Science 56, Dept. Fish. Oceans. Service Occasional Paper No. 75: 86-96.
Sydney, BC, Canada. 535.Verts, B. J. and L. N. Carraway. 1998. Land Mammals of Oregon.
516. Thut, R. N. 1970. Feeding habits of the Dipper in southwestern University of California Press, Berkeley, CA.
Washington. Condor 72:234-235. 536. Wallace, J. B., S. L. Eggert, J. L. Meyer, and J. R. Webster. 1997. Multiple
trophic levels of a forest continuum concept. Canadian Journal of
Fisheries and Aquatic. Sciences 37:130-104.
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 673

537. Watson, J. W. M. G. Garrett and R. G. Anthony. 1991. Foraging ecology 556. Williams, W. R., M. Laramie, and J. J. Ames. 1975. Catalog of
of bald eagles in the Columbia River estuary. Journal of Wildlife Washington Streams and Salmon Utilization—Volume 1 Puget Sound.
Management 55:492-499. Washington Department of Fisheries. Olympia, WA.
538. Warren, C. E. 1971. Biology and water pollution control. The 557. Williams, D. D., and H. B. N. Hynes. 1974. The occurrence of benthos
Department of Fisheries and Wildlife, Oregon State University, deep in the substratum of a stream. Freshwater Biology 4:233-256.
Corvallis, OR. 558. ———, and B. W. Feltmate. 1992. Aquatic insects. CAB International.
538a. Washington Department of Fisheries. 1992. Draft. Goals and United Kingdom.
objectives for stocks and fisheries. Olympia, WA. 559. Willson, M. F., and K. C. Halupka. 1995. Anadromous fish as keystone
538b. ———. 1992. Strategic plan for management of Washington’s species in vertebrate communities. Conservation Biology 9:489-497.
salmon resources. Olympia, WA. 560. Winter, B. D. 1992. Determinate migratory behavior of steelhead
539. Washington Department of Fish and Wildlife and Western (Oncorhynchus mykiss) parr. Ph.D. Dissertation. University of
Washington Treaty Indian Tribes. 1994. 1992 Washington State Salmon Washington, School of Fisheries. Seattle, WA.
And Steelhead Stock Inventory. Appendix One, Puget Sound Stocks. 561. Willson, M. F. , S. M. Gende, B. H. Marston. 1998. Fishes and the
540. Washington Department of Fisheries (WDF) and Oregon Forest: Expanding perspectives on fish-wildlife interactions. BioScience
Department of Fish and Wildlife (ODFW). 1992. Status report. 48:455-462.
Columbia River fish runs & fisheries, 1938-91. Olympia, WA. 562. ———, Halupka K. C. 1995. Anadromous fish as keystone species in
541. ———. 1998. 1998 Washington salmonid stock inventory, coastal vertebrate communities. Conservation Biology 9:489-497.
cutthroat trout appendix. DRAFT. Washington Deptartment of Fish and 563. Wilson, U. W and D. A. Manuwal. 1986. Breeding biology of the
Wildlife, Olympia, WA. rhinoceros auklet in Washington. Condor 88:143-155.
541a. Washington Department of Fish and Wildlife (WDFW). 1998. 564. Wipfli, M. S., J. Hudson, and J. Caouette. 1998. Influence of salmon
Forage fish management plan - A plan for managing the forage fisheries carcasses on stream productivity: response of biofilm and benthic
in Washington. Adopted by the Washington Fish and Wildlife macroinvertebrates in southeastern Alaska, U.S.A. Canadian Journal of
Commission on January 24, 1998. Fisheries and Aquatic Sciences 55:1503-1511.
542. Washington Department of Fisheries, Washington Department of 565. Wissmar, R. C., and C. A. Simenstad. 1988. Energetic constraints of
Wildlife, and Western Washington Treaty Indian Tribes. 1992. 1992 juvenile chum salmon (Oncorhynchus keta) migrating in estuaries.
Washington State Salmon And Steelhead Stock Inventory. Washington Canadian Journal of Fisheries and Aquatic Science. 45:1555-1560.
Department of Fish and Wildlife, Olympia, WA 566. ———, and ———. 1998. Variability of estuarine and riverine
543. Washington Department of Ecology. 1994. Inventory of dams in the ecosystem productivity for supporting Pacific salmon. Pages 253-301 in:
state of Washington. Washington Department of Ecology, Water G. R. McMurray and R. J. Bailey, editors. Change in Pacific Northwest
Resources Program, Dam Safety Division. Revised Edition, January 1994, Coastal Ecosystems. NOAA Coastal Ocean Prog., Decision Analysis
Publication #94-16. Series No. 11, NOAA Coastal Ocean Office. Silver Spring, MD.
544. Washington Department of Fish and Wildlife’s (WDFW). 1997. Wild 567. Wood, C. C. 1986. Dispersion of Common Merganser (Mergus
Salmonid Policy. Washington Department of Fish and Wildlife. Olympia, merganser) breeding pairs in relation to availability of juvenile Pacific
WA. salmon in Vancouver Island streams. Canadian Journal of Zoology
545. Washington Department of Natural Resources (WDNR). 1997. Final 64:756-765.
Habitat Conservation Plan. Washington Department of Natural 568. ———. 1985. Aggregative response of Common Mergansers
Resources, Jennifer Belcher, Commissioner of Public Lands. Olympia, (Mergus merganser): predicting flock size and abundance on Vancouver
WA. Island salmon streams. Canadian Journal of Fisheries and Aquatic
546. Washington Department of Fish and Wildlife and Oregon Science 42:1259-1271.
Department of Fish and Wildlife. 1994. Status Report. Columbia fish 569. ———, and C. M. Hand. 1985. Food searching behavior of the
runs and fisheries 1938-93. Olympia, WA. Common Merganser (Mergus merganser) 1: functional responses to
547. Waters, T. F. 1969. Invertebrate drift - Ecology and significance to prey and predator density. Canadian Journal of Zoology 63:1260-1270.
stream fishes. Pages 121-134 In: T. G. Northcote, editor. H. R. MacMillan 570. ———. 1987a. Predation by the common merganser (Mergus
Lectures In Fisheries. Held Feb. 22-24, 1968. Univ. British Columbia. merganser) on Eastern Vancouver Island. I: Predation during the seaward
Vancouver, BC, Canada. migration. Canadian Journal of Fisheries and Aquatic Sciences
548. Wehle, D. H. S. 1983. The food, feeding, and development of young 44:941-949.
tufted and horned puffins in Alaska. Condor 85:427-442. 571. ———. 1987b. Predation by the common merganser (Mergus
549. Weitkamp, L. A., T. C. Wainwright, G. J. Bryant, G. B. Milner, D. J. Teel, R. merganser) on Eastern Vancouver Island. II: Predation of stream-resident
G. Kope, and R. S. Waples. 1995. Status review of coho salmon from juvenile salmon by merganser broods. Canadian Journal of Fisheries and
Washington, Oregon, and California. U.S. Dept. of Commerce. NMFS- Aquatic Sciences 44:950-959.
NWFSC-24. 572. Wurtsbaugh, W. A., and G. E. Davis. 1977. Effects of temperature and
550. Wendler, H. O., and G. Deschamps. 1955. Logging dams on coastal ration level on the growth and food conversion efficiency of Salmo
Washington streams. Fish. Res. Pap. 1: 27-38, Washington Department gairdneri Richardson. J. Fish. Biol. 11:87-98.
of Fisheries, Olympia, WA. 573. Wydoski, R. S., and R. R. Whitney. 1979. Inland fishes of Washington.
551. White, H. C. 1936. The food of kingfishers and mergansers on the University of Washington Press, Seattle, WA.
Margaree River, Nova Scotia. J. Biol. Bd. Can. 2:299-309. 574.Yoakum, J. 1964. Observations on bobcat—water relationships.
552. ———. 1939. Bird control to increase the Margaree River salmon. Journal of Mammalogy 45:477-479.
Bull. Fisheries Research Board. Can. 58. 575.Young, S. P. 1944. The Wolves of North America. Their history, life
553. ———. 1957. Food and natural history of mergansers on salmon habits, economic status, and Control. The American Wildlife Institute,
waters in the Maritime provinces of Canada. Fisheries Research Board Washington, D.C.
Can., Bulletin No. 116. 576. Zarnowitz, J. E., and K. J. Raedeke. 1984. Winter predation on coho
554. White, R. 1992. Land use, environment, and social change: The fingerlings by birds and mammals in relation to pond characteristics.
shaping of Island County, Washington. University of Washington Press. College of Forest Resources, University of Washington. Service
Seattle,WA. Contract No. 1480. Seattle, WA.
555.Whitman, J. S. 1981. Ecology of the mink (Mustela vison) in west-
central Idaho. M.S. Thesis, University of Idaho.
555a. Wilke, F.A., and K.W. Kenyon. 1957. The food of fur seals in the
eastern Bering Sea. Journal of Wildlife Management 21:237-238.
 674 Wildlife–Habitat Relationships in Oregon and Washington

Appendix I. The nine wildlife species identified as having (or historically had) a strong, consistent relationship
with salmon in Oregon and Washington.

Incubation FreshwaterRearing Saltwater Spawning Carcass

Common Merganser x x x
Harlequin Duck x x
Osprey x x x
Bald Eagle x x x
Caspian Tern x x
Black Bear x x
Grizzly Bear x x
Northern River Otter x x x
Killer Whale x
An “x” identifies the life stage(s) of salmon applicable to the species.

Appendix II. The fifty-eight wildlife species identified as having (or historically had) a recurrent relationship
with salmon in Oregon and Washington.

Incubation Freshwater Rearing Saltwater Spawning Carcass

Cope’s Giant Salamander x x

Pacific Giant Salamander x x
Pacific Coast Aquatic x x
Garter Snake
Red-throated Loonx x x
Pacific Loon x
Common Loon x x
Pied-billed Grebe x
Western Grebe x x
Clark’s Grebe x
American White Pelican x
Brandt’s Cormorant x x
Double-crested Cormorant x x
Pelagic Cormorant x x
Great Blue Heron x x
Black-crowned Night-heron x x
Turkey Vulture x
California Condor h
Common Goldeneye x x x
Barrow’s Goldeneye x x x
Common Merganser x
Red-breasted Merganser x x x
Golden Eagle x x
Bonaparte’s Gull x x x
Heermann’s Gull x
Ring-billed Gull x x x
California Gull x x
Herring Gull x x x
Thayer’s Gull x
Western Gull x x
Glaucous-winged Gull x x x x
Glaucous Gull x x
Common Tern x x
Arctic Tern x x
Forster’s Tern x x
Elegant Tern x
Common Murre x
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 675

Incubation Freshwater Rearing Saltwater Spawning Carcass

Marbled Murrelet x x
Rhinoceros Auklet x
Tufted Puffin x
Belted Kingfisher x x x
American Dipper x x x
Steller’s Jay x
Black-billed Magpie x x
American Crow x x
Northwestern Crow x x x
Common Raven x x x
Virginia Opossum x
Water Shrew x x x
Coyote x
Gray Wolf x x
Raccoon x x
Mink x x x
Bobcat x x
Northern Fur Seal x
Northern (Steller) Sea Lion x x x
California Sea Lion x x
Harbor Seal x x x
Pacific White-sided Dolphin x
An “x” identifies the life stage(s) of salmon applicable to the species currently, “h” represents a historical relationship with salmon.

Appendix III. The twenty-five wildlife species identified as having an indirect relationship with salmon in
Oregon and Washington.

Incubation Freshwater Rearing Saltwater Spawning Carcass

Harlequin Duck xa
Bald Eagle xb xb xb xb
Gyrfalcon xb xb xb
Peregrine Falcon xb xb xb
Killdeer xa
Spotted Sandpiper xa
Snowy Owl xb
Willow Flycatcher xa
Tree Swallow xa
Violet-green Swallow xa
Northern Rough-winged xa
Swallow
Bank Swallow xa
Cliff Swallow xa
Barn Swallow xa
American Dipper xa
Masked Shrew xa
Vagrant Shrew xa
Montane Shrew xa
Fog Shrew xa
Pacific Shrew xa
Water Shrew xa
Pacific Water Shrew xa
Trowbridge’s Shrew xa
Harbor Porpoise xb
Dall’s Porpoise xb
An “x” identifies the life stage(s) of salmon applicable to the species. Species have indirect relationship with salmon either through
consuming insects associated with carcasses (a), or through feeding on gulls, terns, or waterfowl that eat live salmon or salmon
carcasses (b).
 676 Wildlife–Habitat Relationships in Oregon and Washington

Appendix IV. The sixty-four wildlife species identified as having (or historically had) a rare relationship with
salmon in Oregon and Washington.

Incubtion Freshwater Rearing Saltwater Spawning Carcass

Snapping Turtle x
Western Pond Turtle x
Western Terrestrial Garter x
Snake
Common Garter Snake x
Pacific Loon x
Common Loon x
Yellow-billed Loon x
Horned Grebe x x
Red-necked Grebe x x
Western Grebe x
Sooty Shearwater x
Brown Pelican x
Great Egret x x
Snowy Egret x
Green Heron x x
Trumpeter Swan x x x
Mallard x x
Green-winged Teal x
Canvasback x
Greater Scaup x x
Surf Scoter x x
White-winged Scoter x x
Common Goldeneye x x
Barrow’s Goldeneye x
Hooded Merganser x x x
Red-tailed Hawk x
Greater Yellowlegs x
Franklin’s Gull x
Mew Gull x
Black-legged Kittiwake x x
Pigeon Guillemot x
Ancient Murrelet x
Gray Jay x
Winter Wren x
American Robin x
Varied Thrush x x
Spotted Towhee x
Song Sparrow x
Masked Shrew x
Vagrant Shrew x
Montane Shrew x
Fog Shrew x
Pacific Shrew x
Pacific Water Shrew x
Torwbridge’s Shrew x
Douglas’ Squirrel x
Northern Flying Squirrel x
Deer Mouse x
Red Fox x
Gray Fox x
Ringtail x
American Marten x
Fisher x
Long-tailed Weasel x
Wolverine x
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 677

Incubtion Freshwater Rearing Saltwater Spawning Carcass

Striped Skunk x
Mountain Lion x x
White-tailed Deer x
Minke Whale x
Sperm Whale x
Humpback Whale x
Northern Right-whale x
Dolphin
Dall’s Porpoise x
Harbor Porpoise x
An “x” identifies the life stage(s) of salmon applicable to the species.

Appendix V. The sixty wildlife species identified as having an unknown relationship

with salmon in Oregon and Washington.

Northwestern Salamander Fringed Myotis

Long-toed Salamander Keen’s Myotis
Olympic Torrent Salamander Long-eared Myotis
Columbia Torrent Salamander Silver-haired Bat
Southern Torrent Salamander Western Pipistrelle
Cascade Torrent Salamander Big Brown Bat
Rough-skinned Newt Hoary Bat
Dunn’s Salamander Spotted Bat
Van Dyke’s Salamander Townsend’s Big-eared Bat
Tailed Frog Pallid Bat
Western Toad Brazilian Free-tailed Bat
Woodhouse’s Toad Townsend’s Chipmunk
Red-legged Frog Columbian Mouse
Oregon Spotted Frog Townsend’s Vole
Foothill Yellow-legged Frog Water Vole
Northern Leopard Frog Ermine
Bullfrog Western Spotted Skunk
Painted Turtle
Red-eared Slider Turtle
Western Sandpiper
Dunlin
Short-billed Dowitcher
Least Flycatcher
Hammond’s Flycatcher
Dusky Flycatcher
Pacific-slope Flycatcher
Cordilleran Flycatcher
Black Phoebe
Warbling Vireo
Purple Martin
Gray Catbird
European Starling
Brewer’s Blackbird
Northern Waterthrush
Baird’s Shrew
Merriam’s Shrew
Pygmy Shrew
Shrew-mole
California Myotis
Western Small-footed Myotis
Yuma Myotis
Little Brown Myotis
Long-legged Myotis
 678 Wildlife–Habitat Relationships in Oregon and Washington

Appendix VI. List of published and unpublished observations of wildlife predation and scavenging on salmon.

Species Relationship to salmon References Location of study/report

Incubation—Eggs and Alevin

Cope’sGiant Salamander Recurrent Johnson et al. in prep.
Pacific Giant Salamander Recurrent Graf 1949 California
Pacific Coast Aquatic Recurrent Brown et al. 1995 Oregon
Garter Snake
Common Merganser Strong Munro and Clemens 1939, Munro 1932, 1937 British Columbia
Hooded Merganser Rare Reimchen 1994 British Columbia
Red-breasted Merganser Recurrent Munro and Clemens 1939 Canada
Barrow’s Goldeneye Recurrent Willson and Halupka 1995 Alaska
Munro 1938, 1939 British Columbia
Common Goldeneye Recurrent Willson and Halupka 1995 Alaska
Munro 1938, 1939 British Columbia
American Dipper Recurrent Ehinger 1930 Washington
Munro 1923, Obermayer et al. 1999, Piorkowski Alaska
1995, Willson and Halupka 1995
Reimchen 1994, Burcham 1904 British Columbia
Glaucous-winged Gull Recurrent Baird 1990 Washington
Reimchen 1994 British Columbia
Mossman 1958, Moyle 1966 Alaska
Bonaparte’s Gull Recurrent Moyle 1966 Alaska
Harlequin Duck Strong Dzinbal and Jarvis 1982, Obermayer et al. 1999 Alaska
Horned Grebe Rare Palmer 1962 Washington
Munro 1941 British Columbia
Trumpeter Swan Rare Farley 1980 Alaska
Mallard Rare Willson and Halupka 1995 Alaska
Green-winged Teal Rare Ned Pittman pers. comm. Washington
Greater Scaup Rare Munro 1941 British Columbia
Greater Yellowlegs Rare Elphick and Tibbitts 1998 Alaska
Mew Gull Rare Moyle 1966 Alaska
American Robin Rare Willson and Halupka 1995 Alaska
Varied Thrush Rare Ned Pittman pers. comm. Washington
Water Shrew Recurrent Banfield 1974 Canada
Freshwater Rearing—Fry and Parr
Cope’s Giant Salamander Recurrent Antonelli et al. 1972 Washington
Pacific Giant Salamander Recurrent Antonelli et al. 1972 Washington
Parker 1993, Parker 1994 California
Snapping Turtle Rare Johnson and Hasler 1954, Lagler 1943 Michigan
Western Pond Turtle Rare Johnson et al. in prep.
Pacific Coast Aquatic Recurrent Brown et al. 1995a, Fitch 1941, 1984, Oregon, California
Garter Snake Drummond 1983, Hansen 1980
Western Terrestrial Rare Anderson 1977, Tanner 1949 Montana, Utah
Garter Snake
Common Garter Snake Rare Lagler and Salyer 1945 Michigan
Red-throated Loon Recurrent Eriksson et al. 1990 Sweden
Palmer 1962 Labrador
Common Loon Recurrent Palmer 1962, Johnson and Hasler 1954, Michigan
Alexander 1977
Fraser 1972, 1974, Matkowski 1989, Matkowski Canada, British
1984, Barr 1973, Smith 1968 Columbia
Willson and Halupka 1995 Alaska
Munro 1945 British Columbia
Pied-billed Grebe Recurrent Zarnowitz and Raedeke 1984 Washington
Western Grebe Recurrent Modde et al. 1996 Utah
American White Pelican Recurrent Myers and Peterka 1976, Lingle 1977 North Dakota
Palmer 1962 Wyoming
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 679

Species Relationship to salmon References Location of study/report

Brandt’s Cormorant Recurrent Johnson et al. in prep.

Double-crested Cormorant Recurrent Modde et al. 1996 Utah
Mayers and Peterka 1976 North Dakota
Pelagic Cormorant Recurrent Johnson et al. in prep.
Great Blue Heron Recurrent Alexander 1977, 1979, Bent 1926, Michigan
Johnson and Hasler 1954
Fraser 1972, Matkowski 1989, Smith 1968 Canada
Dolloff 1993, Willson and Halupka 1995 Alaska
Zarnowitz and Raedeke 1984 Washington
Henney and Bethers 1971 Oregon
Great Egret Rare Johnson et al. in press
Snowy Egret Rare Johnson et al. in press
Green Heron Rare Jurek 1974 California
Black-crowned Night Heron Recurrent Spanier 1980 Israel
Myers and Peterka 1976 North Dakota
Trumpeter Swan Rare Farley 1980 Alaska
Hampton 1981 Montana
Common Goldeneye Recurrent Beach 1937 Michigan
White 1939 Nova Scotia
Barrow’s Goldeney Recurrent Munro 1938, Munro and Clemens 1939 British Columbia
Hooded Merganser Rare Zarnowitz and Raedeke 1984 Washington
Common Merganser Strong Alexander 1977, 1979, Beach 1937, Johnson and Michigan
Halser 1954, Salyer and Lagler 1940, Shetter and
Alexander 1970, Miegs and Rieck 1958, Senn 1958 Washington
Fraser 1972, Huntsman 1941, Munro and Clemens Canada
1939, Munro 1923, 1939, Smith 1968
Willson and Halupka 1995 Alaska
White 1936, 1957 Nova Scotia
Red-breasted merganser Recurrent Munro and Clemens 1939 Canada
Marquiss and Duncan 1993 Scotland
Osprey Strong Swenson 1978 Wyoming
Steeger et al. 1992 British Columbia
MacCarter 1972 Montana
Johnson and Hasler 1954 Michigan
Van Daele and Van Daele 1982 Idaho
French and Koplin 1977 California
Hughes 1983 Alaska
Lind 1976 Oregon
Franklin’s Gull Rare Myers and Peterka 1976 North Dakota
Ring-billed Gull Recurrent Johnson et al. in prep.
Caspian Tern Strong Johnson et al. in prep.
Herring Gull Recurrent Mendall 1939 Maine
Forster’s Tern Recurrent Ayles et al. 1976 Canada
Common Tern Recurrent Ayles et al. 1976 Canada
Arctic Tern Recurrent Mossman 1959, Willson and Halupka 1995 Alaska
Marbled Murrelet Recurrent Brooks 1928 British Columbia
Carter and Sealy 1984 Alaska
Belted Kingfisher Recurrent Alexander 1977, 1979 Michigan
Willson and Halupka 1995 Alaska
Gould 1934, Eipper 1956 New York
White 1936 Nova Scotia
Elson 1962, Huntsman 1941 Canada
Black-billed Magpie Recurrent Willson and Halupka 1995 Alaska
American Crow Recurrent Willson and Halupka 1995 Alaska
Northwestern Crow Recurrent Willson and Halupka 1995 Alaska
Common Raven Recurrent Johnson et al. in prep.
American Dipper Recurrent Dolloff 1993 Alaska
Loegering 1997 Oregon
Thut 1970 Washington
Kingery 1996, Cottam and Uhler 1937, Munro 1923 British Columbia
 680 Wildlife–Habitat Relationships in Oregon and Washington

Species Relationship to salmon References Location of study/report

Water Shrew Recurrent Lampman 1947 Oregon

Banfield 1974 Canada
Conaway 1952 Montana
Raccoon Recurrent Alexander 1977 Michigan
Mink Recurrent Whitman 1981 Idaho
Dunstone 1993 New York
Banfield 1974, Burgess and Bider 1980, Fraser 1972 Canada
Ben-David et al. 1997 Alaska
Grinnell et al. 1937 California
Alexander 1977, 1979 Michigan
Akande 1972 Scotland
Northern River Otter Strong Zarnowitz and Raedeke 1984 Washington
Banfield 1974, Stenson et al. 1984 British Columbia
Dolloff 1993 Alaska
Alexander 1979 Michigan
Saltwater—Smolt, Immature Adults, and Adults
Pacific Loon Recurrent Mace 1983 British Columbia
Red-throated Loon Recurrent Johnson et al. in prep.
Common loon Recurrent Palmer 1962 Alaska
Mace 1983 British Columbia
Yellow-billed Loon Rare North 1994 Russia
Horned Grebe Rare Mace 1983,Vermeer et al. 1992 British Columbia
Red-necked Grebe Rare Mace 1983 British Columbia
Western Grebe Recurrent Vermeer 1992, Mace 1983 British Columbia
Clark’s Grebe Recurrent Johnson et al. in prep.
Sooty Shearwater Rare Emmett 1997, Bayer 1989 Oregon
Brown Pelican Rare Bayer 1986a, Emmett 1997, McNeil et al. 1991 Oregon
Brandt’s Cormorant Recurrent Ainley and Sanger 1979 California
Bayer 1986, Scott 1973 Oregon
Double-crested Cormorant Recurrent Bayer 1986, Bayer 1989, Erickson 1988, Hoffman Oregon
and Hall 1988, Roby et al. 1998
Ainley and Anderson 1981, Mace 1993, British Columbia
Robertson 1974
Pelagic Cormorant Recurrent Bayer 1986, Scott 1973 Oregon
Jewett et al. 1953 Washington
Mace 1983 British Columbia
Great Blue Heron Recurrent Forbes and Simpson 1982, Mace 1983, Myers 1980 British Columbia
Green Heron Rare Johnson et al. in prep.
Black-crowned Night Heron Recurrent Johnson et al. in prep.
Great Egret Recurrent Johnson et al. in prep., Schlorff 1978 California
Harlequin Duck Strong Cottam 1939, Mace 1983 British Columbia
Surf Scoter Rare Mace 1983 British Columbia
White-winged Scoter Rare Mace 1983 British Columbia
Common Goldeneye Rare Mace 1983 British Columbia
Barrow’s Goldeneye Rare Mace 1983 British Columbia
Common Merganser Strong Elson 1962 Canada
Mace 1983, Macdonald et al. 1988, Wood and British Columbia
Hand 1985, Wood 1985, 1986, 1987a
Red-breasted Merganser Recurrent Mace 1983 British Columbia
Osprey Strong Bayer 1986a, Emmett 1997, Roby et al. 1998 Oregon
Bald Eagle Recurrent Knight et al. 1990, Watson et al. 1991 Washington
Bonaparte’s Gull Recurrent Macdonald et al. 1988, Mace 1983 British Columbia
Heermann’s Gull Recurrent Bayer 1986, 1989 Oregon
Ring-billed Gull Recurrent Bayer 1989, Roby et al. 1998 Oregon
Ruggerone 1986 Washington
California Gull Recurrent Roby et al. 1998 Oregon
Mace 1983 British Columbia
Herring Gull Recurrent Bent 1921 Washington
Mace 1983, Macdonald et al. 1988 British Columbia
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 681

Species Relationship to salmon References Location of study/report

Thayer’s Gull Recurrent Mace 1983 British Columbia

Western Gull Recurrent Bayer 1986, 1986b, Roby et al. 1998 Oregon
Glaucous-winged Gull Recurrent Baird 1990 Washington
Roby et al. 1998, Bayer 1986 Oregon
Mace 1983,Vermeer 1982 British Columbia
Glaucous Gull Recurrent Sanger 1983 Alaska
Black-legged Kittiwake Rare Rowlett 1980, Sanger 1983 Alaska
Simenstad et al. 1979 Oregon
Caspian Tern Strong Smith and Mudd 1978 Washington
Roby et al. 1998 Oregon
Elegant Tern Recurrent Johnson et al. in press
Common Tern Recurrent Johnson et al. in press, Simenstad et al. 1979 Oregon
Forster’s Tern Recurrent Johnson et al. in press
Arctic Tern Recurrent Simenstad et al. 1979 Oregon
Willson and Halupka 1995 Alaska
Common Murre Recurrent Sydeman et al. 1997 California
Bayer 1986, Bayer 1986b, Matthews 1983 Oregon
Common Murre Recurrent Oregon
Ainley et al. 1990 Alaska
Pigeon Guillemot Rare Bayer 1986 Oregon
Marbled Murrelet Recurrent DeGange 1996 Alaska
Mace 1983 British Columbia
Ancient Murrelet Rare Mace 1983 British Columbia
Rhinoceros Auklet Recurrent Wilson and Manuwal 1986 Washington
Burger et al. 1993,Vermeer and DeVito 1986, British Columbia
Vermeer and Westrheim 1984,Vermeer 1979
Gaston and Dechesne 1996, Sydeman et al. 1997 California
Sanger 1983 Alaska
Tufted Puffin Recurrent Baird 1990, 1991b, Wehle 1983 Alaska
Belted Kingfisher Recurrent Bayer 1989 Oregon
Mace 1983 British Columbia
Northwestern Crow Recurrent Mace 1983 British Columbia
Northern Fur Seal Recurrent Antonelis an Perez 1984, Baker et al. 1970, Washington, Oregon
Kajimura 1983
Clemens and Wilby 1933, Spalding 1964 British Columbia
Scheffer 1950, Wilke and Kenyon 1957 Bering Sea
Banfield 1974 Canada
Fiscus and Baines 1966, Imler and Sarber 1947, Alaska
Pitcher 1981
Dalquest 1948 Washington
Riemer and Brown 1997, Roffe and Mate 1984 Oregon
Banfield 1974, Spalding 1964 British Columbia
California Sea Lion Recurrent Gearin et al. 1988, Jeffries 1985 Washington
Jameson and Kenyon 1977, Riemer and Brown Oregon
1997, Roffe and Mate 1984
Baltz and Morejohn 1977, Harvey and Weise 1997, California
Jones 1981, NMFS 1997
Harbor Seal Recurrent Dalquest 1948, Everitt et al. 1981, Scheffer and Washington
Sperry 1931, Scheffer and Slipp 1944
Beach et al. 1985, Brown and Mate 1983, Brown Oregon
et al. 1995, Browne et al. 1997, Graybill 1981,
Jeffries 1985, Riemer and Brown 1997,
Roffe and Mate 1984
Olesiuk 1993, Olesiuk et al. 1990, Spalding 1964 British Columbia
Imler and Sarber 1947, Pitcher and Calkins 1981, Alaska
Pitcher 1977, 1981
Briggs and Davis 1972, Jones 1981, Herder 1983, California
Hanson 1993
Minke Whale Rare Banfield 1974 Canada
Stewart and Leatherwood 1985 Atlantic Ocean
 682 Wildlife–Habitat Relationships in Oregon and Washington

Species Relationship to salmon References Location of study/report

Humpback Whale Rare Johnson and Wolman 1984 Northern Hemisphere

Pacific White-sided Dolphin Recurrent Kajimura 1983, Stroud et al. 1980 Washington
Northern Right-Whale Rare Johnson et al. in prep.
Dolphin
Killer Whale Strong Hall 1986 Alaska
Scheffer and Slipp 1948 Washington
Killer Whale Strong Nichol and Shackleton 1996, Banfield 1974 British Columbia
Balcomb et al. 1982, Heimlich-Boran 1986, 1988, Pacific Northwest
Felleman et al. 1991
Harbor Porpoise Rare Gearin et al. 1994 Washington
Fontaine et al. 1994 Canada
Dall’s Porpoise Rare Norris and Prescott 1961 Oregon
Mizue et al. 1996 Washington
Sperm Whale Rare Leatherwood and Reeves 1983, Pike 1950 British Columbia
Osprey Strong Johnson et al. in prep.
Bald Eagle Strong Hunt et al. 1992, Servheen 1975, Spencer et al. Washington
1989, Stalmaster 1976
Simons 1983 Oregon
Spawning
Bald Eagle Strong Munro 1938a British Columbia
Ofelt 1975 Alaska
Golden Eagle Recurrent McClelland 1973 Montana
Glaucous-winged Gull Recurrent Mossman 1958 Alaska
Belted Kingfisher Recurrent Reimchen 1994 British Columbia
Common Raven Recurrent Reimchen 1994 British Columbia
Gray Wolf Recurrent Young 1944 Alaska
Black Bear Strong Kellyhouse 1975 California
Reimchen 1994 British Columbia
Reinhart and Mattson 1989 Idaho
Chi 1999, Moyle 1966, Piorkowski 1995, Alaska
Wilson et al. 1998
Banfield 1974 Canada
Grizzly Bear Strong Banfield 1974 Canada
Mink Recurrent Eagle and Whitman 1987 Canada
Melquist et al. 1981 Idaho
Hatler 1976 British Columbia
Northern River Otter Strong Melquist et al. 1981 Idaho
Toweill 1974 Oregon
Mountain Lion Rare Ned Pittman pers. comm. Washington
Bobcat Recurrent Yoakum 1964 Washington
Harbor Seal Recurrent Reimchen 1994 British Columbia
Everitt et al. 1981 Washington
Riemer and Brown 1997 Oregon
California Sea Lion Recurrent Jeffries 1985 Oregon
Riemer and Brown 1997, Roffe and Mate 1984 Oregon
Northern (Steller) Sea Lion Recurrent Roffe and Mate 1984, Riemer and Brown 1997 Oregon
Reimchen 1994 British Columbia
Carcasses
Western Pond Turtle Rare Holland 1985a California
Common Loon Rare Reimchen 1994 British Columbia
Western Grebe Rare Reimcher 1994 British Columbia
Pacific Loon Rare Reimchen 1994 British Columbia
Red-necked Grebe Rare Reimchen 1994 British Columbia
Turkey Vulture Recurrent Jewett et al. 1953 Washington
California Condor Recurrent Gabrielson and Jewett 1940, Simons 1983 Oregon
California Condor Recurrent Suckley and Cooper 1860 Washington
Trumpeter Swan Rare Butler 1973 British Columbia
 CHAPTER 26: SALMON–WILDLIFE RELATIONSHIPS 683

Species Relationship to salmon References Location of study/report

Mallard Rare Jewett et al. 1953 Washington

Reimchen 1994 British Columbia
Canvasback Rare Jewett et al. 1953 Washington
Greater Scaup Rare Munro 1941, Reimchen 1994 British Columbia
Surf Scoter Rare Reimchen 1994 British Columbia
White-winged Scoter Rare Reimchen 1994 British Columbia
Common Goldeneye Recurrent Dawson 1909, Jewett et al. 1953, Servheen 1975 Washington
Taverner 1934 Canada
Barrow’s Goldeneye Recurrent Jewett et al. 1953 Washington
Hooded Merganser Rare Reimchen 1994 British Columbia
Common Merganser Recurrent Stalmaster and Gessaman 1984 Washington
Munro and Clemens 1939, Munro 1932, 1937 British Columbia
Bald Eagle Strong Cederholm et al. 1989, Jewett et al. 1953, Knight Washington
and Knight 1983, Skagen et al. 1991, Stalmaster
and Gessaman 1984
Simons 1983 Oregon
Munro 1938a British Columbia
Shea 1973 Montana
Red-tailed Hawk Rare Willson and Halupka 1995 Alaska
Cederholm et al. 1989, Ned Pittman pers. Washington
comm., Stalmaster and Gessaman 1984,
Stalmaster 1980
Golden Eagle Recurrent Stalmaster and Gessaman 1984 Washington
Ring-billed Gull Recurrent Johnson et al. in prep.
California Gull Recurrent Johnson et al. in prep., Reimchen 1994 British Columbia
Bonaparte’s Gull Recurrent Moyle 1966, Willson and Halupka 1995 Alaska
Western Gull Recurrent Johnson et al. in prep.
Herring Gull Recurrent Reimchen 1994 British Columbia
Glaucous-winged Gull Recurrent Reimchen 1994 British Columbia
Simons 1983 Oregon
Moyle 1966, Bent 1921 Alaska
Glaucous-winged Gull Recurrent Servheen 1975, Skagen et al. 1991, Stalmaster Washington
and Gessaman 1984
Glaucous Gull Recurrent Johnson et al. in prep.
Black-legged Kittiwake Rare Reimchen 1994 British Columbia
Gray Jay Rare Cederholm et al. 1989 Washington
Steller’s Jay Recurrent Cederholm et al. 1989 Washington
Willson and Halupka 1995 Alaska
Black-billed Magpie Recurrent Willson and Halupka 1995 Alaska
McClelland 1973 Montana
American Crow Recurrent Cederholm et al. 1989, Skagen et al. 1991, Washington
Stalmaster and Gessaman 1984
Northwestern Crow Recurrent Campbell et al. 1990, Reimchen 1994 British Columbia
Common Raven Recurrent Murie 1959, Willson and Halupka 1995 Alaska
Knight and Anderson 1990, Stalmaster and Washington
Gessaman 1984, Cederholm et al. 1989, Suckley
and Cooper 1860
Winter Wren Rare Cederholm et al. 1989 Washington
Reimchen 1994 British Columbia
American Dipper Recurrent Willson and Halupka 1995 Alaska
Cederholm et al. 1989 Washington
Varied Thrush Rare Reimchen 1994 British Columbia
Spotted Towhee Rare Ned Pittman pers. comm. Washington
Song Sparrow Rare Ned Pittman pers. comm. Washington
Virginia Opossum Recurrent Johnson et al. in prep.
Deer Mouse Rare Cederholm et al. 1989 Washington
Douglas’ Squirrel Rare Cederholm et al. 1989 Washington
Northern Flying Squirrel Rare Cederholm et al. 1989 Washington
Water Shrew Recurrent Cederholm et al. 1989 Washington
Conaway 1952 Montana
 684 Wildlife–Habitat Relationships in Oregon and Washington

Species Relationship to salmon References Location of study/report

Vagrant Shrew Rare Cederholm et al. 1989 Washington

Masked Shrew Rare Cederholm et al. 1989 Washington
Trowbridge’s Shrew Rare Johnson et al. in prep.
Pacific Water Shrew Rare Johnson et al. in prep.
Pacific Shrew Rare Johnson et al. in prep.
Montane Shrew Rare Johnson et al. in prep.
Fog Shrew Rare Johnson et al. in prep.
Coyote Recurrent Young 1944 Oregon
Cederholm et al. 1989, Stalmaster and Washington
Gessaman 1984
Gray Wolf Recurrent Young 1944 Oregon, British
Columbia, Alaska
Red Fox Rare Young 1944 Oregon
Willson and Halupka 1995 Alaska
Hewson 1995 Scotland
Gray Fox Rare Young 1944 Oregon
Black Bear Strong Young 1944 British Columbia
Piorkowski 1995, Chi 1999 Alaska
Cederholm et al. 1989, Hilderbrand et al. 1996 Washington
Grizzly Bear Strong Mattson et al. 1991 Idaho
Hamilton and Archibald 1985, Hamilton and Bunnel British Columbia
1987,Young 1944
Banfield 1974 Canada
Verts and Carraway 1998 Oregon
Ringtail Rare Johnson et al. in prep.
Raccoon Recurrent Cederholm et al. 1989 Washington
American Marten Rare Reimchen 1994, Nagorsen et al. 1989, 1991, British Columbia
Hatler 1976
Striped Skunk Rare Cederholm et al. 1989 Washington
Long-tailed Weasel Rare Cederholm et al. 1989 Washington
Mink Recurrent Ben-David et al. 1997, Ben-David 1997, Willson Alaska
and Halupka 1995
Cederholm et al. 1989 Washington
Hatler 1976, Young 1944 British Columbia
Eagle and Whitman 1987 Canada
Melquist et al. 1981 Idaho
Fisher Rare Young 1944 British Columbia
Wolverine Rare Willson and Halupka 1995, Banci 1987 Alaska
Northern River Otter Strong Cederholm et al. 1989 Washington
Young 1944 British Columbia
Grinnell et al. 1937 California
Bobcat Recurrent Brittell et al. 1979, Cederholm et al. 1989, Knick Washington
et al. 1984, Schwartz and Mitchell 1945,
Sweeney 1978
White-tailed Deer Rare Shea 1973, Cederholm et al. 1989 Washington
Black-tailed Deer Rare Cederholm et al. 1989 Washington
Northern (Steller) Sea Lion Recurrent Reimchen 1994 British Columbia
Willson and Halupka 1995 Alaska
Harbor Seal Recurrent Reimchen 1994 British Columbia
 27
An Introduction to Wildlife-Habitat
Relationships CD-ROM
MarlaTrevithick,Thomas A. O’Neil, & Charley Barrett

Introduction CD-ROM Navigation

The purpose of this chapter is to provide a brief
introduction to the accompanying CD-ROM to help System Requirements
navigate through the Wildlife-Habitat Relationships The Matrixes for Wildlife-Habitat Relationships in Oregon and
(WHR) digital information, and to list the kinds of Washington’s CD-ROM are optimized to operate on IBM
information found on it. The CD-ROM is divided into four compatible Personal Computers (PCs). See the ReadMe.txt
main sections, Introduction, Definitions, Data Queries, and file for Unix and Macintosh. To view this CD-ROM, you
Maps, which can be found on the Main Navigation Bar. will need Microsoft ® Internet Explorer 5.0 or later.
Specifically, the Introduction gives an overview that has 7 Recommended minimum computer requirements are: a
submenus that depict Acknowledgements, Authors who Pentium 100 processor (or its equivalent), 32 megabytes
developed the digital information, CD-ROM Navigation, RAM (64+ is highly recommended), 4 megabyte video
Trouble-shooting, Site Map MetaData, and copyright adapter, 4X CD-ROM drive or faster, and if copying to a
information. The Definitions section gives explanations of hard drive you will need 100 megabytes of free space.
the fields found within each matrix. The detailed
descriptions can actually be found on the CD-ROM. Data Getting Started
Queries allows the user to view the digital information in To view the CD-ROM, insert the disk into your CD-ROM
a pre-determined or canned query format. These canned drive. On PCs with a Microsoft® Windows operating
queries were developed to answer the most commonly system and Auto-Run enabled, the CD-ROM will
asked questions regarding the subject matter and to allow automatically start. On other systems, start Internet
the user to view and print out the results. Finally, the Maps Explorer and then click “File/Open” from the main menu.
section illustrates several maps of wildlife-habitat types In the Open dialog box, either type “CD-ROM Drive Letter
for Oregon and Washington that can be reviewed on screen [i.e. D:]/START.HTM” into the “Open:” box or use the
and also printed. “Browse” button to locate the file “START.HTM” in the
The data presented on the CD-ROM are a compilation root directory of the CD-ROM. Next, click the “OK” button
of 7 matrixes that focus on wildlife species relationships in the Open dialog box and the CD-ROM will start. Please
with: 1) Wildlife-Habitat Types, 2) Structural Conditions, note, if you experience problems, check the
3) Habitat Elements, 4) Key Ecological Functions, 5) Life Troubleshooting section.
History, 6) Salmon-Wildlife Relationships, and 7)
Management Activity Links. Each of these matrixes can
be found as submenus under the Main Navigation Bar
for Data Queries. In-depth discussions and descriptions
about the CD-ROM data (i.e. metadata) actually reside in
specific chapters within this book, and these chapters will
be highlighted in the Guide to the Wildlife-Habitat
Relationships on the CD-ROM.
Information presented on the CD-ROM has undergone
an extensive Quality Control effort that was the equivalent
of two people working on reviewing the integrity of the
data for two years. Thus a significant attempt has been
made to ensure that the information presented is accurate.

686
 Appendix
Scientific and common names
for 743 wildlife species found in
Oregon and Washington

687
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Amphibia Caudata Ambystomatidae Ambystoma tigrinum Tiger Salamander melanosticum

Amphibia Caudata Ambystomatidae Ambystoma gracile Northwestern Salamander
Amphibia Caudata Ambystomatidae Ambystoma macrodactylum Long-toed Salamander “macrodactylum, columbianum,
sigillatum”
Amphibia Caudata Dicamptodontidae Dicamptodon copei Cope’s Giant Salamander
Amphibia Caudata Dicamptodontidae Dicamptodon tenebrosus Pacific Giant Salamander
(Dicamptodon ensatus)
688

Amphibia Caudata Rhyacotritonidae Rhyacotriton olympicus Olympic Torrent Salamander

(Olympic Seep Salamander)
Amphibia Caudata Rhyacotritonidae Rhyacotriton kezeri Columbia Torrent Salamander
(Rhyacotriton olympicus) (Columbia Seep Salamander)
Amphibia Caudata Rhyacotritonidae Rhyacotriton variegatus Southern Torrent Salamander
(Rhyacotriton olympicus) (Southern Seep Salamander)
Amphibia Caudata Rhyacotritonidae Rhyacotriton cascadae Cascade Torrent Salamander
(Rhyacotriton olympicus)
Amphibia Caudata Salamandridae Taricha granulosa Rough-skinned Newt granulosa
Amphibia Caudata Plethodontidae Plethodon dunni Dunn’s Salamander gordoni
(Plethodon gordoni)
Amphibia Caudata Plethodontidae Plethodon larselli Larch Mountain Salamander
Amphibia Caudata Plethodontidae Plethodon vandykei Van Dyke’s Salamander “idahoensis, vandykei”
Amphibia Caudata Plethodontidae Plethodon vehiculum Western Red-backed Salamander
Amphibia Caudata Plethodontidae Plethodon elongatus Del Norte Salamander
Amphibia Caudata Plethodontidae Plethodon stormi Siskiyou Mountains Salamander
(Plethodon elongatus)
Amphibia Caudata Plethodontidae Ensatina eschscholtzii Ensatina “oregonensis, picta”
Amphibia Caudata Plethodontidae Aneides ferreus Clouded Salamander
Amphibia Caudata Plethodontidae Aneides flavipunctatus Black Salamander
Amphibia Caudata Plethodontidae Batrachoseps wrighti Oregon Slender Salamander
Amphibia Caudata Plethodontidae Batrachoseps attenuatus California Slender Salamander
Amphibia Anura Leiopelmatidae Ascaphus truei Tailed Frog
(American Bell Toad)
Amphibia Anura Pelobatidae Scaphiopus intermontanus Great Basin Spadefoot
Wildlife–Habitat Relationships in Oregon and Washington

(Spea intermontana)
Amphibia Anura Bufonidae Bufo boreas Western Toad halophilus
(“Boreal Toad, Northwestern Toad”)
Amphibia Anura Bufonidae Bufo woodhousii Woodhouse’s Toad woodhousii
Amphibia Anura Hylidae Pseudacris regilla Pacific Chorus (Tree) Frog
(Hyla regilla)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Amphibia Anura Ranidae Rana aurora Red-legged Frog aurora

(Northern Red-Legged Frog)
Amphibia Anura Ranidae Rana cascadae Cascades Frog
Amphibia Anura Ranidae Rana pretiosa Oregon Spotted Frog
(Spotted Frog)
Amphibia Anura Ranidae Rana luteiventris Columbia Spotted Frog
(Rana pretiosa) (Spotted Frog)
Amphibia Anura Ranidae Rana boylii Foothill Yellow-legged Frog None
Amphibia Anura Ranidae Rana pipiens Northern Leopard Frog
Amphibia Anura Ranidae Rana catesbeiana Bullfrog None
Amphibia Anura Ranidae Rana clamitans Green Frog
Reptilia Testudines Chelydridae Chelydra serpentina Snapping Turtle
Reptilia Testudines Emydidae Chrysemys picta Painted Turtle belli
(Western Painted Turtle)
Reptilia Testudines Emydidae Clemmys marmorata Western Pond Turtle marmorata
(Northwestern Pond Turtle)
Reptilia Testudines Emydidae Trachemys scripta Red-eared Slider Turtle “elegans, scripta”
(Pseudemys scripta) (Slider)
Reptilia Testudines Cheloniidae Caretta caretta Loggerhead Sea Turtle
Reptilia Testudines Cheloniidae Chelonia mydas Green Sea Turtle
Reptilia Testudines Cheloniidae Lepidochelys olivacea Pacific Ridley Sea Turtle
Reptilia Testudines Dermochelyidae Dermochelys coriacea Leatherback Turtle
Reptilia Squamata Anguidae Elgaria coerulea Northern Alligator Lizard “principis, shastensis”
(Gerrhonotus coerulea)
APPENDIX: SCIENTIFIC & COMMON NAMES

Reptilia Squamata Anguidae Elgaria multicarinata Southern Alligator Lizard scincicauda

(Gerrhonotus multicarinata)
Reptilia Squamata Iguanidae Crotaphytus bicinctores Mojave Black-collared Lizard
689

Reptilia Squamata Iguanidae Gambelia wislizenii Long-nosed Leopard Lizard

Reptilia Squamata Iguanidae Phrynosoma douglassii Short-horned Lizard douglassii
Reptilia Squamata Iguanidae Phrynosoma platyrhinos Desert Horned Lizard platyrhinos
(Northern Desert Horned Lizard)
Reptilia Squamata Iguanidae Sceloporus graciosus Sagebrush Lizard graciosus
Reptilia Squamata Iguanidae Sceloporus occidentalis Western Fence Lizard “occidentalis, longipes”
Reptilia Squamata Iguanidae Uta stansburiana Side-blotched Lizard stansburiana
Reptilia Squamata Scincidae Eumeces skiltonianus Western Skink “utahensis, skiltonianus”
Reptilia Squamata Teiidae Cnemidophorus tigris Western Whiptail tigris
Reptilia Squamata Teiidae Cnemidophorus velox Plateau Striped Whiptail
Reptilia Squamata Boidae Charina bottae Rubber Boa
Reptilia Squamata Colubridae Coluber constrictor Racer mormon
(Coluber mormon)
Reptilia Squamata Colubridae Contia tenuis Sharptail Snake
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Reptilia Squamata Colubridae Diadophis punctatus Ringneck Snake occcidentalis

(Diadophis amabolis) (Northwest Ringneck Snake)
Reptilia Squamata Colubridae Hypsiglena torquata Night Snake
Reptilia Squamata Colubridae Lampropeltis getula Common Kingsnake californiae
Reptilia Squamata Colubridae Lampropeltis zonata California Mountain Kingsnake
Reptilia Squamata Colubridae Masticophis taeniatus Striped Whipsnake taeniatus
Reptilia Squamata Colubridae Pituophis catenifer Gopher Snake “catenifer, deserticola”
690

(Pituophis melanoleucus)
Reptilia Squamata Colubridae Sonora semiannulata Western Ground Snake None
Reptilia Squamata Colubridae Thamnophis atratus Pacific Coast Aquatic Garter Snake hydrophilus
(Thamnophis couchii)
Reptilia Squamata Colubridae Thamnophis elegans Western Terrestrial Garter Snake “vagrans, elegans, terrestris”
Reptilia Squamata Colubridae Thamnophis ordinoides Northwestern Garter Snake
Reptilia Squamata Colubridae Thamnophis sirtalis Common Garter Snake “concinnus, fitchi, pickeringii”
Reptilia Squamata Viperidae Crotalus viridis Western Rattlesnake “oreganus, lutosus”
Aves Charadriiformes Alcidae Synthliboramphus antiquus Ancient Murrelet None
Aves Charadriiformes Alcidae Ptychoramphus aleuticus Cassin’s Auklet None
Aves Charadriiformes Alcidae Aethia psittacula Parakeet Auklet
(Cyclorrhynchus psittacula)
Aves Charadriiformes Alcidae Cerorhinca monocerata Rhinoceros Auklet None
Aves Charadriiformes Alcidae Fratercula corniculata Horned Puffin
Aves Charadriiformes Alcidae Fratercula cirrhata Tufted Puffin None
Aves Columbiformes Columbidae Columba livia Rock Dove None
Aves Columbiformes Columbidae Columba fasciata Band-tailed Pigeon monilis
Aves Columbiformes Columbidae Zenaida asiatica White-winged Dove
Aves Columbiformes Columbidae Zenaida macroura Mourning Dove marginella
Aves Cuculiformes Cuculidae Coccyzus erythropthalmus Black-billed Cuckoo
Aves Cuculiformes Cuculidae Coccyzus americanus Yellow-billed Cuckoo occidentalis
Aves Strigiformes Tytonidae Tyto alba Barn Owl pratincola
(Common Barn Owl)
Aves Strigiformes Strigidae Otus flammeolus Flammulated Owl idahoensis
Aves Strigiformes Strigidae Otus kennicottii Western Screech-owl “kennicottii (brewsteri), bendirei
Wildlife–Habitat Relationships in Oregon and Washington

(macfarlanei)”
Aves Charadriiformes Scolopacidae Heteroscelus incanus Wandering Tattler
Aves Charadriiformes Scolopacidae Heteroscelus brevipes Gray-tailed Tattler
Aves Charadriiformes Scolopacidae Actitis macularia Spotted Sandpiper None
Aves Charadriiformes Scolopacidae Bartramia longicauda Upland Sandpiper
Aves Charadriiformes Scolopacidae Numenius phaeopus Whimbrel hudsonicus
Aves Charadriiformes Scolopacidae Numenius tahitiensis Bristle-thighed Curlew
Aves Charadriiformes Scolopacidae Numenius americanus Long-billed Curlew parvus
Aves Charadriiformes Scolopacidae Limosa haemastica Hudsonian Godwit
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Charadriiformes Scolopacidae Limosa lapponica Bar-tailed Godwit baueri

Aves Charadriiformes Scolopacidae Limosa fedoa Marbled Godwit “fedoa, beringiae”
Aves Charadriiformes Scolopacidae Arenaria interpres Ruddy Turnstone “interpres, morinella”
Aves Charadriiformes Scolopacidae Arenaria melanocephala Black Turnstone
Aves Charadriiformes Scolopacidae Aphriza virgata Surfbird None
Aves Charadriiformes Scolopacidae Calidris tenuirostris Great Knot
Aves Charadriiformes Scolopacidae Calidris canutus Red Knot “rogersi, canutus”
Aves Charadriiformes Scolopacidae Calidris alba Sanderling
Aves Charadriiformes Scolopacidae Calidris pusilla Semipalmated Sandpiper None
Aves Charadriiformes Scolopacidae Calidris mauri Western Sandpiper None
Aves Charadriiformes Scolopacidae Calidris ruficollis Red-necked Stint
Aves Charadriiformes Scolopacidae Calidris minuta Little Stint
Aves Charadriiformes Scolopacidae Calidris subminuta Long-toed Stint
Aves Charadriiformes Scolopacidae Calidris minutilla Least Sandpiper None
Aves Charadriiformes Scolopacidae Calidris fuscicollis White-rumped Sandpiper
Aves Charadriiformes Scolopacidae Calidris bairdii Baird’s Sandpiper
Aves Charadriiformes Scolopacidae Calidris melanotos Pectoral Sandpiper None
Aves Charadriiformes Scolopacidae Calidris acuminata Sharp-tailed Sandpiper
Aves Charadriiformes Scolopacidae Calidris ptilocnemis Rock Sandpiper “tschuktschorum,couesi”
Aves Charadriiformes Scolopacidae Calidris alpina Dunlin pacifica
Aves Charadriiformes Scolopacidae Calidris ferruginea Curlew Sandpiper
Aves Charadriiformes Scolopacidae Calidris himantopus Stilt Sandpiper None
Aves Charadriiformes Scolopacidae Tryngites subruficollis Buff-breasted Sandpiper None
Aves Charadriiformes Scolopacidae Philomachus pugnax Ruff
APPENDIX: SCIENTIFIC & COMMON NAMES

Aves Charadriiformes Scolopacidae Limnodromus griseus Short-billed Dowitcher “caurinus, hendersoni”

Aves Charadriiformes Scolopacidae Limnodromus scolopaceus Long-billed Dowitcher
Aves Charadriiformes Scolopacidae Gallinago gallinago Common Snipe delicata
691

(Wilson’s Snipe)
Aves Charadriiformes Scolopacidae Phalaropus tricolor Wilson’s Phalarope None
Aves Charadriiformes Scolopacidae Phalaropus lobatus Red-necked Phalarope None
(Northern Phalarope)
Aves Charadriiformes Scolopacidae Phalaropus fulicaria Red Phalarope None
(Grey Phalarope)
Aves Charadriiformes Laridae Stercorarius maccormicki South Polar Skua None
(Mccormick’s Skua)
Aves Charadriiformes Laridae Stercorarius pomarinus Pomarine Jaeger None
(Pomarine Skua)
Aves Charadriiformes Laridae Stercorarius parasiticus Parasitic Jaeger None
(Arctic Skua)
Aves Charadriiformes Laridae Stercorarius longicaudus Long-tailed Jaeger None
(Long-Tailed Skua)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Charadriiformes Laridae Larus atricilla Laughing Gull

Aves Charadriiformes Laridae Larus pipixcan Franklin’s Gull None
Aves Charadriiformes Laridae Larus minutus Little Gull
Aves Charadriiformes Laridae Larus ridibundus Black-headed Gull
(Common Black-Headed Gull)
Aves Charadriiformes Laridae Larus philadelphia Bonaparte’s Gull None
Aves Charadriiformes Laridae Larus heermanni Heermann’s Gull None
692

Aves Charadriiformes Laridae Larus canus Mew Gull brachyrhynchus

(Common Gull)
Aves Charadriiformes Laridae Larus delawarensis Ring-billed Gull None
Aves Charadriiformes Laridae Larus californicus California Gull None
Aves Charadriiformes Laridae Larus argentatus Herring Gull smithsonianus
Aves Charadriiformes Laridae Larus thayeri Thayer’s Gull
Aves Charadriiformes Laridae Larus glaucoides Iceland Gull
Aves Charadriiformes Laridae Larus schistisagus Slaty-backed Gull
Aves Charadriiformes Laridae Larus occidentalis Western Gull occidentalis
Aves Charadriiformes Laridae Larus glaucescens Glaucous-winged Gull None
Aves Charadriiformes Laridae Larus hyperboreus Glaucous Gull “barrovianus, hyperboreus”
Aves Charadriiformes Laridae Xema Sabini Sabine’s Gull None
Aves Charadriiformes Laridae Rissa tridactyla Black-legged Kittiwake pollicaris
Aves Charadriiformes Laridae Rissa brevirostris Red-legged Kittiwake
Aves Charadriiformes Laridae Rhodostethia rosea Ross’s Gull
(Ross’ Gull)
Aves Charadriiformes Laridae Pagophila eburnea Ivory Gull
Aves Charadriiformes Laridae Sterna caspia Caspian Tern None
Aves Charadriiformes Laridae Sterna elegans Elegant Tern None
Aves Charadriiformes Laridae Sterna hirundo Common Tern hirundo
Aves Charadriiformes Laridae Sterna paradisaea Arctic Tern None
Aves Charadriiformes Laridae Sterna forsteri Forster’s Tern None
Aves Charadriiformes Laridae Sterna antillarum Least Tern
Aves Charadriiformes Laridae Chlidonias niger Black Tern surinamensis
Aves Charadriiformes Alcidae Uria aalge Common Murre “inornata, californica”
Wildlife–Habitat Relationships in Oregon and Washington

Aves Charadriiformes Alcidae Uria lomvia Thick-billed Murre

Aves Charadriiformes Alcidae Cepphus columba Pigeon Guillemot columba
Aves Charadriiformes Alcidae Brachyramphus perdix Long-billed Murrelet
Aves Charadriiformes Alcidae Brachyramphus marmoratus Marbled Murrelet “marmoratus, perdix”
Aves Charadriiformes Alcidae Brachyramphus brevirostris Kittlitz’s Murrelet
Aves Charadriiformes Alcidae Synthliboramphus hypoleucus Xantus’s Murrelet
(Xantus’ Murrelet)
Aves Strigiformes Strigidae Bubo virginianus Great Horned Owl “saturatus, lagophonus, subarcticus”
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Strigiformes Strigidae Nyctea scandiaca Snowy Owl None

Aves Strigiformes Strigidae Surnia ulula Northern Hawk Owl
(Northern Hawk-Owl)
Aves Strigiformes Strigidae Glaucidium gnoma Northern Pygmy-owl “grinnelli, californicum”
Aves Strigiformes Strigidae Athene cunicularia Burrowing Owl hypugaea
(Speotyto cunicularia)
Aves Strigiformes Strigidae Strix occidentalis Spotted Owl caurina
Aves Strigiformes Strigidae Strix varia Barred Owl varia
Aves Strigiformes Strigidae Strix nebulosa Great Gray Owl nebulosa
Aves Strigiformes Strigidae Asio otus Long-eared Owl tuftsi
Aves Strigiformes Strigidae Asio flammeus Short-eared Owl flammeus
Aves Strigiformes Strigidae Aegolius funereus Boreal Owl richardsoni
Aves Strigiformes Strigidae Aegolius acadicus Northern Saw-whet Owl acadicus
Aves Caprimulgiformes Caprimulgidae Chordeiles minor Common Nighthawk “hesperis, minor”
Aves Caprimulgiformes Caprimulgidae Phalaenoptilus nuttallii Common Poorwill californicus, nuttallii
Aves Apodiformes Apodidae Cypseloides niger Black Swift borealis
Aves Apodiformes Apodidae Chaetura vauxi Vaux’s Swift vauxi
Aves Apodiformes Apodidae Aeronautes saxatalis White-throated Swift saxatalis
Aves Apodiformes Trochilidae Archilochus alexandri Black-chinned Hummingbird None
Aves Apodiformes Trochilidae Calypte anna Anna’s Hummingbird None
Aves Apodiformes Trochilidae Calypte costae Costa’s Hummingbird
Aves Apodiformes Trochilidae Stellula calliope Calliope Hummingbird None
Aves Apodiformes Trochilidae Selasphorus platycercus Broad-tailed Hummingbird platycercus
Aves Apodiformes Trochilidae Selasphorus rufus Rufous Hummingbird None
APPENDIX: SCIENTIFIC & COMMON NAMES

Aves Apodiformes Trochilidae Selasphorus sasin Allen’s Hummingbird sasin

Aves Coraciiformes Alcedinidae Ceryle alcyon Belted Kingfisher caurina
Aves Piciformes Picidae Melanerpes lewis Lewis’s Woodpecker None
693

(Lewis’ Woodpecker)
Aves Piciformes Picidae Melanerpes formicivorus Acorn Woodpecker bairdi
Aves Piciformes Picidae Sphyrapicus thyroideus Williamson’s Sapsucker None
Aves Piciformes Picidae Sphyrapicus varius Yellow-bellied Sapsucker
Aves Piciformes Picidae Sphyrapicus nuchalis Red-naped Sapsucker None
Aves Piciformes Picidae Sphyrapicus ruber Red-breasted Sapsucker “ruber, daggettii”
Aves Piciformes Picidae Picoides nuttallii Nuttall’s Woodpecker
Aves Piciformes Picidae Picoides pubescens Downy Woodpecker “fumidus, gairdnerii, turati, leucurus”
Aves Piciformes Picidae Picoides villosus Hairy Woodpecker “monticola, orius, harrisi, septentrionalis”
Aves Piciformes Picidae Picoides albolarvatus White-headed Woodpecker albolarvatus
Aves Piciformes Picidae Picoides tridactylus Three-toed Woodpecker fasciatus
Aves Piciformes Picidae Picoides arcticus Black-backed Woodpecker None
Aves Piciformes Picidae Colaptes auratus Northern Flicker “cafer, collaris”
(Red-Shafted Flicker)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Piciformes Picidae Dryocopus pileatus Pileated Woodpecker abieticola (picinus)

Aves Passeriformes Tyrannidae Contopus cooperi Olive-sided Flycatcher cooperi
(Contopus borealis)
Aves Passeriformes Tyrannidae Contopus sordidulus Western Wood-pewee “veliei, saturatus”
Aves Passeriformes Tyrannidae Contopus virens Eastern Wood-pewee
Aves Passeriformes Tyrannidae Empidonax traillii Willow Flycatcher “brewsteri, adastus”
(Traill’s Flycatcher)
694

Aves Passeriformes Tyrannidae Empidonax minimus Least Flycatcher None

Aves Passeriformes Tyrannidae Empidonax hammondii Hammond’s Flycatcher None
Aves Passeriformes Tyrannidae Empidonax wrightii Gray Flycatcher None
Aves Passeriformes Tyrannidae Empidonax oberholseri Dusky Flycatcher None
(Empidonax wrighti)
Aves Passeriformes Tyrannidae Empidonax difficilis Pacific-slope Flycatcher difficilis
(Western Flycatcher)
Aves Passeriformes Tyrannidae Empidonax occidentalis Cordilleran Flycatcher helmayi
(Empidonax difficilis) (Western Flycatcher)
Aves Passeriformes Tyrannidae Sayornis nigricans Black Phoebe semiatra
Aves Passeriformes Tyrannidae Sayornis phoebe Eastern Phoebe
Aves Passeriformes Tyrannidae Sayornis saya Say’s Phoebe “saya, yukonensis”
Aves Passeriformes Tyrannidae Pyrocephalus rubinus Vermilion Flycatcher
Aves Passeriformes Tyrannidae Myiarchus cinerascens Ash-throated Flycatcher cinerascens
Aves Passeriformes Tyrannidae Tyrannus melancholicus Tropical Kingbird
Aves Passeriformes Tyrannidae Tyrannus verticalis Western Kingbird None
Aves Passeriformes Tyrannidae Tyrannus tyrannus Eastern Kingbird None
Aves Passeriformes Tyrannidae Tyrannus forficatus Scissor-tailed Flycatcher
Aves Passeriformes Tyrannidae Tyrannus savana Fork-tailed Flycatcher
Aves Passeriformes Laniidae Lanius ludovicianus Loggerhead Shrike “mexicanus (gmbeli), exubitorides”
Aves Passeriformes Laniidae Lanius excubitor Northern Shrike borealis
Aves Passeriformes Vireonidae Vireo griseus White-eyed Vireo
Aves Passeriformes Vireonidae Vireo bellii Bell’s Vireo
Aves Passeriformes Vireonidae Vireo flavifrons Yellow-throated Vireo
Aves Passeriformes Vireonidae Vireo plumbeus Plumbeous Vireo plumbeus
Wildlife–Habitat Relationships in Oregon and Washington

(Solitary Vireo)
Aves Passeriformes Vireonidae Vireo cassinii Cassin’s Vireo None
(Vireo solitarius) (Solitary Vireo)
Aves Passeriformes Vireonidae Vireo huttoni Hutton’s Vireo obscurus
Aves Passeriformes Vireonidae Vireo gilvus Warbling Vireo swainsoni (leucopolius)
Aves Passeriformes Vireonidae Vireo philadelphicus Philadelphia Vireo
Aves Passeriformes Vireonidae Vireo olivaceus Red-eyed Vireo carnividis
Aves Passeriformes Corvidae Perisoreus canadensis Gray Jay “bicolor, obscurus (rathbuni, connexus,
(Canada Jay) griseus)”
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Passeriformes Corvidae Cyanocitta stelleri Steller’s Jay “annectens, fronatlis, paralia”
Aves Passeriformes Corvidae Cyanocitta cristata Blue Jay
Aves Passeriformes Corvidae Aphelocoma californica Western Scrub-Jay “californica (immanis, caurina),
(Aphelocoma coerulescens) (Scrub Jay) superciliosa, woodhouseii = nevadae,”
Aves Passeriformes Corvidae Gymnorhinus cyanocephalus Pinyon Jay cassini
Aves Passeriformes Corvidae Nucifraga columbiana Clark’s Nutcracker None
Aves Passeriformes Corvidae Pica hudsonia Black-billed Magpie
(Pica pica)
Aves Passeriformes Corvidae Corvus brachyrhynchos American Crow hesperis
Aves Passeriformes Corvidae Corvus caurinus Northwestern Crow None
Aves Passeriformes Corvidae Corvus corax Common Raven “sinuatus, clarionensis”
Aves Passeriformes Alaudidae Alauda arvensis Sky Lark arvensis
(Eurasian Skylark)
Aves Passeriformes Alaudidae Eremophila alpestris Horned Lark “strigata, arcticola, alpina, merrilli,
lamprochroma”
Aves Passeriformes Hirundinidae Progne subis Purple Martin “arboricola, subis”
Aves Passeriformes Hirundinidae Tachycineta bicolor Tree Swallow None
Aves Passeriformes Hirundinidae Tachycineta thalassina Violet-green Swallow thalassina (lepida)
Aves Passeriformes Hirundinidae Stelgidopteryx serripennis Northern Rough-winged Swallow serripennis (aphractus)
(Rough-Winged Swallow)
Aves Passeriformes Hirundinidae Riparia riparia Bank Swallow riparia
(Sand Martin)
Aves Passeriformes Hirundinidae Petrochelidon pyrrhonota Cliff Swallow “pyrrhonota, hypopolia, aprophata”
(Hirundo pyrrhonota)
APPENDIX: SCIENTIFIC & COMMON NAMES

Aves Passeriformes Hirundinidae Hirundo rustica Barn Swallow erythrogaster

Aves Passeriformes Paridae Poecile atricapilla Black-capped Chickadee “fortuitus, occidentalis, nevadensis”
(Poecile atricapillus, Parus atricapillus)
695

Aves Passeriformes Paridae Poecile gambeli Mountain Chickadee “baileyae (grinnelli, abbreviatus),
(Parus gambeli) inyoensis”
Aves Passeriformes Paridae Poecile rufescens Chestnut-backed Chickadee rufescens
(Parus rufescens)
Aves Passeriformes Paridae Poecile hudsonica Boreal Chickadee columbianus (cascadensis)
(Poecile hudsonicus, Parus hudsonicus)
Aves Passeriformes Paridae Baeolophus inornatus Oak Titmouse inornatus (sequestratus)
(Parus inornatus) (Plain Titmouse)
Aves Passeriformes Paridae Baeolophus ridgwayi Juniper Titmouse zaleptus
(“Parus inornatus, (Plain Titmouse)
Baeolophus griseus”)
Aves Passeriformes Aegithalidae Psaltriparus minimus Bushtit “saturatus, minimus, californicus,
plumbeus”
Aves Passeriformes Sittidae Sitta canadensis Red-breasted Nuthatch None
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Passeriformes Sittidae Sitta carolinensis White-breasted Nuthatch “aculeata, tenuissima”

Aves Passeriformes Sittidae Sitta pygmaea Pygmy Nuthatch melanotis
Aves Passeriformes Certhiidae Certhia americana Brown Creeper “montana, occidentalis, zelotes”
Aves Passeriformes Troglodytidae Salpinctes obsoletus Rock Wren obsoletus
Aves Passeriformes Troglodytidae Catherpes mexicanus Canyon Wren griseus
Aves Passeriformes Troglodytidae Thryomanes bewickii Bewick’s Wren “calophonus, drymoecus, atrestus”
Aves Passeriformes Troglodytidae Troglodytes aedon House Wren parkmanii
696

Aves Passeriformes Troglodytidae Troglodytes troglodytes Winter Wren “pacificus, salebrosus”

Aves Passeriformes Troglodytidae Cistothorus palustris Marsh Wren “pulverius, browningi, paludicola, plesius”
(Long-Billed Marsh Wren)
Aves Passeriformes Cinclidae Cinclus mexicanus American Dipper unicolor
Aves Passeriformes Regulidae Regulus satrapa Golden-crowned Kinglet “olivaceus, apache”
Aves Passeriformes Regulidae Regulus calendula Ruby-crowned Kinglet “calendula, grinnelli”
Aves Passeriformes Sylviidae Polioptila caerulea Blue-gray Gnatcatcher obscura
Aves Passeriformes Turdidae Oenanthe oenanthe Northern Wheatear
Aves Passeriformes Turdidae Sialia mexicana Western Bluebird occidentalis
Aves Passeriformes Turdidae Sialia currucoides Mountain Bluebird None
Aves Passeriformes Turdidae Myadestes townsendi Townsend’s Solitaire townsendi
Aves Passeriformes Turdidae Catharus fuscescens Veery salicicola
Aves Passeriformes Turdidae Catharus minimus Gray-cheeked Thrush
Aves Passeriformes Turdidae Catharus ustulatus Swainson’s Thrush “ustulatus, swainsoni”
Aves Passeriformes Turdidae Catharus guttatus Hermit Thrush “auduboni, slevini, jewetti, vaccinius,
oromelus = dwighti, nanus = osgoodi,
guttatus, verecundas”
Aves Passeriformes Turdidae Hylocichla mustelina Wood Thrush
Aves Passeriformes Turdidae Turdus migratorius American Robin “caurinus, propinquus, migratorius”
Aves Passeriformes Turdidae Ixoreus naevius Varied Thrush “naevius, godfreii, meruloides, carlottae”
Aves Passeriformes Timaliidae Chamaea fasciata Wrentit “phaea, margra”
Aves Passeriformes Mimidae Dumetella carolinensis Gray Catbird ruficrissa
Aves Passeriformes Mimidae Mimus polyglottos Northern Mockingbird None
Aves Passeriformes Mimidae Oreoscoptes montanus Sage Thrasher None
Aves Passeriformes Mimidae Toxostoma rufum Brown Thrasher
Wildlife–Habitat Relationships in Oregon and Washington

Aves Passeriformes Mimidae Toxostoma redivivum California Thrasher

Aves Passeriformes Sturnidae Sturnus vulgaris European Starling vulgaris
Aves Passeriformes Prunellidae Prunella montanella Siberian Accentor
Aves Passeriformes Motacillidae Motacilla flava Yellow Wagtail
Aves Passeriformes Motacillidae Motacilla alba White Wagtail
Aves Passeriformes Motacillidae Motacilla lugens Black-backed Wagtail
Aves Passeriformes Motacillidae Anthus cervinus Red-throated Pipit
Aves Passeriformes Motacillidae Anthus rubescens American Pipit “pacificus, alticola, geophilus”
(Anthus spinoletta) (Water Pipit)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Passeriformes Bombycillidae Bombycilla garrulus Bohemian Waxwing pallidiceps

Aves Passeriformes Bombycillidae Bombycilla cedrorum Cedar Waxwing larifuga
Aves Passeriformes Ptilogonatidae Phainopepla nitens Phainopepla
Aves Passeriformes Parulidae Vermivora pinus Blue-winged Warbler
Aves Passeriformes Parulidae Vermivora chrysoptera Golden-winged Warbler
Aves Passeriformes Parulidae Vermivora peregrina Tennessee Warbler
Aves Passeriformes Parulidae Vermivora celata Orange-crowned Warbler “lutescens, orestera”
Aves Passeriformes Parulidae Vermivora ruficapilla Nashville Warbler ridgwayi
Aves Passeriformes Parulidae Vermivora virginiae Virginia’s Warbler
Aves Passeriformes Parulidae Vermivora luciae Lucy’s Warbler
Aves Passeriformes Parulidae Parula americana Northern Parula
Aves Passeriformes Parulidae Dendroica petechia Yellow Warbler “morcomi,brewsteri”
Aves Passeriformes Parulidae Dendroica pensylvanica Chestnut-sided Warbler
Aves Passeriformes Parulidae Dendroica magnolia Magnolia Warbler
Aves Passeriformes Parulidae Dendroica tigrina Cape May Warbler
Aves Passeriformes Parulidae Dendroica caerulescens Black-throated Blue Warbler
Aves Passeriformes Parulidae Dendroica coronata Yellow-rumped Warbler auduboni
(Audubon’s Warbler)
Aves Passeriformes Parulidae Dendroica nigrescens Black-throated Gray Warbler “nigrescens, halseii”
Aves Passeriformes Parulidae Dendroica virens Black-throated Green Warbler
Aves Passeriformes Parulidae Dendroica townsendi Townsend’s Warbler None
Aves Passeriformes Parulidae Dendroica occidentalis Hermit Warbler None
Aves Passeriformes Parulidae Dendroica fusca Blackburnian Warbler
Aves Passeriformes Parulidae Dendroica dominica Yellow-throated Warbler
APPENDIX: SCIENTIFIC & COMMON NAMES

Aves Passeriformes Parulidae Dendroica pinus Pine Warbler

Aves Passeriformes Parulidae Dendroica discolor Prairie Warbler
Aves Passeriformes Parulidae Dendroica palmarum Palm Warbler palmarum
697

Aves Passeriformes Parulidae Dendroica castanea Bay-breasted Warbler

Aves Passeriformes Parulidae Dendroica striata Blackpoll Warbler
Aves Passeriformes Parulidae Mniotilta varia Black-and-white Warbler
Aves Passeriformes Parulidae Setophaga ruticilla American Redstart None
Aves Passeriformes Parulidae Protonotaria citrea Prothonotary Warbler
Aves Passeriformes Parulidae Helmitheros vermivorus Worm-eating Warbler
Aves Passeriformes Parulidae Seiurus aurocapillus Ovenbird
Aves Passeriformes Parulidae Seiurus noveboracensis Northern Waterthrush notabilis
Aves Passeriformes Parulidae Oporornis formosus Kentucky Warbler
Aves Passeriformes Parulidae Oporornis philadelphia Mourning Warbler
Aves Passeriformes Parulidae Oporornis tolmiei Macgillivray’s Warbler tolmiei (intermedia)
Aves Passeriformes Parulidae Geothlypis trichas Common Yellowthroat “arizela, campicola, occidentalis
(idahonicola, oregonicola)”
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Passeriformes Parulidae Wilsonia citrina Hooded Warbler

Aves Passeriformes Parulidae Wilsonia pusilla Wilson’s Warbler “pileolata, chryseola”
Aves Passeriformes Parulidae Wilsonia canadensis Canada Warbler
Aves Passeriformes Parulidae Icteria virens Yellow-breasted Chat auricollis (longicauda)
Aves Passeriformes Thraupidae Piranga rubra Summer Tanager
Aves Passeriformes Thraupidae Piranga olivacea Scarlet Tanager
Aves Passeriformes Thraupidae Piranga ludoviciana Western Tanager None
698

Aves Passeriformes Emberizidae Pipilo chlorurus Green-tailed Towhee None

Aves Passeriformes Emberizidae Pipilo maculatus Spotted Towhee “oreganus, falcinellus, curtatus”
(Pipilo erythrophthalmus) (Rufous-Sided Towhee)
Aves Passeriformes Emberizidae Pipilo crissalis California Towhee bullatus
(Pipilo fuscus) (Brown Towhee)
Aves Passeriformes Emberizidae Spizella arborea American Tree Sparrow ochracea
Aves Passeriformes Emberizidae Spizella passerina Chipping Sparrow “stridula, arizonae”
Aves Passeriformes Emberizidae Spizella pallida Clay-colored Sparrow None
Aves Passeriformes Emberizidae Spizella breweri Brewer’s Sparrow “breweri, taverneri”
Aves Passeriformes Emberizidae Spizella atrogularis Black-chinned Sparrow cana
Aves Passeriformes Emberizidae Pooecetes gramineus Vesper Sparrow “affinis, confinis”
Aves Passeriformes Emberizidae Chondestes grammacus Lark Sparrow strigatus (actitus)
Aves Passeriformes Emberizidae Amphispiza bilineata Black-throated Sparrow deserticola
Aves Passeriformes Emberizidae Amphispiza belli Sage Sparrow campicola = nevadensis
Aves Passeriformes Emberizidae Calamospiza melanocorys Lark Bunting
Aves Passeriformes Emberizidae Passerculus sandwichensis Savannah Sparrow “brooksi, nevadensis, crassus,
sandwichensis, anthinus”
Aves Passeriformes Emberizidae Ammodramus savannarum Grasshopper Sparrow perpallidus
Aves Passeriformes Emberizidae Ammodramus leconteii Le Conte’s Sparrow
Aves Passeriformes Emberizidae Ammodramus nelsoni Nelson’s Sharp-tailed Sparrow
(Ammodramus caudacutus) (Sharp-Tailed Sparrow)
Aves Passeriformes Emberizidae Passerella iliaca Fox Sparrow “fuliginosa, schistacea, olivacea,
unalaschcensis, ridgwayi (insularis),
sinuosa, annectens, townsendi,
chilcatensis, megarhyncha”
Wildlife–Habitat Relationships in Oregon and Washington

Aves Passeriformes Emberizidae Melospiza melodia Song Sparrow “merrilli, fisherella, morphna, kenaiensis,
caurina, rufina, inexpectata, cleonensis,
montana”
Aves Passeriformes Emberizidae Melospiza lincolnii Lincoln’s Sparrow “lincolnii, alticola”
Aves Passeriformes Emberizidae Melospiza georgiana Swamp Sparrow ericrypta
Aves Passeriformes Emberizidae Zonotrichia albicollis White-throated Sparrow None
Aves Passeriformes Emberizidae Zonotrichia querula Harris’s Sparrow None
(Harris’ Sparrow)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Passeriformes Emberizidae Zonotrichia leucophrys White-crowned Sparrow “pugetensis, gambelii, oriantha”
Aves Passeriformes Emberizidae Zonotrichia atricapilla Golden-crowned Sparrow None
Aves Passeriformes Emberizidae Junco hyemalis Dark-eyed Junco “oreganus, simillimus, thurberi, shufeldti
(montanus, eumesus)”
Aves Passeriformes Emberizidae Calcarius mccownii McCown’s Longspur
Aves Passeriformes Emberizidae Calcarius lapponicus Lapland Longspur alascensis
Aves Passeriformes Emberizidae Calcarius ornatus Chestnut-collared Longspur
Aves Passeriformes Emberizidae Emberiza rustica Rustic Bunting
Aves Passeriformes Emberizidae Plectrophenax nivalis Snow Bunting Nivalis
Aves Passeriformes Emberizidae Plectrophenax hyperboreus Mckay’s Bunting
Aves Passeriformes Cardinalidae Pheucticus ludovicianus Rose-breasted Grosbeak
Aves Passeriformes Cardinalidae Pheucticus melanocephalus Black-headed Grosbeak “maculatus, melanocephalus”
Aves Passeriformes Cardinalidae Guiraca caerulea Blue Grosbeak
Aves Passeriformes Cardinalidae Passerina amoena Lazuli Bunting None
Aves Passeriformes Cardinalidae Passerina cyanea Indigo Bunting
Aves Passeriformes Cardinalidae Passerina ciris Painted Bunting
Aves Passeriformes Cardinalidae Spiza americana Dickcissel
Aves Passeriformes Icteridae Dolichonyx oryzivorus Bobolink None
Aves Passeriformes Icteridae Agelaius phoeniceus Red-winged Blackbird “caurinus, nevadensis”
Aves Passeriformes Icteridae Agelaius tricolor Tricolored Blackbird None
Aves Passeriformes Icteridae Sturnella neglecta Western Meadowlark “confluenta, neglecta”
Aves Passeriformes Icteridae Xanthocephalus xanthocephalus Yellow-headed Blackbird None
Aves Passeriformes Icteridae Euphagus carolinus Rusty Blackbird
Aves Passeriformes Icteridae Euphagus cyanocephalus Brewer’s Blackbird “minusculus, brewsteri, cyanocephalus
APPENDIX: SCIENTIFIC & COMMON NAMES

(aliastus)”
Aves Passeriformes Icteridae Quiscalus quiscula Common Grackle
Aves Passeriformes Icteridae Quiscalus mexicanus Great-tailed Grackle
699

Aves Passeriformes Icteridae Molothrus ater Brown-headed Cowbird artemisiae

Aves Passeriformes Icteridae Icterus spurius Orchard Oriole
Aves Passeriformes Icteridae Icterus cucullatus Hooded Oriole
Aves Passeriformes Icteridae Icterus pustulatus Streak-backed Oriole
Aves Passeriformes Icteridae Icterus galbula Baltimore Oriole
(Northern Oriole)
Aves Passeriformes Icteridae Icterus bullockii Bullock’s Oriole bullockii
(Icterus galbula) (Northern Oriole)
Aves Passeriformes Icteridae Icterus parisorum Scott’s Oriole
Aves Passeriformes Fringillidae Fringilla montifringilla Brambling
Aves Passeriformes Fringillidae Leucosticte tephrocotis Gray-crowned Rosy-Finch “wallowa,tephrocotis, littoralis”
(Leucosticte arctoa) (Rosy Finch)
Aves Passeriformes Fringillidae Leucosticte atrata Black Rosy-finch None
(Leucosticte arctoa) (Rosy Finch)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Passeriformes Fringillidae Pinicola enucleator Pine Grosbeak “flammula, carlottae, montanus, leucurus”
Aves Passeriformes Fringillidae Carpodacus purpureus Purple Finch californicus (rubidus)
Aves Passeriformes Fringillidae Carpodacus cassinii Cassin’s Finch vinifer
Aves Passeriformes Fringillidae Carpodacus mexicanus House Finch “frontalis (grinnelli), solitudinus”
Aves Passeriformes Fringillidae Loxia curvirostra Red Crossbill “minor group, pusilla group”
Aves Passeriformes Fringillidae Loxia leucoptera White-winged Crossbill leucoptera
Aves Gaviiformes Gaviidae Gavia stellata Red-throated Loon None
700

(Red-Throated Diver)
Aves Gaviiformes Gaviidae Gavia pacifica Pacific Loon
(Black-Throated Diver)
Aves Gaviiformes Gaviidae Gavia immer Common Loon None
(Great Northern Diver)
Aves Gaviiformes Gaviidae Gavia adamsii Yellow-billed Loon None
(White-Billed Diver)
Aves Podicipediformes Podicipedidae Podilymbus podiceps Pied-billed Grebe podiceps
Aves Podicipediformes Podicipedidae Podiceps auritus Horned Grebe cornutus
(Slavonian Grebe)
Aves Podicipediformes Podicipedidae Podiceps grisegena Red-necked Grebe holbollii
(“Holboll’s Grebe, Holboell’s Grebe,
Gray-Cheeked Grebe”)
Aves Podicipediformes Podicipedidae Podiceps nigricollis Eared Grebe californicus
(Black-Necked Grebe)
Aves Podicipediformes Podicipedidae Aechmophorus occidentalis Western Grebe occidentalis
Aves Podicipediformes Podicipedidae Aechmophorus clarkii Clark’s Grebe clarkii
Aves Procellariiformes Diomedeidae Thalassarche cauta Shy Albatross
Aves Procellariiformes Diomedeidae Phoebastria immutabilis Laysan Albatross None
(Diomedea immutabilis)
Aves Procellariiformes Diomedeidae Phoebastria nigripes Black-footed Albatross None
(Diomedea nigripes)
Aves Procellariiformes Diomedeidae Phoebastria albatrus Short-tailed Albatross
(Diomedea albatrus) (Steller’s Albatross)
Aves Procellariiformes Procellariidae Fulmarus glacialis Northern Fulmar rodgersii
Wildlife–Habitat Relationships in Oregon and Washington

(Arctic Fulmar)
Aves Procellariiformes Procellariidae Pterodroma ultima Murphy’s Petrel
Aves Procellariiformes Procellariidae Pterodroma inexpectata Mottled Petrel
Aves Procellariiformes Procellariidae Pterodroma cookii Cook’s Petrel
Aves Procellariiformes Procellariidae Puffinus creatopus Pink-footed Shearwater
Aves Procellariiformes Procellariidae Puffinus carneipes Flesh-footed Shearwater
Aves Procellariiformes Procellariidae Puffinus bulleri Buller’s Shearwater None
(New Zealand/Grey-Backed Shearwater)
Aves Procellariiformes Procellariidae Puffinus griseus Sooty Shearwater
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Procellariiformes Procellariidae Puffinus tenuirostris Short-tailed Shearwater None

(Slender-Billed Shearwater)
Aves Procellariiformes Procellariidae Puffinus ruffinus Manx Shearwater
Aves Procellariiformes Procellariidae Puffinus opisthomelas Black-vented Shearwater
Aves Procellariiformes Hydrobatidae Oceanites oceanicus Wilson’s Storm-petrel
Aves Procellariiformes Hydrobatidae Oceanodroma furcata Fork-tailed Storm-petrel plumbea
Aves Procellariiformes Hydrobatidae Oceanodroma leucorhoa Leach’s Storm-petrel leucorhoa
Aves Procellariiformes Hydrobatidae Oceanodroma melania Black Storm-petrel
Aves Pelecaniformes Phaethontidae Phaethon aethereus Red-billed Tropicbird
Aves Pelecaniformes Sulidae Sula nebouxii Blue-footed Booby
Aves Pelecaniformes Pelecanidae Pelecanus erythrorhynchos American White Pelican None
Aves Pelecaniformes Pelecanidae Pelecanus occidentalis Brown Pelican californicus
Aves Pelecaniformes Phalacrocoracidae Phalacrocorax penicillatus Brandt’s Cormorant None
Aves Pelecaniformes Phalacrocoracidae Phalacrocorax auritus Double-crested Cormorant albociliatus
Aves Pelecaniformes Phalacrocoracidae Phalacrocorax pelagicus Pelagic Cormorant resplendens
Aves Pelecaniformes Fregatidae Fregata magnificens Magnificent Frigatebird
Aves Ciconiiformes Ardeidae Botaurus lentiginosus American Bittern None
Aves Ciconiiformes Ardeidae Ixobrychus exilis Least Bittern hesperis
Aves Ciconiiformes Ardeidae Ardea herodias Great Blue Heron “herodias, treganzai, fannini”
Aves Ciconiiformes Ardeidae Ardea alba Great Egret egretta
(Casmerodius albus)
Aves Ciconiiformes Ardeidae Egretta thula Snowy Egret brewsteri
Aves Ciconiiformes Ardeidae Egretta caerulea Little Blue Heron
APPENDIX: SCIENTIFIC & COMMON NAMES

Aves Passeriformes Fringillidae Carduelis flammea Common Redpoll flammea

Aves Passeriformes Fringillidae Carduelis hornemanni Hoary Redpoll
Aves Passeriformes Fringillidae Carduelis pinus Pine Siskin “vagans, pinus”
701

Aves Passeriformes Fringillidae Carduelis psaltria Lesser Goldfinch psaltria (hesperophilus)

Aves Passeriformes Fringillidae Carduelis lawrencei Lawrence’s Goldfinch
Aves Passeriformes Fringillidae Carduelis tristis American Goldfinch “pallida, jewetti”
Aves Passeriformes Fringillidae Coccothraustes vespertinus Evening Grosbeak brooksi (californica)
Aves Passeriformes Passeridae Passer domesticus House Sparrow domesticus
Aves Ciconiiformes Ardeidae Egretta tricolor Tricolored Heron
Aves Ciconiiformes Ardeidae Bubulcus ibis Cattle Egret ibis
Aves Ciconiiformes Ardeidae Butorides virescens Green Heron anthonyi
(Butorides striatus) (Green-Backed Heron)
Aves Ciconiiformes Ardeidae Nycticorax nycticorax Black-crowned Night-heron hoactli
Aves Ciconiiformes Ardeidae Nyctanassa violacea Yellow-crowned Night-heron None
Aves Ciconiiformes Threskiornithidae Plegadis chihi White-faced Ibis
Aves Ciconiiformes Cathartidae Cathartes aura Turkey Vulture meridionalis
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Ciconiiformes Cathartidae Gymnogyps californianus California Condor

Aves Anseriformes Anatidae Dendrocygna bicolor Fulvous Whistling-Duck
Aves Anseriformes Anatidae Anser albifrons Greater White-fronted Goose “frontalis, gambeli”
Aves Anseriformes Anatidae Chen canagica Emperor Goose
Aves Anseriformes Anatidae Chen Ccaerulescens Snow Goose caerulescens
(Lesser Snow Goose)
Aves Anseriformes Anatidae Chen rossii Ross’s Goose None
702

(Ross’ Goose)
Aves Anseriformes Anatidae Branta canadensis Canada Goose
(Great Basin Canada Goose or
Giant Canada Goose)
Aves Anseriformes Anatidae Branta canadensis moffitti Western Canada Goose
Aves Anseriformes Anatidae Branta canadensis maxima Giant Canada Goose
Aves Anseriformes Anatidae Branta canadensis taverneri Taverner’s Canada Goose
Aves Anseriformes Anatidae Branta canadensis leucopareia Aleutian Canada Goose
Aves Anseriformes Anatidae Branta canadensis minima Cackling Canada Goose
Aves Anseriformes Anatidae Branta canadensis occidentalis Dusky Canada Goose
Aves Anseriformes Anatidae Branta canadensis fulva Vancouver Canada Goose
Aves Anseriformes Anatidae Branta canadensis parvipes Lesser Canada Goose
Aves Anseriformes Anatidae Branta bernicla Brant nigricans
(Black Brant)
Aves Anseriformes Anatidae Cygnus olor Mute Swan None
Aves Anseriformes Anatidae Cygnus buccinator Trumpeter Swan None
Aves Anseriformes Anatidae Cygnus columbianus Tundra Swan columbianus
Aves Anseriformes Anatidae Cygnus cygnus Whooper Swan
Aves Anseriformes Anatidae Aix sponsa Wood Duck None
Aves Anseriformes Anatidae Anas strepera Gadwall None
Aves Anseriformes Anatidae Anas falcata Falcated Duck
(Falcated Teal)
Aves Anseriformes Anatidae Anas penelope Eurasian Wigeon
Aves Anseriformes Anatidae Anas americana American Wigeon None
(Baldpate)
Wildlife–Habitat Relationships in Oregon and Washington

Aves Anseriformes Anatidae Anas rubripes American Black Duck

Aves Anseriformes Anatidae Anas platyrhynchos Mallard platyrhynchos
Aves Anseriformes Anatidae Anas discors Blue-winged Teal discors
Aves Anseriformes Anatidae Anas cyanoptera Cinnamon Teal eptentrionalium
Aves Anseriformes Anatidae Anas clypeata Northern Shoveler None
Aves Anseriformes Anatidae Anas acuta Northern Pintail acuta
Aves Anseriformes Anatidae Anas querquedula Garganey
Aves Anseriformes Anatidae Anas formosa Baikal Teal
Aves Anseriformes Anatidae Anas crecca Green-winged Teal carolinensis
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Anseriformes Anatidae Aythya valisineria Canvasback

Aves Anseriformes Anatidae Aythya americana Redhead
Aves Anseriformes Anatidae Aythya collaris Ring-necked Duck None
Aves Anseriformes Anatidae Aythya fuligula Tufted Duck
Aves Anseriformes Anatidae Aythya marila Greater Scaup mariloides
Aves Anseriformes Anatidae Aythya affinis Lesser Scaup None
Aves Anseriformes Anatidae Polysticta stelleri Steller’s Eider
Aves Anseriformes Anatidae Somateria spectabilis King Eider
Aves Anseriformes Anatidae Histrionicus histrionicus Harlequin Duck pacificus
Aves Anseriformes Anatidae Melanitta perspicillata Surf Scoter None
Aves Anseriformes Anatidae Melanitta fusca White-winged Scoter deglandi
(Velvet Scoter)
Aves Anseriformes Anatidae Melanitta nigra Black Scoter americana
Aves Anseriformes Anatidae Clangula hyemalis Long-Tailed Duck epixanthum
(Oldsquaw)
Aves Anseriformes Anatidae Bucephala albeola Bufflehead None
Aves Anseriformes Anatidae Bucephala clangula Common Goldeneye americana
Aves Anseriformes Anatidae Bucephala islandica Barrow’s Goldeneye
Aves Anseriformes Anatidae Mergellus albellus Smew
Aves Anseriformes Anatidae Lophodytes cucullatus Hooded Merganser None
Aves Anseriformes Anatidae Mergus merganser Common Merganser americanus
(Goosander)
Aves Anseriformes Anatidae Mergus serrator Red-breasted Merganser serrator
Aves Anseriformes Anatidae Oxyura jamaicensis Ruddy Duck rubida
APPENDIX: SCIENTIFIC & COMMON NAMES

Aves Falconiformes Accipitridae Pandion haliaetus Osprey carolinensis

Aves Falconiformes Accipitridae Elanus leucurus White-tailed Kite majusculus
(Elanus caeruleus) (Black-Shouldered Kite)
703

Aves Falconiformes Accipitridae Haliaeetus leucocephalus Bald Eagle alascanus

Aves Falconiformes Accipitridae Circus cyaneus Northern Harrier hudsonius
Aves Falconiformes Accipitridae Accipiter striatus Sharp-shinned Hawk “velox, perobscurus”
Aves Falconiformes Accipitridae Accipiter cooperii Cooper’s Hawk None
Aves Falconiformes Accipitridae Accipiter gentilis Northern Goshawk “atricapillus, laingi”
Aves Falconiformes Accipitridae Buteo lineatus Red-shouldered Hawk elegans
Aves Falconiformes Accipitridae Buteo platypterus Broad-winged Hawk
Aves Falconiformes Accipitridae Buteo swainsoni Swainson’s Hawk None
Aves Falconiformes Accipitridae Buteo jamaicensis Red-tailed Hawk calurus
Aves Falconiformes Accipitridae Buteo regalis Ferruginous Hawk None
Aves Falconiformes Accipitridae Buteo lagopus Rough-legged Hawk sanctijohannis
Aves Falconiformes Accipitridae Aquila chrysaetos Golden Eagle canadensis
Aves Falconiformes Falconidae Falco sparverius American Kestrel sparverius
Aves Falconiformes Falconidae Falco columbarius Merlin “columbarius, suckleyi, richardsoni”
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Falconiformes Falconidae Falco rusticolus Gyrfalcon None

Aves Falconiformes Falconidae Falco peregrinus Peregrine Falcon “pealei, tundrius, anatum”
Aves Falconiformes Falconidae Falco mexicanus Prairie Falcon None
Aves Galliformes Phasianidae Alectoris chukar Chukar chukar
Aves Galliformes Phasianidae Perdix perdix Gray Partridge perdix
(Hungarian Partridge)
Aves Galliformes Phasianidae Phasianus colchicus Ring-necked Pheasant
704

(Chinese Pheasant)
Aves Galliformes Phasianidae Bonasa umbellus Ruffed Grouse “sabini, castanea, affinis, phaia”
Aves Galliformes Phasianidae Centrocercus urophasianus Greater Sage-grouse phaios
Aves Galliformes Phasianidae Falcipennis canadensis Spruce Grouse franklinii
(Dendragapus canadensis)
Aves Galliformes Phasianidae Lagopus leucurus White-tailed Ptarmigan “rainierensis, leucurus”
Aves Galliformes Phasianidae Dendragapus obscurus Blue Grouse “sierrae, fuliginosus, pallidus,
richardsonii”
Aves Galliformes Phasianidae Tympanuchus phasianellus Sharp-tailed Grouse columbianus
Aves Galliformes Phasianidae Meleagris gallopavo Wild Turkey “merriami, intermedia, silvestris”
Aves Galliformes Odontophoridae Oreortyx pictus Mountain Quail “palmeri, pictus”
Aves Galliformes Odontophoridae Callipepla squamata Scaled Quail pallida
Aves Galliformes Odontophoridae Callipepla californica California Quail “californicus, brunnescens, orecta”
(Valley Quail)
Aves Galliformes Odontophoridae Colinus virginianus Northern Bobwhite “virginianus, texanus, taylori”
Aves Gruiformes Rallidae Coturnicops noveboracensis Yellow Rail noveboracensis
Aves Gruiformes Rallidae Rallus limicola Virginia Rail limicola
Aves Gruiformes Rallidae Porzana carolina Sora None
Aves Gruiformes Rallidae Gallinula chloropus Common Moorhen
Aves Gruiformes Rallidae Fulica americana American Coot americana
Aves Gruiformes Gruidae Grus canadensis Sandhill Crane “canadensis, tabida”
Aves Charadriiformes Charadriidae Pluvialis squatarola Black-bellied Plover None
Aves Charadriiformes Charadriidae Pluvialis dominica American Golden-Plover None
(Pluvialis dominicus)
Aves Charadriiformes Charadriidae Pluvialis fulva Pacific Golden-Plover None
Wildlife–Habitat Relationships in Oregon and Washington

(Pluvialis dominica) (Lesser Golden-Plover)

Aves Charadriiformes Charadriidae Charadrius mongolus Mongolian Plover
Aves Charadriiformes Charadriidae Charadrius alexandrinus Snowy Plover nivosus
Aves Charadriiformes Charadriidae Charadrius semipalmatus Semipalmated Plover None
Aves Charadriiformes Charadriidae Charadrius melodus Piping Plover
Aves Charadriiformes Charadriidae Charadrius vociferus Killdeer vociferus
Aves Charadriiformes Charadriidae Charadrius montanus Mountain Plover
Aves Charadriiformes Charadriidae Charadrius morinellus Eurasian Dotterel
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Aves Charadriiformes Haematopodidae Haematopus bachmani Black Oystercatcher None

Aves Charadriiformes Recurvirostridae Himantopus mexicanus Black-necked Stilt None
Aves Charadriiformes Recurvirostridae Recurvirostra americana American Avocet None
Aves Charadriiformes Scolopacidae Tringa melanoleuca Greater Yellowlegs None
Aves Charadriiformes Scolopacidae Tringa flavipes Lesser Yellowlegs
Aves Charadriiformes Scolopacidae Tringa brythropus Spotted Redshank
Aves Charadriiformes Scolopacidae Tringa solitaria Solitary Sandpiper “solitaria, cinnamomea”
Aves Charadriiformes Scolopacidae Catoptrophorus semipalmatus Willet inornatus
Mammalia Didelphimorphia Didelphidae Didelphis virginiana Virginia Opossum
Mammalia Insectivora Soricidae Sorex cinereus Masked Shrew
(Commom Shrew)
Mammalia Insectivora Soricidae Sorex preblei Preble’s Shrew
Mammalia Insectivora Soricidae Sorex vagrans Vagrant Shrew
(Sorex trigonirostris) (Wandering Shrew)
Mammalia Insectivora Soricidae Sorex monticolus Montane Shrew “setosus, obscurus”
(Sorex monticola) (Dusky Shrew)
Mammalia Insectivora Soricidae Sorex bairdi Baird’s Shrew “bairdi, permiliensis”
(Sorex bairdii)
Mammalia Insectivora Soricidae Sorex sonomae Fog Shrew “sonomae, tenelliodus”
Mammalia Insectivora Soricidae Sorex pacificus Pacific Shrew “pacificus, cascadensis”
Mammalia Insectivora Soricidae Sorex palustris Water Shrew navigator
Mammalia Insectivora Soricidae Sorex bendirii Pacific Water Shrew “palmeri, bendirii, albiventer”
(Marsh Shrew)
Mammalia Insectivora Soricidae Sorex trowbridgii Trowbridge’s Shrew “trowbridgii, mariposae, destructioni”
APPENDIX: SCIENTIFIC & COMMON NAMES

Mammalia Insectivora Soricidae Sorex merriami Merriam’s Shrew merriami

Mammalia Insectivora Soricidae Sorex hoyi Pygmy Shrew washingtoni
(Microsorex hoyi)
705

Mammalia Insectivora Talpidae Neurotrichus gibbsii Shrew-mole “gibbsii, minor”

Mammalia Insectivora Talpidae Scapanus townsendii Townsend’s Mole “townsendii, olympicus”
Mammalia Insectivora Talpidae Scapanus orarius Coast Mole “oriarus, schefferi, yakimensis”
Mammalia Insectivora Talpidae Scapanus latimanus Broad-footed Mole dilatus
Mammalia Chiroptera Vespertilionidae Myotis californicus California Myotis “californicus, caurinus”
Mammalia Chiroptera Vespertilionidae Myotis ciliolabrum Western Small-footed Myotis melanorhinus
(“Myotis leibii, Myotis subulatus”)
Mammalia Chiroptera Vespertilionidae Myotis yumanensis Yuma Myotis “saturatus, sociabilis”
Mammalia Chiroptera Vespertilionidae Myotis lucifugus Little Brown Myotis “carissima, alascensis”
Mammalia Chiroptera Vespertilionidae Myotis volans Long-legged Myotis “longicrus, interior”
Mammalia Chiroptera Vespertilionidae Myotis thysanodes Fringed Myotis thysanodes
Mammalia Chiroptera Vespertilionidae Myotis keenii Keen’s Myotis keenii
Mammalia Chiroptera Vespertilionidae Myotis evotis Long-eared Myotis “evotis, pacificus”
Mammalia Chiroptera Vespertilionidae Lasionycteris noctivagans Silver-haired Bat
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Mammalia Chiroptera Vespertilionidae Pipistrellus hesperus Western Pipistrelle hesperus

Mammalia Chiroptera Vespertilionidae Eptesicus fuscus Big Brown Bat bernardinus
Mammalia Chiroptera Vespertilionidae Lasiurus cinereus Hoary Bat cinereus
Mammalia Chiroptera Vespertilionidae Euderma maculatum Spotted Bat
Mammalia Chiroptera Vespertilionidae Corynorhinus townsendii Townsend’s Big-eared Bat “Townsendii, Pallescens”
(Plecotus townsendii)
Mammalia Chiroptera Vespertilionidae Antrozous pallidus Pallid Bat “pacificus, pallidus”
706

Mammalia Chiroptera Molossidae Tadarida brasiliensis Brazilian Free-tailed Bat mexicana

Mammalia Lagomorpha Ochotonidae Ochotona princeps American Pika “brunnescens, fumosa, taylori, jewetti”
Mammalia Lagomorpha Leporidae Brachylagus idahoensis Sylvilagus idahoensis Pygmy Rabbit
Mammalia Lagomorpha Leporidae Sylvilagus bachmani Brush Rabbit “ubericolor, tehamae”
Mammalia Lagomorpha Leporidae Sylvilagus floridanus Eastern Cottontail
Mammalia Lagomorpha Leporidae Sylvilagus nuttallii Nuttall’s (Mountain) Cottontail nuttalli
Mammalia Lagomorpha Leporidae Oryctolagus cuniculus European Rabbit
Mammalia Lagomorpha Leporidae Lepus americanus Snowshoe Hare “washingtoni, klamathensis, oregonus”
Mammalia Lagomorpha Leporidae Lepus townsendii White-tailed Jackrabbit townsendii
Mammalia Lagomorpha Leporidae Lepus californicus Black-tailed Jackrabbit “californicus, wallawalla”
Mammalia Rodentia Aplodontidae Aplodontia rufa Mountain Beaver
Mammalia Rodentia Sciuridae Tamias minimus Least Chipmunk
(Eutamias minimus)
Mammalia Rodentia Sciuridae Tamias amoenus Yellow-pine Chipmunk “abiventris, amoenus, ludibundus,
(Eutamias amoenus) ochraceus, affinis, canicaudus,
luteiventris, felix, caurinus”
Mammalia Rodentia Sciuridae Tamias townsendii Townsend’s Chipmunk
(Eutamias townsendii)
Mammalia Rodentia Sciuridae Tamias senex Allen’s Chipmunk
` (Eutamias senex)
Mammalia Rodentia Sciuridae Tamias siskiyou Siskiyou Chipmunk
(Eutamias siskiyou)
Mammalia Rodentia Sciuridae Tamias ruficaudus Red-tailed Chipmunk
(Eutamias ruficaudus)
Mammalia Rodentia Sciuridae Marmota flaviventris Yellow-bellied Marmot “avara, flaviventris”
Wildlife–Habitat Relationships in Oregon and Washington

Mammalia Rodentia Sciuridae Marmota caligata Hoary Marmot

Mammalia Rodentia Sciuridae Marmota olympus Olympic Marmot
Mammalia Rodentia Sciuridae Ammospermophilus leucurus White-tailed Antelope Squirrel leucurus
Mammalia Rodentia Sciuridae Spermophilus townsendii Townsend’s Ground Squirrel “townsendii, nancyae, canus, vigillis,
(Citellus townsendii) mollis”
Mammalia Rodentia Sciuridae Spermophilus canus Merriam’s Ground Squirrel “canus, vigilis”
(“Citellus canus,
Spermophilus vigilis”)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Mammalia Rodentia Sciuridae Spermophilus mollis Piute Ground Squirrel mollis

(Citellus mollis)
Mammalia Rodentia Sciuridae Spermophilus washingtoni Washington Ground Squirrel
(Citellus washingtoni)
Mammalia Rodentia Sciuridae Spermophilus elegans Wyoming Ground Squirrel
(Citellus elegans) (Richardson’s Ground Squirrel)
Mammalia Rodentia Sciuridae Spermophilus beldingi Belding’s Ground Squirrel “oregonus, creber”
(Citellus beldingi)
Mammalia Rodentia Sciuridae Spermophilus columbianus Columbian Ground Squirrel “columbianus, ruficaudus”
(Citellus columbianus)
Mammalia Rodentia Sciuridae Spermophilus beecheyi California Ground Squirrel douglasii
(“Citellus beecheyi,
Otospermophilus beecheyi”)
Mammalia Rodentia Sciuridae Spermophilus lateralis Golden-mantled Ground Squirrel “chrysodeirus, trinitatis, connectens,
(“Citellus lateralis, trepidus, saturatus”
Callospermophilus lateralis”)
Mammalia Rodentia Sciuridae Spermophilus saturatus Cascade Golden-mantled Ground Squirrel
(“Citellus saturatus,
Callospermophilus saturatus”)
Mammalia Rodentia Sciuridae Sciurus carolinensis Eastern Gray Squirrel pennsylvanicus
Mammalia Rodentia Sciuridae Sciurus niger Eastern Fox Squirrel
Mammalia Rodentia Sciuridae Sciurus griseus Western Gray Squirrel griseus
Mammalia Rodentia Sciuridae Tamiasciurus hudsonicus Red Squirrel “richardsoni, streatori”
Mammalia Rodentia Sciuridae Tamiasciurus douglasii Douglas’ Squirrel “douglasii, mollipilosus, albolimbatus”
APPENDIX: SCIENTIFIC & COMMON NAMES

Mammalia Rodentia Sciuridae Glaucomys sabrinus Northern Flying Squirrel “oregonensis, fuliginosus, klamathensis,
bangsi, columbiensis, latipes”
Mammalia Rodentia Geomyidae Thomomys talpoides Northern Pocket Gopher “columbianus, wallowa, quadratus,
707

douglasii, immunis, limosus, shawi,

yakimensis, aequalidnes, devexus,
fuscus”
Mammalia Rodentia Geomyidae Thomomys mazama Western Pocket Gopher “helleri, niger, hesperus, oregonus,
(Mazama Pocket Gopher) mazama, nascius, couchi, glacialis, buiei,
pugatensis, tacomensis, tumuli,
yelmensis, melanops”
Mammalia Rodentia Geomyidae Thomomys bulbivorus Camas Pocket Gopher
Mammalia Rodentia Geomyidae Thomomys bottae Botta’s (Pistol River) Pocket Gopher “laticeps, detumidus, leucodon”
(Thomomys umbrinus)
Mammalia Rodentia Geomyidae Thomomys townsendii Townsend’s Pocket Gopher “townsendii, nevadensis”
Mammalia Rodentia Heteromyidae Perognathus parvus Great Basin Pocket Mouse “parvus, mollipilosus, yakimensis,
columbianus, lordi”
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Mammalia Rodentia Heteromyidae Perognathus longimembris Little Pocket Mouse nevadensis

Mammalia Rodentia Heteromyidae Microdipodops megacephalus Dark Kangaroo Mouse oregonus
Mammalia Rodentia Heteromyidae Dipodomys ordii Ord’s Kangaroo Rat columbianus
Mammalia Rodentia Heteromyidae Dipodomys microps Chisel-toothed Kangaroo Rat preblei
Mammalia Rodentia Heteromyidae Dipodomys californicus California Kangaroo Rat californicus
(Dipodomys heermanni)
Mammalia Rodentia Castoridae Castor canadensis American Beaver
708

Mammalia Rodentia Muridae Reithrodontomys megalotis Western Harvest Mouse “longicaudus, megalolits”
Mammalia Rodentia Muridae Peromyscus maniculatus Deer Mouse “rubidus, gambelii, sonoriensis,
artemisiae, alpinus, austerus”
Mammalia Rodentia Muridae Peromyscus keeni Columbian Mouse
(Peromyscus oreas) (Forest Deer Mouse)
Mammalia Rodentia Muridae Peromyscus crinitus Canyon Mouse crinitus
Mammalia Rodentia Muridae Peromyscus truei Pinon Mouse “gilberti, preblei”
Mammalia Rodentia Muridae Onychomys leucogaster Northern Grasshopper Mouse “durranti, brevicaudus”
Mammalia Rodentia Muridae Neotoma lepida Desert Woodrat nevadensis
Mammalia Rodentia Muridae Neotoma fuscipes Dusky-footed Woodrat “fuscipes, monochroura”
Mammalia Rodentia Muridae Neotoma cinerea Bushy-tailed Woodrat “fusca, aticola, occidentalis, pulla”
Mammalia Rodentia Muridae Clethrionomys gapperi Southern Red-backed Vole “idahoensis, saturatus, cascadensis,
occidentalis, caurinus, nivarius”
Mammalia Rodentia Muridae Clethrionomys californicus Western Red-backed Vole “californicus, manzama, obscurus”
(Clethrionomys occidentalis)
Mammalia Rodentia Muridae Phenacomys intermedius Heather Vole “intermedius, oramontis”
(Phenacomys olympicus)
Mammalia Rodentia Muridae Phenacomys albipes White-footed Vole
(Arborimus albipes)
Mammalia Rodentia Muridae Phenacomys longicaudus Red Tree Vole “silvicola, longicaudus”
(“Arborimus longicaudus,
Phenacmys silvicola”)
Mammalia Rodentia Muridae Microtus pennsylvanicus Meadow Vole kincaidi
Mammalia Rodentia Muridae Microtus montanus Montane Vole “montanus, nanus, micropus, canescens”
Mammalia Rodentia Muridae Microtus canicaudus Gray-tailed Vole
Wildlife–Habitat Relationships in Oregon and Washington

Mammalia Rodentia Muridae Microtus californicus California Vole eximius

Mammalia Rodentia Muridae Microtus townsendii Townsend’s Vole “townsendii, pugetti”
Mammalia Rodentia Muridae Microtus longicaudus Long-tailed Vole “abditus, angusticeps, longicaudus, halli,
macrurus”
Mammalia Rodentia Muridae Microtus oregoni Creeping Vole “adocetus, bairdii, oregoni”
Mammalia Rodentia Muridae Microtus richardsoni Water Vole “arvicoloides, macropus”
Mammalia Rodentia Muridae Lemmiscus curtatus Sagebrush Vole “pauperrimus, intermedius”
(Lagurus curtatus)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Mammalia Rodentia Muridae Ondatra zibethicus Muskrat “occipitalis, osoyoosensis”

Mammalia Rodentia Muridae Synaptomys borealis Northern Bog Lemming
Mammalia Rodentia Muridae Rattus rattus Black Rat
(Roof Rat)
Mammalia Rodentia Muridae Rattus norvegicus Norway Rat
Mammalia Rodentia Muridae Mus musculus House Mouse
Mammalia Rodentia Zapodidae Zapus princeps Western Jumping Mouse “pacificus, oregonus”
Mammalia Rodentia Zapodidae Zapus trinotatus Pacific Jumping Mouse “trinotatus, montanus”
Mammalia Rodentia Erthizontidae Erethizon dorsatum Common Porcupine
Mammalia Rodentia Myocastoridae Myocastor coypus Nutria bonariensis
Mammalia Carnivora Canidae Canis latrans Coyote “lestes, umpquensis”
Mammalia Carnivora Canidae Canis lupus Gray Wolf “fuscus, irremotus, columbianus”
Mammalia Carnivora Canidae Vulpes vulpes Red Fox “cascadensis, macroura”
(“Vulpes fulvus,Vulpes fulva”)
Mammalia Carnivora Canidae Vulpes velox Kit Fox macrotis
(Vulpex macrotis nevadensis)
Mammalia Carnivora Canidae Urocyon cinereoargenteus Gray Fox townsendi
Mammalia Carnivora Ursidae Ursus americanus Black Bear “altifrontalis, cinnamomum”
(Euarctos americanus)
Mammalia Carnivora Ursidae Ursus arctos Grizzly Bear “horribilis, arctos”
(Ursus chelan) (Brown Bear)
Mammalia Carnivora Procyonidae Bassariscus astutus Ringtail raptor
Mammalia Carnivora Procyonidae Procyon lotor Raccoon “pacificus, excelsus”
Mammalia Carnivora Mustelidae Martes americana American Marten “caurina, vulpina”
APPENDIX: SCIENTIFIC & COMMON NAMES

Mammalia Carnivora Mustelidae Martes pennanti Fisher pacific

Mammalia Carnivora Mustelidae Mustela erminea Ermine “streatori, muricus”
(Short-Tailed Weasel)
709

Mammalia Carnivora Mustelidae Mustela frenata Long-tailed Weasel

Mammalia Carnivora Mustelidae Mustela vison Mink energumenos
Mammalia Carnivora Mustelidae Gulo gulo Wolverine luscus
(Gulo luscus)
Mammalia Carnivora Mustelidae Taxidea taxus American Badger jeffersonii
Mammalia Carnivora Mustelidae Spilogale gracilis Western Spotted Skunk “saxatalis, latifrons”
Mammalia Carnivora Mustelidae Mephitis mephitis Striped Skunk “major, notata, occidentalis, spissigrada”
Mammalia Carnivora Mustelidae Lutra canadensis Northern River Otter pacifica
(Lontra canadensis)
Mammalia Carnivora Felidae Puma concolor Mountain Lion “oregonensis, californica, missoulensis”
(Felis concolor)
Mammalia Carnivora Felidae Lynx canadensis Lynx canadensis
(“Felis canadensis, Felis lynx”)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Mammalia Carnivora Felidae Lynx rufus Bobcat

(Felis rufus)
Mammalia Perissodactyla Equidae Equus asinus Wild Burro
Mammalia Perissodactyla Equidae Equus caballus Feral Horse
(Mustang)
Mammalia Artiodactyla Suidae Sus scrofa Feral Pig
(“Wild Pig, Wild Boar, Feral Swine,
710

Wild Hog, European Wild Boar”)

Mammalia Artiodactyla Cervidae Cervus elaphus roosevelti Roosevelt Elk roosevelti
(Cervus canadensis)
Mammalia Artiodactyla Cervidae Cervus elaphus nelsoni Rocky Mountain Elk nelsoni
(Cervus canadensis)
Mammalia Artiodactyla Cervidae Odocoileus hemionus columbianus Black-tailed Deer columbianus
Mammalia Artiodactyla Cervidae Odocoileus hemionus hemionus Mule Deer
Mammalia Artiodactyla Cervidae Odocoileus virginianus leucurus Columbian White-tailed Deer
Mammalia Artiodactyla Cervidae Odocoileus virginianus ochrourus White-tailed Deer (Eastside) ochrourus
Mammalia Artiodactyla Cervidae Alces alces Moose shirasi alces alces
Mammalia Artiodactyla Cervidae Rangifer tarandus Mountain Caribou caribou
(Woodland Caribou)
Mammalia Artiodactyla Antilocapridae Antilocapra americana Pronghorn Antelope americana
Mammalia Artiodactyla Bovidae Bos bison Bison athabascae
(Bison bison) (Buffalo)
Mammalia Artiodactyla Bovidae Oreamnos americanus Mountain Goat “americanus, missoulae”
Mammalia Artiodactyla Bovidae Ovis canadensis canadensis Rocky Mountain Bighorn Sheep canadensis
Mammalia Artiodactyla Bovidae Ovis canadensis californiana California Bighorn Sheep californiana
Mammalia Carnivora Otariidae Callorhinus ursinus Northern Fur Seal None
Mammalia Carnivora Otariidae Eumetopias jubatus Northern (Steller) Sea Lion None
Mammalia Carnivora Otariidae Zalophus californianus California Sea Lion californianus
Mammalia Carnivora Phocidae Phoca vitulina Harbor Seal richardsi..richardsi
(Halicyon richardsi) (“Hair Seal, Spotted Seal, Common Seal”)
Mammalia Carnivora Phocidae Mirounga angustirostris Northern Elephant Seal None
Mammalia Carnivora Mustelidae Enhydra lutris Sea Otter
Wildlife–Habitat Relationships in Oregon and Washington

Mammalia Cetacea Eschrichtiidae Eschrichtius robustus Gray Whale

(Eschrichtius glaucus) (California Gray Whale)
Mammalia Cetacea Balaenopteridae Balaenoptera acutorostrata Minke Whale
(Little Piked Whale)
Mammalia Cetacea Balaenopteridae Balaenoptera borealis Sei Whale
Mammalia Cetacea Balaenopteridae Balaenoptera musculus Blue Whale musculus
(Sulphur Bottom)
Mammalia Cetacea Balaenopteridae Balaenoptera physalus Fin Whale
(Finback Whale)
Class Order Family Scientific Name Common Name Subspecies Name
(Other Scientific Name) (Other Common Name) Oregon &Washington only

Mammalia Cetacea Balaenopteridae Megaptera novaeangliae Humpback Whale

Mammalia Cetacea Balaenidae Balaena glacialis Northern Right Whale Glacialis
(Eubalaena glacialis) (“Black Right Whale, Pacific Right Whale”)
Mammalia Cetacea Delphinidae Stenella coeruleoalba Striped Dolphin
Mammalia Cetacea Delphinidae Delphinus delphis Common Saddle-backed Dolphin
(“Common Dolphin, White-Bellied
Porpoise”)
Mammalia Cetacea Delphinidae Lagenorhynchus obliquidens Pacific White-sided Dolphin
Mammalia Cetacea Delphinidae Grampus griseus Risso’s Dolphin
(Delphinus griseus) (Gray Grampus)
Mammalia Cetacea Delphinidae Pseudorca crassidens False Killer Whale
(False Pilot Whale)
Mammalia Cetacea Delphinidae Globicephala macrorhynchus Short-finned Pilot Whale
“Globicephala macrorhyncha, (“Pilot Whale, Pothead Whale”)
Globicephala sieboldii”)
Mammalia Cetacea Delphinidae Orcinus orca Killer Whale
(Orca)
Mammalia Cetacea Delphinidae Lissodelphis borealis Northern Right-whale Dolphin
(Pacific Right Whale Porpoise)
Mammalia Cetacea Phocoenidae Phocoena phocoena Harbor Porpoise
(Common Porpoise)
Mammalia Cetacea Phocoenidae Phocoenoides dalli Dall’s Porpoise
(Dall Porpoise)
Mammalia Cetacea Ziphiidae Berardius bairdii North Pacific Bottle-nosed Whale
APPENDIX: SCIENTIFIC & COMMON NAMES

(“Baird’s Beaked Whale, Giant

Bottlenose Whale”)
Mammalia Cetacea Ziphiidae Ziphius cavirostris Goose-beaked Whale
711

(Cuvier’s Beaked Whale)

Mammalia Cetacea Ziphiidae Mesoplodon stejnegeri Bering Sea Beaked Whale
(“Stejneger’s Beaked Whale, North
Pacific Beaked Whale”)
Mammalia Cetacea Ziphiidae Mesoplodon carlhubbsi Arch-beaked Whale
(Hubb’s Beaked Whale)
Mammalia Cetacea Kogiidae Kogia breviceps Pygmy Sperm Whale
Mammalia Cetacea Physeteridae Physeter macrocephalus Sperm Whale
(Physeter catodon)
Mammalia Cetacea Monodontidae Delphinaptrus leucas Beluga Whale
(White Whale)
 Anthropogenic: Of, relating to, or resulting from the influence of
Glossary humans on nature.
Anticyclonic: Referring to an area or system of high atmospheric
Abiotic: Non-living components of an ecosystem; basic elements, and pressure having a characteristic pattern of air circulation which usually
compounds of the environment. induces settled weather conditions. Light winds flow clockwise in the
Abyssal: Pertaining to zones of great depth in the oceans or lakes into northern hemisphere and counterclockwise in the southern
which light does not penetrate; occasionally restricted to depths below hemisphere.
2,000 meters, but more usually used for depths between 4,000 and Aquatic Ecosystem: Any body of water, such as a stream, lake, or
6,000 meters. estuary, and all organisms and non-living components within it, and
Accretion: Deposition of material by sedimentation which increases functioning as a natural system.
land area. Aquatic Integrity: A mosaic of well connected, high-quality water and
Active Layer: A seasonally thawed surface layer of soil in arctic or habitats that support a diverse assemblage of native and desired non-
alpine regions that lies above permanently frozen ground and is native species, the full expression of potential life histories and
between a few centimeters and about three meters thick. taxonomic lineages, and the taxonomic and genetic diversity necessary
Adaptive Radiation: The evolutionary diversification of a taxon into a for long-term persistence and adaptation in a variable environment.
number of different forms, usually as a result of encounters with new Arboreal: Living in the canopies of trees.
resources or habitats. Archaebacteria: A taxonomic kingdom of bacteria, including sulphur-
Adaptation: A genetically determined characteristic that enhances an dependent bacteria, methane-producing bacteria, and halophilic
organism’s chances for survival and reproduction. bacteria.
Adaptive Management: An adaptive approach to management where Areas of Environmental Concern: Areas within the public lands
we use the best scientific knowledge and technologies, clearly where special management attention is required (when such areas are
recognize knowledge gaps, build shared expectations among those who developed or used or where no development is required) to protect
have a stake in ecosystem outcomes, monitor actions, and adjust and prevent irreparable damage to important natural systems or
management actions accordingly. processes, or to protect life and safety from natural hazards.
Adventive Plant: A species of plant that is not native and has been Arthropod: Invertebrate animals with a segmented body and jointed
introduced into the area but has not become permanently established. appendages, for example, spiders, bees, and crabs.
Afforestation: The establishment of forest by natural succession or by Aspect: The direction a slope faces with respect to the cardinal
the planting of trees on land where they did not grow formerly. compass points.
Albedo: A measure of surface reflectivity, usually expressed as a Association: A stable grouping of two or more plant species that
percentage, such as the proportion of solar radiation that is reflected characterize or dominate a type of biotic community.
back into space from the Earth, clouds, and atmosphere without Autecology: A subdivision of ecology that deals with the relationship of
heating the receiving surface. individuals of a species to their environment.
Alcids: Any of the Alcidae family (Order Charadriiformes) of marine Avalanche Chute: An area where periodic snow or rockslides prevent
birds having a stout bill, short wings and tail, webbed feet, a large head the establishment of forest conditions; typically shrub and herb
and heavy body, and thick, compact plumage. Confined to the northern dominated.
parts of the Northern Hemisphere, alcids include auks, guillemots, Avian: Relating to or derived from birds.
murres, and puffins. Avifauna: The birds of a specific region or period.
Alevin: A young fish, particularly a young salmon that is still attached to Barrens: A level area with poor, usually sandy or serpentine soils that is
the yolk sac. sparsely forested or unable to support normal vegetative cover and
Algae: The common name for the relatively simple type of unicellular or that generally has a low level of productivity. Barrens are frequently
multicellular plant which is never differentiated into root, stem, and dominated by specialized groups of endemic plants.
leaves, contains chlorophyll a as its photosynthetic pigment, has no true Bathymetry: The measurement of the depth of the ocean floor from
vascular system, and has no sterile layer of cells surrounding its the water surface; the oceanic equivalent of topography.
reproductive organs. Bathypelagic: Of, relating to, or living in the depths of the ocean,
Alluvial: Relating to river and stream deposits. especially in the area between about 600 and 3,000 meters deep. The
Alluvial Soil: Soil formed in material deposited by the action of running number of species and populations is relatively low in the bathypelagic
water, such as a floodplain or delta. zone, where no light source exists other than bioluminescence,
Alpine Tundra: A treeless region above the treeline of high mountains, temperature is uniformly low, and pressures are great.
characterized by cold winters and short, cool summers and having Benthic: Occurring at the bottom of a body of water, for example, a
permafrost below a surface layer that may melt in summer. seabed, riverbed, or lake bottom.
Altricial: Naked or helpless when hatched; immobile, downless, eyes Benthos: In freshwater and marine ecosystems, the collection of
closed. organisms both attached to or resting on the bottom sediments and
Alvar: A plant community dominated by mosses and herbs, occurring on burrowed into the sediments.
shallow, alkaline limestone soils. Bight: A large indentation in a coastline or continental shelf margin
Amphidromous: Referring to the migratory behavior of fishes moving forming an open bay.
from fresh water to the sea and vice versa, not for breeding purposes Bioaccumulation (Also called Biomagnification): The process by which
but occurring regularly at some stage of the life cycle (such as feeding chemical contaminants become more concentrated in the tissues of
or overwintering). organisms as they pass higher up the food chain. Heavy metals and
Amphipod: Any of a large order of small, usually aquatic crustaceans pesticides such as DDT are stored in the fatty tissues of animals and
with a laterally compressed body, for example, beach fleas. are passed along to predators of those animals. The resulting
Anadromous: Referring to the life cycle of fishes, such as salmon, in concentrations eventually reach harmful levels in predators at the top
which adults travel upriver from the sea to breed, usually returning to of the food chain.
the area where they were born. Biodiversity: The variety of organisms considered at all levels, from
Anaerobic: Referring to an environment in which oxygen is absent, or genetic variants belonging to the same species through arrays of
to a process, which occurs only in the absence of oxygen, or to an genera, families, and still higher taxonomic levels, includes the variety of
organism, which lives, is active, or occurs in the absence of oxygen, ecosystems, which comprise both the communities of organisms within
such as some yeasts or bacteria. particular habitats and the physical conditions under which they live.
Annelids: Any of a phylum (Annelida) of usually elongated, segmented Bioenvironments: Combinations of environmental factors to which
coelomate invertebrates, such as earthworms, various marine worms, the biota responds directly (e.g., temperature), or consume as
and leeches. resources (e.g., nutrients).
Anoxic: Greatly deficient in oxygen; oxygenless. Biogeochemical: The flow and interactions of biological and chemical
processes in relation to organisms and their environments.

712
 GLOSSARY 713

Biogeographic: The spatial distribution patterns of organisms in measurements taken under the canopy, as openings in the branches and
relation to changes through time (paleoecological, historical, current, crowns must be accounted for.
and future). Capability: The potential of an area of land to produce resources,
Biogeographical Region: Any geographical region characterized by supply goods and services, and allow resource uses under an assumed
distinctive flora or fauna (such as a biome or a province). set of management practices and at a given level of management
Biogeography: The science that deals with the geographical distribution intensity.
of animals and plants. Carrying Capacity: The maximum population of a given organism that
Biologic Diversity: The full variety of living organisms and their a particular environment or habitat can sustain; implies continuing yield
assemblages; the genetic variation within and between populations of without environmental damage; often denoted as K.
species, and the many processes that link organisms and their physical Catadromous: An organism, which lives in fresh water and goes to the
environments into ecological systems. sea to spawn, such as some eels.
Biomass: The total mass of all living organisms or of a particular set of Catchment: The area drained by a river or body of water.
organisms in an ecosystem or at a trophic level in a food chain; usually Cetacean: Any of an order of aquatic, mostly marine mammals that
expressed as a dry weight or as the carbon, nitrogen, or caloric include the whales, dolphins, porpoises, and related forms.
content per unit area. Chaetognaths: A group of small, active, transparent marine worms of
Biome: A major regional ecological community characterized by uncertain systemic position with horizontal lateral and caudal fins and a
distinctive life forms and principal plant or animal species, such as a row of moveable, curved spines around the mouth, for example,
tropical rain forest, tundra, grassland, or a desert. arrowworms.
Bioregion: A territory defined by a combination of biological, social, and Changed Biophysical Template: Biophysical systems that have the
geographic criteria, rather than geopolitical considerations; generally, a biotic or physical potentials of the historical range of variability (HRV),
system of related, interconnected ecosystems. but have a different composition, structure, or disturbance regime than
Biota: The plants and animals of a specific region or period, or the total present during HRV.
aggregation of organisms in the biosphere. Channelization: The straightening of rivers or streams by means of an
Biotic Community: Any assemblage of populations living in a artificial channel.
prescribed area or physical habitat; an aggregate of organisms, which Chaparral: A vegetation type dominated by shrubs and small trees,
form a distinct ecological unit. especially evergreen trees with thick, small leaves.
Biotic Climax: A climax caused by a permanent influence or Chironomids: Any of a family (Chironomidae) of midges that lack
culmination of influences caused by one or more kinds of organisms, piercing mouthparts.
including humans. Chlorofluorocarbons (Also called greenhouse gases or CFCs.): A
Biotype: A group of individuals within a population occurring in nature, group of gaseous compounds that contain carbon, chlorine, fluorine,
all with essentially the same genetic constitution. A species usually and sometimes hydrogen, and are aerosol propellants and in the
consists of many biotypes. manufacture of plastic foams.
Bivalve: A mollusk whose body is enclosed by two hinged valves or Cirque: A steep hollow, often containing a small body of water, found at
shells. the upper end of a mountain valley.
Blowdown: An extensive toppling of trees by wind within a relatively CITES: Convention on International Trade in Endangered Species of
small area, which significantly alters the small-scale climate within the Wild Fauna and Flora an agreement between 103 nations to restrict
ecosystem. international commerce involving endangered and threatened species
Boreal Forest: The circumpolar, subarctic forest of high northern of animals and plants, such as tropical birds, rhinoceros horns, orchids,
latitudes that is dominated by conifers. It is found south of the tundra and ivory.
in the Northern Hemisphere and often contains peaty or swampy Cladocerans: Any of an order (Cladocera) of minute, freshwater
areas. brachiopod crustaceans, including the water fleas.
Brackish: Water that is saline but not as salty as seawater. Clear-cut: An area where the entire stand of trees has been removed in
Braided Channel: A stream consisting of a network of interlacing small one cutting.
channels separated by bars, which may be vegetated and stable or Climate: Generalized statement of the prevailing weather conditions at
barren and unstable. a given place, based on statistics of a long period of record. Includes
Breeding Bird Survey: The North American Breeding Bird Survey seasonality of temperature and moisture.
(BBS) begun in 1966 to collect standardized data on bird populations Climax: The final stage of succession in an ecosystem. Also, a community
along more than 3,400 survey routes across the continental United that reached a steady state under a particular set of environmental
States and southern Canada for more than 250 species. conditions.
Broad Scale: Encompassing a wide area. Close-crowned: Descriptive of crowded forests where closely spaced
Broader Geographic Scales: (See landscape). Watershed, river basin, trees have tops that touch or overlap.
or other physiographic region suitable for analyzing management Cluster Analysis: A method grouping those variables within a set of
proposals relative to other proposals or activities; something larger variables that are highly correlated and excluding from clusters those
than a stand, soil mapping unit, local landform, lake or stream. that are negatively correlated or uncorrelated.
Brood Parasitism: (Also called nest parasitism or breeding parasitism.) Coarse Woody Debris (CWD): Portion of a tree that has fallen or
The laying of eggs by one bird species in the nest of another bird been cut and left in the woods. Usually refers to pieces at least 20
species and the subsequent brooding of the egg and raising of the inches in diameter.
young by the parasitized host, usually to the detriment to the host’s Cohort: Individuals all resulting from the same birth-pulse, and thus all
young. of the same age.
Bunchgrass: Any of several grasses, especially of the western United Commensal: Referring to the relationship between two kinds of
States that grow in tufts rather than forming turf, for example, the organisms in which one obtains food or other benefits from the other
genus Andropogon. without damaging or benefiting it.
Bycatch: Nontarget organisms that are caught in fishing or other Community: Any grouping of populations of different organisms that
harvest operations and are usually discarded. live together in a particular environment.
Calcareous: Consisting of or containing calcium carbonate; a soil rich in Connectivity: Condition, in which the spatial arrangement of land cover
calcium salts, derived from limestone or chalk. Also, an organism which types allows organisms and ecological processes (such as disturbance)
has an affinity for such an alkaline or basic soil. to move across the landscape. Connectivity is the opposite of
Candidate Species: A species being considered for listing as a federally fragmentation.
endangered or threatened species. Conservation Biology: The body of knowledge that deals with the
Canopy: A layer of foliage in a forest stand; most often refers to the careful protection, utilization, and planned management of living
uppermost layer of foliage. organisms and their vital processes to prevent their depletion,
Canopy Closure: The degree to which the canopy blocks sunlight or exploitation, destruction, or waste.
obscures the sky. It can only be accurately determined from
 714 Wildlife–Habitat Relationships in Oregon and Washington

Conservation Strategy: A management plan for a species, group of management-induced disturbance stresses operate during and
species, or ecosystem that prescribes standards and guidelines that if immediately after drought. The term is used particularly for plant
implemented provide a high likelihood that the species, groups of species on rangelands.
species, or ecosystem, with its full complement of species and Defoliators: Insects that feed on foliage and act to remove some or all
processes, will continue to exist well-distributed throughout a planning of the foliage from a tree shrub, or herb.
area, i.e., a viable population. Degradation: The breaking down of a substance into smaller or simpler
Conspecific: Relating to the same species. parts, usually by erosion.
Continental Shelf: The shallow, gradually sloping seabed around a Delta: An alluvial deposit at the mouth of a river or tidal inlet. Deltas
continental margin not usually deeper than 200 meters and formed by occur when a sediment-laden current enters an open body of water, at
submergence of part of a continent. which point there is a reduction in the velocity of the current, resulting
Continentality: Tendency of large land areas in mid-latitude and high in rapid deposition of the sediment, as at the mouth of a river where
latitudes to impose a large annual temperature range on the air the river discharges into the sea or a lake.
temperature cycle. Demersal: Living at or near the sea floor but having the capacity for
Copepods: Any of a large subclass (Copepoda) of usually minute active swimming.
freshwater and marine crustaceans that form an important element of Demography: The quantitative analysis of population structure and
the plankton in the marine environment and in some fresh waters. trends; population dynamics.
Corridor: A more or less continuous connection between landmasses Dendrochronology: The science of dating events and variations in the
or habitats; a migration route that allows more of less uninhibited environment by the comparative study of annual growth rings of trees.
migration of most of the animals of one faunal region to another. In Desertification: The process by which an area or region becomes
terms of conservation biology, a connection between habitat fragments more arid through loss of soil and vegetative cover. The process is
in a fragmented landscape. often accelerated by excessive continuous overstocking and drought.
Corridors:The landscape elements that connect similar patches through Detritus: Debris or waste material, usually organic, such as dead or
a dissimilar matrix or aggregation of patches. partially decayed plants and animals, often important as a source of
Corvids: A family of birds that includes the crows, ravens, jays, and nutrients; or, small particles of minerals from weathered rock, such as
magpies. sand or silt.
Cover: Vegetation used by wildlife for protection from predators, to Dewatering/Dewater: The removal of water from a stream/river
mitigate weather conditions, or to reproduce. May also refer to the network, typically for irrigation, industrial, or human use; commonly
protection of soil and the shading provided to herbs and forbs by changes a network that developed by concentrating flows from stream/
vegetation. river branches to main-stems to main-stems branching to canals, which
Crevasse: A breach in a levee along the bank of a river through which reduces the flow in the main-stems.
floodwater may flow and produce sheet-like deposits of gravel or Diel: A 24-hour period, usually encompassing one day and one night.
sandy sediment; or a large, open fissure forming in a glacier as it moves Dinoflagellates: Any of an order (Dinoflagellata) of chiefly marine,
and is deformed. planktonic, usually solitary phytoflagellates (which have many
Critical Habitat: Under the Endangered Species Act, critical habitat is characteristics in common with algae) that includes luminescent forms,
defined as the specific areas within the geographic area occupied by a forms important in marine food chains, and forms causing red tides.
federally listed species on which are found physical and biological Disjunct: Distinctly separate; a discontinuous range in which one or
features essential to the conservation of the species, and that may more populations are separated from other potentially interbreeding
require special management considerations or protection, and specific populations by a sufficient distance to preclude gene flow between
areas outside the geographic area occupied by a listed species, when it them.
is determined that such areas are essential for the conservation of the Dispersal: The movement, usually one-way and on any time scale, of
species. plants or animals form their point of origin to another location where
Crown Fires: Fires that spread from tree crown to tree crown, usually they subsequently produce offspring.
indicative of particularly hot fires in dry conditions. Distributary: A river branch flowing away from the main stream.
Crustacean: Any of a large class (Crustacea) of mostly aquatic Disturbance: An effect of a planned human management activity, or
mandibulate arthropods that have a chitinous of calcareous and unplanned native or exotic agent or event that changes the state of a
chitinous exoskeleton, a pair of often modified appendages on each landscape element, landscape pattern, or regional composition.
segment, and two pairs of antennae; includes lobsters, shrimps, crabs, Disturbance Severity Classes:
wood lice, water fleas, and barnacles. Lethal: Disturbance that causes morality to most of the upper layers
Cultivar: A variety of a plant produced and maintained by horticultural of vegetation and changes the structure.
techniques and not normally found in wild populations. Non-lethal: Disturbance does not cause mortality to a substantial
Cyanobacteria: A large and varied group of bacteria which possess portion of the upper layer, removes susceptible individuals from all
chlorophyll a and which carry out photosynthesis in the presence of layers, and maintains the structure.
light and air, producing oxygen. They were formerly regarded as algae Mixed: Disturbance causes a fine-scale mosaic of lethal and non-lethal
and were called “blue-green” algae. The group is very old and is effects that result in clump/gap mosaics of changed and maintained or
believed to have been the first oxygen-producing organisms on Earth. unaffected structure.
Cyclonic: Referring to a region of low atmospheric sea level pressure; Disturbance Regime: The pattern of intervals between disturbance
or, the wind system around such a low-pressure center that has a and severity of disturbance. For landscapes, this can be for a given
clockwise rotation in the Northern Hemisphere and a disturbance, such as fire, or for a complex of disturbances.
counterclockwise rotation in the Southern Hemisphere. Diurnal: Occurring or active only in daylight.
Debouch: To emerge or issue; often used in reference to rivers or Diversity: The distribution and abundance of different plant and animal
streams. communities and species within the area covered by a land and
Debris torrent: A flood of debris (branches, shrubs, rocks, mud, and so resource management plan.
forth) and water rushing down a stream channel, caused by excessive Doliolids: Any of a small family of oceanic tunicates.
rainfall or snowmelt. Debris torrents have a significant scouring effect Downwelling: The downward movement of surface waters caused by
on the stream ecosystem. the convergence of different water masses or where surface waters
Deciduous: Plants having structures that are shed at regular intervals or flow toward the coast.
at a given stage in development, such as trees that shed their leaves Drawdown: A lowering of the water level in a reservoir or other body
seasonally. of water.
Decreaser Species: plant species of the potential vegetation that will Echinoderms: Any of a phylum (Echinodermata) of radially symmetrical
decrease in relative amount when management-induced disturbance coelomate marine animals including the starfishes, sea urchins, and
stresses (such as excessive livestock grazing pressure or alteration of related forms.
fire frequency and severity) operate during drought and when
 GLOSSARY 715

Ecological Approach: Natural resource planning and management Ecosystem Structure: The physical arrangement of the various
activities that assure consideration of the relationship between all components. Also, trophic structure; measured in standing crop or
organisms (including humans) and their environment. energy fixed per unit area per unit time. May be pyramids of numbers,
Ecological Disequilibria/Disequilibrium: A system that has unequal biomass, or energy flows.
relationships of inputs and outputs that result in erratic (and Ecosystem Viability: The ability to maintain diversity, productivity,
unpredictable) successional patterns and associated responses to resilience to stress health, renewability, and/or yields of desired values,
disturbance. resource used, products, or services from an ecosystem while
Ecological Element: The individual constituent of the whole. For maintaining the integrity of the ecosystem over time.
example: vegetation patch, stream reach, road, city site, or large snag. Ecosystems Approach: The ecosystem approach embodies three
Ecological Function: The activity or role performed by an organism or fundamental concepts: designating the physical boundary of the system
element in relation to other organisms, element in relation to other and its parts; understanding the interactions of the parts as a
organisms, elements, or the environment. functioning whole; and understanding the relation between the system
Ecological Integrity: The maintenance of native and desired non-native and its context (external factors that influence the system and also
species and associated processes. internal information that must be synthesized to be understood at the
Ecological Predictability: Causal agents of disturbance (such as fire, scale of the defined system).
erosion, floods, grazing, harvest, predation, insects, and pathogens) Ecotone: The boundary or transitional zone between adjacent
produce “expected” effects that are consistent with the limitations of communities containing the characteristic species of each, such as the
the biophysical system and inherent disturbance regimes (based on edge of a woodland next to a field or lawn.
native historical range of variability, or altered regime, as appropriate). Ecotype: A locally adapted population of a species which has a
Ecological Process: A series of actions, changes, or functions that distinctive limit of tolerance to environmental factors; a genetically
produce a resulting condition for biota, elements, or the environment. uniform population of a species resulting from natural selection by the
For example succession, decay, photosynthesis, food chain, fire, drought, special conditions of a particular habitat factors.
or flood. Ecotypic Variation: The variation within a species that is adapted to
Ecological Resiliency: Ability of system biota and their environments the variety of different habitats across the range of the species.
to renew the cycle of functions and processes following disturbance. At Ectotherm: A cold-blooded animal, one having a body temperature
a landscape scale the focus is on connections of the hydrologic and determined primarily by the temperature of its surrounding
land system, carbon-nutrient system, food web, and evolutionary environment. Terrestrial reptiles are ectotherms.
systems and their aggregate tendency towards equilibrium or Edaphic: Pertaining to soil or to the physical, chemical, and biological
disequilibrium in response to disturbance. properties of the soil or substratum, which influence associated biota,
Ecological Simplification: Loss of inherent patterns of elements (such such as pH and organic matter content.
as native species, large trees, bunchgrasses, soil, or channel stability), Edge Effect: The tendency for a transitional zone between communities
mosaics (such as vegetation patch patterns, migration routes, or (an ecotone) to contain a greater variety of species and more dense
connected stream pools and runs), and succession/disturbance regimes populations of species than either community surrounding it.
(such as frequent non-lethal fire to infrequent lethal fire or high El Niño (Also called El Niño—Southern Oscillation Event, or ENSO): A
sinuosity streams to low sinuosity). Simplification results in loss of warm water current which periodically flows southward along the
resiliency and associated predictability (or reliability) of system coast of Ecuador, associated with the southern oscillation in the
response. atmosphere, and which affects climate throughout the Pacific region.
Ecological Site: A specific location on the land, that is representative of Elasmobranches: Any of a subclass (Elasmobranchii) of cartilaginous
an ecological type. fishes that have five to seven later gill openings on each side,
Ecological Succession: The chronological sequence of vegetation and comprising sharks, rays, skates, and extinct related fishes.
associated animals in an area; or, continuous colonization, extinction, Electrophoresis: A technique for separating mixtures of organic
and replacement of species’ populations at a particular site, due either molecules based on their different rates of travel in electric fields.
to environmental changes or to the intrinsic properties of the plants Emergent: An aquatic plant having most of its vegetative parts above
and animals. water. Also, a tree which reaches or exceeds the level of the
Ecological Type: A category of land having a unique combination of surrounding canopy.
potential natural community, soil, landscape features, climate, and Empirical: Originating in or based upon observation or experience;
differing from other ecological types in its ability to produce vegetation capable of being verified or disproved by observation or experiment.
and respond to management. Encroachment: Conditions where the succession/disturbance regimes
Ecology: The relationship of species, including humans, and their have been changed to allow transition to dominance by species or
environment. structures that are not adapted to the biophysical succession/
Ecoregion: A continuous geographic area in which the environmental disturbance regime.
complex, produced by climate, topography, and soil, is sufficiently Endemic: Belonging or native to a particular people or geographic
uniform to develop characteristics of potential major vegetation region; a genetically unique life form.
communities. Endotherm: A warm-blooded animal, one that maintains a body
Ecosystem: A community of organisms and their physical environment temperature largely independent of the temperature of the
that interact as an ecological unit. environment. Mammals are endothermic.
Ecosystem Degradation: Reductions in ecosystem sustainability Environment: The complex of climatic, soil and biotic factors that act
because of natural or human effects. upon an organism or ecological community and ultimately determine
Ecosystem Function (Processes): The major processes of its form and survival.
ecosystems that regulate or influence the structure, composition, and Eolian: Pertaining to the action or effect of the wind; wind-borne..
pattern. These include nutrient cycles, energy flows, trophic levels (food Ephemeral streams: Streams that contain running water only
chains), diversity patterns in time/space development and evolution, sporadically, such as during and following storm events.
cybernetics (control), hydrologic cycles and weathering processes. Epipelagic: The oceanic zone extending from the surface to about 200
Ecosystem-based Management: The careful and skillful integration of meters, where enough light penetrates to allow photosynthesis.
ecological, economic, social, and managerial principles to conserve, Epiphyte: A plant that uses another plant (usually a tree) for support or
enhance, and restore ecosystems (including their functions, processes, anchorage but not for water or nutrients.
constituent species, and productive capacities) to maintain their long- Epizootic: An outbreak of disease (an epidemic) in nonhuman animals,
term viability and integrity while seeking desired conditions for uses, or pertaining to such an outbreak.
products, values, and services. Equilibria/Equilibrium: A system that has cyclic successional patterns
Ecosystem Pattern: The structure that results from the distribution of or multiple stable states, and associated response in disturbances.
organisms in, and their interaction with their environment. Includes Ericaceous: Of, relating to, or being a heath or of the heath family of
zonation, stratification, activity, or periodicity, food-webs, reproductive, plants, which are mostly shrubby, dicotyledonous, and often evergreen
social, and stochastic. plants that thrive on open, barren soil that is usually acidic and poorly
drained.
 716 Wildlife–Habitat Relationships in Oregon and Washington

Escapement: The number of fish that are permitted to survive and Guild: A group of species having similar ecological resource
spawn (as by adjustment of fishing season or by provision of fishways). requirements and foraging strategies and therefore having similar roles
Estivation: Dormancy associated with warm/dry period of the year. in the community.
Estuary: A semi-enclosed coastal body of water which has a free Gymnosperm: A plant, such as a cycad or a conifer, whose seeds are
connection with the open sea and where fresh water derived from not enclosed in an ovary (fruit).
land drainage (usually mouths of rivers) is mixed with seawater; often Gyre: A circular or spiral system of movement, especially a giant circular
subject to tidal action and cyclic fluctuations in salinity. oceanic surface current.
Euryhaline: Able to live in waters with a wide range of salinity. Habitat: The place, including physical and biotic conditions, where a
Eutrophication: The process by which a body of water acquires a high plant or an animal usually occurs.
concentration of nutrients, especially phosphates and nitrates, which Habitat Connections: A network of habitat patches linked by areas of
typically promote excessive growths of algae, decomposition of which like habitat. The linkages connect habitat areas within the watershed to
depletes oxygen, causing the death of other organisms. each other and to areas outside the watershed. These connections
Evapotranspiration: Loss of water from the soil both by evaporation include riparian areas, mid-slopes, and ridges. In the case of old-growth
and by transpiration from plants. forest habitat connections, each connection is planned to be sufficiently
Excessive livestock grazing pressure: Grazing pressure that results wide (at least 1,000 feet) to retain interior old-growth associated
in a decline in physiological vigor of plants, typically observed as a species.
decline in reproductive output (for example, tiller production of Habitat Fragmentation: The breaking up of a habitat into
grasses) and below ground (for example, root growth) growth. unconnected patches interspersed with other habitat, which may not
Exotic Species: Species which occur in a given place, area, or region as be inhabitable by species occupying the habitat that was broken up. The
the result of direct or indirect, deliberate or accidental introduction of breaking up is usually by human action, as, for example, the clearing of
the species by humans, and for which introduction has permitted the forest or grassland for agriculture, residential development, or overland
species to cross a natural barrier to dispersal. electrical lines.
Extinction: The dying out of a species, or the condition of having no Habitat Type: Place where an animal or plant normally lives, often
remaining living members; also, the process of bringing about such a characterized by a dominant plant form or physical characteristic.
condition. Halophytic: Referring to a plant that can tolerate or thrive in alkaline
Extirpation: The loss or removal of a species from one or more specific soil rich in sodium or calcium salts; tolerant of saline conditions.
areas but not from all areas. Hard Mast: Fruit of hardwood trees such as beech and oaks.
Fauna: The animal life of a region or geological period. Heavy Grazing: A comparative term, which indicates that the stocking
Fecundity: The potential reproductive capacity of an organism or rate of a pasture is relatively greater than that of other pastures. Often
population. erroneously used to mean overuse.
Fen: A marshy, low-lying wetland covered by shallow, usually stagnant, Heavy Metals: A metallic element of high specific gravity, such as
and often alkaline water that originates from groundwater sources. antimony, bismuth, cadmium, copper, gold, lead, mercury, nickel, silver,
Feral: Relating to plants or animals which have escaped from tin, and zinc. These metals, which are toxic even in low concentrations,
domestication, and to their descendants. persist in the environment and can accumulate to levels that stunt
Fetch: The distance along open water or land over which the wind plant growth and interfere with animal life.
blows; the distance traversed by waves without obstruction. Hectare (ha): A metric unit of measure for area, equal to 2.47 acres.
Fire regime: The characteristic frequency, extent, intensity, severity, and Hematocrit: The percentage of blood volume occupied by red blood
seasonality of fires in an ecosystem. cells.
Flora: Plant or bacterial life forms of a region or geological period. Hermaphroditic: An individual that possesses both male and female
Fluvial: Pertaining to rivers or streams and their action. sex organs.
Food Web: The interlocking pattern of food chains in an ecosystem. A Herptiles: Reptiles and amphibians collectively.
food chain is a transfer of food energy from plants through a series of Heterogeneity: Variation in the environment over space and time.
animals. Heterogeneous: Consisting of diverse or dissimilar parts; having non-
Forb: An herbaceous plant which is not a grass. uniform structure or composition.
Fragmentation: Breaking up of contiguous areas into progressively Heterotrophic: An organism that is unable to manufacture its own
smaller patches of increasing degrees of isolation. food from simple chemical compounds and therefore consumes other
Gallery Forest: A narrow strip of forest along the margins of a river in organisms, living or dead, as its main source of carbon.
an otherwise unwooded landscape. Heterozygous: Having two different alleles at a particular gene locus
Gap Analysis: The process of identifying and classifying components of on a chromosome pair. Provides a measure of genetic variation either
biological diversity to determine which components already occur in in a population or in an individual.
protected areas and which are not present or are under-represented in Hibernation: Dormancy associated with cold period of the year.
protected areas. Historic: The approximate 1,000-year time period prior to Euro-
Gastropod: Any of a large class (Gastropoda) of mollusks, usually with a American settlement (substantial effects in Oregon and Washington
univalve shell or no shell and a distinct head bearing sensory organs, assumed to have begun by the mid 1800s).
such as snails and slugs. Holocene: The present, post-Pleistocene geologic epoch of the
Geographic Information System (GIS): A spatial type of Quaternary period, including the last 10,000 years; the most recent
information management system, which provides for the entry, storage, postglacial period.
manipulation, retrieval, and display of spatially oriented data. Home Range: The geographic area within which an animal restricts its
Geomorphology: The study of landforms on a plant’s surface and of normal, daily activities.
the processes that have fashioned them. Homeothermy: Of being warm blooded, able to maintain body
Glade: An open space in the forest. temperature above that of the surroundings despite large variation in
Graminoids: Grasses and grass like plants, such as sedges. environmental temperature.
Gravid: Carrying eggs or young; pregnant. Human Dimension: An integral component of Ecosystem Management
Grazing System: A specialization of grazing management that defines that recognizes people are part of ecosystems, that people’s pursuits of
the periods of grazing and non-grazing. past, present, and future desires, needs and values have and will
Greenhouse Effect: Heating of the Earth’s atmosphere that is loosely continue to influence ecosystems and must be included in ecosystem
analogous to the glass of a greenhouse letting light in but not letting management.
heat out. Hybridization: Any crossing of individuals of different genetic
Gregarious: Tending to form into groups, which possess a social composition, often belonging to separate species, resulting in hybrid
organization, such as schools of fish, herds of mammals, flocks of birds. offspring.
Groundfish: A bottom-dwelling fish, especially one of commercial Hydric: Characterized by, relating to, or requiring an abundance of
importance such as cod, haddock, pollock, or flounder. moisture.
 GLOSSARY 717

Hydrocarbon: A naturally occurring organic compound that contains Landscape: A spatially heterogeneous area with repeating patterns of
carbon and hydrogen; may be gaseous, solid, or liquid, for example, elements and associated disturbance regimes, with similar climate and
natural gas, bitumens, and petroleum. geomorphology.
Hydrographic: Relating to the characteristic features of bodies of Landscape Connectivity: The spatial contiguity with in the landscape:
water, such as depth and flow. a measure of how easy or difficult it is for organisms to move through
Hydrological cycle: The movement of water from the sea through the the landscape without crossing habitat barriers.
air to the land and back to the sea. Landscape Contrast: The degree to which adjacent landscape
Hydrology: The study of the movement of water from the sea through elements differ from each other, with respect to species composition
the air to the land and back to the sea; the properties, distribution, and and physical attributes.
circulation of water on or below the Earth’s surface and in the Landscape Ecology: The relationships of structure, function, and
atmosphere. change in a heterogeneous land area composed of interacting
Hydromorphic: Descriptive of an intra-zonal soil formed under ecosystems. Structure, function, and change refer to the patterns and
waterlogged or poorly drained conditions. processes of terrestrial, aquatic, hydrologic, social, and economic
Hydroperiod: The duration and frequency of flooding. systems across space and through time.
Hyporheic Zone: The area under the stream channel and floodplain Landscape Edge: The interface between landscape elements of
that contributes to the stream. different composition and structure, for example between an open
Hypoxic: Deficient in oxygen. clear-cut and a closed-canopy forest.
Impoundment. A natural or artificial body of water held back by a Landscape Grain: The average size of landscape elements: the
dam. “texture” of the landscape.
Increaser: Plant species that will increase in relative amount when Landscape Heterogeneity: The variation in aggregations of landscape
management induced disturbance stresses (such as excessive livestock elements across a landscape.
grazing pressure or alteration of fire frequency and severity) operate Landscape Matrix: The most concentrated portion of the landscape,
and when management-induced disturbance stresses operate during that is, the vegetation type that is most contiguous.
and immediately after drought. The term is used particularly for plant Landscape Patches: Areas of vegetation which are relatively
species on rangelands. homogeneous internally (with respect to composition, successional
Indicator Species: An organism whose presence or state of health is stage, etc.) but differ from what surrounds them (the matrix, or other
used to identify a specific type of biotic community or as a measure of patches).
ecological conditions or changes occurring in the environment. Landscape Patchiness: The density of all types of patches within a
Indigenous: A species that occurs naturally in an area; native. landscape; considers the diversity among patches.
Integrated Pest Management: A pest management philosophy based Landscape Porosity: The density of a particular type of patch within a
on an understanding of forest growth and development, forest pest matrix.
dynamics, and the interaction of the two. Landscape Stability: The likelihood a landscape structural element will
Integrated Resources Management: The simultaneous consideration change significantly (in composition, physical features, etc.) over time,
of ecological, physical, economic, and social aspects of lands, waters, and and the rate of that change.
resources in developing and carrying multiple-use, sustained-yield Landscape Unit: A continuous geographic area with fairly consistent
management. landform, and vegetation communities.
Intermediate Host: The host occupied by juvenile stages of a parasite Landscape Use by Wildlife:
prior to the definitive host and in which asexual reproduction often Contrast: Species requires the contrast between two major structural
occurs. stages; uses two major structural stages in close proximity. Typically
Intermittent Stream: Any non-permanent flowing drainage feature forages in open stages and breed/rests in large tree structural stages.
having a definable channel and evidence of scour or deposition. This Generalist: Species use all or many patch types. Patches not
includes what are sometimes referred to as ephemeral streams if they restricted to one structural stage.
meet these two criteria. Mosaic: Species uses aggregates of patches of habitat.
Intertidal: Relating to the littoral zone above the low-tide mark. Patch: Species likely uses only one homogeneous patch during life
Invertebrate: An animal without a backbone, such as snails, worms, and cycle (for residents) or breeding/wintering period (for migrants).
insects. Large Woody Debris (LWD): see Coarse Woody Debris (CWD).
Invertivore: An animal or plant that eats invertebrate animals. Larva (Larvae): The wingless and often wormlike hatchlings of insects;
Isobath: A line on a map or chart that connects all points having the also, the early form of an animal (such as a frog or sea urchin) which at
same depth below the surface of a body of water; also, having constant birth or hatching is fundamentally unlike its parent and must
depth. metamorphose before assuming adult characteristics.
Isopod: Any of a large order of sessile-eyed crustaceans with the body Leaching: The removal of readily soluble components, such as chlorides,
composed of seven free thoracic segments, each bearing s similar pair sulfates, organic matter, and carbonates, from soil by percolating water.
of legs. The remaining upper layer of leached soil becomes increasingly acidic
Isotherm: A line on a map or chart of the Earth’s surface connecting and deficient in plant nutrients.
points having the same temperature at a given time or the same mean Legacy Tree (also called Live Remnant Tree): A live tree remaining from
temperature for a given period. the previous stand. Does not necessarily have to be a mature or old
Iteroparous: Repeat spawning strategy typical of the rainbow and growth tree.
cutthroat trout. Lek: A mating system among birds during which males display
Karst: A limestone landscape that is characterized by sinks, underground communally at a traditional site (one used year after year).
streams, and caverns. Lentic: Related to still waters such as ponds lakes, or swamps.
Keystone Species: Organisms that play dominant roles in an ecosystem Levee: A raised embankment along the edge of a river channel, often
and affect many other organisms. The removal of a keystone predator constructed as protection against flooding. Natural levees result from
from an ecosystem causes a reduction of the species diversity among periodic overbank flooding, when coarser sediment is immediately
its former prey. deposited because of a reduction in river velocity.
Krill: Planktonic crustaceans and larvae that constitute the primary food Lichen: A composite organism consisting of a fungus and algae or
of baleen whales. cyanobacteria living in symbiotic association.
Krummholz: A discontinuous belt of stunted forest or scrub typical of Life History: The significant features of the life cycle through which an
windswept alpine regions close to treeline; a wind-deformed tree at organism passes, with particular reference to strategies influencing
high elevations. survival and reproduction.
Lacustrine: Pertaining to or living in lakes or ponds. Limnic: Pertaining to lakes or to other bodies of standing fresh water;
Lagoon: A shallow water body that is near or connected to a larger often used with reference only to the open water of a lake away from
body of water. the bottom; limnetic.
 718 Wildlife–Habitat Relationships in Oregon and Washington

Linkages: Route that permits movement of individual plant (by Mode: Value occurring most frequently in a series of observations.
dispersal) and animals from a Landscape Unit and/or habitat type to Model: An idealized representation or reality developed to describe,
another similar Landscape Unit and/or habitat type. analyze, or understand the behavior of some aspect of it; a
List of Endangered or Threatened Species: A listing of animals and mathematical representation of the relationships under study. The term
plants administratively determined to meet legal criteria for protection model is applicable to a broad class of representation, ranging from a
under provisions of the U.S. Endangered Species Act. relatively simple qualitative description of a system or organization to a
Littoral Zone: The biogeographic zone in a body of fresh water where highly abstract set of mathematical equations.
light penetration is sufficient for the growth of plants; the intertidal Mollusk: An organism in the phylum Mollusca (for example, snails, clams,
zone of the seashore. or squids), characterized by soft, un-segmented body parts enclosed in
Loess: Unconsolidated sediment deposited by wind. Loess is usually a shell.
composed of unstratified fine sand or silt. Monitoring: A process of collecting information to evaluate whether
Lotic: Relating to or living in moving water, such as a river or stream. objectives of a management plan are being realized.
Macroclimate: Climate that lies just beyond the modifying irregularities Montane: Of, relating to, growing in, or being the biogeographical zone
of landform and vegetation. of relatively moist, cool upland slopes below the timberline, often
Macrofauna: Animals large enough to be seen with the naked eye. dominated by large coniferous trees.
Management Disturbances: Intentional, planned human disturbance Moraine: An accumulation of boulders, stones, or other debris carried
that changes the structure and composition of a landscape element, and deposited by a glacier.
landscape pattern, or regional composition, such as timber harvest, Morphology: The form and structure of organisms.
thinning, range improvements, livestock grazing, prescribed fire planned Mosaic: Heterogeneous ecological conditions on a landscape usually
ignition, fire suppression, etc. produced by the variable, patchy effects of disturbances; a patchwork of
Management Region: The collective delineation of land that are vegetation communities within a landscape as determined by
modeled with similar assumptions relative to management objectives. environmental conditions.
Marine Protected Areas (MPAs): Any area of intertidal or subtidal Mustelid: One of a large, widely distributed family of small, lithe,
terrain, together with its overlying water and associated flora, fauna, carnivorous mammals, including weasels, otters, skunks, wolverines, and
historical, and cultural features, which has been reserved by law or minks.
other effective means to protect part or all of the enclosed Mutagen: Any agent that produces a mutation or enhances the rate of
environment (IUCN 1988). mutation in an organism, for example, x-rays, gamma rays, and certain
Marsh: An ecosystem of more or less continuously waterlogged soil chemicals.
dominated by emersed herbaceous plants but without a surface Mutualism: An interaction between members of two species, which
accumulation of peat. A marsh differs from a swamp in that it is benefits both; in strict terms, obligatory mutualism, in which neither
dominated by rushes, reeds, cattails, and sedges, with few if any woody species can survive under natural conditions without the other.
plants, and differs from a bog in having soil rather than peat at its base. Mycorrhizae: The mutually beneficial association between a fungus and
Matrix: The most extensive and most connected landscape element the roots of a plant; a mycorrhizal root takes up nutrients more
type present, which plays the dominant role in landscape functioning. efficiently than an uninfected root.
Also, a landscape element surrounding a patch. National Environmental Policy Act (NEPA): An act which
Mature Forest: Generally used in an economic sense to indicate that a encourages productive and enjoyable harmony between man and his
forest has attained harvest age. environment; promotes efforts to prevent or eliminate damage to the
Maximum Sustainable Yield: The maximum yield or crop which may environment and biosphere and stimulate the health and welfare of
be harvested year after year without damage to the system, or the man; enriches the understanding of the ecological system and natural
theoretical point at which the size of a population is such as to resources important to the nation; and establishes a Council on
produce a maximum rate of increase. Environmental Quality.
Megafauna: The largest size category of animals in a community. National Forest Management Act (NFMA): A law passed in 1976
Meiofauna: That part of the microfauna, which inhabits algae, rock as amendments to the Forest and Rangeland Renewable Resources
fissures, and superficial layers of the muddy sea bottom. They are Planning Act that requires the preparation of regulations to guide that
smaller than 1 millimeter but larger than 0.1 millimeter. development.
Melanism: A condition in which dark pigment produces dark color or Native: Plants or animals that are indigenous to a given place; the pre-
blackness in scales, skin, or plumage. Euro-American settlement system.
Meristems: The undifferentiated, growing parts of plants, consisting of Natural Conditions: Plant and animal communities where people have
groups of cells capable of actively dividing. not directly impacted either the plant community or the soil by such
Mesic: Neither wet (hydric) nor dry (xeric); intermediate in moisture, activities as logging, grazing, or cultivation.
without extremes. Natural Variability: Range of the spatial, structural, compositional, and
Mesopelagic: The ocean zone from 200 to 1,000 meters deep, where temporal characteristics of ecosystem elements during a period
little light penetrates and the temperature gradient is even and gradual specified to represent “natural” conditions.
with little seasonal variation. This zone contains an oxygen minimum Nekton (Also spelled necton): Free-swimming organisms in aquatic
layer and usually has the maximum concentrations of the nutrients ecosystems; unlike plankton, they are able to navigate at will (such as
nitrate and phosphate. It overlies the bathypelagic zone and is overlain fishes, amphibians, and large swimming insects).
by the epipelagic zone. Nematode: Any of a phylum (Nematoda or Nemata) of elongated
Metabolite: A product of metabolism or a substance that is essential to cylindrical worms parasitic in animals or plants, or free-living in soil or
the metabolism of an organism or to a metabolic process. water.
Metapopulation: A group of populations, usually of the same species, Nemerteans: Any of a phylum (Nemertea) of often vividly colored
which exist at the same time but in different places. marine worms, most of which burrow in the mud or sand along
Microclimate: The climate that prevails in a small area, usually in the seacoast; often called ribbon worms.
layer near the ground. Neotenic: Referring to an organism, which has attained sexual maturity
Midden: A heap of refuse. Also, a pile of seeds or of various items that while retaining juvenile characteristics.
were gathered by a rodent, for example, by a squirrel or packrat. Neotropical Migrant: A bird that nests in temperate regions and
Mitochondria (Mitochondrion): Organelles occurring in the migrates to the Neotropical faunal region, which includes the West
cytoplasm of all aerobic cells of plants and animals and containing Indies, Mexico, Central America, and that part of South America within
enzymes responsible for converting foods to usable energy. the tropics.
Mitochondria also contain double-stranded DNA encoding some of Neritic: Relating to or inhabiting the shallow water, or nearshore
the genes functioning in the processing energy and protein synthesis. marine zone extending from the low-tide level to a depth of 200
Miocene: A geologic epoch within the Tertiary period (about 26 to 5 meters. The neritic zone is populated by benthic organisms because of
million years before the present). the penetration of sunlight to these shallow depths.
 GLOSSARY 719

Nitrogen Fixation: The process of converting inorganic, atmospheric Phenology: The study of the relationship between climate and the
nitrogen into an organic form of nitrogen, ammonia. This process can timing of periodic natural phenomena such as migration of birds, bud
be carried out by lightning, by photochemical fixation in the bursting, or flowering of plants.
atmosphere, or by the action of microorganisms. Also, the chemical Phenotype: The observable manifestation of a specific genetic makeup;
processes used in the manufacture of fertilizers. those observable properties of structure and function of an organism
Nival: Of, relating to, or growing under or in snow. as modified by genetic structure in conjunction with the environment.
Nocturnal: Referring to organisms that are active or functional at night. Philopatry (also referred to as site fidelity or site tenacity): The
Nonindigeneous (Also called exotic, nonnative, introduced, and alien): A tendency to return each season to the same nest or breeding colony.
plant or animal that is not native to the area in which it occurs; it was Photic Zone: The surface zone of the sea or a lake having sufficient light
either purposely or accidentally introduced. penetration for photosynthesis.
Nonpoint: Not from a single, well-defined site. Nonpoint sources are Photoperiod: The length of time an organism is daily exposed to light,
pollution-producing entities not tied to a specific origin, such as an especially with regard to how that exposure affects growth and
individual smokestack; include runoff, which washes pollutants from development.
roads into storm sewers and bodies of water or agricultural chemicals Phylogenetic: Pertaining to the evolutionary history of a group or
from lawns, fields, and golf courses. lineage, or the evolutionary relationships within and between
Nutrient Cycling: Circulation or exchange of elements such as taxonomic levels; the relationships of groups of organisms as reflected
nitrogen and carbon between non-living and living portions of the by their evolutionary history.
environment. Includes all mineral and nutrient cycles involving Physiographic Province: A region of the landscape with distinctive
mammals and vegetation. geographical features.
Obligate: Essential, necessary; unable to exist in any other state, mode, Physiography: Landform; physical geography.
or relationship; restricted to one particularly characteristic mode of Phytoplankton: One of two groups into which plankton are divided, the
life. other being zooplankton. Phytoplankton comprises all the freely floating
Obligate Species: A plant of animal that occurs only in a narrowly photosynthetic forms in the oceans.
defined habitat such as tree cavity, rock cave, or wet meadow. Pinniped: Any of a suborder of aquatic carnivorous mammals with all
Old-growth: Referring to an ecosystem or community, particularly a four limbs modified into flippers; includes seals, sea lions, and walruses.
forest, which has not experienced intense or widespread disturbance Pioneer: The first species or community to colonize or re-colonize a
for a long time relative to the life spans of the dominant species and barren or disturbed area, thereby commencing a new biological
which has entered a late successional stage; usually associated with high succession.
diversity of species, specialization, and structural complexity. Piscivores: Fish-eaters; those organisms that subsist exclusively or
Oligochaetes: Any of a class or order (Oligochaeta) of hermaphroditic primarily on fish.
terrestrial or aquatic annelid worms that lack a specialized head. Plankton: Small aquatic organisms (animals and plants) that generally
Oligotrophic: Waters or soils that are poor in nutrients and have low having no locomotive organs, drift with the currents. The animals in this
primary productivity. category include protozoans, small crustaceans, and the larval stages of
Omnivore: Consuming a variety of plants and animals; neither plant nor larger organisms.
animal food usually comprises less than one-third of diet. Plant Association: Stands of vegetation with similar combinations of
Ontogenetic: Relative to the course of growth and development of an species united into abstract types; a basic unit in plant community
individual organism. classification.
Osmerid: A member of the family of fishes (Osmeridae) to which the Playa: A nearly level area at the bottom of an undrained desert basin,
true smelts belong; smelts and smelt-like fishes. sometimes temporarily covered with water during wet periods. Playas
Overgrazing: Continued heavy grazing that exceeds the recovery are barren and usually saline.
capacity of the plant community and creates a deteriorated range. Pleistocene: The earlier epoch of the Quaternary period or the
Ovigerous: Carrying eggs, or modified for carrying eggs. corresponding system of rocks; 1.6 million-10,000 years ago; the “Ice
Oxbow: A pond or wetland created when a river bend is cut off from Age.”
the main channel of the river. Pluvial: Characterized by abundant rain.
Palustrine: Pertaining to wet or marshy habitats. Polychaetes: Any of a class (Polychaeta) of chiefly marine annelid
Parasite: An organism that is intimately associated with and worms (such as clam worms), usually with paired segmental
metabolically dependent on another living organism (the host) for appendages, separate sexes, and a free-swimming trochophore larva.
completion of its life cycle, and which is typically detrimental to the Polyandry: One female mates with two or more males.
host. Polychlorinated Biphenyls (PCBs): A group of toxic, carcinogenic
Passerine: Of or relating to the largest order (Passeriformes) of birds, organic compounds containing more than one chlorine atom; very
which includes more than half of all living birds and consists primarily stable compounds, fat-soluble; they therefore accumulate in ever-higher
of perching songbirds, whose young are hatched in an immature and concentrations as they move up the food chain.
helpless condition Polymerase Chain Reaction (PCR): A molecular biology technique
Patch: Ecosystem elements (e.g., areas of vegetation) that are relatively allowing small and specific amounts of DNA sequences to be amplified
homogeneous internally and that differ from what surrounds them. by many thousand fold in an automated fashion.
Patch Dynamics: The idea that communities are a mosaic of different Polygamy: Both polygyny and polyandry occur.
areas (patches) within which non-biological disturbances (such as Polygyny: One male mates with two or more females.
climate) and biological interactions proceed. Population: A group of organisms, all of the same species, which
Pathogen: A specific causative agent of a disease, such as a bacterium or occupies a particular area. Also, the total number of individuals of a
a virus. species within an ecosystem, or of any group of similar individuals.
Pelagic: Referring to or occurring in the open sea. Population Distribution by Wildlife:
Percent Cover: In descriptions of plant communities, the proportion of Contiguous: Habitat is broadly distributed over the species’ range
ground, expressed as a percentage that is occupied by the with opportunity for continuous or nearly continuous occupation by
perpendicular projection down onto it of the aerial parts of individuals the species; little or no limitation on population interaction.
of the species under consideration. Gaps: Habitat is broadly distributed over the species’ range, but gaps
Perennial: A plant that normally lives for more than two seasons and exist within this distribution. Disjunct patches of habitat are typically
which, often after an initial period, produces flowers annually. large enough and close enough to other patches to permit dispersal
Perennial Stream: A stream that typically has running water on a year- among patches and to allow species to interact as a meta-population.
round basis. Isolated: Habitat locally distributed as isolated patches, causing strong
pH: A Measure of acidity and alkalinity of a solution, taken by measuring limitations for population interaction among patches, and limited
the relative concentration of hydrogen ions in the solution. opportunity of dispersal among patches. Some local populations may
be extirpated and rates of re-colonization will likely be slow.
 720 Wildlife–Habitat Relationships in Oregon and Washington

Patchy: Habitat exists primarily as patches, some of which are small or Rehabilitation: Returning of land to farm use or to productivity in
isolated to the degree that species interactions are limited. Local conformity with a prior land use plan, including a stable ecological state
subpopulations in most of the species’ range interact as a meta- that does not contribute substantially to environmental deterioration
population, but some populations are so disjunct that subpopulations and is consistent with surrounding aesthetic values.
in those patches are essentially isolated from other populations. Relict: Persistent remnants of a formerly widespread species in certain
Scarce: Habitat is very scarce throughout species’ range with little or isolated areas.
no possibilities for interactions between local populations, strong Remote Sensing: Methods for gathering data on a large or landscape
potential for extirpations, and little likelihood of re-colonization. scale which do not involve on-the-ground measurement, especially
Population Dynamics: The aggregate of changes that occur during the satellite photographs and aerial photographs; often used in conjunction
life of a population. Included are all phases of recruitment and growth, with Geographic Information Systems.
senility, mortality, seasonal fluctuation in biomass, and persistence of Resilience: The ability of an ecosystem to maintain diversity, integrity,
each year class and its relative dominance, as well as the effects that and ecological processes following disturbance.
any or all of these factors exert on the population. Resource Partitioning: Division of some resource or resources
Population Viability: Probability that a population will persist for a among two or more con-occurring species; for example, eating slightly
specified period across its range despite normal fluctuations in different foods.
population and environmental conditions. Restoration:
Potential Natural Community: The biotic community that would be Ecological: The reestablishment of predisturbance functions and
established if all successional sequences of its ecosystem were related chemical, biological, and hydrological characteristics.
completed without additional human-caused disturbances under Passive (or Natural): The discontinuation of those activities that are
present environmental conditions. causing degradation or preventing the ecosystem’s recovery.
Precocial Young: Mobile, downy, follow parents, and find their own Retrogression: The return along the successional path to an earlier
food. successional community or an altered simpler state.
Prenatal: Existing or occurring before birth. Riffles: Shallow rapids where the water flows swiftly over completely or
Prescribed Fire: A fire burning under specified conditions that will partially submerged obstructions to produce surface agitation, but
accomplish certain planned objectives. The fire may result from planned where standing waves are absent.
or unplanned ignitions. Riparian: Relating to, living, or located on the bank of a natural
Primary Producer: An organism capable of using the energy derived watercourse (such as a river) or sometimes of a lake or tidewater.
from light or a chemical substance in order to manufacture energy-rich Riparian Ecosystem: Ecosystems transitional between terrestrial and
organic compounds, mainly green plants. aquatic ecosystems. Also, streams, lakes, wet areas and adjacent
Primary Productivity: The rate at which biomass is produced by vegetation communities and their associated soils, which have free
organisms which synthesize complex organic substances from simple water at or near the surface.
inorganic substrates, such as in photosynthesis and chemosynthesis. Riparian Zone: An area of vegetation adjacent to an aquatic ecosystem.
Primary Production: The biomass produced through photosynthesis It has a high water table, certain soil characteristics, and some
and chemosynthesis in a community or group of communities. vegetation that requires free (unbound chemically) water or conditions
Primiparity: Having a single offspring/pregnancy. that are more moist than normal. This zone is transitional between
Progradation: The outward building of a sedimentary deposit, such as aquatic and upland zones.
the seaward advance of a delta or shoreline, or the outbuilding of an Riprap: A general term for large, blocky stones that are artificially placed
alluvial fan. to stabilize and prevent erosion along a riverbank or shoreline.
Promiscuity: Males and females mate more or less indiscriminately. Risk Analysis: A qualitative assessment of the probability of persistence
Province: An area of land, less extensive than a region, having a of wildlife species and ecological systems under various alternatives
characteristic plant and animal population. and management options; generally also accounts for scientific
Purse Seine: A large seine net designed to be set by two boats around uncertainties.
a school of fish and so arranged that after the ends have been brought Rookery: Breeding or nesting place for some gregarious mammals and
together, the bottom can be closed. birds.
Radiation: In ecology, the spread of a group of organisms into new Runoff: Precipitation on land that runs off to a body of water.
habitats. Runs/Glides: Areas of swiftly flowing water, without surface agitation or
Range (of a species): The area or region over which an organism occurs. waves, which approximates uniform flow and in which the slope of the
Range of Variability: The spectrum of conditions possible in ecosystem water surface is roughly parallel to the overall gradient of the stream
composition, structure, and function considering both temporal and reach.
spatial factors. Salinity: A measure of the total concentration of dissolved salts in
Rangeland: Land on which the native vegetation is predominantly water.
grasses, grass-like plants, forbs, or shrubs. Includes lands re-vegetated Salmonid: Any of a family of elongate bony fishes (such as salmon or
naturally or artificially when routine management of that vegetation is trout) that have the last three vertebrae upturned.
accomplished mainly through manipulation of grazing. Sanitation: The removal of dead or damaged trees, or trees susceptible
Recovery Plan: A plan that lists the actions that must be taken and the to insect and disease attack, such as intermediate and suppressed trees,
objectives that must be reached before an organism is no longer essentially to prevent the spread of pest or pathogens and to promote
endangered or threatened and may be removed from the list of forest health.
endangered and threatened species. Savanna (also spelled savannah): A grassland-woodland mosaic
Recruitment: The influx of new members into a population by vegetation type found in tropical and subtropical regions with long dry
reproduction or immigration. periods and receiving more rain fall than desert areas but not enough
Refugium: An isolated area where extensive changes, typically due to to support complete forest cover.
changing climate (such as glaciation) but also due to large-scale Seamount: An underwater mountain (usually a submarine volcanic
disturbances such as those caused by humans, have not occurred, and mountain peak) rising from the ocean floor whose summit is below the
where plants and animals typical of a region may survive. Such a refuge water’s surface.
is a center of relict forms, from which dispersion and speciation may Secondary Production: The biomass production resulting from the
take place after environmental readjustment. assimilation of organic matter produced by a primary consumer:
Regime: A regular pattern of occurrence or action. production by organisms (mainly animals), which consume primary
Region: The broadest scale of landscape ecology composed of a course- producers (mainly plants).
grained pattern of connected landscapes, with contrasting boundaries, Secondary Productivity: The rate of biomass production resulting
that have a similar macroclimate and sphere of human activity and from the assimilation of organic matter produced by a primary
interest. consumer; production by organisms (mainly animals), which consume
primary producers (mainly plants).
 GLOSSARY 721

Sediment: Materials that sink to the bottom of a body of water or Standard Error: In statistics, the standard deviation of the sampling
materials that are deposited by wind, water, or glaciers. distribution of a statistic; an estimate of the range by which the means
Semelparous: A reproductive strategy that allows only one mating of a number of sets of data deviate about the mean of those means.
prior to death, typical of the Pacific salmon. Standing Stock: Biomass; the total mass of organisms comprising all or
Senescence: The aging process in mature individuals; or, the period near part of a population or other specified group or within a given area;
the end of an organism’s life cycle; in deciduous plants, the process that measured as volume, mass, or energy.
occurs before the shedding of leaves. Steppe: Specifically, the temperate, semiarid areas of treeless grassland
Sensitive Species: A species not formally listed as endangered or in the mid-latitudes of Europe and Asia; more generally, any such
threatened, but considered to be at risk, as evidenced by: a significant grassland.
current or predicted downward trend in population numbers or Stewardship: A land ethic for current and future generations that 1)
density; or a significant current or predicted downward trend in habitat encourages wise use and conservation of resources; 2) sustains and
capability that would reduce a species’ existing distribution. enhances productivity of resources; and 3) protects resources.
Seral: Relating to a phase in the sequential development of ecological Stochastic: Random.
communities formed in ecological succession in a particular habitat and Stressors: Physical or biotic factors that stress individual organisms/
leading to a particular climax association; intermediate communities in communities.
an ecological succession. Structural Stage: A stage of development of a vegetation community
Sere: The series of stages that follow one another in an ecologic that is classified on the dominant processes of growth, development,
succession; a series of biotic communities that follow one another in competition, and mortality.
time on any given area of the Earth’s surface. Structure: The various horizontal and vertical physical elements of the
Serotinous Cones: Pinecones that remain on the tree for many years vegetation.
and are tightly closed until stimulated by the heat of a forest fire to Subaerial: Occurring immediately above the surface of the ground.
open and release seeds. Subalpine: The zone just below treeline on temperate mountains,
Serpentine: A mineral rock consisting essentially of a hydrous usually dominated by a coniferous forest ecologically similar to boreal
magnesium silicate (chrysolite and antigorite and usually having a dull forest. The elevation of this zone increases with a decrease in latitude.
green color and often a mottled appearance; or, the usually infertile, Subbasin: The fourth delineation within the hydrologic unit code
excessively well-drained soil derived from serpentine. system. Provides a delineation generally of a river, or group of rivers,
Sessile: Permanently attached to a substrate or established; not free to that flow into a basin.
move about. Also, attached without a stalk. Sublittoral Zone: The deeper zone of a lake below the limit of rooted
Short-stopping: The process of creating habitat improvements to hold vegetation; the marine zone extending from the lower margin of the
geese or ducks throughout the winter in an area that was historically intertidal (littoral) to the outer edge of the continental shelf at a depth
used only as a migratory stopover point en route to wintering grounds of about 200 meters.
farther south. Submersed: Pertaining to a plant or plant structure growing entirely
Silviculture: The art and science of managing forest stands to provide underwater.
or maintain structures, species composition, and growth rates that Subnivian: Beneath the snow cover; specifically, the interface between
contribute to forest management goals. snow and the surface of the ground where small mammals are active in
Simulation: To project or estimate the future or historic conditions or winter.
outcomes of various attributes. Subsidence: The process of sinking or settling of a land surface or a
Sink: A sinkhole; or, an area with a demand for metabolic substances. For crustal elevation because of natural or artificial causes.
example, growing meristems are sinks for energy compounds from Subspecies: A race of a species that is granted a taxonomic name; rules
photosynthesis, mitochondria are oxygen sinks, and tropical rainforests, for designating subspecies are subjective, but subspecies are generally
or deep oceans may act as carbon sinks, absorbing carbon dioxide geographically distinct and form populations (not merely morphs)
from the atmosphere. which differ to some degree from other geographic populations of the
Site: The classification of land area based on its climate, physiographic species.
(physical geography), edaphic (soil), and biotic factors that determine its Substrate: The surface of medium that serves as a base for something.
suitability and productivity for particular species and silvicultural Subterranean: Under the surface of the Earth.
alternatives. Subtidal: Applied to that portion of a tidal-flat environment which lies
Site-potential Tree: A tree that has attained the maximum height below the level of mean low water for spring tides. Normally it is
possible given site conditions where it occurs. covered by water at all states of the tide. Often used as a general
Slough: A swamp, marsh, or muddy backwater. descriptive term for a subaqueous but shallow marine depositional
Smolt: The stage in the life of salmon and similar fishes in which the environment.
subadult individuals acquire a silvery color and migrate down the river Subtropical: The latitudinal zone between 23.50 and 34.00 in either
to begin their adult lives in the open sea. hemisphere, bordering the tropical zone. Also can refer to vegetation,
Snag: A standing dead tree or stump that provides habitat for a broad organisms, or weather typical of subtropical habitats.
range of wildlife, from beetle larvae (and the birds that feed upon Subwatershed: The sixth delineation within the hydrologic unit code
them) to dens for raccoons. system. Provides a delineation of a group of streams that flow into a
Spawn: The eggs of certain aquatic organisms; also, the act of producing watershed.
such eggs or egg masses. Succession: The development of biotic communities following
Species: A group of organisms formally recognized as distinct from disturbances that produce an earlier successional community.
other groups; the taxon rank in the hierarchy of biological classification Successional Path: The sequence of successional communities; may
below genus; the basic unit of biological classification, defined by the follow one path or multiple paths depending on competition, morality
reproductive isolation of the group from all other groups of organisms. of individuals, and non-lethal disturbances. There may be one endpoint
Species Diversity: See Biological Diversity. or multiple endpoints depending on the degree of change in climate
Species Richness: The absolute number of species in an assemblage or and soil properties.
community. Successional Stage: One in a series of usually transitory communities
Staging Area: A traditional area, usually a lake, where birds that migrate or developmental stages that occur on a particular site or area over a
in flocks rest and feed either immediately before or during migration. period of time.
Many flocks may be gathered in such an area. Succulent: A plant that has a specialized fleshy tissue in roots, stems, or
Stand Composition: The representation of tree species in a forest leaves for the conservation of water. Most succulents are xerophytes,
stand, expressed by some measure of dominance (e.g., percent of plants preferring dry climates, such as cactus or aloe, but some are
volume, number, basal area, cover). halophytes, adapted for living in salty soils where water retention is a
Stand Structure: The physical and temporal distribution of plants in a problem.
stand.
 722 Wildlife–Habitat Relationships in Oregon and Washington

Suitability: The appropriateness of applying certain resource Tundra: A level or rolling treeless plain in the arctic or subarctic regions;
management practices to a particular area of land, as determined by an the soil is black and mucky, the subsoil is permanently frozen, and the
analysis of the economic and environmental consequences and the vegetation is dominated by mosses, lichens, herbs, and dwarf shrubs. A
alternative uses foregone. similar environment occurs in mountainous areas above the timberline.
Sustainability: The ability to sustain diversity, productivity, resilience to Tunicate: Any of a subphylum (Urochordata or Tunicata) of marine
stress, health, renewability, and/or yields of desired values, resources chordate animals that have a thick secreted covering layer, a greatly
uses, products, or services from an ecosystem while maintaining the reduced nervous system, a heart able to reverse the direction of blood
integrity of the ecosystem over time. flow, and a notochord in the larval stage.
Sustainable Development: The use of land and water to sustain Turbid: Having sediment or foreign particles stirred up or suspended;
production indefinitely without environmental deterioration, ideally muddy.
without loss of native biodiversity. Tussock: A compact tuft of grass or sedges, or an area of raised solid
Sustainable Ecological System: Emphasizing and maintaining the ground, which is held together by roots of low vegetation, found in a
underlying ecological processes that ensure long-term production of wetland or tundra.
goods, services, and values without impairing productivity of the land. Understory: The vegetation layer between the overstory or canopy and
Sympatric: Referring to populations, species, or taxa occurring together the groundcover of a forest community, usually formed by shade-
in the same geographical area; they may occupy the same habitat or tolerant species or young individuals of emergent species. May also
different habitats within the same geographical area. refer to the groundcover if no tree or shrub layer is present.
Synergistic: Pertaining to the cooperative action of two or more Ungulate: Any four-footed, hoofed, grazing mammal (such as a ruminant,
agencies such that the total is greater than the sum of the component swine, deer, hippopotamus, horse, antelope, elk, elephant, or hyrax) that
actions; combined action or operation. is adapted for running but is not necessarily related to other ungulates.
Syntopic: Relating to or displaying conditions, as they exist Upwelling: The upward movement of cold, nutrient-rich water from
simultaneously over a broad area, as of the atmosphere or weather. ocean depths, produced by wind or diverging currents.
Talus: Broken rock forming a more or less continuous layer that may or Vascular Plants: A plant with a specialized conducting system (for the
may not be covered by duff and litter. transport of water and nutrients) that includes xylum and phloem;
Taxon (Taxa): Any organism or group of organisms of the same includes familiar high plants such as trees, shrubs, and grasses.
taxonomic rank; for example, members of an order, family, genus, or Vertebrate: An animal with a backbone; includes mammals, birds,
species. reptiles, amphibians, and fishes.
Tectonic Movement: The formation of faults and folds on the crust of Viability: The likelihood of continued existence in an area for some
a planet. specified period of time.
Temporal Niche: The functional position of an organism in its Viable Population: A population, which has adequate numbers and
environment as determined by the periods of time during which it dispersion of reproductive individuals to ensure the continued
occurs and is active there. existence of the species population on the planning area.
Terrestrial Community Type (Also referred to as terrestrial Volant: Flying or capable of flying.
vegetation type): A group of cover types in the same seral stages that Watershed: An area or a region that is bordered by a divide and from
have similar characteristics for interpretation habitat values. which water drains to a particular watercourse or body of water.
Terrestrial Ecosytem: A land-based ecosystem (see ecosystem). An Watershed Analysis: A systematic procedure for characterizing
interacting system of soil, geology, and topography with plant and watershed and ecological processes to meet specific management and
animal communities. social objectives. Watershed analysis is a stratum of ecosystem
Territory: The area that an animal defends, usually during breeding management planning applied to watersheds of approximately 20 to
season, against intruders of its own species. 200 square miles.
Tertiary: The first period of the Cenozoic Era which began about 65 Wetland: A general term applied to land areas, which are seasonally or
million years and lasted to 1.6 million years before the present, marked permanently waterlogged, including lakes, rivers, estuaries, and
by formation of high mountains, the dominance of mammals on land, freshwater marshes; an area of low-lying land submerged or inundated
and angiosperms superseding gymnosperms as dominant plants. periodically by fresh or saline water.
Thaliaceans: Any of a small order of tunicates consisting of various Wilderness: An area designated by congressional action under the 1964
aberrant, free-swimming pelagic forms, including those of the genera Protection Act. Wilderness is defined as undeveloped federal land
Salpa and Doliolum. retaining its primeval character and influence without permanent
Threatened Species: Those plant or animal species likely to become improvements or human habitation. Wilderness areas are protected
endangered species throughout all or a significant portion of their and managed to preserve their natural conditions, which generally
range within the foreseeable future as identified by the Secretary of appear to have been affected primarily by the forces of nature, with the
Interior as threatened, in accordance with the 1973 Endangered imprint of human activity substantially unnoticeable; have outstanding
Species Act. opportunities for solitude or for a primitive and confined type of
Threshold: The boundary between ecological states that, once crossed, recreation; include at least 5,000 acres or a are of sufficient size to
is not easily reversible and results in the loss of capacity of produce make practical their preservation, enjoyment, and use in an unimpaired
commodities and satisfy values. condition; and may contain features of scientific, education, scenic, or
Topography: The natural and constructed relief of an area. historical value as well as ecological and geologic interest.
Transect: A line or narrow belt used in ecological surveys to provide a Woodland: A vegetation community that includes widely spaced large
means of measuring and representing graphically the distributions of trees. The tree crowns are typically more spreading in form than those
organisms across a given area. of forest trees and do not form a closed canopy. Grass, heath, or scrub
Transpiration: The loss of water vapor from a plant to the outside may develop between the trees.
atmosphere, mainly through the stomata of leaves and the lenticels of Xeric: Dry; tolerating or adapted to dry conditions.
stems. Year-class: Fish of a given species spawned or hatched in a given year;
Treeline: The upper limits of tree growth in mountains or at high for example, a three year-old fish caught in 1998 would be a member
latitudes. of the 1995 year-class.
Trophic: Pertaining to nutrition or to a position in a food web, food Zoeae: The free-swimming, planktonic larval forms of many decapod
chain, or food pyramid. crustaceans (especially crabs) that have a relatively large cephalothorax,
Tropical: Referring to the zone between the Tropic of Cancer conspicuous eyes, and fringed antennae and mouthparts.
(23E27’N) and the Tropic of Capricorn (23E27’S); characterized by a Zooplankton: See plankton.
climate with high temperatures, humidity, and rainfall. Also can refer to
vegetation, organisms, or weather typical of tropical conditions.
 Managing Directors Senior Staff
To describe our role in creating this book and CD-ROM we Charley Barrett. Charley has been
use the term “Managing Directors” because of our overall specializing in geographic informa-
involvement in the many facets of how the book and CD- tion systems (GIS) for nine years. He
ROM were designed and developed. Specifically, our roles earned an M.S. degree in Geography
included (1) creating and identifying the structure and roles at Oregon State University in 1998.
of the five scientific teams that guided this project, (2) He studied GIS, cartography, and
developing a concept (and for several cases an outline) for rural planning while also teaching
each of the chapters in the book, (3) identifying the structure computer mapping and GIS labs. As
and core fields of information for the matrixes on the CD- an undergraduate at Towson
ROM, (4) organizing and conducting the 15 expert scientific University, Maryland, Charley
panels to gather key wildlife habitat information, (5) raising majored in geography and economics with an emphasis in
funds to support Senior Staff and contractors for four years, environmental planning. Currently a member of the
and for the publication of this book, (6) fostering the Northwest Habitat Institute, he assists in mapping vegetation,
development of several new wildlife and ecological concepts, habitat, and species at local to regional scales, as well as
(7) developing and publishing the CD-ROM, (8) coordinating developing the Internet site. Charley enjoys motorcycling,
and guiding the involvement of more than 600 people to help bicycling, camping, and picking the five-string banjo in his
build a common understanding for management along with spare time.
coordinating with adjacent states and provinces.
Kelly Anne Bettinger. Kelly
David H. Johnson David has been received her B.S. in Wildlife
involved in the natural resource field Management from Virginia Tech in
for 24 years. He has held forestry, 1989 and her M.S. in Wildlife Biology
wildlife biologist, and habitat from Oregon State University in
scientist positions with the 1996. She has worked on a variety
Minnesota Department of Natural of projects on both U.S. coasts
Resources, BLM, USFWS, ODFW, studying screech owls, spotted owls,
and WDFW. He has Associate red-cockaded woodpeckers, sea
degrees in Natural Resource snakes, sea turtles, and rare plants.
Management and Civil Engineering, Her current interests center on breeding bird community
a B.A. in Biology with a Minor in Archaeology, and an M.S. studies and she has spent the past eight field seasons
in Wildlife Science. He has a passion for owls and is deeply surveying birds in various habitats throughout Oregon. Kelly
involved with international owl conservation. He loves has been involved in most aspects of the project including
traveling and reading historical exploration accounts, and for compiling literature, organizing and moderating expert panel
therapy from it all, builds and paddles sea kayaks and cedar- sessions, filling out life history matrixes, reviewing the
strip canoes. matrixes for quality control, providing literature and database
information to chapter authors, and editing chapters. She also
Thomas O’Neil Tom has spent 25 enjoys participating in Christmas bird counts, running a
years working on ecological issues
breeding bird survey in eastern Oregon, and collecting old
in the Pacific Northwest and has natural history books.
several degrees from the University
of Montana and the University of Pat Doran. Pat earned M.S. degrees
Toledo. Currently, Tom is the in Ecology and Environmental
Director of the Northwest Habitat Science from Indiana University. His
Institute. Previously, he has worked research focused on the relationship
as a Wildlife Ecologist for ODFW, between land cover and the
Research Division; University of reproductive success of songbirds.
Chicago’s Argonne National Laboratory; Montana Power While with our project, Pat was
Company; as a Wildlife Biologist for the USFS, and as an involved in database and chapter
Administrator for the Jeep Corporation. Tom has also co- development, and scientific panels,
authored the book Atlas of Oregon Wildlife, and has written as well as project planning. Pat left
over 40 other publications or documents. Tom is a second the project to pursue a Ph.D. in Ecology at Dartmouth College.
generation Irish-American with three of four grandparents In his free time, Pat enjoys bird watching, hiking and
coming directly from Ireland. His father worked in the early backpacking, traveling, and just about any athletic activity.
stages of the U.S. Space Program. Tom is married and has 2
Eva Greda. Eva graduated from
children. He has been a Certified Wildlife Biologist with The
Wildlife Society for 18 years. In his spare time, Tom enjoys Western Washington Univer-sity
with a B.S. degree in Cell Biology.
the outdoors, traveling, and working on his 1949 Willys Jeep.
While at Western she was involved
Bringing all this together could not have been possible in a research project attempting to
without a dedicated staff who devoted many hours wading find a sequence homology between
through countless reports, articles, databases, and books, the Caudal gene of the fruit fly
assisting with the scientific panels, as well as coordinating Drosophila melanogaster, and cDNA of
and working with numerous wildlife professionals through- the wasp Nasonia vitripenisis. Eva has
out the region. These invaluable people, who helped us from worked for more than a year with
development to completion of this book and digital matrixes the WDFW and previously with the Washington State
and maps, are listed below:
723
 724 Wildlife–Habitat Relationships in Oregon and Washington

Attorney General’s office regarding tobacco litigation. Her computer specialist in technical user support. She was the
contributions to this project included recording and entering data systems and digital technologies co-lead for the project.
the results from the scientific panel process, compiling a Marla’s interests include both indoor and outdoor soccer,
portion of the life history matrix, and co-authoring several downhill skiing, crafts, and traveling. She has also been an
chapters. Her interests are gardening, painting, playing the assistant coach for the WOU women’s soccer team for two
piano, and walking her dog, Dana. years. Having grown up in Alaska gave her an appreciation
for the outdoors, and she enjoys exploring Oregon’s
Teri Guydish. Teri received her
landscapes.
Clerk/Receptionist Certificate at
South Puget Sound Community Madeleine Vander Heyden
College (SPSCC) in Olympia, Madeleine is a wildlife biologist
Washington after twenty-seven currently living in North Bend,
years of being a housewife and Oregon. She graduated with an M.S.
mother of four children. Through the in Wildlife Ecology from Oregon
cooperative work program at State University in 1997 and received
SPSCC, she found a position with her B.S. from the University of
the Washington Department of Fish Wisconsin in 1987. She has worked
and Wildlife. This opened the door for several opportunities on a variety of wildlife species,
in the Habitat Program. She contributed to this project by including research on spotted owls,
copying articles to support the development of the matrixes, small mammals, bald eagles, and black bears. Madeleine’s
doing data entry, and helping out with many other tasks. In experience contributed to the digitial matrixes for habitat
her free time she enjoys spending time with her family, elements, species key ecological functions, and management
gardening, fishing, camping, and baking great cookies. activity links to the habitat elements. Her special areas of
interest include wildlife-habitat relationships, carnivore
Chris Kiilsgaard. Chris received a
ecology, and public lands management.
B.S. in Botany from Montana State
University and an M.S. in While we were initiating this effort, Dr. Laurie Wunder helped
Biogeography from Oregon State tremendously with identifying and defining information to
University. He has eighteen years of be considered in the Wildlife-Habitat types, Structural
experience in mapping vegetation Conditions, Habitat Elements, and Life History matrixes, as well
and wildlife habitats in the Pacific as assisting with the scientific panel process. Additional
Northwest. Chris is the Director for project staff whose efforts were also invaluable are Kelly
Mapping Research for the Cruce, Jackie Enriquez, Connie Iten, Patricia Johnson, Kathryn
Northwest Habitat Institute. LePome, Kathleen Morley, John Morgan, Russell Rodgers,
Presently he is compiling a map of wildlife habitats for the Meg Shaughnessy, Daniel Stein, and Susan Tank.
seven-state Columbia River Watershed and mapping upland A number of individuals who, although they did not work
vegetation in the Oregon Coast Range. His other professional for the project directly (as employees), contributed
interest is habitat restoration. In his spare time he enjoys importantly to key aspects of the effort: Joseph R. Evenson
tennis, vegetable gardening, and scheming to own a vineyard. consistently offered his assistance and insights into marine
mammal and bird aspects. A most talented individual,
Derek Stinson. Derek received his Terrance P. Johnson, provided critical GIS support by
B.S. in Biology at Framingham State
developing many of the species range maps, marine habitat
College, and his M.S. in Zoology type boundary determinations, marine species richness
from Washington State University.
analysis, county occurrence determinations, and essential
His diverse interests and work habitat-type analysis for the project. We are deeply indebted
experiences have included shrews,
to the scientific insights and rigor, and all-around professional
bats, galliforms, raptors, insects, and input of Dr. Bruce G. Marcot, who helped guide us at many
wetland birds. He spent four years
important junctures in the project. We also thank Nikki
working on endangered birds, fruit Derringer, Monica Hoover, Karol McFarlane, Cindy J. Neff,
bats, and migrant shore and
Sharon Newton, Jeff Reams, Joann Smith, Lori Turner, Sue
waterbirds in the Mariana Islands, and wrote the recovery Vance, Sara Vickerman, and Karla Yeager for their important
plan for the Micronesian Megapode. More recently he has
contributions to the success of this project.
worked on rare species and conservation planning of forest
wildlife in Washington. Derek’s expertise was used to assist
Acknowledgments
in compling information for the life history matrix. He enjoys
This book, Wildlife—Habitat Relationships in Oregon and
gardening, birding, painting, and other forms of celebrating
Washington, has been developed through the combined efforts
and caring for creation.
of more than 600 people. The final tally shows that we
Marla Trevithick. Marla received a received input from 246 questionnaire respondents, 225
B.A. in Computer Science from people who attended the landscape modeling workshop in
Western Oregon University (WOU) Olympia, 73 species specialists who participated in the 15
and worked with the ODFW and scientific panels, 40 people who accessed and contributed to
WDFW for three years. Currently, our Internet site, 88 chapter authors, and another 30 people
she is working with U.S. Geological who guided our process by participating on one of five
Survey’s Forest and Rangeland advisory teams.
Ecosystem Science Center as a Special thanks go to Bill Martin, the individual who
designed the Northwest Coast Indian artwork for this book.
 ACKNOWLEDGMENTS 725

Bill is a fisherman, diver, and renowned carver and artist, W. Keegan, Jeffrey C. Lewis, Joseph B. Lint, John P. Loegering,
and specializes in Northwest Coast Indian artwork. Bill is a Michael Marsh, Hal Michael, Russell Morgan, Tim R.
Makah tribal member, and he and his wife, April, and their Mullican, Dave Nysewander, Thomas O’Neil, Joel E. Pagel,
four children live in Neah Bay, Washington. His work can be Mike Patterson, Dr. Mary Poss, Gary J. Roloff, Derek W.
found in many locations, including the Smithsonian Institute. Stinson, Rebecca Thompson, Cynthia Trombino, Matthew
A recent example of his talent is reflected in the eight-pole Vander Haegen, Madeleine Vander Heyden, David Vesely,
carving at the new marina in Neah Bay. During discussions James W. Watson, and Simon Wray.
of the artwork for this project, Bill was overheard saying that People who participated in the 15 scientific panels were:
he can draw the Northwest Coast designs as easily as he Amphibians and Reptiles: Doug Calvin, Marc Hayes, Richard
breathes. Hoyer, Bill Leonard, Kelly McAllister, Richard Nauman,
During the course of this project, project staff acquired Chuck Peterson, and Alan St. John; Bats: Ralph Anderson,
over 9,000 hard copies of journal articles and related Burr Betts, Steve Cross, James Hallett, Barry Keller, Pat
publications. The Washington State Library staff heroes on Ormsbee, Steve West, and Laurie Wunder; Small Mammals:
this monumental task were Diane Mitchell, Shaun Fuller, Don Ralph Anderson, Brian Biswell, Charles Drabeck, Jim Hallett,
Gulliford, Ron Howard, Ingrid Morley, David Pope, Marina Barry Keller, Tom Manning, Margaret O’Connell, Steve West,
Rodriguez, Heidi Schroder, and Becky Stewart. Heartfelt Laurie Wunder, and Eric Yensen; Meso Mammals: Robert
thanks go to Diane for her undying support and peppy Anthony, Keith Aubrey, Terry Ferrell, Tom O’Neil, Dale
attitude, and to Heidi, who contributed more than two Toweill, and Don Whittaker; Ungulates: Ed Arnett, Lou
summers of her time working on the project’s literature Bender, Richard Pedersen, Dale Toweill, Madeleine Vander
requests. Heyden, Gary Witmer, and Don Whittaker; Marine Mammals:
We are indebted to the primary contacts we had from our John Calambokidis, Greg Green, Jan Hodder; Loons, Grebes
thirty-four Project Partners and Contributing Sponsors. Extra and Waterfowl: Kevin Blakely, Robert Jarvis, Dave
efforts came from the following folks: Robert Anthony, Kate Nysewander, and Tom O’Neil; Pelicans, Herons, and Cranes:
Benkert, Tim Bodurtha, Jim Bottorff, Kay Brown, Charlie Ken Popper, Joe Engler, Gary Ivey, Deborah Jaques, and Don
Bruce, Joe Buchanan, Eric Campbell, Alan Christensen, Pete Norman; Shorebirds: Joe Buchanan, Pat Doran, Jeff Gilligan,
Comaner, Katie Distler, Bob Falkenstein, Kelly Cassidy, Ron and Nils Warnock; Seabirds: Ken Warheit, Kim Nelson, Chris
Escaño, Randy Fisher, Guy Green, Craig Hansen, Leslie Thompson, and Craig Strong; Raptors: Eric Forsman, Rick
Lehmann, Dave Marshall, Lisa Norris, Rose Owens, Peter Gerhardt, Denver Holt, Frank Isaacs, David Johnson, John
Paquet, Russ Peterson, Claire Puchy, Jim Rochelle, Paul F. Marzluff, Joel Pagel, and Brian Woodbridge; Gallinaceous
and Teresa Roline, J. Michael Scott, Tom Toman, Julie Birds, Pigeons, and Doves: Ralph Anderson, John Crawford,
Thompson, Laura Todd, Bill Tweit, Cheryl Quade, Paul Kathy Martin, Alan Sands, and Mike Schroeder; Passerine and
Wagner, Jeff Waldon, Tom Williams, and Lenny Young. Maria Near Passerine Birds (Group 1): Paul Adamus, Bob Altman,
A. Hug consistently played a crucial and supportive role in Kelly Bettinger, Dan Gumtow-Farrior, Rick Lundquist, Dennis
the fiscal aspects of the project. Vroman, Kim Mellen, Catherine Raley, and Russell Rogers;
The process to create and develop the wildlife habitat and Passerine Birds (Group 2): Bob Altman, Kelly Bettinger,
information was guided by five teams. Team members gave Chris Chappell, Steve Dowlan, Joan Hagar, Matt Hunter, Rex
freely of their time and the insights from their many years of Sallabanks, and Matt Vander Haegen. Management Activities
experience proved essential to the integrity of the final and Links: Susan Tank, Marla Trevithick, Thomas O’Neil,
products. Science Team members were Richard Holthausen, David H. Johnson, Pat Chapman, Curt Leigh, Randy Carman,
USFS; Donavin Leckenby, ODFW (retired); Charles Meslow, Cheryl Friesen, Donavin Leckenby, and Doug Runde, Derek
Wildlife Management Institute (retired); Martin Raphael, Stinson, and Kelly Bettinger, Cheryl Broyles, Paul Wagner,
USFS; and James Rochelle, Weyerhauser Company (retired). and E. Charles Meslow.
Species Habitat Team members were Steve Kohlmann, Prairie We are deeply indebted to the chapter authors (their names
Wings, Inc.; Eric Larsen, WDFW; Kim Mellen, USFS; Tom are published with their respective chapters in this book),
Mumford, WDNR; Doug Runde, Weyerhaeuser Company; who brought forward their scientific best for this publication.
and Matt Vander Haegen, WDFW. Digital Products Team We are also grateful to Sally Olson-Edge, who was a technical
members were Donavin Leckenby, ODFW (retired); Tom editor for many chapters, and to the staff at Oregon State
Owens, WDFW; Jon Sadowski, BLM; Barbara Wales, USFS; University Press, especially Jo Alexander and Jeff Grass.
and Tim Young, WDNR. Management Application Team Lastly, the Managing Directors take full responsibility for
members were Heather Ballash, Washington Community, any errors found within this publication and CD-ROM.
Trade, and Economic Development; Bonita Cleveland and
Trevin Taylor, Quileute Indian Tribe; Cheryl Friesen, USFS; Credits for Habitat Photographs
Cheryl Gruenthal, Boise Cascade; Joe Lint, BLM; Terry Luther, (Chapter 2)
Confederated Tribes of the Warm Springs; Russ Peterson,
USFWS; Steve Smith, ODFW; and Paul Wagner, Washington Agriculture (color)
1. Near Samish Bay, Washington. Photo: Joseph B. Buchanan
Department of Transportation. Marine Team members were 2. Sinlahekin Valley, Washington. Photo: Reid Schuller
Helen Berry, WDNR; Ken Warheit, WDFW; Jeff Skriletz, 3. Ellensburg area, Washington. Photo: Andrew Kratz
WDFW; Dave Nysewander, WDFW; and Joseph Evenson, 4. Wheat Field, Southeast Washington. Photo: Rollin Geppert
WDFW. 5. Christmas Tree farm, Benton County, Oregon. Photo: Kelly A. Bettinger
People who contributed information via the Internet were 6. Scotch Creek Wildlife Area, Okanogan County, Washington. Photo:
Burr J. Betts, Ed Bowlby, R. Mark Brigham, Brian L. Biswell, Jerry Benson
Anna M. Bruce, Eric K. Cole, Michael G. Cope, L. Morris Alpine Grasslands and Shrublands (color)
Eiffert, Joseph Engler, Joseph R. Evenson, Terry Farrell, 1. Panhandle Gap, Mt. Rainier National Park, Washington. Photo: Chris
Howard Ferguson, John H. Guetterman, Lisa A. Hallock, Chappell
Aaron L. Holmes, Ronald J. Jameson, Martha Jordan, Thomas
 726 Wildlife–Habitat Relationships in Oregon and Washington

2. Steens Mt., Oregon. Photo: Chris Chappell Eastside Grasslands (color)

3. Berkeley Park, Mt. Rainier National Park, Washington. Photo: Chris 1. Gibraltar Mt., Ferry County, Washington. Photo: Rex Crawford
Chappell 2. Palouse River, Washington. Photo: Rex Crawford
4. NE Olympics, Washington. Photo: Chris Chappell 3. Dalles Mts., Washington. Photo: Rex Crawford
Alpine Grasslands and Shrublands (black and white) 4. Hanford area, Washington. Photo: Chris Chappell
Example of Alpine grasslands and shrublands. Photo: Jeff Reams Eastside Grasslands (black and white)
Bays and Estuaries (color) Example of Eastside Grasslands. Franklin County, Washington. Photo:
1. Estuary, Washington. Photo: WDFW Andy Kratz
2. Dogfish Pt., Skagit County, Washington. Photo: Chris Chappell Eastside Mixed Conifer Forest (color)
3. Niawiakum River, Willapa Bay, Washington. Photo: Linda Kunze 1. Wenatche Mts., Kittitas County, Washington. Photo: Chris Chappell
4. Nisqually Delta, Washington. Photo: Linda Kunze 2. Alice Mae Mt., Stevens County, Washington. Photo: Chris Chappell
5. Niawiakum River, Willapa Bay, Washington. Photo: Mark Sheehan 3. Alice Mae Mt., Stevens County, Washington. Photo: Chris Chappell
6. Newport, Oregon. Photo: David H. Johnson 4. Rainbow Creek, Research Natural Area, Blue Mts., Washington. Photo:
Bays and Estuaries (black and white) Chris Chappell
Bays and Estuaries. Skokomish River, Hood Canal, Washington. Photo: Eastside Mixed Conifer Forest (black and white)
Steve Jeffries. Example of Eastside Mixed Conifer Forest. Photo: Jeff Reams.
Ceanothus Manzanita Shrublands (color) Eastside Riparian Wetlands (color)
1. Southwestern Oregon. Photo: James F. Harper 1. Northrup Canyon, Washington. Photo: Reid Schuller
2. South of Shady Cove, Oregon. Photo: James F. Harper 2. Little Pend Oreille River, Stevens County, Washington. Photo: Reid
3. Southwestern Oregon. Photo: Jeff Reams. Schuller
4. South of Shady Cove, Oregon. Photo: James F. Harper 3. Douglas Creek, Douglas County, Washington. Photo: Chris Chappell
Ceanothus Manzanita Shrublands (black and white) 4. Myers Creek, Okanogan County, Washington. Photo: John Gamon
Example of Ceanothus Manzanita Shrublands, southwestern Oregon. 5. Eastern Klickitat County, Washington. Photo: DNR Natural Heritage
Photo: James F. Harper. Program
6. Crimm’s Creek, Washington. Photo: Cheryl Quade
Coastal Headlands and Islets (color)
1 Near Cape Perpetua, Oregon. Photo: Jeff Ream Eastside Riparian Wetlands (black and white)
2 Heceta Head, Oregon. Photo: Jeff Ream 1. Example of Eastside Riparian Wetlands. Crab Creek, Lincoln County,
3. Near Yaquina Head, Oregon. Photo: David H. Johnson Washington. Photo: Rex Crawford.
4. Near Cape Perpetua, Oregon. Photo: Jeff Reams HerbaceousWetlands (color)
5. Coastline near Quinalt River, Washington. Photo: Chris Chappell 1. Leuton Flat, Okanogan County, Washington. Photo: Elisabeth Rodrick.
Coastal Headlands and Islets (black and white) 2. Methow Marsh, Washington. Photo: Rex Crawford.
Example of Coastal Headlands and Islets. Photo: Jeff Reams. 3. Pacific County, Washington. Photo: Robin Woodin
4. Lincoln County, Washington. Photo: Rex Crawford
Coastal Dunes (color) 5. Linn County, Oregon. Photo: Jeff Reams
1. Florence, Oregon. Photo: Madeleine Vander Heyden
2. Dugualla Bay, Washington. Photo: DNR Aquatic Lands Program Herbaceous Wetlands (black and white)
3. Ocean Shores, Washington. Photo: David H. Johnson Example of Herbaceous Wetlands. Photo: Jeff Reams
4. Rocky Pt., Whidbey Island, Washington. Photo: Chris Chappell Inland Marine Deeper Waters (color)
5. Rocky Pt., Whidbey Island, Washington. Photo: Chris Chappell 1. Squaxin Island, South Puget Sound, Washington. Photo: David H.
Coastal Dunes (black and white) Johnson
Example of Coastal Dunes. Photo: Jeff Reams 2. San Juan Islands, Washington. Photo: Coastal and Ocean Resources, Inc.
Desert Playa (color) Inland Marine Deeper Waters (black and white)
1. Alvord Desert, Oregon. Photo: Jeff Rems Example of Inland Marine Deeper Waters. March Point, Skagit County,
2. Alvord Desert, Oregon. Photo: Chris Chappell Washington. Photo: DNR, Nearshore Habitat Program.
3. Alvord Desert, Oregon. Photo: Chris Chappell Lodgepole Pine Forest andWoodlands (color)
4. Harney Basin, Oregon. Photo: Joseph B. Buchanan 1. Loomis State Forest, Washington. Photo: Cheryl Quade
5. Alvord Desert, Oregon. Photo: Chris Chappell 2. Loomis State Forest, Washington. Photo: Cheryl Quade
Desert Playa (black and white) 3. Loomis State Forest, Washington. Photo: Cheryl Quade
Example of desert playa and salt scrub shrublands. Photo: Jeff Reams. 4. Loomis State Forest, Washington. Photo: Cheryl Quade
5. Loomis State Forest, Washington. Photo: Cheryl Quade
Dwarf Shrub-Steppe (color)
1. Castle Rock, Grand Coulee, Washington. Photo: Andy Kratz Lodgepole Pine Forest and Woodlands (black and white)
2. Umtanum, Kittitas County, Washington. Photo: Chris Chappell Example of Lodgepole Pine Forest and Woodlands. Loomis, Washington.
3. Saddle Mts., Washington. Photo: Chris Chappell Photo: Cheryl Quade
4. Sheldon N.R.W., Nevada. Photo: Michael Gregg Marine Nearshore (color)
Dwarf Shrub-Steppe (black and white) 1. Ben Ure Island, Island County, Washington. Photo: DNR, Nearshore
Example of Dwarf Shrub-Steppe. Moses Coulee, Douglas County, Habitat Program
Washington. Photo: Rex Crawford 2. Cypress Island, Skagit County, Washington. Photo: DNR, Nearshore
Habitat Program
Eastside Canyon Shrublands (color) 3. Guemes Island, Washington. Photo: DNR, Nearshore Habitat Program
1. Barker Mt. Okanogan County, Washington. Photo: Chris Chappell 4. Rock North of Biz Pt., Skagit County, Washington. Photo: DNR,
2. H. J. Experimental Forest, Oregon. Photo: Chris Chappell Nearshore Habitat Program
3. H. J. Experimental Forest, Oregon. Photo: Chris Chappell 5. Near Cascade Head, Lincoln County, Oregon. Photo: Jeff Reams
4. Smith Canyon, Methow Valley, Washington. Photo: Chris Chappell
5. Klickitat County, Washington. Photo: Mark Sheehan Marine Nearshore (black and white)
Example of Marine Nearshore. North Samish Island, Skagit County,
Eastside Canyon Shrublands (black and white) Washington. Photo: DNR, Nearshore Habitat Program
Example of Eastside Canyon Shrublands. Steens Mt., Oregon. Photo:
Madeleine Vander Heyden Montane ConiferousWetlands (color)
1. Arlecho Creek, Washington. Photo: Chris Chappell
2. Roger Lake, Washington. Photo: Rex Crawford
 ACKNOWLEDGMENTS 727

3. Mt. Rainier, Washington. Photo: Chris Chappell Upland Aspen Forest (black and white))
4. Boulder Creek, Washington. Photo: Chris Chappell Example of Upland Aspen Forest. Sun Mt., Winthrop, Washington. Photo:
Montane Coniferous Wetlands (black and white) Rollin Geppert
Example of Montane Coniferous Wetlands. Photo: Jeff Reams. Urban (color)
Montane Mixed Conifer Forest (color) 1. Example of high density urban, Seattle, Washington. Photo: Rollin
1. Boulder Creek, Washington. Photo: Chris Chappell Geppert
2. Pend Oreille County, Washington. Photo: Reid Schuller 2. Example of medium density urban, Washington. Photo: DNR
3. Bald Mt., Mt. Pilchuck, Washington. Photo: Chris Chappell Nearshore Habitat Program
4. Arlecho Creek, Washington. Photo: Chris Chappell 3. Example of low density urban, Benton County, Oregon. Photo: Kelly a.
Bettinger
Montane Mixed Conifer Forest (black and white) 4. Example of low density urban, Wenatchee, Washington. Photo: Rollin
1. Example of Mixed Conifer Forest. Photo: Jeff Reams Geppert
OpenWater Urban (black and white)
1. Lake Ozette, Washington. Photo: Linda Kunze Example of Urban. Maple Valey, Washington. Photo: Rollin Geppert
2. Oxbow in lower Soleduck River, Washington. Photo: Chris Chappell
3. Grant County, Washington. Photo: James Hannah Westside Grasslands (color)
4. Snake River, Washington. Photo: Rollin Geppert 1. Scatter Creek, Washington. Photo: Chris Chappell.
5. Willamette River, Linn County, Oregon. Photo: Jeff Reams 2. Scatter Creek, Washington. Photo: Chris Chappell.
3. Bald Hill Natural Area Preserve, Washington. Photo: Mark Sheehan
Open Water (black and white) 4. Burrows Island, Washington. Photo: Chris Chappell.
Example of Open Water. Columbia River, Washington. Photo: Rollin 5. Washington. Photo: Dave Hayes.
Geppert.
Westside Grasslands (black and white)
Ponderosa Pine Forest andWoodlands (color) Example of Westside Grasslands. Smith Prairie, Washington. Photo: Chris
1. Barker Mt., Okanogan County, Washington. Photo: DNR, Natural Chappell.
Heritage Program
2. Near Sisters, Oregon. Photo: Cathy Rose Western Juniper and Mountain MahoganyWoodlands (color)
3. Turnbull Wildlife Refuge, near Spokane, Washington. Photo: Cheryl 1. Sheldon National Wildlife Refuge, NV. Photo: Michael A. Gregg
Quade 2. Sheldon National Wildlife Refuge, NV. Photo: Michael A. Gregg
4. Indian Ford, north of Sisters, Oregon. Photo: Chris Chappell 3. Jefferson County, Oregon. Photo: Jeff Reams
5. Briske Canyon, Washington. Photo: Rollin Geppert 4. Klamath County, Oregon. Photo: Jeff Reams.
6. Badger Gulch Natural Area Preserve, Klickitat County, Washington. Western Juniper and Mountain Mahogany Woodlands
Photo: Mark Sheehan (black and white)
Ponderosa Pine Forest andWoodlands (black and white)) Example of Western Juniper and Mountain Mahogany Woodlands. Photo:
Example of Ponderosa Pine. Photo: Jeff Reams Jeff Reams.
Shrub-Steppe (color) Westside Lowlands Conifer-Hardwood Forest (color)
1. Yakima, Washington. Photo: Chris Chappell 1. Goodman Creek, Oregon. Photo: David H. Johnson.
2. Horse Heaven Hills near Prosser, Washington. Photo: Matt Vander 2. H. J. Andrews Experimental Forest, Oregon. Photo: Chris Chappell
Haegen 3. Hoypus South, Washington. Photo: Chris Chappel
3. Yakima, Washington. Photo: Chris Chappell 4. Capitol Forest, Olympia, Washington. Photo: Rollin Geppert
4. Steens Mt, Oregon. Photo: Chris Chappell 5. Upper Dungeness, Washington. Photo: Chris Chappell
5. Varnita, Washington. Photo: Rex Crawford 6. Lake Quinalt, Washington. Photo: David H. Johnson
Shrub-Steppe (black and white)) Westside Lowlands Conifer-Hardwood Forest (black and white)
Example of Shrub-Steppe. Goose Hill, Benton County, Washington. Photo: Example of Westside Lowlands Conifer-Hardwood Forest. Skokomish,
Reid Schuller Washington. Photo: Rollin Geppert.
Southwest Oregon Mixed Conifer-Hardwood Forest (color) Westside Oak and Dry Douglas-Fir Forest andWoodlands (color)
1. Southwestern Oregon. Photo: James F. Harper 1. Cormorant Bay, San Juan Islands, Washington. Photo: Chris Chappell
2. Jackson County, Oregon. Photo: Jeff Reams 2. James Island, San Juan Islands, Washington. Photo: Chris Chappell
3. Ruch, Oregon. Photo: Jeff Reams 3. Cady Mt., Orcas Islands, Washington. Photo: Chris Chappell
4. Argonne Forest, Fort Lewis, Washington. Photo: Chris Chappell
Southwest Oregon Mixed Conifer-Hardwood Forest
(black and white) Westside Oak and Dry Douglas-Fir Forest and Woodlands
Example of Southwest Oregon Mixed Conifer Forest. Southwestern (black and white)
Oregon. Photo: James. F. Harper Example of Westside Oak and Dry Douglas-Fir Forest and Woodlands.
Photo: Jeff Reams
Subalpine Parkland (color)
1. Grand Park, Mt. Rainier National Park, Washington. Photo: Chris Westside RiparianWetlands (color)
Chappell 1. Middle Stequaleho, WA. Photo: Chris Chappell
2. Strawberry Mt., Oregon. Photo: Madeleine Vander Heyden 2. Cranberry Creek, WA. Photo: Linda Kunze
3. Bald Mt., Mt. Pilchuck, Washington. Photo: Chris Chappell 3. Maxfield Creek, WA. Photo: Chris Chappell
4. Eagle Cap, Oregon. Photo: Madeleine Vander Heyden 4. Cranberry Creek, Grays Harbor, WA. Photo: Linda Kunze
5. Goat Rocks, Lewis County, Washington. Photo: Chris Chappell 5. Quinalt River, WA. Photo: David H. Johnson
Subalpine Parkland (black and white)) Westside Riparian Wetlands (black and white)
Example of Subalpine Parkland. Eagle Cap, Oregon. Photo: Madeleine Example of Westside Riparian Wetlands. Tacoma Creek, Pierce County,
Vander Heyden. Washington. Photo: Chris Chappell.
Upland Aspen Forest (color)
1. Sum Mt., Winthrop, Washington. Photo: Rollin Geppert
2. Hart Mt., Oregon. Photo: Joseph B. Buchanan
3. Steens Mt., Oregon. Photo: Chris Chappell
4. Hart Mt., Oregon. Photo: Madeleine Vander Heyden
728
WILDLIFE IN WESTSIDE AND HIGH MONTANE FORESTS 729
 Index species species richness Waterbank Program
antelope, pronghorn Rocky Shores Strategy Wetland Reserve
adaptive management at-risk species Program Program
agricultural, criteria for identifying strategies continental shelf
grasslands, defining risk bioyear precipitation corridors,
case study methods for determining bison, American greenways
composition risk analysis and management bittern, American migration
elements auklet, Cassin’s least coot, American
and exotic wildlife species rhinoceros blackbird, Brewer’s cormorant, Brandt’s
and neotropical migrants autochthonous organic matter red-winged double-crested
buildings and hibernacula avocet, American tricolored pelagic
ephemeral lands/amphibian badger, American yellow-headed cottontail, eastern
breeding sites bat, big brown bluebird, mountain Nuttall’s (mountain)
field borders as filter strips Brazilian free-tailed western cougar, see mountain lion
hedge and fencerows California myotis boa, rubber, see snake cowbird, brown-headed
roadsides fringed myotis bobcat coyote
shelterbelts and windbreaks hoary bobolink crane, sandhill
fertilizers Keen’s myotis bobwhite, northern creeper, brown
pesticides little brown myotis brant crossbill, red
animal damage long-eared myotis brown bear, see grizzly bear white-winged
ecological traps long-legged myotis buffalo, see American bison crow, American
edges pallid bufflehead, see duck northwestern
wells and water developments silver-haired bugler, see trumpeter swan cryptogammic crust
landowner assistance program spotted bunting, lazuli cuckoo, yellow-billed
(table) Townsend’s big eared snow cultivated cropland, defined
managing for wildlife western small-footed myotis burro, wild curlew, long-billed
field borders yuma myotis bushtit DDT and eggshell thinning in
food plots Bays and Estuaries California Current California condor
hedge and fencerows acreage canvasback, see duck merlin
odd areas habitat description caribou, mountain DecAID (wood decomposition
reducing pesticide impacts beaches catbird, gray model)
roadsides drift logs Ceanothus-Manzanita Shrublands components of
shelterbelts and windbreaks tidal regime acreage quantitative objectives
point-count censusing (table) see Coastal Dunes and Beaches habitat description decaying wood,
research needs types of cetaceans abundance, factors
structure bear, black chaparral habitat, birds in influencing
wildlife benefits (table) grizzly chat, yellow-breasted biological agents of
wildlife diversity beaver, American chickadee, black-capped classification system for
Agriculture, Pastures, and Mixed mountain boreal ecological function of
Environs biodiversity, conservation of chestnut-backed fragmentation
acreage current approaches mountain functions of
habitat description aquatic diversity areas chipmunk, Allen’s
albatross, black-footed at-risk species least human disturbance,
Laysan conservation opportunity areas red-tailed effect of
short-tailed current conservation network Siskiyou in streams and riparian
allelic diversity data layers Townsend’s forests
allozyme analysis ecoregional analyses yellow-pine in wetlands
alpine and sub-alpine habitats land ownership chukar in wilderness areas
air and water quality multiple use protected areas Coarse Wood Dynamics Model management of
avalanche chutes as cover national gap analysis program (CWDM) output rates
climate change, global warming Oregon biodiversity project coarse woody debris processes
conservation and management priority areas coastal bluffs restoration techniques
krummholz priority habitats Coastal Dunes and Beaches role in salmon habitat
passerines priority species acreage successional patterns of
plant phenology gradients salmon core areas habitat description stages and structures
predator density SLOSS model Coastal Headlands and Islets status, current and
refugia from predators statewide analysis acreage historical
seasonal migration to Man and the Biosphere habitat description deer, black-tailed
snow-melt and food for herbivores Reserve birds Columbian white-tailed
thermal belts maps mammals mule
threats to wildlife current vegetation Coastal Landscape Analysis and Desert Playa and Salt Scrub
ungulates historic vegetation Management System (CLAMS) Shrublands
wildlife adaptations prioritizing areas coastal upwelling acreage
wildlife and winter recreational species distribution Coastal Zone Management Act habitat description
activities units of analysis (CZMA) dipper, American
wildlife conflicts multiple use protected Columbia River plume disturbance (in managed forests)
winter residents of areas condor, California coarse-scale
Alpine Grasslands and Shrublands quantitative methods conservation, incentives density
acreage complementary or iterative Conservation Reserve Program fine-scale
habitat description analyses (CRP) frequency
alpine-tolerant species scoring techniques and indices strategies intensity
altitudinal migrations of alpine

730
 WILDLIFE IN WESTSIDE AND HIGH MONTANE FORESTS 731

pattern ecological restoration of riparian sharp-tailed grouse dusky Canada

DNA fingerprinting habitats trumpeter swan giant Canada
dolphin, northern-right whale active restoration upland sandpiper greater white-fronted
Pacific white-sided passive or natural restoration Wyoming ground lesser Canada
Risso’s ecosystem based management squirrel Ross’s
donor populations definition yellow-billed cuckoo snow
dove, mourning ecosystem approach extralimital strandings Taverner’s Canada
rock filter concept falcon, gyrfalcon Vancouver Canada
dowitcher, long-billed models, analytical peregrine western Canada
short-billed sustainable development prairie genetic diversity
down wood, classification of ecotones Farm Bill, The gnatcatcher, blue-gray
decay class ecotypes Federal Clean Water Act goat, mountain
in riparian areas edge effects finch, black rosy godwit, bar-tailed
in upland areas effective population size Cassin’s hudsonian
duck, American black egret, cattle gray-crowned rosy marbled
American wigeon great house goldeneye, see duck
Barrow’s goldeneye snowy purple golden-plover, American
blue-winged teal El Niño, events defined fire, and small mammals Pacific
bufflehead and marine birds range goldfinch, American
canvasback and marine mammals wildfires, effects on lesser
cinnamon teal effects on salmon habitat breeding birds gopher, Botta’s (pistol river) pocket
common goldeneye elevational migration fisher Camas pocket
common merganser elk, Rocky Mountain flicker, northern northern pocket
Eurasian wigeon Roosevelt flycatcher, ash-throated Townsend’s pocket
gadwall Endangered Species Act (ESA) Cordilleran western pocket
greater scaup ensatina dusky goshawk, northern
green-winged teal environmental policy requirements gray grasslands
harlequin Environmental Quality Incentives Hammond’s avifauna in westside
hooded merganser Program least reptiles, a link between
lesser scaup ermine olive-sided Grays Harbor Estuary as a stopover
mallard estuaries, Pacific-slope in Pacific Flyway
northern pintail beaches, locations of (figure) willow grazing, effects of livestock on
northern shoveler channel reconfiguration forest, amphibian populations
oldsquaw description of (see Bays and and rangeland grebe, Clark’s
red-breasted merganser Estuaries) conservation laws eared
ring-necked erosive flooding management horned
ruddy habitat components regeneration pied-billed
wood raptor hunting grounds harvest red-necked
dunlin shorebirds structural classes, western
Dwarf Shrub-steppe types of closed canopy grosbeak, black-headed
acreage turbidity maxima old forest, multistory evening
habitat description exotic species, single story pine
eagle, bald accidental introductions stand initiation ground squirrel, see squirrel
golden of stem exclusion open canopy grouse, blue
Eastside Canyon Shrublands Ad Hoc Federal Invasive understory reinitiation ruffed
acreage Alien Species Task Force young forest multistory sage
habitat description amphibian fox, gray sharp-tailed
Eastside Forest and Woodland annuals, invasion of kit spruce
fire suppression galliforms red guillemot, pigeon
habitat features extinction freshwater gull, Bonaparte’s
closed-canopy forest rate physical characteristics California
down wood risk water quality and Franklin’s
grass/forb stages stochastic extinction processes quantity glaucous
shrub/seedling stages demographic cover factors glaucous-winged
snags stochasticity biological factors Heermann’s
management environmental frog, bullfrog herring
natural approach stochasticity Cascades mew
natural disturbance regime genetic stochasticity Columbia spotted ring-billed
status, threats, and trends interactive effects on foothill yellow-legged Sabine’s
timber harvest populations green Thayer’s
wildlife systematic extinction processes northern leopard western
research needs extirpated species in Oregon and/ Oregon spotted habitat,
Eastside Grasslands or Washington Pacific chorus connectivity
acreage American bison Pacific treefrog defined
habitat description bighorn sheep red-legged effectiveness models
Eastside Mixed Conifer Forests California condor tailed elements,
acreage gray wolf fulmar, northern anthropogenic
habitat description grizzly bear gadwall, see duck data matrix (and CD
Eastside Riparian-Wetlands merlin galliforms Rom)
acreage pronghorn antelope geese, Aleutian Canada defined
habitat description sea otter cackling Canada ecological
 732 Wildlife–Habitat Relationships in Oregon and Washington

fire broad-tailed knot, red Marine Shelf

forest/woodland vegetative Calliope La Niña events description
elements rufous lagomorphs acreage
freshwater riparian and aquatic hyporheic zone lake fertilization programs marmot, hoary
bodies ibis, white-faced landscape by design Olympic
major categories and definitions improved pasture, defined land management planning yellow-billed
(table) inbreeding depression lark, horned marten, American
marine Inland Marine Deeper Waters sky martin, purple
energy acreage legacy trees meadowlark, western
forms habitat description lemming, northern bog melanism (in alpine habitats)
intertidal Integrated Pest Management (IPM) life history merganser, common
non-vegetative, abiotic, terrestrial practices adjustments to high altitude hooded
rock crevices intertidal mudflats attribute red-breasted
rocky outcrops and ridges introduced wildlife species, (see also data matrix, (see also CD Rom) metapopulation theory
ranking value of exotic species) lion, mountain migration corridors
for rare species adverse effects of lizard, desert horned migratory land birds
for all species ecological consequences green minimum viable population
for economically of long-nosed leopard mink
important species habitat use by Mojave black collared mockingbird, northern
structural conditions, in Oregon and Washington northern alligator modeling,
association with wildlife, management of sagebrush community and ecosystem
levels defined potential benefits short-horned simulation
closely purposeful introductions side-blotched community structure
associated regulation of southern alligator correlation
confidence invasive species western fence gap analysis
rating irrigation canals Lodgepole Pine Forests and habitat preference
generally jackrabbit, black-tailed Woodlands hierarchy
associated white-tailed acreage life history
present jaeger, long-tailed habitat description statistical classification
data matrix (see also parasitic longspur, lapland optimal foraging
CD Rom) pomarine loon, common defined
defined jay, gray Pacific future directions in modeling
forest pinyon red-throated multiple species approach
shrubland and grassland Steller’s yellow-billed patches
urban western scrub jay lynx rank model
agricultural jetty magpie, black-billed rule induction approach
wildlife, 32 types, see chapter 2 junco, dark-eyed mallard, see ducks spatially explicit
hare, snowshoe kestrel, American mammals, in riparian areas spatially implicit
hawk, Cooper’s key ecological functions, (see also in chaparral habitats trend analysis
ferruginous CD Rom) management activities theoretical Bayesian
northern goshawk classification of (table) land use coarse filter
red-shouldered community patterns managed forests, fine filter
red-tailed critical function management guild
rough-legged defined of habitat capability
sharp-shinned extinct and extirpated species even-age effectiveness
Swainson’s exotic species uneven-age evaluation procedures
Hells Canyon Initiative, The functional, attenuation non- suitability index
hemimetabolous invertebrates bottlenecks or cold spots traditional pattern recognition
Herbaceous Wetlands diversity wood species-habitat matrices
acreage keystones legacies modified grassland habitat
habitat description linkage or hot spots stepwise assessment process defined
herbivore exclosure profiles quantitative objectives species associated with (table)
heron, black-crowned night redundancy marine environments (tables) mole, broad-footed
great blue resilience environmental constraints coast
green richness migration/dispersal capability shrew-mole
herpetofauna diversity roles of species potential and actual threats Townsend’s
in riparian areas web seasonal occurrence of birds and Montane-Coniferous Wetlands
in westside grasslands geographic patterns mammals acreage
of chaparral habitats imperiled functions species adaptations to habitat description
of the Southern Puget Trough management implications marine gardens Montane Mixed Conifer Forest
(table) trophic marine habitats, acreage
herptiles as indicators of the health wildlife species conservation habitat description
of aquatic systems killdeer mechanisms moose
heuristic programming technique kingbird, eastern description of mouse, canyon
high elevation habitats in western seasonal species residency (table) Columbia
Washington and Oregon kingfisher, belted wildlife communities dark kangaroo
holometabolous invertebrates kinglet, golden-crowned Marine Protected Areas (MPAs), deer
horse, feral ruby-crowned defined Great Basin pocket
hummingbird, Allen’s kingsnakes, see snakes in Oregon (figure) house
Anna’s kite, white-tailed in Washington (figure) little pocket
black-chinned kittiwake, black-legged Marine Sanctuaries northern grasshopper
 WILDLIFE IN WESTSIDE AND HIGH MONTANE FORESTS 733

Pacific jumping phalarope, red restoration, passive energy source

pinon red-necked stream enhancement fisheries
western harvest Wilson’s activities habitat
western jumping pheasant, ring-necked Restriction Fragment Length estuary
mudflats and tidal regime Szechwan Polymorphism (RFLP) freshwater
multiple use protected areas phoebe, black ringtail loss/
murre, common Say’s riparian habitat, degradation
murrelet, ancient pig, feral biological diversity
marbled pigeon, band-tailed downed wood agricultural land clearing
muskrat pika, American ecosystem perspective
myotis, see bat pinnipeds flooding channel clearing
National Estuarine Research pintail, see duck functional attributes of (table)
Reserve (NERR) pipistrelle, see bat functions and values development activities
native ungulates pipit, American groundwater chemistry
Natural Area Preserve (NAP) plant alliances human influence on persistent toxin release
Nearshore Marine plant-animal interactions dams
acreage plover, American golden flow regulation splash damming
habitat description black-bellied livestock grazing ocean
waterfowl winter densities (table) Pacific golden logging riparian forests riparian
newt, rough-skinned semipalmated upland land use key ecological functions
nighthawk, common snowy water diversions of
northern harrier, see hawk pocket gopher, see gopher hydrological features nitrogen input
nutcracker, Clark’s polymerase chain reaction (PCR) importance to wildlife and fish nutrient cycling
nuthatch, pygmy Ponderosa Pine-Eastside Oak instream biota phosphorus input
red-breasted Woodlands and Forests keystone species life history stages
white-breasted acreage large woody debris (LWD) carcasses
nutria habitat description litter input freshwater rearing (fry/parr)
Oceanic poorwill, common management considerations incubation (eggs/alevin)
acreage population, microclimate saltwater
habitat description defined nurse logs spawning
oldsquaw, see duck dynamic carrying nutrient dynamics pink
Open Water-Lakes, Rivers, and capacity nutrients input productivity
Streams bottleneck obligates of redd construction
lakes, ponds, and reservoirs viability organic matters, inputs of sockeye
acreage viable population establishment preservation spawning escapement goals
habitat description vulnerability analysis restoration wildlife species relationship with
rivers and streams porcupine, common risk analysis management (figure) (see also CD Rom)
acreage porpoise, Dall’s score cluster (table) salt marsh and roosting shorebirds
habitat description harbor sediment control salvage logging
opossum, Virginia pronghorn, see antelope soils/geomorphology features sand dunes
orca, see killer whale protein electrophoresis streambank stabilization sanderling
orchards/vineyards/nurseries, ptarmigan, vegetation sandpiper, Baird’s
defined white-tailed zone, defined buff-breasted
Oregon silverspot butterfly willow robin, American least
oriole, Bullock’s puffin, tufted ruff pectoral
osprey quail, California salamander, black rock
otter, northern river mountain California slender semipalmated
sea scaled Cascade torrent sharp-tailed
outbreeding depression rabbit, brush clouded solitary
owl, barn European Columbia torrent spotted
barred pygmy Cope’s giant stilt
boreal raccoon del norte upland
burrowing racer, see snake Dunn’s western
flammulated rail, Virginia larch mountain sapsucker, red-breasted
great gray yellow long-toed red-naped
great horned rain bird, see yellow-billed cuckoo northwestern Williamson’s
long-eared random genetic drift Olympic torrent scaup, see duck
northern hawk owl rank model Oregon slender scoter, black
northern pygmy-owl rat, black Pacific giant surf
northern saw-whet California kangaroo Siskiyou Mountains white-winged
short-eared chisel-toothed kangaroo southern torrent sea lion, California
snowy Norway tiger northern (Steller)
spotted Ord’s kangaroo Van Dyke’s seabird nesting colonies
western screech-owl rattlesnake, western, see snakes western red-backed in Oregon (table)
oystercatcher, black raven, common salmon in Washington (table)
Pacific Decadal Oscillation (PDO) redhead as food source for seal, harbor
partridge, gray redpoll, common vertebrates northern fur
pelican, American white redstart, American chinook northern elephant
brown reintroduction programs chum ribbon
petrel, fork-tailed storm resident land birds coho ringed
leach’s storm residential areas core areas semelparity
 734 Wildlife–Habitat Relationships in Oregon and Washington

semi-arid environments, recommendations Columbian ground translocation programs

constraints, consequences, removal of Douglas’ trout, cutthroat
adaptations to (table) restoration techniques eastern fox rainbow
shearwater, black-vented stressors and indicators eastern gray steelhead
Buller’s terrestrial and marine golden-mantled ground turkey, wild
flesh-footed considerations Merriam’s ground turnstone, black
pink-footed Snag Recruitment Simulator (SRS) northern flying ruddy
short-tailed Snag Dynamics Projection Model piute ground turtle,
sooty (SDPM) red green
sheep, California bighorn snake, Townsend’s ground leatherback
Rocky Mountain bighorn California Mountain tri-colored loggerhead
shoveler, northern, see ducks kingsnake Washington’s ground North American soft shell
shrew, Baird’s common garter western gray Pacific ridley
fog common kingsnake white-tailed antelope painted
masked common viper Wyoming ground red-eared slider
Merriam’s gopher starling, European snapping
montane night steelhead, see trout western pond
Pacific northwestern garter stilt, black-necked unimproved pasture, defined
Pacific water Pacific Coast aquatic garter stochastic extinction processes, see Upland Aspen Forests
Preble’s plateau striped whipsnake extinction acreage
pygmy racer storm petrel, black habitat description
Trowbridge’s ringneck Subalpine Parklands Urban and Mixed Environs
vagrant rubber boa acreage acreage
water sharptail habitat description cemeteries
shrew-mole, see mole western ground surfbird composition
shrike, loggerhead western rattlesnake sustainable ecosystems ecological features of
northern western terrestrial garter swallow, bank effects of management and
Shrub-steppe western whiptail barn anthropogenic impacts
acreage snipe, common cliff environmental conditions/
bird communities solitaire, Townsend’s northern rough-winged constraints (table)
composition sora tree fragmentation
conversion of herb layer to Southwest Oregon Mixed Conifer- violet-green fringe areas
cheatgrass Hardwood Forest swan, mute golf courses
effects of management and acreage trumpeter gradient concept
anthropogenic impacts habitat description tundra growth management
fire/cheatgrass cycle spadefoot, Great Basin, see toad swift, black clustering
habitat description sparrow, American tree Vaux’s controlling urban sprawl
habitat fragmentation black-throated white-throated easement rights
livestock grazing Brewer’s talus educational programs
mammal communities chipping tanager, western greenways and corridors
management clay-colored tatanka, see American bison landscape by design
natural disturbance regime fox tattler, wandering land-use planning tools
organizing principles golden-crowned taxonomic classifications, see Oregon Statewide Planning Goal 5
reptile and amphibian communities grasshopper chapter 1 planning for open spaces
research needs Harris’s teal, see ducks protecting fish and wildlife habitat
species associated with (table) house tern, Arctic transfer of development rights
silvertip, see bear, grizzly lark black Washington Growth Management
silvicultural methods Lincoln’s Caspian Act (GMA)
green tree retention sage common habitat description
long rotations savannah elegant high-density zone
structural retention song Forster’s description
thinning swamp Territorial Sea Plan food
variable retention harvest system Vesper thermocline vegetation
siskin, Eurasian white-crowned thrasher, sage wildlife
pine white-throated threatened species history of
skink, western Species At Risk Advisor (SARA) thrush, hermit low density zone
skua, South Polar species, known to be imperiled song description
skunk, striped likely very secure Swainson’s food
western spotted occurrence status categories varied vegetation
skylark, Eurasian accidental northern waterthrush wildlife
snag and down wood extirpated timber wolf, see gray wolf management,
and wildlife species introduced titmouse, juniper active vegetation
data sources occurs oak regulatory buffer
ecological processes reintroduced toad, stormwater detention facilities
example queries richness in marine habitats (table) Great Basin spadefoot medium density zone
geographical extent and with intermediate security western description
context spirit bird, see California condor Woodhouse’s food
habitat assessment squirrel, Arctic ground towhee, California vegetation
habitat elements Belding’s green-tailed wildlife
in riparian forests California ground spotted native plants propagation
management Cascade golden-mantled ground toxicological risk assessments of
 WILDLIFE IN WESTSIDE AND HIGH MONTANE FORESTS 735

parks/open spaces tree size (table) Lewis’s orange-crowned

research needs wildlife assemblages pileated palm
roads Westside Grasslands red-cockaded Townsend’s
status of acreage three-toed Wilson’s
urbanization, effects of agricultural development white-headed yellow
channel instability endemics, extirpations, and woodrat, bushy-tailed yellow-rumped
ecological consequences (table) peripheral species desert garden
faunal diversity declines exotic species dusky-footed water regime alteration
habitat loss fire suppression wren, Bewick’s waxwing, Bohemian
on raptors habitat description canyon cedar
reduction in water quality habitat types house weasel, long-tailed
stream channel degradation or loss chaparral marsh Western Juniper and Mountain
water level fluctuations grasslands rock Mahogany Woodlands
woody debris loss livestock grazing winter acreage
veery management and conservation of wrentit biotic integrity
viability goals urbanization yellowlegs, greater ecoregions
viper, see snakes wildlife communities lesser habitat description
vireo, Cassin’s birds egetation habitat elements (table)
Hutton’s herpetofauna wildlife management
plumbeous mammals management, timber harvest
red-eyed Westside Lowlands Conifer- active vegetation alternative silviculture
warbling Hardwood Forest regulatory buffer downed wood recruitment
vole, California acreage stormwater detention facilities stream buffers
creeping habitat description medium density zone organismal relationships (table)
gray-tailed Westside Oak and Dry Douglas-Fir description provincial patterns
heather Forests/Woodlands food snags
long-tailed acreage vegetation species occurrences (table)
meadow habitat description wildlife trophic relationships (table)
montane western prairie chicken, see sharp- native plants propagation wildlife species association with
red tree tailed grouse of tree size (table)
sagebrush Westside Riparian-Wetlands parks/open spaces wildlife assemblages
southern red-backed acreage research needs Westside Grasslands
Townsend’s habitat description roads acreage
water Wetland Reserve Program status of agricultural development
western red-backed wetlands, defined urbanization, effects of endemics, extirpations, and
white-footed whale, blue channel instability peripheral species
vulture, turkey gray ecological consequences (table) exotic species
warbler, black-throated humpback faunal diversity declines fire suppression
garden killer habitat loss habitat description
hermit minke on raptors habitat types
Macgillvray’s North Pacific bottle-nosed reduction in water quality chaparral
Nashville short-finned pilot stream channel degradation or loss grasslands
orange-crowned sperm water level fluctuations livestock grazing
palm whimbrel woody debris loss management and conservation of
Townsend’s white bear, see grizzly bear veery urbanization
Wilson’s white swan, see trumpeter swan viability goals wildlife communities
yellow whiptail, see snakes viper, see snakes birds
yellow-rumped wigeon, see ducks vireo, Cassin’s herpetofauna
garden wildfire, see fire Hutton’s mammals
water regime alteration wildlife, plumbeous Westside Lowlands Conifer-
waxwing, Bohemian conservation (coarse filter concept) red-eyed Hardwood Forest
cedar ecosystem approach warbling acreage
weasel, long-tailed fine filter concept vole, California habitat description
Western Juniper and Mountain habitat, defined creeping Westside Oak and Dry Douglas-Fir
Mahogany Woodlands goals gray-tailed Forests/Woodlands
acreage non-spatial heather acreage
biotic integrity spatial (adjacency requirements) long-tailed habitat description
ecoregions types, 32 defined, see chapter 2 meadow western prairie chicken, see sharp-
habitat description minimum patch size montane tailed grouse
habitat elements (table) multiple species approach red tree Westside Riparian-Wetlands
management willet sagebrush acreage
timber harvest willow grouse, see sharp-tailed southern red-backed habitat description
alternative silviculture grouse Townsend’s Wetland Reserve Program
downed wood recruitment wolf, gray water wetlands, defined
stream buffers wolverine western red-backed whale, blue
organismal relationships (table) wood decay, see decaying wood white-footed gray
provincial patterns wood pewee, western vulture, turkey humpback
snags woodpecker, acorn warbler, black-throated killer
species occurrences (table) black-backed garden minke
trophic relationships (table) downy hermit North Pacific bottle-nosed
wildlife species association with hairy Macgillvray’s short-finned pilot
Nashville sperm
 736 Wildlife–Habitat Relationships in Oregon and Washington

whimbrel woody debris loss livestock grazing

white bear, see grizzly bear veery management and conservation of
white swan, see trumpeter swan viability goals urbanization
whiptail, see snakes viper, see snakes wildlife communities
wigeon, see ducks vireo, Cassin’s birds
wildfire, see fire Hutton’s herpetofauna
wildlife, plumbeous mammals
conservation (coarse filter concept) red-eyed Westside Lowlands Conifer-
ecosystem approach warbling Hardwood Forest
fine filter concept vole, California acreage
habitat, defined creeping habitat description
goals gray-tailed Westside Oak and Dry Douglas-Fir
non-spatial heather Forests/Woodlands
spatial (adjacency requirements) long-tailed acreage
types, 32 defined, see chapter 2 meadow habitat description
minimum patch size montane western prairie chicken, see sharp-
multiple species approach red tree tailed grouse
willet sagebrush Westside Riparian-Wetlands
willow grouse, see sharp-tailed southern red-backed acreage
grouse Townsend’s habitat description
wolf, gray water Wetland Reserve Program
wolverine western red-backed wetlands, defined
wood decay, see decaying wood white-footed whale, blue
wood pewee, western vulture, turkey gray
woodpecker, acorn warbler, black-throated humpback
black-backed garden killer
downy hermit minke
hairy Macgillvray’s North Pacific bottle-nosed
Lewis’s Nashville short-finned pilot
pileated orange-crowned sperm
red-cockaded palm whimbrel
three-toed Townsend’s white bear, see grizzly bear
white-headed Wilson’s
woodrat, bushy-tailed yellow
desert yellow-rumped
dusky-footed garden
wren, Bewick’s water regime alteration
canyon waxwing, Bohemian
house cedar
marsh weasel, long-tailed
rock Western Juniper and Mountain
winter Mahogany Woodlands
wrentit acreage
yellowlegs, greater biotic integrity
lesser ecoregions
egetation habitat description
wildlife habitat elements (table)
management, management
active vegetation timber harvest
regulatory buffer alternative silviculture
stormwater detention facilities downed wood recruitment
medium density zone stream buffers
description organismal relationships (table)
food provincial patterns
vegetation snags
wildlife species occurrences (table)
native plants propagation trophic relationships (table)
of wildlife species association with
parks/open spaces tree size (table)
research needs wildlife assemblages
roads Westside Grasslands
status of acreage
urbanization, effects of agricultural development
channel instability endemics, extirpations, and
ecological consequences (table) peripheral species
faunal diversity declines exotic species
habitat loss fire suppression
on raptors habitat description
reduction in water quality habitat types
stream channel degradation or loss chaparral
water level fluctuations grasslands