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Milton W. Weller-Wetland Birds - Habitat Resources and Conservation Implications - Cambridge University Press (1999)
Milton W. Weller-Wetland Birds - Habitat Resources and Conservation Implications - Cambridge University Press (1999)
Habitat Resources
and Conservation
Implications
Milton W. Weller
Wetland Birds
Wetland Birds
Habitat Resources and Conservation Implications
milton w. weller
Contents
Introduction []
. Classification of wetlands []
. Factors influencing wetland formation []
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Water variability []
Temporal changes in wetland vegetation []
Plant succession []
Bird responses to short-term fluctuations in water regimes []
Seasonality of bird responses to water and vegetation []
Bird responses to long-term changes in water and vegetation []
. Wetland losses []
.Human impacts on existing wetlands []
. Direct disturbance of waterbirds in remaining wetlands []
.Unwanted biodiversity []
Outlook []
Epilogue []
Appendix Scientific names of birds and bird groups []
Appendix Scientific names of animals and animal groups other than birds []
Appendix Scientific names of plants and plant groups []
Index of birds and bird groups []
Subject index []
Plates
Plate I
Plate II
Plate III
Plate IV
Plate V
Plate VI
Preface
Wetland birds have long attracted the attention of the public and scientists
because of their beauty, abundance, visibility, and social behavior, as well as for
their recreational and economic importance. Recently, they have become of
interest as indicators of wetland quality, and as parameters of restoration
success and regional biodiversity. Recognition of the importance of wetland
habitat resources as the dominant influence on the presence and repeated use
of wetlands by birds has come slowly, but sufficient information is now available that it seems timely to summarize what we know, and to speculate on what
we think we know, as a means of seeking direction for future work. This book is
partly a product of my own interest in patterns of wetland habitat use by birds
in relation to wetland dynamics. In addition, nonspecialists have been seeking
help in using birds as indicators of wetland function or as measures of success
in wetland management, restoration, and creation. I have tried to make the
book understandable to students of various ages and stages who are interested
in wetland birds and their conservation, but some may find more detail than
they like. Others with greater knowledge of certain topics will find it advantageous to skip from the known to the less well-known subject matter. The literature on wetland birds now is enormous, but to avoid constant interruption of
the text, I have cited literature in some chapters less than some may prefer.
However, some chapters list further reading material not cited in the text; in
both sections I have tried to include classics in the subject area and more recent
papers or books that provide summaries of pertinent literature. Words or
phrases that are important to the understanding of the topic and that are in
common use in the field are in bold-face type and usually are defined directly
or by example.
The focus of the book is on how wetland habitat resources influence its use
by birds in general rather than on any specific taxonomic group of birds, or
their breeding biology a separate but massive subject in itself. I have used the
term wetland birds in the title to provide a habitat focus that considers birds
adapted to a wide range of relatively shallow waters through to those in wetsoil habitats. The term waterbirds is often used for these various groups, commonly identified by habitat- or resource-related descriptors: divers (especially
loons), waterfowl (wildfowl in Europe, but also used broadly like waterbirds),
Preface
xi
waders (herons in Europe), shorebirds (waders in Europe), littoral or wateredge birds, aquatic birds, coastal birds, and estuarine birds. Some of these
groups of birds avoid saline wetlands but may use coastal or inland wetlands as
long as they are freshwater; others use mainly saline wetlands. Some species
may use either fresh or saline wetlands but use is influenced by factors other
than salinity: for example, life cycle function such as breeding or nonbreeding
periods. The book does not treat species of the deep-ocean (i.e. oceanic,
pelagic, or seabirds) except as they come to coastal wetlands to nest, feed, or
roost in shallow and food-rich waters. The last usually nest in groups and have
attracted the attention of ecologists and ornithologists, who have focused on
the social behavior of colonial waterbirds, a term that includes many freshwater species as well. Overlap in categorizing such birds by habitat characteristics
reflects both the dynamic nature of wetlands and the evolution of great adaptability to exploit wetland resources wherever they are.
To aid the interested but less experienced, I have included a chapter
describing the various taxonomic groups that use wetlands, with emphasis on
habitat adaptations, foods, and habitat-influenced breeding biology. The
reader will find a wide range of birds, from those groups where most species
are obligate users of wetland habitats through taxonomic orders or families
where only a few species are so restricted to those that are regular users but do
so facultatively. Obviously, almost any bird that happens to be at a wetland
edge may occasionally exploit wetland resources such as water or food opportunistically. This range of variation may disturb some readers who search for
simple patterns, but it reflects the dynamics of many wetland habitats and the
amazing flexibility of birds. Although photographs throughout the book
provide examples of activities and adaptations of various wetland bird species,
the book does not deal with identification. Several bird guides covering
specific taxonomic groups or geographic areas have been included in the bibliographies. To save space and simplify reading, scientific names are given in
Appendix for birds, Appendix for other animals, and Appendix for plants.
Most books written about the taxa of birds that regularly use wetlands have
been taxonomically structured even when emphasizing life-history information. Moreover, habitat descriptions have been limited or lacking in many such
books partly because of the absence of information on species and adaptations
to resources such as food, but also because of the difficulty of finding widely
recognized descriptive terms. This is rapidly changing because of current
interest in the conservation of natural habitats as the essential approach to preserving threatened species and maintaining regional biodiversity. This muchneeded habitat focus has increased the need for information on the
environmental resources likely to be found within the habitat. Several recent
books on wetland birds reflect this growing habitat interest, and hopefully this
discussion as well as the publications cited will encourage further descriptive
as well as integrative work.
After an introduction to habitat concepts essential to provide an equal
xii
Preface
footing for all readers, subsequent chapters of the book will outline wetland
diversity and classification, review the major groups of bird that use wetlands,
consider how wetland features influence bird biology and adaptation, elucidate how birds can influence wetlands, examine methods of describing potential wetland microhabitats, and identify how we might relate changes in
wetland bird communities to the dynamics of habitat resources over time and
space. Opportunistically throughout the text, I shall try to relate patterns to
potential application in the conservation and management of wetland birds,
and additional issues will be summarized in several chapters near the end of
the book.
This book reflects my biases by virtue of experience with certain taxonomic
groups, geographic areas, and literature, but I hope the examples reflect a view
of species as part of a community and consider the features and dynamics of
habitats as driving influences on the evolution of bird groupings as well as on
species attributes. For groups I know poorly, I have tried to incorporate examples documented in the literature. I suspect this will satisfy few specialists in
those areas, but I hope there is enough information to allow us to focus on
general patterns. Many patterns will serve only as hypotheses for future
testing, which should help determine not only what occurs but how and why
such patterns exist. Therefore, it is especially important for readers to evaluate
general statements of apparent patterns analytically.
Asking general questions important in wetland ecology may be relatively
easy but answering them is not! What is it about wetlands that produces such
concentrations of birds, both in breeding and nonbreeding periods? Unlike
our expectation of terrestrial birds, why are they often present in one year and
not another? Why are many of the groups so widely distributed and among the
most mobile animals in existence, whereas other species have modest ranges
and are so scarce that the list of endangered wetland species is long? How has
this habitat influenced breeding biology and life-history strategies? Why are
some birds not represented among wetland users, and what does this tell us
about the wetland ecosystem? How can we apply our knowledge, even when
minimal, in a conservation strategy of protection, management, or restoration?
Persons wanting to preserve wetland bird communities, enhance bird populations, preserve species diversity through management, or to use birds as an
index for assessing habitat quality usually want to know which single habitat
feature is most important to birds, how data can be gathered quickly, and how
this knowledge can be applied in a simple and practical way. Unfortunately,
ecological problem-solving is neither simple nor conclusive. The answer
requires not only a significant amount of life-history information about each
bird species or group in question, but consideration of the quantity, quality,
and dispersion of biological resources such as food, vegetation, and other
animals; additionally, a knowledge of physical features such as water, ice, and
geomorphology are essential. Issues of spatial scale (size, shape, and disper-
Preface
xiii
sion of components) and temporal scale (daily, seasonal, annual, irregular but
still time-related influences) complicate the picture further. Therefore, most
problems are, as in all natural ecosystems, multivariate and difficult to test and
to use to draw indisputable conclusions. Much of the book will emphasize
understanding such difficult questions, but I suspect that I will answer too few
of them. That is, however, part of the strategy of the book: most generalizations
induce questions on the basis of personal experience to the ultimate good of
science. To that end, I do not hesitate to generalize with the hope that such
statements result in challenge, additional insights, observation, and, ultimately, new information.
xiv
Preface
Acknowledgments
Acknowledgments
xv
1
Introduction
Introduction
Specific foods
Feeding tactics
Feeding flights
Roosting patterns
Water-resistant plumage
Behavioral adaptations of preening and drying
Specialization for general types of food
Physiology: respiratory, digestive tract
Breeding strategies
Social behavior: spacing, aggression,
Water depth adaptations
Wetland types, water dynamics and salinity
Long-range mobility/resource exploitation
systems must be geared to acuity and protection both in water and air, and to
eye accommodation of as much as -fold, which allows a quick change of
focus from near to far in seeking food, and to binocular vision, which occurs in
herons and kingfishers (Campbell and Lack ). Birds that feed at night, like
skimmers, have eyes that are better adapted for nocturnal vision compared
with diurnal species (Rojas, McNeil and Cabana ). Respiratory physiology
differs dramatically in those species that engage in long-term and deep diving.
Although amateurs by comparison with large penguins, long-tailed ducks and
loons have been recorded in fishermans nets at and feet ( and m),
respectively, during winter when they may leave wetland fringes for deep water
(Schorger ). Morphologically, adaptations include bills that strain, peck,
spear, store, and grab, and feet that allow swimming, diving, walking on
mudflats, wading, or grabbing and holding fish. Not only do body parts differ
in general form , but size of bills, legs, and flight patterns differ across a gradient of wetland edges, as noted in both North American and European peatlands (Niemi ). Additionally, special feathers and plumage designs ensure
waterproofing under the most severe conditions.
Food types also are diverse and demand highly specialized digestive
systems: some wetland birds specialize on plant diets, some on animal, some
use both, and most switch foods seasonally as induced by needs for breeding or
energy for migration. Water also provides protection from predators, and
many waterbirds capitalize on the protection combined with the rich latesummer resources to molt all their wing feathers simultaneously. Most terrestrial birds or birds that feed on-the-wing must do this annual molt a few
feathers at a time, retaining flight capability while reducing energy demands.
As a result of these adaptations, birds are better equipped as a group to
exploit wetland resources wherever they occur than any other animals,
except some mobile insects that use similar approaches but on a smaller scale.
Because birds are larger, more colorful, fewer in number of species and, therefore, better known, they are more often used as indicators of conditions within
a wetland ecosystem.
Introduction
layering in herbaceous wetlands includes vegetation that has less actual height
change but is of no less important to water-level birds; also it includes physical
components like shoreline, vegetation layers, water column depth and structure, and wetland substrate (Fig. .) Habitat diversity is a product of various
biological and physical features that make multiple sub- or microhabitats
available in a small area which are attractive to different species or groups of
birds. Here, we will most commonly denote this diversity as species richness
(i.e., number of species found in an area) and discuss other elements of diversity later.
The habitat concept also infers that this spatial site meets the psychological perception by a bird species, termed umwelt by German workers to infer
the animals view of its environment as influenced by its special senses; Klopfer
() pointed out that this means mostly visual clues in birds. A simple model
of some features of habitat recognition by birds was developed to demonstrate
the role that habitat plays in separating closely related species such as pipits, in
which one species used wet meadows and others select more vegetated and elevated sites (Svardson ). These perspectives resulted in the concept of
habitat patterns as a way of describing features of habitat structure that we
humans recognize as important to birds but have difficulty quantifying (e.g.,
water, open horizon, plant height or density, etc.)(Williams, Russell and Seitz
). Such clues to suitable habitat presumably are perceived from the air as
well as on-water, in-water, and underwater and consider both visual appearance and feel. One clue to such visual components comes from accidents made
by night-time migrants that land on wet streets and are stranded until daylight
and even then may have difficulty taking off.
Based on studies of the behavior of young waterbirds, we also expect that
some of this response to habitat is innate (i.e., instinctive in newly hatched
birds) and some may be learned in early imprinting to environmental features.
For example, some workers have noted that young, captive ducklings
imprinted on humans still frequent species-oriented habitat preferences when
possible (Fabricius ). The influence of learning was suggested to me early
in my studies when I found the nest of a banded Redhead Duck that had been
hand-reared and released the previous year but returned to nest in flooded
Introduction
forest rather than in the typical open marsh where the egg had been collected
and where adults normally breed. Did this forest canopy provide the habitat
image acquired during its early life in an incubator and covered rearing pen?
It is obvious from the above that birds make decisions about which habitat
to use based on innate and learned behavior and by trial-and-error testing of
features and resources. They go through a process of selection and ultimately
use of a chosen habitat. But habitat use may not infer habitat preference
unless a bird has choices including the best (i.e., where they are most successful) a most difficult thing to demonstrate unless one knows all of the birds
requirements.
Habitat is a general, descriptive term often used without stated scales; it is
enhanced by the addition of temporal scales essential to reflect annual lifecycle stages, and by spatial scales to denote structural differences or geographical coverage. Considerable effort now is directed toward larger scales termed
landscape because of current technological advances both in computer-based
imagery and in statistical methodology (Forman and Godron ). Several
spatially separated habitat units may be necessary to complete the annual
cycle, even in nonmigratory but mobile species, and all contribute to reproductive success and population maintenance. However, the habitat structure
of these sites may differ markedly, as may food and other resources. Compare,
for example, the similarity of wooded feeding and resting sites used by migratory North American Wood Ducks during either breeding or wintering with
the habitats of Redhead ducks, which may dive for invertebrate foods in
densely vegetated prairie potholes during the breeding season but dabble for
seagrass rhizomes in hypersaline estuaries during winter.
Regardless of the temperature, probably fewer wetland birds reside yearround in small home ranges than is common to terrestrial species, because seasonal changes in wetlands are so dramatic. Water levels rarely remain the same
in most wetland types: vegetation dies and decomposes; foods such as seeds,
foliage, and invertebrates vary in abundance and distribution; water temperatures influence oxygen levels and thereby aquatic animal abundance; and
freeze-ups severely affect what birds can do. Often, the changes are simply less
conspicuous and reflect distribution of resources on a smaller scale, demanding that investigators achieve more precise assessment.
Whereas the term habitat deals with spatial and structural components of
the living space, the word niche (and especially ecological niche) is a concept
describing the role of a species in a community of organisms living together.
The meaning of this term was broadened by a multidimensional perspective of
factors that reflect the environmental requirements of the species (Hutchinson
). Despite many attempts to clarify the term, uncertainty and confusion
still exist (Begon et al. , Brown , Odum , Patten and Auble ,
Whittaker et al. ), but that is not uncommon with theoretical concepts. For
our purposes, its importance is mainly to denote what the species does and
how it lives in the community. When used in reference to food-chains and
Introduction
could talk about Muskrats, Common Carp, and swans as a single guild grazing
in shallow water on submergent vegetation, but that is perhaps too gross to be
useful. More often, one identifies a group of more similar species that exploit a
resource such as food or nest sites in a prescribed habitat such as a stand of
emergent vegetation. Thus, we can focus on patterns of use based on how they
operate (e.g., swimming invertebrate strainers or wading piscivores) rather
than their taxonomic relationship (e.g., ducks, herons, sparrows).
Having given the background of essential tools used in the book as well as
clarifying terms, I will next introduce the habitat of focus, wetlands. It is
important, however, that we do not fall into the trap of ignoring the many
other influences on the evolution of the features and species of birds in these
habitats and which will influence their success over time. Birds that cannot
succeed at maintaining a population do not use a habitat for long, but such
deterministic influences may be genetically based morphological and anatomical features of importance and other environmental influences that are rarely
discussed in relation to habitat. Several simple models have been used by
various workers to place these issues in perspective in the life history of the
species and to outline a framework for the analysis of interactions and relative
importance of habitat features that determine the presence and success of the
species (e.g., Karr , Price and Bock )(Fig. .). But few facets of the life
history of obligate wetland birds are not influenced by water and wetland
resources.
References
Aldrich, J. W.(). Birds of deciduous forest aquatic succession. Wilson Bulletin ,
.
Allen, A. A. (). The red-winged blackbird, a study in the ecology of a cat-tail
marsh. Proceedings of the Linnaean Society of New York , .
Allen, R. P. (). The roseate spoonbill. Research Report No. . New York: National
Audubon Society.
Bailey, R. G. (). Descriptions of the ecoregions of the United States, nd edn,
Miscellaneous Publication No. . Washington, DC: US Forest Service.
Beecher, W. J. (). Nesting birds and the vegetative substrate. Chicago, IL: Chicago
Ornithological Society.
Begon, M., Harper, J. L., and Townsend, C. R. (). Ecology, rd edn. Oxford:
Blackwell Science.
Bennett, L. J. (). The blue-winged teal: its ecology and management. Ames, IA:
Collegiate Press.
References
Introduction
References
2
Wetlands: what, where, and why
. Classification of wetlands
Wetlands are not easily defined because they range from near-terrestrial to the
aquatic, and because many are dynamic in water regime and, therefore, variable in vegetation patterns and bird use. Some vary rather predictably and
dramatically by season or year, and many also are subject to long-term variation owing to large-scale climate patterns and cycles. Therefore, wetlands are
sometimes not recognizable by standing water, and their vegetation can be
equally deceiving. In addition to the presence of shallow water or wet soil
periodically, two other key features of wetlands are the periodic presence of
water-adapted plants (hydrophytes), which range from mosses to giant trees,
and hydric soil, with biochemical features influenced by anaerobic conditions
of flooding (Cowardin et al. ).
Our interest here is mainly in wetlands as biological communities, for
which the Cowardin et al. () classification is mandated in regulatory situations in the United States, and is also now popular worldwide. It involves five
systems (not to be confused with ecosystem) delimited in most cases by
Classification of wetlands
uplands on the shallow side and by prescribed water depth (m), source of
salinity (coastal or inland), and vegetation patterns in the deeper portion. The
five systems are Marine (shallow coastal saltwater), Estuarine (brackish
coastal water), Lacustrine (relatively shallow, open, freshwater lakes or their
sparsely vegetated margins), Palustrine (marshy fresh or inland saline waters
or the more dense shoreward vegetated margins of larger water bodies), and
Riverine (used in this system to mean those wetlands found only within the
river channel, as opposed to river-formed wetlands). Hierarchical subdivisions within each system incorporate terms that help describe the habitat
structure or vegetation of the wetland type (Table. .). For example, a
Palustrine wetland could be dominated by mosslichens (as in a bog), by
herbaceous, emergent vegetation (e.g., cattail marsh), by shrubs or scrub
(shrub swamp), or by water-tolerant trees (wooded or forested swamp or
bottomland forests). Examples of some of these systems and classes are evident
in photos throughout the book particularly in those of Plate I. These dominant
vegetation types are products of several factors: the presence of water over time
(hydroperiod), water depth, seasonality, temperature regimes in the locale,
and other factors that will become obvious later. But in all cases, they form
diverse habitats attractive to different bird and other animal species.
The Canadian wetland classification system is more strongly influenced by
cold-climate bog wetlands but uses commonly recognized terms such as bog,
fen, swamp, marsh, and shallow water (National Wetland Working Group
). Regional denotations such as arctic, coastal, and similar terms help to
focus on the extensive boreal areas so dominated by wetlands. Similar interests
are evident in several Scandinavian countries, where limnological terms are
favored that reflect long-term succession from deep, clear lakes poor in nutrients (oligotrophic) to the more shallow, vegetation-and nutrient-rich lakes of
marshes (eutrophic)(e.g., Kauppinen and Vaisanen ). Several of these
publications trace the history of the development of various classification
systems devised, and the terms are many. In contrast, the Cowardin system was
designed to eliminate earlier and often conflicting terms and create a hierarchical approach that allows any subcategories which may be necessary for the
description and differentiation of wetlands.
The Cowardin system considers as wetlands areas that are not in a discrete
basin but which are wet long enough to induce wetland plant or animal
response. While most wetlands are basins that capture water from precipitation, flooding, or underground aquifers, there are flat areas that hold water for
a long time or hillside slopes in hilly regions or where underground water percolation reaches plant root systems sometimes without even appearing wet
on the surface. Thus, water and the geomorphic setting drive the origin of
wetland communities, and these causal and formative processes are best elaborated in the Hydrogeomorphic Method (HGM)(Brinson ). There are
five categories: depression or basin wetlands, which may receive water from
surface runoff or groundwater; fringe wetlands, which develop along large
Table . The division of wetlands into five systems with subsystems where they exist and
examples of classes useful as habitat descriptors
System
Subsystem
Classes
Marine
Subtidal
Rock
Intertidal
Unconsolidated
Aquatic bed
Reef
Estuarine
Subtidal
Rock
Intertidal
Unconsolidated
Aquatic bed
Reef
Emergents
Shrubscrub
Forested
Riverine
Tidal
Rock
Lower perennial
Unconsolidated
Upper perennial
Aquatic bed
Intermittent
Emergent
Streambed
Lacustrine
Limnetic
Rock
Littoral
Unconsolidated
Aquatic bed
Emergent
Palustrine
(None)
Rock
Unconsolidated
Aquatic bed
Moss-lichen
Emergent
Shrubscrub
Forested
lakes or oceans; and riverine wetlands (typically linear), which are a product of
running water (differing from the Cowardin System, which includes only the
stream channel). Less well-known are groundwater slope wetlands, which
have wet soils or actual discharge areas where the water induces hydrophyte
growth even on hillsides, and flatlands, where high rainfall runs off very slowly
and is lost by permeating the substrate but also by evaporation and plant transpiration.
Classification of wetlands
For clarification, ponds and lakes are common and often poorly defined
terms for more open water-bodies with abrupt banks that may have little or no
emergent vegetation. Cowardin et al. () separated ponds from lakes on the
basis of size, using ha ( acres) as the breaking point. The term pond is often
used for small, water-bodies with abrupt shorelines, deep basins, and open
water (only for constructed water areas in some regions). Lakes typically differ
from wetlands in having a higher percentage of open water, little emergent
vegetation, and well-defined shorelines produced by water- or ice-formed
boundaries or banks.
The boundary between a wetland and the upland is important ecologically
because it forms a transition zone of varying water regimes and plant communities. At this point, various microhabitats are created and selected for by
wetland birds and by those species that tap resources of both upland and
wetland vegetation (Fig. .). In wetlands with dynamic water regimes, edges
often change seasonally and annually, and birds favoring that edge select
habitat upslope or downslope to follow features or resources important to
them. Boundaries are of concern in a conservation and regulatory sense
because they influence decisions on land-use, and there are well-described
techniques for delineation that involve ecological measures of plant species
richness and composition change over the edge gradient (Federal Interagency
Committee for Wetland Delineation ).
and lagoons. This same patterns closes passes and channels made by other
forces such as hurricanes and severe winter storms (Chabreck ). Over
time, and often with sea-level change and climate change, particles may be
deposited as beaches. Old beach rows and declining seas levels result in what in
western Louisiana and extreme eastern Texas are called Chenier Marsh, named
after the wooded ridges or cheniers that really are ancient beach ridges; these
parallel barriers result in a series of fringing marshes that become more
shallow and fresher in more shoreward areas. Still more ancient river outpourings resulted in ancient depositions that subsequently are moved far
inland by coastal winds, forming sand/silt dune and swale areas, so that freshwater ponds and lakes may be common even along rather dry coastlines.
Such beach/ridges structures also are major influences on wetland formation at various scales along large lakes such as the Great Lakes of North
America (Prince and DItri ). In the bordering states and provinces, much
of which was once a part of post-glacial Lake Agassiz, lake decline left ancient
beaches that major highways cross without most drivers being aware of their
existence, history, or formation, and major wetland areas between these
beaches (Glaser ).
Marine coasts may demonstrate different profiles dependent upon shoreline materials and gradient (e.g., cliff versus mudflat), which affect how long
tidewater remains. Shoreline mudflats and sandbars are especially crucial to
shorebirds, gulls, and terns. Substrate characteristics influence how long water
is retained in the sand, silt, or mud, and, therefore, how suitable it is for prey
such as the clams and worms that are so important to birds. These features of
coastal wetlands strongly influence bird species composition, bird density, and
timing of foraging and roosting/resting activities.
A unique wetland region, the coastal tundra of the Arctic, shows the importance of physical influences on wetland development. These open but shallow
wetlands are formed on permafrost by solar melting of the substrate wherever
water collects and gathers heat (Bergman et al. ). Once formed, they tend
to enlarge both by melting of the substrate and by wind erosion. But these wetlands can be huge but very shallow because of the vertical temperature equilibrium between the water (in summer) and the underwater ice base that remains
all year. With precipitation of less than cm per year, water in the shallow
basins is derived mostly from snowfall driven by winds. Once in the pools,
evaporation results in drying so few plants grow in the water, but peat
accumulation forms substrates rich in invertebrates, and thereby attractive to
skimming and probing birds.
The geological processes that form wetlands and lakes also include tectonic
forces such as major earth-shaping forces; earthquakes and volcanic action are
localized but dramatic examples. Several large inland cypress swamps
(Reelfoot Lake of northwestern Tennessee and Caddo Lake of eastern Texas
and western Louisiana) may have been formed by earthquakes as recently as
the s. Earthquakes have modified extensive wetland areas in the Copper
River delta of Alaska. Lakes with associated wetlands areas have been formed
by volcanic actions in the Galapagos Islands and lost and reborn in the Mount
St. Helens earthquake in southern Washington.
Biological forces also influence and, in some cases, actually form locally
abundant and important wetlands. The most common animal that forms wetlands is the beaver, which is widespread over North America and can have
major impacts on small stream systems. They are unpopular because they kill
trees through flooding and cutting, but this is part of a process of succession
that produces other wetland plant species in the resulting flooded areas. They
are regarded as major stream-flow regulators by some hydraulic engineers.
Alligators in southern marshes create openings in herbaceous vegetation and
dig holes and burrows that strongly influence the presence of open water and
the potential survival of fish, snakes and amphibians, and insects in times of
drought (Kushlan ).
Plant production itself provides a water-holding mechanism through the
development of wet, organic soils. Bogs grow on water-holding substrates,
sometimes forming islands with surrounding moats of water and often
forming patterned ground where slope and peat are involved (Damman and
French , Glaser ). A similar but more dynamic process is the development of floating islands of grass mats, common at flooding stages of herbaceous marshes or swamps, tree islands, which also include herbaceous plants
(e.g., the Florida Everglades), and tropical mangroves, which may establish on
such islands (Odum, McIvor, and Smith ).
In these ways, wetland habitats are products of many physical and biological processes and events at various scales, and they are continuous. It is mainly
our short-term and often static view that leaves an impression of wetlands as
fixed entities. The impact of human development has often changed many of
the influential parameters, and especially the rates at which such changes take
place.
Once wetlands have formed, many factors influence the integrity,
resilience, and existence of these biotic communities; their productivity
influences which regional bird species use them, in which season of the year,
and for how long. We will discuss a number of examples later, but it will be
helpful to generalize the water regimes typical of the wetland types mentioned
above because these patterns and mean depths are reflected in the responses of
plants that influence bird use (Fig. .). These are hypothetical estimates of the
average ranges of water fluctuation in various wetland types in a season. At any
one time, water levels could be at one extreme or the other, but it is the average
depth and seasonal timing of this water that dictates the characteristic vegetation of the site. Therefore, lakes are lakes because they tend to be wet most of
the time and not conducive to the establishment of emergent vegetation.
Generally, lakes tend to be more stable than constructed reservoirs but both
tend to be deeper than m and to lack emergent vegetation (Cowardin et al.
). Floodplain wetlands are characterized by extreme water-level
fluctuations, resulting from seasonal river flows. Newly created bars and
shallow shorelines are frequented by annual plants, animals with short lifecycles, and more adaptable birds geared to feeding under those shallow conditions. Shallow but semipermanently flooded wetland basins are rich in more
persistent and perennial emergents and even in aquatic plants, and, as a result,
with birds favoring deeper water. Long-lived woody plants more often develop
in backwater basins that capture and hold water for part of the year. These
water dynamics and substrate disturbance patterns should not be construed as
reducing productivity of the system, which often is greatest in seasonally
dynamic wetlands. Tidal systems could be illustrated in any magnitude of
extremes, but in (Fig. .) a median point is shown with a range from tidal
highs to lows. This range differs by latitude and geomorphic settings from the
extremes observed in the Bay of Fundy and Tierra del Fuego (over feet,
m) to a modest inches (cm) in the Gulf of Mexico. These ranges
influence vegetation development and bird use because they require adaptations to extremes and daily and seasonal timing. At any site, the celestial
influences are predictable but wind tides are an especially important influence
on water depths in lesser tide ranges.
To understand how these general wetland characteristics influence avian
habitat selection, we will next examine life-history characteristics and adaptations of the birds using these diverse and often variable habitats.
References
Bergman, R. D., Howard, R. L., Abraham, K. F., & Weller, M. W. (). Waterbirds and
their wetland resources in relation to oil development at Storkerson Point, Alaska.
Resource Publication . Washington, DC: US Fish & Wildlife Service.
Brinson, M. M. (). A hydrogeomorphic classification of wetlands. Wetlands
Research Program Technical Report WRP-DE-. Vicksburg, MS: US Army
Corps of Engineers.
Chabreck, R. H. (). Coastal marshes: ecology and wildlife management.
Minneapolis, MN: University of Minnesota Press.
Cowardin, L. M., Carter, V., Golet, F. C., & LaRoe, E. T. (). Classification of
wetlands and deepwater habitats of the United States. FWS/OBS /.
Washington, DC: US Fish & Wildlife Service, Office of Biological Services.
Damman, A. W. H. & French, T. W. (). The ecology of peat bogs of the glaciated
Northeastern United States: a community profile. Biological Report (.).
Washington, DC: US Fish & Wildlife Service.
Duebbert, H. E. (). The ecology of Malheur Lake. Refuge Leaflet No. .
Washington, DC: US Fish & Wildlife Service.
Federal Interagency Committee for Wetland Delineation (). Federal manual
for identifying jurisdictional wetlands. Cooperative Technical Publication.
Washington DC: US Army Corps of Engineers, US Environmental Protection
Agency, US Fish & Wildlife Service, & US Soil Conservation Service.
Glaser, P. H. (). The ecology of patterned boreal peatlands of northern Minnesota:
a community profile. Biological Report (.). Washington, DC: US Fish &
Wildlife Service.
Gosselink, J. G. (). The ecology of delta marshes of coastal Louisiana: a community
profile. FWS/OBS-/. Washington, DC: US Fish & Wildlife Service.
Kadlec, J. A. (). Rising Great Salt Lake inundates marshes. National Wetlands
Newsletter , .
Kauppinen, J. and Vaisanen A. (). Ordination and classification of waterfowl
communities in south boreal lakes. Finish Game Research , .
Kushlan, J. A. (). Wading bird predation in a seasonally fluctuating pond. Auk
,.
Maxwell, G. (). The people of the reeds. New York: Pyramid Books.
National Wetland Working Group (). Wetlands of Canada. Montreal, Quebec:
Polyscience.
Odum, W. E., McIvor, C. C., & Smith III, T. J. (). The ecology of the mangroves of
South Florida: a community profile. FWS/OBS- /. Washington, DC: US Fish
& Wildlife Service.
Odum, W. E., Smith III, T. J., Hoover, J. K., & McIvor, C. C. (). The ecology of tidal
freshwater marshes of the United States east coast: a community profile. FWS/OBS/. Washington, DC: US Fish & Wildlife Service.
Prince, H. H. & DItri, F. M. (). Coastal wetlands. Chelsea, MI: Lewis.
Weller M. W. (). Freshwater marshes: ecology and wildlife: management. rd edn.
Minneapolis, MN: University of Minnesota Press.
Wharton, C. H., Kichens, W. M., Pendleton, E. C., & Sipe, T. W. (). The ecology of
bottomland hardwood swamps of the Southeast: a community profile. FWS/OBS/. Washington, DC: US Fish & Wildlife Service, Biological Services Program.
Zimmerman, J. L. (). Cheyenne Bottoms, wetland in jeopardy. Lawrence, KS:
University Press of Kansas.
Further reading
Cowardin, L. M. & Johnson, D. H. (). A preliminary classification of wetland plant
communities in north-central Minnesota. Special Scientific Report-Wildlife .
Washington, DC: US Fish & Wildlife Service.
Fredrickson, L. H. & Taylor, T. S. (). Management of seasonally flooded impound-
Further reading
3
Major groups of birds that use wetlands
Many books have been written about the taxonomic groups of birds that frequent shallow water or the waters edge, and readers who do not know these
birds should use a field guide in conjunction with this text. However, to help
the nonspecialist, I have tried to list the major groups worldwide that are obligate waterbirds and I have included other orders less dedicated to water but with
examples of wetland representatives (Table .). To appreciate this diversity, it
may be helpful to provide a framework for bird distribution at various scales.
Table . Major taxonomic orders and families or other lower taxa of birds that regularly use wetlands, with general
breeding distribution. Examples are given where only a few species of the group are obligate or regular usersa
No. speciesb
Gaviiformes
Podicipediformesc
Grebes (Podicipedidae)
Worldwide
Pelecaniformesc
Frigatebirds (Fregatidae)
Tropical oceanic
Cormorants (Phalacrocoracidae)
Cormorants (Phalacrocoracinae)
Shags (Leucocorboninae)
Circumtropical
Pelicans (Pelecanidae)
Anseriformesc
Screamers (Anhimidae)
South America
Australia
Worldwide
Whistling Ducks
White-backed Duck
Africa
Geese
Swans
Freckled Duck
Australia
Spur-winged Goose
Africa
Steamer-ducks
Worldwide
Worldwide
Seaducks
Stifftail Ducks
Phoenicopteriformesc
Flamingos (Phoenicopteridae)
Circumtropical
Ciconiiformesc
Herons, egrets, and bitterns (Ardeidae)
Worldwide
Africa
Africa
Table . (cont.)
No. speciesb
Storks (Ciconiidae)
Falconiformes
Kites, eagles, harriers, hawks (Accipitridae)
Worldwide
Ospreyc
Harriers
Worldwide
Kites, e.g., Snail, Slender-billed, Brahminy, Swallow-tailed Mostly tropical and warm temperate
Hawks, e.g., Red-shouldered, Black-collared,
Americas (only)
Common Black-Hawk
Eagles, e.g., Bald Eagle, Stellers Sea-eagle,
Worldwide
Chimango Caracara
South America
Gruiformesc
Rails, crakes, gallinules and coots (Rallidae)
Worldwide
Sunbittern (Eurypygidae)
Cranes (Gruidae)
Limpkin (Aramidae)
Charadriiformesc
Jacanas (Jacanidae)
Painted-snipe (Rostratulidae)
Worldwide
Northern hemisphere
Beach Stone-Curlew
Oystercatchers (Haematopodidae)
Worldwide
Worldwide
Crab-plover (Dromadidae)
Europe, Asia
Worldwide
Lapwings (Vanellinae)
Plovers (Charadrinae)
Skimmers (Rynchopidae)
Worldwide
Table . (cont.)
No. speciesb
Gulls (Larinae)
Worldwide
Terns (Sterninae)
Worldwide
Cucuculiformes
Mangrove and Swamp (or Pheasant) Coucal
Hoatzin
South America
Strigiformes
Barn owl (Tytonidae)
Worldwide
Worldwide
Fishing-Owls
Barred Owl
North America
Short-eared Owl
Coraciiformes
Kingfishersc (Alcedinidae)
Piciformes
Ivory-billed Woodpecker (presumed extinct)
Australia
Mangrove Warbler
Crows (Corvidae): Fish Crow, Northwestern Crow
Vireos (Vireonidae): White-eyed, Mangrove
Dipperc (Cinclidae):
Worldwide
Africa
or Flycatcher
Starlings (Sturnidae): Slender-billed Starling
Babblers (Muscicapidae): Marsh, Chestnut-capped
Wrens (Troglodytidae): Sedge, Marsh
North America
Europe
Table . (cont.)
No. speciesb
Worldwide
Eurasia
Cape Wagtail
Africa (only)
Africa (only)
Papyrus Gonelek
Old World Weavers (Plocidae), e.g., Red Bishop, Fan-tailed
Africa (only)
Africa (only)
Sharp-tailed, Le Contes
New World Warblers (Parulidae), e.g., Prothonotary,
Americas (only)
Gill () was used as a basis for taxonomic ordering except for loons.
Number of species varies because of various interpretations in group treatments (see Appendix for scientific
names).
adaptation to wet areas and most nest in association with freshwater. The
various species of rail demonstrate a range from swimmers and divers (coots)
to shoreline or upland-edge nesters (King Rail), but all frequent wetlands
regularly and are regarded as obligate users of these resources. Many other
orders and families have few obligate wetland bird species, most often reflected
in nestsites or food specializations. In the Americas, many blackbirds (family
Icteridae) build nests in or near wetlands and rear their young there. Some
species are obligate (Yellow-headed Blackbird) and others range from the very
wet areas to the dry upland (Red-winged Blackbird) (facultative); one
typically nests in the uplands near a wetland but feeds its young mainly on
aquatic insects (Brewers Blackbird). Other blackbirds are terrestrial or upland
but may nest near water or wherever food and cover are suitable (opportunistic). In Eurasia, Old-World warblers seem more likely to use wetland vegetation (there are no Icterids or marsh-nesting wrens there) than do New World
warblers. However, several New World warblers favor wet bottomland woods
where they nest in tree holes or in cane and other grasses of the understory or
small gaps in the forest.
Shorebirds show similar segregation by species, although it perhaps is less
conspicuous in migration and wintering areas where most of us see them.
Their habitat focus tends to be on mudflats and other shoreline vegetation,
with species use of a microhabitat influenced by the amount of water and
vegetation as well as by food distribution. Some shorebird species of the drier
areas (Killdeer and Grey or Black-bellied Plover) are adaptable to drier sites as
well but usually occur near water; yet we always associate the birds with water
and consider them at least facultative.
Many species use woody vegetation of the shallows or waters edge, like
shrubby willows and alders or trees like mangroves; often these same species
use plants of similar lifeform in drier areas. Such habitat use by many birds is
more opportunistic; almost all species of swallows use wetlands as feeding and
roosting areas, and some (Tree Swallows) nest in holes in snags over water.
Kingbirds and some woodpeckers may also nest in similar sites, but it is more
an opportunity than a necessity.
Diurnal predators like harriers and Short-eared Owls are even less clearly
tied to water; they lack specific adaptations to water but frequent wet areas
such as marshes, wet prairie, or flooded meadows, perhaps because of food
availability, and they commonly nest there. However, they are not restricted to
wetlands and may nest in upland prairie. Still more extreme opportunism is
shown by the Red-billed Oxpecker, which picks leeches off a hippopotamus in
water but also feeds on ticks on land-based ungulates and nests in upland tree
holes and rock crevices.
Some upland birds use wetlands only when they are not wet, and are not
considered wetland birds at all: a meadowlark, dove, or Ring-necked Pheasant
may nest in a dry wetland and feed there. Pheasants also use wetland cover in
winter when the water is frozen, mainly because of the thermal cover the plants
or trapped snow provide, but the pheasant has no adaptations for this specific
plant community.
ornithology texts (Gill ) and most field guides. The number of species in a
taxon may vary significantly with the authority and timing of such taxonomic
reviews, some of which are listed among the references. Common English
names and scientific names are given in Appendix . Later, these birds will be
regrouped into community-oriented categories such as resource guilds,
wetland types, or microhabitats.
3.2.1 divers
Divers (or loons as they are known in North America) of the order
Gaviiformes are large, northern forest (Common Loon) or Arctic tundra
species (four other species) well adapted to feed on fish, amphibians, and large
invertebrates by long, spear-like bills and webbed feet placed far back on the
body. They often feed in deep water, can dive to depths of more than m, and
can change body density by compression of body and feathers so they can sink
slowly.As a result of diving adaptations, they walk poorly on land and rarely do
so. Feeding and nesting tend to be solitary, although Red-throated Loons are
social. Nest sites are selected on fairly solid substrates immediately adjacent to
water, sometimes enhanced with vegetation from the nest area along shores of
islands in freshwater lakes. Red-throated Loons often build nests on small wetlands or pools on islands and fly to adjacent larger bodies of water for food,
which is carried in their bill to the young at the nest area. Clutches of eggs tend
to be small, and the young are precocial and down-covered and able to swim
shortly after their down is dry. Loons are day-time migrants and move long
distances from northern breeding areas to warmer wintering areas on large
freshwater lakes, marine bays, or coastal waters. After breeding and probably
after fall migration, a complete wing molt occurs on lakes selected for
sufficient size and food resources to ensure survival during the flightless
period.
3.2.2 grebes
Grebes (order Podicipediformes), like the loons, are skilled diving specialists
but are a more diverse group of about species worldwide in distribution.
However, they are most abundant in cold-temperate climates occurring
through either latitude or altitude. All have lobed toes for swimming underwater with great maneuverability, and their terminally placed legs make it necessary for them to crawl on their bellies on land. Like loons, they can sink
underwater without active jump-diving, as is common in ducks and coots.
Most species of grebe favor more shallow and often well-vegetated wetlands
rather than great expanses of open and deep water used by loons. Their bills
vary from short and sharp to long and spear-like, dependent upon their
aquatic prey. They nest mainly in freshwater wetlands, sometimes in sizable
colonies, and may build nests of submergent vegetation in open water or of
decomposing to fairly fresh emergent vegetation in quite dense marsh vegetation. The precocial young tend to spend a few days on or near the nest, where
they are fed by the parents; some are fed until nearly grown and feeding independently. Some species may be double-brooded (i.e., attempt to rear two
clutches per breeding season) where temperature and water conditions
permit. Adults and young eat fish, amphibians, and large invertebrates. They
migrate long distances at night to winter in large freshwater lakes or marine
bays. Three high-altitude Central/South America species (or subspecies)
endemic to large lakes are essentially flightless.
3.2.3 pelicans
The Order Pelecaniformes contains a diverse group of large-bodied waterbirds
most commonly associated with nearshore marine systems. However, many
are freshwater species and a few are found almost entirely there. Their feet are
uniquely adapted for swimming, with webbing between all four toes (totipalmate), rather than between three (palmate), which is common to divers and
ducks.
The seven or eight species of pelicans occur on all major continents and are
the best known of the order Pelecaniformes because of their large bill and
gular pouch. Most species capture fish, amphibians, or other aquatic prey in
water while surface swimming or tipping up. The relatively smaller Brown
Pelican is the only one that regularly fishes by aerial plunge-diving with closed
wings from m or more above the water, like the related oceanic boobies (six
species) and gannets (three species). Brown Pelicans frequent tropical to warm
temperate nearshore marine waters whereas the other species nest in inland
fresh or saline lakes in the Americas, Africa, and Australia, but they may winter
along seacoasts in highly saline waters. Most nest socially on the ground on
islands, although Brown Pelicans may nest in shrubs or trees such as mangroves. The young are altricial and naked and are fed in or near the nest until
feathered and able to swim. Adults may travel up to km to gather food
(Findholt and Anderson ).
3.2.4 shags and cormorants
Shags (about species) and cormorants ( species) are worldwide members
of the Pelecaniformes; the former subfamily is especially well represented in
marine and fresh cold waters at high latitudes whereas the cormorants are
more tropical and freshwater (Johnsgard ). One tropical cormorant of the
Galapagos Islands is flightless. All species feed mainly on fish, are skilled
aquatic divers, and often feed in mixed flocks with pelicans and gulls. Most
species are social nesters, often mixed with herons, egrets, ibises, and anhingas
in colonies in tree, shrub, or marshy nest sites. Coastal birds nest on cliffs
socially or in mixed flocks of seabirds. The young are altricial, naked at hatch-
ing, and remain in the nest when small. Most are strongly migratory, often
moving in goose-like wedges even in local moves from overnight roosts to
feeding areas.
3.2.5 anhinga and/or darters
The four species of anhinga and/or darters are tropical or subtropical birds of
dominantly freshwater wetlands (Portnoy ); they are shag-like in shape
but with longer necks, straight bills, and long, fan-shaped tails. They are sometimes called snakebirds because they are skilled divers and can sink in water
like grebes, with only the head and neck exposed. Solitary feeders, they take
mostly fish by impaling with one or both mandibles rather than by grabbing
(Johnsgard ); amphibians and large aquatic invertebrates also are eaten.
They frequent shallow and often vegetated wetlands but seem to favor shrub or
open-forest areas where they can readily perch. They have difficulty in getting
airborne and may climb a perch first, but they then fly strongly with a distinctive flap-and-glidetechnique. Like pelicans and frigatebirds, they readily
soar. Nests typically are over water in shrubs, small trees, or snags and are
found with those of other colonial waterbirds such as herons, ibises, and
storks. The altricial young are fed by both parents on regurgitated food. After
nesting, adults move to larger water bodies (Meanley ), presumably where
they undergo a complete and simultaneous wing molt (the only pelecaniform
to become flightless during molt)(Palmer ). Birds that nest in warm-temperate areas seem to migrate to tropical wintering areas (Palmer ).
3.2.6 frigate birds
The five species of frigatebirds are mainly tropical oceanic seabirds that often
nest on remote islands but also may nest or roost in shrubs and small mangrove trees along coastlines and, therefore may be considered birds of coastal
wetlands. They are the ultimate aerialists, gracefully surface-skimming for
flying fish or snatching squid, rarely if ever landing on water and may even
drink from freshwater pools on-the-wing. They commonly pirate food from
boobies, gulls, and other fish or squid feeders. They nest colonially in bushes or
low mangroves, commonly in areas attracting other species of nesting
seabirds.
3.2.7 waterfowl: ducks, geese, and swans
The waterfowl (wildfowl in Europe) or ducks, geese, and swans of the order
Anseriformes are a large and cosmopolitan group widely distributed in tropical as well and extreme north and south latitudes (Delacour ,
Johnsgard ). They are more conspicuous than many abundant birds
because of size, their day as well as night migration, and their large and
conspicuous flocks. Species that live at high latitudes are strongly migratory
and are, therefore, excellent pioneers at exploring and exploiting new habitats.
Several endemic species of subantarctic and cold-temperate islands are
flightless (Weller ).
Nest sites vary from cliffs and tree holes to highly aquatic but well-vegetated situations. The young are precocial, downy, and leave the nest to feed
with other brood members as soon as they are dry. Young birds are strongly
insectivorous except in the true herbivores like geese and swans. Parental care
(brooding and predator defense) ranges from long term (geese and swans) to
none (the parasitic Black-headed Duck). After breeding, males in particular
but sometimes females as well engage in molt-migration, flying a few to hundreds of miles to suitable feeding areas with protective water depths where they
lose their flight feathers prior to autumn migration (Hochbaum ). A few
species molt after post-breeding migration.
Waterfowl, and especially the ducks, are best grouped by larger taxa for discussion because such groupings tend to reflect common names of the groups
and describe their behavior and resource use. The term tribe, a taxonomic category between subfamily and genus, will be used here (Table .) because it has
been popular and functional and is still a basis for the latest classification
although subfamily status has been given to some tribes (Livezey ().
Screamers of South America (three species) are large, goose-like birds with
chicken-like bills and semipalmated feet; they graze on meadows, wet grasslands, and shallow marshes. The Southern Screamer nests over water in robust
emergent vegetation, and the precocial young feed on aquatic invertebrates
and plants until able to feed safely in drier areas. They are taxonomically distinct from all other members of the order and are usually placed in a separate
suborder (Anhimae) and family (Anhimidae).
Although a unique-looking bird, the single species of Magpie or Pied
Goose is more closely related to the suborder of typical waterfowl but is placed
in a separate family (Anseranatidae). It occurs in tropical Australia where it
feeds on plant tubers and other vegetation by wading or upending; it is highly
social in feeding as well as nesting and is unique among the suborder Anseres
in having a serial wing molt. Frith () considered this an adaptation to the
drying of the marshy swamps in which it nests.
Whistling ducks number about nine species; they are closely related to
swans and are of tropical to warm-temperate distribution around the world in
diverse habitats. Some field-feed in farm crops and nest in tree holes in semidesert (Black-bellied) and others dive for invertebrates in densely vegetated
subtropical pools (Fulvous); this results in seasonal and regional variation in
food and in foraging tactics. The White-backed Duck of Africa often is placed
with or near the whistling ducks as a separate tribe or subtribe, but it is a highly
aquatic diving specialist. The seven or so swan species (sometimes separated as
the tribe Cygnini) are well known as usually white aquatic birds that feed on
submergent vegetation as first choice but will also graze in meadows. The Black
Swan of Australia, and the Black-necked Swan and Coscoroba Swan (with
black wing tips) of temperate South America, contrast with the all-white swans
of the northern hemisphere. Marine and estuarine vegetation is used by swans
in both hemispheres.
Geese (tribe Anserini) number about species throughout the northern
hemisphere and are specialized as grazers in meadows and uplands but may
feed in wet areas on tuberous plants. The more coastal Brant feed on eelgrass or
algae in estuaries during migration and in winter and may use grasses and
sedges during breeding on freshwater lakes. Some wild species have been introduced widely for hunting, and domesticated varieties of Eurasian geese are
important human food resources worldwide.
A complex tribe (or subfamily (Livezey ()) includes shelducks and
sheldgeese (commonly called geese because of body shape and feeding behavior but not taxonomically the same as northern hemisphere geese). These
show a great range of habitat selection and specialization: alpine meadows
(Andean Goose), clinging marine algae (Kelp Goose), and tropical savannah
(Egyptian Goose and South American Orinoco Goose). The shelducks are
more duck-like in body form but also may graze in moist grasslands or feed on
estuarine or freshwater invertebrates as well (Common Shelduck).
The four species of steamer-ducks found at the tip of South America often
are classified with shelducks but are massive diving specialists forming an
ecological equivalent of the northern hemisphere eiders. The three flightless
species are found only on marine coasts, whereas the flying species frequents
large inland lakes.
Dabbling ducks (Anatini) number between and species and are considered the most ubiquitous and adaptable of all waterfowl and among the
most widespread of all waterbirds. Species complexes such as mallard- and
pintail-like ducks span both hemispheres and are successful in a great variety
of habitats from cold latitudinal and alpine tundra to the tropics. They are
flexible in foods, pioneers in new habitats of even temporary water, and are
persistent in rearing young under tenuous conditions. Foods of these omnivores include seeds and other plant reproductive parts during nonbreeding
seasons and invertebrates during pre-breeding, growth, and molting periods.
Nests commonly are on the ground near water, but clutches are large and renesting is common.
Bay ducks or pochards (Aythyini) include species and are nearly as widespread as dabblers but require more permanent waters, where they feed
seasonally on invertebrates and submergent plant material. Breeding areas
usually are on freshwater, but wintering birds may use either saline or freshwater. Some members of this group nest over water in emergent vegetation,
but ground nests near water are common in other species.
Sea ducks (Mergini) such as eiders and goldeneyes number about
species and are found mainly in the northern hemisphere. They are well
designed for cold water regardless of salinity; most eiders nest near tundra
ponds and some winter and feed among Arctic ice floes. Other sea ducks like
goldeneyes and mergansers nest in tree holes in coniferous forests and winter
in large lakes and river systems, often in warmer climates. Harlequin Ducks,
goldeneyes, and mergansers are well adapted to fast-moving streams where
they feed on invertebrates (Harlequin) or fish (goldeneyes and mergansers).
Long-tailed Ducks rival loons in their diving ability.
The Stifftail ducks (Oxyurini) include nine species. They are skilled divers
of worldwide distribution and frequent shallow but often open water
(Johnsgard and Carbonell ). Their long and often upright tail is used in
social displays but undoubtedly is also a useful diving tool. They feed mainly
on benthic invertebrates for much of the year, although submergent foliage
and seeds are used during winter in some areas. The young are among the most
precocious of all birds; the young of the parasitic Black-headed Duck leave the
hosts nest a day or two after hatching and are thereafter independent (Weller
).
3.2.8 flamingos
Flamingos (order Phoenicopteriformes) are widely known because of their
typically vivid pink coloration, unique bill shape, feeding behavior, unusual
body form yet graceful flight, and their highly social behavior. The Lesser
Flamingo specializes on algae (Spirulina spp.)(Zimmerman, Turner and
Pearson ), the Greater Flamingo seems to favor microcrustaceans and
mollusks, but the American Flamingo also uses some bulbs and seeds of emergent plants (Arengo and Baldassare ). These are obtained from bottom
oozes by straining through their sieve-like bill with the head upside-down in
the water or mud; the bill is flattened laterally rather than dorso-ventrally as in
spoonbills. Foot-stirring is common to enhance food intake, as is true in a
slightly different form in fish-feeding egrets. Flamingos build unique cone-like
nests of mud often quite high to avoid flooding on mudflats in highly saline
and sometimes caustic lakes (pH of up to .). African populations have
demonstrated highly nomadic patterns in search of suitable breeding areas
after traditional areas have been flooded or totally dried (Crowther and Finlay
). Others remain in certain areas and often face nest failure or juvenile
mortality because of drying habitat and lack of freshwater. They rear few
young per nesting effort but seem to live long lives ( or more years). The
young are precocial and down-covered but tend to remain in the nest for a
week or more and are fed with a regurgitant until fledged. Their bill shape
changes rapidly with growth, and they are able to strain water for food in an
adult-like manner in about a month. Except for island populations, flocks may
engage in both migration to wintering areas and more local molt-migration
prior to becoming flightless (as in ducks and geese and a few other families or
orders).
tree snags and even chimneys as nest sites, and White Storks and Marabou
Storks supplement their food from refuse dumps (Blanco , Zimmerman et
al. ). The unique African Hamerkop feeds in both water and on land but
builds a stick nest, which it enters through an opening in the side.
3.2.10 hawks and eagles
Hawks and eagles (order Falconiformes) are mostly facultative or opportunistic wetland users, but several that specialize on aquatic foods are almost
restricted to wetlands: Snail Kites of North and South America feed almost
exclusively on Apple Snails and nest over water in small shrubs. Ospreys,
nearly worldwide in coastal areas and large rivers and lakes, feed mainly on
fish in shallow waters but build nests in snags and artificial structures near
water. The African Fish Eagle, Bald Eagle of North America, and several seaeagles of Eurasia are adept at taking fish and not afraid of getting wet;
although they are somewhat flexible in habitat, they seem to favor seacoasts,
large lakes, or rivers, where they often feed on dead, stunned, or vulnerable live
fish. Like Ospreys, eagles often build large nests, which may be used year-afteryear; the young of all members of the order are altricial. A few hawks favor
open marshy wetland edges and nest on the ground in fairly wet places (harriers) but also hunt in and occasionally nest in upland grasses and forbs. Some
forest hawks favor wet shrub and bottomland wetlands (South American
Black-collared Hawk, North American Swallow-tailed Kites and Red-shouldered Hawks) or mangroves but are not restricted to these habitats (Howell
and Chapman ). Similar responses to water are noted among other species
of kites worldwide.
3.2.11 rails, crakes, coots, and cranes
The rails and crakes, coots, and cranes of the large, diverse, and worldwide
order Gruiformes range from the more conspicuous cranes and coots to the
habitat-shielded rails of emergent vegetation. Many rails and crakes frequent
the shoreward zone in dense vegetation, where they are more commonly heard
than observed. Moorhens, gallinules, and coots use deeper and even open
water. Foods are highly variable, with seasonal shifts from animal foods (especially invertebrates) to seeds, tubers, and foliage in fall and winter. Coots regularly use open water when feeding on submergent vegetation, and they dive for
both vegetation and invertebrates especially during pre-nesting egg formation and when feeding young. They are equipped with lobate toes effective for
diving as well as for walking on mudflats. Coots and probably other members
of this family lose the power of flight during their complete and simultaneous
wing molt. Rails fly little except when pressed but are capable long-distance
migrants and inhabit some of the most remote islands of the world. Several
island species are flightless or nearly so, as is the famous Takahe or Notornis of
New Zealand, which occupies tussock swamps and steep but soggy slopes
above m elevation with little standing water.
Cranes switch from nesting areas in marshes, where they eat amphibians
and other small vertebrates, to open grasslands (tubers) or fields in winter
(grains). Some cranes (Whooping) use estuarine areas and feed on crabs and
other marine invertebrates but also take freshwater crayfish and acorns opportunistically. Most species are found in the northern hemisphere, where they
nest in bogs and marshes after long daytime migrations in goose-like flocks,
vocalizing as they travel. Sandhill Cranes rest during migration on river islands
and sheetwater, where night-time roosts are adjacent to fields in which highenergy foods are readily available. The more terrestrial species seem to have
prospered as a result of current grain crops, but southern forms have lost
nesting and roosting sites to intensive agriculture and water developments.
The Limpkin of Florida and Central and South America has some ibis-like
features such as a decurved bill and general stature but is a close relative of the
cranes and rails. It is larger than ibises, is typically well-hidden in marsh
vegetation, and its screaming call alerts one to its unseen presence. It feeds
mainly on Apple Snails and other large invertebrates. Nests are in marshy areas
or low shrubs, and the young are precocial and downy, leaving the nest soon
after hatching as in other members of the order. Although commonly solitary,
it may occur in flocks.
Although the three Finfoots seem to be related to the rails, they have grebelike bills and feet. One species is often called the Sungrebe. Both the species
found in tropical America and Africa favor slow-moving streams (Alvarez del
Toro , Zimmerman et al. ), swim well, and are more fish-oriented than
others in this order; they also eat crustaceans and large insects. Most nest sites
are elevated over water or at the waters edge; the young are nearly precocial
and are fed in the nest before ranging afield. Adults apparently walk well on
land and wade as well as dive in shallow water. They take flight with difficulty,
favor well-vegetated areas, may roost in trees at night, and are not very
conspicuous. The single species of Sunbittern of Central and South America is
a stream-edge wader, nesting in solitary pairs along streams, and feeding on
aquatic resources. The young are precocial, as are others of this order.
3.2.12 charadriiformes
The order Charadriiformes is a large, diverse, complex, and important group
involving about mostly obligate taxa (families, subfamilies or tribes) of
wetland birds best identified by common group names. Best known are the
gulls and terns, sandpipers and associates like bog-country snipe, forestdwelling woodcock, phalaropes, stilts and avocets, and plovers. The auks and
allies also are members of this order but are truly seabirds and nest colonially
on rocky islands and cliffs. One member of the group, the Marbled Murrelet,
nests on the horizontal branches of old-growth conifers growing near the
northwest coast of North America and eastern Asia. Most members of the
order are strong fliers and interhemisphere long-distance migrants; the spectacular migration and complex routes of Red Knots and Golden and Grey
Plovers between Arctic nesting areas and southern continents are renowned
(Harrington ).
Plovers, especially the three Golden Plovers, share with sandpipers the
strategy of long-distance migrations; which allows them to tap global
resources. They are more terrestrial than the sandpipers, feeding and nesting
in drier and shorter vegetation but commonly use wetland shorelines. The
common North American Killdeer and the threatened Piping Plover are well
known: the former tolerating human development to the point of nesting on
gravel rooftops; the latter frequenting temporary habitats often eliminated by
development and reservoirs designed for water management. The young are
precocial as is common to species of dynamic habitats. The Glareolidae of
Eurasia includes the coursers and swallow-plovers or pratincoles. They are
plover-like and nest on the ground. The Collared Pratincole nests on dry
meadows or cropland (Calvo and Furness ) and hawks for aquatic insects
over wet meadows and marshes (Calvo ). The coursers are more terrestrial, but one member of that group, the Egyptian Plover, nests and feeds
along sandbars of large rivers. The Crab-plover, the only member of the
family Dromadidae, is strictly a coastal species; its name describes its food
(Fasiola, Canora and Biddar ) but it is unique in its petrel-like behavior of
nesting in burrows in sand dunes, often in huge colonies (Campbell and Lack
).
The sandpiper family is large and diverse and often separated into tribes.
Sandpipers are birds of shallow, open or grassy sheetwater, mudflat, or sandy
shore; they feed on live invertebrates, spawn, or small fish or amphibians most
of the year. Many are habitat-specific, with preferences for substrate, water
depth, wave height, wind, etc., so that adaptations of leg length, neck length,
bill length, and shape differ by species and sometimes by sex. They glean, peck,
pick, drill, bore, probe, and sweep, and they demonstrate efficient adaptations
in behavior as well as in structure. Their nesting habitats often differ markedly
from their winter or migration feeding areas, and one sandpiper, the Solitary
Sandpiper, nests in old songbird nests in stunted trees in the northern forest
edge that is interspersed with grassy feeding pools (Kaufman ). The young
are precocial. Curlews are larger species with long, down-curved bills and
longer legs; some feed on mudflats and other species favor uplands. They may
nest in wet-meadows or tall grasses and winter in fresh or saline coastal wetlands where at times they feed like Willets or avocets while wading. Woodcock
specialize in moist forested or shrubby bog wetlands, where they probe for
earthworms and other burrowing invertebrates of the northern hemisphere.
The oystercatchers number six or so species of truly coastal shorebirds that
are recognized by their black or black and white plumage and straight, sturdy,
and brightly colored chisel-like bills. They are worldwide in distribution
except for high latitudes; they frequent rocky to sandy shores and feed on
limpets, clams, and other shellfish.
The avocets and stilts are considered either as a separate family or as a tribe
of shorebirds. They are worldwide in distribution and are characterized by
long legs, useful in shallow open water. They favor saline/soda wetlands and
lake shores or estuaries. Stilts have long straight bills and seem more commonly to be running on mudflats than wading, but that tactic depends on
feeding opportunities. Avocets have recurved (i.e., upturned) bills ideally
suited to sweeping the substrate while wading in shallow water up to bellydeep; in fact they sometimes swim. They often nest on sites with little vegetation, and the precocial young are well camouflaged for their habitat.
The phalaropes are among the most aquatic of shorebirds; they may
behave like other walking or wading shorebirds but more commonly they
swim like feathered whirligigs, feeding on tiny crustaceans by bill dipping.
The food seems to be held on the bill by water tension, and the mouth is hardly
opened to take in the food (Rubega and Obst ). They nest by or in freshwater wetlands in temperate or Arctic areas, using relatively short emergent
vegetation like water-tolerant grasses or sedges, near and sometimes over
water and, therefore, have food nearby. This group is characterized by reverse
sexual plumage colors and sexual roles, with males doing most of the incubation and care of young. They are strongly migratory, with the Red (or Grey in
Europe) and Red-necked Phalaropes often wintering at sea. All three species
regularly migrate in autumn to winter in tropical and southern hemisphere
habitats and are very tolerant of saline waters.
Jacanas are the tropical or subtropical water-walkers; they have light
frames, long toes to disperse the body weight, and can gracefully flit across
even submergent vegetation and especially over lily pads and similar floatingleaf vegetation. They feed mainly on snails, insects, and other invertebrates.
Rather small and shallow nests are made of plant material on floating or rooted
vegetation, and eggs are incubated by one of several males associated with the
same laying female. The eggs are incubated by the males, and the precocial
young leave the nest shortly after drying.Young may be chased or killed by new
females entering the mating group (Emlen, Demong and Emlen ).
Many of the common gull-like birds are grouped as one family but make up
at least four subfamilies. Gulls and terns, while closely associated and often
nesting together or on similar sites, represent two subfamilies that reflect
different feeding strategies involving adaptations of flight and feeding apparatus. Terns are predators on live fish or aquatic invertebrates taken on or near
the surface of fresh- or saltwater, and a few capture insects in flight over land.
Most are skilled at hovering, plunge-diving, and devouring their prey aloft.
Black Terns skim marsh water during breeding or dry fields in migration for
insects.
Gulls are more commonly scavengers or kleptoparasites (hijackers),
harassing and taking food from other birds. During the breeding season,
marsh-nesting species like Franklins Gulls in North America and Brownhooded Gulls in Argentina feed dominantly on marsh flies (Chironomidae).
Marsh-nesting terns like the Whiskered Tern may nest on water-lily pads,
whereas the Black Tern typically nests on soggy and rotting plant debris but
may build a nest of fresh plants on a floating structure. Others, especially along
large lakes or seashores, nest on islands or shorelines on sand or rocks with
little nest material. The Forsters Tern does both, varying with region and habitats (marsh, lakeshore, or seashore). The young of gulls and terns typically are
precocial but stay in or near the nest while being fed insects or bits of regurgitated fish, crabs, and squid.
Jaegers and skuas are gull-like but are placed in a separate family or subfamily; they are typically coastal marine birds but they often nest on land near
freshwater areas. They are large and more predators than scavengers, taking
mammals, fish, or young birds or eggs during the breeding season.
Skimmers are placed in a separate family because of their unique adaptations for surface feeding, including an extended lower bill and elegantly
designed wings for surface flight. They feed on fish and large invertebrates by
skimming with the lower bill and grabbing on contact, by day or night.
Although considered tactile feeders, they seemingly are adapted for nocturnal
vison by a larger than usual number of cones in the retina (Rojas, McNeil and
Cabana ). They commonly nest on bare to mat-covered sandbars along the
coast (Burger and Gochfield ) or on large lakes and river systems, where
they can easily take flight. As a result, where locally abundant, they are commonly seen at nests, and some populations even use gravel rooftops for
nesting.
3.2.13 cuckoos
There are few cuckoos that frequent wetlands but several parasitic species do
lay eggs in over water nests of other birds. The Old World Swamp Cuckoo or
Pheasant Coucal and the Mangrove Cuckoo in subtropical America (sometimes considered a race of the Yellow-billed Cuckoo) were named for the
common habitat selection. But one unique species, the South American
Hoatzin, was once taxonomically linked with gallinaceous birds but now is
considered an aberrant cuckoo by most taxonomists. It nests over water in
small trees along slow-moving streams and feeds on coarse vegetation of
wetland trees, absorbing their nutrients through a foregut fermentation
system (Gralal ).Young remain in the nests until able to clamber around in
the branches, aided by vestigial claws on the wings.
3.2.14 owls
Although few people in America and Europe think of Owls as wetland birds, a
few are well adapted for preying on animal food resources of rivers and basin
wetlands. There are three species of fishing-owls of Africa and four fish-owls of
Asia, which specialize on fish much as Ospreys and eagles do, but they often
feed at dusk or night. They lack extensive feathering on the legs and have gripping feet to snatch fish from the surface. They also eat frogs, crabs, and other
large invertebrates. The widespread and mostly high-latitude Short-eared Owl
frequents marshy meadows of grass and sedge as well as grasslands and nests
on the ground, sometimes in habitats as wet as that of harriers. Its food is
mainly rodents and small birds, and its association with water sometimes
makes it a predator of other wetland birds (Holt ). Although the widespread Short-eared Owl does not nest in Africa, a close relative, the African
Marsh Owl, nests in wet vegetation and seems to take even more wetland prey.
A similar habitat is used by a member of the Barn Owl family, the African Grass
Owl, which nests on the ground in wet or damp places. The Barred Owl of the
southern USA favors mature and wet bottomland woods along large rivers,
where it feeds on amphibians, fish, and crayfish in addition to mammals and
birds.
3.2.15 kingfishers
Kingfishers are a worldwide group of about species (Fry, Fry and Harris
) that are far more diverse in habitat choice and feeding habits than generally appreciated. There are three families, of which only two are perch-anddive fish eaters. The largest family of about species comprises mainly forest
hunters like the Australian Laughing Kookaburra and the Shovel-billed
Kingfisher or Kookaburra of New Guinea, which gropes on the forest floor in
search of grubs and worms. The traditional wetland kingfishers ( species in
two families) specialize on fish or large invertebrates of various sizes, with prey
body size often related to bird body size (Remsen ). They use a variety of
wetland types, including saltwater, where perches allow. Many are stream-oriented, and artificial ditches or canals are favored sites, especially when wires are
conveniently situated. Others frequent pools of all sizes, and although they
seem to favor perches of certain heights (Remsen ), they can hover
effectively some distance from shore; for example, the Pied Kingfisher of Africa
has been recorded some km from shore on large lakes (Fry et al. ). Like
many fish-feeders, they capitalize on prey trapped during declining water
levels so will use roadside ditches and tiny pools far from other water. Most
nest in burrows or crevices, rear few young per brood, and vary greatly in
migration patterns.
3.2.16 songbirds
Compared to most of the taxa mentioned above, few of the perching birds or
songbirds (order Passeriformes) are obligate wetland specialists with anatomical, behavioral, and physiological adaptations, but many tap water resources
and nest at the waters edge. Most probably evolved from adjacent upland
species that successfully reared young from nests built in shoreward wetland
vegetation because of the protection from predators afforded by the water, but
also because of the richness of invertebrate foods essential to growing nestlings
and common to wetlands. Most have altricial young in cup, dome, hole, or
crevice nests, and feeding by parents is essential until young are flying.
Several passerines are adapted behaviorally to streams or rivers. In the
mountainous parts of the northern hemisphere and in South America, there
are five dippers that walk, swim, dive, and feed in, and nest along, flowing
mountainous streams. They seem tied to several insect groups like caddisflies
that frequent these areas. A number of South America ovenbirds favor marshy
areas, and the genus Cinclodes uses, streamsides or seashores. The African
Slender-billed Starling is associated with waterfalls, hopping around and
feeding in the spray, and nesting behind the falls even in groups (Zimmerman
et al. ).
Several wrens of North America are well adapted to fresh and saline
marshes, with the Marsh Wren found in no other habitats and the Sedge Wren
using shallow marshes of less-robust, grassy vegetation or wet prairie. This
habitat-use pattern applies also to several New World grackles and smaller
blackbirds, mentioned above. Additionally, each area of the world seems to
have a few marsh, rush, reed, or shoreline specialists among flycatchers, vireos,
finches, and warblers. The last range in North America and include
Prothonotary Warblers nesting in tree-hole cavities in bottomland swamps,
Palm Warblers of northern bogs, Bachmanns (Remsen ), Swainsons, and
Hooded Warblers of southeastern cane breaks, and Yellow Warblers, which
seem always near water but use diverse sizes of shrubs and trees for nesting.
The New World warblers called waterthrushes and the ovenbirds use small
habitat patches along stream courses or lowland woods. A number of warblers
use mangroves either for feeding or wintering, but little is known about their
relative use of these versus adjacent habitats. The closely related Common
Yellowthroats also frequent water areas and may nest in emergent plants like
cattail but most often use upland shrubs and thickets. As noted elsewhere,
there seem to be many Eurasian and African warblers (family Sylviidae) that
specialize in emergent marshes where no Icterids or wrens compete.
Flycatchers are fairly common residents of shrubs and emergents at the
waters edge; the Alder and Willow Flycatchers of North America and the
Swamp Flycatcher in Africa are essentially restricted to such habitats.Although
the Kiskadees of subtropical areas use a wide range of habitats, they generally
frequent water areas where they feed on dragonflies or large invertebrates and
fish.
Few seed-eating sparrows have obvious morphological specialization for
wetlands, but many use them and a few seem limited by habitat patterns and
feeding behavior (see Rising ). Some, like Song Sparrows, seem most
abundant in floodplains, where they use small trees for song perches and brush
for nesting, but they have wide latitude in habitat use. Several sparrows use
damp to wet vegetation of grass and sedge meadows (LeContes and Henslows
Sparrows). Lincolns Sparrow favors northern boggy habitats. Only a few actually nest in wet, marsh-edge emergent herbaceous plants or small trees
(Swamp Sparrow), but more species feed in wet or shallowly flooded emergent
vegetation (Nelsons Sharp-tailed Sparrow) even far from shore (Seaside
Sparrows). Populations of several species of both more upland (Savannah)
and wetland-edge species (Song and Swamp Sparrows) have races or sibling
species that nest in tidal saltmarsh (e.g., Powell ).
Swallows feed over and may nest over water but despite a strong association
with wetland insects lack specializations that restrict them to wetlands. Tree
Swallows and Mangrove Swallows feed mainly over water and wetland vegetation, presumably because of the abundance of food (St Louis, Breebart and
Barlow ), and tend to nest in holes in snags over or near water.
3.2.17 summary
The use of wetlands and their resources is widespread among many diverse
bird taxa of the world, with some using them to the exclusion of other habitats
via specializations that maximize exploitation of the rich resources but that
limit use of alternative habitats. Many groups adapted to other habitats have
few wetland representatives occurring only regionally or locally and are
limited in adaptation to behavioral responses such as selection perhaps as a
product of reproductive success.
Hopefully, this sketchy coverage will introduce readers to the elaborate
adaptations and gradients of wetland use demonstrated by many taxonomically diverse birds of the world and encourage additions to this list.
References
Acosta, M., Mugia, L., Mancina, C., & Ruiz, X. (). Resource partitioning between
Glossy and White Ibises in a rice field system in Southcentral Cuba. Colonial
Waterbirds , .
Alvarez del Toro, M. (). On the biology of the American Finfoot in Southern
Mexico. Living Bird , .
Arengo, F. & Baldassarre, G. A. (). Effects of food density on the behavior and distribution of nonbreeding American Flamingos in Yucatan, Mexico. Condor ,
.
Blanco, G. (1996). Population dynamics and communal roosting of White Storks foraging at a Spanish refuse dump. Colonial Waterbirds .
Burger, J. & Gochfield, M.(). The black skimmer: Social dynamics of a colonial
species. New York, NY: Columbia University Press.
Calvo, B. (). Feeding habitats of breeding Collared Pratincoles (Glareda pratincola) in Southern Spain. Colonial Waterbirds (Spec. Publ. No. ): .
Calvo, B. & Furness, R. W.(). Colony and nest-site selection by Collared
Pratincoles (Glareda pratincola) in southwest Spain. Colonial Waterbirds , .
References
Further reading
Daiber, F. C. (). Animals of the tidal marsh. New York: Van Nostrand Reinholt.
DeSchaunsee, R. M. (). A guide to the birds of South America. Edinburg: Oliver
and Boyd.
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management of Swainsons warbler in Southern Illinois. Wildlife Society Bulletin
, .
Erlich, P. R., Dobkin, D. S., and Wheye, D. (). The birders handbook. New York:
Simon and Schuster.
Erskine, A. J. (). Buffleheads. Monograph Series No. . Ottawa: Canadian Wildlife
Service.
Evans, P. R., Goss-Custard, J. D., and Hale, W. G. (eds.) (). Coastal waders and
wildfowl in winter. Cambridge: Cambridge University Press.
Faaborg, J. (). Habitat selection and territorial behavior of the small grebes of
North Dakota. Wilson Bulletin , .
Falla, R. A., Sibson, R. B., and Turbott, E. G. (). A field guide to the birds of New
Zealand. Boston, MA: Houghton Mifflin.
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Press.
Gibb, J. (). Food, feeding habits and territory of the Rock Pipit. Ibis , .
Glover, F. A. (). Nesting ecology of the Pied-billed Grebe in northwestern Iowa.
Wilson Bulletin , .
Gonzales, J. A. (). Breeding biology of the jaribu in the llanos of Venezuela.
Wilson Bulletin , .
Gorena, R. L. (). Notes on the feeding habits and prey of adult Great Kiskadees.
Bulletin of the Texas Ornithological Society , .
Hamerstrom, F. (). Harrier, hawk of the marshes. Washington, DC: Smithsonian
Press.
Hancock, J. A. (). Storks, ibises, and spoonbills of the world. San Diego, CA:
Academic Press.
Hancock, J. and Kushlan, J. (). The herons handbook. New York: Harper Row.
Harrison, C. (). A field guide to the nests, eggs, and nestlings of North American
birds. London: Collins.
Hori, J. (). The breeding biology of the Shelduck Tadorna tadorna. Ibis ,
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Johnsgard, P. A. (). Plovers, sandpipers, and snipes of the world. Lincoln, NB:
University of Nebraska Press.
Johnsgard, P. A. (). Cranes of the world. Bloomington, IN: Indiana University
Press.
Johnsgard, P. A. (). Diving birds of North America. Lincoln, NE: University of
Nebraska Press.
Johnsgard, P. A. (). Hawks, eagles, and falcons of North America. Washington,
DC: Smithsonian Institution Press.
Kear, J. & Duplaix-Hall, N. (). Flamingos. Berkhamsted, UK: T & AD Poyser.
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by Hooded Warblers in bottomland hardwoods of South Carolina. Wilson
Bulletin , .
Kirsh, E. M. (). Habitat selection and productivity of Least Terns on the lower
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Ortega, B., Hamilton, R.B., and Noble, R.E. (). Bird usage by habitat types in a
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.
Owen, M. (). Wildfowl of Europe. London: Macmillan.
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Further reading
4
Water and other resource influences
Habitat needs for all birds include similar resources: water for drinking and
bathing, food, cover for protection from the elements and potential predators,
and undisturbed space for meeting social and other life functions. But each
group of birds adapts to general features of their typical habitat and exploits its
particular resources. Obviously, the unique and dominant feature for wetland
birds is water, and long-term adaptations include genetically selected
modifications in anatomy, morphology, and physiology (Table .). Shorterterm adjustments involve mostly behavioral changes, but we know little about
these and they are difficult to study and test experimentally. Birds, like many
wetland animals, often are not truly water adapted but retain terrestrial
adaptations that allow them to survive with extremes of water regimes and also
to exploit resources of both habitats along the wetland edge.
Migratory, or at least mobile, birds obviously have greater flexibility in
their search for habitat than do flightless, terrestrial vertebrates. It has been
suggested that birds logically should follow a hierarchy of selection from large
to smaller scale. First, on the largest scale, they either reside in or migrate to
large geographic regions with climatic regimes and general landscape or
vegetation features attractive to them (either via instinctive preferences or
prior experience). Second, a bird could select from various landscape alternatives within this geographic region, such as areas showing certain types, spatial
relationships, or sizes of wetlands. Third, a single wetland or complex that provided suitable habitat might be the focus of the birds search for that season or
function (e.g., breeding, nonbreeding, feeding, roosting), and, finally, selection of microhabitats, foraging sites, nest site, etc. within the chosen wetland or
wetland complex will be made to meet its needs. Both the general region and
the specific area presumably would meet some innate habitat image, possibly
influenced by experience, but some trial-and-error exploration may be
involved especially in cases of resource shortages.
General needs will be considered first here and more detailed examples will
be given throughout the text. I will start with water because wetland birds are
influenced by this constantly, and in ways that few of us may be aware.
Figure .. Relationship
between typical water regularity (here termed hydroperiod)
and hydrologic energy in the
three major wetland types in
the hydrogeomorphic classification method (Brinson
).
food. For example, avocets are among the more aquatic shorebirds, usually
wading in water deep enough that they can sweep their upturned bills from
side to side in search of food. But when the water levels are deep as a result of
flooding or tides, they swim and probe head-under like a duck. When that
occurs, it is not unusual for other shorebirds like Greater Yellowlegs and
Willets to be feeding in deeper-than-normal water too, presumably induced by
food resources.
Although water attracts waterbirds and provides both food and protection,
it has its dangers because it can be a powerful force. It may directly harm or kill,
or stress birds through energy demand, thus influencing the species adaptations and ultimately the species composition of birds using certain wetland
types. Based on general observations, it seems that the hydrologic forces of the
three major types of wetland (Fig. ., Brinson ) typically differ in their
bird species richness. During the breeding season in North America, it is
common to find to obligate species in productive basin wetlands, perhaps
to species in fringe wetlands, but many fewer species in river channels. This
pattern presumably results both from energy demands of feeding in such
systems and from the nature and abundance of suitable resources present,
which in turn is the product of the physical stresses that influence the basic
productivity of each system: current in streams and wave action in larger lakes
and oceans. These stresses affect water regimes, which in turn influence plant
physiology (Riemer , Schulthorpe ) and plant succession processes
(which will be elaborated below).
Except for periodic inflow from sheetwater or stream overflow, basin wetlands lack current and have the least wave energy except as produced by wind.
Obviously, the larger the wetland the greater the wind fetch and potential for
wave energy to build. Although such wetlands have vertical water movement
width, e.g., small and intermittent, narrow and high velocity, torrent and riffle,
pool and riffle, rapid and broad, meandering and slow, etc. Obviously, all rivers
act like torrents during flood events, but torrents and other high-energy riverine systems regularly would be the most energy demanding locality for birds;
therefore, we would expect few bird species (or fish or invertebrates for that
matter) because of the adaptations required. It is not surprising that most of
the birds that use streams feed on caddisflies and similar stream insects, but
they also eat fish eggs. Once such adaptations are acquired, stream specialists
are essentially limited to such habitats, which is less true of birds of basin wetlands. We will discuss in Chapter the implications of human-induced
changes in stream speed and configuration as it might affect dippers in alpine
areas (Tyler and Ormerod ) or ducks and herons in slower stream systems
(Weller ).
Adaptations of many birds to feeding in swift water include a laterally compressed body, powerful legs placed well back on the body, and often long bills
suitable for probing between gravel and rocks, and perhaps auditory adaptations to water noise, which otherwise could induce higher predation. Many of
these birds reduce the impacts of current by diving and feeding underwater,
and the current against the back and long tails of Torrent Ducks, Common
Mergansers, and Harlequin Ducks provide downward and forward thrust
(comparable to putting your flat hand out of the car window to experience this
free energy in an air medium). Obviously, tail length for use in water must be
optimized with use in flight. The only diving passerines of high-energy
streams, the dippers, do not have long tails, but some authors point out that the
entire back functions as a driver when the head is down. In fact, dippers seem
to have very few adaptations for what they do so well! Dippers seemingly use
their wings to swim underwater and then walk or swim according to need.
They also avoid the high-velocity portion of streams by feeding along the
shoreline, and several South American species dive little. I have also seen
Andean Torrent Ducks swim up-current along the edges of a flooded stream,
where current was less but where no human could stand, but even Torrent
Ducks may fly short distances to avoid this effort.
There are other stream specialists among ducks that offer an interesting
comparison: the Blue Duck of New Zealand (Eldridge , Kear ) probably is the nearest to the Torrent Duck in current tolerance, but it tends to avoid
streams with great volume of high-speed flow. The Spectacled Duck of the
lower Andes chooses slower, broader, and more shallow streams and may also
use small basin lakes, as does the African Black Duck (McKinney et al. ,
Siegfried ). The North American Wood Duck is considered a riverine bird,
but it favors backwater areas where it nests in large trees in bottomland woods;
most feed and rear their young in quiet, backwater areas or even ponds
(Bellrose and Holm ). Finally, Canada Geese may graze along large, slowmoving streams or streams with pools, but they tend to nest on islands or adjacent cliffs away from floods and predators.
Ice is a form of water that often is used as a resting area briefly or overnight
and for escape cover, but it creates problems for divers in pools and lakes,
inhibits feeding by dabblers when the bottoms are still frozen (which can last
for weeks or months after surface water melts in northern wetlands), and
limits access to food. Starvation has been documented among coots during
early spring migration when weather changes reduced food access and stressed
the fat reserves of the birds (Fredrickson ). Recently, satellite-tracking of
radio-tagged eiders has shown that they use pack ice areas in the north much as
penguins do in the southern hemisphere, loafing on ice and feeding by diving
in the icy waters between the floes (Petersen, Douglas and Mulcahy ).
Undoubtedly, we have much more to learn about winter life at sea for the many
species of ducks and phalaropes that use freshwater wetlands during the
breeding season.
Tidal action in coastal wetlands is a product of celestial gravitational
influences but also of meterological conditions such as wind and barometric
pressure. These influence water depth and food availability both in saltwater
areas and in estuarine and freshwater tidal marshes, often in unpredictable
and unexpected ways. Tidal action in saltwater areas influences availability of
foods in fringing wetlands such as rock substrates (Weller ), seagrass
meadows (Cornelius ), or mudflats (Withers and Chapman ). As a
result, birds either do not feed during high-water periods or move into alternative habitats such as ponds or truly upland feeding areas for New Zealand
Brown Ducks (Weller ), or flooded agricultural fields for shorebirds
(Rottenborn ). Few of us think about the upslope influence of tides upon
freshwater flowing into the sea, but a discrete plant community is formed
there, and birds in that area are fairly diverse but relatively low in population
density compared with saltwater areas (Odum et al. ). Water levels in
these wetlands are influenced by the coastal tidal regime, the distance to the
sea, and the volume of freshwater inflow. Such levels have a major impact on
nesting waterbirds that have water-level nests, such as rails and ducks, but
also influence passerines that nest higher in tall emergent grasses like
Spartina spp. For example, Seaside Sparrows not only suffer high levels of
nest loss during high spring tides but also have high re-nesting potential,
which results in the synchrony of nesting populations with lower tidal
regimes later in the season, and, in ultimately, higher nest success (Marshall
and Reinert ).
. Salinity
Although most birds seem to have salt-removal glands as well as suitable excretory processes, birds vary in their ability to tolerate high salt intake, ranging
from those that perhaps can but rarely do to those oceanic birds that rarely
drink freshwater or have salt-free foods (albatrosses, petrels, and auks).
Moreover, there is considerable evidence that increased use improves the functioning of salt glands under high salinity (Bradley and Holmes , Cornelius
).
North America White and Brown Pelicans often segregate by use of nesting
islands in inland freshwater or saline lakes versus coastal saltwater. Degree of
salinity may be a factor but prey availability is more important. Predator protection also must be important, however, because the colony of White Pelicans
in the Great Salt Lake in Utah nests where there are few large prey items available and makes long flights to food sources to feed their young. White Pelicans
of African soda lakes must get along with no or limited freshwater, but they and
young flamingos do seek out and use freshwater for drinking and bathing,
which may well be essential for their survival. While many flamingos use only
high-saline waters, they may favor saltier water for some functions and avoid it
for others; patterns of use demonstrate that factors such as food, water depths,
and substrate influence this choice (Espino-Barros and Baldassarre ).
Salinity gradients seem easily recognized by birds as Laysan Ducks often go to
the seashore to bathe rather than using a hypersaline lagoon (Moulton and
Weller ).
Among those groups of birds that commonly nest in or near freshwater
during the breeding season, several eiders use only freshwater or estuary nest
sites, but Common Eiders will use full-strength seawater (about parts per
thousand (p.p.t.)). Although adults ducks regularly drink saltwater, they seem
more able to tolerate this heavy salt load than can juveniles, and therefore,
breeding often is on freshwater rather than salt to a point that it may well
limit the distribution of many species (Schmidt-Nielsen and Kim , Weller
). In alkaline lakes in arid regions, efficient salt glands are important survival mechanisms even though we think of the birds using these lakes as freshwater birds (Cooch ). Many waterfowl species in marine environments
nest at stream outlets where they can get freshwater rather than in other seemingly suitable situations. Wintering birds feeding in hypersaline areas often
make regular flights to freshwater ponds to drink (Adair et al. , Woodin
). Moreover, studies of body condition in Northern Shovelers showed that
birds wintering on saltwater sites were less fat than those of most freshwater
sites (Tietje and Teer ).
Recent studies of White Ibis (Bildstein and others cited therein) have
demonstrated that birds nesting in coastal areas may feed many miles inland to
obtain foods that are less salt-laden than those found along the coast. If young
are fed excessive salt loads, growth is reduced and their condition is generally
less healthy (Figure .). It is clear that salt costs the bird energy in physiological removal of salt or in flight energy to seek freshwater. In cases where the
physiology of the species is not well adapted to high salt intake, the alternative
is selection of food with a reduced salt content. Obviously, the latter may
increase adult vulnerability to predators and other dangers arising from
mobility.
Salinity
. Food
Food is so important to birds on a continuing basis that we will devote the next
chapter to this topic, but for perspective, it is important here to identify patterns by group and to clarify that food needs differ greatly by stage of the life
cycle, making generalizations difficult. Except for those birds specialized as
herbivores (specialized gut structure and complex fermentation mechanisms), most omnivores need higher animal protein during egg development,
growth of the young, and body molt (Eldridge and Krapu , Heitmeyer
; Krapu ) They need high-energy carbohydrates during the physical
stress of migration and during thermoregulation at low winter temperatures.
As a result, their diets differ seasonally and by function, and birds move, seek,
and vary food choices to meet these needs. Food for young often differs from
that of adults unless the adults are in molt or body-growth phases. Young
omnivores gradually shift from animal protein in early growth to more seeds
and then foliage as they mature. Carnivores or piscivores show shifts more in
size or species of prey, often dictated by selection made by the parents during
the feeding phase.
Getting food of the correct type and at the right time is one issue, but some
birds are not effective until they have the proper feeding conditions. Hence, a
feeding perch is preferred or needed by most kingfishers or flycatchers.
Aerialists like swallows or plunge divers like Ospreys or Brown Pelicans have
greater flexibility, but even these require areas large enough to permit flight
maneuvers, and water of sufficient depth to reduce potential injury. For most
species, there are habitat-related issues such as water depth, water clarity, and
vulnerability of the prey. Lack of disturbance by potential predators (including
people) also may be important.
. Resting
All birds need undisturbed places for rest both during the day and at night to
avoid predation or stress. If one observes at a site where various waterbirds fly
in during the day, some land in water and start to drink, some start to feed, and
some land on a mudflat or snag, often disturbing other individuals or species
to find a place to rest. Some species have very specific requirements for rest
sites, presumably based on genetic makeup and social structure. Many swimmers like ducks simply float on open water, but losses could occur through predation by large fish or alligators, and, therefore, birds are always testing the
waters. Many rest along bare shorelines or mudflats, preferably on islands
where they are protected from ground predators at night, and the benefits of
social tolerance produce alertness for predators. Vulnerability probably is
greater at nights, but vocalizations of geese, cranes, and ducks suggest that
sleep is not as sound as in humans. Some species like coots apparently sit on
mats of submergent vegetation where alligators cannot reach them effectively.
Cranes (Tacha, Nesbitt and Vohs ) and many geese and Southern
Screamers in South America need open sheetwater to roost overnight. For
wintering Lesser Snow Geese on the Gulf of Mexico coastal plain, use of a
region seems to depend upon the availability of such a roost area, and birds
may come and go at night if feeding conditions or disturbance patterns
demand.
Many opportunistic wetland users such as swallows or blackbirds perch on
emergent vegetation or trees over water, but they are subject to predation by
owls, mink, and perhaps snakes. Ground roosters like Northern Harriers may
be subject to predation by Great-horned Owls (Weller, Adams and Rose ),
but still larger perchers like herons and Ospreys use snags or posts in conspicuous places but are large enough to escape aerial predators.
. Thermoregulation
Because of their high body temperature, thermoregulation may seem of less
importance to birds than to other animals, but there is strong evidence that, in
addition to migrating to warmer regions during cold periods, they select
microhabitats to reduce chilling winds, overtopping waves, or other stressful
situations (Burger et al. ). Observers may chill at the sight of ducks splashing around in a pool in an ice-edged wetland, forgetting that water temperatures are above freezing within the water and are, therefore, a comparatively
warmer site in what may be a very cold ambient temperature. Waterfowl are
especially able to winter in cold areas because of their compact feathering and
heavy down layer. Canada Geese can even manage for weeks or months
without water if they have high-energy food resources like grains in the northern USA. Energetic costs of various types of activity are better understood now
(Wooley and Owen ) and are related to how birds maintain body temperature. Mallard ducks may remain buried in snow or other heat-preserving shelters rather than go out to feed on severely cold days presumably saving more
energy than they could take in (Jorde et al. ). Black Ducks wintering in
Thermoregulation
coastal Maine conserve heat behaviorally by tucking their feet into flank feathers and their bill into scapular feathers (Brodsky and Weatherhead ), and
by selecting sheltering landforms (Longcore and Gibbs ).
. Escape cover
The need for and use of predator escape cover or conditions vary greatly
among groups and species. Most of us immediately think of the swimming
waterbirds that can and do dive when pursued from above or within the water,
but waders and walkers gain surface protection only by the presence of water,
and the ability to fly from ground predators. Anyone who has observed waterbirds for long has seen bobcats, coyotes, or foxes attempt such predation, but
their success at catching ducks seems low except during nesting hence the
high mortality of females, especially common in ground-nesting species. It is
particularly important that rearing areas have water and food to sustain the
young during their flightless period, which can be quite long in geese and
swans.
attract birds to build nests there (see Knopf and Sedgwick ). Among
wetland birds, nest sites typically are found in microhabitats ranging from
highly aquatic to terrestrial: (i) completely floating nests of buoyant vegetation (small grebes); (ii) in water but essentially resting on some substrate
(some rails and ducks); (iii) above water and remote from shore (Least
Bitterns, herons); (iv) near shore but at wet-to-damp sites (some rails,
American Bitterns, and ducks); (v) underground burrows of mammals or
natural crevices (some goldeneyes population); (vi) dry ground with varying
degrees of short, herbaceous cover, at various distances from, but associated
with, water (Common Yellowthroats, Sedge Wrens, some ducks); (vii) at bases
of tall emergent vegetation, over land or water (sparrows, some New World
blackbirds); (viii) mid-level in robust herbaceous vegetation or small trees
that can support the weight of nest, eggs or young, and the incubating parent
(New World blackbirds); (ix) at the top of sturdy vegetation such as trees or
snags (Ospreys, certain eagles, herons); (x) old nests in trees (Speckled Teal in
Monk Parakeet nests); (xi) tree holes created by woodpeckers (Bufflehead)
(Erskine ), small tree cavities (Prothonotary Warbler), larger tree cavities
or crevices (several sheldgeese, Hooded Mergansers, Wood Ducks), cliff faces,
or solid soil banks (kingfishers); and (xii) cliff ledges (several species of
geese).
Dependent upon their precocity at hatching and their rate of growth,
young of some species like Moorhens (Greij ) and coots (Frederickson
) use the hatching nest or special brooding ramps over water to provide
predator protection. Others like ducks and geese are so precocious that they
rarely return to the nest after the first day or two.
References
Schmidt-Nielsen, K. and Kim, Y. T. (). The effect of salt intake on the size and
function of the salt glands in ducks. Auk , .
Schulthorpe, C. D. (). The biology of aquatic vascular plants. New York: St
Martins Press,
Siegfried, W. R. (). The Black Duck in the South-western Cape. Ostrich , .
Stanley, E. H., Fisher, S. G., and Grimm, N. B. (). Ecosystem expansion and
contraction in streams. Bioscience , .
Stresemann, E. and Stresemann, V. (). Die Mauser der Vogel. Journal fr
Ornithologie Sonderheft , .
Tacha, T. C., Nesbitt, S. A., and Vohs, P. A. (). Sandhill crane. In Migratory shore
and upland game bird management in North America, eds. T. C. Tacha and C. E.
Braun, pp. . Washington DC: International Association Of Fish and
Wildlife Agencies.
Thompson, D. Q. and Person, R. A. (). The eider pass at Point Barrow, Alaska.
Journal of Wildlife Management , .
Tietje, W. D. and Teer, J. G. (). Winter body condition of Northern Shovelers
on freshwater and saline habitats. In Waterfowl in winter, ed. M. W. Weller,
pp. . Minneapolis, MN: Univ. of Minnesota Press.
Tyler, S. and Ormerod, S. (). The dippers. London: T and AD Poyser.
Weller, M. W. (). Habitat selection and feeding patterns of Brown Teal
(Anas castanea chlorotis) on Great Barrier Island. Notornis , .
Weller, M. W. (). The island waterfowl. Ames IA: Iowa State Univ. Press.
Weller, M. W. (). Use of two waterbird guilds as evaluation tools for the
Kissimmee River restoration. Restoration Ecology , .
Weller, M. W., Adams, I. C., and Rose, B. J. (). Winter roosts of marsh hawks and
short-eared owls in Central Missouri. Wilson Bulletin , .
Williams, M. (). The moult gatherings of Paradise Shelduck in the Gisborne-East
District. Notornis , .
Withers, K. and Chapman, B. R. (). Seasonal abundance and habitat use of shorebirds on an Oso Bay mudflat, Corpus Christi, Texas. Journal of Field Ornithology
, .
Woodin, M. C. (). Use of saltwater and freshwater habitats by wintering redheads
in southern Texas. In Hydrobiologia, Vol. : Aquatic birds in the trophic web
of lakes, ed. K. K. Kerekes, pp. .
Wooley, J. B. Jr and Owen, R. B. (). Energy costs of activity and daily energy
expenditures in the black duck. Journal of Wildlife Management , .
Woolfenden, G. E. (). Selection for a delayed simultaneous wing molt in loons
(Gavidae). Wilson Bulletin , .
5
Foods, feeding tactics, strategies, and guilds
Food resources within wetlands can be diverse and vary temporally and spatially. Birds are unique among vertebrates in their ability to use wetlands dispersed over hundreds or thousands of miles in their annual range. However,
they are most restricted in movement during the nesting and rearing season or
when they are flightless owing to age or annual wing molt. In addition to flight,
they may have the flexibility of swimming, running, or walking as needed and
efficient, and this reflects the end-products of long years of trial and error. This
should not infer that all wetland birds are wandering long distances in search
of food, but that they could and sometimes do. Typically, they are bound by a
series of historical and evolutionary events that have resulted in the development of successful patterns of movement and breeding strategies for a given
region, which is intimately tied to food resources.
Finding accurate information on foods used by birds seasonally and at
various life-cycle stages and ages is difficult. An old but still valuable summary
of food use by birds in North America is Martin, Zimm and Nelson (), but
this and papers of this period contain some errors resulting from the examination of gizzards rather than esophagi alone for some bird groups. This underestimates soft animal matter and exaggerates hard seeds. More available but
less quantitative summaries of general food uses are in Erlich, Dobkin and
Wheye () and Kaufman () in the USA and in similar treaments of bird
life histories in each region or country. In addition, there are numerous
detailed studies of selected species; however review papers for various taxa,
where they exist, are the best source.
forces that drive vegetation. Annual plants are especially common to temporary floodwaters and non-flowing waters; they produce large seed crops useful
to pioneering species. Plants of longer hydroperiods tend to produce tubers or
rhizomes, but some also produce seed concurrently or under certain stressful
conditions such as drought. Vegetation structure is influenced by adaptations
to water permanence, timing, current, and climatic and salinity regimes,
resulting in microhabitats with differing vegetation structures such as nonpersistent emergents, persistent or robust emergents with sufficient structure that
they survive into the next growing season or longer, woody forest or shrub
species that may grow continually for many years, or aquatic beds made up of
diverse annual or perennial underwater plants that tend to reoccur annually
when conditions permit (Fig. .).
Aquatic vertebrates like amphibians and reptiles (herps), fish, or wetland
mammals like rice rats are major foods of predaceous wetland birds and other
vertebrates. But as consumers, they also are linked directly to vegetation, invertebrates, or other smaller vertebrates for food; directly to water for protection;
and indirectly to water as the driving force for wetland plant and animal
succession over time. Vertebrates of temporary waters, like amphibians, have
periodic population bursts based on time-limited water, plant, and invertebrate resources: suddenly they are gone many eaten by wetland predaceous
birds! More aquatic vertebrates like fish, while geared to longer hydroperiods
and more permanent areas, include species such as the mosquito fish, which
populate, thrive, and die in temporary waters of shallow basins or intermittent
streams. These are also eaten by wetland birds when vulnerable. Fish eggs often
are laid in shallow waters and are a favorite food of many birds, especially in the
spring before the egg-laying period.
Although smaller, the mass and diversity of invertebrates makes them the
choice food for many breeding birds and for some species all year. Many if not
most wetlands are rich in plant biomass and, therefore, are dominated by
herbivores (mostly invertebrates but some mammals and birds) and detrivores (mostly invertebrates). A recent summary of data on use of invertebrates
by shorebirds showed over genera of phyla, with significant flexibility in
use (Skagen and Oman ). In some species, birds at stopovers may double
their body weight in a few weeks while feeding on invertebrate eggs and larva
(Harrington ). Invertebrates also are regulated by annual hydroperiods
and year-to-year water permanence: short-cycle invertebrates vary dramatically in abundance according to timing and duration of water; long-lived
invertebrates with long developmental stages like dragonflies are typical of
longer hydroperiods and even more permanent waters. Birds are well adapted
to exploit this range of wetland habitats. They move easily, test the resources,
and make a choice to establish breeding areas or wintering sites; when they
have accomplished their functional goals, or when food resources or water
regimes change, they move on.
What foods are rare or absent in wetlands that might influence which bird
species can survive? There are few flowering plants among the truly aquatic
plants of lake systems (Philbrick and Les ), but more probably occur in
wetlands. Few, however, have large and prominent flowers except those that
float (e.g., water lilies); therefore, it is not surprising that there seem to be few
nectar-feeders among wetland birds. Hummingbirds do use jewelweed and
other plants of wetland margins that have flowers placed at a suitable and safe
height for feeding, but getting nectar from a water lily could be tricky.
Moreover, some water lilies have time-regulated, spiraling stems that pull the
flower head beneath the waters surface, where the seeds can mature in a protected environment (Riemer ). However, diving seed eaters like wintering
Ring-necked Ducks tap such underwater resources as a major food in southeastern USA wetlands. There also seem to be few plants that produce large
berries and other fleshy fruits in herbaceous and shrub wetlands; as a result,
birds like tanagers are not prominent except in forested areas, and they tend to
avoid water. Orioles may feed on nectar or fruits, and while they use tall trees
adjacent to wetlands, they seem to avoid feeding in wetlands or nesting overwater. Seed production can be impressive, and ground-feeding birds like sparrows, doves and pigeons, quail, and pheasant may use shallow wetlands when
they dry in autumn and early winter, but dense vegetation is a danger to larger
ground-feeding species.
A model of these habitat/food/bird relationships (Fig. .) may provide
some perspective on the dynamics and interrelationships and provide a basis
for the topics that follow. How the bird feeds, what it eats, how efficient it is,
and how it might compete with other species are strongly influenced by the
nature of the wetland it chooses. Its food choice is based on its trophic niche
and its morphological and physiological adaptations, all steeped in evolutionary history. Therefore, wetland choice, and the precise habitat or microhabitat
within the wetland, and food needs for specific stages in the annual life cycle
influence how the bird forages: its overall strategy and its immediate tactic for
food intake (Table .).
. Foraging behavior
Before examining factors that influence food use by waterbirds, it is essential to
think in terms of survival needs and adaptations that impose restrictions on
how and why birds feed as they do. Most center about energy efficiency in
obtaining food. The searching for and intake of food is termed foraging and
obviously must be efficient as well as effective if the animal is to survive. We
often assume that foods are abundant and unlimited for birds, but starvation
and poor health is common, and many studies of food habits of birds have
noted individuals that have spent long periods foraging but had near-empty
digestive tracts when collected. I have seen several cases where waterbirds are
limited in access to foods and show intention movements and push the limits
to get to food quickly. Kelp Geese of Patagonia and offshore islands feed at low
tide by walking among the rocky shore to gather sea lettuce and other large and
leafy but often heavily grazed seaweeds. But tides are extreme at that latitude
and the wait is often long, and some individuals will arrive at feeding areas and
swim above the rocks, even tipping-up like ducks, in an effort to obtain food
quickly. Another example of time pressurers is where light limits availability:
Microhabitat
Basin substrate
Epibenthic
Endobenthic
Dry (epibenthic)
Mudflat
Water column
Surface
Above water
Shoreline
Wet (endobenthic)
Planktonic (drifters)
Daphnia, cyclops
Neustonic (surface)
Nekton(underwater swimmers)
Waters surface
Seeds, invertebrates
Herbaceous plants
Woody plants
White Pelicans in the Gulf of Mexico capture fish that at night are attracted to
pier lights. These birds fly from daytime roosts, often arriving before capture is
efficient, and loaf until darkness arrives and the lights go on. In both examples,
one assumes that hunger triggers the feeding response and that feeding is less
successful elsewhere.
The optimization of effort or energy expenditures in the search for food
has been termed the optimal foraging theory (Stephens and Krebs ); it
assumes that birds follow the most efficient strategies and tactics that produce
the needed food. It helps us to understand why the same species of birds do not
all feed in the same way in all areas or at all times. They are doing what it takes
in that particular situation to gain the best quality food in the shortest time and
with the least effort. This also infers evolutionary adaptations and selection
toward certain feeding strategies, and specializations may be the end-product.
However, the roles of habit, learning experiences, and innate flexibility are
more difficult to evaluate.
mobility patterns that ensure security, health, and survival of the species
during one or more life-history stages (more in the sense of diet selection, e.g.,
Ellis et al. ). Such patterns are not simply a matter of choice by the bird but
may be fixed by traditional or genetic adaptations that reflect an evolutionary
history which maximizes efficiency and ensures resources in these variable
environments. For each species, and probably for some regional populations,
the complex of habitat resources required and patterns of mobility to accomplish this result from a long evolutionary history that has pre-adapted the bird
for what we consider typical habitat, resulting in preference for and selection
of that structural habitat pattern. Hence, there is a scale of foraging dictated by
general foods and strategies: a fish-eating cormorant, a duck diving for invertebrates that are hidden in rocks, and a shorebird moving along the shore to
opportunistically grab clams or worms exposed by the waves all have different
scales and rates of movement as well as different anatomical specializations.
These adaptations may be sufficiently unique to create species termed specialists, such as the Snail Kite of the Everglades (Sykes ). These kites seem
limited by their food, Apple Snails, which are characteristic of semipermanent
wetlands; this potentially creates a crisis for the species when that habitat is
impacted naturally or by human intervention. Other species have been termed
generalists: feeding on whatever is most available and using diverse nesting
and resting sites. Such species have been able to fit into a variety of available
habitats, perhaps through exposure to dynamic conditions over areas and
time, and may be so widespread and abundant as to be a nuisance (e.g., many
New World blackbirds).
Several examples of strategies may indicate the complexity and diversity of
approaches required to meet annual needs. (i) Dabbling ducks feed on invertebrates in summer ponds and on seeds or foliage during wintering periods, thus
tapping the greatest abundance of each at the peak seasons. Some move to
marine habitats, but most favor wintering habitats that are similar to their
breeding areas. (ii) Redhead ducks use inland freshwaters for breeding and
molting; here, they and their young feed by diving in shallow water for benthic
invertebrates. In winter, however, they frequent mainly estuaries or large freshwater lakes, using quite different foods. (iii) Red Phalaropes extend such
habitat shifts further by moving from small freshwater tundra ponds to wintering well out at sea where they feed on invertebrates derived from upwelling
waters. (iv) The northward migration of Red Knots to tundra for breeding
areas is timed to spawning by Horseshoe Crabs on limited beach sites of the
Atlantic coast of the USA and to similar invertebrate pulses elsewhere. After
breeding, they use the resources of vast tidal mudflats rich in small clams and
other marine invertebrates, especially in migration and wintering in the
southern hemisphere. (v) Sedge Wrens seem to come and go as a breeding bird
of the Prairie Pothole Region and even breed at varying times of the summer
dependent on the geographic and seasonal distribution of rainfall. (vi) Wood
Storks adjust their breeding times to drying conditions, which create opportu-
Table . Examples of food and foraging tactics employed by some wetland birds
Guild (as denoted by
habitat/locomotion/
food taken)
Foods
Tactics
Species examples
Surface swimmers
Strain/sweep/grab
Dabble/grab
Blue-winged Teal
Invertebrates
Search/dipping
Phalarope
Inverted straining
Flamingos
Fish/invertebrates
Visual search/grab
Grebes, ducks
Fish
Search/spear
Anhinga
Invertebrates
Benthic straining
Tubers
Digging/rooting
Flight-feeders
Insects
Aerial hawking
Fish
Surface skim
Skimmer
Carrion
Surface grab
Many gulls
Fish
Surface grab
Fish/invertebrates
Shallow dive
Terns
Fish
Plunge-diver
Osprey, Pelican
Perch-divers
Fish/invertebrates
Plunge-diver
Kingfishers
Walkers/runners
Invertebrates
Search, grab
Sandpipers, plovers
Insects
Runsnatch
(out-of-water)
Waders/waiters
(in-water)
Impact feeders
Fish
Search/strike
Many herons
Fish/invertebrates
Probe/grab/sweep
Fish/mice
Wait
Fish/insects
Stalk
Fish
Wing-flashing
Reddish Egret
Fish
Wing-shading
Black Heron
Invertebrates
Probe
Dowitcher
Minnows
Baiting
Clams/shellfish
Gulls
Large eggs
Use of rocks
Bristle-thighed Curlew
nities for access to fish and other prey needed for nestlings. To accomplish this,
they shift breeding times and breeding areas.
Tactics and techniques (the latter I think of as more simple that the former)
of obtaining food are mechanistic but efficient actions that are still fairly consistent but do vary by food and microhabitat conditions. I have not attempted
here to define, differentiate, or describe and name these variations but rather
to indicate the array of techniques one can find (Table .). Ashmole ()
described many techniques used by seabirds that are similar to those of
. Guilds
Those species (related or not) that exploit the same resources in a similar way
can be termed guilds. Although this term is not used consistently, ecologists
have found it useful in describing important resource-seeking attributes when
comparing different species of birds or animals as diverse as birds and bees.
Some of the terms in common use for various guilds and others that I have
coined are listed in Table .; these are merely examples and not intended as a
descriptive system. It may be, in fact, better to maintain flexibility in terms but
to structure these hierarchically, to allow comparison at scales from large to
small. Feeding sites or microhabitats (Table .), i.e., the physical position in
the wetland, induce physical limitations on food gathering and thus influence
behavioral tactics and the use of locomotion, or the design of bill, leg, and
sometimes neck.
In attempting to apply information on guilds to assess characteristics of a
wetland or bird community, a common mistake is to use a taxonomic category
such as waterfowl, assuming that all species in the group are similar in foods
because they have generally similar body shapes, bills, feet, etc. In fact, there are
over species of waterfowl worldwide (Table .) and they range from pure
herbivores that walk on land most of the time to pure carnivores that eat
animal foods all year. Most are omnivores, and their foods differ by season, so
temporal scale as well as taxonomic level is vital in making realistic comparisons. Even comparing dabbling ducks with bay ducks is not a safe approach
because each tribe has a range of foods, microsites, and diving depths that segregate the species (e.g., Smith et al. ). For example, three dabbling ducks
are commonly seen in open water (shoveler, wigeon, and Gadwall) along with
coots, whereas most other dabbling ducks favor vegetated or otherwise protected areas. Shovelers typically sweep their bills side-to-side while straining
for plankton whereas wigeon and Gadwall are grabbing and cutting leaves
while feeding bill-under or upending. The coots (as well as bay ducks) may
dive to grab submergent vegetation such as wigeongrass or pondweeds, which
then is pirated by the widgeon.
At some times and places, shovelers may be followed by phalaropes, which
are not filter-feeders but are tapping the same invertebrates by their unique
probing action, sometimes swimming in circles (hence the name whirligig,
after the water beetle). The shovelers apparently are acting as beaters by stirring up invertebrates by their swimming actions. Shovelers use the same tactic
following swans, which are seeking submergent vegetation rather than invertebrates.
Another example showing how foods might be misleading in classifying
guilds is that of the Virginia Rail and Blue-winged Teal. Both are often omnivores during summer and both might have snails and seeds in their digestive
tract. However, because their feeding techniques (walking versus swimming)
and microsites (tall and dense vegetation versus semiopen, shallow water)
differ, most biologists would not consider them in the same guild, if categorizing at a fine scale. Some workers have noted significant variation in foraging
sites seasonally or through the day (Kelly and Wood ), part of which could
be a result of targeting available foods or selecting foods on the basis of needed
nutrition or energy. Therefore, limited observations at one time could be misleading and one must recognize such potential influences.
Krapu and Reineke ) and coots, but the degree to which this is necessary
for laying seems to vary by species (Arnold and Ankney ) and probably
habitat conditions at the site. During spring migration, most omnivorous dabbling ducks seem to fatten and also to increase their intake of invertebrates
prior to arrival at breeding marshes and, therefore, are less dependent on local
food conditions; those that nest later may fatten in the breeding marsh on local
foods (Northern Shovelers), and South African Red-billed Teal seemingly are
able to lay with minimal intake of invertebrates (Petrie ).
During nesting, the precocity of the young determines where foods are
obtained: colonial nesters like egrets, herons, pelicans, cormorants (see review
in Weller ) carry food many miles to feed young in the nest. Coots, rails,
and grebes swim within a few feet of the nest until the young are able to join
them. Young grebes seem more inclined than most swimmers to ride on the
back of adults for longer water travel. Although some ducklings can walk
several miles (Hochbaum ) and swim strong river currents when newly
hatched (Bellrose and Holm ), it is an obvious efficiency to have abundant
foods nearby.
Protein-rich feeding areas also are important for birds that shed their flight
feathers simultaneously, and it is not surprising that post-breeding moltmigrations of waterfowl take place at a time for that species that ensures both
food and water. Most ducks do this in mid to late summer, and movement to
more permanent waters is common, some even flying northward to ensure
protection from predators when small water areas might dry up (Bailey and
Titman , Bergman ). Loons (Palmer ) and some eiders delay wing
molt until reaching wintering or at least fall staging areas (Hohman, Ankney
and Gordon ). After molt, migrant and wintering birds are more likely to
use high-energy sources like carbohydrates (seeds, foliage, fruits) to maintain
body condition and sustain normal activities (Weller ).
Body condition, as judged by weight and fat storage, is a major requirement for migration in many bird species, especially those that fly long distances between stops (e.g., Ruddy Turnstones and Red Knots, Harrington
). It is fairly common for a few shorebirds to remain in wintering areas all
year, presumably because they are diseased, injured, or their body condition
has not reached a level essential for the long flight to Arctic areas (Johnston and
McFarlane ). Some short-range migrants do not seem to show this need to
fatten on long but few stopovers (Skagen and Oman ), but undoubtedly
many factors influence this strategy.
Many factors influence where foods are found within a wetland (i.e., spatial
distribution): bottom substrate materials, vegetation at all levels (layers),
water depth, tidal regimes, water temperature, or oxygen supply in the water.
For example, bottom sediments of sand might be suitable for clams and not for
midges. Submerged vegetation provides substrates for countless invertebrates
of varying sizes and food requirements. Emerging vegetation such as cattail
provides insect emergence sites for organisms like damselflies and dragonflies
that live underwater for months or years and crawl up vegetation to dry and
emerge from their juvenile exoskeleton as flying adults. Dead and dying
vegetation is suitable for detritivores when herbivores would not flourish, but
detritivores typically are dominant organisms in energy cycles of wetlands
and, therefore, are likely to be major food items for many species of bird. Water
depth influences temperature and oxygen levels in water, and thereby the fish
or invertebrate species than can thrive there. Declining water depth and low
oxygen levels make invertebrates and fish vulnerable to predation by ibises,
egrets, and herons. Mudflats are rich places for many invertebrates, and shorebirds focus their search there. These microsites strongly influence where birds
are found in a wetland. Table . provides a partial list of such feeding locations
within the wetland, more specific feeding sites, food types, and examples of
fairly common bird foods.
Foods used by birds vary with their body size, and although many have bills
and tactics designed for essential flexibility, they can be more efficient by
taking large prey (if of equal quality) and spending less energy chasing many
small items. For example, large herons tend to eat larger fish than small herons
(see Weller for review); large ducks take larger maximum animal prey
than small ones (Pehrsson , Weller ). Curlews take larger clams at
greater depths than do oystercatchers (Zwarts and Wanink ). (Fig. .),
which take larger and older clams than do Sanderlings (Myers, Williams and
Pitelka ). In fact, oystercatchers do not waste time with small clams, and
big ones are too deep to reach. Hence, there is an impact on one age-class of
clams, and, reciprocally, if the suitable age-class were unavailable, this would
decrease habitat (food) quality. This sort of optimization in bird/food size is
closely related to specialization, because a suitable bill may increase efficiency
of taking prey but decrease flexibility in food options, and may avoid extensive
overlap in foods and often habitat. Even specialists like shoveler ducks and
spoonbills can strain for multiple small items like zooplankton through the
specially designed straining lamellae along the sides of the mandible, or use the
nail on the tip of the bill to grab large items like snails when they are conspicuously abundant in late summer. In tidal areas, prey availability varies with tide
level, and tide-chasers like Sanderlings feed where and when food resources
are most available and efficiently taken (Connors et al. ).
Vulnerability of prey may be affected by numerous factors, such as lighting,
underwater plant cover, distraction behavior, or speed of the chase. Piscivores
like the Reddish Egret of North America and the Black Heron of Africa use
their wings to shade the waters (although in quite different ways), and several
species of smaller egrets and the Black Herons have yellow feet, which are
thought to attract fish. Black-crowned and Yellow-crowned Night Herons
commonly feed at dusk or night, seemingly finding crayfish and crabs especially vulnerable then. Night feeding by ducks has been associated with predator avoidance (Tamisier ) and human disturbance (Thompson ,
Thornburg, ), but such foraging is limited to strainer-feeders like shovelers
or Green-winged Teal, bottom dabblers or divers that need less visual information to obtain food, and probing mudflat or shallow-water birds like dowitchers, Willets, and curlews. Herons and cormorants seem to be more successful
in catching rough fish that are slow-moving.
Use of certain habitats that are potentially more stressful than others suggests the long-term evolution of suitable foraging strategies and adaptations in
morphology and physiology for specific foods, which make the bird successful
in this environmental setting. For example, most if not all waterbirds have salt
glands, but they vary by age and species in how large and efficient they are.
Some birds (e.g., Redheads) breed in freshwater probably because the young
cannot tolerate the salt, but adults and immatures later switch to higher salinity water in wintering areas; their salt glands change accordingly. Other species
are still better adapted and can tolerate salty conditions all year, where they not
only survive but reproduce (e.g., Common Eiders, avocets, stilts). Clearly,
those species less-well adapted avoid such habitats.
segregation (e.g., Holmes and Pitelka ), (Fig. .). Other birds use the same
foods in the same wetland but not always at the same time (Connors, Myers
and Pitelka ) or at the same habitat site (Holmes and Pitelka ). In a taxonomic array, adaptation of various taxa demonstrates reduced overlap in foraging sites and foods, as shown in waterfowl that are dominantly swim-feeders
(Krapu and Reineke ) (Fig. .). For perch-feeders like kingfishers, feeding
sites are important in segregation, as shown in tropical species, which may
avoid competition by selecting different perch heights from which to fish (Fig.
.).
Forsters and Black Terns both nest in prairie potholes but seem segregated
by nest sites and cover-water conditions, as well as by foods. They may nest
close together in the same wetland, but the Forsters Tern favors relatively elevated structures like muskrat lodges near large and open water where fish are
prevalent. In contrast, Black Terns usually feed on insects and are solitary
nesters in small pools surrounded by emergent vegetation. They use mats of
floating vegetation or muskrat feeding ramps for nests where available.
Some workers feel that the great abundance of foods in some wetland sites
and their availability have allowed some closely related species to feed on the
same resources at the same time as with mixed flocks of herons and egrets
catching fish or crayfish stranded in pools caused by declining water, macroinvertebrates at their population peak sought by many species of migrant
ducks and shorebirds, and feeding by many species at all habitat layers exploiting full grown and emerging marsh flies (Chironomidae) in northern
marshes. Water levels, near-freezing water, hot water with reduced oxygen
levels, which all influence vulnerability of invertebrates, frogs, and fish, and
plant-cover conditions are important in making some resources available. As a
result, interpreting species richness or predicting use by different species
requires an evaluation of birds in the area physically and behaviorally adapted
to efficiently using those foods under the prescribed conditions.
There are still further influences of foods on the presence of waterbirds that
will be detailed below: food density, influence on breeding seasons, and
impacts of birds on their resources. An assemblage of wetland birds is not a
fixed entity, but rather a collection of species with common adaptations that
are commonly seen together under certain water or weather conditions. They
are limited by food resources, and, therefore, we expect to see a gross relationship between food types, abundance, and populations of each species or guild.
Data pooled on Falkland Island waterfowl (Fig. .) schematically show the
relationship of abundance to food or trophic niches and to habitat.
References
Alisauskas, R. T. and Ankney, C. D. (). The cost of egg laying and its relationship
to nutrient reserves in waterfowl. In Ecology and management of breeding waterfowl, eds. B. D. J. Batt, A. D. Afton, M. G. Anderson, C. D. Ankney, D. H. Johnson, J.
A. Kadlec, and G. L. Krapu, pp. . Minneapolis, MN: University of Minnesota
Press.
Arnold, T. W. and Ankney, C. D. (). The adaptive significance of nutrient reserves
to breeding American Coots: a reassessment. Condor , .
Ashmole, N. P. (). Seabird ecology and the marine environment. In Avian Biology,
Vol. I, eds. D. S. Farner and J. R. King, pp. . New York: Academic Press.
Bailey, R. O. and Titman, R. D. (). Habitat use and feeding ecology of postbreeding Redheads. Journal of Wildlife Management , .
Bellrose, F. C. and Holm, D. J. (). Ecology and management of the wood duck.
Washington, DC: Wildlife Management Institute.
Bergman, R. D. (). Use of southern boreal lakes by postbreeding canvasbacks
and redheads. Journal of Wildlife Management , .
Burnett, J. (). A further observation of a feeding association between the
Platypus and the Azure Kingfisher and a discussion of feeding associations
between birds and mammals. Sunbird , .
Connors, P. G., Myers, J. P., Connors, C. S. W., and Pitelka, F. A. (). Interhabitat
movements by sanderlings in relation to foraging profitability and the tidal cycle.
Auk , .
Connors, P. G., Myers, J. P. and Pitelka, F. A. (). Seasonal habitat use by Arctic
Alaskan shorebirds. Studies in Avian Biology , .
Croft, P. (). Grebe foraging with jacanas. Australian Birds , .
Ellis, J. E., Weins, J. A., Rodell, C. F., and Anway, J. C. (). A conceptual model of
diet selection as an ecosystem process. Journal of Theoretical Biology , .
Erlich, P. R., Dobkin, D. S., and Wheye, D. (). The birders handbook. New York:
Simon and Schuster.
References
Douthwaite, R. J. (). Seasonal changes in the food supply, numbers and male
plumages of pigmy geese on the Thamalakane river in northern Botswana.
Wildfowl , .
Emlen, S.T. and Ambrose III, H. W. (). Feeding interactions of Snowy Egrets and
Red-breasted Mergansers. Auk , .
Eriksson, M. O. G. (). Lake selection by Goldeneye ducklings in relation to the
abundance of food. Wildfowl , .
Esler, D. (). Avian community responses to hydrilla invasion. Wilson Bulletin
, .
Evans, P. R. and Dugan, P. J. (). Coastal birds: numbers in relation to food
resources. In Coastal waders and wildfowl in winter, eds. P. R. Evans, J. D. GossCustard, & W. G. Hale, pp. . Cambridge: Cambridge University Press.
Evans, P. R., Goss-Custard, J. D., and Hale, W. G. (eds.) (). Coastal waders and
wildfowl in winter. Cambridge: Cambridge University Press.
Glazener, W.C. (). Food habits of wild geese on the Gulf Coast of Texas. Journal
of Wildlife Management , .
Goodman, D. G. and Fisher, H. I. (). Functional anatomy of the feeding apparatus
in waterfowl. Carbondale, IL: Southern Illinois University Press.
Gray, L. J. (). Response of insectivorous birds to emerging aquatic insects in
riparian habitats of a tallgrass prairie stream. American Midland Naturalist ,
.
Joyner, D. E. (). Influence of invertebrates on pond selection by ducks in Ontario.
Journal of Wildlife Management , .
Kaminski, R. M., and Prince, H. H. (). Dabbling duck activity and foraging
response to aquatic macroinvertebrates. Auk , .
Kaminski, R. M. and Prince, H. H. (). Dabbling duck-habitat associations during
spring in the Delta Marsh, Manitoba. Journal of Wildlife Management , .
Kantrud, H., Krapu, G. L., and Swanson, G. A. (). Prairie basin wetlands of the
Dakotas: a community profile. Biology Report (.). Washington, DC: US Fish
& Wildlife Service.
Krapu, G. L. (). Feeding ecology of pintail hens during reproduction. Auk ,
.
Kushlan, J. A. (). Quantitative sampling of fish populations in shallow, freshwater
environments. Transactions of the American Fisheries Society : .
Lingle, G. R. and Sloan, N. F. (). Food habits of White Pelicans during and
at Chase Lake National Wildlife Refuge, North Dakota. Wilson Bulletin ,
.
Maltby, E. (). Waterlogged wealth. London and Washington, DC: International
Institute for Environment and Development, Earthscan.
Murkin, H. R., Kaminski, R. M., and Titman, R. D. (). Responses by dabbling
ducks and aquatic invertebrates to an experimentally manipulated cattail marsh.
Canadian Journal of Zoology , .
Nelson, J. W. and Kadlec, J. A. (). A conceptual approach to relating habitat structure and macroinvertebrate production in freshwater wetlands. Transactions of
the North American Wildlife and Natural Resource Conference , .
Orians, G. H. (). Some adaptations of marsh-nesting blackbirds. Princeton, NJ:
Princeton University Press.
Further reading
Orians, G. H. and Horn, H. S. (). Overlap in foods and foraging of four species
of blackbirds in the Potholes of central Washington. Ecology , .
Pehrsson, O. (). Food and feeding grounds of the goldeneye Buchephala
clangula (L.) on the Swedish west coast. Ornis Scandinavia , .
Remsen, J. V. Jr and Robinson, S. K. (). A classification scheme for foraging
behavior of birds in terrestrial habitats. Studies in Avian Biology , .
Root, R. B. (). The niche exploitation pattern of the blue-gray gnatcatcher.
Ecological Monographs , .
Rosine, W. N. (). The distribution of invertebrates on submerged aquatic plant
surfaces in Muskee Lake, Colorado. Ecology , .
Rundel, W. D. and Fredrickson, L. H. (). Managing seasonally flooded impoundments for migrant rails and shorebirds. Wildlife Society Bulletin , .
Shearer, L. A., Jahn, B. J., and Ienz, L. (). Deterioration of duck foods when
flooded. Journal of Wildlife Management , .
Shull, G. H. (). The longevity of submerged seed. Plant World : .
Swanson, G. A. and Meyer, M. I. (). The role of invertebrates in the feeding
ecology of Anatidae during the breeding season. In The Waterfowl Habitat
Management Symposium, pp. . Moncton, New Brunswick: The Wildlife
Society.
Swanson, G. A. and Meyer, M. I. (). Impact of fluctuating water levels on feeding
ecology of breeding blue-winged teal. Journal of Wildlife Management , .
Telfair II, R. C. (). The cattle egret: a Texas focus and a world view. College Station,
TX: Texas A&M University Press.
Thayer, G. W., Wolfe, D. A., and Williams, R. B. (). The impact of man on seagrass
systems. American Scientist , .
Tietje, W. D. and Teer, J. G. (). Winter feeding ecology of Northern Shovelers on
freshwater and saline wetlands in South Texas. Journal of Wildlife Management
, .
Voigts, D. K. (). Food niche overlap of two Iowa marsh icterids. Condor , .
Voigts, D. K. (). Aquatic invertebrate abundance in relation to changing marsh
vegetation. American Midland Naturalist , .
van Eerden, M. R. (). Waterfowl movements in relation to food stocks. In Coastal
waders and wildfowl in winter, eds. P. R. Evans, J. D. Goss-Custard, & W. G. Hale,
pp. . Cambridge: Cambridge University Press.
6
Bird mobility and wetland predictability
COLD
Seasonal
use
Figure .. A simple model of
the water regimes and climatic
factors that seem to drive
various types of movement of
wetland birds: regular latitudinal migrants with varying
stopover patterns, nondirectional nomadic movements,
and permanent residency.
MIGRATION:
MANY
STOPS
Permanent
Residency
LONG
MOVEMENTS:
FEW STOPOVERS
Periodic use
Nomadism
WET
WARMER
DRY
Resources: water/food/cover
B
Unpredictable
Ephemeral
Year
movement, flexibility in ability to lay eggs over a long period, and by having a
large clutch (Briggs ).
Changes in waterbird migration in some species have challenged long-held
ideas that these movements are all ancient patterns. Cattle Egrets first appeared
in Florida in the early s, and by the s had established breeding populations widely from southern Canada to Mexico and Florida. They now even
move out of south Texas in winter to Central America (Telfair ), where presumably they have more predictable water, temperature, and accompanying
food supplies. A similar extension of breeding range of Northern Pintails from
drought-impacted or drained prairie wetlands to Alaska has been documented
over several time periods, often based on banded (ringed) birds (Derksen and
Eldridge ). There also have been changes in southward migration routes of
Canvasbacks and other diving birds, which have shifted from areas of reduced
food supplies in polluted impoundments to richer but more distant areas
(Bellrose ). Reservoir construction along the Missouri River has resulted
in changes in migration routes of Canada Geese, in their choice of wintering
areas, and in delayed flights to final wintering sites while they exploit assured
resources (Simpson ). Other species of long-range migrants have
extended their ranges: the Pacific Black Brant, which moves from Alaskan or
Siberian nesting areas to coastal estuaries of the west coast of North America,
now moves into Mexican estuaries in winter. Its resources are marine,
characteristic of fringing wetlands, and its migration is linear along the coast
but with long overseas flight as necessary to reach its resources. Stops are relatively few but probably fairly long, like those of some shorebirds, making the
protection of these sites crucial for the survival of the species. Most studies of
west coast estuaries suggest that the extension to Mexico has resulted from
human disturbance in estuarine feeding areas along the west coast (Henry
) and reduced availability of eelgrass caused by pollution and humanrelated disturbance (Wilson and Atkinson ). More recently still, the species
seems also to be using estuaries in Japan (Derksen et al. ).
dynamic waters, as drying and aeration result in nutrient cycling and natural
fertilization. Dependent upon the local climatic regime, many wetlands tend to
be wet in spring and early summer and dry later; thus becoming less suitable for
waterbirds in autumn or for overwintering; even where temperature regimes
would permit use year-round. Movements of birds like ducks, geese, rails, and
shorebirds utilize many of these wet areas and often favor small areas in spring
and the larger ones that still hold water in the autumn. The regularity of these
water regimes probably has been an important influence on strategies of
migration, as suggested by shorebirds that stop at the same wetlands but come
various distances from breeding and wintering areas (Skagen and Knopf ).
Not only do some northerly shorebirds use these arid-land wetlands during
migration, but residents often are birds like avocets, Willets, and Black-necked
Stilts, which can tolerate and even favor shallow saline (i.e. akaline or soda) wetlands. Other migrants follow a loop strategy of migration that exploits the
resources of the Great Plains wetlands in spring (White-rumped Sandpiper,
Hudsonian Godwit, and Lesser Golden Plover) but not in the autumn, when
instead they move from breeding areas to the northeast coast where water and
food may be more reliable than in the dry plains (Zimmerman ), and from
where they leave on long overseas flights to South America. A few species follow
a reverse pattern (Western Sandpiper), presumably because resources for
spring migration are better on the western coast (Zimmerman ).
Many shorebirds and ducks make short flights between residual wetlands
of extensive complexes, these areas becoming major resting and feeding areas
at times (Bellrose and Crompton , Skagen and Knopf , ).
Presently, there is great concern about human-induced modification of the
shallow but interior wetlands that once formed major stopovers for migrant
shorebirds and now are drying as a result of alternative water uses
(Zimmerman ) or drainage (Krapu , Skagen and Knopf ).
Obviously, society must learn that a wetland which is used only a few weeks or
months a year is an essential element of the life cycle of millions of birds
treking thousands of miles in a constant search for resources.
It is more than chance that waterbird wintering areas tend to have seasonal
rainfall regimes that provide water resources, and that many are near coasts or
large interior water bodies that ensure still more dependable resources. The
resource tracking pattern of interior breeding waterbirds also involves movements of significant distance and direction as needed. For example, waterfowl
from the Prairie Pothole Region or large and saline Great Basin marshes move
to winter in coastal freshwater ponds or saline bays along the Gulf of Mexico
(Fig. .). Blue-winged Teal move into Central America or northern South
America (Botero and Rusch , Saunders and Saunders , Weller ),
but despite their powerful flight capabilities, few cross the Tropic of Cancer. I
suspect that many of the wetlands beyond this latitude are less predictable in
their suitability because of the reverse seasonal patterns, which influence
whether they can serve as migration stops and wintering areas.
. Nomadism
Birds have been termed nomadic when they are geographically and temporally
unpredictable; these birds seek out suitable conditions for nesting, feeding,
and other functions (Briggs ). Such shifts of populations and assemblages
are renowned in Australia (Frith ) but also occur in Africa (Douthwaite
, Milstein ) and India (Ali , Breeden and Breeden ). In some
cases, drying conditions induce birds to move to new wetlands within the
region that are more suitable for nesting. In larger areas and where temperature is nonlimiting, some species seem merely to wait until rainfall results in
suitable conditions. Droughts are common to the southern USA where
reduced late winter and spring rainfall causes potential breeders such as coots,
grebes, and ducks to move out of drying wetlands because they cannot swim
and find food and security. Nomadic shifts of breeding ducks and presumably
other wetland birds also are common in the western basin and northern USA
glacial potholes, with populations moving long distances but often after
breeding when both young and adults may capitalize on foods elsewhere.
Based on my personal experience in large emergent marshes in Argentina, and
discussion with residents and other biologists who have worked there, this
type of nomadic response is common. Both in northern Argentina and in the
southern USA, movements and breeding of the Masked Duck reflect a species
that is unpredictable in nesting location because its habitat requirements are
precise. In some way, it seems to locate suitable newly flooded and densely vegetated wetlands or ricefields. When conditions are suitable, the bird may
appear, as it has several times recently in southern Texas (Blackenship and
Anderson ).
The term nomadism has also been applied to regional shifts in breeding
colonies of White Ibis, which like Wood Storks have moved out of Florida and
northward into Georgia and the Carolinas (Frederick et al. ).
Concurrently, heron and egret populations have decreased in Florida and
Texas but increased in Louisiana, possibly because of wetland resources produced by the predictability of extensive crayfish aquaculture (Fleury and
Sherry ). In such cases, one can only speculate whether this represents
growth of newly established populations with concurrent decline of old
colonies or actual shifts in populations. Sedge Wrens seemingly have small territories and meet their nesting needs in very small areas, but they must be very
mobile to find these because they use such short-lived habitats produced by
flooding and drying. They, therefore, are unpredictable (on the basis of our
present understanding) in their presence in a prairie wetland and may be
breeding in June in one area and in August in another (Bedell ). They seem
more nomadic within their breeding range but are still long-distance
migrants. However, this may well be more the norm than the exception for
many inconspicuous birds of shallow and temporary wetlands.
Drought during late summer also can attract post-breeding waterbirds
Nomadism
such as Wood Storks, ibises, and egrets to move into drying wetlands to take
vulnerable fish and amphibians essential to their breeding cycle. Winter
droughts in the same areas mean poor stopovers for migrants and poor wintering areas.
. Local movements
Movements also can be viewed at the scale of a single wetland, as in a swamp
where habitat heterogeneity is great or in fringing wetlands along large lakes or
seashore. After reaching such an area following long-distance movement, the
establishment of small and resource-rich breeding territories, defended or
undefended feeding sites (varying by species or group), or the more extensive
home range may require only localized movements even when powers of flight
are strong (wrens, blackbirds, warblers, and sparrows). It is probably a matter
of efficiency but also of body size and resource distribution. Other species,
perhaps those more dependent upon water, often build the local habitat unit
around a wetland complex that provides various needs but also may act as a
backup in event of catastrophic change. In breeding areas, some species seem
to have minimal mobility (coots, grebes, loons) so selection of the wetland is
crucial; in fact, some water-adapted species like grebes may temporarily lose
their powers of flight owing to muscle degeneration. Waterbirds that are
shoreline and mudflat-walkers or that wade in shallow water are highly mobile
because foods are so closely linked to precise water depths; a centimeter can
make major differences in invertebrate distribution.
Krapu and Reineke ). Examples of the three trophic niches may help us to
examine similar relationships in other waterbirds. Specialized, year-long carnivores (typically invertivores or piscivores) are species like eiders and scoters,
which eat mussels and clams, and mergansers, which eat fish or invertebrates;
plant food seems rare. These birds may shift geographic location and habitat
but seem to have minimal major changes in foods. Herbivores are fairly
common among waterfowl, and their digestive system seemingly allows little
use of animal matter. Black Brant feed on foliage of either seagrasses or sedges
and grasses at breeding sites and almost exclusively on seagrasses during
migration and in winter. Interior freshwater herbivores like geese feed on
grasses and sedges and are prominent where these foods are common, even in
winter. Southern Screamers seem to feed on floating aquatics during nesting
and on upland grasses during nonbreeding periods (Weller ).
Omnivory is more common among waterfowl presumably because it is
efficient in fulfilling nutrient and energy needs, and in capitalizing on the most
abundant food at the most convenient time of year (Fig.e .). Redhead Ducks
were cited above as birds of moderate to larger wetlands and more open waters.
The diminutive dabbling teal (such as the Green-winged Teal of North
America, the Common Teal of Eurasia, and the Speckled Teal of South
America) generally favor smaller wetlands for breeding, or the muddy edges of
large ones. Teal are small bodied and have tiny lamellae along the edges of their
bills that allow them to sort out the tiniest of seeds and invertebrates, making
them suitable to use small wetlands of diverse types over a wide range of
vegetation and salinity. Their upper mandible is equipped with a nail on the tip
and apparently works on large food items too, as they feed on snails and clams
and have been observed floating down the Columbia River on dead salmon
eating all the way!
In other orders of waterbirds (Table .), similar ranges of food use and
adaptation occur. First, many are carnivores and lack dramatic seasonal food
change (e.g., grebes, loons, cormorants, pelicans, herons, shorebirds,
flycatchers, etc.). Some change in food size and taxon probably occurs during
feeding young, and adults are more flexible in feeding sites because they are
less restricted to the nesting area. Their movements naturally are geared to the
availability of open (i.e., unfrozen) water of suitable depths for capturing
aquatic prey.
As noted among the ducks, many if not most wetland birds seem to be
omnivores, and shifts in food are complicated by sites and wetlands types (e.g.,
coots, rails, warblers, sparrows, blackbirds, etc.). Herbivores seem less
common, presumably because use of pure plant foods requires considerable
specialization of the digestive system. Perhaps the most specialized is a marginal wetland bird, the Hoatzin, which eats coarse leaves of tropical trees, but
its mobility seems to be limited. Some flamingos feed on algae rather than
macroinvertebrates, which may explain some of their massive if not long-distance movements to track suitable water conditions. Cranes demonstrate
considerable variation in movements based on season and functional needs:
many use marshes during nesting, wet meadows for tubers during migration,
and upland areas in nonbreeding periods.
Although mobility is common and expected, it differs among groups and
much is still unknown. Patterns are especially difficult to measure and interpret when birds are adaptable and opportunistic. It is important, therefore,
that interesting observations of even single incidents or suspected relationships be published so that they can be synthesized for pattern identification.
Mobility presumably has developed for many reasons, only some of which
have been addressed here, and these examples from wetland birds may provide
clues to causes of mobility in other birds where the driving forces may be more
obscure.
References
Alerstam, T. (). Strategies of migratory flight, illustrated by Arctic and Common
Terns, Sterna paradisea and Sterna hirundo. In Contribution in Marine Science
Vol. : Migration: mechanisms and adaptive significance, ed. M. A. Rankin,
pp. . Austin, TX: University of Texas .
Ali, S. (). A book of Indian birds. Bombay: Bombay Natural History Society.
Bedell, P. A. (). Evidence of dual breeding ranges for the Sedge Wren in the
Central Great Plains. Wilson Bulletin , .
Bellrose, F. C. (). A spectacular waterfowl migration through central North
America. Illinois Natural History Survey Biological Notes , .
Bellrose, F. C. (). Waterfowl migration corridors. Illinois Natural History Survey
Biological Notes , .
Bellrose, F. C. (). Ducks, geese and swans of North America, rd edn. Washington,
DC: Wildlife Management Institute.
Bellrose, F. C. and Crompton, R. D. (). Migration behavior of mallards and black
ducks as determined from banding. Illinois Natural History Survey Bulletin ,
.
Blakenship, T. L. and Anderson, J. T. (). A large concentration of Masked Ducks
References
Further reading
7
Other behavioral and physical influences
on wetland living
hatch and growth of young before many of the freshwater ponds dry.
Therefore, the two species can coexist for part of the year, using slightly
different microhabitats, and show little or no overt aggression where they
overlap. An interesting but dangerous experiment was the release of large
numbers of hand-reared but wild-stock Mallards in the range of the Mottled
Duck in South Texas (Kiel ). After many years, Mallards have failed to
survive in free-living, viable populations, suggesting that they probably do not
do the right thing at the right time in this semiarid area. Because the typical
water regimes in northern wetlands involves spring breeding by Mallards,
those trying to breed in South Texas in spring and early summer often would
meet with drying freshwater ponds, and young would be placed in jeopardy
seeking new foraging areas. Mallards do use estuarine areas in some places, but
most populations seem to avoid saltwater when possible. Perhaps young
Mottled Ducks may be better adapted to saltwater than Mallards, but they do
seem to favor freshwater habitats until they can better tolerate salt after the first
few weeks of life.
The Wood Stork, a species often thought of in terms of lush and wet tropical wetlands, actually is linked to declining water regimes. It feeds its
young on fish and invertebrates, often obtained at long distances from the nest.
Its breeding time and location are dictated by water regimes in wetlands,
regardless of rainfall. Drying water areas make prey vulnerable to capture,
whereas flooded areas make prey less available (Kahl ). Nest success is
highly variable because of these wetland dynamics, and shifts in populations
have been attributed to excessive water in feeding areas. Like many colonial
waders, they commonly engage in post-breeding movements of hundreds of
miles or more northward to suitable feeding areas, exploiting the foods of
drying wetlands.
. Geographical range
Although local and regional habitat dictates presence or absence of a bird
species, there are evolutionary (historical) origins and other environmental
aspects of regions that influence the distribution of major taxa of birds, which
are treated in detail elsewhere (e.g., Vuilleumier ). Obviously, there are relatively few higher taxa (orders) that are worldwide, more families or genera
that are found on several continents, but few species that are worldwide among
birds. Without attempting a detailed analysis, waterbirds must rank high in
average size of the geographic range and in distribution on several continents
because they are strong fliers, can use small and isolated habitat islands (wetlands), and often are adaptable to a wide range of foods and breeding sites
(Weller ). However, owing to speciation during geographical isolation,
closely related species of the same genera do cover extensive areas of the world
as breeding habitat: pintail and Mallard ducks, Black-headed or Brown-
hooded Gulls, and several terns and shorebirds. Numerous shorebirds breed
widely in the northern hemisphere and winter widely in the extreme southern
latitudes.
Analyses of various taxa and continents from an ecological perspective
have been attempted for Anseriformes (Weller , , ) and
Podicipidiformes (Fjeldsa ), and a still wider ecological and taxonomic
analysis of waterbirds of South America has revealed some interesting patterns
of higher species richness of some groups in the low latitudes (tropical fisheating birds) and of omnivorous waterfowl at subtropical or temperate latitudes (Reicholf ). From a habitat perspective, these patterns can be
analyzed in only rather general ways because we have too little information on
individual species and because we are unable to assess the historical influence
that camouflages present habitat issues. They are of special interest, however,
because we can see how similar wetland habitats in different areas are used by
different taxonomic groups, such as the seeming ecological equivalents in
northern and southern hemispheres: Black Brant and Kelp Goose, and
Common Eiders and Flightless Steamer Ducks. Additionally, we need to consider interfamily and interorder competition for resources, which probably
occurs in some groups of wetland birds and may influence community
composition (Weller ).
Geographical patterns of bird distribution in wetlands lead one to speculate on the importance of historical events and dominance of highly competitive bird groups in the origin of current distribution. In North America,
blackbirds (Icteridae), and wrens and sparrows dominate the songbird community of emergent marshes, with highly social blackbirds being most abundant and conspicuous. In South America, a greater diversity of blackbirds
occurs (but few are colonial), but other species (representing several other
families not found in North America) nest over water in emergent vegetation.
In Europe, Icterids do not occur and there are at least ten warblers
(Muscicapidae and Silviidae) bearing such common names as swamp,
marsh, reed etc. because they nest and feed in emergent vegetation. In
North America, distribution patterns within wetlands range from that of the
near-shore or shallow-marsh species like Sedge Wrens or Swamp Sparrows to
that of species that like more open and deeper water like Yellow-headed
Blackbirds building nests in robust plants like cattail and bulrushes (Fig. .).
Clearly, different groups have exploited these rich wetland habitats all over the
world; if one taxon or specialist does not use a habitat, another does.
displays, mating, and identification of the nest site is possible. Courtship displays and copulation in the more aquatic waterbirds tends to take place on
water. Duck and grebe courtships are renown for their fascinating but often
confusing display signals, interesting vocalizations, and beauty of form and
color. Shorebirds use more terrestrial sites, have rapid, moving, and agressive
actions, and some have aerial displays like the bleating or winnie of the
Wilsons Snipe (a sound created by the tail feathers) or the aerial breaking
sound of American Woodcock (resulting from the special wing feathers).
Some shorebirds have elaborate display feathers like the collar of the Ruff.
Waders and coast-nesting seabirds commonly call and display from potential
nest sites to attract a mate. In such species, copulation occurs on the platform
or nest site and is tied to nest construction activities such as the ceremonial displays when nest material is brought to the nest. Males of more terrestrial areas
and marsh-edge birds like Red-winged Blackbirds or Swamp Sparrows typically establish territories by use of song perches, but the females check out the
vegetation, the general locale, and probably the food supply carefully before
establishing a site or pair attachment.
Multiple mating systems increase in wetland birds compared with other
terrestrial species. According to Verner and Willson (), only of
North American passerines are polygynous, and of these are wetland birds.
There are numerous cases in other taxonomic groups as well and these have
been the focus of major studies on the general aspects of breeding in birds. In
most species of polygamous marsh songbirds that have been studied in detail,
males defend large territories and attract and mate with several females (e.g.,
North American Marsh Wrens (Verner and Engelson ) and Red-winged
and Yellow-headed Blackbirds (Orians )). In such territorial species, those
with the highest quality territory (i.e., habitat resources) seem to attract the
most females, which has been attributed mainly to food availability. Studies in
South and Central America suggest that the lower density of food resources
may explain the reduced coloniality of blackbirds there compared with birds
nesting in North America (Orians ).
Although virtually all types of mating system are known in shorebirds,
polyandry, the mating of one female with more than one male, is especially
common. It may not be influenced by habitat, but it is at least known mainly
from this group; however, strategies vary. In some species of sandpipers and
plovers, the male of the pair incubates the first clutch and the female the
second, presumably taking advantage of optimal resources in a short time
frame. Female Spotted Sandpipers, which feed on small, temperate-zone
mudflats, lay a clutch of eggs, which a male then incubates. Later, she may lay
other clutches if conditions are suitable, and males remain ready to take over
the incubation duties. Presumably, locating males in such dispersed habitat
patches is difficult and females defend territories around the males. It is
assumed that resources are very important, especially the number of invertebrates, which is essential for egg-laying (Oring and Knudson ).
Among jacanas, females defend a territory with several males,which incubate the eggs she lays in nests built by the males (Emlen, Demong and Emlen
). In Red and Red-necked Phalaropes, females are more colorful in
plumage, have prominent courtship displays, and defend territories. Males
incubate the eggs and females devote no time to protection of the highly precocial young.
protection from ground predators and will reduce the travel time to take their
young for foraging. Only down is added to the decomposing tree cavity to
enhance an already ideal nesting environment (Breckenridge ).
Cormorants, herons, egrets, and ibises may build nests in large colonies on elevated sites (islands, shrubs, snags, trees, cliffs), carrying material and fighting
over it and stealing it from neighbors.
The more aquatic swans, geese, and ducks that nest over water in emergent
plants all use materials at hand (obviously selecting sites with suitable materials) and those nests on land are built of fine grasses and sedges and are downlined. Those over water may be raised in the vegetation, more by the bending
down of supporting plants which criss-cross over the water. As the bird sits on
this sparse foundation, it pulls material into the center and molds it into an
egg-holding as well as insulating structure. Some build an enclosing canopy
over the top by reaching up and pulling down tall emergent vegetation as it
grows (Redheads and Ruddy Ducks).
Highly aquatic grebes may build floating nests of wet plant debris derived
from the previous years growth, or sometimes living submergent vegetation.
Such nests are quite often in open areas, and coot and grebe nests may suffer
high failure rates through large or continuous waves (Boe , Fredrickson
).
One of the most unique nests and nest sites among waterbirds is a source of
curiosity not only in its origin but also in its functions. Nests of Horned Coots
are built of submergent vegetation on slightly submerged mounds of rocks in
water about half a meter ( ft) deep in Andean lakes at elevations of over m
( foot) (Ripley ). Birds also have been seen carrying large rocks from
the shoreline to deposit on ft-high rock islands, which subsequently serve as
sites to deposit vegetation and lay eggs. In shallow water, natural elevations of
the basin may be used. After nesting, other bird species like grebes, gulls, and
ducks may use these constructed sites. The closely related Giant Coot (actually
a little smaller than the Horned Coot) is essentially flightless and nests in still
higher lakes (up to m or ft) (McFarlane ). Nests are build of
floating masses of aquatic plants, which are protected from drifting by placement on underwater island-like undulations of the basin.
. Nest microhabitat
What influences the location of the nest in relation to habitat features? At the
proximate level, feeding success by the potentially nesting female must be of
primary concern. Availability of nesting materials, structural support, or cover
would be important in selecting the general area and the specific nest site.
These are all part of resource quality or habitat suitability. The role of microclimate has not been carefully explored for many wetland birds, but technological advances now allow detailed monitoring of climatic conditions.
The nesting strategy varies with the social structure, as territorial birds
protect their nests and often feeding areas.Where polygamy is involved, several
nest sites may be protected, but presumably it is the nest builder (whether
female or male) that selects the specific nest site within the territory (e.g., in
blackbirds, sparrows, and shorebirds). Nesting females with solitary nests and
which incubate alone presumably pick and protect their nest sites, often with
little involvement of the mates (e.g., ducks). Species with strong pair bonds
that share in incubation (Killdeer, stilts, avocets) share in alarm and defense.
Our question here is whether these patterns relate to habitats, directly or indirectly, or are merely reflections of a general reproductive strategy common to
the ancestral stock and subsequently adapted to wetter habitats. One probably
cannot be separated from the other (Southwood ), but I suspect the
influence of reproductive and foraging strategies blend in some optimized
pattern that works for the species or group: feeding strategy seems to dictate
territorial and social behavior, with seasonally carnivorous birds like songbirds
and shorebirds defending feeding areas that provide for young. But even some
herbivores are highly protective of nest sites (e.g., geese). Waterbirds feeding
on dispersed food resources generally seem less aggressive. Body size must be a
factor in terms of protection from potential predators and hence placement of
nests, for a Canada Goose would be less endangered by nesting on the ground
than a duck or shorebird. Egg size and size of the young must have a similar
influence in relation to the size of the predator, e.g., a small garter snake is
serious for a blackbird or sparrow but it would take a crocodile to influence a
goose nest site. Many studies of ground-nesting birds have shown a preference
for dense over light vegetation cover; the latter may result in greater visibility.
One study reported differences among two species of duck in avoiding sun or
favoring humidity but found other features of greater importance in dictating
nest site (Gloutney and Clark ).
. Body size
The significance of body size in birds is a current interest in ecology as it relates
to niche segregation (Nudds and Wickett ), distribution and rarity of
species (Gaston and Blackburn ), and population size (Blackburn and
Lawton ). Its importance here is the need to search for resource-related
patterns. Body size of waterbirds varies enormously, from fish eaters like pelicans (up to kg) and storks to large herbivores like screamers, swans, and geese
(>kg) to minute rails and passerines (<g). This range and complexity
allows for little analysis except between ecosystems, but patterns within taxonomic groups may be more meaningful. Among waterfowl, herbivores like
swans and geese typically exceed the size of the carnivorous sea ducks, which
in turn are larger than most omnivorous ducks. There are some general
Body size
relationships between body size and prey size in herons (see reviews in Weller
) and ducks (Nudds and Bowlby , Pehrsson ) (Fig. .), and probably in shorebirds (Holmes and Pitelka ) although opportunism strongly
influences this pattern. Although similar relationships are known in land
birds, the patterns are often quite dramatic among wetland birds because of
the diverse food sizes and types, and resulting specializations of bill, feet, and
foraging behavior.
tions of foods under certain conditions that make feeding less competitive, less
defensible, and mutually advantageous.
7.11.1 feeding
Clearly, freshwater habitats are rich and diverse in food resources and nesting
sites. Concentrated food is difficult to protect from congeneric competitors,
and such effort probably is not energy efficient except when nesting requires
the insuring of a food supply for the young. Moreover, food production in
many wetlands seems to be one of seasonal and/or hydrologic pulses, with
invertebrates available during mid to late summer and seed levels high during
autumn and winter. Vertebrate foods like amphibians and fish often are
vulnerable after the production of young and the decline of water levels: this
varies seasonally and regionally.At these times of abundant resources, flocks of
the same species and multispecies flocks may result.
It is difficult to identify patterns among these flocking species because of
regional variability and because of exploitation of opportunities for concentrated resources. Several studies have focused on such patterns by comparing
food habits of specific taxa. In Anseriformes, flocking tendencies are highest in
those species that are dominantly herbivores like geese, which roost together
and feed in large flocks, whereas carnivores are more likely to roost in smaller
groups or feed alone (Ydenberg and Prins ). Researchers concluded that
solitary search for mobile prey and individual competition influence these
social patterns. Such patterns vary according to resource need; for example
terns feed alone because of food dispersion but are social in nesting or roost
areas. There are many alternatives, but feeding tactics and food resources must
both play important roles in the evolution of such flocking behavior.
Of the birds listed above, many of the species that lead single life styles tend
to be carnivores, feeding by predation on insects, snails, mice, fish, frogs, or
birds all year, where individual action is essential and foods may be less concentrated. Many of the other flocking species are of three types: omnivores that
switch trophic niches seasonally (e.g., many ducks, blackbirds); herbivores
that eat plant foliage and seeds most of the year (geese, sheldgeese, and a few
ducks); and carnivores that feed as groups on vulnerable schooling fish,
amphibians, or large invertebrates (egrets, herons, cormorants, pelicans,
mergansers). But there are many exceptions induced by opportunity. For
example, many migrant shorebirds are social in feeding and are carnivorous,
feeding mostly on small, abundant, and dense invertebrate prey. Normally
solitary Snail Kites, Boat-tailed Grackles, and Limpkins occasionally are seen
in company with White Ibis and Great Egrets at low water levels, essentially
using them as beatersto disturb prey for hunting (Bennetts and Dreitz ).
Mixed-species flocks of feeding waterbirds are most common and best
known among pelicans, cormorants, ducks, sandpipers, egrets and herons,
and gulls. Pelicans and cormorants seem regularly to feed in groups and are
joined by several species of gulls. Often they are in masses of mixed species of
individuals feeding independently; others seem more organized forming
semicircles, rows, and wedges that may drive fish into more vulnerable positions. Presumably, this cooperative effort aids all in locating the schools and
perhaps in capture. Sometimes they are linked to porpoise activity, suggesting
that all are eating from the same school of fish. Moreover, when fish are large,
the leaders seem to be the cormorants while the White, and occasionally
Brown, Pelicans are stealing fish from the cormorants.
It is obvious that mixed flocks may be a human perception, and that
although the bird species are close together they are in small clusters keeping
visual and perhaps vocal contact, and often even take flight in groups by
species. This is especially evident among sandpipers and other shorebirds
feeding side by side. Mixed flocks also occur among aerial predators like swallows, which feed over marshes.
Gulls and terns seem often to be closely associated in feeding, but their
feeding tactics differ greatly, and the degree of food overlap probably is
minimal except among gulls that hawk for insects. Small terns feed on smaller
fish than do large ones, and some feed on surface or emerging invertebrates.
Large gulls seem more flexible and scavenge or take prey of all sizes in competition with smaller gulls, but large prey seem most likely to be preferred.
Swan or shoveler and phalarope associations occur commonly, in which
case they presumably are competing for the same but very abundant foods
only with shoveler. Snowy Egrets and Tricolor Herons have been observed
feeding symbiotically with Red-breasted Mergansers or Pied-billed Grebes
(Emlen and Ambrose ); clearly the diving bird could be taking deeper fish
or invertebrates disturbed by the waders searching for surface foods.
Relationships between birds and aquatic mammals were mentioned in
Chapter .
Another interspecies mix involves food stealing (piracy or kleptoparasitism) and is common among waterbirds. Gulls and terns commonly rob food
from other species as well as their own, and frigatebirds obtain much of their
food that way. A well-studied relationship between Black-headed Gulls stealing from Golden Plovers and Lapwings is common in wet meadows (Barnard
and Thompson ). Surface-feeding wigeon and Gadwall ducks often harass
diving coots as soon as they surface with herbaceous foods and retrieve the
dropped items (LeSchack and Hepp ).
7.11.2 nesting
During the breeding period, colonies of some species of gulls, ibises, egrets and
herons, and flamingos may contain thousands of nests with spacing dictated
by vegetation or other structural support and their territorial pecking distance. However, nesting coloniality does vary regionally, as many South
American blackbirds are less social that those in North America (Orians ),
herons, swallows, and blackbirds, and vegetated sites like water lilies or lotus
beds, flooded shrubs, or forest canopy for swimmers like Wood Ducks (Hein
and Haugen ; Martin and Haugen ). Other duck species may join
Wood Ducks in these roosts (Bellrose and Holm ). Open water is preferred
by Common Goldeneyes and other rafting sea ducks (Sayler and Afton ).
Cranes, geese, and Southern Screamers tend to use roosts as single species, but
mixes of species sometimes result by virtue of the habitat features and availability. Roost sites are an essential ingredient of the home range of such species
and may be related to the location of food, ideally spaced at a minimal distance.
In Sandhill Cranes, such roost sites are among major habitat features considered in management recommendations (Tacha, Nesbit and Vohs ).
Herons and egrets, blackbirds, swallows, and often gulls and dabbling ducks
commonly roost as mixed species. Protection from predators and location at
an energy-saving distance from foraging sites are among the obvious advantages of such roosts. There are hundreds of papers on the topic, including
many studies designed to resolve the problem these flocks may create for
people: aircraft strikes on landing fields, mixed flocks of blackbirds that move
into urban area and whitewash sidewalks and streets, and concentrations of
egrets and herons that kill trees through excess nutrient deposition on root
systems.
One of the more interesting and controversial ideas about roosts and other
flocking is the idea that members of a roost gain visual or vocal clues to the
location of food resources and seek those out as they depart the roost. This
theory was first applied to terrestrial granivorous finches of Africa (termed the
Information-Center Hypothesis), but some workers have concluded that
herons and egrets gain information and even follow successful birds to
daytime feeding areas (Krebs ). It also has been suggested that Cattle Egrets
that lag behind at roosts may be looking for signals rather than reflecting good
feeding the previous day (Siegfried ). Other species seem to leave in flocks
but disperse to different feeding sites; nevertheless, some white birds like
Snowy Egrets are conspicuous and are followed by others to feeding sites
(Smith ). In Barnacle Geese, post-roosting gatherings (i.e., pre-feeding
staging) were influenced by departure time from feeding areas the previous
day. Rather than being an indication of good feeding, investigators concluded
that it was organization to seek out new feeding areas because of poor results
earlier (Ydenberg and Prins ). Establishment of nesting colonies of some
seabirds seems to precede the obvious need for information on feeding but
does result in subsequent opportunities to do so (Buckley ). Clearly, these
patterns are important indicators of social organization; but detecting potential causes is a complex and difficult task.
In cold climates, winter roost sites often are selected to minimize loss of
body heat by use of sheltered areas and warm waters (Albright, Owen and Corr
, Brodsky and Weatherhead ). Birds also use a variety of body postures
and feature postures to minimize heat loss (Wooley and Owen ) and,
under extreme conditions, may stay in the roost rather than fly to foraging
areas, presumably because heat conservation is more important than food
intake for a short period (Jorde et al. ).
References
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the behavior and energy reserves of black ducks in Maine. Transactions of the
Northeast Section Wildlife Society , .
Anderson, M. G. and Titman, R, D. (). Spacing patterns. In Ecology and management of breeding waterfowl, eds. B. D. J. Batt, A. D. Afton, M. G. Anderson, C. D.
Ankney, D. H. Johnson, J. A. Kadlec, and G. L. Krapu, pp. . Minneapolis,
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Anderson, W. (). Waterfowl production in the vicinity of gull colonies. California
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Ball, I. J., Frost, P. G. H., Siegfried, W. R., and McKinney, F. (). Territories and
local movements of African black ducks. Wildfowl , .
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DC: Wildlife Management Institute.
Bellrose, F. C. and Holm, D. J. (). Ecology and management of the Wood Duck.
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Bennetts, R. E. and Dreitz, V. J. (). Possible use of wading birds as beaters by Snail
Kites, Boat-tailed Grackles, and Limpkins. Wilson Bulletin , .
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Loons at Storkerson Point, Alaska. Arctic , .
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nesting Forsters and black terns. Wilson Bulletin , .
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8
Spatial and structural patterns
Wetlands are especially known for their abundance of some species or groups
of birds as well as for their diversity of species not found elsewhere. The major
reason for this abundance presumably is the availability of resources produced
by the great primary productivity of wetlands. The diversity of species
undoubtedly is linked to diversity of resources produced in the many microhabitats resulting from water-depth gradients and fluctuations. The range of
spatial scales we can examine was outlined above, but I would like here to focus
on factors that influence bird use within and among wetlands. Although the
emphasis is on geographical, and physical or structural patterns in wetlands,
this includes structural aspects of live components such as plants (Table .).
Plants vary in life form or physical stature, which, with water distribution,
creates visual patterns useful to us as indicators of wetland types and conspicuous to birds as indicators of various essential resources. We will first consider
the range from homogeneity to heterogeneity that can exist in certain wetland
types, forming within-wetland variation of habitats (microhabitats), and then
consider larger-scale relationships among wetlands or wetland complexes or
regimes, which can best be appreciated by first understanding the patterns
found in individual wetlands.
Table 8.1 Wetland structural and physical features that influence habitat classification and
segregation of bird species and groups
Basin size
Basin shoreline configuration (development)
Basin substrate materials (clay, silt, sand, etc.)
Basin shoreline slope (rate of change)
Water depth and variability
Water clarity, salinity, icing, etc.
Water column structure (plant layering and distribution underwater)
Hydrologic influences: current, waves, and tides
River gradient and volume
Pools or openings in vegetation (size, dispersion)
Vegetation zonation and other distribution patterns
Coverwater ratios
Plant density
Plant height
Plant sturdiness or robustness
Plant aerial coverage based on life form: graminoid, herbaceous, woody, bush, tree, etc.
Vertical layering or strata
have evolved different preferences, which suggests how habitat diversity can
result in habitat sharing of different species in a small area (Weller and
Spatcher ). Such zones can change rapidly over time through catastrophic
events such as flooding or fire, which cause plant mortality and also create new
plant growth sites through soil exposure or sedimentation. Subsequent invasion occurs via underground rhizomes, mass seedings, or major germination
events from the seed bank. Even without catastrophic events, more gradual
biotic change is common. These dynamic patterns result in changes in availability of resources as well as in structure, potentially attracting different bird
species at different times.
The most aquatic plants of shallow wetlands are those that are submersed
but often rooted at some stage and are, therefore, associated either with
shallow waters or with the littoral zone of lakes or ponds where light penetration is greatest. These often have filiform leaves (e.g., milfoils and pondweeds)
that are able to capture light underwater more effectively. Some submersed
species may survive floating in the water column when they are torn free from
their roots; still other submergents are unrooted. Different species of submergent also are specialized for different depths and soil substrate and form
zonal patterns that vary in value to different bird species. Dense mats can
inhibit use by diving birds; others can walk on and hunt from such dense mats
of surfacing submergents.
Plants that float on the waters surface (floating plants, e.g., Lesser
Duckweed and Water-hyacinth) or occasionally in the water column (Star
Duckweed) and take their nutrients directly from the water via suspended
roots or the osmotic processes of individual cells are less restricted in distribution in a wetland when water is present. Larger species of floating vegetation
will support the weight of small walking birds like jacanas, Purple Gallinules,
or small egrets and bitterns, but such mats often inhibit swimming by larger
birds.
Often found in water of variable depths such as riverine backwater oxbows
and swamps, floating-leaf or pad plants (e.g., water-lilies) can flourish in
fluctuating or turbid water because they send up long stalks from often huge
and buried tubers (which are in themselves a valuable food for vertebrates).
There are intermediate plant growth strategies of course, and the Yellow Lotus
is one that may start its growth as a pad plant and mature as a robust emergent
as water support declines and the stalk strengthens.
Much if not most of the great productivity of wetlands comes in the form
of rooted herbaceous emergent plants, which live in the air/sunlight medium
that is ideal for photosynthesis but also can tap the nutrients of the basin soil.
Examples are cattail, sedges, bulrushes, rushes, spike-rushes, and the more
water-tolerant grasses such as cordgrasses in shallow water or damp soil.
Although they often grow by emerging from the water, woody plants like
scrub, shrubs, and trees are separated from herbaceous emergent plants in
classifications because of the dramatic difference in physical structure and
mass. In North America, they include water-tolerant shrubs like Buttonbush,
which germinate in shallow water or moist soil but survive as flooded plants
for many years. Other woody shrubs include alder and shrub willow which
occur along the shoreline slopes of basin wetlands and in stream riparian zone
sites where periodic water enables germination and survival. Similar scrub or
shrub species occur in slope wetlands such as the seeps from snow fields at high
altitude, even though there is rarely standing water. Several species of small tree
like willows are common in wet sites around the world, being very successful
mudflat pioneers. Many species of larger trees establish in wet sites of stream
riparian zones or floodplains that are seasonally flooded (cottonwood), nearly
continuously flooded (some oaks and gums), or continuously flooded (Baldcypress). Mangroves of many species dominate warm-climate tidal wetlands
around the world and vary from single shrubs to massive forests.
. Horizontal patterns
Wetland plant variety produces variation in habitat structure through species
differences in heights and density, and through patterns of plant establishment, competition, and interaction. In basin and fringe wetlands, such patterns reflect plant adaptation to different and often varying water depths in the
Horizontal patterns
. Vertical structure
Perhaps the most obvious structural habitat component that influences bird
use is vertical structure, first recognized by forest ecologists. Descriptive
measurements include not only height and number of vegetation layers, but
foliage volume and leaf form (MacArthur and Wilson ). Such factors are
equally important for waterbirds, regardless of what may seem like minor
height differences in emergent marshes; in addition, waterbirds confront
another layer, the water column, that terrestrial birds do not use. Moreover,
this is complicated by the fact that food resources may be at any height above
the water, float on the waters surface, be distributed at any level within the
water column, be mainly on the surface of the wetland bottom (e.g., epibenthic
. Wetland size
Some wetland diversity often is a product of size. It has been known for many
years that increasing the size of sample plots in plant communities results in
increased number of species (i.e., the speciesarea curve). Increased island size
results in increased number of vertebrates, including birds (Duebbert , He
and Legendre , MacArthur and Wilson ), and wetlands seem to function in the same way (Fig. .). Increased species richness with wetland size has
been reported for ducks in small prairie and forested wetlands (Nudds ),
forested bog wetlands in Maine (Gibbs et al. ), (Figs. . and .), diverse
waterbirds in lakes and smaller wetlands in Italy (Celada and Boglianai ),
and Finland (Lampolahti and Nuotio ), and estuarine tidal wetlands
(Craig and Beal ). Hirano and Higuchi () reported that broader riparian areas along large streams also demonstrated increased bird richness over
smaller streams in winter, mainly because of the extensive habitats for waders
produced by expanded bars and mudflats (Fig. .). The influence of river
Wetland size
width on dipper populations is less clear, but lower stream gradients seem to be
favored in some areas, inferring use of wider streams with more diverse habitats (Tyler and Ormerod ). Studies of streamside or riparian habitats suggests that broad areas hold more species of neotropical migrants than more
narrow ones (Hodges and Krementz , Keller, Robbins and Hatfield ),
resulting in the recommendation for a m minimal buffer zone, which is
especially important for these neotropical migrants.
Within-wetland vegetation heterogeneity clearly influences bird species
richness and the abundance of breeding birds in the vegetation of freshwater
marshes (Beecher , Kaminski and Prince , Weller and Spatcher ).
In one study of small tidal and nontidal wetlands (Craig and Beal ), the
influence of size exceeded that of heterogeneity for breeders, but other bird
users showed a greater but not major relationship. An important observation for conservation of species diversity was that larger wetland units were
most important in providing habitat for those species that occurred in low
density, a conclusion in agreement with observations by Burger et al. () for
tidal salt marsh, where diversity was less than in impoundments.
. Wetland diversity
Because many birds use several wetlands to accomplish goals like pairing or
rearing young, and because diversity of wetlands creates habitat diversity
Figure .. Relationship of
species richness to pond size for
ducks in small forested beaver
ponds of Eastern Canada compared with the more productive
prairie basin wetlands. Data
recorded by water hardness (a)
and invertebrate biomass (b).
(From Nudds , with permission.)
(a)
(b)
Wetland diversity
(a)
(b)
. Wetland configuration
Shoreline configuration is one of the more conspicuous features that would
seem to influence bird use, especially during the breeding season when aggressive behavior is prominent. Meaningful parameters of such edges are difficult
to relate to birds and other mobile vertebrates, but most workers use correlations with the shoreline index, a ratio of shoreline length to water area. Based
on studies such as those in Sweden, dabbling and bay ducks and coots show a
significant relationship to shoreline index (Fig. .), whereas Great-crested
Grebes do not (Nilsson ). Similar relationships have been observed by
other workers (see Kaminski and Weller ) and have been demonstrated
experimentally (Kaminski and Prince ).
Figure .. Relationship
between species richness of
wading birds and river width
(a) and extent of gravel areas
(b) ( are m2) (Hirano
and Higuchi , with permission).
Figure .. Relationship
between species richness and
wetland size for individual
wetlands and wetland complexes. (From Brown and
Dinsmore () with permission The Wildlife Society.)
Figure .. Relationship
between the density of breeding
ducks and shoreline development in some eutrophic
Swedish lakes (Nilsson ,
with permission).
Tree, shrub, bush, forb, and graminoid forms all influence whether the
foliage layer is high above the water or low, the robustness and obscurity produced by the size of leaves, stems, stalks, and trunks, and the dispersion and
density of vegetation in relation to water (semiopen or densely vegetated).
These factors influence the birds ability to walk or swim through the vegetation for food or cover, to build nests in it, to sun, or to see its neighbors. These
are difficult parameters to identify and measure, and subsequently a statistical
correlation of bird use with specific vegetation characters is necessary.
Moreover, these parameters are often highly variable by time and place. This
type of research is becoming more common, and each attempt identifies some
Table . Wetland habitat patterns that serve as descriptors of wetland types and habitats for bird assemblages
Deep, open-water
Submerged aquatic beds
Floating vegetation and pad plants
Robust and persistent herbaceous emergents (graminoids)
Nonpersistent, meadow-like emergents
Bog mats and organic islands
Shrubs
Forested wetlands
Unvegetated shallow water and mudflats
Tidal flats and salt marsh shores
Riverine sandbars, islands, and depositional areas
Boulders, rock substrate and other hard-shores
Constructed pond, lake, and reservoir
Riverine shore, substrate, and pools
periods, long-legged waders feed there, such as Black-necked Stilts and avocets
in saline areas, and dowitchers, yellowlegs, and sometimes ibises and egrets if
large invertebrate foods are present in temperate areas. Grazing waterfowl
such as White-fronted Geese, Brant, or American Wigeon forage on new
shoots resulting from spring flooding of meadow or tundra vegetation (Fig.
.). Seed-eating finches, plovers, and shorebirds also feed there and may nest
in the developing cover as the water recedes.
Birds that prefer persistent emergent plants and shorter and less robust
nonpersistent vegetation also are conspicuous at the shallow edges of wetlands
or the perimeter of large islands within a wetland. These areas tend to be dominated by walkers rather than swimmers, including King and Clapper Rails,
American Bitterns, Snowy Egrets, and Little Blue Herons. Red-winged
Blackbirds prefer the edges and are less abundant in deeper water. Some
species spend as much time in the adjacent uplands as at the wetland edge, such
as aerial foragers like Willow Flycatchers and water or ground feeders like
Swamp Sparrows and Northern Harrier (McCabe , Weller and Spatcher
). Small trees and shrubs on unflooded upland sites are used for song
perches or nest sites. Tree Swallows and Eastern Kingbirds may nest in trees
over water near the shore. Other species find concentrations of important
foods like insects, crayfish, and amphibians in this moist interface.
8.7.6 birds of bog mats and organic islands
Buoyant organic and living vegetation forms mats where other vegetation
becomes established. Moss, sedge and grassy tussock mats create areas of low
life form in forests; these tend to become higher and drier over time and form
sites suitable for shrubs and even trees. But these areas change slowly and
provide different habitats for birds that are characteristic of boreal areas and
often have greater species richness than surrounding forested habitats
(Damman and French ). Vegetation structure and diversity most affect
bird species richness, and species use is influenced by latitude and tree
composition. Taxa represented include several ducks common mainly to those
areas and a variety of songbirds (Fig. .), as well as one unique shorebird, the
Wilsons Snipe (Tuck ). Bogs and peat mats also are common in south-
Figure .. Distribution
patterns of lakeshore bogmat wetlands and adjacent
shrub and forest birds in
two northeastern North
American forest zones: (a)
boreal and coastal sprucefir
zone; (b) Appalachian oak
zone (Damman and French
).
V EGETATION LAYERS
R ED - BELLIED W OODPECKER
R ED - EYED V IREO
P ROTHONOTARY
W ARBLERS
Y ELLOW- BELLIED
S APSUCKER
2
S PARROWS /W OOD DUCK /M ALLARDS /G REEN - WINGED T EAL /W HITE I BIS
50
100
150
200
250
eastern slope and basin wetlands where plant production exceeds decomposition partly because of saturation and reduced oxygen; these areas are best
known for their pitcher plants. Glade areas in Georgia and Florida have even
more extensive mats of grasses and other herbaceous plants that are used for
nesting, feeding, and resting by rails, coots, and other walkers and by some
songbirds, which feed on the mat or from aerial perches. Forest communities
in the tropics also have bog areas, but bird species seem to be less specific to
those sites, perhaps because of small size.
8.7.7 birds of shrub and forested wetlands
Wetland shrubs occur around a variety of wetland types in the seasonally flooded
zone or in saturated soils in riparian or alpine habitats. Streamsides or drainages,
where wetland vegetation is apparent or where terrestrial vegetation is more
abundant and diverse owing to the presence of water, are well known as important riparian areas for birds (Hehnke and Stone , Taylor , Wauer ).
Tree- or shrub-nesting passerines and flycatchers are prominent there because of
the availability of foods from the stream and from the several layers of vegetation.
Water-induced shrubs also occur as extensive stands or patches in coldtemperate climates at high altitude or latitude around the world. Wild and
domesticated herbivores browse these shrubs, and they are important habitats
for Willow Flycatchers, Yellow Warblers, and numerous other passerines that
exploit insect life of wetland edges or patches. More water-tolerant shrubs like
Buttonbush and certain species of willow and alder can survive flooded conditions even during the growing season and are among the plant dominants in
areas termed shrubscrub swamps (Fig. .). Such habitats often are used by
herons and egrets as well as by passerines for nest sites. These plant zones commonly are distributed along stream banks, especially in western areas where
extensive vegetation is present only because of the increased water table associated with streams.
Forested wetlands include bottomland hardwoods that are seasonally
flooded during the winter dormant period. These are characteristic of backwaters of floodplains along major southeastern river systems, or of naturally
impounded and trapped waters that are semipermanent and form true
swamps. Birds of larger wetland trees commonly include not only the herons,
which may nest at the tops of dead trees in or near water, but also woodpeckers,
which make and use cavities. Large cavities in large trees may later be used by
Wood Ducks and Hooded Mergansers, whereas Prothonotary Warblers and
Tree Swallows use smaller holes in old snags. Such dead trees and snags must be
preserved if the species are to be successful in the area, but all these species will
respond to artificial nest boxes, where the expense and effort is justified.
Understory plants like plume grasses and cane or bamboo form dense habitats
used by Swainsons Warbler and Bachmans Warbler (which is probably now
extinct); this is now a rare microhabitat because of the intense use of forested
areas. Open areas are used by sparrows and wrens favoring moist and seed-rich
habitats with lower herbaceous cover. As can be seen, diversity of birds here is
strongly influenced by forest diversity and successional stage (Buffington et al.
, Swift, Larson and DeGraaf .)
8.7.8 birds of unvegetated shallow water, mudflats,
and sandbars
Shallow sheetwater over soil, sand, or other nonvegetated or sparsely vegetated
substrate may result from rainfall, stream, or tidal flooding of mudflats and
shorelines. Plowed fields and other agricultural areas also may be flooded
under such circumstances, which often makes available a rich supply of foods.
Open-water areas within wetlands form similar new habitats for edge birds
when water levels decline below the emergent and submergent zones and
expose bare basin bottoms. These offer special habitats for American Avocets,
ibis and teal, which specialize in feeding in shallow water, usually by feel or
straining rather than sight. Waders often walk such flats in search of fish,
amphibians, reptiles, or large invertebrates in the water. Such sheetwater areas
may be used also during migration or in wintering areas as overnight roosts by
cranes and geese; protection of these habitats is a vital management approach
to capitalizing on waste grain and other adjacent food resources attractive to
such granivores (Hobaugh ).
Mudflats are moist and usually unvegetated areas generally not appreciated
by those who do not recognize the abundance and distribution of invertebrates that occur in these habitats and the role the invertebrates play as detritivores in the cycling of nutrients within wetland systems. Thus, the mudflats
offer a unique habitat for carnivorous foragers like Killdeers and numerous
other plovers and sandpipers (Capen and Low ). More fluid mud is the
favorite habitat for a number of groups of tiny crustaceans, nematodes and
annelids, which are eaten especially by Green-winged Teal, a species often
termed mudder for its walking and sifting behavior in soupy mud, and by
shorebirds.
8.7.9 tidal flats
Tidal flats and shores differ from mudflats in basin wetlands because of water
hydraulics, forming areas that are flooded periodically by celestial or wind
tides. Therefore, some areas are available for bird use for some types of feeding
only part of the time because of fluctuations in water level. Moreover, when
nesting occurs in these areas, invasion by ground predators may be possible.
These sites also differ in soil texture, varying from clay and silt to sand and shell
deposits, which influences water drainage rates and benthic organisms and
therefore, the bird species that use the habitat. Sandy or gravel (shingle) shores
are common in specific areas because particle separation is a product of wave
and current action on suspended or substrate materials. Therefore, mudflats
commonly occur near river mouths in estuaries, with larger sand particles
being deposited and regularly moved by active surf or tidal action (Britton and
Morton ). Wind also plays a role in distribution of sand and influences
resting sites, drinking areas, and even endangers bird nests. Birds that use
sandy shores regularly include the well-named Sanderlings, several peeps, Red
Knots, Ruddy Turnstones, and oystercatchers for feeding on clams and other
burrowing invertebrates, and various gulls, terns, and skimmers for resting
and nesting (Burger et al. , Whitlack ). Zonation of bird use is a
common product of water depth, wave forces, and food distribution
(Whitlack ), and similar diversity occurs in salt marshes and other tidal
plant communities (Daiber ) that develop in more stablized depositional
areas. Many species forage microsites in tidal areas based on consistency of the
substrate and move seaward with the ebbing and shoreward with the rising
tide.
8.7.10 emergent salt marsh
Herbaceous saltmarsh vegetation is common on stabilized sediments where
seeds or rhizomes persist, forming parallel zonation similar to basin wetlands.
This grades into shrubs in some areas, but this zone may be dominated by
mangrove trees in tropical waters. This diversity of vertical structure creates
foraging and nesting sites attractive to various bird species and functions as a
segregating mechanism (Fig. .). During breeding, some unique saltwater
species are found there, such as the Clapper Rail and Seaside Sparrow, but also
ubiquitous species like Red-winged Blackbird and Least Bittern. Wintering
species include many freshwater, alkaline water, and wetland-edge birds from
interior breeding areas (Sharp-tailed Sparrows, avocets, Willets, coots, and
Northern Shovelers).
8.7.11 riverine pools, substrates and shores
In their linearity and gradual increase from seasonal trickles of water to raging
torrent and eventually to wide and permanent water at lower levels near their
terminus in lake or sea, streams and rivers provide diverse shoreline
configuration, vegetation, substrate and water characteristics, and water
sources that are tapped by a great variety of birds. However, some stretches are
nearly inhospitable at times because of the current while other stretches are
characterized by flooding regimes that make them unpredictable. In higher
areas, dippers and other passerines exploit adjacent vegetation and foods in
and out of the water. Pools and backwater areas that hold fish and herps are
favored by wait-feeders like Green Herons and night-herons, and pool and
riffle areas are used by mergansers and other stream ducks mentioned above.
Larger and slower streams have greater habitat diversity,which attracts a wider
range of birds. River meanders are noted for moving basin materials of all sizes
during high water periods and depositing them by size categories according to
current and basin resistence. Thus, rivers cut against high ground and create
vertical soil and sand banks suitable for kingfisher burrows, swallows, and
other waterside birds. Depositional flats are formed on the inner courses,
resulting in shallow water gradients suitable for waders and drier vegetated
areas suitable for songbirds. Sandbars, islands and peninsulas are ideal for
nesting by early succession species like Least Tern, Piping Plover, and Killdeer
even when sparsely vegetated or bare. As these areas age, plant succession
moves toward annual herbaceous vegetation and then to perennial plants,
forming good areas for ground birds. Those areas grown up in shrubs, willows,
and cottonwoods may form roosting areas for migrants, nest sites for colonynesting egrets and herons, and perches for scavengers like eagles or piscivores
like Ospreys.
8.7.12 birds of boulder, rock substrate,
and other hardshores
Marine and large lakes can have large rocks and boulders deposited by forceful
waves or flooded rock strata where aquatic organisms (plant or animal) occur;
these areas attract birds that are specialists at probing and freeing attached
foods. Constructed jetties, levees, and other stone or block structures simulate
such natural sites and form substrates for invertebrates that attract birds.
Many of the birds using rocky areas are strictly coastal, and they tend to be
invertivores (especially limpet and mussel eaters), like Rock and Purple
Sandpipers, Surfbirds, turnstones, oystercatchers, Harlequin Ducks, and
eiders. Herbivorous birds that feed in these areas include Black Brant or their
ecological counterpart in Tierra del Fuego, Kelp Geese, which use leafy sealettuce.
8.7.13 birds of constructed ponds, lakes, and reservoirs
Typically, constructed waters are impounded waters that are deep enough that
emergent vegetation is either absent or restricted to a narrow fringe, and they
sometimes have extensive submerged plant beds in the shore zone that are
valuable for waterbirds. These artificial lakes may be important foraging sites
for birds that feed on insects over the water, or for swimming birds that eat submergent foliage or resident animals; as a result, they add avian diversity especially in arid regions (Evans and Kerbs ). Larger reservoirs may have
extensive littoral zones in the upstream area, which may be excellent waterbird
habitat if they are not subject to rapid and extreme water fluctuations.
However, wave action often seriously limits shoreline plant establishment.
Because of the abundance of fish, many are excellent areas for cormorants, pelicans, herons, and mergansers. Geese and similar grazers around the world use
them as rearing areas in summer because they feed in the uplands and use the
water for brood protection. Large reservoirs also may be favored by wintering
flocks of Canada Geese and Mallards, because they are deep and remain open
during most winters and the birds can feed in grain fields nearby (Simpson
).
8.7.14 summary
The habitat patterns described above are based on observed associations;
although many have been quantified at study sites, fewer have been related to
vegetation composition and structure by statisical tests. Patterns vary regionally, but they form a starting point for grouping microhabitats in many types of
wetland and in wetland regions that should serve well in descriptive studies.
References
Bartonek, J. C. and Hickey, J. J. (). Food habits of canvasbacks, redheads, and
lesser scaup in Manitoba. Condor , .
Beecher, W. J. (). Nesting birds and the vegetative substrate. Chicago, IL: Chicago
Ornithological Society.
Bergman, R. D. (). Use of southern boreal lakes by postbreeding canvasbacks
and redheads. Journal of Wildlife Management , .
Britton, J. C. and Morton, B. (). Shore ecology of the Gulf of Mexico. Austin,
TX: University of Texas Press.
Brown, M. and Dinsmore, J. J. (). Implications of marsh size and isolation
for marsh management. Journal of Wildlife Management , .
Brown, M. and Dinsmore, J. J. (). Area-dependent changes in bird densities
in Iowa marshes. Journal of the Iowa Academy of Sciences , .
Buffington, J. M., Kilgo, J. C., Sargent, R. A., Miller, K. V., and Chapman, B. R. ().
Comparison of breeding bird communities in bottomland forests of different
successional stage. Wilson Bulletin , .
Burger, J. (). Habitat selection in temperate marsh-nesting birds. In Habitat
selection in birds, ed. M. L. Cody, pp. . Orlando, FL: Academic Press.
Burger, J., Howe, M. A., Hahn, D. C., and Chase, J. (). Effects of tide cycles on
habitat selection and habitat partitioning by migrating shorebirds. Auk ,
.
Burger, J., Shisler, J., and Lesser, F. H. (). Avian utilisation on six salt marshes
in New Jersey. Biological Conservation , .
Capen, D. E. and Low, J. B. (). Management considerations for nongame birds in
western wetlands. In Workshop proceedings: Management of western forests and
grasslands for nongame birds, pp. . USDA Forest Service General Technical
Report INT- . Washington DC: US Forestry Service.
Celada, C. and Bogliani, G. (). Breeding bird communities in fragmented wetlands. Bolettino Zoologia , .
Colwell, M. A. and Dodd, S. L. (). Environmental and habitat correlates of
pasture feeding by nonbreeding shorebirds. Condor , .
Cowardin, L. M., Carter, V., Golet, F. C., and LaRoe, E. T. (). Classification of wetlands and deepwater habitats of the United States. FWS/OBS-/. Washington,
DC: US Fish and Wildlife Service.
Craig, R. J. and Beal. K. G. (). The influence of habitat variables on marsh bird
communities of the Connecticut River estuary. Wilson Bulletin , .
Daiber, F. C. (). Animals of the tidal marsh. New York: Van Nostrand Reinhold.
Damman, A. W. H. and T. W. French. (). The ecology of peat bogs of the glaciated
References
Keller, C. M., Robbins, C. S., and Hatfield, J. S. (). Avian communities in riparian
forests of different widths in Maryland and Delaware. Wetlands , .
Lampolahti, J. and Nuotio, K. (). Umpeenkasvu koyhdyttaa lintuvesia (with
English summary). Linnut , .
MacArthur, R. H. and Wilson, E. O. (). The theory of island biogeography.
Princeton, NJ: Princeton University Press.
McCabe, R. A. (). The little green bird: ecology of the willow flycatcher. Madison,
WI: Rusty Rock Press.
Murkin, H. R., Kaminski, R. M., and Titman, R. D. (). Responses by dabbling
ducks and aquatic invertebrates to an experimentally manipulated cattail marsh.
Canadian Journal of Zoology , .
Murkin, E. J., Murkin, H. R., and Titman, R. D. (). Nectonic invertebrate abundance and distribution at the emergent vegetation-open water interface in the
Delta Marsh, Manitoba, Canada. Wetlands , .
Nilsson, L. (). Breeding waterfowl in eutrophicated lakes in south Sweden.
Wildfowl , .
Nudds, T. D. (). Patterns in breeding waterfowl communities. In Ecology and
management of breeding waterfowl, eds. B. D. J. Batt, A. D. Afton, M. G. Anderson,
C. D. Ankney, D. H. Johnson, J. A. Kadlec, & G. L. Krapu, pp. . Minneapolis,
MN: University of Minnesota Press.
Orians, G. H. (). Some adaptations of marsh-nesting blackbirds. Princeton, NJ:
Princeton University Press.
Patterson, J. H. (). The role of environmental heterogeneity in the regulation of
duck populations. Journal of Wildlife Management , .
Simpson, S. G. (). Use of the Missouri River in South Dakota by Canada Geese
in fall and winter, . In Waterfowl in winter, ed. M. W. Weller , pp. .
Minneapolis, MN: University of Minnesota Press.
Skagen, S. K. and Knopf, F. L. (). Migrating shorebirds and habitat dynamics
at a prairie wetland complex. Wilson Bulletin , .
Spence, D. H. N. (). The zonation of plants in freshwater lakes. Advances in
Ecological Research , .
Swanson, G. A., and Meyer, M. L. (). Impact of fluctuating water levels on feeding
ecology of breeding blue-winged teal. Journal of Wildlife Management , .
Swift, B. L., Larson, J. S., and DeGraaf, R. M. (). Relationship of breeding bird
density and diversity to habitat variables in forested wetlands. Wilson Bulletin ,
.
Taylor, D. M. (). Effects of cattle grazing on passerine birds nesting in riparian
habitat. Journal of Range Management , .
Tuck, L. M. (). The Snipes: a study of the genus Capella. Monograph Series No. .
Otttawa: Canadian Wildlife Service.
Tyler, S. J. and Ormerod, S. J. (). The dippers. London: T & A D Poyser.
Wauer, R. H. (). Significance of Rio Grande riparian systems upon the avifauna.
In Importance, preservation, and management of riparian habitat: a symposium,
Technical Coordinators R. R. Johnson & D. A. Jones, pp. . General
Technical Report RM-. Washington, DC: US Forest Service.
Weller, M. W. (). Birds of some Iowa wetlands in relation to concepts of faunal
preservation. Proceeding of the Iowa Academy of Science , .
References
9
Habitat dynamics:
water, plant succession, and time
. Water variability
The dynamics of water over time, whether seasonal, annual, or longer term,
dictates the chemical and physical character of wetland water, the resulting
vegetation, and the use of wetlands by birds and other aquatic or semiaquatic
life. Numerous authors have summarized the consequences of variable water
regimes on vegetation, plant succession, and size and depth of various types of
wetlands (Bellrose, Paveglio and Steffeck , Chabreck , Golet and
Parkhurst , Gosselink , Kantrud, Krapu and Swanson , Kushlan
, Stanley, Fisher and Grimm , Weller and Fredrickson ). Here we
examine the general patterns of water influences on vegetation and the physical aspects of the wetland habitat that influence bird use. Examples also are
given that show the similarities of these patterns worldwide.
. Plant succession
A wetland plant community may be established or re-established when suitable water conditions and biochemical properties of the soil induce germination of seeds or growth of plant propagules (e.g., tubers and rhizomes) of
hydrophytes (van der Valk ). The viability, mobility, and response of seeds
to suitable germination conditions have been well documented (Crocker ,
van der Valk and Davis ). This is perhaps best demonstrated in small
wetland basins or along fringing shorelines of larger wetlands where gradually
deepening water depth determines which species germinate, spread, and
survive according to their individual adaptations for particular water and soil
conditions. As discussed earlier, the structure or life form of various species
tends to vary with water depth, but this is strongly influenced by the taxonomic
groups dominant in the locale. Such plant zones also are clear-cut in the submerged vegetation of shallow lakes, where it influences accretion rates and
increases lake eutrophication (Carpenter ), and in shallow wetlands, where
competition as well as physiological responses to conditions at those sites may
influence plant species competition and success (van der Valk and Welling
, Wilson and Keddy ).
Coastal estuarine and salt marsh wetland habitats have similar patterns of
zonation, which are induced and influenced by daily and seasonal variability in
tidal regimes (Allison , Niering and Warren , Penfound and Hathaway
). Drying in higher zones increases salt concentration, which may reduce
plant growth or influence which species can survive; however, in lower tidal
areas, freshwater inflows may dramatically change plant survival and species
composition (Allison ). Wave action influences plant success in deeper
zones, and as in all plant communities, competition may influence the plant
species that are present and their success in addition to the effects of water
regimes (Pennings and Callaway ).
The resulting vegetation structure along shorelines is an important one for
foraging and nesting birds, and for those seeking shelter and protective cover.
The vegetation can also be important because of differences in the potential
Habitat dynamics
foods produced, e.g., annuals tend to produce more seeds and perennial
species tend to produce more tubers or rhizomes (Chapter ). As a result,
shoreline gradients that induce zonation of structure and food resources
provide diverse resources to which birds respond dramatically (Allen ,
Beecher , Weller , Weller and Spatcher ), resulting in segregation
through habitat selection by various bird species. But keep in mind that what is
an attractive structure for some bird species will discourage others, for
example those that favor bare mudflats (shorebirds) or short, flooded vegetation (yellowlegs and certain teal). Presumably, these structural patterns are
signals to bird species about the habitat resources.
The distribution of vegetation (i.e., pattern) in more central areas of a
wetland are especially relevant to swimming birds and others that favor more
open water, like the large waders. The development of such patterns are
complex because they represent historical seeding or other short-term plant
establishment events that themselves reflect various rates of survival dependent on water regimes, and wave and ice action.
Clearly, patterns of plant distribution are not fixed but change over time
based on the wetland type, water regimes, and plant life-history strategies. To
appreciate the role of such changes for birds, conceptual models of biotic
change have been useful (Begon, Harper and Townsend ), and two have
been especially relevant to wetlands (van der Valk and Davis ). These differ
mainly in temporal scale but also in their assessment of the variability of individual sites and the influence of chance events on the long-term history of
vegetation at a site.
The traditional view of long-term changes is based mainly on work by
Clements (and, therefore, termed Clementsian succession) and suggests that
water-driven plant (hydrarch) succession proceeds from wet to dry in a series
of stages (called seres). It is considered very long-term (hundreds to thousands of years), directional, and fairly predictable. Examples of long-term
succession in northern bogs (where water is more stable) have been based on
analysis of pollen or planktonic fossils, which show changes over tens of thousands of years. At this scale, models of changes in bird species over time have
been based on the synthesis of observations on concurrently observed examples of various seres (Aldrich ). But even these habitat areas are not stable
and often experience reversals reflecting long-term climate change or other
variables like salinity and sedimentation. Lake succession has been viewed in
this long-term scale, reflecting eutrophication via sedimentation and plant
organic deposits within the basin; submergent aquatic plants are especially
important in sediment trapping (Carpenter , Robel ). These plants
influence use by aquatic birds and even serve as indicators of these lake stages
(Kauppinen and Vaisanen ). Ultimately, lakes may fill in with sediments
and bog plant growth, creating sites for pioneering shrubs and trees (Fig. .).
Other plant ecologists, especially Gleason (hence, Gleasonian succession),
felt that succession was much more influenced by specific site conditions and
Plant succession
by chance environmental events, which result in differing patterns of vegetation establishment even within the same area or site (van der Valk ).
Wetland basins that dry periodically present random opportunities for
different seeds to germinate, and hence can, by chance, result in strikingly
different vegetation in the same basin over a series of years. A series of observations by different workers studying basin wetlands demonstrated dramatic
short-term changes in plant succession induced by drawdowns (also termed
drydowns in some areas) caused by drought (or water-level manipulations for
management purposes); these resulted in ideal germination conditions from
the buried seed bank or residual rhizomes (Harris and Marshall ) (Fig.
.). Animal responses to these patterns also were noted over several years
(Kadlec , Meeks , Weller and Spatcher ). But there are also records
Habitat dynamics
of this same type of semipermanent wetland that seems to have had the same
vegetation for thousands of years. Many invertebrates also have residual and
drought-tolerant propagules or egg banks that allow a quick response and
recolonization when suitable water conditions occur. Others must fly in and
lay eggs (dragonflies), as is true of birds.
Increases or decreases in water depth, changes in hydroperiod (the duration of flooding), or water characteristics such as turbidity influence the
success of plants established during low water. These form a somewhat predictable pattern that varies with water, timing, and herbivory (Fig. .).
The plant reproductive strategies discussed earlier are changed in these
ways and some species decline while others increase over several growing
seasons. Submergent plants are very dependent on water depth and clarity,
and modest changes can affect their success. In both fresh- and saltwater wet-
Plant succession
Habitat dynamics
birds (e.g., stilts, sandpipers, and plovers), dabbling ducks, egrets, herons,
storks, ibises, and shoreline passerines. All are tapping foods that are available,
but each group favors different foods suitable to diverse feeding tactics.
Shorebirds and passerines may be competing for crustaceans and insects
exposed by or attracted to the mudflat or vegetation. Teal may be straining on
the mudflats with the probing or pecking shorebirds but taking still smaller
prey. The walking fish feeders like egrets, ibises, and storks (and sometimes
large Icterids like grackles) are exploiting fish or herptiles concentrated in
drying pools. Birds that favor feeding in vegetation tend to be reduced (rails,
bitterns, and some sparrows and ducks).
In situations when wetlands dry completely, vegetation invasion influences
use, and wetland birds are more likely to be replaced by species that prefer
damp sites (plovers) and upland shorelines (doves, passerines). When water
levels rise beyond the basin shoreline, flooding shoreline vegetation and adjacent meadows and open fields, many species find a diversity of foods available,
which they may pursue as flocks even though they are more typically solitary
birds. These include some some larger shorebirds like stilts and curlews, ibises,
storks, dabbling ducks, egrets, and small herons.
Habitat dynamics
Figure .. Generalized
diagram of common seasonal
water patterns of various
wetland types and areas in
North America as they
influence water availability and
vegetation for migratory birds.
Seasonality of responses
herptiles, or fish typical of larger and more open pools of water. Large waters
like lakes and reservoirs may have few birds, and those will be ones dependent
upon large fish (gulls, loons, cormorants, and Ospreys).
In any given season, birds will respond to water conditions initially, probably more than to vegetation; residual plants are more important to perching
birds establishing a breeding territory than to swimming birds searching for
food and for loafing sites. If typical water regimes do not exist because of the
extremes of drought or heavy rains, nesting may be delayed (Custer, Hines and
Custer , Weller and Spatcher ), may not occur (Breeden and Breeden
, Frith , Rogers ), or some species may move to where conditions
are more suitable to their preferences (supported by the re-capture of banded
ducks and observations of conspicuous colonial species like storks and egrets).
Therefore, populations may decline (Weller, Wingfield and Low ) and the
species composition of nesting birds may change to those characteristic of the
new water regime (Weller ).
Why do few wintering birds stay and nest in southern wetlands rather than
engage in hazardous migration twice a year? If we examine the hypothetical
pattern of deep marsh and southern basins or bottomland wetlands (Fig. .),
it is obvious that birds that do not move from southern to northern breeding
marshes may face drying wetlands too early to rear young.A few species associated with coastal tidal waters (e.g., Black-necked Stilts and Willets) are more
likely to be successful. These typical patterns can be changed drastically by
heavy late summer or autumn rains resulting from hurricane or tropical
storms, and sufficient water may carry over to keep wetlands flooded for
several years. In that situation, many species may remain and breed, although
the numbers probably are modest compared with the total migrant population. Even then, most birds are more likely to find the essential requirements
for breeding by moving from areas of few wetlands to one of more numerous
and diverse wetlands, where some wetlands within a complex of various-sized
basins are likely to be wet.
In autumn, southern basins may be near dry and unsuitable for autumn
migrants especially those birds that require specific water conditions such as
mudflats for shorebirds or flooded areas for birds that feed by swimming.
Migrants then move through quickly on northern storm fronts, and little
wintering would be expected. If flooded by autumn rains, these areas are suitable for late migrants and presumably are more heavily used in those wet
winters.
Seasonality of nest initiation seems to be influenced by pre-migration and
pre-nesting resources but are modified by temperature and other stressors.
Winter body condition seems to influence timing of migration, readiness for
pairing and laying eggs, and survival during this more stressful period
(Heitmeyer and Fredrickson ). These activities are mostly a product of
wetland habitat conditions, including presence and timing of open water
(thawing at high latitudes) and availability of suitable foods and nest sites. For
Habitat dynamics
any given region, data are available that show the general chronology (i.e.,
timing) of nest initiation, and this presumably reflects those long-term patterns of pre-migratory preparation, migration, finding suitable nesting
cover/sites, and pair formation, where that is done on breeding areas. But if the
phenology (climatically influenced events) differs, and the season is late(i.e.,
cold and wet) or early, nesting often is delayed or advanced accordingly in
many species. Habitat suitability clearly plays a major role in regulating these
events.
Pothole Region are impressive, with the highest populations recorded in many
years (Caithamer and Dubovsky ).
9.6.2 south texas coastal ponds
South Texas is known as a dry place even along the coast. The long-term mean
rainfall is, however, about cm ( inches) per year, coming mainly in the
periodic deluges that result from hurricanes and similar tropical summer
storms. Between these flood years, it is very dry. Even most knowledgeable naturalists would be surprised to find that there are numerous wetlands there,
sometimes termed hurricane lakes, because they are not there all the time.
Even during relatively dry periods, they are favored grazing areas for livestock
and deer because they catch and hold whatever rain falls and stimulates winter
plant growth. But when the rains come, or even more inches (cm) fall in
hours, and the rains will come repeatedly over a few weeks or months, the area
rivals the Prairie Pothole Region of the northern USA and Canada. Terrestrial
grasses are no more. Sedges and other moist-soil plants seem to spring up, and
even a small Blue Water-lily seems to explode in shallow, freshwater wetlands
after drought and re-flooding regardless of whether grazing has occurred in
previous years.Wintering waterfowl increase in the area, driven by the need for
sources of fresh water after feeding on saline seagrasses. Ducks such as Bluewinged Teal, Northern Shovelers, and Ruddy Ducks and American Coots,
Moorhens, and rails settle in and nest. Some wetland mapping systems may
not regard these as wetlands because of the long interval between hydric plant
communities, but clearly they have the residual seed banks to respond when
water is present.
9.6.3 larger freshwater basin wetlands
Several of our studies of wetlands (Weller and Fredrickson , Weller and
Spatcher ) of to acres ( to ha) in central and northwest Iowa
demonstrated the impact of vegetation change on species composition and
abundance of wetland bird assemblages. Species richness was greatest when a
: up to : ratio of cover:water existed, as was abundance of most birds
that favored robust emergents. Species such as Black Terns and Red-winged
Blackbirds that are attracted to shallow and dense marsh vegetation lost out to
Forsters Terns and Yellow-headed Blackbirds under the changes to deeper
water and more robust emergent plants.
In still larger basins of thousands of acres, I had an opportunity to spend
some time observing vegetation and birds of the managed wetland called Mud
Lake on the Agassiz National Wildlife Refuge in northern Minnesota, where
Harris and Marshall () developed one of the earliest models of seed-based
succession. They recommend that drawdowns be done periodically as needed
to maintain the variety and dispersions of emergents needed to attract nesting
Habitat dynamics
Habitat dynamics
season. The flooding may have been caused by water from several years of high
rainfall and high creek levels and a small beaver dam. After the kill, the herbaceous vegetation flourished as it cannot do under the dark canopy and
included typical wetland shallow-water or edge species, such as annual smartweeds, and seeded perennials such as grasses, sedges, and rushes.
Concurrently, Water Elms, a deeper-water scrub species about feet (m) in
height, found an ideal seed bed for its annual crop, and seedlings developed
widely. These seedling rapidly became dominants that eventually will shade
out the herbaceous species. The result is that the deeper-water vegetation zone
has moved up slope as well as the birds (Fig. .).
9.6.8 stream-side or riparian wetlands
An example of stream modifications such as dam construction exists in northern Iowa where we followed vegetation periodically for nearly years (Weller,
Kaufmann and Vohs ). Damming upstream has had two major effects.
First, pools formed behind several dams became rather sterile lakes with few
marginal emergents and relatively few aquatic submergents. However, use by
bay diving ducks and geese increased. Second, reduced downstream flow
modified vegetation of temporary pools from diverse mudflat and marsh-edge
annual plants to more perennial grasses characteristic of more shallow and
less-flooded stream banks. Reversion is possible with time and return to traditional small-stream water regimes.
On a larger scale, the Platte River of Nebraska has been impacted by
upstream dams in Colorado and Nebraska build for urban water supplies and
agricultural irrigation, with the result that a traditional steady and shallow
flow has been replaced by a reduced and less predictable flow. As a result,
vegetation along the river and on sand and gravel islands has changed drastically toward drier streamside and woody types. Sparse herbaceous cover on
islands has given way to willows and cottonwoods, completely changing the
character of the wildlife habitat. As a result, shallow-water roosting islands are
dry, vegetated, and unattractive to the Sandhill Cranes during early spring
migration periods. Flooding of backwater and other marginal wetlands also
has been reduced, thus drying favored feeding areas of cranes on tuber-producing sedges. During summer, the islands are exposed but are no longer
habitat for Piping Plovers and Least Terns, which need open, sandy islands.
Only restoration of the stream flow will allow potential recovery, and the area
will then require management such as cutting of trees and brush, which is
costly.
9.6.9 tidal estuarine wetlands
Many people have concluded that coastal estuarine systems show less variability in plant communities on short time scales and are slower to change on long
Habitat dynamics
time scales because sea levels change slowly. But where tidal fluctuations are
low, as in the Gulf of Mexico, estuarine wetlands experience dynamic water
regimes, often as a product of the amount of freshwater inflow and local rainfall, and they can have a major impact on vegetation. The brackish water
characteristic of estuary wetlands is a mix of freshwater and saltwater. The
heavier saltwater sinks and is overridden by the lighter freshwater in a pattern
termed the saltwater wedge. But water often enters the estuary as sheet flow
and percolation everywhere along the coast, and blending of fresh- and saltwater is gradual. Where freshwater volume is high and continuous, the salinity
is low. A reduction in rainfall or a major storm surge from the sea changes this
ratio, and plants may be killed from the sudden high salinity, creating a
germination site not unlike that of shallow, freshwater basin wetlands.
Seasonal variation in salinity and water depths (tide height) occurs as a
result of shifts in lunar/sun gravitational pull, which creates different tide
levels at different seasons (e.g., spring tide). Higher tidal areas well flooded
early in the year often dry out in summer, and obviously their salt concentration increases unless there is continuing freshwater inflow. That rainfall
pattern varies by area of the country, and from year to year. Therefore, the
more saline areas along the coast are not the areas regularly flooded by saltwater (which normally is less than p.p.t.) but the irregularly flooded and
seasonally drying areas that form salt flats, where concentrations commonly
exceed p.p.t. This is an important influence on plant succession, and has
important implications for restoration of wetlands lost along the coast for
various reasons. These tidally or wind-influenced saltflats have their own avifauna, mainly because they also have a unique invertebrate infauna,and such
areas generally are unappreciated by non-ecologists.
At the San Bernard National Wildlife Refuge southwest of Houston, Texas,
we have observed bird use of wetlands over three different summers reflecting
wet, dry, and average years. The presence of a few inches of water in the areas
above the tidal reach produces amazing differences in plant and bird communities, and the freshwater inflow has made one tidal creek near-fresh in some
years and full-strength seawater in others. One might classify the area as estuarine in some years and freshwater (palustrine) wetland in others (Weller
).
Bird assemblages changed with the water and perhaps with the plants.
During water declines, the more aquatic birds declined and presumably
moved elsewhere, and the mudflats were invaded by shorebirds and more
upland species. When water returned, so did the swimming and diving birds.
These examples all reflect the natural variability of wetlands over various
time scales. Frankly, we are not very adept at dealing with such variation in
processes over long time periods, or with the lack of constancy in the product.
Most people expect wetlands to be constant. We expect them to have discrete
edges and stay out of our fields and buildings. We expects plants and soils and
animals to stay put and to have characteristics that will clearly delineate what is
a wetland and what is terrestrial.We expect them to remain either fresh or salty.
They do not. It is true that not all wetlands are as variable as the examples
above. Among those are the more lake-like systems where the water is more
available and the plant species tend to be perennial aquatics. Another more
stable wetland type is the bog, which may float on the surface of a stable water
body or may have water upwelling. In either case, saturation is nearly constant,
and plants are probably more stable and predictable for bird use.
References
Aldrich, J. W. (). Birds of deciduous forest aquatic succession. Wilson Bulletin
, .
Allen, A. A. (). The red-winged blackbird, a study in the ecology of a cat-tail
marsh. Proceedings of the Linnaean Society of New York , .
Allison, S. K. (). The influence of rainfall variability on the species composition
of a northern California salt marsh plant assemblage. Vegetatio , .
Beecher, W. J. (). Nesting birds and the vegetative substrate. Chicago, IL: Chicago
Ornithological Society.
Begon, M., Harper, J. L., and Townsend, C. R. (). Ecology, rd edn. Oxford:
Blackwell Science.
Bellrose, F. C., Paveglio, F. L. Jr, and Steffeck, D. W. (). Waterfowl populations and
the changing environment of the Illinois River Valley. Illinois Natural History
Survey Bulletin , .
Bertness, M. D. and Ellison, A. M. (). Determinants of pattern in a New England
salt marsh plant community. Ecological Monographs , .
Bildstein, K. L., Post, W., Johnston, J., and Frederick, P. (). Freshwater wetlands,
rainfall, and the breeding ecology of White Ibises in coastal South Carolina.
Wilson Bulletin , .
Breeden, S. and Breeden, B. (). The drought of at the Keoladeo Ghana
Sanctuary, Bharatpur, Rajasthan. Journal of the Bombay Natural History Society
, .
Briggs, S. V. (). Movement patterns and breeding adaptations of arid zone ducks.
Corella , .
Browder, J. A. (). A modeling study of water, wetlands, and wood storks. In
Research Report No. : Wading birds, ed. A. Sprunt, pp. . New York:
National Audubon Society.
Caithamer, D. F. and Dubovsky, J. A. (). Waterfowl population status, .
Washington, DC: U S Fish & Wildlife Service.
Carpenter, S. R. (). Submersed vegetation: an internal factor in lake ecosystem
succession. American Naturalist , .
Chabreck, R. H. (). Coastal marshes: ecology and wildlife management.
Minneapolis, MN: University of Minnesota Press.
Crocker, W. (). Life span of seeds. Botanical Review , .
Crome, F. H. J. (). To drain or not to drain? Intermittent swamp drainage and
waterbird breeding. Emu , .
Custer, T. W., Hines, R. K., and Custer, C. M. (). Nest initiation and clutch size
of Great Blue Heron on the Mississippi River in relation to the flood.
Condor , .
Habitat dynamics
Damman, A. W. H. and French, T. W. (). The ecology of peat bogs of the glaciated
Northeastern United States: a community profile. Report (). Washington,
DC: US Fish and Wildlife Service.
Fredrickson, L. H. and Reid, F. A. (). Impacts of hydrologic alteration on management of freshwater wetlands. In Management of dynamic ecosystems, ed. J. M.
Sweeney, pp. . West Lafayette, IN: The Wildlife Society North Central
Section.
Frith, H. J. (). Waterfowl in Australia. Honolulu, HI: East-West Press
Golet, F. C. and Parkhurst, J. A. (). Freshwater wetland dynamics in South
Kingstown, Rhode Island, . Environmental Management , .
Gosselink, J. G. (). The ecology of delta marshes of coastal Louisiana: a community
profile. FWS/OBS-/. Washington, DC: US Fish & Wildlife Service.
Gunderson, L. H., Light, S. S., and Holling, C. S. (). Lessons from the Everglades.
BioScience Supplement : SS.
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Harris, S. W. and Marshall, W. H. (). Ecology of water-level manipulations of a
northern marsh. Ecology , .
Heitmeyer, M. E. and Fredrickson, L. H. (). Do wetland conditions in the
Mississippi Delta hardwoods influence mallard recruitment? Transactions of the
North American Wildlife & Natural Resources Conference , .
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Dakotas: a community profile. Biological Report (.). Washington, DC: U S
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communities in south boreal lakes. Finish Game Research , .
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habitats of nesting ducks in North Dakota. Journal of Wildlife Management ,
.
Kushlan, J. A. (). Avian use of fluctuating wetlands. In Freshwater wetlands and
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. Jaipur, India: National Institute of Ecology and International Scientific
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ducks and aquatic invertebrates to an experimentally manipulated cattail
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References
Habitat dynamics
10
Population consequences of wetland
abundance and quality
Population consequences
150
Nests
120
y = 18.595 + 5.759 x
r = 0.86
p < 0.05
90
60
30
0
0
12 16
20
24
Population consequences
Birds (100)
Habitat (ha)
12
6
0
2
23
14A
10B
WET MUD
SHALLOW WATER
DEEP WATER
extremely wet or dry conditions. Several bird species linked to prairie pothole
edges also were included in grassland models by Cody (), who analyzed the
relationships between rainfall variation and spring temperatures with bird
density (Fig. .)(Cody ). The positive effects of high rainfall is more
conspicuous in the birds of the wetland edge, like Bobolinks, Red-winged
Blackbirds, and ducks, than with more upland species that also nest in warmer
periods. Dry periods affect especially the quality of basin wetlands for swimming and fish-eating birds like ducks and herons (Fig. .) (Derksen and
Eldridge ; Weller et al. ; Weller ). But the other extreme, flooding,
may disrupt breeding even by swimming birds that nest over water like ducks,
and sometimes without measureable increases in what seem like suitable adjacent areas (Foote ). The effect of floods on riverine wetland and floodplain
birds has been studied less intensively, presumably because it is unpredictable.
In the major Mississippi River flood of , species richness and species abundance of floodplain birds declined after the flood either because of reduced
habitat quality or because of poor reproductive success, but some water birds
were less affected (Miller and Nudds , Knutson and Klaas ). In this
case, increases in abundance were noted in adjacent unflooded areas, presumably caused by displacement during the flood period. However, impacts on
individual species vary with food resources and availability (Custer, Hines and
Custer ).
Presence of breeding birds has been correlated with the number of water
bodies in a landscape region for Canvasbacks (Fig. .)(Sugden ) and for
ducks by taxonomic groups (Johnson, Nichols and Schwartz ). Wetland
formation by beavers is important in a number of areas of North America, and
harvesting of beavers potentially could reduce the number of beaver ponds
and thereby wetland bird habitat. However, studies of beaver harvest and duck
populations suggest that minor beaver harvest is not significant but continued
harvest over to years does result in reduced waterfowl populations (McCall
Population consequences
0.8
0.6
Red-winged
Blackbird
0.4
Yellow Warbler
Savannah Sparrow
0.2
Eastern Kingbird
Western Meadowlark
Common Yellowthroat
0.2
0.4
Willow Flycatcher
Bairds Sparrow
Clay-colored Sparrow
Grasshopper Sparrow
0.6
0.8
0.8 0.6 0.4 0.2
0.2
Correlation coefficient
(density spring temperature)
et al. ). Few studies seem to have been conducted on more stable wetlands
where we might expect variation in numbers to be more a result of continentwide population changes or changes in resources other than water.
Vegetation structure and food resources are inseparably linked to wetland
selection or to within-wetland habitat selection on a breeding area. Returning
adults are forced to abandon a former or potential breeding area if nesting
cover and food are not available. Hence, risk of mobility and search would confront them even if nest or roost sites were available. In both American Coots
and Yellow-headed Blackbirds, we have observed but been unable to quantify
the arrival, temporary occupancy, and then sudden population decline that
suggested that birds had tested quickly and moved elsewhere in search of better
resources. Loss of food resources during nesting can cause abandonment of
the nests and young. This is especially obvious in colonial species such as terns,
egrets, and, probably, pelicans. Terns may remain and nest if at least some food
is available but reproductive success as measured by number of young per pair
of terns is related to food biomass in a sigmoid fashion (Suddaby and Ratcliffe
). Brown and Urban () noted abandonment of nests of African
Great White Pelicans caused by a collapse of the fish population. And Wood
Storks often abandon nests when food resources are unavailable because of
high water.
Population consequences
Figure .. Relationship of
various grassland and wetland
birds to climatic regimes in the
North American prairie (Cody
, with permission by
Academic Press).
Figure .. Numbers of
Canvasback pairs established
in eight wetland complexes in
Saskatchewan during various
water conditions (Sugden ).
Population consequences
Figure .. Regression of
species richness against
number of pools of open water
in emergent marsh vegetation.
The numbers represent observation points from different
sample years, and the two clusters reflect low richness when
few pools exist (i.e., dense vegetation) and high richness when
vegetated is opened by interspersed pools (based on data in
Weller and Fredrickson ).
(Wilcove ). This may be compensated in part by the protection of overwater nest sites, but many edge-nesters are exposed to greater snake and
mammal predation.
Physical features like wetland size were discussed earlier and are important in explaining population size where species overlap to a limited degree by
preferences for a particular wetland size. In the case of grebes, this is linked
with vegetation: Horned Grebes use the smaller more open areas, Eared
Grebes use medium to large areas that also lacked vegetation, and Pied-billed
Grebes use medium to large areas that are more vegetated (Boe , Faaborg
). Some extensive studies of Finish lakes with a wide range of nutrients,
vegetation, and sizes has provided evidence of strong linkages of waterbird
species to certain limnological characteristics such as (i) fish eaters in oligotrophic lakes (loons, mergansers); (ii) certain typical species in eutropic
lakes (small grebes, coots, and teal, shovellers, and bay ducks); and (iii)
species using shallow, nutrient-rich, but stressed lakes (termed mixotrophic),
such as Eurasian Wigeon, Northern Pintail, and Tufted Duck (Kauppinen
).
Other physical features are important, often because they affect or are
affected by water. Nesting White-throated Dippers in Britain seem to prefer
lower stream gradients where currents are slower and pools are more common
(Tyler and Ormerod ). This also seems true of feeding Spectacled Ducks of
South America, which feed and nest along slower streams, whereas Torrent
Ducks of the same areas of the Andes use smaller, higher streams with strong
currents (Weller ). I suspect that feeding Blue Ducks of New Zealand fall
somewhere between these extremes (Eldridge ), as do Harlequin Ducks of
North America (Bellrose ).
Tidal regimes strongly affect shoreline exposure (Evans, Goss-Custard and
Hale ), especially of mudflats, impacting the availability of feeding sites of
many species, like shorebirds that feed on the wet mudflats of very shallow
water (Fig. .,) (Burger et al. ; Evans et al. ). Current telemetry
methods have allowed precise measurement of spatial distribution and seasonal timing of mudflat use in Western Sandpipers and other small shorebirds
Population consequences
Population consequences
Figure .. Relationships
between habitat resources and
use by waders generalized for
application to restoration of
the Kissimmee River Basin of
Florida (Weller ). Use of
two waterbird guilds as evaluation tools for the Kissimmee
River restoration. Restoration
Ecology (), . Reprinted
by permission of Blackwell
Science, Inc.).
Population consequences
Population consequences
Population consequences
Figure .. A semiquantitative
model of eelgrass, fish, and
waterfowl trophic relationships
based on early studies of annual
production of seagrasses in
Denmark (numbers in
metric tonnes) (redrawn from
Thayer, Wolfe and Williams
, with permission).
years until recovery of the plant food. Recently, the western Pacific population
has experienced major declines attributed to eelgrass losses. These are blamed
on estuarine pollution and perhaps oyster mariculture, as well as boating disturbance that prevents the use of the food resources available (Einarsen ,
Wilson and Atkinson ).
The above observations suggests a strong and logical relationship
between the number, size, and quality of wetlands with species richness,
population density, and community structure. But in our zeal to protect wetlands, it is common for conservationists to use the loss of wetlands (now over
% in the contiguous USA according to Tiner ()) as a crude measure to
estimate the decline in numbers if not species of birds in the USA since
European settlement. But when we consider the fact that all wetlands vary in
their suitability for birds, we would need an estimate based not only on the
average number of wetlands available for use over the recent history of each
species but also on their quality. The index that best approximates conditions
for waterfowl breeding in the North American prairie potholes is an aerial
survey where numbers (usually without measures of quality or size) of basin
wetlands have been counted concurrently with duck populations. At this
scale of tens of thousands of wetlands across the breeding range of the more
common ducks, populations tend to decline after a series of years when more
wetlands are dry, and enormous increases in populations can occur after a
series of wet years when more basin wetlands contain water (Caithhamer and
Dubovsky , Cowardin and Blohm , Dzubin ). There are many
deviations from the expected, and a time lag of a year or more seems
common owing to variation in breeding patterns and maturation of the
various species.
This does not infer that the decline in ducks and other waterbirds over long
periods is not proportional to wetland losses, but that we lack accurate background bird population data and information on wetland quality.
Additionally, waterbirds are adaptable and movements mask some changes.
Obviously, this also differs by bird group and life-history strategy: sparrows
Population consequences
and wrens with small home ranges and high fidelity to breeding sites would be
expected to suffer serious population losses even with declines of small wetlands. Ducks, rails, herons, and other waterbirds seemingly seek out better
habitats (i.e., locally nomadic behavior induced by climatic conditions), and
ducks and herons probably explore new breeding and wintering areas when
pressed by poor conditions in previously used sites. But how far can the system
be pushed? Most optimal habitat ranges have been populated, and marginal
ones presumably can be used at some times but not at others because of variation in water regimes and climate. There are few opportunities to get this kind
of information on wetland birds except perhaps on colonial breeders (e.g.,
ibises, egrets and herons), which are conspicuous by their large nesting
colonies. This does not really make them easier to count because of layering of
nests as well as tree cover, but they have been of concern because of dramatic
declines and, therefore, are surveyed periodically. Several recent analyses of
waders have demonstrated that colonies seem to shift geographically, i.e.,
abandoning long-used sites and appearing suddenly in other areas, states, or
regions.We assume they move to better conditions, although we can only speculate on why.Water conditions are among the most important influences, with
high water levels often as detrimental for some species as drought is for others
because it reduces availability of prey (Kushlan ). Declines have been especially prominent in Florida and increases in Georgia and South Carolina;
recently, increased numbers of species have been noted westward from
Georgia to Louisiana and Texas concomitant with declines in Florida populations.
Another type of population change linked to wetland habitat quality or
dynamics is the change in numbers of birds observed (or shot in the case of
ducks and geese) at favoring migratory stopovers and wintering areas. One of
the earliest of such observations resulted from kill records of the Winous Point
Hunting Club along the shores of Lake Erie in Ohio, where Redhead ducks
declined drastically from to . Although this was cited as evidence of
the serious decline of the species, I suspect it was a product of habitat change in
the managed area, which favored dabbling ducks. Another was the case of the
loss of the Canvasbacks at Lake Christina in Minnesota, which was attributed
to increased turbidity and the decline of the favored submergent vegetation,
sago pondweeds. Similar causes and impacts were documented in the
Chesapeake Bay, where restoration efforts have included reduction of fertilizer
and other nutrient flowing into the bay. Decline in water quality in pools of the
Illinois River caused by upstream siltation and pollutants resulted in losses of
invertebrate populations and pondweeds, with major impacts on use by
migrant and wintering duck populations (Mills, Starrett and Bellrose ).
Moreover, migratory routes have changed for Canvasbacks and perhaps for
other plant-feeding ducks because they have shifted to other areas with more
available resources.
Clearly, more data are needed on this important topic because it is the key
Population consequences
(e) Floating nest of a Piedbilled Grebe made of wet emergent vegetation of the previous
year. Eggs are normally covered
by the incubating bird before
departure (Ruthven, Iowa).
to the evaluation of conservation efforts for individual species and assemblages. Because of the great expense of conducting this kind of research, theoretical and mathematical models will be essential to synthesize the
information we have and to use it to identify priorities in research. However,
this can only work if we have better information on the strategies of resource
use by birds, e.g., how and why bird use certain nest sites, what restricts use of
roost sites, and how flexible birds are in changing migration routes.
References
Alonso, J. C., Alonso, J. A., and Veiga, J.P. (). Flocking in wintering Common
Cranes: influence of population size, food abundance and habitat patchiness.
Ornis Scandinavia , .
Arengo, F. and Baldassarre, G. A. (). Effects of food density on the behavior and
distribution of nonbreeding American Flamingos in Yucatan, Mexico. Condor ,
.
Baldassare, G. A. and Bolen, E. G. (). Waterfowl ecology and management. New
York: Wiley.
Banko, W. E. (). The trumpeter Swan: its history, habits, and population in the
United States. North American Fauna . Washington, DC: U S Fish & Wildlife
Service.
Bateman, H. A., Joanen, T., and Stutzenbaker, C. D. (). History and status of
Midcontinent Snow Geese on their Gulf Coast winter range. In Waterfowl in
winter, ed. M. W. Weller, pp. . Minneapolis, MN: University of Minnesota
Press.
Bellrose, E C. (). Ducks, geese and swans of North America rd edn. Washington,
DC: Wildlife Management Institute.
Bellrose, F. C., Paveglio, F. L. Jr and Steffeck, D. W. (). Waterfowl populations and
the changing environment of the Illinois River Valley. Illinois Natural History
Survey Bulletin , .
Bengtson, S. A. (). Variations in clutch-size in ducks in relation to the food
supply. Ibis , .
Boe, J. S. (). Colony site selection by Eared Grebes in Minnesota. Colonial
Waterbirds , .
Bolen, E. G. and Rylander, M. K. (). Whistling ducks: zoogeography, ecology,
anatomy. Special Publication . Lubbock, TX: Texas Technical University
Museum.
Brown, L. H. and Urban, E. K. (). The breeding biology of the Great White
Pelican Pelecanus onocrotalus roseus at Lake Shale, Ethiopia. Ibis , .
Bryant, D. M. (). Effects of prey density and site character on estuary usage by
overwintering waders (Charadrii). Estuarine and Coastal Marine Science ,
.
Burger, J., Howe, M. A., Hahn, D. C., and Chase, J. (). Effects of tide cycles on
habitat selection and habitat partitioning by migrating shorebirds. Auk ,
.
Caithamer, D. F. and Dubovsky, J. A. (). Waterfowl population status, .
Washington DC: U S Fish & Wildlife Service.
References
Cargill, S. M. and Jeffries, R. L. (). The effects of grazing by lesser snow geese
on the vegetation of a sub-Arctic salt marsh. Journal of Applied Ecology ,
.
Cody, M. L. (). Habitat selection in grassland and open country birds. In Habitat
selection in birds, ed. M. L. Cody, pp. . Orlando, FL: Academic Press.
Cottam, C., Lynch, J. J., and Nelson, A. L. (). Food habits and management of
American sea brant. Journal of Wildlife Management , .
Cowardin, L. M. and Blohm, R. J. (). Breeding population inventories and measures of recruitment. In Ecology and management of breeding waterfowl, eds.
B. D. J. Batt, A. D. Afton, M. G. Anderson, C. D. Ankney, D. H. Johnson, J. A.
Kadlec, and G. L. Krapu, pp.. Minneapolis, MN: University of Minnesota
Press.
Custer, T. W., Hines, R. K. and Custer, C. M. (). Nest initiation and clutch size of
Great Blue Herons on the Mississippi River in relation to the flood. Condor
, .
Danell, K. and Sjoberg, K. (). Seasonal emergence of chironomids in relation to
egg laying and hatching of ducks in a restored lake. Wildfowl , .
Derksen, D. V. and Eldridge, W. D. (). Drought-displacement of pintails to the
Arctic Coastal Plain, Alaska. Journal of Wildlife Management , .
Dzubin, A. (). Comments on carrying capacity of small ponds for ducks and
possible effects of density on Mallard production. In Saskatoon wetlands seminar,
pp. . Report Series, No. . Ottawa: Canadian Wildlife Service.
Eadie, J. M. and Keast, A.. (). Do Goldeneye and perch compete for food?
Oecologia , .
Einarsen, A. S. (). Black brant, sea goose of the Pacific coast. Seattle: University
of Washington Press.
Eldridge, J. L. (). Territoriality in a river specialist, the Blue Duck. Wildfowl ,
.
Eriksson, M. O. G. (). Lake selection by Goldeneye ducklings in relation to the
abundance of food. Wildfowl , .
Evans, P. R., Goss-Custard, J. D., and Hale, W. G. (eds.) (). Coastal waders and
wildfowl in winter. Cambridge: Cambridge University Press.
Faaborg, J. (). Habitat selection and territorial behavior of the small grebes of
North Dakota. Wilson Bulletin , .
Fleury, E. and Sherry, T. W. (). Long-term population trends of colonial wading
birds in the southern United States: the impact of crayfish aquaculture on
Louisiana populations. Auk , .
Foote, A. L. (). Response of nesting waterfowl to flooding in Great Salt Lake
wetlands. Great Basin Naturalist , .
Fredrickson, L. H. (). The breeding biology of American coots in Iowa.
Wilson Bulletin , .
Heitmeyer, M. E. and Fredrickson, L. H. (). Do wetland conditions in the
Mississippi Delta hardwoods influence mallard recruitment? Transaction
of the North American Wildlife and Natural Resource Conference , .
Hill, D., Wright, R., and Street, M. (). Survival of Mallard ducklings Anas
platyrhynchos and competition with fish for invertebrates in a flooded gravel
quarry in England. Ibis , .
Population consequences
Mitsch, W. J. and Gosselink, J. G. (). Wetlands, nd edn. New York: Van Nostrand
Reinholdt.
Muehistein, L. K. (). Perspectives on the wasting disease of eelgrass Zostera
marina. Diseases of Aquatic Organisms , .
Murkin, H. R., Kaminski, R. M., and Titman, R. D. (). Responses by dabbling
ducks and aquatic invertebrates to an experimentally manipulated cattail marsh.
Canadian Journal of Zoology , .
Myers, J. P., Williams, S. L., and Pitelka, F. A. (). An experimental analysis of prey
availability for Sanderlings. (Aves: Scolopacidae) feeding on sandy beach crustaceans. Journal of Zoology , .
Nilsson, L. (). Breeding waterfowl in eutrophicated lakes in south Sweden.
Wildfowl , .
References
Pehrsson, O. (). Egg and clutch size in the mallard as related to food quality.
Canadian Journal of Zoology , .
Powell, G. V. N. (). Food availability and reproduction of Great Blue White
Herons, Ardea herodias: a food additive study. Colonial Waterbirds ,
.
Quammen, M. L. (). Predation by shorebirds fish and crabs on invertebrates
on intertidial mudflats: an experimental test. Ecology , .
Rahmani, A. R. and Shobrak, M. Y. (). Glossy ibises (Plegadis fascinellus) and
Black-tailed Godwits (Limosa limosa) feeding on sorghum in flooded fields in
southwestern Saudi Arabia. Colonial Waterbirds , .
Rogers, J. (). Low water and lesser scaup reproduction near Erickson, Manitoba.
Transactions of the North American Wildlife and Natural Resource Conference ,
.
Rottenborn, S. C. (). The use of coastal agricultural fields in Virginia as foraging
habitat by shorebirds. Wilson Bulletin , .
Skagen, S. K. and Knopf, F. L. (). Migrating shorebirds and habitat dynamics
at a prairie wetland complex. Wilson Bulletin , .
Stiles, F. G. and Skutch, A. F. (). A guide to the birds of Costa Rica. Ithaca,
NY: Cornell University Press.
Straw, J. A. Jr, Krementz, D. G., Olinde, M. W., and Sepek, G. F. (). American
Woodcock. In Game bird management in North America, eds. T. C. Tacha, and
C. E. Braun, pp. . Washington, DC: International Association of Fish &
Wildlife Agencies.
Suddaby, D. and Ratcliffe, N. (). The effects of fluctuating food availability on
breeding Arctic Tern (Sterna paradisea). Auk , .
Sugden, L. G. (). Canvasback habitat use and production in Saskatchewan
parklands. Canadian Wildlife Service Occasional Papers , .
Thayer, G. W., Wolfe, D. A., and Williams, R. B. (). The impact of man on sea
grass systems. American Scientist , .
Thompson, D. B. A. (). The abundance and distribution of intertidal invertebrates, and an estimate of their selection by Shelduck. Wildfowl , .
Tiner, R. W. Jr (). Wetlands of the United States: current status and recent trends.
Newton Corner, MA: US Fish and Wildlife Service.
Tyler, S. and Ormerod, S.(). The dippers. San Diego, CA: Academic Press.
Warnock, S. E. and Takekawa, J. Y. (). Habitat preferences of wintering shorebirds
in a temporally changing environment: Western Sandpipers in the San Francisco
Bay. Auk , .
Weller, M. W. (). Distribution and migration of the redhead. Journal of Wildlife
Management , .
Weller, M. W. (). Notes on some Argentine anatids. Wilson Bulletin , .
Weller, M. W. (). Habitat selection and feeding patterns of Brown Teal (Anas
castanea chlorotis) on Great Barrier Island. Notornis , .
Weller, M. W. (). Habitat selection by waterfowl of Argentine Isla Grande. Wilson
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Weller, M. W. (). Density and habitat relationships of blue- winged teal nesting
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Weller, M. W. (). Estimating wildlife and wetland losses due to drainage and
Population consequences
Further reading
ponds on an East Texas coal mine. Masters thesis. College Station, TX: Texas
A&M University.
Schaffer, W. M. (). Optimal reproductive effort in fluctuating environments.
American Naturalist , .
Utschick, H. (). Die Wasservogel als Indikatoren fr den okologischen Zustand
von Seen (with English summary). Verhandlungen der Ornithologischen
Gesellschaft in Bayern , .
Weller, M. W. (). Seasonal dynamics of bird assemblages in a Texas estuarine
wetland. Journal of Field Ornithology , .
Population consequences
11
How birds influence wetlands
Although our major goal throughout the book has been to relate bird habitat
use to wetland resources and features, birds that can directly affect a wetland
ecosystem by their influence on vegetation and thereby bird habitat have an
impact on other birds directly or alter ecosystem processes that may induce
change within the entire system (Table .).
Because of their mobility, birds facilitate movement of invertebrates to
established wetlands that have lost their population via drought (Swanson
), as all invertebrates do not have eggs that can survive long periods
without water (Wiggins, MacKay and Smith ). Birds also transport invertebrate eggs to newly created wetlands such as strip-mine ponds or riverformed wetlands. Common herbivorous crustaceans like amphipods have
been especially prominent in studies of the plumage of hunter-killed or live
waterfowl in both Europe and North America, but snails, snail eggs, and other
macrocrustaceans have also been noted in plumage. One cannot wade in wetlands without experiencing invertebrates and seeds clinging to boots, so it is no
surprise that birds (and also beavers and muskrats) carry them as well. Plant
materials such as seeds and foliage of submergent plants are commonly found
in or on the plumage of herbivorous ducks and coots. One study reported
seeds of saltmarsh plants, most of which had special adhesive devices or
systems, carried by ducks (Vivian-Smith and Stiles ). This helps to explain
a gradual increase in richness that occurs over time in wetland developed on
strip-mined lands, where plant seed banks are minimal (McKnight, ). In
such cases, wetland seed banks from - to -year-old ponds are commonly used
to establish vegetation in newly created wetlands. Although some seeds may be
wind-carried (willows, cattail, or fine seeds like sedges), most seeds are
unlikely to be moved in this way. Birds also transport algae (Proctor ),
seeds (DeVlaming and Proctor ), and invertebrates (Malone ) in their
digestive tracts. Planktonic algae are passed through intact, although viability
is doubtful for most (Atkinson ), but about % viability has been noted in
seeds found in duck digestive tracts (Powers, Noble and Chabreck ).
Although this figure seems small, it would only take a few seeds in the suitable
sites used by ducks to establish a plant species.
The second major way that birds affect wetlands is as users. Herbivores like
Lesser Snow Geese in fresh or saline marshes (Glazener , Smith and Odum
), and Greylag Geese (Kvet and Hudek ) and White-fronted Geese in
fresh marshes (personal observations) can be as devastating to even robust
emergent vegetation as are Muskrats, Nutria, and feral pigs. In this way, they
influence the many aspects of plant succession mentioned earlier and thereby
have ecosystem-level consequences. Brant, swans, wigeon, and some bay ducks
feed heavily on eelgrass foliage, although the impact is often modest and
difficult to measure (Nienhuis and van Ierland ). Many duck species and
coots in both coastal and inland saline settings feed heavily on the more delicate foliage of widgeongrass, presumably to the point of stripping areas and
inducing birds to move elsewhere. Coots measurably impact Hydrilla sp., a
favored, exotic submergent that grows in deep water and reaches to the surface
(Esler ). Redhead ducks concentrate on and impact the abundance of rhizomes and tubers of seagrasses in winter (Cornelius ), and Sandhill Cranes
eat sedge tubers in meadows during migration and breeding (Reineke and
Krapu ). In all cases, their flock sizes and density at feeding sites suggest
that they must have measurable consequences on the density and distribution
of plant propagules. African flamingos, known for their concentration, which
numbers in the hundreds of thousands, reportedly devour tons of planktonic
organisms such as algae or invertebrates per day in interior saline lakes. While
these levels of consumption are impressive, their impact on the system probably is more seasonal than permanent, but the long-term consequences of
movement of this nutrient base away from the lake via nomadic or migratory
movements requires detailed study.
Carnivorous birds remove major biomass in the form of invertebrates, fish,
small mammals, and birds, and several meaningful estimates have been made.
Studies of the use of mudflat invertebrate in fauna have been made in connection with the shorebird studies mentioned above (e.g., Quammen ). By
use of captive Red Phalaropes, both reduction and size selection of crustacean
prey has been suggested in Arctic wetlands (personal communication from S.
I. Dodson). Herons and other fish-eating birds undoubtedly take enormous
numbers of fish, but much of this feeding seems to be most efficient when
water levels decline, oxygen is reduced, or fish are stunned by cold. An estimated take of % of the fish of a declining pond was noted compared with a
% loss during total drying (Kushlan ). However, Smith et al. () were
unable to demonstrate significant diving duck impact on most invertebrates
except for clams. Studies of oystercatchers, mentioned above also suggest that
impact on large and relatively sedentary prey items may be greatest, but major
reduction of the invertebrate community has not yet been demonstrated.
Mixed flocks of cormorants and pelicans do take many large fish by their
efficient cooperative feeding efforts, but the real impact on the fish populations is very difficult to assess. Mergansers (and other sea ducks) that use eggs
(i.e. spawn) in streams or in freshwater inflows to bays have been controlled in
various places because of their potential competition with what are viewed as
human resources. Consumption of small or young birds, herptiles, and
mammals by predatory birds such as hawks or herons obviously are an established part of the food chain, moving energy from low to higher consumer
levels.
Waterbird predation may influence ecosystem-level processes via impact
on major herbivores that directly influence the vegetation base for all consumers. Several examples of heavy predation have been mentioned, but such
impacts are difficult to assess in complex systems with many layers of consumers. In a more simple system, the role of sea ducks (eiders) on sea urchins,
which feed on kelp has been investigated, and although measurable consequences were complicated by structural aspects of the vegetation and the technical difficulty of assessing foods in kelp forests, strong evidence was
obtained that eiders can affect sea urchin populations in small patches, and
thereby potentially influence the survival of patches of kelp (Bustnes and
Lonne ).
Less direct but perhaps more widespread influences of waterbirds on
wetland occurs through nutrients added to the water column from defecation
and sometimes from carcasses of flocking birds in breeding colonies and
roosting sites. Among egrets and herons, this could have longer-term implications for the plant community, as mortality of trees has been attributed to
heavy fecal deposition. Interestingly, dead trees are probably better nest sites
for most herons and egrets, and the opening in the canopy must create new
plant germination sites that ultimately influence local plant succession.
Franklins Gulls concentrate in enormous breeding colonies and potentially
can have a major nutrient input to marsh emergents (McColl and Burger )
as can blackbirds and starlings roosting in cattail marshes (Hayes and Caslick
). Flocks of free-flying but site-attracted waterbirds have been shown to
deposit significant nutrients with the potential to influence algal growth, thus
entering the nutrient cycle, for example Canada Geese in Michigan (Manny,
Wetzel and Johnson ), mixed species of ducks and coots in New York
(Have ), marine birds in estuaries (Ganning and Wulff ), and Redhead
Figure .. Hypothetical
influence of hillside nesting of
Antarctic seabirds on nutrients
of lower freshwater wetlands
that are rich in fairy shrimp and
other invertebrates and are used
for breeding by South Georgia
(S. G.) Pintails (L. M., Lightmantled; BL., Black; A.,
Antarctic) (Weller , with
permission).
References
12
Conservation implications
Wetland birds differ widely in their species composition and relative abundance within a community, but we lack thorough studies of communities in
many wetland types and at various temporal scales. In addition to helping us to
understand how such systems function, these data provide a basis for measuring change and evaluating impacts of natural and societal influences and,
therefore, are important in conservation efforts. The purpose of this chapter is
to summarize some of the patterns of habitat use by wetland birds and their
potential biological properties, outlined above, and generalize on their relationship to conservation and management. Conserving wetland birds also
requires an understanding of their relationships to society, because human
populations do not have an impact on all groups of wetland birds uniformly
(Weller a). Moreover, some wetland types are more at risk than others
because of alternative human uses, which will have a secondary effect on the
success of birds there (Krapu et al. ).
Interest in wetland bird conservation often has focused on single species or
taxonomic groups that have been of interest or importance for esthetic, recreational, or food values, and efforts to maintain populations have been funded
by user-fees (e.g., hunting, bird watching). The use of general tax appropriations is less common except for those species that are especially rare; this has
been an area of increasing concern because of the role that society has played in
extirpations and extinctions. Although initial efforts at conservation of such
species were aimed at legal protection, conservation strategies now tend to
focus on preservation of suitable habitat or the assurance of continued functioning of existing habitat as the first step in protection of threatened populations, species, and communities.
The following concepts and commentaries are divided by general topics
and include some issues and strategies of importance in wetland bird
conservation. These are examples rather than a complete cook book of
approaches. As with all generalizations, there may be major variation among
bird groups, types of wetland, and geographical location, but they should help
to identify patterns applicable in the development and evaluation of a
conservation strategy for a species, assemblage, or community of wetland
birds. While some of these ideas may seem to lack immediate application
Conservation implications
Conservation implications
. Although a birds habitat needs may seem simple, species preferences for
different vegetation structure, foods, and water depth and dynamics often
are poorly known and, therefore impossible to manage for or replicate
without extended observations and interpretation based on similar
species elsewhere.
. Where either specific habitat needs or patterns for groups are known,
success at creating suitable living conditions has been great, but it can be
costly in time and money. Prominent among direct and relatively inexpensive measures are the provision of nest structures that encourage
selection in a particular wetland where the species has bred previously.
These species include hole-nesting birds like North American Wood
Ducks or Prothonotary Warblers, platform-nesting birds like Ospreys,
Peregrine Falcons, and Canada Geese, and sandbar birds like Least Terns.
Artificial structures are regarded negatively by many, and their use should
be minimized to periods before natural structures have developed, such
as in restored or created wetlands.
. Planted food patches and artificial feeding attract field-feeding, seedeating birds like sparrows, ducks, geese, and cranes but also less desirable
groups like blackbirds (Icterids). Direct feeding is of course very costly
and only a short-term solution often used to divert birds from agricultural
crops. In most cases, natural seed banks provide adequate food when
water regimes are properly managed.
. Management of invertebrate foods is more difficult but becoming better
understood and more feasible in wetlands with controlled water regimes.
Typically, they are incidental to drawdowns designed to regulate vegetation, but water regulation to create mudflats for shorebirds or reduce
water depths for duck feeding is more common (e.g., Helmers ).
water levels. Where wetland tracts are large and dispersed enough to provide
for both species but in different areas, the long-term values of increased
wetland productivity that results from the dynamic water regimes and the
natural mobility of the birds probably produces the most natural and self-sustaining strategy for both.
Temporal and spatial scales commonly are not considered. A wetland area
that serves one migratory or mobile bird species at one season will also serve
many others at other times, and resource needs may differ drastically, thus
requiring considerable knowledge of avian biology and potential management
strategies.
Wetland birds sometimes respond quickly to habitat management, creating the illusion that all wetland birds are easily managed for and populations
enhanced at will. In fact, we often are attracting local birds and have no quantitative measure of the population consequences of the action and may not be
achieving goals for whole assemblages or communities in jeopardy. Such
efforts also may attract species like Red-winged Blackbirds, locally considered
as nuisances or agricultural pests.
Although the abundance of many species of wetland birds seem seriously
affected by one or several years of drought, their reproductive strategies are
products of such evolutionary stresses and most seem to have compensatory
mechanisms of mobility to alternative sites, large clutches of eggs during good
years, and multiple nesting attempts to ensure a high reproductive rate when
conditions allow. Therefore, we should not expect annual constancy in
populations when habitats are naturally unstable at various time scales.
Conservation implications
Conservation implications
. Extreme periodic drought, while harmful to bird use in the short term, is
essential for (i) cycling of nutrients deposited in anaerobic wetland substrates, and (ii) re-vegetation of shallow wetlands that otherwise may
remain as open-water or submergent aquatic beds in both saline and
freshwater sites. Extended droughts may result in increased salinity in
some areas and even salt deposits; these may change the vegetation drastically, resulting in lengthy periods when the area is attractive only to those
species of birds especially tolerant of these conditions (e.g., avocets and
stilts in salt pans).
. From a conservation standpoint, severe drought is of greater concern
because most wetlands are permanently drained during this event for
alternative uses.
. Bird diversity alone is not the best measure of the value of a wetland; for
example, Everglade Kites live in wetlands that are simple, relative to some
others, but we would not encourage the elimination of these wetlands in
favor of other more complex or species-rich ones.
.We must evaluate species richness and diversity of assemblages on large
spatial and temporal scales. All birds will not be equally abundant at one
time or under all wetland conditions, but their dynamics will change over
time and area.
12.4.2 among-wetland comparisons
. Bird species diversity of two wetlands of the same size will be greater in
the wetland with the greatest habitat diversity.
. Abundance and probably biomass of birds in two wetlands of the same
size will be greater in the wetland with the greatest nutrient riches (e.g.,
more eutrophic) because the potentially greater productivity is reflected
in invertebrates and other community components that may be important resources.
. The likelihood that two wetlands will have the same bird species composition is greater in wetlands of the same type that are close together rather
than in those that are far apart. In mobile species, two wetlands that are
close together may be used by the same individuals.
Conservation implications
. Vegetation dynamics
Understanding wetland plant succession by type and region is essential to providing quality habitat for wetland birds to use for specific needs such as cover,
nesting sites, and foods. Results of managing plant succession always are
influenced by chance but most birds seem sufficiently adaptable in use of foods
or cover that the plant species itself is less important than the general type of
food produced (e.g., seed size, nutritional quality, and location. Basin
configuration and shoreline slope are important both in vegetation management and in bird use because they affect food availability and hence bird foraging tactics. Upslope management for water quality and quantity directly
dictates plant communities in the wetland and thereby its attractiveness to
birds and other wildlife.
References
Diamond, J. M. (). The island dilemma: lessons of modern biogeographic studies
on the designs of natural preserves. Biological Conservation , .
Helmers, D. L. (). Shorebird Management Manual. Publication No. . Manomet,
MA: Wetlands for the Americas.
Krapu, G. L., Greenwood, R. J., Dwyer, C. P., Kraft, K. M., and Cowardin, L. M. ().
Wetland use, settling patterns, and recruitments in mallards. Journal of Wildlife
Management , .
Weller, M. W. (a). The influence of hydrologic maxima and minima on wildlife
habitat and production values of wetlands. In Wetland Hydrology, eds. J. Kusler
and G. Brooks, pp. . Technical Report . Berne, NY: Association of State
Wetland Managers.
Weller, M. W. (b). Issues and approaches in assessing cumulative impacts on
wildlife habitat in wetlands. Environmental Management , .
References
13
Measures of bird habitat use and quality
To apply some of the concepts and principles noted in previous chapters, biologists typically measure habitat quality for birds for a variety of purposes
(Adamus et al. , , Bookout ): (i) ornithologists or ecologists studying the biology and ecology of individual species or groups; (ii) ecologists
studying how birds are distributed in space and time; (iii) wildlife managers
measuring habitat use or habitat quality to assess the effectiveness of some
form of management; (iv) consultants selecting alternatives for habitat creation or restoration projects; (v) conservation biologists setting priorities for
purchase or lease options to protect species, habitats, or communities; (vi)
conservation or regulatory agencies evaluating the effectiveness of a wetland
mitigation (i.e., replacement) project and subsequent monitoring of success in
meeting goals; and (vii) regulatory biologists and population managers following trends in populations, measuring effects of harvest regulations, or
evaluating environmental impacts,
Population size and density often are used as indicators of good or poor
habitat, but such data have numerous biases and are expensive to obtain.
However gathered, it is also essential to have measurements of habitat features
to correlate with the population indices to provide some relative scale of
quality. Measurement of habitat condition for a particular animal commonly
is termed habitat suitability, but some workers advise against use of the word
suitabilityas redundant (i.e., it either is or is not habitat) and recommend the
word quality instead (Hall, Krausman and Morrison ). However, suitability is an integral part of a widely used evaluation system (see below) that
describes a range of features attractive to and indicative of habitat quality for
the species in question. Habitat use infers the presence or relative occupancy
rate of birds in a particular habitat as measured by presence and absence, index
of abundance, or complete census of sampling areas. Selection is sometimes
used as a synonym for use but it is better used to describe the process of choosing from available options (Johnson ). If one also has data on availability
of potential habitat or a specific resource such as food or nest sites, preference
may be expressed as a percentage of that present and feasible to use (Alldrege
and Ratti ). This measurement is not a simple task because presence of a
habitat or food does not necessarily mean that it is available to the bird. For
many habitats, and countless feeding tactics dictated by the wetland and temporal variations in habitat use. However, such approaches may provide data
useful for comparison with other high-order groups in multispecies analyses,
such as wading or walking birds like herons, which feed on different foods in
different ways.
13.1.2 multispecies level assessment
Community-level assessments usually are based on population data by species
recorded on surveys, as mentioned above, from a particular habitat type. By
expressing the percentage that each species constitutes of the total, a composite
view termed species composition is derived. This is an old method that has
regained popularity because it reflects diversity of species and relative
numbers with little mathematical bias (James and Rathbun ). Perhaps the
most simple measure of the variety of species present in a specific habitat or
area is a species list on which the number of species (species richness) is used
for comparisons between areas (Pielou ). It often is correlated with habitat
features and food resources, but without additional and often detailed
information on species or habitats, it can mask major habitat differences. For
example, one could have species of shorebirds using the invertebrates in one
wetland that was mostly mudflat and species of ducks and other swimming
waterbirds using different foliage and seeds in another wetland of more
permanent water. These are not equivalent and will not provide valuable data
unless the analysis addresses the question asked.
The species diversity index is a mathematical approach developed as an
index to diversity of plants that has been very popular among animal ecologists as well. It includes a measure of both the diversity of bird species (richness
above) and relative abundance of each species (evenness). If only this index is
published or used in statistical analyses, species identity is lost as is information on resource use that can be inferred from knowing the natural history of
those birds.
13.1.3 examples of some common bird surveys
Because of their economic and recreational importance, or because of threatened status, waterbirds have figured prominently in the development of census
techniques for all birds. Moreover, volunteers have played a major role in providing information to aid in bird conservation and population management.
Nationwide teams of observers were organized by Wells W. Cooke of the US
Biological Survey to make counts and observations on timing of migration in
shorebirds and waterfowl in the late s and early s (e.g., Cooke ).
These formed the basis for understanding many patterns of distribution on a
continent-wide basis and for concepts of bird flyways (Lincoln ) and corridors (Bellrose ).
Reduced populations of egrets, herons, and other waders have been cause
for concern, and a variety of techniques have been developed and used to
assess population changes as well as shifts in distribution. Traditionally,
nesting colonies have attracted the most attention and best records (e.g., the
Colonial Bird Register). Many colonies of nesting birds are in conspicuous sites
in trees or shrubs, and annual or periodic surveys have occurred to assess
long-term trends (Lange , Nesbitt et al. , Portnoy , Runde et al.
). Broad-scale but systematic censuses by plane and on ground have been
used to provide data on colony location, feeding areas, species richness, and
general population size. In colonies where thousands of birds nest and are at
several levels in the vegetation and, therefore, cannot be counted precisely
from the air, ground strip-transects through colonies have been used to estimate numbers of nests (King ). Comparisons of helicopter and fixedwing aircraft for colony surveys show that helicopters are much more accurate
(<% error for white species but much greater for dark birds) than fixedwing flights (>% even for white birds)(Kushlan ). However, use of
systematic, transect-based aerial surveys reportedly underestimates the true
wader population by to %, which is comparable to the variation for
waterfowl surveys. Comparison of aerial and water-level surveys (via airboat)
also demonstrates an advantage for colony discovery by aerial methods, espe-
cially for white birds like egrets as opposed to dark herons or ibises (Frederick
et al. ).
Numerous individuals and groups worldwide have established long-term
surveys of species culminating in detailed avian atlases, and more are now
assessing birds by habitat as well as in geographical units. A state-level survey
established in North Dakota formed the basis for a habitat-oriented bird book
that provided species composition and species density estimates for some
groups (Stewart ). Re-visits to these sites have resulted in comparative
trends for many species and reflect long-term declines for wetland and grassland species (Igl and Johnson ). Some waterfowl surveys at the national
level show migration routes as well as population density estimates along these
routes (Bellrose ).
International migrants have been much more difficult to trace, but recent
efforts of several shorebird study groups have resulted in the International
Shorebird Survey and many subgroups have a long history of effort for smaller
areas. Numerically, these bird groups represent one of the greatest challenges
because of numbers running into the millions at some sites like the Bay of
Fundy and areas of similar tidal extremes in Tierra del Fuego, with a complex
mix of species as well.
At the other extreme are small-scale but habitat-specific censuses that have
been done by boat along lakeshores to associate wader species with specific
habitats (Esler ), or along coastal islands for species and population
counts, but such access is not always available in backwater parts of riverine
systems. Counts from boats often are less disturbing than ground surveys and
can be repeated regularly with little apparent population effect. Ground
counts of nests have been used for some waders, especially those that are solitary or less visible; belt transects often are useful to minimize the labor but
assure coverage of various vegetation types (Weller and Frederickson ).
Radio-tracking of individual birds in flocks using Global Positioning Systems
now allows pinpointing of microsites, temporal data on individuals, and
meaningful estimates of turnover time, which is essential to estimate real
numbers of birds using a site.
The above very incomplete arrays of techniques and issues may be both
complicating and discouraging for those who are searching for simple techniques to answer important questions; unfortunately, such techniques do not
exist. Choice of technique is dependent upon the questions asked, and the time
and money available to gather the information at a satisfactory level. Some
census techniques have been refined for years or more and are little more
accurate than when first developed, but they now form important indices.
Each study is different, and the suggestions I outlined earlier can help to avoid
useless data. Statistical requirements complicate data gathering even more and
may result in a failure to address the specific question asked. Knowledge of the
bird life histories and habitat requirements are essential first steps for ensuring
good questions, the development of practical techniques, and thereby useful
answers; however these answers are not always at the level of detail needed or
have the accuracy desired. Whatever is done, measurement of population and
species diversity without information on habitats and their status will weaken
the understanding of the community in question and the opportunity to apply
such population information to conservation assessments.
these features, and it is especially frustrating that one can measure many things
and still not find the suitable correlate! Moreover, what is important in one
season may be meaningless in another.
Horizontal distribution of vegetation and other structural features is the
most simple and currently easiest parameter to identify because of the availability of aerial photographs or satellite imagery, computer scanning, and
Geographic Information Systems, but these measurements may not demonstrate the expected correlation. Typically, one seeks to measure heterogeneity
in horizontal or spatial patterns of coverwater, plant diversity, etc. that
influences habitat structure. Size of used habitats (patch size) and their distribution or dispersion are important components used as correlates for species
or community distribution and abundance. These data have been most often
used in calculating coverwater or covercover ratios, amount of edge, and
other structural indices that can be correlated with species presence and
density (Beecher , Weller and Spatcher ). These do not necessarily
indicate the cause of the relationship but do lead one closer to finding it.
Measurements and analyses of data on vertical structure of habitat diversity are more difficult. These usually have compared things like nest height in
relation to plant structure (e.g., height, stem thickness and density, foods,
water, etc.) compared with that in unused plants. Rarely have these analyses in
wetlands incorporated the features of the water column or substrate
characteristics.
Habitat patterns derived from a few parameters of simple systems like
urban and forest-edge habitats (DeGraaf ) combined with computerbased models of pattern recognition have been used to assess and predict the
presence and relative abundance of selected bird species for forest birds
(Grubb , Williams, Russel and Seitz ). Such patterns have not been
fully exploited for wetland birds except in relation to a diversity index
(Adamus ), but the generalized habitat patterns used earlier to describe
wetland birds have potential for such conceptual and perhaps mathematical
models. Features other than structure have been used to explain the presence
and density of selected species, such as food (Chapters and ), water depth,
and water quality (Utschick ).
The Habitat Evaluation Procedure (HEP) is used by US federal agencies
such as the Fish andWildlife Service in assessing wetland habitat loss or damage.
The relative quality of the habitat is based on a simple model called the Habitat
Suitability Index (HSI),which uses a few parameters that seem to correlate with
a species presence, abundance, and reproductive success. Such indices have
been created for wetland birds as well as for fish, mammals, and invertebrates;
these are index ratios with a maximal value of .. These quality estimates then
can be used to specify the amount of replacement habitat to compensate for lost
habitat but based on quality. If, for example, acres of wetland were to be
destroyed by a highway development project which had an HSI of . for a
certain species (or pooled ratio for several key species), . = habitat
units (HU) of HSI . value would be required to replace this habitat. If lower
quality habitat (HSI) is used for mitigation (HSI = .), it would require more
acreage to replace that lost: HU/. HSI = acres. One can also use this
system to document changes over time, or compare two communities, but the
system has been used more for single species than for groups.
An analysis based upon the incidence of bird species derived by surveys
which are then clustered according to the type and manner in which they use
resources (i.e., guilds) stresses the importance of habitat structural diversity,
food resources,and water regimes (Short and Burnham ).Life-history data
of members of an assemblage or community are analyzed as a guild matrix
according to a series of habitat features viewed as resources for some function
like nesting plotted against another resource use such as feeding. Occupied
blocks in the matrix reflect species using those guilds, allowing a visual assessment of the complexity of a community based on the diversity of resources.
The resulting guild matrix also is a very useful approach to documenting
changes over time resulting from either natural events or human impacts and
results in a conceptual model of the habitat pattern that allows prediction of
species composition on the basis of habitat features (Short ) (Fig. .).
Several broader and less bird-oriented techniques have been developed
and used by various agencies to assess the overall ecosystem or community
components, at least on a relative scale. The system developed for the US
Federal Highway Administration (Adamus et al. , ) is among the best
because it qualitatively judges or so parameters of the wetland and its components; it uses waterbirds in particular because they indirectly reflect important habitat quality features that are important to wetland functions other
than wildlife habitat (e.g., water purification, sediment trapping, erosion
control, fish habitat). The Adamus evaluation system is hierarchical, allowing
the user to incorporate any level of detail desirable, based on available knowledge or for which funds are available.
New approaches are constantly being developed, often to reduce time and
costs, but it should be apparent that these will fail if not based on a thorough
understanding of the manner in which various bird species use such habitats
for food or nest sites, etc. and of the influence of habitat diversity and dynamics on populations and species composition. From a practical standpoint, it
may be impossible to gather all the desired information, but one often can
exploit existing biological data, perhaps gathered for other purposes, that may
be useful in conservation and other ecological applications.
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Smith, R. D. (). Wetland Evaluation Technique (WET), Vol. I. Technical
Report WRP-DE-. Vicksburg, MS: U S Army Corps of Engineers, Waterways
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York: Wiley.
Beecher, W. J. (). Nesting birds and the vegetative substrate. Chicago, IL: Chicago
Ornithological Society.
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Colwell, M. A. and Cooper. R. J. (). Estimates of coastal shorebird abundance:
the importance of multiple counts. Journal of Field Ornithology , .
Cooke, W. W. (). Distribution and migration of North American ducks, geese and
swans. Biological Survey Bulletin No. . Washington, DC: U S Department of
Agriculture.
Cowardin, L. M. and Blohm, R. J. (). Breeding population inventories and measures of recruitment. In Ecology and management of breeding waterfowl, eds.
B. D. J. Batt, A. D. Afton, M. G. Anderson, C. D. Ankney, D. H. Johnson, J. A.
Kadlec, and G. L. Krapu, pp.. Minneapolis, MN: University of Minnesota
Press.
DeGraaf, R. M. (). Urban bird habitat relationships: application to landscape
design. Transactions of the North American Wildlife & Natural Resources
Conference , .
Esler, D. (). Habitat use by piscivorous birds in a power plant cooling reservoir.
Journal of Field Ornithology , .
Frederick, P. C., Towles, T., Sawicki, R. J., and Bancroft, G. T. (). Comparison
of aerial and ground techniques for discovery and census of wading bird
(Ciconiiformes) nesting colonies. Condor , .
Gibbs, J. P. and Melvin, S. M. (). Call-response surveys for monitoring breeding
waterbirds. Journal of Wildlife Management , .
Glahn, J. F. (). Study of breeding rails with recorded calls in North-Central
Colorado. Wilson Bulletin , .
Grubb, T. G. (). Pattern recognition a simple model for evaluating wildlife
habitat. US Forest Service Rocky Mountain Experiment Station Research Note
RM-.
Hall, L. S., Krausman, P. R., and Morrison, M. L. (). The habitat concept and a
plea for standard terminology. Wildlife Society Bulletin , .
Igl, L. D. and Johnson, D. H. (). Changes in breeding bird populations in North
Dakota: . Auk , .
James, F. C. and Rathbun, S. (). Rarefaction, relative abundance, and diversity of
avian communities. Auk , .
Johnson, D. H. (). The comparison of usage and availability measurements for
evaluating resource preference. Ecology , .
King, K. A. (). Colonial wading bird survey and census techniques. In Wading
birds, eds. A. C. Sprunt IV, J. C. Ogden, and S. Winckler, pp. . Research
Report . New York: National Audubon Society.
Kushlan, J. A. (). Effects of helicopter censuses on wading bird colonies. Journal
of Wildlife Management , .
References
Tacha, T. and Braun, C. (). Migratory shore and upland game bird management
in North America. Washington, DC: International Fish and Wildlife Agencies.
Utschick, H. (). Die Wasservogel als Indikatoren fr den okologischen Zustand
von Seen (with English summary). Verhandlungen der Ornithologischen
Gesellschaft in Bayern (/), .
van Horne, B. (). Density as a misleading indicator of habitat quality. Journal of
Wildlife Management , .
Vickery, P D., Hunter, M. L. Jr, and Wells, J. V. (). Use of a new reproductive index
to evaluate relationship between habitat quality and breeding success. Auk ,
.
Weller, M. W. (). Use of two waterbird guilds as evaluation tools for the
Kissimmee River restoration. Restoration Ecology , .
Weller, M. W. and Fredrickson, L. H. (). Avian ecology of a managed glacial
marsh. Living Bird , .
Weller, M. W. and Spatcher, C. E. (). Role of habitat in the distribution and abundance of marsh birds. Iowa State University Agriculture & Home Economics
Experiment Station Special Report , .
Williams, G. L., Russell, K. R. and Seitz, W. (). Pattern recognition as a tool in the
ecological analysis of habitat. In Classification inventory, and analysis of fish and
wildlife habitat: proceedings of a national symposium; Phoenix, AZ, pp. .
Washington, DC: U S Fish & Wildlife Service.
Further reading
Bellrose, F. C. (). Waterfowl migration corridors. Illinois Natural History Survey
Biological Notes , .
Cooke, W. W. (). Distribution and migration of North American rails and their
allies. Biological Survey Bulletin No. . Washington, DC: US Department of
Agriculture.
Furness, R. W. and Greenwood, J. J. D. (eds.) (). Birds as monitors of environmental change. London: Chapman & Hall.
Johnson, D. H. and Grier, J. W. (). Determinants of breeding distribution of
ducks. Wildlife Monographs , .
Maurer, B. A. (). Predicting habitat quality for grassland birds using
densityhabitat correlations. Journal of Wildlife Management , .
Mikol, S. A. (). Field guidelines for using transects to sample nongame bird
populations. Publication /. Washington, DC: U S Fish & Wildlife Service
Office of Biological Services.
Reichholf, J. H. (). The possible use of the aquatic bird communities as indicators
for the ecological conditions of wetlands. Landschaftsrerband Stadt , .
Root, R. B. (). The niche exploitation pattern of the bluegray gnatcatcher.
Ecological Monographs , .
Schamberger, M. and Farmer, A. (). The habitat evaluation procedures: their
application in project planning and impact evaluation. Transactions of the
North American Wildlife and Natural Resources Conference , .
Weller, M. W. (). Issues and approaches in assessing cumulative impacts on
wildlife habitat in wetlands. Environmental Management , .
14
Current status and some
conservation problems
. Wetland losses
To appreciate the current status of wetland habitats for birds, we need to
examine briefly how society has dealt with wetland habitats in most of the
world. A number of early cultures learned to live with and in wetlands for their
resources, for example the marsh Arabs of Iraq, where massive living quarters
once were built from reeds (Maxwell ); the Uro Indians of Lake Titicaca in
Peru, who built small houses on floating mats and constructed their famous
Wetland losses
canoes of bulrushes; the boat dwellers of Pakistan marshes who fish and also
use mounted waterbirds on their heads to hide their underwater approach to
living birds, much as the ancient Egyptians did; the Calusa Indians of the
Everglades (Douglas ) and other coastal tribes of the Gulf of Mexico, who
used clams, fish, and water plants as their main food; and the tribes of the oncevast Stillwater Marsh and Carson Sink in the Great Basin of the western USA
(Larson and Kelly ), well-known for their beautiful duck decoys made of
emergent marsh plants like California bulrush or tule (Earnest ). Many
river societies of South America, Asia, and Africa have devised mobile lifestyles dictated by flood waters, and housing designs to minimize loss or
damage. All societies have used vertebrate resources of wetlands and some still
do: hunting, trapping, fishing, clamming, using ducks domesticated for egg or
meat production, and some have used birds as watchdogs (Southern
Screamer of South America). Fish-dependent cultures like those of the Orient
and Middle East have for centuries used trained cormorants or herons to
capture food rather than using the less dependable arrow, spear, or fish-hook.
These reflect the early origins of humans as functional entities in natural community relationships, an example of which is generalized in a food web of the
Kafue River flats of Zambia (Fig. .) (Maltby ).
Most well-developed societies have tried to eliminate wetlands through
drainage ditches and have modified water flow via dams or diversion structures for reasons of convenience, disease control, and re-direction of the water
for use in irrigated agriculture, urbanization, and industry (Dugan ).
Wetland loss worldwide has been calamitous but it is poorly documented in
most countries. Considering that wetlands constituted only % of the land
area in most regions of the contiguous USA, why are wetlands so in demand
and why are we so often in conflict with such natural land features? And why,
after eliminating about % of the original wetlands, are we still exerting pressure to change or eliminate wetland? Moreover, some parts of the world exceed
this figure, having lost to % (Dahl , Dahl, Johnson and Frayer ,
Maltby , Tiner ). In terms of major bird habitats, only grasslands have
suffered greater loss in area or quality in North America and probably around
the world.
Pressures on wetlands varies by region, land-use, and human perspective.
In agricultural areas around the world, it is often an economic need by individuals and families to maximize earnings from every acre of available land, but
these efforts vary in their ecological and economic efficiency. In major urban
areas, losses result from the association of developments with water: rivers,
lakes, and coastlines. As cities grow, choice land for continued development is
no longer available, areas considered of low-valueare used for commercial or
industrial developments or built-up for housing by filling. Even if maintained
as wet areas by necessity, they often are modified by impoundments for water
storage or dedicated as intensively managed city and county parks, no longer
performing the multiple functions of dynamic natural systems. Therefore,
Figure .. Relationships
between primary production
of hydrophytes and waterbirds,
grazing ungulates, and humans
in the Kafue River floodplain,
Zambia (Maltby , with
permission).
many of our urban and suburban nature preserves are relicts of a once
common habitat type.
Appreciation of wetland values has come slowly to most developed
nations, initially as part of the conservation movement and more recently
through recognition of the ecological and economic values of wetland ecosystems (Good, Whigham and Simpson , Gosselink, Odum and Pope ,
Greeson, Clark and Clark , Maltby ). Interest in preservation of
unique lands in North America arose during the late s and early s after
recognition and concern over the loss of rare species. Much of the motivation
was prompted by loss of large mammals and terrestrial birds, for example
Passenger Pigeons, which concerned hunters. However, declines of waterbirds
such as Eskimo Curlew, Great Auk, Labrador Duck, Trumpeter Swan,
Whooping Crane (Forbush ) and the Ivory-billed Woodpecker became of
special interest with the potential damage threatened by passage of the United
States Reclamation Act in . This Act was directed toward a national survey
Wetland losses
of watery wastelands that could be re-claimed, mainly to encourage drainage, regulation, and impounding of streams, and for water use in irrigation.
One secondary result was an information base on the extent of wetlands in that
period, from which we now estimate losses. Similar actions have occurred
around the world, often much earlier than in North America.
Drainage of larger and deeper wetlands has required major engineering
efforts, suitable topography, or pump technology. Yes, some wetland drainage
schemes actually have failed because people tried to drain them uphill! Much
of this technology was developed in the Netherlands and the United Kingdom
for land reclamation or in Asia for rice production but has been transported
around the world. In many cases, drainage water has been moved via rivers and
into the sea, but in more inland settings, drainage typically flows from smaller
areas to large collecting basins, lakes, or sumps. Therefore, enlargement of wetlands or stabilization of lakes as reservoirs have been the products of drainage
or irrigation programs, all enhanced by the development of huge earthmoving equipment allowing massive land changes heretofore impossible.
Losses also have been facilitated by our failure to develop national land management goals that recognize the long-term wetland values of flood control
and aquatic resources compared with immediate or short-range economic
benefits. There are numerous examples of how drainage systems have worsened riverine and urban flooding regimes and have been destructive to wetlands (Miller and Nudds ), and how their present maintenance continues
to have an impact on wetland quality and predictability with significant effects
on waterbirds (Krapu ).
The Florida Everglades, perhaps the largest wetland in North America, is a
product of rivers like the Kissimmee or a river-like sheetwater flow from north
to south. It has been modified by impoundments for water supply or alternative uses, and the northern segment now is a series of three large impoundments that protect nearby developments from flooding and ensure urban
water supplies and agricultural development. In some areas, this has changed
the character of the wetland from seasonally flooded to semipermanent water
and changed all the organisms and birds therein. Other areas were exposed for
sugar cane and other crops and have demonstrated major subsidence because
of the decomposition of organic matter. Enrichment and sedimentation have
been major problems on large river impoundments, many of which function
as wetlands in the more shallow portions. In addition, poor land management
upslope has exacerbated the problem, seriously affecting all aquatic resources
and waterbirds in riverine wetlands (Bellrose, Paveglio and Steffeck ) and
estuaries (Perry and Deller ).
Although river impoundments currently are less commonly supported by
government agencies in developed nations, the economies of some nations
seem focused on construction of ever-more massive impoundments (China,
Canada), reflected also in long-range planning (South America and Africa).
This may bring more water, electrical power, and perhaps food for people,
agriculture, and industry, but it will flood wetlands and other lowland natural
habitats and will stabilize water levels with a strong probability of a net reduction in productivity of most water resources.
Coastal tidal impoundments dedicated to expanding reclamation of land
for agriculture or for energy projects are increasing as land values rise and
technology allows; these will have major impacts on saltmarsh and mudflats
vital to many local and global bird species (Goss-Custard and Yates ).
There have been some major and many seemingly minor but ubiquitous
dangers that confront waterbirds daily. Highly toxic and long-lived pesticides
were early recognized as serious in waterbirds because of their concentration
especially in fish of coastal marine systems. As a result, of this concentration
Brown Pelicans in North America suffered major losses and were nearly eliminated from the Gulf of Mexico and the Pacific Coast breeding areas, whereas
interior- nesting White Pelicans in non-agricultural areas were less affected.
Although many of the more serious pesticides have been banned or closely
regulated in many countries, others are widespread in third-world nations.
Interior basins fed by large streams, such as the worlds largest freshwater
wetland, the Pantanal of Brazil (Alho, Lacher and Gonclaves ), have
suffered from heavy use of agricultural pesticides, which has resulted in
modified fish and invertebrate communities.
Oil and gas development continues, especially in remote and often delicate
ecosystems around the world, and there are numerous cases of small but
locally serious impacts on birds (see Burger ). Waste deposition of oil,
caustic salts, chemicals, and other substances harmful to birds are surprisingly
widespread and poorly controlled and very attractive to waterbirds when in a
fluid state.
Line fishing with exposed hooks is deadly to diving birds in fresh and saltwater, and net fishing can be serious for diving birds like loons, grebes, and
ducks (Bartonek , Schorger ). Because of nocturnal migration, often
over land, collisions with human structures is common, including fences and
telephone lines (Cornwell ) and powerlines (Malcolm ).Although lead
shotgun pellets have been banned for hunting waterfowl over water in the
USA, the residual lead in the substrates still accounts for % of waterfowl
mortality in the whole country, and more in localized areas (Anderson and
Havera , Sanderson and Bellrose ). Any species that strains bottom
oozes for grit is subject to lead intake, as recorded in flamingos in Mexico and
Europe (Schmitz et al. ). Moreover, the use of lead for shotguns is still
widespread in many areas of the world.
Botulism and other natural diseases are still poorly understood but strike
unexpectedly, with huge and uncontrollable mortality, as in the loss of
pelicans in the Salton Sea of California (Anonymous ), an area where in
past years major losses of grebes have occurred. Birdaircraft strikes are still
more common than they should be and often involve waterbirds because of
their flock and roosting behavior (Holden ), or because they become concentrated in certain areas in response to societal strategies like landfills (gulls)
or protected breeding areas (Canada Geese, Cooper ).
Fish production, sport fishing, and introduction of exotic fish have
resulted in direct impacts on habitat through introduction of herbivores like
carp, and through competition for food of native birds (Bouffard and Hanson
, Weller ) from exotics and trout reared in wetlands. A related topic is
the intentional persecution of cormorants and other fish-eating birds
. Unwanted biodiversity
Society has had another major impact on avian diversity and plant composition in wetlands, as in other habitats, by directly importing foreign (exotic)
birds and other animals and plants, or by inducing unwanted mobility or
range expansion via drainage and other land-use changes. As a result, some
wetland birds occur now where they once did not. Most conservationists
strongly oppose intentional releases of exotic birds because of the potential for
interspecific competition with impact on native species, such as the classic
examples of the Starling, House Sparrow, and Common Pigeon. But such
events have occurred with wetland species, sometimes with the approval of
well-meaning but poorly informed conservationists. Most intentional releases
have been of waterfowl intended for either hunting or esthetic purposes. For
example, introduction of Mallard ducks occurred for hunting purposes in
many areas: in New Zealand they hybridized with the native Grey Duck, in
Hawaii they interbred with Hawaiian Ducks (Weller ), and in the eastern
USA interbreeding with local Black Ducks is widespread. Fortunately,
Mallards introduced in south Texas, where potential interbreeding with local
Mottled Ducks might have been a problem, have not survived well and now are
rare (Kiel ). They also have disappeared from the Falkland Islands, where
competition with other dabbling ducks might have been a problem, but they
have been introduced into Argentina.
For esthetic purposes, Mute Swans from Europe have been introduced or
escaped from captivity widely around the world because of their use in cemetery lakes for atmosphere. Free-living populations now occur in Toronto, the
northwestern and eastern coasts of the USA, and in New Zealand, to mention
but a few. At least one study has found no apparent negative impacts on local
species (Conover and Kania ). Black Swans have been introduced from
Australia to New Zealand and have been very successful and are now abundant
there. A similar situation holds for Canada Geese intentionally introduced and
now widespread in New Zealand, but opinion varies between hunters, farmers,
and conservationists as to their merits. In North America, Canada Geese
released in urban areas and refuges for restoration and esthetic purposes have
been protected by lack of hunting and artificial feeding so that they become
nuisance birds. Moreover, such introductions have often resulted in mixing of
genetic strains, which has confounded the original distribution patterns of
geographic races.
The desire of aviculturists to keep and rear exotic waterfowl has had mixed
blessings. Birders now constantly report accidental or very rare ducks more
commonly, and escapes from captivity are suspected in many cases. The
magnificent Mandarin Duck of the Orient has been a favorite show bird in
many collections and escaped from captivity in England long ago. It now is
free-living over a wide area there and more recently has been free-living in
California as well (Shurtleff and Savage ). Fortunately, it does not seem to
hybridize with its closest relative, the North American Wood Duck. A more
serious recent example of the potential consequence of escapees is the North
American Ruddy Duck in England. Its beautiful colors and unique behavior
made it an attractive bird to show; the challenge of rearing young resulted in
captive birds being imported to the UK, where they subsequently escaped in
. By , the Ruddy Duck had spread to European countries and
populations are still steadily increasing. Unfortunately, it is an aggressive
species and cross-mates with the European White-headed Duck (another
stifftail), which is already in trouble because of habitat loss (Johnsgard and
Carbonell ).
As mentioned above, the natural range of the Cattle Egret is now nearly
worldwide in tropical and warm-temperate areas. Apparently, efforts have
been made to introduce it on islands like Hawaii and elsewhere, but most of its
success seems to result from its own pioneering behavior. As its name implies,
it feeds at the feet of feeding livestock and large wild ungulates, where it gathers
insects and other small prey stirred by their movement. It also feeds in newly
plowed fields, often following the tractor like gulls and harriers. Therefore, it
occupies a new or expanded habitat created by agriculture. It has since
expanded into heronries throughout the USA and makes an annual migration
into Mexico, a pattern than has evolved since the early s. Although it does
not seem to compete directly for food (at least with wetland species), some
observers speculate that its dominance of nest sites could adversely affect other
species in mixed colonies.
It is obvious that introductions can produce serious direct results and have
the potential of carrying diseases and competing in ways we cannot easily
measure. Moreover, it modifies natural and long-established bird communities in ways that we have not yet adequately assessed. Although such releases
may be interesting experimentally, they are poorly controlled and should not
be condoned. The International Treaty on Biodiversity incorporates a statement on limits and controls of invaders and is encouraging prompt and meaningful policies and control methods (Baskin ).
Indirectly, introductions of other exotic species can have important
Unwanted biodiversity
Cooper, J. A. and Keefe, T. Managing urban Canada Geese: policies and procedures.
Transactions of the North American Wildlife & Natural Resources Conference ,
.
Cornwell, G. (). Collisions with wires a source of anatid mortality. Wilson
Bulletin , .
Custer, C. M. and Custer, T. W. (). Food habits of diving ducks in the Great Lakes
after the zebra mussel invasion. Journal of Field Ornithology , .
Dahl, T. E. (). Wetland losses in the United States s to s. Washington DC:
US Fish & Wildlife Service.
Dahl, T. E., Johnson, C. E., and Frayer, W. E. (). Wetlands status and trends in the
coterminous United States mid-s to mid s. Washington DC: US Fish and
Wildlife Service.
Dahlgren, R. B. and Korschgen, C. E. (). Human disturbance of waterfowl: an
annotated bibliography. Resource Publication . Washington DC: U S Fish &
Wildlife Service.
Des Granges, J. L. and Hunter, M. L. Jr. (). Duckling response to lake
acidification. Transactions of the North American Wildlife & Natural Resources
Conference , .
Douglas, M. S. (). The everglades: river of grass. St Simons, GA: Mockingbird
Books.
Drennan, S. R. (). Horseshoe crab decline in Delaware Bay. National Audubon
Society Field-Notes , .
Dugan, P. J. (ed.) (). Wetland conservation, a review of current issues and required
action. Gland, Switzerland: World Conservation Union, IUCN.
Earnest, A. (165). The art of the decoy: American bird carvings. New York: Bramhall
House.
Esler, D. (). Waterfowl habitat use on a Texas reservoir with Hydrilla. Proceedings
of the Annual Conference of the Southeastern Association of Fish and Wildlife
Agencies : .
Forbush, E. H. (). A history of the game birds, wild-fowl and shore birds of
Massachusetts and adjacent states, nd edn. Boston: Massachusetts State Board
of Agriculture.
Gilbert, D. W., Anderson, D. R., Ringelman, J. K., & Szymczak, M. R. (). Response
of nesting ducks to habitat management on the Monte Vista National Wildlife
Refuge, Colorado. Wildlife Monographs , .
Good, R. E., Whigham, D. F., and Simpson, R. L.(eds.) (). Freshwater wetlands,
ecological processes and management potential. New York: Academic Press.
Goss-Custard, J. D. and Yates, M. G. (). Towards predicting the effect of saltmarsh reclamation on feeding bird numbers on the Wash. Journal of Applied
Ecology , .
Gosselink, J. G., Odum, E. P,. and Pope, R. M. (). The value of a tidal marsh. Baton
Rouge, LA: Center for Wetland resources, Louisiana State University.
Greeson, H., Clark, J., and Clark, J. (eds.) (). Wetland functions and values: the
state of our understanding. Minneapolis, MN: American Water Resources
Association.
Hamilton, D. J. and Ankney, C. D. (). Consumption of zebra mussels Dreissena
polymorpha by diving ducks in Lakes Erie and St Clair. Wildfowl , .
References
Harrington, B. (). The flight of the Red Knot. New York: W. W. Norton,
Holden, C. (). Eco-solution to airport bird pests. Science , .
Johnsgard, P. A. and Carbonell, M. (). Ruddy ducks and other stifftails. Norman,
OK: University of Oklahoma Press.
Kiel, W. H. Jr (). A release of hand-reared mallards in South Texas. Publication
MP. College Station, TX: Texas A&M University Agricultural Experiment
Station.
Klein, M. L. (). Waterbird behavioral response to human disturbances. Wildlife
Society Bulletin , .
Klein, M. L., Humphrey, S. R., and Percival, F. (). Effects of ecotourism on
distribution of waterbirds in a wildlife refuge. Conservation Biology , .
Krapu, G. L. (). Effects of a legal drain clean-out on wetlands and waterbirds:
A recent case history. Wetlands , .
Larson, C. S. and Kelly, R. L. (). Bioarchaeology of the Stillwater Marsh, prehistoric human adaptations in the western Great Basin. American Museum of
Natural History, Anthropology Papers , .
Ludyanskiy, M. L., McDonald, D., and MacNeil, D. (). Impact of the zebra
mussel, a bivalve invader. BioScience , .
Luo, H., Smith, L. M., Allen, B. L., and Haukos, D. A. (). Effects of sedimentation
on playa wetland volume. Ecological Applications , .
Malcolm, J. M. (). Bird collisions with a power transmission line and their relation to botulism at a Montana wetland. Wildlife Society Bulletin , .
Maltby, E. (). Waterlogged wealth. London: Earthscan. International Institute
for Environment and Development.
Maxwell, G. (). People of the reeds. New York: Harper and Row.
Miller, M. W. (). Route selection to minimize helicopter disturbance of molting
Pacific Black Brant: a simulation. Arctic , .
Miller, M. W. and Nudds, T. D. (). Prairie landscape change and flooding in the
Mississippi River Valley. Conservation Biology , .
Myers, J. P., Morrison, R. G., Antas, P. Z., Harringon, B. A., Lovejoy, T. E., Sallaberry,
M., Senner, S. E., and Tarak, A. (). Conservation strategies for migratory
species. American Scientist , .
National Research Council (). Irrigation-induced water quality problems.
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biology, conservation and management. Colonial Waterbirds , .
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Further reading
15
Conservation and management strategies
The existing problems that wetland birds face and the potential applications of
ecological data must be considered in the light of current structure and
approaches in wildlife conservation. Historically, conservation and management efforts have focused on legal protection, habitat management, population management, and human dimensions, and all must be used concurrently
to be effective. Our focus here is habitat management, which has been used for
different purposes but is often aimed at species management when, at least for
wetlands, most management influences the entire community.
Early efforts of habitat preservation in the USA, and probably in most
countries, focused on wetland habitat protection. Direct purchase for refuges,
sanctuaries, and preserves has been common, but leasing and legally binding
conservation easements and agreements have been more common recently as
a means of controlling more areas at lower cost per unit area, especially when
multiple ownership is involved. Protection of small basins or patches of
habitat has worked well for wetlands because many birds are habitat-island
species that use several habitat units and fly between them. In the USA, there
are over National Wildlife Refuges operated by the US Fish and Wildlife
Service involving over million acres, most of which include major wetlands
(Riley and Riley ). Other federal agencies such as the Forest Service and
Bureau of Land Management also hold title to and manage similar areas, and
the Natural Resource Conservation Service program accomplishes similar
protection on private lands through the Soil Conservation Reserve Program
(Johnson and Igl ). Many states, counties and cities have protected additional and generally smaller but no-less valuable areas through refuges, public
hunting areas, nature centers, and green belts. Similar strategies have occurred
around the world, but, in most areas, these constitute a small percentage of the
original area that once supported wetland bird diversity and numbers.
However, these wetlands often lack surrounding upland vegetation so important to water quality (e.g., sediment control) and for nesting habitat for birds
that use wetland margins. Moreover, they tend to represent the typical or
expected examples rather than the diversity of habitats essential to attracting
different species of bird.
Land acquisition for conservation of wildlife sometimes has involved land-
more important than putting all our eggs in one or a few baskets. However, it is
essential to preserve major migration stops for large numbers of long-range
migrants as well, and the international Shorebird Reserve Program has
identified and is working toward such measures. Other examples exist
throughout the world, reflecting the interest and zeal of amateur as well as professional bird students.
The term biodiversity has been applied broadly to an overall conservation
goal for society and as a technique for identifying key areas based on species
richness of all taxonomic entities and the supporting ecosystem structure
(Noss and Cooperrider ). Computer-based geographic information
systems and data bases on species distribution allow the identification of
species-rich areas that have few or no preserves, an approach termed gap
analysis (Kiester et al. , Scott et al. ). This system is also being used on
wetland animals, but it seems better suited to larger scales (Noss and
Cooperrider ). A currently popular index for selecting important areas to
conserve is based on species-rich areas (often termed hotspots) (Dobson et
al. ), but such areas are more difficult to identify when dealing with variable environments like wetlands and migratory or even nomadic species. Not
everyone is convinced that simple measures of biodiversity are adequate for
major conservation decisions that could affect worldwide conservation policies. Part of the weakness of the system seems to be that ecosystems are not consistent in the relationship of number of species to uniqueness, and with
wetlands, they are not constant over time. Several studies have concluded that
assessing the priority of differing wetlands for preservation based on species
richness may not identify the important habitats for those bird species that
have narrow or precise requirements, and for which no other habitats are suitable (Burger, Schisler and Lesser , Weller ). Moreover, because many
wetlands vary markedly in water regimes, they may be under-valued if assessed
by a few surveys at a poor time when they actually fill an important need such as
for roosting habitat, with other functions such as feeding or breeding occurring elsewhere. Hence, our analyses often must treat wetlands as separate
habitat units that serve a bird species or group, recognizing the spatial and temporal aspects of these components of an annual habitat that fulfill different
functions and in different areas. Additionally, some workers stress that preserving a natural area of any size is still an ideal goal when no other such natural
area exists (Shafer ) as is often the case for isolated or urban wetlands.
Once an area of habitat is acquired, management too-often fails because
the processes that drive the system often are unknown, or management strategies are applied in a rigid manner. In my experience, wetlands with watercontrol structures tend to have water gauges, whereas naturally controlled
ones rarely do even if under study.Without being able to relate wetland vegetation and bird use to water regimes, we cannot understand regulators in the
system, we have no measure of suitable and optimal conditions, and, therefore,
we cannot set realistic goals or measure our success at meeting these. Another
failing has been what I call the pick-up truck syndrome, which induces a
view from afar when one needs a hip- boot approach. Regular monitoring
and corrective action are essential. Moreover, with wetlands, this requires a
longer-term view than in terrestrial systems, because water regulation must be
planned well in advance to induce successional changes that lead to changes in
bird habitat (Weller ). Such adaptive management must be dynamic,
changing as needs demand to meet stated goals. However, it involves carefully
planned objectives, experimental approaches to ensure reasonable answers,
and a monitoring system than provides feedback and allows change toward a
better final product (Holling , Walters ).
15.1.1 avian habitat management in existing wetlands
Wetland management for birds and other wildlife must take a community and
ecosystem perspective, recognizing the influences of most actions on species
that are not major targets. Moreover, process-oriented management involving
drivers of the system (e.g., water regime and fire) that influence nutrient
dynamics and natural plant succession is the most ecologically and economically sound approach. In addition to the manipulation of physical processes,
grazers like Muskrats, Nutria, geese, Beaver, Marsh Deer, Water Buffalo, and
domestic livestock can have major impacts (Weller ). Large numbers of
herbivorous fish alter vegetation, and predatory fish modify invertebrate
populations and thereby bird use, but this is less measurable than are the positive effects of large numbers of fish available as food for egrets and herons.
Undoubtedly, these food-web interactions are essential when devising techniques needed to manipulate wetlands for optimal diversity and production.
15.1.2 restoration, enhancement, and creation of wetlands
Much of the management of wetlands today, regardless of objective and
wetland type, is an effort to improve habitat within and around the wetland for
maximal production and diversity of birds or other wildlife. This is termed
enhancement (Weller ) and may involve manipulating water regimes,
water quality, and vegetation through methods described above. Such actions
may be legally demanded in cases of mitigation for wetland losses, but in most
cases, the approach is used by private, state, and federal managers attempting
to compensate for lost or impacted wetlands nearby. The techniques may be
less drastic than in restoration, but the degree of improvement also is less dramatic and, therefore, not easily measured.
The most dramatic and measurable results are seen when returning a
former wetland to its previous productive state, termed restoration. In some
cases, the heritage of the area may only be discernable via the presence of a
wetland seed bank, a pool of long-lived residual seeds in the substrate that will
germinate when water is returned and maintained at a suitable level (van der
CHANNELIZED
RESTORED/NATURAL
DEEP CHANNEL
NARROW FLOODPLAIN
LOW HABITAT DIVERSITY
REDUCED FOOD DIVERSITY
VARIABLE DEPTH
BROAD FLOODPLAIN
HIGH HABITAT DIVERSITY
HIGH FOOD DIVERSITY
EDGE SPECIES
STABLE WATER DEPTH
OPEN WATER/SPARSE EMERGENTS
ers often are willing to pay the high price to mitigate for wetlands destroyed or
damaged as a result of other uses. Creation also occurs when someone wants to
develop a wetland habitat at a specific site to attract birds or to perform other
wetland functions such as water purification. It requires the establishment of a
suitable water source and hydraulic regime that will support artificially seeded
or planted vegetation. Successful developments are those where the water
problem is resolved before the structure is built. In strip-mine areas, where we
have studied plant response to areas virtually free of seed banks (owing to soilhandling techniques), natural seeding of common wetland annuals, submergent vegetation, and shoreline willows requires from to years for
establishment of the most minimal wetland plant assemblages; similar timing
of plant and bird occupancy has also been noted in restored wetlands in Iowa
(Van Rees-Siewert and Dinsmore ). Efforts to create small basins in
Wisconsin showed very slow invasion of native plant, and seeding was used to
enhance diversity within years (Reinartz and Warne ). Some invertebrates invade in a matter of weeks or months, and herptiles and fish are highly
variable and dependent on connectivity to other wetlands (Weller ). These
in turn dictate whether, how, and when they will be used by waterbirds.
Despite our good fortune in that many wetlands can be managed, restored,
and even created, we should not be too complacent nor feel we can be liberal in
destroying the natural and old in place of the new and the young.There is much
we do not know, but it is certain that the complexity of the more sustained
wetland system is great and that the pioneering successional stages are those we
most readily duplicate. Birds are but one component of this complex system
and are clearly successful because they are so adaptable.Insects and crustaceans
may be equally so but many other animals are not. Moreover, we typically deal
with a fragment of the long time scale or the vast geographic sphere of mobile
waterbirds.From all that has been observed and measured,we assume that each
wetland makes a difference, that each influences birds locally, and that often
birds move long distances. Future studies will need to determine the reality of
these and similar assumptions, but until we know more, conservation of all
types of wetland in all places is the only safe strategy for the conservation of
wetland birds. As in most things, what is good for natural systems is good for
society; it just takes a little longer to understand and appreciate it!
the world. However, the usual situation was not prediction but rather
demonstration of the effects for legal purposes, such as a demand for mitigated
replacements, and this was only possible in those rare cases where pre- and
post-impact data were available. The meandering and unique Kissimmee
River in central Florida is a classic example of a river channelized to minimize
flooding (Blake ) during the period when environmental awareness was
increasing. The result was an environmental impact assessment that predicted
devastating effects on the abundance and diversity of wildlife and on human
recreational use. However, the project had been popular and well-financed and
was too far along to stop. Predictions were realized immediately because of
valuable pre-channelization data on wetland animals (Perrin et al. ), and
actions were set in motion to reverse the damage. Millions of dollars were
spent in the modifications, and millions more will be spent in the restoration
one of the largest ever to be undertaken (Koebel ). Prediction of waterbird
response to the return of seasonal flows and restoration of the diverse wetlands
used a modified guild approach to estimate changes in species composition
based on habitat diversity (Fig. .).
Such approaches have the potential to prevent destruction of natural areas
that provide multiple benefits to society, and recreation and human enjoyment
of natural values carry more weight than they once did.In most cases,the greatest alarm has been registered where endangered or threatened species have been
involved.Depending on the country,such legal protection has been responsible
for saving not only breeding areas for the species but also important habitat for
the entire animal community. However, species-oriented legislation may not
clearly address the community or focus on the essential habitat. In the USA this
resulted in the development of Habitat Conservation Plans to promote
recognition and early resolution of an obvious and rising problem. Such plans
have been developed where one or several species of concern are intertwined
with major commercial developments or resource uses and allow advance
negotiation and sometime mitigation to ensure that all activities can proceed.
Obviously, this is not the same as total preservation of the species or community, and the quality of the plan is dependent upon the scientific underpinning
of the requirements and the effectiveness of the negotiators (Luoma ).
Moreover, because multiple habitats may be involved, some types of natural
area (small wetlands) may be sacrificed to save those of greater importance for
the species of focus. Realistic mitigation and replacement of these may be
impossible, particularly when a bottomland is permanently flooded.
. Population management
Because of the scientific advances in rearing of endangered wildlife and of
releasing them into the wild, and the necessity of ensuring their subsequent
survival, a few comments are required to link these efforts to habitat. Except for
References
Riley, L. and Riley, W. (). Guide to the National Wildlife Refuges. New York:
Macmillan.
Scott, D. A.(ed.) (). Managing wetlands and their birds. Slimbridge, UK:
International Wildfowl Research Bureau.
Scott, J. M., Davis, F., Csuti, B., Noss, R., Butterfield, B., Groves, C., Anderson, H.,
Caicco, S., DErchia, F., Edwards T. C. Jr, Ulliman, J., and Wright, R. G. (). Gap
analysis: a geographic approach to protection of biological diversity. Wildlife
Monographs , .
Shafer, C. L. (). Values and shortcomings of small reserves. BioScience , .
Stanley, T. R. Jr (). Ecosystem, management and the arrogance of humanism.
Conservation Biology , .
van der Valk, A. G. and Davis, C. B. (). The role of seed banks in the vegetation
dynamics of prairie glacial marshes. Ecology , .
Van Rees-Siewert, K. L. and Dinsmore, J. J. (). Influence of wetland age on bird
use of restored wetlands in Iowa. Wetlands , .
Walters, C. J. (). Adaptive management of renewable resources. New York:
McGraw-Hill.
Weller, M. W. (). Management of freshwater marshes for wildlife. In Freshwater
wetlands, ecological processes and management potential, eds. R. E. Good, D. F.
Whigham, and R. L. Simpson, pp. . New York: Academic Press.
Weller, M. W. (). Birds of some Iowa wetlands in relation to concepts of faunal
preservation. Proceedings of the Iowa Academy of Science , .
Weller, M. W. (). Waterfowl management techniques for wetland enhancement,
restoration and creation useful in mitigation procedures. In Wetland creation and
restoration, eds. J. A. Kusler and M. E. Kentula pp. . Washington DC: Island
Press.
Weller, M. W. (). Seasonal dynamics of bird assemblages in a Texas estuarine
wetland. Journal of Field Ornithology , .
Weller, M. W. (). Use of two waterbird guilds as evaluation tools for the
Kissimmee River restoration. Restoration Ecology , .
Weller, M. W. (). Birds of rangeland wetlands. In Rangeland Wildlife, ed.
P. Krausmann, pp. . Denver, CO: Society for Range Management.
References
16
Outlook
Those who have observed wetland birds and noted changes in wetland habitats
over time share a great concern about whether these birds can and will survive.
Obviously, they cannot survive without wetlands. Fortunately, wetlands are
amazingly resilient systems because they are communities of diverse organisms adapted to dynamic water regimes; some one of the many possible species
whether algae, forb, invertebrate or bird seems able to succeed when others
do not, and the system seems to function at some biochemical and trophic
level with these alternatives. Birds show even more flexibility than most
members of the community because of their great mobility, but because they
are dependent on water and food resources, they are no less sensitive to external impacts on the system. While some bird species usually can find resources
for survival, fewer can fulfill the needs for successful reproduction in stressed
wetlands, which results in a community with reduced species richness caused
by reduced habitat diversity, poor conditions for food organisms, and reduced
water quality.
We find amazing examples of wetland birds that live with society: Killdeer
and Black Skimmers nesting on rooftops and feeding elsewhere; Ospreys,
geese, storks, and herons nesting on various artificial structures and flying long
distances for food; Brown Pelicans, Great Egrets, and Great Blue Herons gathering at boat docks to mooch food from incoming fishermen; White Pelicans
feeding at night by pier lights; gulls feeding behind fishing boats; pipits, harriers, egrets, plovers, and gulls and terns feeding behind the plow or the stillburning fire in agricultural fields; and mass migrations of ducks, geese, swans,
cranes, and pelicans along their traditional river routes that now take them
directly over sprawling and spewing cities.
Many people have had the experience of visiting countries with human
populations at levels that have seriously affected wetland habitats and bird
resources. Yet, we are surprised at the numbers if not the variety of waterbirds
seen. A few species use constructed wetlands designed for various alternative
uses: ducks may find submergent vegetation or invertebrates in gravel pits
(Svedarsky and Crawford ); many birds species feed and nest in rice fields
(Fasola and Ruiz ), and similar agricultural fields are used in the nonbreeding season; flamingos use salt-ponds (Espino-Barros and Baldassarre
Outlook
); and ducks and shorebirds find rest and water in livestock ponds (Evans
and Kerbs ), or food, cover, and even nest sites in any unused low-lying
areas or those used for light grazing (e.g., Killdeer).
Wetlands now are built or modified as tertiary sewage treatment systems,
producing enormous quantities of invertebrates favored by swimming waterbirds that strain for their foods (Swanson ). Minnows are an adaptable and
diverse group common to many types of wetland and attract bitterns, egrets,
and herons. I have visited several such treatment units that have become
favored bird observation sites, with an impressive array of birds and with
wonderful opportunities to observe foraging behavior. Northern Shovelers
and Ruddy Ducks find the enriched waters a great place for invertebrate foods;
Tricolored Herons, several egrets, and the seemingly shy Least Bitterns and
Clapper Rails also demonstrate their feeding tactics. Not all come to eat;
Redheads and Northern Pintails come to drink, rest, and avoid the wind on
open bays. At one such unit, Frigatebirds briefly scoop up fresh water and sail
on. American Coots, Moorhens, and Pied-billed Grebes nest with little
obvious concern for those who pass by.
In many cases, use of such areas is temporary and by migrants and therefore, is the product of breeding successes elsewhere, which makes us wonder
whether that source also will fade under future human stresses. But some
populations breed successfully and maintain themselves for many years in
areas that seem to us unsuitable, presumably because the essential habitat
resources are there despite the seeming stresses of society (e.g., Ospreys, Bald
Eagles, skimmers, terns, herons, and kites). Some have maintained free-living
populations within major cities for many years, such as the breeding Tufted
Ducks, Common Pochards, and Mallards of London parks (Gilham ), and
Mallards of the northeastern USA (Figley and van Druff ). Others, like
Mute Swans, have been domesticated and managed as decorative flocks for
centuries (Ticehurst ). In some cases, however, we have succeeded too well
because we, in effect, have created a predator-free habitat, a non-hunting zone,
and excellent food or water supplies that may lead to overpopulations of
attractive but often nuisance species like Canada Geese now widely distributed around the world (Cooper and Keefe , Hanson ).
Addition of water areas for livestock or water supplies and associated agricultural crops often brings new species into an area, such as Maned Ducks in
Australia (Kingsford ) and Black-bellied Whistling Ducks in south Texas
(Bolen and Forsythe , Bolen and Rhylander ). Other examples of the
adaptability of selected species to large and permanent water are species like
cormorants, which seem to have increased in areas where we have created
ponds and lakes, and then we complain because they eat ourfish! In the USA,
the rate of wetland loss should be declining because of a declared policy of nonet loss, but our wetland surveys show an increasing proportion of openwater types and do not detect degradation of wetlands, which tend to have
lesser quality because of extremes in water volume and water quality, and
Outlook
Outlook
cultures and functional values of the biosphere. Where such values are esthetic
or recreational rather than monetary, wetlands have been preserved through
outright purchase and lease, but is this feasible in third-world nations as
human populations continue to rise (Errington )? As Aldo Leopold recognized many years ago (Leopold , ), conservation of remaining wildlife
habitats and restoration of those recently lost probably is feasible only as individuals develop a vested interest and conservation ethic that recognizes
resource values, have some access to the resources, and accept responsibility
for them.
Certainly, pristine wetlands of original size, density, and characteristics
would be an ideal goal and we should establish that goal for major wetland
types in each region of each country or political unit. Such areas have been set
aside to some degree in refuges and preserves, but many of these have traditional grazing and mineral rights and other land-uses. Moreover, many have
modified water regimes outside the preserve boundaries that seriously
influence how the protected system functions. Perhaps the only National
Park in the USA that is totally a wetland system is Everglades National Park in
Florida, but external uses and diversion of water has made that a highly
impacted area made up of patches of modified wetlands that no longer function as a unit (see Douglas ). Recent state, local, and national agreements
to repair the plumbing in this system and to reduce agricultural pollution
hold real promise for this unique wetland. Natural preserves like Big Cypress in
Florida and the Big Thicket of east Texas have major wetland components.
As the public becomes concerned and more knowledgeable about ecological approaches to conservation, there are dangers of focusing the public on the
wrong goals and values and improper temporal and spatial scales to simplify
and to gain support. This is especially true if guidelines are based on a pattern
of terrestrial systems, which operate on a more annual time frame. On a still
broader basis, the concept of biodiversity is very popular, and a variety of other
richness-based methods of delineation are in use (e.g., Williams et al. ),
but we must ensure that the identification of clear goals precedes development
of rigid approaches and strategies. There are some frightening interpretations
of richness data inferring that we can save many of the species of the world by
acquiring only % of the land areas (US News and World Report ); this
includes mostly tropical forests where species richness of breeding residents is
high and tropical islands where endemic species tend to have low populations
and therefore, are threatened or endangered. Such a plan could rob society of
a high percentage of the high-latitude wetlands and wetland birds of the world,
and unique examples of habitat exploitation strategies by a single species that
spans the extremes of both hemispheres! Concurrently, a strategy of this type
would minimize protection of many unique wetland habitats dispersed elsewhere on the planet but which may not have high concentrations of endemic
species and yet are productive and valuable resources with multiple functions
serving humans as well as other diverse and unique animals (Maltby ).
Outlook
References
Outlook
17
Epilogue
Wetlands are exciting places for many reasons, but birds are among the more
prominent attractions. Birds are perhaps the most conspicuous component of
a diverse biotic community with complex physical drivers interwoven in an
ecosystem that is both unique and important. Wetlands have played an important role in supplying the resources needed for many societies and still function in that way for some groups today, but they also have provided an ideal
setting for developing the scientific understanding of how such complex
ecological systems function. But many wetlands are in jeopardy, and birds will
be among the first indicators of dangers ahead for an individual wetland or for
a wetland type or region. Despite the amazing resilience of many wetland types
(which we can and do exploit to our benefit), others are delicate and probably
irreplaceable. Moreover, they are sensitive to seemingly unrelated events both
inside and outside the system that can change so much and so fast, and often to
the detriment of waterbirds. Few habitats demonstrate the importance of one
major driver, water, on the complexity, organization, and dynamics of an
ecosystem. Although all habitats respond to water, wetlands have short-term
and long-term consequences that dictate the biodiversity and trophic structure of the entire community sometimes for hundreds of years through
plant and animal succession. Although temperature can also be very important, it is always secondary to the availability and timing of water.
It should be clear by now that habitat and its diverse resources determine in
many direct and indirect ways the evolution of individual species and the relationships between species of a community. As we have seen, individual and
species adaptations include anatomy, morphology, physiology, and behavior.
Assemblages involve interrelationships such as predation, competition, and
diet specializations. While all birds respond to their habitat in this way, it is
especially dramatic in wetland species because of the richness of the system,
which creates dense populations, complex social behavior, diverse interspecies
interactions, and close ties to the biological and physical influences of the
habitat. Through their structural diversity and water regimes, wetland types
dictate assemblages of birds (obviously influenced by geographic area), and
these function as communities that may be skewed in species composition by
the immediate characteristics of the vegetation and water. The uniqueness and
Epilogue
integrity of such communities are reason for amazement, in part because they
are ever changing and are remarkably successful in view of the dynamics of
their habitat.
It is this uniqueness that makes us anxious to preserve more than relicts of
these diverse wetland habitats. Without doubt, we can have simple wetland
systems under stressed conditions that satisfy the needs of a small number of
tolerant species living together as a community, but we can retain the complex
systems of dynamic species only with the preservation of the environmental
features that induce and maintain their integrity. When most water is intensively managed for alternative uses, when basins are filled, and when streams
can no longer flood and scour, fewer and more simple wetland types will result,
and lower diversity and less adaptive and resilient communities of birds can be
expected. We will all be the poorer for it.
Part of our conservation problem is that we lack a clear-cut goal for wetlands. Moreover, because of the multiplicity of interests represented, agreement on such targets will always be a process of optimization, with the
probability that the majority will be unhappy. However, the alternative of total
elimination is far more serious. Not only must these individuals and peoples
work together toward somewhat evasive goals, they also chase a moving target
that is rarely understood by laypersons and politicians. Therefore, we often
focus on the short term and small scale and rarely can achieve funding to
pursue research or management efforts that are essential.
Few habitats demand more diversity of scientific expertise, or more
integration of system functions and interactions than do wetlands. The wetlands survive only by cooperative efforts with political and socioeconomic
entities, with businesses on the land and in the cities, and with primitive societies eking out a subsistence-level way of life. Wetlands demand the attention
of wildlife biologists, plant ecologists, limnologists, stream ecologists, marine
biologists, physical oceanographers, soil specialists, geologists, anthropologists, urban planners, city- to national-level politicians, and global economists
and financiers.
Ecosystem-level research at this scale is more expensive than can be imagined and much less predictable than building a defense arsenal or funding
research on a new food resource for starving people. It is understandably
difficult for politicians to spend vast sums on the nebulous outcome of such
research, but the ultimate outcome of ecosystem-level research has potential
application far beyond birds or bird habitat.
Variability, instability, and unpredictability are all characteristics of most
productive wetland habitats for wildlife. Nevertheless, it is important that we
understand and learn to live with this variability. It does not make the
conservation challenge any easier because it demands that sophisticated
educational efforts precede group agreement and it makes it dangerous to
make promises or encourage expectations about complex and variable
systems. Although we are doing better at getting people to appreciate wetland
Epilogue
Epilogue
The scientific names of birds or bird groups mentioned in this book are listed
alphabetically by English name. Most nomenclature is derived from Monroe
and Sibley (), American Ornithologists Union Check-list of North
American Birds () and its supplements to , and regional works cited
elsewhere.
Albatross (Diomedeidae)
Albatross, Black-browed Diomedea
melanophris
Albatross, Grey-headed Diomedea
chrysostoma
Albatross, Light-mantled Sooty
Phoebetria palpebrata
Alseonax, Swamp Muscicapa aquatica
(Africa)
Anhinga Anhinga anhinga
(Americas)
Auk, Great Pinguinus impennis
(extinct)
Avocet Recurvirostra spp. (most
continents)
Avocet, American Recurvirostra
americana
Babbler, Chestnut-capped Timalia
pileata
Babbler, Marsh Pellorneum palustre
Bishop, Red Euplectes orix
Bittern, American Botaurus
lentiginosus
Bittern, Least Ixobrychus exilis
Blackbirds, New World (Icteridae)
Blackbird, Brewers Euphagus
cyanocephalus
Appendix 1
Appendix 1
Appendix 1
Appendix 1
The scientific names of animal species or groups other than birds mentioned
in this book are listed alphabetically by English name.
Alligator Alligator mississippiensis
Amphipoda: scuds and sideswimmers
(Crustacea)
Arachnida: spiders and relatives
Barbus (a genus of common African
fish) (Cyprinidae)
Beaver Castor canadensis
Bobcat Lynx rufous
Buffalo, Water Syncerus caffer
Caddisfly (Tricoptera)
Carabidae (a family of common
beetles)
Carp Cyprinus carpio
Cattle, Domestic Bos taurus
Chironomid, Midge (Chironomidae)
Cladocera: water fleas, e.g., Daphnia
Clam, Fingernail Sphaerium spp.
Clarius (African air-breathing catfish)
Clarius gariepinus
Copepod, e.g., Cyclops spp.
(Copepoda)
Coyote Canis latrans
Crab, Horseshoe Limulus polyphemus
Crayfish (Decapoda)
Crocodile, Nile Crocodilus niloticus
Cyclops (Copepoda)
Damselfly (Odonata)
Daphnia or water fleas (e.g., Daphnia
spp. Cladocera)
Appendix 2
The scientific names of plants or plant groups mentioned in this book are
listed alphabetically by English name or genus when used as a common name.
Algae Spirulina spp.
Arrowhead Sagittaria spp.
Azalea, Swamp Rhododendron
viscosum
Baccharis (Groundsel-tree/bush)
Baccharis spp.
Beggarstick Bidens spp.
Bladderwort Utricularia spp.
Blueberry, Highbush Vaccinium
corymbosum
Bog-laurel Kalmia polifolia
Bulrush, Alkali Scirpus paludosus
Bulrush, Hardstem Scirpus acutus
Bulrush, River Scirpus fluviatilis
Bulrush, Softstem Scirpus validus
Burreed Sparganium spp.
Buttonbush Cephalanthis occidentalis
Nemopanthus (Mountain-Holly)
Nemopanthus mucronata
Appendix 3
brant, , , , , , , , , ,
,
blackbird, IVa, , , , , , , , , ,
, , , , , , , ,
, , , ,
Bobolink, ,
Bufflehead, ,
Canary, Papyrus,
Canvasback (duck), , ,, , ,
Caracara, Chimango,
Catbird, Grey,
chickadee,
Cinclodes,
coots, , , , , , , , , , , ,
, , , , , , , , , ,
American , , , , , , , , ,
, , , ,
Common, IIh
Giant,
Horned,
cormorants, , , , , , , , , , ,
, , , , , , , , ,
, , , , , , ,
Double-crested, Vd
Neotropic, IIi
Coucal, Pheasant, ,
Crab-plover, ,
cranes, , , , , , , , , , , ,
, , , ,
Sandhill, , , , ,
Whooping, ,
Crossbill, White-winged,
Crow, Fish,
Cuckoo
Mangrove,
Swamp
Yellow-billed,
curlews, , , , , ,
darters,
dipper, , , , , , , , , ,
divers or loons, , , , , , , , , , ,
, , ,
doves,
dowitcher,
Duck
African Black, ,
American Black, ,
Black-bellied Whistling,
Black-headed, , , ,
Blue, , , ,
Brown, Ie
Fulvous Whistling, ,
Grey,
Harlequin, , ,
Hawaiian, Id
Labrador,
Laysan, IIc,
Long-tailed or Oldsquaw, , ,
Mandarin,
Maned (or Maned Goose),
Masked,
Mottled, , ,,
Ring-necked, , , ,
Ruddy, , , , , , , ,
Spectacled,
Steamer-, , ,
Stifftail, , ,
Duck (cont.)
Torrent, , ,
Tufted, VIc, ,
White-backed, ,
White-headed,
Wood, Ic, , , , , , , , ,
ducks, bay or pochard, , , , , ,
ducks, dabbling, , , , , , , , ,
, , , , , ,
ducks, sea, , , , , , , , ,
Dunlin,
eagles, , , , ,
Bald, , , , ,
Fishing,
Stellars Sea-,
eiders, , , , , , , , , , , ,
, , , ,
egrets, , , , , , , , , , , ,
, , , , , , , , , ,
, , , , , , , , ,
Little,
Reddish, ,
Snowy, IIg, IVj, Vd, , , ,
Falcon, Peregrine, ,
finches, , ,
flamingos, Ik, , , , , , , , , ,
,
flycatchers, , , , ,
Acadian,
Alder,
Great-crested,
Olive-sided,
Swamp,
Willow, ,
Yellow-bellied,
frigatebirds, , , ,
Gadwall, , , , ,
gallinule, ,
grebes, , , , , , , , , , , , ,
, , , , , , , , ,
, , , , , ,
Great-crested
Eared,
Horned,
Pied-billed, IVe, , , ,
Western,
godwit, ,
goldeneye, , , , , , ,
Goose
African Pygmy or Nettapus,
Andean,
Barnacle,
Canada, IVc, , , , , , , , ,
Egyptian,
Greylag,
Kelp (or Kelp Sheldgoose), IIe, , ,
Magpie, , ,
Orinoco,
Snow, , , , ,
Spur-winged, ,
Upland (or Upland Sheldgoose), IVg,
White-fronted, ,
grackle, , ,
Grouse
Ruffed,
Spruce,
gulls, , , , , , , , , , ,
, , , , , , , , ,
Franklins,
Grey-headed,
Herring,
Laughing, IIj, Vd,
Harrier, , ,
hawks, , ,
Black-collared,
Common Black-,
Hamerkop, ,
herons, , , , , , , , , , , ,
, , , , , , , , ,
, , , , , , , , , ,
, , , , , , , ,
Black, ,
Black-crowned Night-, Ig, ,
Great Blue, IVj, , , , , ,
Great White,
Green, , , ,
Grey, ,
Little-blue, , ,
Tricolored, , , , , ,
Yellow-crowned Night-, ,
Hoatzin, , ,
hummingbirds,
ibis,
Glossy,
Puna, IIIb
White, If, IVj, , , , , , , ,
jays,
jacanas, ,
jaeger,
Killdeer, , , , , , , , ,
Kingbird, Eastern, , ,
kingfishers, , , , , ,
Kite
Hook-billed,
Snail, , , , , ,
Knot, Red, IIj, , ,
Kookaburra, Laughing,
Lapwing, ,
Limpkin, , , , , ,
loons or divers, , , , , , , , , , ,
, , ,
Mallard, Ig, VIc, , , , , , , , ,
, , , , ,
Murrelet, Marbled, ,
Marshbird, Brown and Yellow,
Meadowlark, , ,
mergansers, , , , , , , , ,
Moorhen, Common, , , , , , ,
Notornis or Takahe,
oriole,
Osprey, , , , , , , ,
Ovenbird,
Owl
Barred, ,
Fishing,
Grass, ,
Great-horned,
Marsh, ,
Short-eared , ,
Oxpecker, Red-billed,
Oystercatcher, , , , , ,
Painted-snipe,
Parakeet, Monk,
Passseriformes, , ,
Pewee, Wood,
pelicans, , , , , , , , , , , ,
, , , , , , , , ,
Brown, , , , , , ,
African White,
American White, IVh, , , ,
Penguin, Gentoo,
Petrels, ,
phalarope, IVk, ,
Pheasant, Ring-necked,
Pintail
Northern, , ,
South Georgia, Ij,
pipits, IIm, , ,
plovers, ,
Black-bellied or Grey, Vd, ,
Crab-, ,
Egyptian, ,
Golden, ,
Piping, , ,
Semipalmated,
Upland (or Upland Sandpiper),
Pochard or bay duck, , , , ,
African or Southern,
Common, VIc,
pratincole, ,
Prion, Antarctic,
quail, ,
rails, , , , , , , , , , , ,
, , , , , , , ,
,
Clapper, , , ,
King, , ,
Virginia, ,
Redhead (duck), , , , , , , , , ,
, , , ,
Redstart, American,
Ruff,
Rushbird, Wren-like,
Rush-Tyrant, Many-colored,
Sanderling, ,
sandpipers, , , , , , , , , , ,
,
Bairds,
Pectoral,
Purple,
Semipalmated,
Solitary,
Spotted,
Upland (or Upland Plover),
Western, ,
White-rumped,
Sapsucker, Yellowed-bellied,
scaup, , , , ,
scoters, , , , ,
Screamer, IId, IVb, , , , , , ,
Shag,
Shelduck, Common, ,
Sheldgoose
Ashy-headed, Ih
Egyptian, Vc
Kelp, IIe, , ,
Upland, IVg,
Shoveler, VId, , , , , , , ,
Skimmer, Black, IVd, ,
Skua, ,
Snail Kite, ,
Snipe, Wilsons, ,
songbirds, , , , , , , , ,
Sora, ,
sparrows, , , , , , , , , , , ,
, , , , , , ,
Bairds,
Clay-colored,
Grasshopper,
Henslows,
House
LeContes,
Lincolns, ,
Nelsons Sharp-tailed,
Savannah, ,
Seaside,
Song,
Swamp, , ,
spoonbills, , , ,
Starling, Slender-billed, ,
stiff-tail (ducks), , ,
stilts, IIl, , , , , , , , ,
Stone-Curlew or Thick-knee, Beach,
storks, If, Vd, , , , ,
Sunbittern, ,
Sungrebe,
Surfbird,
swallows, , , , ,
swans, , , , , , , , , , ,
, , ,
Black,
Black-necked
Coscoroba,
Mute,
Trumpeter, ,
Tundra,
Whooper, IIb
Takahe or Notornis,
Tchagra, Marsh,
Teal
Auckland Island Flightless, IIa, IIIc, Va, ,
Blue-winged, , , , , , , , ,
,
Brown (or Brown Duck), Ie,
Common,
Green-winged, , , , ,
Speckled, ,
terns, , , , , , , , , , , , ,
, , , , , , , , ,
, , , ,
Antarctic,
Arctic, ,
Black, ,
Caspian, Vb
Common,
Forsters, Vb,
Least, ,
Royal, Vb
Sandwich, Vb
Whiskered,
Thick-knee or Stone-Curlew, Beach,
Thrush, Swainsons,
Tit, Marsh,
Turnstone, Ruddy, Iij,
Veery,
Vireo
Red-eyed, ,
White-eyed, Ib,
wading birds, , ,
Warbler
Bachmans
Black-throated Green,
Fairy,
Little Rush,
Mangrove,
Palm,
Prothonotary, Ib, IVi, , , ,
Swainsons,
Yellow, , ,
Water-Tyrant,
Whalehead,
Whimbrel,
Widowbird
Fan-tailed,
Marsh,
wigeon, Ih, , , , ,
Willet, , ,
Woodpecker
Black-bellied,
Downy,
Red-headed,
Ivory-billed, ,
Woodcock, American, ,
Wren
Marsh, , , , , ,
Sedge, , , , ,
Yellowlegs,
Yellowthroat, Common, , , ,
Subject index
Subject index
Central America, , ,
channelization,
chemicals, ,
Chesapeake Bay, ,
Chironomidae, , ,
chronology, , , , ,
classification, xiv, , , , , ,
Clementsian succession,
climate, , , , , , , , , ,
cluster,
coastal, xii, , , , , , , , , ,
, , , , , , , , , , , ,
, , , , , , , , , ,
, , ,
colonial, xii, , , , , , , , , ,
, , ,,
Columbia River,
community structure, , , , ,
compensatory,
competition, , , , , , , , , , ,
, , , , ,
configuration, , , , , , , ,
conservation, xi, xii, xiv, , , , , , ,
, , , , , , , , ,
, , , , , ,
cottonwood,
counts, ,
cover, nesting,
cover-water, , , , ,
Cowardin, , , ,
coyote,
crab, , , ,
crayfish, , , , , , , ,
crocodile, ,
Cyclops,
Daphnia,
decomposition, , ,
density, , , , , , , , , , , ,
, , , , , , , , ,
, ,
descriptors, xi, , ,
desert, , ,
detritivores, , ,
diet, ,
die-off, ,
dispersion, xiv, , , , ,
disturbance, , , , , , , , , , ,
diving, , , , , , , , , , , ,
, , , , , , , , , , ,
, , , , , , ,
dragonflies, , , ,
drainage, , , , , , , , ,
drinking, , , , ,
driver, , , , , ,
drought, , , , , , , , , , ,
, , , , , , ,
duckweed, ,
dynamic, xii, , , , , , , , , , ,
, , , , , , , , , ,
, ,
East Coast,
ecological integrity,
ecosystem, xiv, , , , , , , ,
, ,
ecosystem management,
eelgrass, , , , ,
elephant, IIn
egg laying, , , ,
eggs, , , , , , , , , , , , ,
, , , , , , , , ,
embryo,
energy, , , , , , , , , , ,
, , , , , , , , , ,
Subject index
Florida, , , , , , , , , , , ,
, , , , ,
fluctuations, , , , , , , ,
food
availability, , , , , , , , ,
, , ,
density, , ,
distribution, ,
use, , , , , , , , , ,
foot-stirring,
foraging
behavior, Ian, , , ,
strategies, Ian, , , ,
tactics, Ian, , , , , ,
theory,
forb, ,
forested wetlands, Ib, , , , , ,
fox,
future, xi, xiv, ,
gap analysis,
geographic information system, ,
geographic range, , ,
geomorphology, xiv, ,
Gleasonian succession,
graminoid, ,
gravel, Ih, , , , , , , , ,
grazing, , , , , , ,
Great Plains, ,
global positioning system,
growth
animal, , , , , , , , , ,
plant, , ,, , , , , , ,
, ,
bird population, , ,
guild, , , , , , , , ,
guild analysis, ,
habitat
diversity, , , , , , , , ,
, , , , ,
dynamics, ,
evaluation procedure,
management, , , ,
pattern, , , , , ,
predictability, , , , , , , , ,
preference, ,
quality, xiv, , , , , ,
requirements, , , ,
segregation, IIg, , ,
selection, , , , , , , , ,
shift, ; see also mobility
structure, , , , , ,
suitability index,
unit, ,
Subject index
use, xi, , , , , , , , , ,
, , ,
variability, , ,
hardshores,
Hawaii, ,
herbaceous plants, , , , , , , , ,
, , , , , ,
herbivores, , , , , , , , , , ,
, , , , , , , ,
hippopotamus, Vc,
home-range, , , , , , ,
horizontal patterns,
Horseshoe Crab, IIj,
human dimensions, Vb, VIae, , , , ,
, , , ,
human impacts, , , ,
hunting, , , , , , , , ,
,
Hydrilla, ,
hydrogeomorphic, ,
hydrology, ,
hydroperiod, , ,
hydrophyte,
ice, xiv, , , , , , , , ,
impact assessment, ,
impoundment,
India, ,
innate behavior, , , ,
instability, ,
integrity, , , ,
interbreeding,
interspecies, , ,
interspersion,
invertebrate, , , , , , , , , , , ,
, , , , , , , , , ,
, , , ,
succession,
Iowa, , , ,
Iran,
Iraq, ,
irrigation, , , , ,
islands, , , , , , , , , , ,
, , , , , , , , , ,
, , , , , , ,
isolation, , , , , ,
jewelweed,
Kafue, ,
Kissimmee River,
kelp, IIa,e, Va, , , , , ,
lacustrine, , ,
lake, , , , , , , , , , ,
, , , , , , , ,
landscape, , , , , , , , ,
land-use,
latitude, , , , , , , , , , , ,
,
learned behavior, , ,
legal aspects, , , , ,
lek,
life cycle, xii, , , , , , ,
life form, , , , , , ,
life history, ,
local movements,
London,
Lotus, American or Yellow, Ic,
Louisiana, , , , ,
mangrove, If, IIIa, , , , , ,
marine, , , , , , , , , , , ,
, , ,
marsh, , , , , , , , , , , ,
, , , , , , , , , ,
, , , , , , , , , ,
, , , , , , ,
Marsh Deer,
matrix,
mats, , , , , , , , ,
meadow, , , , ,
microhabitat, , , , , , , , ,
migration, , , , , , , , , , ,
, , , , , , ,
, , , , , , , , ,
, , , , , ,
mink, ,
minnows, ,
Mississippi River, , , , ,
Missouri River,
mitigation, , , ,
mixed-species flocks, IIe, i, j, ,
mobility, , , , , , , , , , ,
, , , , , , , , , ,
modeling, ,
molting, , , ,
molt-migration, ,
morphology, , , , ,
mosquito,
mudflat, , , , , , , , , , ,
, , ,
muskrat, IVc, , ,
mussel, ,
Namibia,
nature preserves,
Nearctic,
Neotropical, ,
nest, xii, IVak, , , , , , , , , , ,
, , , , , , , , , ,
, , , , , , , , ,
, , , , , , , , , ,
, , ,
construction, IVb,
desertion,
structures, ,
New World, , , , , ,
New Zealand, , , , ,
niche, , , , , , , , , ,
niche shift,
no-net loss,
nomadism, ,
North America, , , , , , , , , ,
, , , , , , , , , , ,
, , , , , , , , ,
,
North Dakota, ,
northern hemisphere, , , , , , , , ,
, , ,
Nutria, , ,
nutrient cycling,
nutrients, , , , , , , , , ,
obligate, xii, , , , , , , ,
ocean, xii, ,
oceanic, xii, , , , , , , ,
oil,
Okovango,
oligotrophic, ,
omnivore, , ,
open water, , , , , , , , , , , ,
, , , , , , , , ,
openness,
opportunistic, , , , , , ,
optimal foraging theory,
organization, ,
otter,
outlook,
overflow,
oxygen, , , , , , , ,
oyster,
pair-space,
palustrine, Iac, , ,
Pantanal,
parks, , ,
patch, , ,
peat, Ij, , ,
pelagic, xii
perch, , , , , , ,
phenology, , ,
physiology, , , , , , , ,
pig,
planning, , ,
Subject index
plant succession, , ,
plankton, ,
plant life form, , , , , , , ,
platypus,
plumage, IIIac, , , , , , , ,
plunge-diving, , ,
pollution, ,
pond, Ic, j, , , , , , ,
pool, IIg, , , , , , , ,
population, , , , , ,
management, , , ,
prairie, Ia, , , , , , , , , ,
, , , , ,
precipitation, , , , ,
precocial, , , , , , , ,
predation, , , , , , , , , , ,
, , , ,
predictability, , , , , , , ,
preference, , , , , , ,
presence, xii, , , , , , , , , , , ,
, , , , , , , , , ,
, ,
prey, , , , , , , , , , , , , ,
, , , , , , , , , ,
, , ,
production, , , , , , , , , ,
, , , , , ,
productivity, , , , , , , , , , ,
,
protection, xi, xiv, , , , , , , , , ,
, , , , , , , , , ,
, , , , , , ,
ptilopedic,
rainfall, , , , , , , , ,
, , , ,
range
geographic, , ,
home, , , , , , ,
recreation, ,
reservoir, , , ,
resource
quality,
tracking,
use, , , , , ,
rest and resting, Vad, , , , , , , , ,
, , , , , , , , , ,
river, Ii, , , , , , , , , , , ,
Subject index
, , , , , , , ,
, , , , , , , ,
riverine, Igi, , , , , , , , , ,
, , , , ,
rock, , , , , , ,
rooftops, , , ,
roost, xii, , , , , , , , , ,
, ,
route,
runoff, , ,
rush, , , ,
salinity, xii, Ik, , , , , , , , ,
, ,
salt marsh, , , ,
saltwater, , , , , , , ,
sandbar, IVd,
savannah, , ,
seagrass, ,
seal, ,
seasonality, , , , , , , , , ,
,
seasons, , , , , , , , , , ,
,
seawater, ,
sea-lettuce,
sea-level,
sedge, IVk, , , , , , , , , ,
, , ,
seed, , , , , , , , , , ,
, , , , , ,
seed bank, , , , ,
segregation, , , , , , , , , ,
, ,
selection, , , , , , , , , , , ,
, , , , , , , , , ,
shallow water, , , , , , , , , , , ,
, , , , , , , , , ,
sheetwater, , , , , , , ,
shoreline, , , , , , , , , , ,
, , , , , , , , , ,
, ,
shoreward, , , , , ,
shrub wetlands, IVj, ,
siltation, ,
single species, , , , , , , , , ,
, , ,
slope, , , , , , , ,
Snail, Apple, ,
snowfall,
social, xi, xii, , , , , , , , , , , ,
, , , , , , , ,
social nesting,
social-space,
songbird, ,
South Georgia, If, IIm, ,
spatial, xiv, , , , , , , , , , ,
, , , ,
specialist, , ,
species composition, , , , , , ,
, , , , , , , , ,
,
species diversity, xiv, , , , , , ,
,
species richness, , , , , , , , ,
, , , , , , , , ,
, , , , ,
spikerush, ,
Spirulina spp.,
stability, , , , , ,
status, , , , ,
stopover, , , , , ,
strata, , , ,
strategy, xiv, xv, , , , , , , , , ,
, , , , , , , , , ,
,
stream flow, ,
streamside, ,
stressed, , , , , , ,
structural diversity, , ,
submersed,
substrate, , , , , , , , , , , , ,
, , , , , , ,
succession, , , , , , , , ,
, , , , , , , , , ,
surface-skimming,
surveys, , , , , ,
swamp, , , , , , , , , , , ,
,
Sweden, ,
tactic, , , , , ,
taro, Id
tax, ,
taxonomy, ,
technique, , , , ,
temperature, , , , , , , , , , , ,
, , , , , , ,
temporal, xv, , , , , , , , , ,
, , , , , ,
territory, IVa, , , , , , , , , ,
Texas, , , , , , , , , , ,
, , ,
thermoregulation, , ,
tidal, , , , , , , , , , , , ,
, , , , , , , , , ,
, , ,
tide, , , , , , , , ,
tidewater,
TigrisEuphrates,
tree, , , , , , , , , , , , , ,
, , , , , , , , , ,
trophic niches, , , , ,
tropical, , , , , , , , , ,
, , , , , , , ,
trout,
tuber, , ,
tundra, , , , ,
turtle,
Umwelt,
underwater, , , , , , , , , , , ,
, ,
United Kingdom, IIb, IIIb, VIc,
United States, , , , , , , , , , ,
, , , , , , , , , ,
, , , , , , ,
unvegetated, ,
upslope, , , , , ,
urbanization, ,
Utah,
vegetation layers, , , , , , ,
vegetation management,
vulnerability, , , , , ,
water
clarity,
declining levels, , , , , , ,
depth, , , , , , , , , , , ,
, , , , , , ,
influences, , , , , , , ,
level, , , , , , , , ,
regimes, , , , , , , , , , , , ,
, , , , , , , , ,
, , , , , , , , ,
, , , , , , , ,
use,
variability, IIg, , , , , , , , ,
,
Water-hyacinth,
water-lily,
waves, , , , , , ,
wetland
classification, xiv, , , , , ,
complexes, , , , , , , , ,
, , ,
configuration,
diversity, xiv, ,
formation, , ,
isolation,
losses, , ,
Subject index
management, xi, ,
quality, xi, , ,
size, ,
type, , , , , , , , , , ,
, ,
variability, ,
vegetation, , , , , , , , , ,
whirlygig, ,
Wigeongrass, ,
Subject index
willow, , , , ,
wintering, , , , , , , , , , , , ,
, , , , , , , , , , ,
, , , , , , , , , ,
, , ,
wooded wetlands, ,
Zambia,
zonation, , , , , ,
zoogeography,