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Regression models for spatial prediction: Their role for biodiversity and
conservation

Article  in  Biodiversity and Conservation · December 2002

DOI: 10.1023/A:1021354914494

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 Biodiversity and Conservation 11: 2085–2092, 2002.
© 2002 Kluwer Academic Publishers. Printed in the Netherlands.

Regression models for spatial prediction: their role

for biodiversity and conservation

A. LEHMANN1,∗ , J.McC. OVERTON2 , and M.P. AUSTIN3

1 Swiss Centre for Faunal Cartography, Terreaux 14, CH-2000 Neuchâtel, Switzerland; 2 Landcare
Research, Private Bag 3127, Hamilton, New Zealand; 3 CSIRO Sustainable Ecosystems, GPO Box 284,
Canberra City ACT 2601, Australia; *Author for correspondence
(e-mail: [email protected]; fax: +41-32-717-7969)

Received 14 June 2002; accepted in revised form 9 July 2002

Abstract. This paper is an introduction to a Special Issue on ‘regression models for spatial predictions’
published in Biodiversity and Conservation following an international workshop held in Switzerland in
2001 (http://leba.unige.ch/workshop). This introduction describes how the exponential growth in comput-
ing power has improved our ability to reach spatially explicit assessment of biodiversity and to develop
cost-effective conservation management. New questions arising from these modern approaches are listed,
while papers presenting examples of applications are briefly introduced.

Key words: Community composition, Conservation, Generalized additive models, Geographic informa-
tion systems, Generalized linear models, Hotspots, Reserve network, Spatial predictions, Species distribu-
tion, Species richness.

Abbreviations: GAM – generalized additive models, GLM – generalized linear models, GIS – geographic
information systems.

Introduction

Ecologists are increasingly being asked to produce detailed spatial estimates of natu-
ral resources, particularly biodiversity attributes (e.g. Prendergast et al. 1993; Dobson
et al. 1997; Scott and Jennings 1997; Reid 1998; van Jaarsveld et al. 1998; Austin
1999; Margules and Pressey 2000; Ferrier 2002). The exponential growth in com-
puting power has allowed both conceptual and operative advances in biodiversity
assessment and the possibility of cost-effective conservation management. The ad-
vent of suitable software for geographic information systems (GIS), remote sensing,
statistical modelling and database management has changed the way ecologists think
about data on species distribution (e.g. Franklin 1995; Bio et al. 1998; Leathwick
1998; Guisan and Zimmermann 2000; Overton et al. 2000). Ecologists are redefining
the questions they ask using their new analytical tools. These questions are becom-
ing much more directed towards providing answers to problems of conservation and
ecosystem sustainability.
2086

Examples of recent questions asked in the literature are:

• What is the potential distribution of species? (e.g. Augustin et al. 1996; Lehmann
1998; Heegard and Hangelbroek 1999; Elith 2000; Frescino et al. 2000; Osborne
et al. 2001; Texeira et al. 2001)
• How high is the biological diversity of this pond, of ponds in this region, of
ponds in this country? (Oertli et al. 2002)
• Can we use biodiversity surrogates in regional conservation planning? (Ferrier
and Watson 1997)
• How can we import expert opinion into the modelling of faunal distribution?
(Pearce et al. 2001)
• Is the observed species diversity of this region higher or lower than its potential?
(e.g. Wolgemuth 1998)
• How can we improve forest inventory? (Moisen and Edwards 1999)
• What is the ecological condition of this ecosystem and is it sustainable?
• What was the composition of the primary forest that previously grew in this
pasture? (e.g. Leathwick 2001)
• What will be the impact of climate change on vegetation patterns? (e.g. Guisan
and Theurillat 2000)
• How damaging are these weeds and pests for agricultural production in our
country?
• What is the most suitable area for a new nature reserve? (e.g. Margules and
Pressey 2000)
• Are these remnant patches of forest sufficiently connected to ensure the survival
of endangered species?
Overton et al. (this volume) examine how best to provide answers to questions
such as these that span a wide range of detail and spatial extent. Policy-makers need
answers to these questions and estimates of other biodiversity attributes which are
transparent, reliable, spatially explicit and inexpensive. Numerous methods of eco-
logical modelling are being used to address these requirements at the present time.
The proceedings of a recent conference ‘Predicting Species Occurrences. Issues of
accuracy and scale’ provide an extensive overview of modelling methods and issues
(Scott et al. 2002).
The combination of regression modelling and GIS is a powerful tool for biodi-
versity and conservation studies (e.g. Lehmann et al. 2002), and is seen in all of the
papers of this special issue. A model used for biodiversity assessment should have a
number of characteristics:
(1) The model should not only be precise, but should also be ecologically sensible,
meaningful and interpretable. In order to achieve this, it is important to develop
modelling procedures in agreement with current theory in ecology (Austin and
Gaywood 1994). The nature of environmental predictors (direct, resource or in-
direct gradients), the shape of response curves, the effect of species interactions,
are all important considerations in developing environmental models.
 2087

(2) The model should also be general, which means applicable in other regions or
different times (e.g. Leathwick et al. 1996). This is a crucial issue with regression
models because their results can only be generalized to predict within the envi-
ronmental conditions found in the data used to calibrate a model. In other words,
it is relatively safe to interpolate within the environmental space considered, but
generally dangerous to extrapolate outside of known boundaries.
(3) The model should be fully data-defined, so that all predictions are fully specified
from empirical data, with no parameters or modules that have been guessed or
defined as having convenient values.
(4) Finally, a model used for biodiversity and conservation should be expressed in a
spatial framework (e.g. Austin and Meyers 1996). In order to produce spatially
explicit outputs from point observations using regression models, the environ-
mental predictors selected in a model must correspond to GIS maps, defined at a
suitable grid resolution. By replacing predictors in a model by their correspond-
ing maps, we can, not only derive spatial predictions of the response we are in-
terested in, but also predictions of the errors we make.
Two particular methods of regression analysis are subject to intense use and evalu-
ation. Generalized linear regression (GLM; McCullagh and Nelder 1997) is an exten-
sion of classical regressions that allows modelling of responses (dependent variables)
such as species richness, abundance classes and presence–absence data. These types
of data are usually not normally distributed like measures of biomass or body size, and
should not be modelled in the classical Gaussian framework. With GLM, predictors
(independent variables) are introduced in a model essentially in a linear or curvilinear
form as in classical regressions. Generalized additive models (GAMs; Hastie and
Tibshirani 1990; Yee and Mitchell 1991) are a non-parametric extension of GLM that
allow introduction of non-linear responses to predictors. The shape of response form
is driven by the data and is not predefined, allowing the study of response shape and
potentially improving predictions by modelling closer to the actual data.
Regression analysis and the resulting spatial prediction is intrinsically objective
because it is explicit, consistent and repeatable, and creates models that are fully
data-defined. However, numerous subjective decisions are necessary when building
a model, from the choice of a sampling strategy, the nature of what is measured
in the field, through to the method used to build a model. For these reasons, it is
important to determine the effects that these modelling decisions make on the models
and predictions, and to define a set of best practices. Continuous improvement in our
current best practice in using these methods is necessary if ecologists are to contribute
to management of our natural resources.

Context

The edition of this special issue (Lehmann, Austin and Overton, eds.), together with a
parallel issue for Ecological Modelling (Guisan, Hastie and Edwards, eds.) was initi-
2088

ated by the organizers of a workshop held in August 2001 at Riederalp in Switzerland

(http://leba.unige.ch/workshop). The workshop and the two special issues share the
same aims and philosophy: bringing together ecologists, statisticians and GIS spe-
cialists in order to discuss and improve the state of the art of generalized regression
modelling for spatial predictions in ecology. A major concern was to improve current
practice by achieving a synthesis of ecological and statistical approaches to statistical
modelling as applied to biodiversity.
The workshop was divided in six themes corresponding to logical steps in the
process of building spatial predictions and used as introductory talks available on
http://leba.unige.ch/workshop/video.htm:
• links with theory in ecology (by M. Austin),
• data sampling (by T. Edwards),
• model selection (by T. Hastie),
• model evaluation (by J. Elith),
• spatial predictions (by R. Aspinall),
• from spatial predictions to environmental management (by J. Overton).
The focus of the workshop was oriented towards methodological aspects except
for the last theme. It was decided therefore that a special issue for ecological mod-
elling would concentrate on the first five themes, whereas the present special issue
would concentrate on applications for biodiversity and conservation, though these
methods and examples have much wider application in natural resource management.
Our aim with this special issue is to provide a vision and examples for the use
of statistical modelling for conservation management. We hope that these papers will
stimulate and convince potential users that these methods have now reached a level
of quality where they should be considered as valuable tools for conservation plan-
ning, especially when spatially explicit information is needed. This field of ecology
provides many opportunities and challenges along with much promise to deliver a
quantitative basis for biodiversity management and conservation policy.

Special issue content

The first paper presents a vision for the way in which information is integrated and
generalized into forms that can be used to support conservation decisions. Jake Over-
ton and co-authors use information pyramids as a metaphor for a general paradigm for
informed ecosystem management. The pyramidal metaphor is, however, based upon
a rigorous foundation of quantitative and explicit methods of data generalization.
Regression analyses and their use in spatial predictions provide an important method
for integrating and generalizing information. The paper uses examples of efforts in
New Zealand as a case study to demonstrate the concepts. Information pyramids, and
their characteristics, provide a powerful perspective for viewing the rest of the papers
in this special issue.
 2089

The remainder of the papers are organized roughly in increasing order of number
of species and complexity of modelling. We begin with papers that investigate single
species, then work up to multiple species, species interactions, emergent properties,
communities, and biodiversity hotspots. Each of these steps includes a number of
conceptual and methodological problems and solutions.
Ramona Maggini and co-authors use GLM to predict the distribution of a threat-
ened ant species in the Swiss National Park and throughout Switzerland. This single
species approach leads to numerous potential applications for the conservation of the
species and to a better understanding of its ecology. Population management within
the Park, reasons for the decline in Switzerland, a sampling design for new field
surveys and the study on the effect of habitat fragmentation are among the possible
outcomes discussed by the authors.
Habitat fragmentation is the central topic of the paper presented by Nicolas Ray
and co-authors. Its aim is to model the spatial distribution of amphibian populations
based on the connectivity between breeding sites. They develop new methods of as-
sessing connectivity using detailed land use maps that generally improve the ability to
predict amphibian occurrence. Methods such as these show promise for understand-
ing and managing amphibian populations in areas with intensive human use.
José Teixeira and J. Arntzen predict the potential impact of climate warming on
the distribution of a vulnerable species of salamander in Portugal and Spain. This pa-
per examines methods for assessing changes in habitat suitability and degree of frag-
mentation under climate change scenarios. The approach should help in conservation
planning by identifying stronghold areas and potential habitat corridors to maintain
this species in the future.
John Leathwick addresses more directly the complex species interactions that are
most often ignored in species distribution modelling. New Zealand forest inventories
provide here a perfect dataset to test our understanding of the niche concept in differ-
ent competitive contexts. Results indicate strong competitive interactions that under-
line the difficulties of using current species distribution models to predict distributions
under future climatic conditions.
Ana Bio and co-authors provide an example where an entire community of river
valley plant species in Belgium was studied and modelled. Their paper discusses lim-
itations in data quantity and quality that can affect the applicability of the ecological
modelling procedures used. However, they conclude that despite a restricted number
of observations, non-randomly collected and spatially correlated data, ecologically
sound models were obtained that predicted most species well.
Anthony Lehmann and co-authors present a species and community level ap-
proach for assessing fern biodiversity in New Zealand. The aim of this paper is to in-
vestigate broad scale distributions of species in relationship to climatic and landform
variables and to identify hotspots of fern diversity. This approach also provides the
basis for setting targets for a biodiversity assessment and restoration program. This is
the first of several papers that models many species, and has developed streamlined
2090

procedures for analyzing and predicting each species, as well as methods for organiz-
ing and integrating the individual species predictions into higher level assessments of
biodiversity.
Margaret Cawsey and co-authors present a concrete example from Australia of
how species and community level modelling can provide a basis for the assessment
of conservation priority to be attached to any particular vegetation remnant using the
predicted original distribution of that vegetation type and the percentage remaining.
The approach can also provide a basis for the setting of priorities for revegetation
to connect and expand existing forest remnants in an area that has been extensively
cleared since European settlement. This paper presents also an excellent example
of practical difficulties that must be expected in applying modern approaches based
on species and community modelling to underpin conservation planning in heavily
human-influenced ecosystems.
We end with two papers by Ferrier et al. that describe what we believe is the
most extensive and longest running application of species and community modelling
to regional conservation planning. The first paper by Ferrier et al. describes the role
of statistical modelling and spatial prediction of species distribution in regional con-
servation planning. The state of New South Wales in Australia has benefited from
this work in a series of major land-use planning decisions allocating public land to
forestry and conservation uses, and in the design of a network of nature reserves.
Streamlined species modelling, dealing with spatial autocorrelation, and evaluation
of model results are among the new approaches detailed in this paper.
The second paper by Simon Ferrier and co-authors follows on from the first paper
to discuss community level modelling approaches. Current approaches to community
level modelling are discussed, and new approaches such as canonical classification
and the modelling of compositional dissimilarity are outlined.
In collection, these papers demonstrate a range of methods and solutions for the
application of regression modelling to conservation management. We hope that you
will find the approaches and results as compelling as we do.

Acknowledgements

As guest editors we gratefully acknowledge the enthusiasm and help received by Alan
Bull from the early stages of the preparation of this special issue. The contribution of
Kluwer Academic Publishers to the cost of colour figures was also greatly appreciated
by the authors. We also acknowledge the collaborative efforts made with Antoine
Guisan, Trevor Hastie and Thomas Edwards, guest editors of the related special is-
sue for ecological modelling. As organizers of the GLM/GAM modelling workshop,
we thank the staff at Villa Cassel in Riederalp for their wonderful hospitality and
efficiency that greatly contributed to the success of the workshop. The workshop
 2091

was supported by the Swiss Academy of Science and the Swiss National Science
Foundation, as well as private sponsors ESRI and Insightful.

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