WI LDLI FE HAB I TAT C ONS E RVATI ON

Wildlife Management and Conservation

Paul R. Krausman, Series Editor

WI LDLIFE
Habitat Conservation
Concepts, Challenges, and Solutions

EDITED BY

MICHAEL L. MORRISON
& HEATHER A. MATHEWSON

Published in Association with THE WILDLIFE SOCIETY

j ohns hopkins u niv ers it y p ress | b a lt i m o r e

© 2015 Johns Hopkins University Press
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Library of Congress Cataloging-in-Publication Data

Wildlife habitat conservation : concepts, challenges, and

solutions / edited by Michael L. Morrison and Heather A.
Mathewson.
pages cm. — (Wildlife management and conservation)
Includes bibliographical references and index.
ISBN 978-1-4214-1610-6 (hardcover : alk. paper) — ISBN
978-1-4214-1611-3 (electronic) — ISBN 1-4214-1610-7
(hardcover : alk. paper) — ISBN 1-4214-1611-5 (electronic)
1. Habitat conservation. I. Morrison, Michael L.
II. Mathewson, Heather A. (Heather Alexis), 1974–
QH75.W529 2015
333.95'4—dc23 2014020806

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 Contents

List of Contributors vii 8 The Impact of Invasive Species on Wildlife

Preface ix Habitat 102
Acknowledgments xi Julie L. Lockwood and J. Curtis Burkhalter

Part I • Foundation Part III • Research and Conservation

1 The Misunderstanding of Habitat 3 9 Thoughts on Models and Prediction 117
Heather A. Mathewson and Michael L. Bret A. Collier and Douglas H. Johnson
Morrison
10 Manage Habitat, Monitor Species 128
2 Exploration and Critique of Habitat and Habitat Michael K. Schwartz, Jamie S. Sanderlin, and
William M. Block
Quality 9
Fred S. Guthery and Bronson K. Strickland
11 The Effects of Disturbance and Succession on
3 Demographic Consequences of Habitat 19 Wildlife Habitat and Animal Communities 143
Amanda D. Rodewald Kevin S. McKelvey

4 Managing Habitats in a Changing World 34 12 Wildlife Habitat Restoration 157

Beatrice Van Horn e and John A. Wiens Kathi L. Borgmann and Courtney J. Conway

Conclusion: Synthesis for Advancing Useful

Part II • Habitats in Peril
Knowledge of Habitat: Unifying Themes or Many
5 Habitat Loss and Degradation: Understanding Directions? 169
Anthropogenic Stressors and Their Impacts on Michael L. Morrison and Heather A.
Individuals, Populations, and Communities 47 Mathewson
Cl inton D. Francis

Index 175
6 Population Genetics and Wildlife Habitat 63
Lisette P. Waits and Clinton W. Epps

7 Habitat Fragmentation and Corridors 84

K. Shawn Smallwo od
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 Contributors

William M. Block Douglas H. Johnson

U.S. Department of Agriculture Forest Service U.S. Geological Survey
Rocky Mountain Research Station Northern Prairie Wildlife Research Center
Kathi L. Borgmann Julie L. Lockwood
Arizona Cooperative Fish and Wildlife Research Unit Ecology, Evolution, and Natural Resources
School of Natural Resources and the Environment Rutgers University
University of Arizona
Heather A. Mathewson
J. Curtis Burkhalter Department of Wildlife, Sustainability, and Ecosystem
Ecology, Evolution, and Natural Resources Sciences
Rutgers University Tarleton State University
Bret A. Collier Kevin S. McKelvey
Texas A&M Institute of Renewable Natural Resources U.S. Department of Agriculture Forest Service
Texas A&M University Rocky Mountain Research Station
Courtney J. Conway Michael L. Morrison
U.S. Geological Survey Department of Wildlife and Fisheries Sciences
Idaho Cooperative Fish and Wildlife Research Unit Texas A&M University
Department of Fish and Wildlife Sciences
Amanda D. Rodewald
University of Idaho
Cornell Lab of Ornithology and Department of
Clinton W. Epps Natural Resources
Department of Fisheries and Wildlife Cornell University
Oregon State University
Jamie S. Sanderlin
Clinton D. Francis U.S. Department of Agriculture Forest Service
Department of Biological Sciences Rocky Mountain Research Station
California Polytechnic State University
Michael K. Schwartz
Fred S. Guthery U.S. Department of Agriculture Forest Service
Department of Natural Resource Ecology and Rocky Mountain Research Station
Management
K. Shawn Smallwood
Oklahoma State University
Davis, California
viii contributors

Bronson K. Strickland John A. Wiens

Department of Wildlife, Fisheries and Aquaculture School of Plant Biology
Mississippi State University University of Western Australia
Crawley, Western Australia, Australia
Beatrice Van Horne
Point Blue Conservation Science
U.S. Department of Agriculture Forest Service
Pacific Northwest Research Station
Lisette P. Waits
Department of Fish and Wildlife Sciences
University of Idaho
 Preface

There are a plethora of books that describe the many tat and ranging to how recent insights in demograph-
impacts that humans have on the environment, and ics, spatial and temporal heterogeneity, and behavioral
there are many books that provide very technical (e.g., processes have increased the complexity of the concept
modeling) ways to analyze complex environmental of habitat. It considers how the goals of resource mana-
impacts. Missing are books that blend an understand- gers oftentimes rely on the simplified, traditional con-
ing of the impacts on wildlife and wildlife habitat with cept of habitat, resulting in tension with the habitat
ways to address and hopefully start to ameliorate nega- concept when applied to complex processes.
tive impacts. We chose the topic for each chapter in this Part II delves into specific factors impacting wild-
volume to address a major issue confronting wildlife life habitat through more than just the removal of
habitat. Hence the book presents in-depth coverage of vegetative cover. The traditional view of habitat as a
individual topics while also presenting a broad cover- vegetation type is complicated by loss or degradation
age of topics overall. via sources other than just changes in land cover. An-
The purpose of Wildlife Habitat Conservation is to thropogenic effects such as lights and sounds can de-
deliver to a broad audience an understanding of the grade the environment without removing vegetation
influences on wildlife and wildlife habitats, including cover per se, and genetic and demographic processes
an evaluation of the state-of-the-art and recommenda- are altered or accelerated with changes to land cover.
tions for a path forward that will advance management Furthermore, the invasion of nonnative plants and
and conservation. Each chapter provides information animals alters habitat structure and processes within
in a form accessible to a broad audience, including but the system. Each chapter discusses how improving our
not limited to advanced undergraduate and graduate understanding of these impacts will contribute to more
students in natural resource management and conser- effectively managing wildlife habitat.
vation (e.g., wildlife, range, conservation biology, rec- Part III emphasizes solutions, including how to
reation and parks); resource managers at local, state, predict future changes, monitoring and planning, and
and federal levels (e.g., state wildlife and parks depart- restoration. These chapters investigate how wildlife
ments, USFWS, BLM, Army Corps); and private land managers and researchers can become more proactive
managers. in how they think about planning, monitoring, manag-
The book is organized into three parts. Part I lays the ing, and restoration in wildlife science. The final chap-
foundation for all that follows, starting with a discus- ter synthesizes the major messages from the book.
sion of the current and traditional use of the term habi-
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 Acknowledgments

We thank Paul Krausman, University of Montana Natural Resources assisted with final manuscript
and Editor of the Wildlife Society book series, for preparation.
inviting us to assemble the contributors for this An initial draft of each chapter was reviewed by
volume. Vincent Burke, Executive Editor, Johns two or more independent referees and then returned
Hopkins University Press, substantially improved the to the chapter lead author for revision. We then
outline and direction of this project. We also thank evaluated the response of the authors to review com-
Catherine Goldstead, Editorial Assistant, JHUP, for ments and asked for additional revision. We list each
helping to guide the project to completion. Ross reviewer here and sincerely thank them for their time
Anderson at the Texas A&M Institute for Renewable and efforts in improving this book.

Chapter Reviewers Dylan Kesler Brett Sandercock

Jonathan Ballou University of Missouri–Columbia Kansas State University
Smithsonian
David King Ted Shear
Mark Boyce University of Massachusetts North Carolina State University
University of Alberta
Ashley Long Nova Silvy
Anna Chalfoun Texas A&M University Texas A&M University
University of Wyoming
R. William Mannan Dan Simberloff
Melanie Colon University of Arizona University of Tennessee
Texas A&M University
Bruce Marcot Katy Smith
Bridgette Hagerty U.S. Forest Service Texas A&M University
York College of Pennsylvania Portland, OR
John Swaddle
Dick Hutto Scott Mills College of William and Mary
University of Montana University of Montana
Peter Weisberg
Bill Jensen Abby Powell University of Nevada–Reno
Emporia State University U.S. Geological Survey
University of Alaska–Fairbanks Patrick Zollner
Matt Johnson Purdue University
Humboldt State University C.C. St. Clair
University of Alberta
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PA R T I • FOUNDATION
This page intentionally left blank
 1
Heather A. Mathewson
The Misunderstanding
of Habitat
and Michael L. Morrison

O ver the past century, humans have integrated habi-

tat into our daily usage, creating a broadly defined
term that refers to the physical location and associated
well as where we think we need to focus if we are to ad-
vance ecological knowledge and the pursuit of species
conservation. Much of what we present here has been
conditions that an animal experiences. Oftentimes presented elsewhere; we feel the need to re-state some
the term is qualified in various ways to express some often rather basic concepts because, unfortunately, the
level of suitability (e.g., good habitat, bad habitat, old message is not getting through to the majority of users.
habitat). It is even used in reference to people, such as And by users we mean not just academic and agency
the respectable Habitat for Humanity effort that builds scientists but also the legion of practicing biologists
homes for people, or to express the state of living con- and ecologists that work diligently on our public and
ditions. Habitat is certainly one of the most frequently private lands.
used terms in ecology. As the field of ecology has ma- At the most fundamental root, habitat is a binary
tured, so too has the application of the term, resulting determination. An area, and its associated conditions,
in an apparent discord within the overall discipline of either is or is not habitat. In this sense, a phrase such as
ecology. As research advances, through improved sta- “unsuitable habitat” has no meaning (Hall et al. 1997;
tistical, technological, and theoretical approaches, we Garshelis 2000). If a location is unsuitable for occu-
have improved our understanding of the processes and pancy by an organism, then it, simply, is not habitat.
mechanisms associated with an organism’s habitat, yet Frequently, habitat is used to describe an area sup-
the unfortunate result is a large collection of literature porting a particular type of vegetation or aquatic type.
full of muddled terminology. This use apparently grew from the term habitat type,
As thoroughly reviewed elsewhere, the term habitat coined by Daubenmire to refer to “land units having ap-
is widely misunderstood and misused throughout eco- proximately the same capacity to produce vegetation”
logical and management applications (see reviews by (Daubenmire 1976, 125). Some researchers promote
Hall et al. 1997; Morrison and Hall 2002). The failure this definition because of the simplicity in delineation
of the scientific community to agree upon a common or measurements (Garshelis 2000). Furthermore, this
nomenclature contributes to misunderstandings and is the definition often adopted for use in legislation or
misapplication of other concepts such as metapopula- policy. This broad definition regarding “habitat” simply
tion theory (Dennis et al. 2003; Baguette and Menne- as space or ecological / vegetation zones leads to over-
chez 2004; Shreeve et al. 2004) or leads to conflicts use of the term in place of more suitable and better
in methodology (Diaz et al. 2004), creating a plethora defined terms such as vegetation association, resource,
of unnecessary, modified habitat-based terms (Roun- and landscape (see Hall et al. 1997, table 2). For ex-
tree and Able 2007). Thus, in this opening chapter, we ample, Rountree and Able (2007) point out that part
thought it prudent to discuss habitat terminology, as of the ambiguity of the term arises because of the per-
4 foundation

spective of the investigator. Habitat often is applied to then, is a concept associated with a particular species,
reference an area with similar ecological conditions, and sometimes even to a particular population, of plant
such as coral reefs or mud flats, thus referencing the or animal. Habitat is thus species specific; this is a key
community of organisms reliant upon the collection of concept that is lost on many, many users. One cannot
resources provided within the area. This is in contrast walk to a vista and proclaim that “this is a good” or “this
to an organism-focused perspective of habitat. Roun- is a bad” habitat without, at a bare minimum, including
tree and Able (2007) provided a list of additional termi- the specific species under consideration.
nology that differentiates between “ecological habitat Given our definition of habitat as species-specific
type” and “organismal habitat.” We argue that instead and encompassing necessary resources and conditions
of tacking on modifiers to the term, one should adhere for occupancy, there are only a handful of habitat-
to a single concept for habitat and then use more ap- related terms that need to be defined. The list of accept-
propriate terms to represent mud flats and tidal pools able terms is short because only a few key terms and
(e.g., biome, ecosystem; Ricklefs and Miller 2000). concepts are actually needed to understand, quantify,
For application to conservation and management and manage a species’ habitat (our terminology follows
of organisms, the simplicity of a location in space, Morrison et al. 2006 and Morrison 2009). Unfortu-
or ecosite or vegetation type definition, ignores the nately, numerous authors have applied modifiers to the
complexity of the biological world. For management basic habitat term, including “high quality,” “marginal,”
of populations, this definition is essentially useless. It “transitional,” “optimal,” “suitable,” and many, many
eliminates consideration of spatial and temporal varia- more (see Hall et al. 1997). Many of these modified
tion of resources and biotic interactions and how they terms arise from a failure to assign habitat to a single
contribute to population limitation and control (Den- organism (Rountree and Able 2007), or they are inap-
nis et al. 2003; Diaz et al. 2004; Mitchell 2005). Man- propriately used when population demographics are
aging vegetation and other environmental features in not considered (Hall et al. 1997). “Suitable” or “unsuit-
this broad manner will likely fall short of producing the able” habitats are most egregious in that the terms blur
desired assemblage of wildlife by failing to adequately the fact that a lack of suitability simply means that it
account for the needs of the individual species (Lin- is not habitat in the first place. Habitat-modified terms
denmayer and Hunter 2010). Different taxa use re- create a jargon that is unnecessary when alternate
sources differently across a single day, season, or life terms are available that are defined in basic ecology
cycle, and these dependencies must be quantified in textbooks (e.g., biome). This explosion of terminology
order to understand management for a single species has led to rampant confusion. Additionally, even casual
(or population). Failing to simultaneously manage for reading of published papers shows a continual switch-
plants and animals is a hit-or-miss strategy for animals; ing of terms within even the same article. Confusion
managing vegetation and other environmental features reigns.
will address habitat for some animal species but will
ignore—all or in part—habitat for other species (some
Terminology
of which might not be desired).
We want a concept that allows us to tackle issues We define habitat use as the way an animal uses (or
such as how organisms respond to alterations in the consumes) physical and biological components (i.e.,
landscape, changes in the climate, or responses to sto- resources) in a habitat. Habitat use focuses on how an
chastic events. Thus, we follow the definition adopted organism uses their habitat, not if they use an area or
by Morrison et al. (2006) in which habitat is an area resource (because then it is not actually habitat). Either
with a combination of resources (e.g., food, cover, “habitat use” is a description of how resources are used
water) and environmental conditions (e.g., tempera- or often it is quantified as the proportion of time that
ture, precipitation, presence or absence of predators an animal spends within various components of the
and competitors) that promotes occupancy by indi- environment (Beyer et al. 2010).
viduals of a given species (or population) and allows Habitat selection is the hierarchical process of in-
those individuals to survive and reproduce. Habitat, nate and learned decisions by an animal about where
 the misunderstanding of habitat 5

it should be across space and time in order to persist based on demographics (reproduction and survival) of
(Johnson 1980; Hutto 1985; Piper 2011). Habitat se- individuals or populations, which fluctuate over space
lection is an evolutionary process based on fitness and time. Thus, habitat per se has no inherent “quality.”
consequences that vary with differential resource use Authors discuss habitat quality in paper after paper,
(Morris 2011). Habitat preference is restricted to the when what they really mean, at best, is the ability or
consequence of the habitat selection process, result- probability of an area to result in quality (i.e., the pro-
ing in the disproportional use of some resources over duction of young or survival of individuals). One might
others. As we define these terms, habitat use and habi- identify a plot of land as high quality in one year, but
tat preference differ from habitat selection in that the because of myriad factors the same plot is identified as
former are based on quantifiable patterns using mea- low quality in the following year (e.g., annual drought
sures of habitat availability. Unfortunately, authors leads to low arthropod production). Furthermore, Van
oftentimes use the term habitat selection to represent Horne (1983) brought to our awareness that animal
the outcome of statistical interpretation of habitat use density can be a misleading indicator of habitat qual-
compared to habitat availability (i.e., habitat selection ity. Although animal abundance can be correlated with
models; Mannan and Steidl 2013, 233). We argue that habitat quality in some cases, the quality itself should
the term habitat selection should be restricted to the be based on demographics of individuals or populations
behavioral process (Hutto 1985; Morris 2003; Beyer (Garshelis 2000; Railsback et al. 2003; Johnson 2007).
et al. 2010). The distribution of animals is intimately tied to the
Quantifying habitat use or preference requires concept of niche. The concept of niche has been de-
evaluation of how much habitat occurs in the environ- fined in multiple ways over time and continues to be
ment. Abundance, when applied to characteristics of the subject of much discussion. As reviewed in Mor-
a habitat or a resource, is restricted to the quantity of rison et al. (2006, chapter 1), Grinnell (1917) intro-
that particular component, irrespective of the number duced the term niche when explaining the distribution
of organisms present in the habitat (Wiens 1984, 402). of a single species of bird. His assessments included
Abundance of a resource is not adequate for quantify- spatial considerations (e.g., reasons for a close associa-
ing the availability of the various components of a spe- tion with a vegetation type), dietary dimensions, and
cies’ habitat. Habitat availability must consider the ac- constraints placed by the need to avoid predators. Thus,
cessibility of physical and biological components of the in this view, the niche included both positional and
environment to an individual. An individual’s physical functional roles in the community. Elton (1927) later
limitations plus multiple interspecific, environmental, described the niche as the status of an animal in the
or anthropogenic factors might limit accessibility, mak- community and focused on trophic position and diet.
ing habitat availability a difficult measure for investi- Other, more complicated views of the niche also arose
gators to quantify (Beyer et al. 2010). It is, of course, following the pioneering work by Grinnell, Elton, and
difficult to assess resource availability from an animal’s others (e.g., Hutchinson 1957).
perspective. For example, we can measure the abun- Although habitat is a core concept for developing
dance of food for a particular species, but we cannot general descriptors of the distribution of animals, it can
know that an animal can obtain all of the food in the only provide limited insight into factors responsible
habitat because there are many factors that restrict its for animal survival and fitness, and thus population re-
availability. A simple example is plant material (leaves, sponses to changing environments. It seems apparent
fruit) that is beyond the reach of an animal; it is thus to us that the proliferation of terms used by authors to
unavailable for it to feed on. modify the basic habitat term, as discussed previously,
Habitat quality is the ability of the environment to is because of this failure to even think about the niche
provide conditions appropriate for individual and popu- concept when studying how and why animals are dis-
lation persistence, based on the provision of resources tributed as they are. This limited insight from the habi-
for survival, reproduction, and population persistence, tat concept is because habitat alone does not usually
respectively (Morrison 2009, 62). Habitat quality is an describe the underlying mechanisms that determine
assessment of a continuous gradient that necessarily is survival and fecundity. Other factors, including some
6 foundation

often related to an animal’s niche, must be studied to critical habitat designation that legislation and policy
more fully understand the mechanisms responsible rely on is an oversimplification that can result in mis-
for animal survival and fitness (Morrison et al. 2006, management of resources and wildlife.
56–57). Research has shown that a number of environ- We emphasize the need for fitness covariates to
mental factors can restrict survival and productivity of support habitat quality assessments because habitat
a species (across its range or a portion thereof), and the selection does not necessarily correlate with habitat
influence of any single factor is not necessarily additive quality. Too often studies used for direct management
to the influence of any other factor. That is, usually only or conservation purposes default to using data (i.e.,
one factor is limiting in any particular location and density of individuals) or statistical models of habitat
time, and it is unlikely that the same factor will always selection to assign value to properties. The assumption
be limiting because natural variation causes shifting of that highly selected locations are the most beneficial
the quality and quantity of resources. locations is, in reality, an iterative evolutionary process.
Thus, habitat and niche are both related and core Evolutionary habitat selection is determined by habitat
concepts in ecology and therefore wildlife and habitat quality (i.e., variation in fitness controlled by variation
management. As developed by Morrison (2009, 64– in the environment), but habitat selected by individu-
65), focusing on habitat alone is problematic because als will vary in how productive or how well it survives
the environmental features we measure can stay the (i.e., habitat quality). The assumption that these pro-
same while use of important resources by an animal cesses are in equilibrium such that a linear correlation
within that habitat can change. For example, changes between habitat selection and habitat quality exists is
in the species or size of prey taken by a bird foraging ridiculous. Garshelis (2000) pointed out fundamental
on shrubs: the shrubs (“habitat”) do not need to change flaws in the reasoning behind assumptions of habitat
in physical dimensions or appearance. If we describe preference modeling, which multiple research studies
habitat only as structural or floristic aspects of vege- have supported and expanded upon (Van Horne 1983;
tation, we will often fail to predict organism health Railsback et al. 2003). The concepts of ecological traps
because we did not recognize constraints on exploita- or source / sink habitats are in direct opposition of habi-
tion of other resources that are critical limiting factors tat selection as a reflection of habitat quality. Simula-
(Dennis et al. 2003). tions of trout populations and habitat preference mod-
els revealed multiple reasons why density was low in
areas with high fitness potential (Railsback et al. 2003).
What We Have Learned: It Is Time to
We must conclude that the field of wildlife-habitat
Move On
studies has progressed little over the past fifty years or
Habitat is not a delineation of a single vegetation type more. The primary advances in habitat studies have
in a single location across time. Although we cannot involved technology and statistics but not anything
disconnect spatial location from conditions, we can fundamental in the way studies are conceptualized,
and need to recognize that this is not fixed. Habitat planned, or conducted. This is due in part to a con-
describes the conditions surrounding the location of an tinued misuse of the habitat concept and, thus, misap-
animal (Morrison 2001). It is ephemeral and it changes plication of associated theories. But we can improve
with variations in the environment. Landscape ecolo- and promote progress in this field of study. We can rec-
gists regard habitat as a continuum across the landscape ognize that biological systems are complex, and that
and emphasize the importance of considering the “ma- it is an oversimplification to assume that relationships
trix” between patches of vegetation or ecosites. Multi- with host plants, vegetation types, or environmental
scale approaches are needed to characterize habitat substrates are sufficient for species persistence (e.g.,
relationships. Unfortunately, the lingering prevalence Dennis et al. 2003; Diaz et al. 2004). We can encourage
of habitat as a single- dimension, spatially delineated identification of quality habitat only when linked with
area hinders our advancement of understanding re- population dynamics that vary across time and space.
quirements for species or populations. For example, the We can address habitat selection theory by including
 the misunderstanding of habitat 7

consideration of behavioral processes, such as site fa- biological population and hence appropriate selection
miliarity or the importance of public information (Bou- of sampling locations (see Morrison 2012 for a thor-
linier and Danchin 1997; Morris 2011; Piper 2011). ough discussion).
Habitat studies are stagnant because of what Mor- Advancing studies of habitat involves far more than
rison (2012) referred to as our current habitat sampling terminology. Yes, we must start with a basic set of terms
and analysis paradigm. The scientific community per- that recognizes the limitations that the basic concept
petuates this paradigm because we have become com- of habitat can provide, but the ongoing proliferation of
placent in our construction and evaluation of habitat terms is an obvious indication of our broad failure to
relationship studies. Reviewers and editors tend to come to grips with the limitations of “habitat” alone.
publish habitat relationship studies with limited scru- Regardless of the terms used, an improperly designed
tiny because studies often follow the generic prescrip- study does not advance understanding in a broad and
tion of how we have always conducted them and the in- scientific manner. As mentioned in this chapter and
herent limitations of such work. The majority of these extensively reviewed elsewhere (Anderson 2001; Mor-
studies are doomed from the start because we compro- rison 2012), much has been written about our failure
mise our data collection by failing to establish a cogent to properly design studies of wildlife. It is our hope
framework for sampling (Morrison 2012). Oftentimes that eventually all of us will carry these messages to
our studies are largely driven by logistical constraints, our students and other colleagues and begin the task of
related funding limitations, and limited focus at local advancing studies of wildlife and their habitats.
management areas (Morrison 2012). Unfortunately,
this typically results in reliance upon “convenience
sampling,” results of which only allow weak conclusive LITERATURE CITED
Anderson, D. R. 2001. “The Need to Get the Basics Right in
statements about the sample itself rather than the de- Wildlife Field Studies.” Wildlife Society Bulletin 29:1294–97.
sired inductive inference concerning the population Baguette, M., and G. Mennechez. 2004. “Resource and Habitat
of interest (Anderson 2001). As noted many years ago Patches, Landscape Ecology and Metapopulation Biology: A
by Romesberg (1981), we often use statistics to try to Consensual Viewpoint.” Oikos 106:399–403.
compensate for a lack of clear scientific thinking when Beyer, H. L., D. T. Haydon, J. M. Morales, J. L. Frair,
M. Hebblewhite, M. Mitchell, and J. Matthiopoulos. 2010.
formulating our studies. Most of our habitat studies “The Interpretation of Habitat Preference Metrics Under
have little relevance to the target species with regard Use—Availability Designs.” Philosophical Transactions of the
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sistence. Thus, attempts to translate our published re- Boulinier, T., and E. Danchin. 1997. “The Use of Conspecific
sults into management guidelines for a species tend to Reproductive Success for Breeding Patch Selection in Ter-
restrial Migratory Species.” Evolutionary Ecology 11:505–17.
produce one-size-fits-all documents that have little to
Daubenmire, R. 1976. “The Use of Vegetation in Assessing the
do with long-term viability and, thus, persistence. Productivity of Forest Lands.” Botanical Review 42:115–43.
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we are faced with a desire to provide general, holistic Butterfly Biology Viewpoint.” Oikos 102:417–26.
management prescriptions and conservation initia- Diaz, R. J., M. Solan, and R. M. Valente. 2004. “A Review of
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tives (Lindenmayer and Hunter 2010). However, this ing Habitat Quality.” Journal of Environmental Management
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8 foundation

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Wildlife Society Bulletin 25:173–82. ———. 2012. “The Habitat Sampling and Analysis Paradigm
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Ecology 61:65–71. Morrison, M. L., B. G. Marcot, and R. W. Mannan. 2006.
Johnson, M. D. 2007. “Measuring Habitat Quality: A Review.” Wildlife-Habitat Relationships: Concepts and Applications. 3rd
Condor 109:489–504. edition. Washington, D.C.: Island Press.
Kearney, M. 2006. “Habitat, Environment and Niche: What Piper, W. 2011. “Making Habitat Selection More ‘Familiar’: A
Are We Modelling?” Oikos 115:186–91. Review.” Behavioral Ecology and Sociobiology 65:1329–51.
Lindenmayer, D. B., and M. Hunter. 2010. “Some Guiding Railsback, S. F., H. B. Stauffer, and B. C. Harvey. 2003. “What
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Journal of Wildlife Management 65:613–23.
 2
Fred S. Guthery and
Exploration and Critique
of Habitat and Habitat
Bronson K. Strickland

Quality

H abitat is considered a fundamental, unifying con-

cept in wildlife biology. Yet the word seems to
have suffered meaning proliferation that has sapped it
Introduction
Habitat is a set of concepts rather than a single, unique
of rhetorical heft. We explore and critique habitat and concept. It is difficult to write about because of its
habitat quality based on theoretical definitions from the many meanings, some of which have become ambig-
literature, operational definitions in the minds of prac- uous. Thus, every mention of the word may, at least
ticing biologists, and implicit definitions deduced from tacitly, entail clarification unless authors have explic-
statements in literature. Classically, habitat is a natural itly defined it and adhered to that definition. We have
home or living area with resources (e.g., food) and con- noticed that authors who define the concept usually
ditions (e.g., temperature range) that permit the exis- abandon the stated definition.
tence of an organism in an area (classical outlook). To experience the polysemous nature of habitat,
Operationally, habitat is an area with demographically consider its use as an adjective. What does habitat
or behaviorally meaningful subcomponents, which are mean in various adjective forms? Is habitat a synonym
also called habitats, and that also may have subcom- for landscape in the phrase habitat fragmentation? What
ponents called habitats (hierarchical outlook). Cover is habitat in habitat matrix? In habitat quality? In habitat
types, plant communities, physiognomic types, vege- deterioration? A number of other words modified by the
tation types, habitat types, microhabitat, and patches adjective habitat include condition, type, preference,
are, among others, operational synonyms for sub- avoidance, variable, elements, units, and components.
subcomponent habitats. These meanings (area, sub- What does habitat mean when modified by adjec-
component, sub-subcomponent) often are used with- tives such as nesting, roosting, resting, foraging, and
out restraint and, accordingly, the word habitat may thermal? What is the habitat in low-condition habitat?
issue forth in technical prose like an involuntary plati- Is it generically the same habitat as in quality habitat or
tude. To improve clarity and consensibility of a mes- thermal habitat?
sage in papers dealing with habitat concepts, authors What is habitat in Van Horne’s (1983) mathematical
can define habitat and related terms early in a paper definition of habitat quality? Is it the same habitat as in
and then adhere to those definitions. We recommend Wiens (2002)? “In one way or another, predicting the
that authors conduct a word search for habitat when occurrence of species in space and time comes down
they complete a paper. Then, where the word occurs, to dealing with habitats. The issues we face in predict-
usage can be checked against stated definitions and an ing occurrences therefore are issues of habitat—how
explicit synonym can be substituted if the definition we define [habitat]; how we measure, map, and model
does not hold. [habitat]; how we analyze [habitat]; what [habitat]
10 foundation

means to organisms; and, ultimately, how we can use articles and surveyed research and management biolo-
knowledge about habitat to manage natural resources gists for the operational definitions they hold. We also
in a sustainable and balanced way” (739; emphasis deduced meanings of habitat from wording and context
added). when the term appeared in the literature. We discuss
Or consider: “One objective of habitat relationship habitat per se relative to the concept of habitat quality.
research is to identify biologically important variables Based on these activities, we formulate recommenda-
that have the ability to predict species occurrence . . . tions on use of the word habitat in the ecological litera-
Habitat quality for many species contains a spatial com- ture. We regard such use as optimal if it parsimoniously
ponent related to the arrangement and amount of habi- preserves habitat as a powerful, traditional concept
tat elements across the landscape. If the spatial scale of (Block and Brennan 1993) and maximizes clarity and
measurement alters the values of these variables, this consensibility of a message in written and verbal ex-
influences our ability for predicting species occur- changes among biologists.
rence and for assessing habitat conditions” (Trani 2002,
151; emphasis added). What does habitat mean in this
Definitions of Habitat
paragraph? Does it mean the same thing each time it
Dictionaries and Articles
is mentioned? Is it the same habitat as in Van Horne
(1983) or Wiens (2002) or both? As a classical concept, habitat generally is defined in
These preliminary comments suggest that habi- reference to “home” or “living area”: “The locality in
tat may have experienced meaning proliferation and which a plant or animal naturally grows or lives” (Ox-
overuse, such that it may be in danger of losing (or ford online dictionary); a place where a plant or animal
has already lost) rhetorical heft. Wildlife biologists say lives (Smith and Smith 2001, 778); the natural abode
habitat the way politicians say the American people. Yet of an organism (Grange 1949).
these problems do not override the value of the habitat In a slightly more complex version of the defini-
concept as a fundament in the ecology and manage- tion (requisites added), habitat is defined as a set of
ment of wildlife (Block and Brennan 1993). resources and conditions necessary for sustained oc-
Before stating objectives, we note that our subject cupancy (time considered) of an area by an organism:
matter is extensive. To constrain this chapter to a rea- “The resources and conditions present in an area that
sonable length, we limited surveys to include only what [permit] occupancy . . . by a given organism” (Hall
we believe is the most relevant information. Further, et al. 1997, 175); “[t]he type of place where an animal
we focus on the concepts of habitat and habitat qual- normally lives or, more specifically, the collection of
ity and give short shrift to related topics. We will not resources and conditions necessary for its occupancy”
address microhabitat concepts such as nesting, roost- (Garshelis 2000, 112); “[t]he physical space within
ing, bedding, foraging sites, and so forth, though these which the animal lives, and the abiotic and biotic enti-
are important topics. Neither will we address regions, ties . . . in that space” (Morrison and Hall 2002, 51).
biomes, or other vast quantities of space. Except for Safriel and Eliahu defined habitat as “the environ-
passing comments, we do not discuss selection and ment of a community confined to a portion of the land-
avoidance of entities called habitats nor do we compare scape. Each habitat is made of three species-specific
and contrast habitat and niche. Rather, we focus on the components: 1. ‘Environmental’ physical and chemical
concept at a mesoscale that is meaningful to wildlife features (e.g., temperature, salinity), 2. Resources (e.g.,
managers and researchers: areas the size of ranches, food, space), [and] 3. Interacting organisms (other than
farms, study areas, watersheds, wildlife refuges, and those functioning as resources, e.g., competitors, pred-
other areas that serve as living spaces for organisms ators, mutualists)” (1991, 349).
seasonally or annually. Organisms that use space at a Garshelis expanded his definition of habitat to in-
continental scale (e.g., neotropical migrants) also use clude a “set of specific environmental features that,
space at mesoscales in daily and seasonal activities. for terrestrial animals, is often equated to a plant com-
We surveyed the literature for definitions of habitat munity, vegetative association, or cover type” (2000,
that arise in or from dictionaries, glossaries, and review 112). In this sense, alfalfa fields, corn fields, timothy
 exploration and critique of habitat and habitat quality 11

(Phleum pratense) meadows, marshes, bluestem (Andro- a softball from one shrub the size of a Volkswagen to
pogon sp.) prairies, and mesquite (Prosopis glandulosa) the next.” The softball metric insures that requisites are
brushland are habitats. readily accessible by bobwhites, given their mobility. If
These examples stray from the classical concept of the distance to a shrub exceeds a softball throw, space
“living space with requisites” but show that habitat has may become unusable, leading to a decline in average
sundry meanings in practical use. We turn to those density on an area.
meanings now, including ones held explicitly and im- “I would define bobwhite habitat as a set of patches
plicitly by biologists. within a landscape that contains the features bob-
whites require with the availability and spacing of the
features set to support bobwhite life throughout the
Practicing Biologists
year” (C. B. Dabbert, Texas Tech University, personal
For this section, we informally contacted practicing re- communication).
searchers and managers and asked them to answer two “Habitat (for anything) is the place where [it] lives
queries: “What is habitat for [your choice of taxon]?” and includes the resources needed for survival and
and “How do you judge the quality of [taxon] habitat?” reproduction” (A. T. Pearse, US Geological Survey,
Habitat quality will be dealt with in the next major Northern Prairie Wildlife Research Center, personal
section. communication). Here is a detailed example using
Regarding “What is habitat?,” we received this com- a passerine bird: Breeding habitat for a cliff swallow
ment: “This question is like ‘Explain World War II— (Petrochelidon pyrrhonota) includes “relatively open
use back of page if necessary.’ You could write books areas of fields, pasture, prairie, or agricultural land
on this . . . and many have” (K. K. Karrow, Marais des in relative proximity to a nesting structure (cliff face,
Cygnes Wildlife Area, personal communication). True. bridge, highway or railroad culvert, or building) that
Nevertheless, we present a sampling of responses. offers a vertical wall and horizontal overhang on which
The northern bobwhite (Colinus virginianus) pro- the mud nests can adhere. Habitat must also include
vides an example of variability in the definition of habi- a permanent or ephemeral water source that creates
tat for biologists dealing with the species. “I view [bob- mud for nest building within [3–5 km] of the nesting
white] habitat as a residence. Just like our homes have structure” (C. R. Brown, University of Tulsa, personal
many components (living room, bedroom, kitchen, communication).
bathroom, garage), bobwhite habitat has many types Consistent with the definitions in the preceding sec-
of cover (nesting, feeding, brood-rearing, loafing, es- tion, time may be involved in operational concepts of
cape). Just like in our homes, each cover type fulfills a habitat. Habitat “is an area or location that is able to
different need. All cover types are in close proximity to provide for the specific life history needs of a given . . .
one another” (S. J. DeMaso, US Fish and Wildlife Ser- species at a given time. I say ‘at a given time’ because
vice, personal communication). what constitutes useful habitat changes through time
“Bobwhite habitat may be viewed as circumstances and in accordance with the life history stage of the
or areas with characteristic microstructure (bare individual” (R. O’Shaughnessy, Southern Illinois Co-
ground, vegetation height, plant species composition operative Wildlife Research Laboratory, personal com-
to meet food and cover requirements, etc.) and mac- munication).
rostructure (distribution of woody and herbaceous “Habitat is an area that meets the needs for an
plants)” (T. V. Dailey, Northern Bobwhite Conserva- animal’s existence. I used the term existence instead
tion Initiative, personal communication). of occurrence to exclude circumstances when an area
D. Rollins (Texas A&M University, personal com- is not habitat but an animal may be found temporar-
munication) defines bobwhite habitat as “a landscape ily in it (e.g., quail on [a] football field). Thus, there
frequented by quail, either past or present; such is is an implied element of time in my use of existence”
characterized by a plant community interspersed with (F. Hernández, Texas A&M University–Kingsville, per-
woody plants (especially shrubs), grasses, and early sonal communication).
successional forbs. Ideally, one should be able to throw “Habitat is an area that supplies all of the necessary
12 foundation

components . . . for an animal to survive. Habitat com- “habitats,” and “managers” are plural in this sentence
prises food, cover, [and] water in a suitable arrange- fragment. It is unclear whether the statement applies
ment with sufficient space for a particular species. Thus, to different animals on different areas or different ani-
habitat is not necessarily a vegetation type, and any ref- mals on the same area with multiple habitats. If the
erence to habitat as vegetation (e.g., ‘habitat type’) is implication is different animals on different areas,
confusing and should be avoided” (C. A. Harper, Uni- then habitat would seem to be a homogeneous area in
versity of Tennessee, personal communication; empha- space that somehow differs, perhaps arbitrarily, from
sis in original). other areas in space. Here we mean “area” in the sense
Here we introduce usable space because this term of study area, ranch, watershed, management area,
could be used as a synonym for habitat, and the con- and so on.
cept has explicit consideration of time. “Any . . . area If the implication is different animals of a single spe-
can be envisioned as a set of points (e.g., Cartesian co- cies on the same area, the typical study circumstance,
ordinates) surrounded by [biotic and abiotic features then habitat is possibly synonymous with subdivisions
with various properties]. To be fully usable, a point of the area (e.g., patch, cover type, or element). How-
must by definition be associated with [biotic and abi- ever, it could in this context be synonymous with struc-
otic features] compatible with the physical, behavioral, tural properties of vegetation (small scale) that were
and physiological adaptations of [a target species] in a similar in different patches. If the implication is ani-
time-unlimited sense” (Guthery 1997, 294). The usable mals of different species on the same area, then habitat
space concept posits maximization of usable space in could be the classical definition (i.e., natural abode).
time (space–time) as the goal of management. Space– In any case, the statement is ambiguous because habi-
time is the product of usable space and time (e.g., ha- tat could mean a place where animals live, an area, a
days). The concept also introduces a binary circum- physiognomic type within an area, or a circumstance at
stance—space is usable or not usable—which may be a small scale (e.g., a perch).
viewed as a severe constraint on the concepts of crude
(density on an area) and specific density (density on
Example 2
that portion of an area that provides usable space). The
classical definition of habitat also creates a binary cir- “Demographic information on survival and fecundity
cumstance (is habitat, is not habitat). will aid greatly in establishing the quality of a particu-
lar habitat” (Stauffer 2002, 58). The phrase “a partic-
ular habitat” is telling because it implies one of a set
Deductions
of habitats, which could imply subdivisions in an area.
For this section, we searched articles in journals and However, the demographic variables imply, but do not
chapters in books for the word habitat. Then, we at- confirm, an area of sufficient size to sustain at least a
tempted to deduce the meaning of habitat intended segment of a population. Here again, habitat seems to
(implicitly) by the author(s) based on the context in be an area that differs somehow (perhaps arbitrarily)
which the word was used. For brevity, we will use the from other areas in space. It could also apply to a set of
term natural abode to denote implicit meanings that natural abodes. However, to the extent that a natural
match the classical meaning. The following examples abode includes “the set of requisites of an organism,” it
are not exhaustive but rather provide an overview of is illogical to expect variation in habitat quality except
deduced meanings. in a binary sense: is, is not. That is, all requirements
are consummated when habitat is a set of requisites.
No further need exists.
Example 1
As researchers were seeking “the truth underlying
Example 3
how animals were relating to their habitats, managers
were seeking ways in which to use such information “The landscape metrics examined here are valuable de-
to address their needs” (Stauffer 2002, 56). “Animals,” scriptors of wildlife habitat” (Trani 2002, 151). We have
 exploration and critique of habitat and habitat quality 13

landscape meaning area and wildlife habitat implying We see the word habitat used to specify a landscape
a suite of species. The metrics could be viewed as sum- (area), a subcomponent of that landscape (element),
mary indices with or without explicit subcomponents and a property of the landscape (conditions) or its ele-
(e.g., patches). For example, canopy coverage of trees ments relative to an assumed optimal condition. We are
has a single basis in the landscape. Simpson’s diver- uncertain about the last meaning.
sity of patches deals with multiple components of the
landscape. Trani (2002) seems to mix habitat as area
Example 5
of interest, habitat as unique subcomponents of that
area, and habitat in the classical sense (natural abode). “The distribution of a species may be limited by the be-
havior of individuals in selecting their habitat” (Krebs
1972, 29). Selection implies two or more habitats (se-
Example 4
lection is not an issue with one habitat). If we view
“One objective of habitat relationship research is to selection from the standpoint of habitat as a natural
identify biologically important variables that have the abode, then we can reduce selection opportunities to
ability to predict species occurrence . . . Habitat quality two types: natural abode and not a natural abode. In
for many species contains a spatial component related this case, selection is moot because we know tautologi-
to the arrangement and amount of habitat elements cally that all use will, by definition, be in the natural
across the landscape. If the spatial scale of measure- abode. Krebs might be implying that natural abodes
ment alters the values of these variables, this influences may be classified according to their properties, e.g., oak
our ability for predicting species occurrence and for (Quercus sp.) forest natural abode, hickory (Carya sp.)
assessing habitat conditions” (Trani 2002, 151). forest natural abode, and so on. Such classification is a
In this paragraph (sixty-six words), we have habi- product of human caprice, not animal adaptation. An
tat relationship, habitat quality, habitat elements, and animal might view oak and hickory abodes more gen-
habitat conditions. erally as deciduous tree abodes. However, if we view
landscapes as containers for a mixture of not-natural-
• Habitat relationship involves relations between abode elements and distinct-natural-abode elements,
wildlife and one or more habitat elements. So, we would see selection behavior that limits the dis-
the “habitat” in “habitat relationship” refers to tribution of a species. At face value, Krebs’s statement
elements that could be synonymous with patches, seems ambiguous, but perhaps that is a problem of its
plant communities, cover types, or components. interpreters.
The “habitat” in “habitat elements” probably refers
to the landscape (area). For example, we could say
Example 6
“landscape elements” instead of “habitat elements”
with no change in meaning when landscape and “Semiannual population counts . . . suggest that fall
habitat are synonyms. habitat conditions on [the study area] may have de-
• Habitat quality involves “a spatial component,” so clined relatively more than did late-winter habitat . . .
because of a single component we assume habitat This is not to imply that autumn conditions were more
refers to the landscape (area) and some property of severe than those of late winter, only that habitat de-
the habitat elements in the landscape (e.g., fractal terioration was first apparent during the season of
dimension). highest density . . . Likewise, Robinson . . . considered
• Habitat conditions (plural) perhaps imply the bobwhite carrying capacity . . . to be primarily a func-
condition of habitat elements, but this does not tion of habitat available in late autumn” (Roseberry and
make sense because our interest is in a property of Klimstra 1984, 91–92).
the landscape (habitat quality). Perhaps the phrase In the last sentence, Robinson’s statement could
“habitat conditions” refers to some unstated apply to habitat defined as usable space, i.e., area (ha)
assumption(s) about existing conditions relative to compatible with occupation, given the adaptations of
optimal conditions, however that is defined. bobwhites. If this conclusion is acceptable, then habitat
14 foundation

conditions possibly mean the proportion of available tations thereof do not seem to hold because how can
space that is usable (habitat). Habitat deterioration im- a conceptual entity vary in quality if it supplies all the
plies loss of usable space because of concomitant loss of needs of an organism? Fortunately, Van Horne’s equa-
essential cover features such as hedgerows. We could tion 1 resolves the problem because the equation gives
also deduce from these statements that Roseberry and a mathematical definition of relative habitat quality
Klimstra described trends in the area (ha) of the birds’ (Q i) for area i.
natural abode. Consider the numerator in Van Horne’s (1983)
equation 1, which may be stated as
Example 7 Di = ((survival plus production for all age-classes on area
i) / (ai Ni)) ~ λi / ai ~ fitness density in habitat i
“Habitat fragmentation is usually defined as a
landscape-scale process involving both habitat loss and where ai is the area in the ith habitat (A = ∑ai), λi is the
the breaking apart of habitat” (Fahrig 2003, 487). We growth multiplier for the ith habitat, and Ni is the to-
are given a landscape dimension, but it would make tal population for the ith habitat. Van Horne’s analysis
little sense in context to say “landscape fragmentation”; implies that both the area of a subcomponent (ai) and
habitat adds the wildlife consideration. Moreover, we the total area (∑ai) are called habitat. Examples of total
deduce that the landscape must once have existed in areas could be a ranch, management area, study area,
a relatively unmodified state or a state with modifica- national wildlife refuge, and so on. This outcome (a
tions of minor and innocuous extent. The statement habitat composed of habitats) is consistent with the de-
seems to imply that habitat may be defined as areas fault and deduced definitions given previously, but not
within a landscape that supported sustaining wildlife with the classical definition of habitat (natural abode).
populations in the absence of fragmentation.
Definitions of Habitat Quality
Example 8
Van Horne (1983) provides a natural segue from habi-
Storch (1993, 256) studied habitat selection by caper- tat to habitat quality (P < 0.05). Because of the various
caillie (Tetrao urogallus) in summer and autumn. Habi- meanings of habitat, we will point out what we deduce
tat to Storch was a distinct patch based on vegetation to be the referenced authors’ meaning in the following
and topographic variables (subjectively designated discussion. This protocol will help to insure common
physiognomic type) on a large area (50 km2). We can understanding of concepts.
further define habitat according to the twenty-five vari- The word quality has at least nine meanings, includ-
ables listed in Storch’s table 1. Presumably, the variables ing the configuration of the oral cavity when stating a
listed are known or assumed components of habitat vowel. We will use the word as an index of superiority
as classically defined (natural abode). “[W]e almost or excellence based on human value. In the absence of
always provide a long list of variables for measurement human value, the words excellence and superiority have
that are hopefully related to what the animal perceives no meaning.
as habitat” (Morrison 2012, 8). Undoubtedly, there are Let Di = fitness density in subcomponent habitat i as
hundreds of papers in the ecological literature that defined previously. Then relative habitat (area) quality
consider habitat to be a list of variables that somehow as defined by Van Horne (1983) is
characterize the natural abode of an organism.
Q i = Di / ∑Di.
Relative habitat quality has no general meaning be-
Example 9
cause it is specific to any particular sample. However,
Van Horne’s (1983) classic paper is titled “Density as useful information may be in subcomponent estimates
a Misleading Indicator of Habitat Quality.” How might of fitness density (Di). Use of the word “fitness” in this
we define habitat to portray the meaning of the word in definition is unfortunate because, with the exception
this paper? Again, the classical definition and permu- of rare circumstances, humans can only measure sur-
 exploration and critique of habitat and habitat quality 15

rogates of fitness (e.g., survival and fecundity in instan- the asymptotic limit of population growth under the
taneous time scales relative to the time scale of fitness). logistic model. Such variation seems plausible for large
Before discussing other concepts of habitat quality, herbivores in response to variation in soil fertility and
we want to further comment on Van Horne’s (1983) plant nutrient concentration (Midgley 1937; Strick-
paper. The title is slightly ambiguous because of the land and Demarais 2000; Jones et al. 2008).
phrase “density as a misleading indicator” (emphasis Another way for carrying capacity density to vary is
added). The word as is quite complex (at least twenty- for the proportion of space that is usable on subcom-
one meanings including idioms). The Van Horne title ponent ai to vary among subcomponents. Suppose Ki / ai
could be construed as “density is a misleading indica- = pC = C if the proportion of space usable is p = 1.0.
tor” or “density may be a misleading indicator.” In the So, in general
first case, people would never knowingly use density
Ki / ai = piC
as a correlate of habitat quality (habitat here defined in
the Van Horne sense as an area of interest composed where pi = the proportion of space usable on area i.
of subareas). In the second case, which Van Horne in- Put differently, the aforementioned equation states that
tended, one would have to justify using density as an crude density is proportional to specific density. It also
indicator of habitat quality because it could be mislead- states that carrying capacity density is proportional to
ing. Some people think in terms of the first example, usable space on habitat subcomponent i. This is tan-
i.e., density is a misleading indicator. In fact, density tamount to saying that the quality of habitat subcom-
may be the only reasonable estimator of habitat (some ponent ai is proportional to usable space in the sub-
arbitrary area) quality. component, and this would generalize to habitat as an
Van Horne’s (1983) equation implies that habitat arbitrary area of interest (e.g., ∑ai).
(subcomponent) quality increases with survival and / or In discussing measures of habitat quality, Johnson
fecundity. This may indeed be the case in trending pop- (2007) used patch and habitat as synonyms. He also
ulations over a defined period of time. Examples may used arbitrary area consisting of patches as a synonym
include population growth following disease outbreaks for habitat. This outlook is analogous to Van Horne’s
and extreme environmental events that decreased view (habitat as a collection of habitats).
population numbers. In addition, sustainable harvest Johnson (2007) defined habitat quality at the level of
rates may be an indicator of habitat (subcomponent) an individual bird as “per capita contribution to popu-
quality for game species. Yet if density-dependent pro- lation growth expected from a given habitat.” What is
cesses influence these demographic variables, which implied by habitat in this definition is not clear, but
seems to be a universal circumstance (Brook and habitat as a collection of patches (essentially an area
Bradshaw 2006), then we encounter a dilemma. In composed of habitat types) seems to be a reasonable
a sustaining, nontrending population, higher survival guess. The definition also raises a question about habi-
entails lower fecundity and higher fecundity entails tat quality in a sustaining population (λ = 1). We would
lower survival. If these adjustments do not occur, the deduce from Johnson’s definition that habitat quality
finite annual growth rate exceeds 1.0 (λ > 1.0) and the does not vary for sustaining populations, and we would
population increases exponentially (Guthery and Shaw conclude that whether a population sustains tells us
2013). If we constrain interpretation to sustaining all we need to know about the quality of its habitat
populations, then the only useful demographic index (natural abode or arbitrary area).
of habitat (subcomponent) quality is density given λ = There seems to be general, somewhat rote agree-
1.0. So density may be not only a misleading indicator ment that habitat (area or subcomponent) quality is
in some settings but the only useful indicator of habitat poor in the case of declining populations and good in
(area) quality in other settings. the case of sustaining and growing populations. Of
This outcome begs the question, how can fitness course, such a perception of quality is subject to many
density (Di = λi / ai) vary across habitat subcomponents confounding factors such as time, weather, predation,
(ai)? One way is for carrying capacity density (Ki / ai) competition, standing density, and so on. Further, the
to vary among subcomponents where Ki is similar to perceived quality of an area (called habitat) may vary
16 foundation

ecologically and tribally with properties such as inter- sition (i.e., annual seed producers, perennials, etc.),
spersion and diversity. (By tribal variation, we mean [and] vegetation structure (for escape cover, thermal
variation imposed by human values and allegiance to cover, visual isolation, etc.)” (K. K. Karrow, Marais des
like-thinkers. For example, diversity is a desired prop- Cygnes Wildlife Area, personal communication). In
erty of human social systems, but more diversity is not this statement, habitat means an area of interest and
necessarily a meaningful property of ecosystems be- also patches within the area (types).
cause population responses to diversity may become
asymptotic [Guthery 2008].) We discuss some of these
Discussion
broader aspects of habitat (area) quality in the follow-
ing text. As is evident from the preceding material, the word
One respondent reported that he judges “the qual- habitat has multiple meanings in theory and practice.
ity of . . . habitat [area] based upon [his] experience To recapitulate, biologists use it primarily to mean
as a predator (hunter) and hence based on a search
image . . . for where [he has] found [the quarry] in pre- • where an organism lives, which includes biotic
vious outings.” Actually, the respondent has developed and abiotic requisites and time.
an ad hoc habitat suitability index that scores woody • an arbitrary area of interest (habitat) that contains
cover, nesting cover, food supplies, and interspersion demographically or behaviorally unique subareas
on a 0–10 scale. Habitat suitability models are quanti- (habitats) that may in turn contain descriptively
tative methods of assessing habitat (area) quality. unique subareas (habitats). For the latter compo-
“Bobwhite habitat quality should be judged by bob- nent, vegetation type, ecotype, physiognomic type,
white density given average environmental conditions habitat type, habitat (when used as synonym for
for the site” (C. B. Dabbert, Texas Tech University, habitat type), plant community, element, patch,
personal communication). This definition relates to an and so on are used interchangeably with habitat.
area that is called a habitat. The definition implicitly • gibberish (i.e., meaning apparently indeci-
invokes the concept of usable space. pherable).
“Ultimately, [habitat] quality comes down to the
fitness of individuals within the habitat. We use prox- These definitions break into three natural classes:
ies for this: survival and yearly reproductive output. At classical, hierarchical, and irrational. The classical
certain times of the year, density also might be a good outlook includes definitions with meanings similar
proxy for quality” (A. T. Pearse, US Geological Survey, to “natural abode.” The hierarchical outlook is an ar-
Northern Prairie Wildlife Research Center, personal bitrary area (level 1) with (demographically, behavior-
communication). This communication also seems to ally) meaningful subdivisions (level 2) that are in turn
imply that habitat is a synonym for area of interest (e.g., subdivided (level 3), which may in turn be subdivided
density is number / area). (level 4), and so on. We see no reason why higher level
“I would argue that wetland (read: habitat) of 1 ha subdivisions might not be meaningful (e.g., a wetland
capable of providing one hundred duck use days is bet- plant community habitat could have open water and
ter quality than a 1-ha wetland capable of providing fifty vegetated components that could be called habitats).
duck use days” (R. O’Shaughnessy, Southern Illinois The irrational outlook includes any use of habitat in an
University, personal communication). Here habitat is indecipherable manner.
a type of plant community or substrate, and the quality Note that a unit composed of population-supporting
of that community or substrate is based on the food subunits in the Van Horne (1983) sense would seem
supplies it holds. Judging habitat (area) quality based to be uncommon in the study of hierarchical habitat.
on food abundance is common in wildlife biology. Rather, the most common expression is an arbitrary
Variables for evaluating habitat quality for mallards area of interest composed of subunits that are unique
(Anas platyrhynchos) include “diversity (i.e., the array in some respect. Such subunits are essential for habitat
of habitat types and condition, and their juxtaposition (subunit) preference studies, as is evident from John-
within the area of interest), vegetation species compo- son (1980), who dealt with habitat types.
 exploration and critique of habitat and habitat quality 17

Does the classical definition hold sway over the hi- support for spatial or temporal differences in habitat
erarchical definition, or vice versa, as the appropriate (subcomponent) quality, but only for a defined area
model for research and management? Who is to say? and span of time. Moreover, many studies ignore the
Both seem to have a place and, besides, both will be dilemma of density dependence (Guthery and Shaw
with us forever. 2013): in sustaining populations, increases in produc-
Logically, the classical version of habitat seems to tivity entail decreases in survival and vice versa. Thus,
be rather a dead- end concept except for descriptive higher nest success in area A could have no influence
science. But this descriptive science is by no means on year-to-year trends in abundance because of the
trivial. Imagine how difficult it would be to identify the compensatory mechanisms of density dependence.
minimal set of factors that would describe an organ- Over time, the dilemma of density dependence holds
ism’s habitat in the classical sense. Guthery (2002), for with annual variation in survival and production.
example, posited that bobwhite habitat could be fully Habitat quality is a dearly held concept based on
described with five variables: canopy coverage of woody human value. It apparently sprang into use as an un-
vegetation, visual exposure to ground predators, visual questioned property of nature that is based on fitness,
exposure to aerial predators, coverage of bare ground, something that is difficult to measure except for its soft
and operative temperature. Imagine how research to correlates. The concept needs rigorous theoretical jus-
reveal minimum sets of factors that characterize a spe- tification and clarification if it is to be useful in build-
cies’ habitat (classical sense) could lead to original and ing knowledge about wildlife.
meaningful hypotheses about behaviors and processes
in the field. The main purpose of science is to extract
Recommendations
simplicity from complexity (Cohen and Stewart 1994).
The hierarchical view of habitat fits research in that Because the word habitat is intrinsically ambiguous, as
it represents sequential parsing—the study of an entity its classical, hierarchical, and irrational forms show,
composed of parts that are in turn composed of parts authors should state their intended meaning early in a
and so on. This approach provides a natural means of paper and then religiously apply that meaning through-
studying how animals partition time and occurrence in out the paper. This would serve the goal of making the
third-level units that we call habitats or the synonym concepts in papers consensible. In other words, authors
habitat types. and readers would have a common understanding of
Now let us turn to the concept of habitat quality. meaning.
Under the classical definition of habitat, habitat quality To forestall overuse and misuse of the word, we rec-
is a meaningless concept to which we have alluded to ommend that authors do a word search for habitat after
the point of redundancy. Classical habitat is an exis- finishing a paper. Then, at each encounter of the word
tential matter—it either exists or it does not. Thus, in he or she can check to see whether any particular usage
a spatial sense, there is habitat (quality = 1) and not- matches the stated definition. If it does not, perhaps
habitat (quality = 0). authors can substitute a term that is more explicit than
In the hierarchical sense, a consensus prevails that habitat wherever possible. For example, vegetation
habitat (area) quality varies with some index of fit- types are synonyms for habitat in technical parlance, so
ness as determined by studies that are instantaneous to be precise one could use prairie, oak forest, meadow,
relative to the grand sweep of time over which fitness cattail community, and other explicit synonyms instead
prevails. One might conclude that habitat A (area or of minestrone.
subcomponent) is better than habitat B (area or sub-
component) because nest success was higher in habi- ACKNOWLEDGMENTS
tat A. This inference is weak regardless of the P-value We thank T. E. Fulbright and C. A. Davis for reviewing
or model weight because it provides no information on a draft of this chapter. J. D. Stafford and V. A. O’Brien
other demographic variables such as survival of various contributed ideas and information. We recognize that
age-classes. Studies that document the corresponding these colleagues do not necessarily agree with all the
values for recruitment and adult survival may provide views expressed here.
18 foundation

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Block, W. M., and L. A. Brennan. 1993. “The Habitat Concept ogy: Towards a Common Language to Advance Ecological
in Ornithology.” Current Ornithology 11:35–91. Understanding and Application.” In Predicting Species Oc-
Brook, B. W., and C. J. A. Bradshaw. 2006. “Strength of Evi- currences: Issues of Accuracy and Scale, edited by J. M. Scott,
dence for Density Dependence in Abundance Time Series P. J. Heglund, M. L. Morrison, J. B Haufler, M. G. Raphael,
of 1,198 Species.” Ecology 87:1445–61. W. A. Wall, and F. B. Sampson, 43–52. Island Press, Wash-
Cohen, J., and I. Stewart. 1994. The Collapse of Chaos. Penguin ington, D.C., USA.
Books, New York, USA. Oxford online dictionary. http: // www.oed.com / view / Entry
Fahrig, L. 2003. “Effects of Habitat Fragmentation on Biodi- / 82988 (accessed June 25, 2014).
versity.” Annual Review of Ecology, Evolution, and Systematics Roseberry, J. L., and W. D. Klimstra. 1984. Population Ecology of
34:487–515. the Bobwhite. Southern Illinois University Press, Carbon-
Garshelis, D. L. 2000. “Delusions in Habitat Evaluation: dale, USA.
Measuring Use, Selection, and Importance.” In Research Safriel, U. N., and M. N. Ben-Eliahu. 1991. “The Influence of
Techniques in Animal Ecology, edited by L. Boitani and T. K. Habitat Structure and Environmental Stability on the Spe-
Fuller, 111–64. Columbia University Press, New York, USA. cies Diversity of Polychaetes in Vermetid Reefs.” In Habitat
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ner’s Sons, New York, USA. Mushinsky, 349–69. Chapman and Hall, New York, USA.
Guthery, F. S. 1997. “A Philosophy of Habitat Management Smith, R. L., and T. M. Smith. 2001. Ecology and Field Biology.
for Northern Bobwhites.” Journal of Wildlife Management San Francisco: Addison Wesley Longman.
61:291–301. Stauffer, D. F. 2002. “Linking Populations and Habitats: Where
———. 2002. The Technology of Bobwhite Management. Iowa Have We Been? Where Are We Going?” In Predicting Species
State University Press, Ames, USA. Occurrences: Issues of Accuracy and Scale, edited by J. M.
———. 2008. A Primer on Natural Resource Science. Texas A&M Scott, P. J. Heglund, M. L. Morrison, J. B Haufler, M. G. Ra-
University Press, College Station, USA. phael, W. A. Wall, and F. B. Sampson, 53–61. Island Press,
Guthery, F. S., and J. H. Shaw. 2013. “Density Dependence: Washington, D.C., USA.
Applications in Wildlife Management.” Journal of Wildlife Storch, I. 1993. “Habitat Selection by Capercaillie in Summer
Management 77:33–38. and Autumn: Is Bilberry Important?” Oecologia 95:257–65.
Hall, L. S., P. R. Krausman, and M. L. Morrison. 1997. “The Strickland, B. K., and S. Demarais. 2000. “Age and Regional
Habitat Concept and a Plea for Standard Terminology.” Differences in Antlers and Mass of White-Tailed Deer.”
Wildlife Society Bulletin 25:173–82. Journal of Wildlife Management 64:903–11.
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Johnson, M. D. 2007. “Measuring Habitat Quality: A Review.” and Scale, edited by J. M. Scott, P. J. Heglund, M. L.
Condor 109:489–504. Morrison, J. B Haufler, M. G. Raphael, W. A. Wall, and F. B.
Jones, P. D., S. Demarais, B. K. Strickland, and S. L. Edwards. Sampson, 141–55. Island Press, Washington, D.C., USA.
2008. “Soil Region Effects on White-Tailed Deer Forage Van Horne, B. 1983. “Density as a Misleading Indicator of Hab-
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 3 Demographic
Consequences of Habitat
Amanda D. Rodewald

C onserving wildlife populations often warrants a

habitat-based focus, given that the amount, dis-
tribution, and quality of habitat can strongly mediate
as well as the overall quality of habitat. Habitats are
generally considered to be of “high quality” when they
support and promote health, survival, and / or repro-
population demography. The manner in which demog- duction of individuals within a population. However,
raphy is affected by habitat is determined via numerous many approaches to characterizing wildlife-habitat re-
behavioral, physiological, environmental, and stochas- lationships often fail to tightly link habitat parameters
tic processes. In this chapter, I discuss how the tradi- to population dynamics and viability.
tional view of habitat-population relationships is being The apparent disconnect between habitat and popu-
challenged by new insights from studies that illustrate lation dynamics may result from several shortcomings
the need to better (1) distinguish between individual of our traditional view of the wildlife population–hab-
and population-level measures of habitat quality as they itat link. First, a failure to distinguish between indi-
relate to population ecology, (2) address habitat needs vidual and population-level measures of habitat quality
and demographic connectivity across the full life cycle, can obscure identification of the highest quality habitat
(3) recognize how behavioral processes guiding habitat and, thus, fail to identify the management approach
selection influence population dynamics, and (4) elu- that best achieves desired outcomes. Demographic
cidate heterogeneity in the demographic consequences measures framed from the two perspectives can lead
of habitat use over a wide range of spatial and temporal to seemingly contradictory conclusions about habitat
scales. Addressing these needs will strengthen wildlife quality and its impact on populations. Second, descrip-
science and management by improving our ability to tions of habitat still have a heavy bias toward a single
understand wildlife-habitat relationships and predict life stage or season—usually the breeding season. This
the demographic consequences of habitat use. tendency overlooks the now-rich literature demonstrat-
ing that habitat requirements can vary across different
life stages and that events operating across the full life
Introduction
cycle govern population dynamics. Third, the ways
The identification, protection, and restoration of suit- in which population dynamics can be influenced by
able habitat are essential components of many, if not individual behaviors related to habitat selection are
most, wildlife conservation and management efforts. generally underrecognized. The “if you build it, they
Because habitat directly or indirectly mediates the rates will come” view of restoration and management fails
of birth, death, immigration, and emigration for any to recognize that a diverse suite of environmental, bio-
species, most biologists and managers consider demo- logical, and social cues are used by animals to select
graphic information to be the gold standard for deter- habitats (Morrison 2009). For example, if the pres-
mining which elements of habitat are most important, ence of conspecifics is the primary cue used by a spe-
20 foundation

cies to select habitat, newly created high- quality but Demographic measures at individual and popula-
vacant habitats may not be occupied. In cases where tion levels may prove contradictory when densities do
cues for settlement are lacking, even the best habitats not align with fitness measures. Even in the simplest
may have weak demographic signals because they are cases where life stages do not vary in fitness, demo-
less preferred, settled later, and / or occupied by lower graphic rates as measured by per capita fecundity may
quality individuals. Fourth, spatiotemporal variation in not parallel population growth rates, or the product of
habitat-specific demography is often overlooked or not density and per capita fecundity (Johnson 2007; Ska-
sufficiently examined because many studies of wildlife- gen and Yackel Adams 2011). This lack of concordance
habitat relationships occur at limited spatial and tem- between individual and population metrics is not a
poral scales and fail to fully capture interpopulation trivial issue from a management perspective. Should
variation in the ways animals interact with their envi- a manager give preference to a habitat that supports
ronment. Understanding the causes and consequences the greatest individual fecundity or to that which pro-
of spatiotemporal variability in demography is essential duces the greatest number of young from the local
to developing effective strategies that ensure the long- population—something that also is function of density
term viability of populations. Permeating throughout (fig. 3.1)? For example, suppose Site A has high quality
each of these points is the fact that most habitat stud- resources and produces higher per capita fitness (2.3
ies give weak consideration to ecoevolutionary links young per female) than Site B, which has lower quality
and feedbacks that are especially relevant in systems resources and fitness (1.6 young per female). If Site A
where anthropogenic change has altered selective supports fewer individuals (e.g., 10 breeding pairs ×
environments. 2.3 young / pair = 23 young per year), then it would
This chapter provides an overview of recent devel- contribute less to population viability than the lower
opments that are changing our understanding of the quality Site B with higher densities (e.g., 20 pairs ×
relationship between habitat quality and population dy- 1.6 young / pair = 32 young produced annually). The
namics. I organize the discussion around four key top- contradiction is, then, that from an individual’s per-
ics where recent advances have provided new insights spective, Site A would be of better quality, whereas at
into the interrelationship of habitats and populations. a population level, Site B would better support a large
local or regional population.
Empirical evidence shows that the relationship
Individual- versus Population-Level
between individual and population measures can vary
Measures of Habitat Quality Can Provide
widely. For example, density was positively associated
Different Information, and Sometimes
with recruitment per capita (individual level) for 72%
Contrasting Perspectives, About
and with “reproduction per land area” (population
Population Dynamics
level) for 85% of 109 studies reviewed (Bock and Jones
The idea that density does not necessarily indicate 2004). Negative relationships between density and
habitat quality became both widely recognized and fitness, on the other hand, can result from regulatory
well accepted after Van Horne’s (1983) classic paper mechanisms like inverse density dependence (Brook
suggested that habitat quality should reflect mean in- and Bradshaw 2006). In other cases, the negative rela-
dividual fitness per unit area, which was defined as a tionship can signal the presence of an ecological trap,
function of density, production of young, and survival. where individuals show preferences for habitats that
Taken at face value, use of demographic parameters result in the poorest recruitment. Thus, the variable
to indicate habitat quality is intuitively attractive and relationship between density and fitness can obscure
straightforward. However, as discussed in previous efforts to identify the best quality habitats. For ex-
papers (Chalfoun and Martin 2007; Johnson 2007; ample, Pidgeon et al. (2006) empirically demonstrated
Mortelliti et al. 2010), descriptions of habitat quality that fecundity was the best indicator of individual-level
can be vague, assessed using the wrong fitness metrics habitat quality for black-throated sparrows (Amphispiza
or at inappropriate spatial scales, and often conflate bilineata), but the most robust indicator of habitat qual-
individual and population perspectives. ity at the population level was the combination of nest
 demographic consequences of habitat 21

amounts of low quality food and had higher carrying

capacities (Hobbs and Swift 1985).
In terms of management, biologists need to explic-
itly consider both levels, as the most relevant likely
depends upon the objective. If one is studying habitats
with the aim to guide future management and resto-
ration choices by describing resources that are most
advantageous to a focal species, then individual-level
measures of quality will best elucidate those “ideal” re-
sources or conditions. After all, fitness is an individual
measure and has strong evolutionary components, and
managers need to identify the relative contribution to
the population by individuals occupying a given habi-
tat, also known as habitat fitness potential (sensu Wiens
1989a). Knowing the fitness potential of various habi-
tats makes it possible to discern the effects of landscape
composition on population dynamics and to identify
optimal configurations (Pulliam 2000; Griffin and
Mills 2009 Runge et al. 2006). However, if one aims to
identify which existing sites or locations are most im-
portant to long-term persistence, then population-level
measures, or population mean fitness, are most likely
to indicate where management can have the greatest
impact on population viability (fig. 3.2).

Figure 3.1 Logistic population growth curves for two hy- Population Dynamics Are Influenced by
pothetical sites: site A with few high-quality resources and Habitat Used across the Full Life Cycle
site B with abundant lower-quality resources. Differences
between habitats result in a higher rate of population Many species require multiple habitats to meet their
growth in A (rA = 0.12, rB = 0.03) and a higher carrying ca- needs across the full life cycle or across life stages,
pacity in B (KA = 500, KB = 1000). Johnson (2007) simulated and changing needs complicate efforts to link popu-
both populations with initial population sizes of 100 and lation responses to habitat attributes. Trying to under-
run for 100 time intervals. If habitat quality is considered stand how habitat alteration contributes to population
purely from an individual bird’s perspective, then Habitat changes for species that rely upon multiple habitats,
A is the better habitat until time 23. In contrast, if habitat
or even multiple locations of the same habitat type, re-
quality is measured as the current population size, then
quires determining at which location or stage habitat
Habitat A remains better until time 74. If habitat quality
might limit the population. This challenge cuts across
is considered the maximum sustained population size, as
may be the perspective of many conservationists, then
taxa, as 80% of animals have complex life cycles that
Habitat B is better because it has the higher carrying involve major ecological transitions between life stages
capacity. From Johnson 2007 (Werner and Gilliam 1984). Even nonmigratory ani-
mals may require different habitats. Consider pelagic
seabirds that spend most of their lives at sea but re-
success (or fecundity) and nest density. Another ex- quire terrestrial habitats, often on islands, for nesting.
ample relates to mountain shrub communities, where For other species, disparate habitat needs are met in
recently burned habitats provided small amounts of ex- heterogeneous habitats, as with lesser prairie chickens
tremely high quality food to a lower carrying capacity of (Tympanuchus pallidicinctus) for which various repro-
mule deer, whereas unburned habitats provided large ductive behaviors, such as courtship displays, nesting,
22 foundation

Figure 3.2 An experiment on the effects of canopy reduc- stages (Wilbur and Collins 1973; Wilbur 1980). The
tion on forest-breeding birds showed that sites reduced
red-spotted newt (Notophthalmus viridescens) is aquatic
from ~28 m2 / ha residual basal area (73% canopy cover) (A)
during larval and adult stages, requiring ephemeral
to 14 m2 / ha residual basal area (45% canopy cover) (B)
supported 115% higher densities of Cerulean Warblers com-
vernal pools and streams. However, its juvenile stage,
pared to unharvested control stands (Boves 2011). Although the red eft, is terrestrial and uses upland forest. Many
reproductive output per breeding pair was 40% lower on dis- birds that breed in mature forest, such as wood thrush
turbed sites than on unharvested controls, the higher densi- (Hylocichla mustelina), ovenbird (Seiurus aurocapilla),
ties resulted in a much greater contribution of individuals and worm- eating warbler (Helmitheros vermivorum),
to the local and regional population from disturbed sites shift to using early-successional or shrubby habitats
than from undisturbed sites. Photos by Amanda Rodewald during postbreeding and postfledging periods (An-
ders et al. 1998; King et al. 2006; Vitz and Rodewald
and brood rearing, are each associated with different 2006). Vitz and Rodewald (2006) captured nearly 90%
structural attributes of rangelands (Fuhlendorf and of birds that bred in mature forests within regenerat-
Engle 2001). Likewise, flying foxes in Australia use fruit ing clearcuts during the postbreeding period in Ohio,
resources in rainforests as well as nectar and pollen in with numbers of mature-forest species rivaling those
coastal heath, swamps, and Eucalyptus forest (Law and of early-successional specialists like the prairie warbler
Dickman 1998). (Setophaga discolor) and eastern towhee (Piplio eryth-
rophthalmus). Such observed shifts in habitat use are
thought to result from selection for dense cover to re-
Variation in Habitat Use and Demography
duce risk of predation and / or abundant fruit resources
across Life Stages
to facilitate foraging (Vitz and Rodewald 2007). In-
Most studies define habitat in terms of a single stage deed, use of habitats with dense vegetation promoted
in the annual or life cycle, with breeding being the survival of fledgling ovenbirds and worm-eating war-
most common. However, habitat use can vary widely blers (Vitz and Rodewald 2011). Thus, managers are
across annual and life stages, particularly for migratory increasingly recognizing the need to address habitat
species whose movements span continents. Another requirements of both breeding and postbreeding / post-
striking habitat shift is for amphibians that undergo fledging individuals in conservation strategies for for-
metamorphosis through aquatic and terrestrial life est birds.
 demographic consequences of habitat 23

Many species use multiple habitats across the an- condition of individuals in subsequent seasons. Sea-
nual cycle, and for migratory species the spatial scale sonal interactions at the individual level are oftentimes
over which these habitats occur can be staggering. mediated by body condition, which tends to be closely
Neotropical migratory birds densely occupy disturbed related to habitat quality. However, there can also be
or second-growth forests, including agroforestry habi- population-level interactions that result from changes
tats, in Central and South America (Greenberg et al. in population size in one season that affect per capita
1997; Petit and Petit 2003). Shade-coffee farms are rates in subsequent seasons (Norris and Marra 2007).
agroforestry habitats that appear to provide high qual- From a practical standpoint, the presence of seasonal
ity habitat and are associated with high overwinter interactions means that demographic parameters mea-
survival, site fidelity, and energetic condition (Johnson sured during one season may not indicate habitat qual-
et al. 2006; Bakermans et al. 2009). Large-scale migra- ity if there are strong carryover effects from previous
tions obviously require conservation of multiple habi- seasons and locations.
tats contiguously distributed across large regions, as is Carryover effects have been described for a wide
the case for barren-ground caribou (Rangifer tarandus) range of taxa (reviewed in Harrison et al. 2011), includ-
that can migrate over five thousand km, pronghorn ing mammals (Festa-Bianchet 1998; Perryman et al.
(Antilocapra americana) over five hundred km, moose 2002; Descamps et al. 2008), reptiles (Broderick
(Alces alces) nearly four hundred km, and mule deer et al. 2001), and fish (Bunnell et al. 2007; Kennedy et al.
(Odocoileus hemionus) over two hundred km in their 2008). In seven of eight ungulates in Kruger National
round-trip movements (reviewed in Berger 2004). Al- Park, South Africa, juvenile survival was strongly re-
though breeding seasons are known to profoundly af- lated to rainfall, and therefore food supply, during the
fect population dynamics (e.g., Hoekman et al. 2002), previous dry season (Owen-Smith et al. 2005). In roe
the demographic consequences of nonbreeding habi- deer (Capreolus capreolus), resources encountered in
tat use can be equally important. Sillett and Holmes the spring strongly affected body mass during winter
(2002) showed that over 85% of the apparent annual (Pettorelli et al. 2003). The size of winter food cache,
mortality occurred during migration for black-throated in part, determined breeding condition of wolverines
blue warblers, which suggests that conservation efforts (Gulo gulo; Persson 2005). Black-throated blue war-
directed at migratory periods are likely to strongly in- blers (Setophaga caerulescens) overwintering in high-
fluence population viability. Fortunately, population quality forest were in better breeding condition than
ecology tools, such as structured population models individuals occupying lower quality scrub habitats
and sensitivity analyses (see Mills 2012), can be used (Bearhop et al. 2005). Mass of nestling savannah spar-
to identify those stages and, by association, those habi- rows (Passerculus sandwichensis), which was related
tats that most strongly affect population dynamics. to fledging date, affected an individual’s ability to ac-
In addition, new approaches can quantify per capita, cumulate fat prior to migration (Mitchell et al. 2011).
habitat-specific contributions to population growth Food availability during stopover periods influenced
and, thus, allow for the examination of source-sink testis growth and reproductive behavior of garden war-
dynamics for animals using heterogeneous habitats or blers (Sylvia borin) in ways that can affect reproduction
landscape mosaics (Griffin and Mills 2009). (Bauchinger et al. 2009).
Not only can choices in one season have demo-
graphic consequences in subsequent seasons, but
Seasonal Interactions
habitat choice in one season can constrain choices
Just as a habitat can strongly affect performance, re- in the next by affecting the timing and speed of mi-
production, and survival, there also can be large ef- gration in ways that impact territory settlement and
fects that lead to seasonal interactions. Seasonal reproduction (Norris and Marra 2007). One classic
interactions, also known as carryover or lag effects, example is American redstarts (Setophaga ruticulla),
are distinguished from direct impacts on survival or for which individuals that overwinter in high-quality
reproduction within a single season by the fact that mangrove habitats arrive to breeding grounds earlier,
processes or events from one season affect the state or in better condition, and fledge more young than indi-
24 foundation

viduals that overwintered in poorer-quality scrub habi- fect populations in subsequent seasons depends on how
tats (Marra et al. 1998; Norris et al. 2004). Indeed, the they are regulated. In the case of density-dependence
winter-habitat model created by Norris et al. (2004) due to crowding effects, increases in population size
predicted that females occupying high- quality winter promote negative interactions among individuals (e.g.,
habitat would produce two additional young fledging a competition for food or mates) that depress reproduc-
month earlier compared to females from poor- quality tion (Fretwell and Lucas 1969). Alternatively, with
winter habitat. Thus, the consequences of resources site dependence, increases in population size result in
available to birds in their nonbreeding habitat car- some individuals being pushed into lower-quality terri-
ried over to influence their subsequent reproductive tories that, in turn, depress reproduction (Rodenhouse
success. et al. 1997). In contrast, individual-level effects happen
The likelihood of carryover effects may vary with life when events in previous seasons influence individual
history strategies. Populations of species with slow life performance in subsequent seasons via nonlethal
histories, such as being long-lived with low reproduc- mechanisms. Recent work shows how individual- and
tive output, should be more sensitive to events during population-level carryover effects can be incorporated
the nonbreeding season than fast life history (e.g., low into population models (Runge and Marra 2005; Nor-
survival but high reproductive output) species, which ris 2005; Norris and Taylor 2006).
should be most sensitive to breeding season events (Sa-
ether et al. 1996). Similarly, “capital” breeders finance
Population Dynamics Are Influenced
reproduction from energy stores accumulated in pre-
by Behavioral Processes Guiding
vious months (Drent and Daan 1980; Houston et al.
Habitat Selection
2007) and should face more carryover effects than “in-
come” breeders (Harrison et al. 2011). One example Most habitats are heterogeneous and variable, and in-
of this is gray whales (Eschrichtius robustus), for which dividuals lack complete knowledge about the state of
winter calving success depends upon fat stores accu- current and future conditions. Some important deter-
mulated during the summer (Perryman et al. 2002). minants of fitness may not be predictable, such as pre-
In one of the few carryover studies focusing on a year- dation risk (Doran and Holmes 2005), or recognized,
round resident species, blue tits (Cyanistes caeruleus) in the case of novel alterations to habitats (e.g., invasive
that received supplemental food during winter initi- plants; Lloyd and Martin 2005). Further complicating
ated breeding earlier in the season and with greater matters, the consequences of settlement decisions may
reproductive success than unsupplemented individuals not be realized until later in the season and long after
(Robb, McDonald, Chamberlain, and Bearhop 2008; the decision was made (e.g., when young fledge, at time
Robb, McDonald, Chamberlain, Reynolds, Harrison, of departure for migration). Nevertheless, individuals
and Bearhop 2008). In addition, for species that have have access to many types of information from which
individuals traveling between the same specific breed- they can base their choice of the most appropriate
ing and nonbreeding areas (Esler 2000), or high mi- strategies and / or habitats to occupy. Heterogeneous
gratory connectivity (Webster et al. 2002), seasonal environments create the opportunity for individual be-
interactions should be strong. haviors to influence population-level processes because
In their review of carryover effects across taxa spatiotemporal variation provides options from which
ranging from marine mammals to ungulates to birds, individuals choose. Choices have fitness consequences
Calvert et al. (2009) found that nonbreeding season that ultimately can affect population dynamics. The
events affected fitness in subsequent seasons at both ecology of information is the study of how organisms
the individual and population levels. For example, at acquire and use information and how this affects pop-
the population level, density dependence can manifest ulations, communities, landscapes, and ecosystems
across seasons when mortality or low reproductive (Schmidt et al. 2010).
rates affect the proportion of individuals able to occupy Access to information that is both public (i.e., acces-
higher-quality sites in subsequent seasons (Norris and sible to all individuals) and private (i.e., undetectable
Marra 2007). The ways in which changes in density af- to others and oftentimes based on the past experience
 demographic consequences of habitat 25

of individuals, such as prior reproductive success; Wag- example, are attracted to light from beachfront devel-
ner and Danchin 2010; Schmidt et al. 2010) can shape opment and move inland where they face greater risk
habitat-population links. Common cues used in habitat of mortality (Witherington 1997). Bobolinks (Doli-
selection include specific environmental conditions, chonyx oryzivorus) and other grassland species are at-
presence of conspecifics or heterospecifics, and repro- tracted to hayfields for breeding, despite the fact that
ductive success of conspecifics, such as fledgling calls harvesting of hay occurs before young fledge, thereby
or the presence of predators. When cues are related to reducing reproductive success (Bollinger et al. 1990).
attributes of the population itself, as with conspecific Decision rules can reduce or enhance the likelihood of
density and nest success, rather than specific habitat ecological traps. Through modeling, Kokko and Suther-
attributes, there is the potential for positive feedback land (2001) showed that natal imprinting, philopatric
that can dramatically affect population viability and preferences, or win-stay, lose-switch strategies tended
restoration potential (Schmidt et al. 2010). Therefore, to reduce the impact of ecological traps because indi-
from a practical standpoint, cues matter. For instance, viduals adjusted their habitat preferences. In general,
metapopulation persistence in patchy environments learned cues provide more opportunity for individuals
that have temporally correlated or predictable condi- to adjust their preferences, whereas cues with strong
tions was several-fold greater when fidelity was based genetic components promote the persistence of traps.
on previous successful experience than when disper- Because the cues and decision rules used in habitat
sal was uniform (Schmidt 2004; Schmidt et al. 2010). selection have important consequences for population
Conversely, reliance upon social cues like conspecific dynamics, they can provide insight into which conser-
attraction may reduce the likelihood that birds disperse vation and restoration strategies are most appropriate
to and colonize new or restored high-quality habitats and improve wildlife habitat models. For example, the
(Reed and Dobson 1993). absence of cues used in habitat selection can result in
The optimal dispersal and settlement strategies otherwise high-quality habitats going unused. Conspe-
are expected to vary based on pattern of spatial het- cific attraction can be a useful tool for conservation,
erogeneity and temporal predictability (Doligez et al. particularly when there is a need to attract individu-
2003; Schmidt et al. 2010). For sites with fine-scale als to unoccupied, newly created, or restored habitats
spatial heterogeneity (i.e., high variation among indi- (Reed and Dobson 1993), as has been well established
vidual sites within a patch), individuals should either for seabirds (Podolsky and Kress 1992). Experimental
(1) show high fidelity to sites if temporal predictabil- playbacks of conspecific vocalizations also showed that
ity is high or (2) disperse from unsuccessful sites and the federally endangered black-capped vireo (Vireo at-
prospect within the same habitat patch. For sites with ricapilla) was strongly attracted to conspecifics, and
coarse-scale heterogeneity (i.e., high variation among birds drawn to experimental sites often established
patches), individuals should prospect for information territories and bred, even in subsequent years (Ward
on patch quality and base decisions on average patch and Schlossberg 2004). Incorporating conspecific at-
quality when predictability is high but should prospect traction into wildlife-habitat models can improve the
during the prebreeding season when predictability is predictive ability of habitat models (Campomizzi et al.
low. Conspecific attraction, or the tendency of individ- 2008), especially when forecasting the outcome of re-
uals to settle near other individuals of the same species, introductions. Mihoub et al. (2009) showed that per-
is only favored when variation among patches is greater sistence of reintroduced populations depends heavily
than variation among sites and temporal heterogeneity upon habitat selection strategies when there is high
is high. heterogeneity among habitat patch qualities. Failure
An ecological trap, now a familiar phenomenon of reintroductions was more likely when species used
to scientists and managers, also is rooted in the ecol- conspecific presence or avoidance strategies, both of
ogy of information. Ecological traps form when once- which tended to aggregate individuals in suboptimal
reliable cues of habitat quality are no longer reliable habitats. Direct assessment of environmental cues or
due to altered environments (Schlaepfer et al. 2002). attraction to areas with high conspecific reproductive
Loggerhead sea turtle (Caretta caretta) hatchlings, for success was associated with higher success rates. Given
26 foundation

Table 3.1 Despite a fourfold difference in density, fitness metrics (± SE) for Northern cardinals (Cardinalis cardinalis)
were comparable in seven urban and seven rural forests in central Ohio, USA (Rodewald and Shustack 2008).
In this way, cardinals appear to be resource matching and conforming to ideal free distribution. The pattern is
consistent with predictions from IDF because densities are greatest in urban forests that contain greater levels of
resources used and preferred by cardinals, such as fruit, bird feeders, and understory woody vegetation (Leston
and Rodewald 2006).

Urban Rural

Density (birds per 2-ha grid) 1.30 (0.15) 0.31 (0.11)

Apparent survival, male 0.67 (0.05) 0.58 (0.06)
Apparent survival, female 0.53 (0.06) 0.63 (0.08)
Number of fledglings per pair 2.40 (0.18) 2.10 (0.18)
Nestling mass (7–9 days old; g) 24.80 (0.36) 24.70 (0.50)

these patterns, the authors suggest that reintroductions negative interactions increasing with density. In pop-
should favor release of adults, including females with ulations conforming to the ideal free distribution,
dependent young, and create artificial social informa- density should indicate habitat quality (for empirical
tion for use as cues. demonstration, see table 3.1). With ideal despotic dis-
Decisions about habitat selection, settlement, and tribution (Parker and Sutherland 1986), individuals
dispersal are among the most important behaviors that vary in competitive ability and the highest quality habi-
can regulate populations. Indeed, models of habitat se- tats are occupied by the strongest competitors, such
lection nicely illustrate how behaviors governing habi- that at equilibrium, fitness is lower in poorer quality
tat selection can have population-level consequences habitats. Competition and displacement are important
(Morris 2003). aspects of the ideal despotic distribution model. Under
this model, density is likely to be greater in poorer qual-
ity habitats and, thus, a misleading indicator of habitat
Random Settlement
quality. Consequently, before using density or distribu-
When breeding sites are settled at random (i.e., not re- tion to indicate habitat quality, one should know the
lated to their quality), population growth mirrors the extent to which a population conforms to ideal habitat
average quality of the habitat (Pulliam and Danielson selection.
1991; McPeek et al. 2001). Although average per-capita
fecundity of the population is not related to the number
Site Dependence
of sites occupied, the variation in average per-capita fe-
cundity increases as population size declines (McPeek Population regulation also can be accomplished solely
et al. 2001). One consequence is that small populations due to settlement behaviors and without any changes
might remain small for longer periods and be vulner- in individual demographic rates, and even by way of
able to demographic or environmental stochasticity. noninteracting individuals (Pulliam and Danielson
1991; Rodenhouse et al. 1997, 1999; McPeek et al.
2001). With site dependence, the negative relationship
Ideal Free and Ideal Despotic Distributions
between average per-capita fecundity of the population
Under the ideal free distribution (Fretwell and Lucas and population size results from the sequential use of
1969; Pulliam 1988), competitors are equal and select successively poor sites, rather than from declining pro-
habitats that maximize fitness with the equilibrium ductivity of a site as other sites are occupied or as popu-
distribution, resulting in equivalent fitness of indi- lations increase in size (McPeek et al. 2001). Popula-
viduals across habitats of variable quality. Population tions are regulated despite a lack of negative density
regulation is a consequence of vital rates of individuals dependence at the individual level. Site dependence
changing with population size, with fitness-depressing requires variation in the suitability of sites in terms of
 demographic consequences of habitat 27

demography, exclusive use of sites, and adaptive choice this means is that habitats of similar quality can differ
of sites (i.e., sites are settled sequentially from best to widely in terms of demography, simply as a function of
worst or “preemption”; sensu Pulliam and Danielson population size. Small populations are more likely to
1991) (McPeek et al. 2001). Because the best sites are have positive correlations between density and popu-
always occupied first, smaller populations should have lation growth rate due to a variety of mechanisms such
lower variance in population growth rates than larger as failure to find a mate, reduced foraging efficiency,
populations. In cases where both population size and and inability to defend against predators, which col-
variance in demographic rates are small, sequentially lectively are termed the Allee effect (Skagen and Yackel
settling sites from highest to lowest quality can result in Adams 2011). If populations are in a density-dependent
rapid population growth (McPeek et al. 2001). Species growth phrase, density and per-capita fecundity may
that conform to site- dependent regulation therefore not be correlated if settlement patterns among patches
should be (1) less vulnerable to stochasticity associated should equilibrate fitness via ideal free or ideal free
with small populations, (2) more likely to recover from despotic distribution. If populations are regulated by
small population size, and (3) less impacted by habitats events occurring in different seasons and / or habitats
where average suitability is relatively lower than spe- for species with high migratory connectivity, then the
cies that randomly settle sites. association between single-season (or stage) habitat
and population dynamics may be weak due to carry-
over effects, whereby demographic measures at one
Demographic Consequences
location reflect the quality of habitats used in previous
of Habitat Use Can Show High
seasons. For species subject to strong carryover effects,
Spatiotemporal Heterogeneity
demographic consequences of habitat use may not be
Though habitat can strongly govern population demog- evident until individuals transition across seasons or
raphy, a wide variety of other ecological factors can years. Carryover effects can mask or create patterns
create highly variable relationships between habitat in the apparent demographic consequences of habitat
and demographic parameters—even in the absence use and selection. For instance, if high-quality winter
of differences in the habitat itself. Some heterogene- habitat is limiting and induces carryover effects on fit-
ity may be temporary if there are time lags following ness, then there may be large demographic differences
environmental change, as can be the case with species among populations or individuals occupying similar
showing high levels of site fidelity (Davis and Stamps breeding habitats.
2004). However, heterogeneity is an inherent part of Population demography also is expected to reflect
many systems, and this means that focusing squarely the manner in which animals interact with the abiotic
on habitat as the key driver will result in a limited un- and biotic components of habitat, which can change
derstanding of population ecology (Morrison 2001). with countless ecological factors, including life stage,
Population size and regulatory mechanisms can ob- condition, density, food availability, and presence of
scure the relationship between habitat quality and popu- other species. In other words, even if the habitat re-
lation demography when density- dependent mecha- mains the same, the demographic consequences of that
nisms operate. If populations in high-quality habitats habitat might vary with changes in the way a species
approach carrying capacity, then population growth uses a habitat or interacts with other species (i.e., as
rates should decline. For example, long-term work with its niche changes; Morrison 2001, 2012). The possi-
black-throated blue warblers at Hubbard Brook LTER bility that niche-based changes produce variation in
shows that both fecundity and the condition of young the habitat–demography link are especially likely for
were negatively correlated with adult density within ecotypes, or a distinct geographic variety, population,
a season (Sillett and Holmes 2005), probably due to a or race within a species that is adapted to local envi-
combination of crowding (Sillett et al. 2004) and site ronmental conditions. Optimal habitat conditions are
dependence related to preemptive occupancy of ter- likely to differ among ecotypes, which are poorly de-
ritories that vary in quality (Rodenhouse et al. 1997; fined and understood for most species. Recent work
McPeek et al. 2001; Rodenhouse et al. 2003). What suggests that ecotypes can form more rapidly than pre-
28 foundation

viously thought in cases where there is strong habitat landscape (Dunning et al. 1995; Belisle et al. 2001),
matching. Habitat matching is a behavioral mechanism serve as a source of species and individuals invading
whereby individuals select habitats that best match habitat fragments, especially exotic species (Pysek et al.
their phenotype and ability to use that habitat (Edelaar 2002), and even determine the extent of edge, area,
et al. 2008). Because habitat matching can result in and isolation effects on wildlife (Donovan et al. 1997;
spatial aggregation of individuals with similar ecologi- Hartley and Hunter 1998). Differing landscape con-
cal traits, rapid adaptation can follow. texts, or characteristics of the landscape surrounding
When a species’ niche is considered in terms of the habitat, may explain why similar silvicultural treat-
interspecific interactions, it becomes apparent how ments might induce strong edge-related nest predation
differences in community organization and species in- in one landscape and no detectable effect in another
teractions also can result in spatiotemporal variation (Hartley and Hunter 1998). Area requirements of the
in demography in otherwise similar habitats. Not sur- same species, too, may vary widely with the amount of
prisingly, nest predation rate varies with abundances fragmentation in the landscape or region (Rosenberg
of many rodent nest predators, such as red squirrels et al. 1999) or the amount of residential development
(Tamiasciurus hudsonicus) and Eastern chipmunks surrounding the forest (Friesen et al. 1995; Rodewald
(Tamias striatus) (Sloan et al. 1998; Clotfelter et al. and Bakermans 2006). In this way, landscape context is
2007; Schmidt et al. 2010), which may, in turn, be a an important source of spatiotemporal heterogeneity in
function of mast or seed production by dominant tree demographic responses to habitat attributes.
species. So, too, can the foraging behavior of preda-
tors change with food abundance (e.g., functional re-
Conclusions
sponses like prey-switching). Even the fear of preda-
tion can elicit demographic responses, as illustrated While biologists have long known that the quantity,
by the reduced reproductive output of song sparrows quality, and distribution of habitat directly or indi-
(Melospiza melodia) subjected to experimental preda- rectly affect wildlife populations, recent developments
tor calls (Zanette et al. 2011). In some cases, demo- have improved our understanding of the relationship
graphic outcomes may reflect the entire network of between habitat quality and population demography.
species interactions. Rodewald et al. (2014) showed First, habitat-population relationships need to be more
that the structure of multispecies networks of interac- explicit about the use and relevance of individual- and
tions between birds and nest plants was more closely population-level measures of habitat quality as they
associated with avian nesting success than direct mea- relate to population dynamics and management ob-
sures of habitat or landscape. Recent work suggests that jectives. Second, research and management strategies
species within communities (i.e., species assemblages) must address habitat needs and demographic connec-
do not respond similarly to human-induced rapid en- tivity across the full life cycle and be especially atten-
vironmental change. Some species are more likely to tive to carryover effects that may create or obscure
retain ancestral ecological attributes and preferences habitat-fitness relationships. Third, when interpreting
(i.e., niche conservatism), and this can affect the rate distributional patterns and habitat-demography links,
at which new communities form (Wiens and Graham we need to better recognize how behavioral processes
2005). Because many species respond individualisti- guiding habitat selection influence population dynam-
cally, predicting future range distributions and popu- ics. Fourth, only by describing and investigating the
lation sizes is particularly challenging, given that there causes and demographic consequences of spatiotem-
are likely to be many no-analog communities. poral heterogeneity in habitat use will we be able to
Demographic responses to habitat also can be dra- implement effective management across the range of a
matically affected by landscape context, or the configu- species. By moving beyond the traditional approaches
ration and composition of the landscape. In particular, to characterizing wildlife-habitat relationships in these
the landscape matrix surrounding habitat patches can ways, wildlife science and management will improve
provide alternative habitat (Norton et al. 2000), affect its capacity to link habitat parameters to population
movements of individuals and dispersal through the dynamics and viability.
 demographic consequences of habitat 29

ACKNOWLEDGMENTS Resources before Spawning Influence Gonadal Investment

I am grateful for the helpful reviews provided by Brett of Female, But Not Male, White Crappie.” Journal of Fish
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 4
Beatrice Van Horne and
Managing Habitats in a
Changing World
John A. Wiens

H abitats are the foundation of natural-resource

management and conservation. Without habitats
of the right sorts, the right sizes, the right variety, and
have been written about habitat and its many defini-
tions and permutations (see, e.g., Cody 1985; Morrison
and Hall 2002; Guthery and Strickland, this volume).
in the right places, natural systems cannot exist. Rather, we want to explore how simple conceptualiza-
But habitats the world over are in peril. The ero- tions of habitat can be compromised by the realities of
sive effects of human actions on natural landscapes nature. We conclude on a more hopeful note by consid-
concerned people decades ago (e.g., Leopold 1949; ering how the effectiveness of habitat-based manage-
Osborn 1953; Thomas 1956). These effects accelerated ment and conservation can be strengthened as the pace
as burgeoning human populations converted vast areas of landscape change quickens. First, however, we con-
of native terrestrial habitats into agricultural monocul- sider how the aims of conservation or natural-resource
tures or urban sprawl and transformed natural aquatic management may influence how habitat is perceived.
habitats into plumbing systems. Nearly all of the North
American tallgrass prairie has been converted to farm-
Approaches to Managing Habitat
land, and over 90% of the native vegetation in the
wheatbelt of Western Australia has been replaced by Habitat is generally taken as shorthand for the local
grain production, with the few small remnants widely environmental conditions in which a species of inter-
scattered (Hobbs et al. 1993). The Sacramento–San est lives. Understanding the relationships between
Joaquin Delta in California, which historically was a organisms and habitats has long been an objective of
vast area of tidal wetlands laced by over sixteen hun- scientific research, the foundation of conservation and
dred km of tidal channels, has been engineered into resource management, and the entry point for public
a labyrinth of levees; less than 3% of the wetlands appreciation of nature. It is by closely investigating the
remain (Whipple et al. 2012). Habitat loss and the factors that influence survival and reproductive success
fragmentation of the remaining habitat have been la- that we gain a window into the processes that influence
beled the greatest threats to biodiversity (Wilcove et al. the value of a habitat in successfully supporting a spe-
1998) and are the major causes of the declines of many cies (Van Horne 1986).
imperiled species (Scott et al. 2010; Wiens and Gar- Employing science to understand the processes
dali 2013). that directly influence species of concern, however,
The magnitude and extent of native habitat loss requires focus. If we fully understand these processes,
make it increasingly important to think about how best we should be able to predict how differences or changes
to manage and conserve what remains. It is not our in- in habitat will affect these species. Using this approach,
tent here to review the panoply of papers and books that the objectives are to learn more about a species and
 managing habitats in a changing world 35

to understand which factors influence its selection Despite this interplay, however, conservation and
of habitats, how differences in habitats affect mate management frequently emphasize a focal species or
choice or individual fitness, how the genetic structur- a community / ecosystem approach, rather than both.
ing of populations varies with habitat conditions, and There is an inherent tension between the approaches.
so on. This approach is often reductionist, emphasiz- There is also tension between the urge to simplify
ing individuals, single species, or subdivisions within and the need to acknowledge the complexity of nature.
species, about which more detailed knowledge on the Every situation is different in its details, but manage-
subject of habitats is always sought. More knowledge, ment or conservation that attempts to incorporate all
in turn, generates more questions, which fuels further the details will inevitably be so situation-specific that
investigations. The goal is to minimize uncertainty. In it has no general applicability. Some simplification is
the conservation-management arena, the emphasis is necessary. But simplification carried too far can result
often on species that are protected by laws or regula- in generalities that are appealing but lack the reality
tions (e.g., the Endangered Species Act, the Marine to inform on-the-ground management. The challenge
Mammal Protection Act, the Migratory Bird Treaty) is to find a balance. To do this, one must consider
or are economically or recreationally important (e.g., several factors that, unacknowledged, can confound
salmon, deer). habitat-based research and management.
In recent years, attention has been shifting to fo-
cus on multiple species, communities, ecosystems,
Some Realities
and ecological processes. As much as we need more
information on how habitats influence species viabil- Whether the target of conservation or management
ity, to extend a species-by-species approach beyond is a particular species, a multispecies community, or
a few focal taxa is unrealistically costly in time and a multitrophic ecosystem, habitat is often regarded as
money. More to the point, by emphasizing single spe- a fixed property of the target. Hence, the association
cies, important trophic dynamics and ecosystem pro- with habitat is considered to be sufficiently tight that
cesses such as water balances or nutrient flows will the species, community, and so forth can be charac-
be ignored. Efforts to conserve a species, even if based terized by its habitat. This is the basis for managing
on detailed knowledge of its ecology and life history, or conserving habitat for particular targets: protect or
may fail if the ecological processes that support its restore the right habitat conditions and the desired tar-
habitat are allowed to degrade. Broadening the per- gets will thrive. Simplification is achieved by assuming
spective, however, inevitably means that knowledge of constancy.
the multiple species is less detailed—one can’t know In truth, what we most often manage in the real
everything about everything. Consequently, the goal world is a complex of areas that are being disturbed
frequently is to know enough to reduce uncertainty to naturally or by humans at some frequency and spatial
an acceptable level for implementing management ac- extent while simultaneously changing as a result of
tions; more detailed knowledge may be unnecessary or growth, aging, and interrelationships of the vegetation
difficult to justify, given the additional cost. or the actions of the organisms living there (e.g., geese
Of course, there is a middle ground where the grazing in the tundra, predators affecting herbivore
species- centered and systems-centered approaches populations). Given such dynamics, it is unlikely that
meet and blend. Because conservation and manage- management will be able to maintain the full suite of
ment are often directed toward particular species of habitat conditions that would be optimal for preserv-
concern, the information gathered in single-species ing biodiversity writ large. Management is further
studies can be critically important in framing actions. challenged because conditions in a focal landscape
And because management and conservation frequently are influenced by factors that may originate outside its
consider large, complex systems, they can generate boundaries—fire, acid rain, dust, water flow, invasive
and help to frame the questions (and often provide the species, and the like—all of which are in a continuing
funding) for the more detailed basic investigations. state of flux. And, as if this weren’t enough, managing
36 foundation

habitats is also confounded by variations within species mised by inappropriate study designs. By conducting
and how they respond to habitat, variations in time, studies in an area arbitrarily selected because it is con-
and variations in space. All of these factors thwart at- venient, measuring habitat parameters because they
tempts at simplification. But how important are they? have been used in previous studies, bludgeoning the
data with statistics, and then generalizing the results
to a species over much or all of its distribution, the re-
Variation within Species and Their Responses
sults and conclusions may tell us little about the factors
to Habitat
that really affect a species’ viability. Morrison suggests
In part because of laws, regulations, and the nature of that we would learn more that is important by identi-
legal challenges, investigations are often focused on one fying biologically defined subsets of a species for study
or several species of concern in areas that are directly (he suggests using ecotypes that are associated with a
under management influence or control. The empha- specific type of habitat), conducting studies in areas
sis on species is understandable. Species are (usually) that encompass key population processes, describing
well-defined biological entities. Most vertebrate and habitats using parameters that are biologically relevant
plant species, and many invertebrate species, are eas- to the target organisms, and bounding the scope of in-
ily identified. They are useful units for management ference to what was actually studied. In other words,
and conservation, and they resonate with the public. we will learn more about habitat by incorporating ad-
The value of charismatic species such as African ele- ditional information about the biology of species into
phants (Loxodonta africana), giant pandas (Ailuropoda our study designs so that we measure the right variables
melanoleuca), polar bears (Ursus maritimus), or Cali- at the right places.
fornia condors (Gymnogyps californianus) in mobilizing Even as the term species encompasses variations in
public support for conservation, or the importance of life-history attributes among individuals, populations,
desired tree or fish species such as Douglas-fir (Pseu- and ecotypes, necessitating a more nuanced approach,
dotsuga menziesii) or various salmonids (Oncorhynchus habitat is also subject to multiple sources of variation
spp.) as targets for management, cannot be ignored. A and interpretational challenges that make its defini-
good deal of scientific research carried out by gradu- tion elusive in any but general terms. Habitat is gener-
ate students in biology or wildlife departments or by ally thought of as an attribute of species. Field guides
single investigators rather than teams of scientists is usually include a section on “habitat” in their species’
directed at uncovering important details of the genet- accounts—things like “moist coniferous forests; adja-
ics, physiology, behavior, ecology, evolution, or simply cent oaks, shade trees” or “moist woods of oaks, pines,
the natural history of particular species. This knowl- and Douglas fir” (descriptions for chestnut-backed
edge is essential to conduct species-level management chickadee, Poecile rufescens, in Peterson 1990 and Sib-
or conservation. ley 2003, respectively). Such descriptions usually fo-
Studies of individual species almost always reveal cus on major vegetation types. Habitat designations for
fascinating and often idiosyncratic details about their species of conservation concern, such as those listed
relationships with habitat. Aided by technological de- under federal or state endangered species acts, likewise
velopments (e.g., satellite-linked GPS tags, light-level usually begin (and often end) with documentation of
geolocators; Recio et al. 2011; Lisovski et al. 2012; the vegetation types and “critical habitats” needed to
McKinnon et al. 2013) and advances in statistical ap- maintain the species. “Old-growth coniferous forest”
proaches (e.g., AIC, occupancy models; Burnham and for northern spotted owls (Strix occidentalis carunia)
Anderson 2002; MacKenzie et al. 2006), knowledge is an example. Thus, even though habitat is formally
about the habitat occupancy and use by targeted spe- considered to include all important life requisites for a
cies has become increasingly detailed. Morrison (2012) species, vegetation types are often used as surrogates.
has argued, however, that this greater detail may not be This simplification is understandable since general (or
telling us what we really need to know about species even specific) vegetation types are easily recognized
and their habitats because the information is compro- by many people (there is little need to explain what a
 managing habitats in a changing world 37

“moist coniferous forest” is to someone looking for a to other times. The effectiveness of conservation and
place to find chestnut-backed chickadees). Vegetation management actions will therefore be time-insensitive.
is also more easily managed to create the habitat de- So much of what we know about species and habitats
sired for a species than are other life requisites such as is based on short-term studies that do not record tem-
food, protection from predators, appropriate thermal poral variations, which reinforces this assumption. All
regimes, or suitable breeding sites. By using vegetation environments vary in time, however, and these varia-
as a simplifying proxy for habitat, however, the link- tions can confound attempts to define the habitat con-
ages to habitat processes that actually affect a species ditions favoring one species or another. For example,
can be lost. the expansion of barred owls (Strix varia) into forests
It is frequently assumed that places where a spe- of the Pacific Northwest in recent years has altered
cies occurs contain good habitat and places where it the demography and habitat distribution of northern
is absent do not. This assumption is the basis for us- spotted owls where the species co-occur (J. D. Wiens
ing correlations between environmental features and 2012). Habitat quality may also change between years,
the presence or abundance of a species to describe its depending on weather. Thus, a drought in the shrub-
habitat. Species-distribution models, which are used to steppe of western Idaho caused a higher proportion of
predict current or future distributions based on such ground squirrels (Urocitellus townsendii) to survive in
correlations, assume that where a species occurs de- patches of native shrub habitat than in parched native
fines suitable habitat and that the available habitat is bunchgrass habitat, where they had fared better than
fully occupied (“saturated”) (Wiens et al. 2009). It has in shrub habitat in years of average rainfall (Van Horne
long been known, however, that the abundance or den- et al. 1997).
sity of a species in an area can influence the range of Temporal dynamics over shorter periods can also
habitats occupied; Fretwell and Lucas (1969) modeled influence the assessment of a species’ habitat. The
this in their “ideal free distribution” (see Rodewald, habitat associations of many species (especially migra-
this volume). If habitat occupancy varies with density, tory ones) change seasonally (Rodewald, this volume),
then recording the features of an area that is occupied or even within a season. In a Wisconsin grassland, for
at a particular time to characterize a species’ habitat example, the features of habitats differed for the initial
will be sensitive to its density at that time. Moreover, territories established by grasshopper sparrows (Ammo-
whatever the density of a species in an area at a given dramus savannarum) and savannah sparrows (Passercu-
time, it may not say much about the relationships with lus sandwichensis) when birds returned in the spring.
habitat that influence individual fitness (“habitat qual- As the season progressed, however, more individuals
ity”). This is the basis for Van Horne’s argument (1983) established territories, filling the study area—habitat
that density is a misleading indicator of habitat quality characteristics of the two species converged (Wiens
or of the concept of “ecological traps” (individuals are 1973). In all of these examples, the determination of
attracted to and occupy unsuitable habitats; Donovan a species’ habitat would differ depending on when it
and Thompson 2001). In populations having a source- was assessed.
sink structure (Liu et al. 2011), as many likely do, densi- Time lags, legacy effects, or carryover effects (Rode-
ties may sometimes be greater in the less suitable sink wald, this volume) may also erode the match between
habitats, with population numbers maintained only habitat occupancy and habitat quality. Individuals that
by continuing immigration from the better source exhibit fidelity to previous breeding locations may
habitats. continue to occupy those areas even after the original
habitat has been drastically altered (Wiens and Roten-
berry 1985). Unless appropriate measures of fitness
Variation in Time
(e.g., survival, reproduction) are also recorded, assess-
To simplify things, considerations of habitat in con- ments of habitat made only after the conditions have
servation and management often assume that habi- changed will yield incorrect documentations of habitat.
tat relationships documented at one time will apply This is especially likely to occur for long-lived organ-
38 foundation

isms that are sedentary or philopatric, such as trees or and scale effects but to spatial context as well. Another
many birds. simplifying assumption is that all important relation-
ships between individuals and their habitats are con-
tained within the defined study area. With the excep-
Variation in Space
tion of oceanic islands, study areas are not isolated,
Just as all times are not the same, all places are not the stand-alone places but exist within a broader landscape
same. While few would suggest that studies of habi- mosaic. The boundaries of a study area are permeable
tat relations in a woodland could be extrapolated to to influences from beyond the boundaries (Hansen and
a grassland, it is easier to assume that patterns docu- di Castri 1992), especially if the study area is small.
mented in one oak woodland in one area might apply The features of the surrounding landscape can affect
to a similar oak woodland in another area. If habitat is what occurs within the study area, and thus the infer-
defined only by the dominant vegetation (e.g., “moist ences that are made about habitat. Three decades ago,
coniferous forest” for chestnut-backed chickadees), Janzen (1983) called attention to how the surroundings
this simplification is easier still. of a protected area could alter the survival and repro-
As Morrison (2012) observed, documentations of duction of individuals within the area due to the ac-
species’ habitats are usually made by measuring param- tions of predators or competitors living in the adjacent
eters within an arbitrarily defined (and usually small) landscape. Linkages with the landscape are part of the
study area. Extrapolating results to a larger area as- habitat within an area.
sumes that the habitat preferences and relationships of Much of what we have said about the problems
a species remain constant over a range of spatial scales. with study areas applies also to management that is de-
This is unlikely to be the case. Studies of sage sparrows signed to conserve habitat in a reserve area or system
(Artemisiospiza belli), for example, illustrate how the as- of reserves. Uncertainties about the appropriate units
sociations between abundance and vegetation features to represent a species, flexibility in habitat selection
change with changes in the scale of analysis (Wiens within one or several species, variations in density and
et al. 1987; Wiens 1989). At a broad, biogeographic occupancy of habitat that may be unrelated to habitat
scale, for example, densities were correlated with high suitability, time lags and temporal dynamics, and the
shrub coverage and low grass coverage, while at a re- effects of spatial variation, scale, and landscape con-
gional scale densities were greatest in areas with more text confound habitat management. It is important to
grass and fewer shrubs. These changes occurred over consider the larger landscape, including multiple own-
a broad range of scales encompassing different ecore- erships, in management for conservation, even if re-
gions; within a narrower range (e.g., a few hectares to search is based on simplified portions of the landscape.
tens of square kilometers within a region), habitat rela- Habitat management for conservation, however, is
tionships might be less sensitive to scaling differences. often restricted to areas designated for biodiversity pro-
The limits to how far one can extrapolate among tection, especially of imperiled species that have small
areas or scales depend on one’s objectives. The more populations and severely restricted distributions. In-
general the objectives, the less important the differ- side such reserves or protected areas, habitat manage-
ences among areas or across scales may be. But the in- ment for species or communities may be the primary
ferences, and the guidance for management and con- goal, while outside, other resource-management goals
servation, also become more tenuous. At some point, or human activities (e.g., forestry, grazing, recreation)
differences in key factors are missed, compromising or usually have primacy. Because of natural succession,
invalidating extrapolations. Identifying the bounds of climate change, or processes or threats moving across
extrapolation requires that we know enough about how the reserve boundary, conditions within the reserve
key habitat factors affect fitness to be confident that will likely change in ways that cannot be managed cost-
they remain relatively unchanged in their effects from effectively. A small reserve established for a focal spe-
place to place or among scales. cies may change in ways that no longer benefit the spe-
Documentations of the habitat associations of spe- cies; the invasion of spotted owl habitat by barred owls
cies are sensitive not only to place-to-place variations or changes in the fire regimes that maintain habitat for
 managing habitats in a changing world 39

Kirtland’s warbler (Setophaga kirtlandii) (Bocetti et al. economic forces, and changing societal demands for
2012) are examples. Conversely, the larger landscape resources and commodities, will alter landscapes over
may also contribute to habitat conditions and connec- multiple scales.
tivity that benefit species within the reserve. In some cases, ecological responses to these changes
may be gradual, as species expand, contract, or shift
distributions. Species-distribution modeling for a vari-
The Challenge of a Changing World
ety of taxa (e.g., Iverson et al. 2008; Lawler et al. 2009;
Habitat management generally assumes that, absent Stralberg et al. 2009) has shown how extensive these
major disturbances, the habitat within an area will per- distributional changes may be. The effects will differ
sist more or less unchanged once the area is protected among species, disrupting the composition of local
or managed or the habitat restored. Although succes- communities and the trophic webs of ecosystems. In
sional processes may move the habitat away from (or other cases, the changes may be sudden, as systems are
toward) a desired state, these changes are generally pushed beyond thresholds of resilience or tolerance to
thought to be gradual and at least to some degree pre- drought, temperature, disturbance, or other factors.
dictable. Spatial variation may create heterogeneity in Such changes have already occurred, for example, in
successional states at multiple scales, but this is often the shift from shrub dominance to grass dominance
viewed as variation about a long-term equilibrium, as in shrubsteppe ecosystems (West and Hassan 1985)
envisioned in shifting-mosaic steady-state concepts or from grassland to mesquite (Prosopis glandulosa)
of succession (Borman and Likens 1979; Turner et al. shrubland in the Chihuahuan Desert (Buffington and
1993). Of course, ecologists and managers recognize Herbel 1965). In both situations, grazing by domestic
that environments vary over time in ways unrelated to livestock, the invasion of exotic vegetation, and / or
ecological succession; neither still believes in a strict increased fire frequency and extent have shifted the
equilibrium view of nature. The expectation that such ecosystems into an alternative state, as envisioned
variation occurs around a stable, long-term mean, how- by state-and-transition models (Bestelmeyer 2006;
ever, is encapsulated in the concepts of stationarity (in Bestelmeyer et al. 2011). The habitat changes are po-
aquatic environments) or historical range of variation tentially irreversible.
(in terrestrial systems). Both concepts have been dis- Whether the changes are gradual or sudden, how-
credited (Milly et al. 2008; Wiens et al. 2012); even the ever, they will produce a continuing disassembly and
average conditions are not stable and unvarying over reassembly of biological assemblages, resulting in novel
any time scale relevant to conservation or manage- combinations of species and “no-analog” ecosystems
ment. Consequently, many of the factors determining that neither we nor the species have seen before (Wil-
“habitat” vary over multiple scales in time and space, liams and Jackson 2007; Stralberg et al. 2009; Hobbs
burdening habitat management with cascading uncer- et al. 2013). Over the ecological time scales that are
tainties. Unpredictable extreme events such as hurri- relevant to conservation and management, bedrock
canes and floods (Dale et al. 1998; CCSP 2008) only geology may not change (unless there are major tec-
add to the uncertainty. tonic events), but everything else that we associate
This is the current situation. While the future is by with habitat—vegetation composition and structure;
definition uncertain, the changes now underway are the composition of prey, predator, and competitor as-
sure to create additional uncertainty in habitat-based semblages; food-web dynamics; and even the soil or
management and conservation by altering not only the hydrology—will change, creating new mixtures and
environmental context of habitats but the very nature new habitats. The new habitats, in turn, will provide
of the habitats that people wish to manage, whatever opportunities for the establishment of species new to
the targets. Changes in global climate are projected to the area, some of which may further disrupt the eco-
have profound effects on regional and local precipita- logical systems while diminishing prospects for other,
tion, temperature regimes, and the frequency and mag- desirable species. The novel habitats and ecosystems of
nitude of extreme events. Changes in land use, driven the future will require novel approaches to conserva-
by the combination of climate change, local and global tion and management.
40 foundation

Does the uncertainty about what habitats will exist the broader landscape. This work is messy, involving
in the future mean that the age of habitat-based man- many societal and legal agendas and goals. Our cur-
agement and conservation is past? Not necessarily. rent institutions (state and federal land management
The habitats that exist in the future will still be habi- agencies, conservation organizations) are not generally
tats—places in which organisms can find the essential organized to support and facilitate such approaches.
resources to survive and reproduce, populations can The ideal solution for protecting habitat would en-
persist, and ecological processes can continue to oper- compass large landscapes with multiple owners and
ate. They will be habitats for something; we just may with multiple management goals. It would affirm the
not know what. importance of habitat and biodiversity conservation
goals and support the use of science and modeling to
establish and project habitat conditions over time and
A Way Forward for Habitat Research
space, while applying decision support approaches
and Management
(Marcot 2006; Heaton et al. 2008) to manage compet-
In this chapter, we have discussed some of the diffi- ing goals for the landscape. Tools in remote sensing,
culties in simplifying habitat research and manage- geographic information systems, and statistics now
ment to assure that investments are justifiable while enable us to incorporate more of the complexity of
still providing what is needed to maintain habitat and nature into management and conservation practices.
prevent further loss. The array of confounding factors Cooperative approaches to habitat management (e.g.,
we have discussed is daunting. Although not all need Collaborative Forest Landscape Restoration Program,
be included in all research designs or management Title IV of the Omnibus Public Land Management Act
plans, their potential impacts on habitat assessments of 2009; Bagstad et al. 2012) are also increasing. The
should be acknowledged and given careful thought, in- best approach would be to combine the focused, set-
stead of falling prey to the desire for simplification. For aside reserve approach with a cooperative approach to
example, once one accepts that a landscape includes habitats over entire landscapes.
places that are changing at various rates in ways that To conclude, we suggest several elements of an ap-
alter habitat quality for species or communities, simple proach that may help to resolve the tension between
management rules that place bounds on economic and simplification and complexity in habitat research and
recreational activities, prescribe habitat restoration management. Although this is not a comprehensive
practices, or detail treatments for recognized threats list, we hope it will provide a useful starting point for
such as fire must be adapted to a variety of current identifying what we should be thinking about as we
and expected conditions. Effective management may work to understand and conserve the elusive entity we
require using complex models to project future habitat call “good habitat.”
conditions in landscapes at multiple scales, based on
the gradual and / or sudden changes that are expected. 1. Acknowledge that creating a mix of dominant
Such models are always imperfect, but they may pro- vegetation types across a large landscape does not
vide useful estimates of what is likely to take place in by itself ensure a diversity of animals. Additional
the landscape. information about ecological processes and
Resource managers and conservationists are faced species requirements is needed to evaluate the
with an increasingly complex and demanding task. potential of habitat to support current and future
As managers pursue their larger mission, which may biodiversity.
involve managing lands for national defense, mining, 2. Maintain multiple trophic levels, including
energy development, recreation, hunting and fishing, desired vegetation, herbivores, and predators. As
or other activities, they must also manage habitat to Aldo Leopold observed long ago (1949), a func-
avoid harming species of concern. In addition, mana- tioning ecosystem requires all the parts.
gers must work with diverse landowners with multiple 3. Implement management over broad areas and
interests, which requires using an array of manage- multiple ownerships to consider processes occur-
ment practices that address habitat conditions across ring at broader scales while also addressing the
 managing habitats in a changing world 41

objectives of the multiple parties involved. Fuel ing few years, while you ensure that the man-
buildup in western and Rocky Mountain forests, agement design includes appropriately targeted
declining water tables in the desert southwest scientific monitoring to allow you to learn from
or the upper Columbia River basin, or the lack management activities and adapt to this new
of early successional habitats for grassland birds knowledge.
in the northeastern United States are examples.
Search for the shared objectives! Including these considerations in cost-effective re-
4. Consider how plant and animal species move search and management won’t be easy, but it will go a
across a landscape that is changing and create long way toward ensuring that “habitats” will persist
movement pathways. Habitat linkages need to into an uncertain future.
be designed to facilitate seasonal and dispersive
movements of a wide variety of species, while LITERATURE CITED
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PA R T I I • HABITATS IN PERIL
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 5 Habitat Loss and
Degradation
Clinton D. Francis
Understanding Anthropogenic Stressors
and Their Impacts on Individuals, Populations,
and Communities

H uman activities are drastically changing Earth’s

landscapes, and the magnitude of human impacts
on geophysical and biological processes fuels debate
through mini-reviews of three aspects of human distur-
bance that deserve additional attention: anthropogenic
noise, artificial lighting, and recreational activities. I
on whether we are entering a new epoch coined the focus on these aspects because they are ubiquitous,
Anthropocene (Crutzen 2002; Ellis 2011). Acknowl- growing faster than the human population, and be-
edging this fundamental change, Ellis and coworkers cause we still know relatively little about their impacts.
retooled the standard biome classification system and Throughout the chapter, I also point out how we could
mapped the anthropogenic biomes of the world (Ellis do a better job of collecting meaningful data that can
and Ramankutty 2008; Ellis et al. 2010). The results inform effective conservation policy and highlight the
are stark; less than a quarter of ice-free land remains benefits of approaches that work across several levels
wild. That few landscapes across the globe remain un- of biological organization by examining how individual
touched by humanity should give wildlife profession- responses scale to population and community-level
als and conservation biologists pause. Should conser- patterns. This includes measuring wildlife responses
vation efforts focus primarily on preserving “natural” to habitat degradation differently, using the organ-
habitats and should research efforts concentrate on ism’s sensory systems to guide our choice of relevant
how wildlife persist in these habitat islands? With so and functional environmental variables to measure,
little natural habitat left, these areas will undoubtedly and studying communities rather than single species.
prove insufficient for the preservation of most species. I will start by describing how wildlife habitat loss has
Instead, biologists are increasingly tasked with under- typically been viewed, then provide details on how this
standing how wildlife and their supporting ecological view must be expanded in several ways.
systems function in the context of sustained direct and
indirect interactions with humans.
When Is Habitat Lost?
Throughout this book, readers are exposed to a va-
riety of threats to wildlife. In this chapter, I will briefly Habitat loss occurs when an area is converted from one
describe more traditional views of “habitat loss and state to another. Traditionally, we have almost always
degradation” and place these concepts into the context viewed habitat loss in terms of changes in land cover
that most wildlife already live in areas affected by hu- (i.e., vegetation structure or type). Take, for example,
man activities. In many ways, this is not a description the clear-cut of a once-continuous forest to make way
of particular habitats in peril but an overview of some of for agriculture, grazing, or a housing development. Im-
the ways humans make numerous landscapes perilous portantly, habitat loss is species specific: an area that
for wildlife. As such, I aim to explain the way we should becomes uninhabitable for one species might become
think about sources of habitat loss and degradation a new inhabitable area for another.
48 habitats in peril

Figure 5.1 (A) A once-contiguous area (grey) undergoes both habitat loss and fragmentation, resulting in a landscape with
four remaining patches (B). The use of corridors to maintain connectivity among habitat fragments (C ) could be one
strategy to maintain wildlife populations but may not work for all species (see main text).

cies: A & B. Individuals of Species A require a larger

Habitat Loss versus Habitat
territory than individuals of Species B. Species A can
Fragmentation
also disperse or move farther across unfavorable en-
Habitat loss and habitat fragmentation are terms often vironments than Species B. When a once-contiguous
used to describe land cover change. Although these area (fig. 5.1A) undergoes both habitat loss and habitat
processes are different, they are not always clearly fragmentation (fig. 5.1B), the abundance of both spe-
distinguished from each other in practice and have of- cies declines. Habitat loss might be the mechanism
ten simply been called “habitat fragmentation” in the explaining a reduction in the abundance of Species A,
literature (reviewed in Fahrig 2003; Ewers and Did- simply because the proportion of original habitat is
ham 2006; Collinge 2009). Habitat loss refers to the reduced. In contrast, this smaller amount of habitat
reduction in the overall area of a particular land cover may be sufficient to maintain the abundance of Spe-
type, and fragmentation entails the spatial arrangement cies B, but the isolation of remnant patches precludes
and shape of remaining habitat patches. Why is it that individuals from moving among them, also leading to
habitat loss and fragmentation have proven difficult to a decline in population size. This is but one example
disentangle? The short answer is that the process of of how habitat loss and fragmentation differ, and these
habitat fragmentation is always accompanied by habi- details matter for management. For example, manage-
tat loss, but habitat loss can occur without fragmenta- ment efforts that create travel corridors linking sepa-
tion (Fahrig 2003; Collinge 2009). That is, a habitat rate habitat fragments might help Species B but would
patch could be reduced in size without dividing it into be ineffective for Species A (fig. 5.1C).
separate pieces (i.e., fragmented), but it cannot be di- Thus, both habitat loss and fragmentation can func-
vided without losing some of the original area. These tion in different ways to harm wildlife populations, but
distinctions may seem trivial at first, but because wild- which mechanism is more common? Fahrig (2003)
life biologists are interested in the viability of popula- summarized the difference nicely. In general, habitat
tions and integrity of ecological communities that sup- loss has strong, negative effects on biodiversity, whereas
port them, understanding the mechanisms responsible the influence of habitat fragmentation alone appears
for observed shifts within a biological community in to be much weaker and may range from negative to
response to habitat change is imperative. Only by un- positive. Importantly, one must carefully consider that
derstanding why shifts occur might we be able to imple- the evidence for the influence of habitat fragmentation
ment strategies to halt or slow population declines. in the absence of habitat loss has come from relatively
Here is a hypothetical scenario that demonstrates few studies. More details on habitat fragmentation
why the distinction between the effects of habitat loss can be found in chapter 7 of this volume, where it is
and habitat fragmentation is important. Take two spe- discussed in detail (Smallwood, this volume), plus sev-
 habitat loss and degradation 49

eral other good sources (Forman 1995; Collinge and

Forman 1998; Fahrig 2003; Ewers and Didham 2006;
Collinge 2009).

Rethinking Measurements of
Environmental Change and Wildlife Figure 5.2 Habitat suitability should often be viewed
Responses: Wildlife’s Perspective along a continuum of quality rather than a paradigm that
categorizes areas as inhabitable versus uninhabitable.
The traditional view of habitat loss and fragmentation
Vertical lines represent the point at which a decrease in
that I outlined earlier neither reflects the range of envi-
habitat quality could result in site abandonment for two
ronments experienced by wildlife nor the complexities different species (a and b) and individuals of the same spe-
that wildlife researchers need to consider when study- cies (denoted with subscript numbers). These thresholds
ing wildlife responses to human-caused environmental will be context specific: they will vary among species,
change. Aspects of the human enterprise other than among individuals of the same species, but also according
land cover change can cause areas to become uninhab- to the precise combination of environmental conditions
itable, and many landscapes are impacted by human in the area.
activities in ways far more subtle than obvious cases
like the transformation of a forest stand to a parking reasons. The reverse is also true; occupancy of an area
lot. Take, for example, an increase in trail density for does not necessarily mean that an area is better habitat
recreation or shrub encroachment due to fire suppres- than areas where a species is not detected. Instead, the
sion or grazing. In these and other cases, it may not quality of the habitat could be manifested after settle-
be an issue of habitat simply being “habitable” versus ment. For example, individuals may settle in degraded
“uninhabitable.” Instead, habitat quality ranges along and high-quality areas at similar rates, but then experi-
a continuum from high quality to low quality. At some ence different rates of survival or reproductive success
point along this continuum, an area becomes “unin- (fig. 5.3). Worse still, individuals may actually prefer
habitable,” but this will depend not only on the species degraded or altered habitat where reproductive success
but also on the individual. Considerable research has is very low (sometimes referred to as attractive sinks or
shown that individuals of the same species can differ ecological traps) relative to high-quality habitat where
greatly in their responses to the same stimulus or envi- reproductive success is high (Robertson and Hutto
ronmental cue (fig. 5.2), and this variation is thought to 2007). These highly preferred but reproductively costly
contribute to the degree to which different individuals habitats can be the worst kind to preserve. For these
cope or fail to cope with environmental change (re- reasons, it is unfortunate that much research involv-
viewed in Sih et al. 2004; Blumstein and Fernández- ing habitat loss and degradation focuses primarily on
Juricic 2010). These individual responses have clear the very coarse measures of presence / absence or abun-
consequences for population persistence and commu- dance, especially because we know that abundance
nity patterns (reviewed in Francis and Barber 2013; does not necessarily reflect habitat quality in terms
Rodewald, this volume). Less well known, however, is of reproductive success (e.g., Van Horne 1983; John-
how nonrandom selection among individuals with dif- son and Temple 1986; Rodewald, this volume). These
fering responses to habitat degradation can influence measures do tell us that there is a pattern of concern,
the evolutionary trajectories of populations and either but they usually tell us little about why a population
increase or decrease a population’s viability. declines or thrives due to a particular type of environ-
It is also difficult to determine whether an area mental change.
is “bad habitat” based only on where individuals are Part of understanding why organisms respond as
encountered, because the context of the environment they do to habitat degradation will require rethink-
matters greatly. An area may be unoccupied simply ing about which environmental variables should be
because better habitat exists nearby, because animals measured in addition to quantifying distributional
cannot readily access the patch, or for a variety of other changes and variables reflecting reproductive success
50 habitats in peril

Figure 5.3 Declining habitat quality will not always cause reductions in abundance. For example, Species X (long dashed
line) could decline in occupancy with degradation by avoiding degraded areas when making settlement decisions, but
Species Y (short dashed line) and Z (solid line) may fail to respond to degradation while settling (A). In contrast, Species
Y might experience decreased survival with degradation, whereas Species Z does not (B). Still, increased degradation may
negatively affect reproductive success for Species Z (C). Importantly, over time these three different responses could all
lead to the same pattern of population declines, but the mechanistic reasons responsible for the declines are different
and matter for management.

and fitness. We typically measure habitat structure strength of conspecific (i.e., social) attraction for habi-
as it is perceived visually by humans, often failing to tat selection. Surprisingly, playbacks in areas with little
consider sensory systems used by other organisms that canopy cover, which was thought to be low-quality hab-
differ from our own (Van Dyck 2012). This bias is prob- itat, resulted high clustering of territories. Moreover,
lematic because the way we perceive the structure and measures of breeding and reproductive success among
composition of a particular land-cover type does not males was not related to canopy cover but instead
necessarily reflect an area’s functional significance for positively related to breeding density. This example
organisms under study. Instead, we must try to under- illustrates that the cues used by organisms to make
stand how organisms perceive their surroundings and settlement decisions can differ quite drastically from
frame research efforts with this knowledge to reveal what was assumed to be important. More importantly,
how organisms respond to functional components of however, this study reveals that a federally endangered
both natural and human-altered environments. So how species can successfully occupy a broader environmen-
might we make this shift? Because all organisms use tal range than previously thought and that playback ex-
sensory systems to obtain and respond to information periments could be used in management to reestablish
in their environment, adopting a research approach populations in areas where they no longer exist.
grounded in sensory ecology can help reveal how and A second example comes from birds’ abilities to use
why organisms respond to natural variation in environ- acoustic cues from natural enemies to make settlement
mental conditions and anthropogenic environmental choices. Playback of a common nest predator’s vocal-
changes. izations causes ground-nesting songbirds to avoid nest-
A clear example illustrating the benefit of research ing in areas near the playback source (Emmering and
emphasizing the way in which wildlife perceive their Schmidt 2011). Similarly, several songbirds appear to
environment comes from recent research on habitat respond in the same manner to playback of vocaliza-
selection in birds. For example, Farrell et al. (2012) tions from brown-headed cowbirds (Molothrus ater),
used experimental playback of golden- cheeked war- which commonly reduce the reproductive success of
bler (Setophaga chrysoparia) vocalizations throughout many passerines by parasitizing their nests (Forsman
an area that varied in habitat quality to determine the and Martin 2009).
 habitat loss and degradation 51

These examples illustrate how organisms use sen- as mortalities from collisions with vehicles or the dif-
sory systems to perceive their environment in ways ference in vegetation near roads relative to compari-
different from human visual perception and standard son sites away from roads. Recently, however, several
scientific habitat quantification. Additionally, they studies have dealt with the influence of confounding
emphasize the power of focusing research efforts on variables in their analyses or in their study designs and
variables that are closely tied to function. This focus shown that noise alone can explain declines in bird
on function is especially important for anthropogenic species abundances and species richness (Bayne et al.
stressors. 2008; Francis et al. 2009; Goodwin and Shriver 2011;
Blickley et al. 2012). Interestingly, there appears to be
considerable variation among species with some com-
Anthropogenic Noise, Light, and
pletely absent and others more common in noisy ar-
Recreation: Habitat Loss without Loss
eas (Francis et al. 2009; Francis et al. 2012). Although
of Habitat
we now know that noise does seem to change species
Wildlife now face countless threats from human activi- distributions, it is less clear why some species appear
ties that vary both in type and severity. Three threats to be more sensitive than others. For birds, the in-
are discussed here because many others are covered in ability to communicate in noisy areas appears to play
detail elsewhere in this book, and because these illus- a large role. Several studies have shown that species
trate the importance of research that strives to under- with low-frequency vocalizations tend to avoid noisy
stand how organisms obtain and respond to informa- areas more than species with higher frequency calls
tion in their environment. and songs (Rheindt 2003; Francis et al. 2011b; Good-
win and Shriver 2011). This pattern is directly related
to the functionality of vocalizations in areas degraded
Anthropogenic Noise
by noise. Anthropogenic noise is dominated by low-
Human-generated noise is now common not only in frequency energy, thus species with low-frequency
and around cities but in rural settings exposed to in- vocalizations experience more interference from the
dustrial agriculture, dendritic transportation networks, noise than do species with high-frequency vocaliza-
aircraft overflight, and resource extraction and recre- tions, making it that much more difficult for them to
ational activities (Barber et al. 2010). Collectively, these communicate (fig. 5.4). Because vocal frequency is neg-
sources of noise amount to unprecedented ecological atively related to body size, one generalization might
acoustic conditions for wildlife throughout the world be that larger species could be more sensitive to noise
and only recently has the effects of noise on wildlife than smaller species.
gained heightened attention by ecologists and conser- Why else might species avoid noisy areas? In addi-
vation biologists (Francis and Blickley 2012). Several tion to interfering with communication, continuous
recent reviews highlight some known effects of noise noise can also impair animals’ abilities to use acous-
for taxonomically diverse wildlife but also emphasize tic cues in a number of ways, such as interfering with
that many potential consequences of noise have not yet an organism’s ability to hunt (Schaub et al. 2008;
been investigated (Patricelli and Blickley 2006; Bar- Siemers and Schaub 2011), detect predators (Quinn
ber et al. 2010; Kight and Swaddle 2011; Ortega 2012; et al. 2006), and navigate throughout a landscape by
Francis and Barber 2013). In the following paragraphs, interfering with sounds used for orientation at a large
several ways noise affects wildlife and some key areas scale (Francis and Barber 2013). However, habitat
that warrant more attention are discussed. degradation due to noise can also affect wildlife after
A long history of road ecology studies suggests that they settle in noisy areas. For several bird species, in-
noise reduces habitat quality for wildlife, especially dividuals breeding in noisy areas have reduced repro-
birds (e.g., Reijnen et al. 1995; Kuitunen et al. 1998; ductive success compared to individuals in quiet areas
Forman et al. 2002). However, this evidence has been (Halfwerk et al. 2011; Kight et al. 2012; Schroeder et al.
viewed with skepticism because many other factors 2012), suggesting that noise avoidance cannot be the
might explain population declines near roads, such only metric of focus for biologists seeking to quantify
52 habitats in peril

Figure 5.4 Most anthropogenic noise has increasing energy at lower frequencies (grey background in spectrograms), such
that higher-frequency songs experience less acoustic interference from noise than lower-frequency songs. Additionally,
because body size is negatively correlated with vocal frequency, larger species tend to be more intolerant to noise than
are smaller species.

impacts from noise. Importantly, in all of these studies link between decreased reproductive output and the
the reduction in reproductive success occurred in the influence of noise on parent-offspring communication
absence of the influence of predation, which can com- has yet to be tested in this system. However, research
plicate determining the net effect of noise on reproduc- on tree swallows (Tachycineta bicolor) found that nest-
tive success (see the following discussion). lings exposed to noise beg less to the sounds made by
Halfwerk et al. (2011) proposed four nonmutually parents arriving at the nest than did nestlings not ex-
exclusive mechanisms that could explain reduced re- posed to noise (Leonard and Horn 2012). Apparently,
productive success for birds in noisy areas. First, noise by masking the sounds of arriving parents, noise causes
may interfere with a female’s ability to assess male qual- nestlings to miss feeding opportunities, which could
ity and cause her to invest less energy in egg production compromise growth and survival. Finally, a third study
and / or maternal care of chicks. Second, lower-quality provides limited support for a communication break-
individuals may settle in noisy areas, thus the effect of down between parents and offspring. Schroeder et al.
traffic noise on reproductive success would be governed (2012) studied the breeding biology of a house spar-
by a nonrandom distribution of the quality of individu- row (Passer domesticus) population exposed to chronic
als with respect to noise. Third, noise may increase noise from a large generator. Similar to the study with
physiological stress in individuals occupying noisy ar- bluebirds, they found nests in noise- exposed areas
eas, which can cause behavioral changes that reduce to fledge fewer young than nests in quiet areas. Ad-
fitness and can also have direct deleterious effects. And ditionally, chicks reared in noisy areas were smaller
finally, noise may compromise parent-offspring com- than those in quiet areas, and the authors linked this
munication such that nestling provisioning is altered trend to low provisioning rates by females. Although
in some way. other explanations that could explain the inadequate
Of these possibilities, there is some empirical sup- provisioning rates provided by females are plausible,
port for a breakdown in parent-offspring communi- such as elevated stress, the authors suggest that females
cation. First, Swaddle et al. (2012) found noise sig- provision less because they are unable to hear their
nificantly reduces the signal-to-noise ratio (i.e., how offspring’s begging calls. Collectively, these separate
“hearable” a sound is compared to background noise) studies suggest that impaired parent-offspring commu-
of chick begging calls at nearby perches used by nesting nication may be one mechanism that explains lower
eastern bluebirds (Sialia sialis). In this same study sys- reproductive success in noisy areas. Whether the other
tem, Kight et al. (2012) reported that bluebirds fledge mechanisms are at work remains to be seen.
fewer young with increases in noise levels. A direct Finally, noise can also interfere with breeding at-
 habitat loss and degradation 53

tempts well before nestlings hatch and begin to com- nual rhythms (Rich and Longcore 2006). Importantly,
municate with parents. Two studies have shown that the volume illustrates quite clearly that relatively few
territorial males inhabiting noisy areas are less success- studies have documented the existence and severity of
ful in attaining mates than are males in less noisy areas the many potential consequences of artificial lighting.
(Habib et al. 2007; Gross et al. 2010). This may be due Perhaps the most well-known effect of light pol-
to several different factors, including a nonrandom dis- lution involves sea turtles. Emerging hatchlings on
tribution in the quality of individuals with respect to beaches use light over the horizon of the ocean as
noise or due to females’ decreased ability to detect and one of several cues to find and move toward the surf
discriminate signals masked by noise to make mate- (Tuxbury and Salmon 2005). However, artificial night
choice decisions. Another possibility lies in how birds lighting emitted from structures along the beach can
that regularly inhabit noisy areas respond to noise. A cause hatchlings to travel away from the ocean where
growing number of birds have been shown to adap- survival is unlikely. In this case, light pollution can be
tively alter their songs and calls in response to noise viewed as an ecological trap (Schlaepfer et al. 2002)
(e.g., Gross et al. 2010; Francis et al. 2011a). Although because it mimics what had been a reliable environ-
this flexibility has been viewed as an important behav- mental cue used for navigating toward required habi-
ioral adaptation that permits species to overcome the tat. Fortunately, in-depth research on the sensory ecol-
acoustic conditions of cities and roadways that make ogy of turtle hatchlings has permitted managers with
communication difficult, the consequences of these targeted strategies to reduce inland movements. These
altered songs are unknown. Because small changes strategies include shading lights from beach areas, or
in song features can influence female preference for even using lights with wavelengths that do not attract
a song or even species recognition (Nelson and Mar- hatchlings (Witherington 1997).
ler 1990), it is possible that noise-dependent behav- Unfortunately, however, light management strate-
ioral changes to songs could render individuals unat- gies targeted for one species may not work for others.
tractive to females or, worse yet, unrecognizable as Lighting used along beaches in Florida that minimizes
species-specific signals. In other words, the flexibility orientation problems for hatchling sea turtles also re-
that outwardly appears to be a mechanism used to cope duces habitat use and foraging activity in beach mice
with human noise may actually represent an evolution- (Peromyscus polionotus) (Bird et al. 2004). This contrast
ary trap (Schlaepfer et al. 2002). Although the mecha- highlights the need to understand responses to anthro-
nisms responsible for reduced pairing success in noisy pogenic stressors from many species (see community-
areas remain to be sorted out, the conservation impli- level section later on). Single-species management so-
cations of a smaller proportion of the population breed- lutions have the potential to fail to accommodate the
ing has consequences of real concern to biologists. needs of other community members or, even worse,
be counterproductive by creating additional problems.
Artificial night lighting can alter signaling behavior,
Artificial Night Lighting
too. Male green frogs (Rana clamitans) call less when
As with anthropogenic noise, ecologists and conserva- exposed to artificial lighting (Baker and Richardson
tion biologists have only recently begun to explore the 2006). Both this decrease in calling activity and mice’s
range of wildlife and ecological responses to artificial avoidance of illuminated areas may be explained by
lights. This is surprising given that roughly 30% of the same mechanistic response—a heightened sense
vertebrates and 60% of all invertebrates are nocturnal of predation risk with increases in ambient light. This
(Holker et al. 2010)—clearly a large swath of diver- is a likely explanation because natural light levels can
sity that may be especially sensitive to how we change influence actual predation risk across many species
nighttime conditions. An early edited volume exploring (reviewed in Bird et al. 2004). Whether choosing not
the ecological consequences of artificial light revealed to call for mates or minimizing foraging activities to
that light has the potential to alter species distributions, reduce predation risk, these behavioral adjustments
deplete populations through direct mortality, alter spe- in response to artificial lighting could have relatively
cies interactions, and alter animal circadian and circan- straightforward fitness consequences in the form of
54 habitats in peril

lowered breeding success or creating energy deficits, in which urban and nonurban male blackbirds were
respectively. experimentally treated with light, the authors show
In contrast, predator responses to artificial lighting that urban males treated with light developed their
appear mixed. Several early studies found some bat spe- reproductive system far earlier than nonurban males,
cies to exploit the high density of insects that are at- suggesting that urban light conditions have selected for
tracted to and trapped by streetlamps (e.g., Rydell and different physiological phenotypes than in nonurban
Racey 1995). However, more recent studies have found birds. How these changes impact fitness is unknown,
other bat species to avoid artificially lit areas (Stone but these studies suggest that artificial lighting has the
et al. 2009; Stone et al. 2012). How other nocturnal potential to serve as a source of selection shaping the
predators respond to artificial lighting should be a re- physiology of birds. Whether female physiology is co-
search priority. ordinated with male physiology is unclear, and, if not,
Although not habitat per se, artificial lights lo- these changes could create mismatches that could af-
cated throughout avian migratory corridors contribute fect reproductive success.
to countless bird mortalities each year. During their A final potential consequence of artificial light-
nocturnal migration, birds are drawn to and collide ing involves the indirect effect on wildlife mediated
with structures ranging from buildings to radio tow- through the effect of lighting on invertebrate com-
ers because they are attracted to their artificial lights munities, which can be essential to trophic chains and
(Gauthreaux and Belser 2006). The precise reasons for transferring materials from aquatic to terrestrial
for birds’ attraction to night lighting is far from clear, ecosystems (e.g., Perkin et al. 2011). The attraction
but explanations range from artificial lights interfering and trapping of insects by artificial lighting is a well-
with birds’ ability to orient through magnetism (Poot known phenomenon (e.g., Eisenbeis 2006). Regardless
et al. 2008), temporarily blinding birds, or through of why insects are drawn to artificial lights, it remains
“capture” by disorienting them and altering their flight unclear to what extent insect mortalities associated
paths (Gauthreaux and Belser 2006). Regardless of with artificial lights have larger consequences for the
the mechanism, in efforts to reduce birds’ attraction population or community. One recent study exam-
to lit structures, researchers have tested whether birds ined the attraction of moths to different wavelengths
respond differently to various light colors. Poot et al. of artificial light (van Langevelde et al. 2011). Lamps
(2008) found birds to be most attracted to white and with shorter wavelengths attracted more individuals
red light and less attracted to blue and green light. This and species than lamps with longer wavelengths. Ad-
initial study provides hope that mitigation efforts will ditionally, larger species with large eyes were especially
become available to decrease bird collisions with man- attracted to lamps with shorter wavelengths, presum-
made structures. ably because they are more attracted to ultraviolet light
Although artificial lighting may pose a serious than smaller species. Another study focused on ground-
threat to birds migrating long distances, its influence dwelling invertebrates found artificial lighting to alter
on birds during other life-history stages is less clear. the composition of these communities (Davies et al.
Several breeding European songbirds sing earlier when 2012). The authors report that predatory and scaveng-
their territories are exposed to night lighting and at ing taxa were in higher abundances near streetlights
least one species’ breeding biology changes drastically both during the day and at night, suggesting that they
(Kempenaers et al. 2010). Blue tit (Cyanistes caeruleus) were not attracted to the stimulus energy of the lights
females lay eggs a few days earlier and males are two but to other changes in the community structure. It is
times more successful in attracting extra-pair mates unknown whether the increase in these taxa was due
in lit territories than nonlit areas. A separate study to an increased abundance of prey species that are at-
confirmed these temporal advances in reproductive tracted to lights or due to other factors. These two stud-
physiology in the European blackbird (Turdus merula) ies provide a first glimpse of how insect communities
but also that they molt earlier (Dominoni et al. 2013). change in response to night lighting and raise impor-
Critically, however, in a creative experimental design tant questions regarding the extent of cascading effects
 habitat loss and degradation 55

through mutualistic interactions, such as pollination, (Haematopus palliatus) spend less time incubating with
or predator-prey interactions that could reverberate increased off-road motorized traffic (McGowan and Si-
through trophic levels. mons 2006). Typically, behavioral responses like these
have been assumed to have fitness costs. However, in
this particular case, McGowan and Simons (2006)
Recreation and Presence of Humans
found no evidence for reduced reproductive success
Management of protected areas is increasingly chal- with less time incubating. More studies need to try to
lenged with balancing the needs of wildlife and pro- quantify whether the behavioral changes we measure
viding recreational opportunities for citizens. For translate into actual costs for the individual and the
example, in the United States, the number of people population.
participating in outdoor recreation appears to be in- Perhaps more troublesome is that forms of recre-
creasing exponentially (Reed and Merenlender 2008). ation that lack the sounds and speeds of motorized
Research on the influence of these activities on wild- forms of recreation might also have serious conse-
life, and especially wildlife behavior, has a long history quences for wildlife. For example, desert bighorn sheep
(e.g., Knight and Gutzwiller 1995), but much of it has (Ovis Canadensis nelsoni) in Canyonlands National
fallen short of documenting population-level impacts. Park, Utah, respond more strongly to hikers than to
Consumptive uses, such as unregulated hunting and vehicle or mountain-biking traffic (Papouchis et al.
fishing, have obvious negative consequences by directly 2001). Elsewhere in Utah, the flushing responses of
reducing wildlife population sizes, but nonconsump- bison (Bison bison), mule deer (Odocoileus hemionus),
tive recreation can also create problems. For example, and pronghorn antelope (Antilocapra americana) did
the onset of hiking in a wildlife refuge in New England not differ when responding to hikers versus mountain
triggered a sharp decline in two wood turtle (Clemmys bikers (Taylor and Knight 2003), suggesting that they
insculpta) populations (Garber and Burger 1995), sug- perceived both of these intrusions as a threat worth
gesting that even outwardly benign activities can have escaping. Recreationists may not recognize flushing
negative consequences for wildlife. responses as problems for wildlife, but such responses
Some types of nonconsumptive recreation appear could cause breeding animals to leave young unpro-
more likely to affect wildlife. For example, animals tected from harsh environmental conditions or preda-
might respond to snowmobiles, off-road motorized ve- tors or have energetic consequences by forcing animals
hicles, or boat traffic by fleeing or hiding, which could to spend less time foraging or resting.
result in less time foraging, contributing to declining The presence of people might also render areas un-
health or condition (Frid and Dill 2002). Several stud- suitable for many species and cause dramatic shifts in
ies have shown wildlife to experience elevated baseline the composition of communities. For example, Reed
stress hormones in response to motorized recreation and Merenlender (2008) used scat surveys to compare
and vehicles. For example, both wolves (Canis lupus) carnivore communities in protected areas that permit-
and elk (Cervus elaphus) experience heightened physi- ted hiking relative to those that did not in Northern
ological stress in response to snowmobile traffic (Creel California. Areas where hiking was not permitted had
et al. 2002). Elevated stress has also been associated five times the native carnivore density than areas where
with road density for elk (Millspaugh et al. 2001). From hiking was permitted. Scat of domestic dogs was not
these studies, it is not clear whether elevated stress is detected in areas without recreation, but it was found
due to the physical presence of vehicles, the association in high densities in areas that permitted hiking. Out-
between vehicles and humans, or the noise they create. wardly, one might suspect that the presence of dogs
Elevated stress resulting from these responses to hu- might be responsible for the large differences in carni-
man activities could lead to higher risk of disease and vore communities between sites, and this perception
declines in reproductive success or even survival. Mo- is reflected in many management practices that aim to
torized recreation might also influence reproduction limit the potential impacts of dogs by excluding them
in other ways. For example, American oystercatchers from protected areas or implementing leash laws (For-
56 habitats in peril

rest and St. Clair 2006). However, Reed and Meren- activities could play an important role in the evolution
lender (2011) conducted a follow-up study involving of populations. Whether anthropogenic forces less ob-
dog management practices and found no difference vious than road kill can also serve as strong sources of
in the density or species richness of native carnivore selection should be a priority in future research.
species in areas where dogs were permitted off-leash, Collectively, these studies suggest that obviously
permitted on-leash, or not permitted at all. Instead, the detrimental and outwardly harmless forms of recre-
authors argue that the presence and abundance of hu- ation can have consequences for wildlife that span
man visitors drive these patterns. behavioral adjustments, population demographic, and
Although dogs may not influence the abundance evolutionary change, plus community-level changes.
and diversity of mammalian carnivores in protected Most of these examples demonstrate changes in be-
areas, birds appear to be more sensitive. A study in havior, such as fleeing or hiding, and some document
Australia found that bird detections and estimates of changes in the communities present in areas used by
species richness were more severely affected by ex- recreationists. To what degree the findings from the
perimental treatments of humans walking with dogs first set of studies are linked to the second remains
on-leash than humans walking alone (Banks and Bry- unclear, but it has often been assumed that behavioral
ant 2007). These temporary disturbances that displace responses to avoid humans adequately reflect vulner-
birds or alter their behavior might not always have ability to human activity (Gill et al. 2001). Testing this
long-lasting consequences, but they can cause wildlife link in a quantitative manner remains an important
to abandon otherwise suitable habitats in some circum- goal for wildlife-disturbance studies (Blumstein and
stances. For example, a recent study in Finland com- Fernández-Juricic 2010). Still needed are studies that
pared breeding bird communities along hiking trails determine at what threshold flushing responses to hu-
to those in undisturbed control areas within Oulanka mans translates into permanent site abandonment or
National Park (Kangas et al. 2010). They found that the measurable declines in individual fitness due to ener-
composition of the bird community differed between getic trade-offs.
these two areas but that total species richness did not.
The difference appeared to be driven by high sensitivity
A Community-Level Lens of Habitats in
of ground-nesting species to hikers.
Peril: Interactions and Indirect Effects
Finally, recreation and other human activities have
the potential to impact the evolutionary trajectories of In most studies involving habitat loss or degradation,
populations. For example, estimates of bird mortali- we infer that the decrease in abundance of wildlife is
ties from collisions with vehicles are expected to ex- the direct result of the change in habitat quality. Yet
ceed eighty million per year in the United States (Er- we know natural communities are usually more com-
ickson et al. 2005), and a recent study suggests that plicated and interactions among community members
these mortalities may not only be nonrandom within a can make it difficult to simply draw a straight line
population but also declining, which is suggestive of an between a measure of degradation and a documented
evolutionary response (Brown and Brown 2013). Spe- response. Instead, many responses documented in de-
cifically, Brown and Brown (2013) reported that cliff graded areas are likely indirect effects that are medi-
swallows (Petrochelidon pyrrhonota) killed by vehicles ated by interactions among species. Additionally, re-
accessing a popular recreation site have longer wings gardless of whether a species’ distribution changes as a
than the population at large and that over a thirty-year direct or indirect result of habitat degradation, it is also
period both wing length and number of road-killed important to ask what the consequences of that change
swallows has declined. Although the authors could will be for the larger ecosystem. Community-level ap-
not control for additional variables that could explain proaches to understanding the ecological responses to
some of these patterns—such as the density of scav- habitat degradation will provide much greater insights
engers that remove killed birds from the road and bias on not only the mechanisms driving ecological change
road-kill estimates or temporal changes in the sizes and but also the consequences for organisms with no direct
shapes of vehicles—this study illustrates that human link to the disturbance of interest.
 habitat loss and degradation 57

Figure 5.5 Pathway by which noise directly (solid lines) and indirectly (dashed lines) affects wildlife species, predatory-
prey dynamics, and interactions among species and plants. See text for full description.

I highlight this community-level approach with the avian community based on vocal frequency should
work from my own long-term research involving the explain negative (avoidance) or neutral (persistence)
effects of noise on natural communities. Starting in responses but should not explain positive (attraction)
2005, my colleagues and I have isolated chronic an- responses to noisy areas.
thropogenic noise as a single experimental stimulus A possible explanation for the positive responses by
by sampling woodlands surrounding natural gas wells some species comes from detailed study of the avian
with and without noise-producing compressors. Initial community’s nesting success and nest-predation exper-
findings revealed that noise, in the absence of other iments using artificial nests. In contrast to the pattern
anthropogenic stimuli, reduces the breeding bird com- of declining reproductive success with noise discussed
munity’s species richness and alters the composition of earlier, in our study system nest success increases with
the community such that communities in noisy areas noise levels (Francis et al. 2009; Francis et al. 2012b;
were not simply subsets of communities in quiet areas fig. 5.5B). This pattern is evident both among and
(Francis et al. 2009; fig. 5.5A). For example, some spe- within species and is driven by decreases in predation
cies, such as the house finch (Carpodacus mexicanus) pressure with increases in noise levels because a key
and black-chinned hummingbird (Archilochus alexan- nest predator, the western scrub-jay (Aphelocoma cali-
dri), nest almost exclusively at noisy sites. This filter- fornica), tends to avoid noisy areas (figs. 5.5C, D). Given
ing appears to be largely explained by species-specific recent evidence that birds use acoustic cues represen-
vocalizations (Francis et al. 2011b), as discussed previ- tative of predation risk to make settlement decisions
ously. Importantly, however, this nonrandom sorting of (e.g., Emmering and Schmidt 2011), it is possible that
58 habitats in peril

the positive responses to noise observed in finches, studies that adopt a more rounded research approach
hummingbirds, and other species actually represent that measures responses by multiple organisms simul-
settlement decisions made based on the absence of taneously and explicitly considers relationships among
acoustic cues representative of predation risk by jays. community members.
Noise and other stressors can also have indirect
effects on ecological systems with long-lasting conse-
Key Research Needs for Moving Forward
quences. The western scrub-jay and black-chinned hum-
Measuring Noise and Light and Quantifying
mingbird each provide important ecological services for
the Magnitude of Their Effects
plants in the form of seed dispersal and pollination, re-
spectively. In our study area, noise appears to have both Although research on artificial lighting and noise
positive and negative indirect effects on plants receiv- pollution is still rather new, the early work on these
ing these services (Francis et al. 2012a). Experiments anthropogenic stimuli clearly shows that they can be
using artificial flowers suggest that the high abundance problematic for many wildlife species and communi-
of hummingbirds in noisy areas leads to more visits to ties. To fully understand the mechanisms responsible
flowers and a greater movement of pollen among them for various responses and unravel the cascading eco-
relative to flowers in less noisy areas (figs. 5.5E–G). In logical consequences of their effects, future research
contrast, experiments focused on piñon pine (Pinus edu- on the impacts of these stimuli must be grounded in
lis) seed dispersal suggest that the scrub-jay’s avoidance the sensory biology of the organism(s) and functional
of noisy areas appears to disrupt seed dispersal services aspects of the stimulus. Why is this so? Because spe-
and lead to reduced seedling recruitment in noisy areas cies have unique sensitivities to different wavelengths
(Francis et al. 2012a; figs. 5.5C, H, I). An interesting sup- of light and sound, and artificial lighting and noise
plement to this pattern is that mice (Peromyscus) prey stimuli also vary drastically in their spectral content.
upon piñon seeds more often in noisy relative to quiet These stimuli can also vary in their predictability, in-
areas (figs. 5.5I–K), perhaps representing an indirect ef- tensity, and duration (Francis and Barber 2013). This
fect of noise mediated by reduced competition for food variation guarantees that species will differ in their re-
resources or less predation pressure from acoustically sponses to these stimuli. For this reason, it is critical
oriented nocturnal predators that may be sensitive to that studies properly characterize noise or light stimuli
noise (e.g., owls). While these mechanistic causes for but also pay special attention to the features that will
mice and other species’ responses to noise still must be be of functional significance to the organism. In doing
sorted out, the long-term consequences to ecosystem so, there is substantial promise in finding appropriate
structure resulting from declining seed recruitment of a and cost-effective mitigation techniques to minimize
dominant tree is of considerable conservation concern. the effects of these stimuli on wildlife and the ecologi-
Collectively, this example illustrates that anthropo- cal systems that support them. Additional information
genic stressors can have indirect effects that reverber- on properly characterizing noise and wildlife responses
ate throughout natural communities. For our study can be found in several recent papers (Pater et al. 2009;
system, the importance of the scrub-jay’s response to Ellison et al. 2012; Francis and Barber 2013).
noise for other organisms suggests that research on Also important is describing the spatial footprint
wildlife with important ecological roles within a com- of these two stimuli and linking their variation across
munity could be particularly important to understand- space to actual ecological impacts. Most research to
ing the full ecological costs of environmental degra- date, including some of my own work, has evaluated
dation. Thus, additional research focused on predator the influence of these stimuli using study designs in
responses could be especially revealing; their responses which the stimulus is either present or absent (i.e., lit
could cause the disruption of top-down effects, which versus dark, quiet versus noisy). Many of these studies
can trigger comprehensive changes to the structure of have been instrumental in opening our eyes to the fact
both heterotrophic and autotrophic organisms within a that these stimuli can have important consequences
community (reviewed in Estes et al. 2011). Clearly, un- for wildlife, but they are of little use to management
derstanding these cascades will only come from careful and policy because they say nothing about how wildlife
 habitat loss and degradation 59

responses change with incremental changes in noise we assume that species that remain in areas exposed
or light conditions. Because policy makers and mana- to human activities or other forms of anthropogenic
gers are interested in questions like “how much is too environmental change are doing well and likely need
much?,” future studies need to adopt study designs no form of protection. This assumption is problematic
from research groups that describe how responses vary because distributional changes may only occur if indi-
along gradients of noise and light exposure (e.g., Fran- viduals have alternative areas to go to (Gill et al. 2001)
cis et al. 2011c; Halfwerk et al. 2011; Kight et al. 2012). and, as illustrated through several examples discussed
Doing so has the potential to provide meaningful data previously, known impacts beyond site abandonment
that reflects how wildlife respond to subtle changes in are plentiful and will undoubtedly grow given the
these stimuli—information that will be essential to set- number of probable effects yet to be investigated. It
ting regulatory limits. may be that the most vulnerable populations are those
in which site abandonment is unlikely because other
suitable habitats are unavailable or the costs associated
When Do Avoidance Responses to
with moving to them are too high (Gill et al. 2001). For
Recreational Activities Translate into
populations in decline, but for reasons other than site
Negative Consequences for the Individual
abandonment, we certainly want to know why, and ef-
or Population?
forts need to clarify whether it is an issue of decreased
Research involving wildlife responses to recreation will survival, reproduction, or both. Critically, we also want
greatly benefit from explicitly linking observed behav- to know whether responses are direct or mediated
ioral responses to recreational activities to fitness con- through other organisms in the community. Measuring
sequences. There are now countless examples of wild- multiple responses by many species will be logistically
life fleeing from approaching hikers, snowmobiles, or challenging and more expensive than many current
other common forms of recreation (reviewed in Knight efforts. However, these measurements are unambigu-
and Gutzwiller 1995), and perhaps one generalization ously important to informing conservation policies that
is that human presence that is both unpredictable and have the potential to work.
close to wildlife elicit stronger behavioral responses
(Kight and Swaddle 2007; Blumstein and Fernández-
Summary
Juricic 2010). Although many examples of these re-
sponses exist, we still need a greater understanding Land cover change is just one way that habitats can be
of whether these and other behavioral responses de- degraded or lost to wildlife, and research studies high-
crease an individual’s ability to survive or reproduce. lighted in this chapter emphasize why focusing solely
Ultimately, we want to know whether there are costs on the amount, type, and spatial arrangement of land
to individuals and if they lead to population-level con- cover is inadequate in explaining animal responses to
sequences. Frameworks exist that should aid research- habitat degradation. Instead, investigators must con-
ers in studies aiming to link behavioral responses to sider forces that contribute to habitat loss and degra-
individual costs. For example, the risk-disturbance hy- dation without the removal or change in vegetation,
pothesis explicitly views animal responses to human such as noise and light pollution and the presence of
activities as analogous to their responses to predation humans and our pets. Similarly, single-species studies
risk (Frid and Dill 2002). Whether this framework is focused on the presence or abundance of a species pro-
used or not, investigators will need to think about these vide an incomplete picture of how wildlife and their
responses in terms of energetic trade-offs and how in- supporting ecological systems respond to human activi-
dividuals maximize fitness. ties. Future work can benefit from an organism-centric,
sensory-based approach that links the proximate mech-
anisms governing responses to anthropogenic stressors
Moving beyond Occupancy and Abundance
to their individual-, population-, and community-level
More studies should go beyond simple measures of site consequences. As we learn more about the severity and
occupancy or abundance. The greatest risk is when extent of these stressors’ impacts on natural systems,
60 habitats in peril

we may also discover unforeseen sensory-based solu- Society A: Mathematical Physical and Engineering Sciences
tions that could help maintain wildlife populations in 369:1010–35.
habitats once thought to be lost. Ellis, E. C., K. K. Goldewijk, S. Siebert, D. Lightman, and
N. Ramankutty. 2010. “Anthropogenic Transformation of
the Biomes, 1700 to 2000.” Global Ecology and Biogeography
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Evolution 17:474–80. Sea Turtles and Other Nocturnal Animals.” In Behavioral
Schroeder, J., S. Nakagawa, I. R. Cleasby, and T. Burke. 2012. Approaches to Conservation in the Wild, edited by J. R. Clem-
“Passerine Birds Breeding under Chronic Noise Experience mons and R. Buchholz, 303–28. Cambridge University
Reduced Fitness.” Plos One 7:e39200. Press, Cambridge.
 6
Lisette P. Waits and
Population Genetics and
Wildlife Habitat
Clinton W. Epps

W orldwide, wildlife habitat has been lost, de-

graded, and fragmented as humans have trans-
formed the landscape for agriculture, transportation,
and genetic drift (Hedrick 2009; Allendorf et al. 2012).
Mutation is the process that creates new alleles due to
errors in DNA replication. Gene flow occurs as a re-
and urbanization. The amount, quality, and configura- sult of migration (dispersal and reproduction), and this
tion of the remaining wildlife habitat can greatly affect process moves alleles between populations. Selection
genetic and evolutionary processes and the viability of impacts genetic variation by increasing the frequen-
these populations. Decreases in the size and quality of cies of alleles that are favorable in a particular environ-
habitat fragments lead to decreases in effective popula- ment and decreasing the frequencies of alleles that are
tion size, increases in genetic drift, and loss of genetic less favorable in that environment. Genetic drift is the
diversity, while increased isolation of fragments leads change in allele frequencies due to random sampling
to decreases in gene flow and loss of genetic diversity effects as alleles are passed on from one generation to
(fig. 6.1). Loss of genetic diversity is of great concern to the next (Wright 1931). Genetic drift is thus influenced
wildlife managers because this loss increases the proba- by the number of breeders and the variance in repro-
bility of inbreeding depression, decreases the adaptive ductive success among breeders.
potential of populations, and ultimately threatens popu- Sewall Wright (1931, 1938) introduced the concept
lation viability. This chapter will begin with a review of of effective population size (Ne) to mathematically and
important population genetic theories, methods, and conceptually represent the effects of genetic drift. He
terms as they relate to habitats in peril. Specifically, defined Ne as “the number of breeding individuals in an
we (1) describe the impacts of habitat degradation and idealized population that would show the same amount
fragmentation on population genetic processes; (2) dis- of dispersion of allele frequencies under random ge-
cuss the implications for population viability, hybrid- netic drift or the same amount of inbreeding as the
ization, and population persistence; and (3) highlight population under consideration.” The Ne of a popula-
the role of molecular methods in assessing the impacts tion has a large effect on genetic diversity. Populations
of habitat alteration on wildlife populations. Since with larger Ne will have more genetic variation because
genetic terms may be unfamiliar to some readers, we more new alleles are created by mutation due to a larger
provide a glossary of key terms in box 6.1 and highlight number of breeding events, and fewer alleles are lost to
each glossary term in bold when it is first used. genetic drift since there is a larger sample of breeders
each generation. Ne also influences the relative impacts
of genetic drift and selection in natural populations
Population Genetic Methods and Theory
(Fisher 1930; Wright 1931). For example, as Ne de-
Genetic variation in natural populations is influenced creases, the effect of selection decreases relative to the
by four main processes: mutation, gene flow, selection, effect of genetic drift and important adaptive genetic
64 habitats in peril

Figure 6.1 Impacts of habitat fragmentation on

gene flow, effective population size (Ne ), and
genetic diversity.

variation can be lost (Hedrick 2009). One benchmark 1994). Noninvasive genetic sampling is achieved with-
for conservation biology is the “50 / 500 rule” (Franklin out capturing or handling individual animals and in-
1980), which refers to minimum Ne required to prevent cludes the use of a diversity of sources including hair,
inbreeding (50) and offset the consequences of long- feces, feathers, urine, owl pellets, sloughed and shed
term genetic drift (500). Although this rule may have skin, eggshells, and saliva (Waits and Paetkau 2005).
little utility as a prescription for minimum viable popu- When examining genetic variation, researchers may
lation sizes (Templeton 1995), it serves as a reasonable select a location (locus) or locations (loci) in the mi-
guide to determining when genetic factors should be tochondrial DNA (mtDNA) or nuclear DNA (nDNA)
considered (Jamieson and Allendorf 2012); thus, an Ne genomes. MtDNA is a circular DNA molecule of the
<500 may be regarded as small, and an Ne <50 is re- mitochondrion, an organelle that is the energy power-
garded as very small. house of cells. MtDNA generally has a haploid mode of
When studying genetic variation and gene flow in inheritance as it is passed only from mother to offspring
natural populations, researchers use a diversity of meth- (Birky et al. 1989). In wildlife genetics, this DNA mol-
ods and DNA types. The collection of DNA samples ecule is important for species identification, phyloge-
from wildlife can generally be classified into three main netic and phylogeographic analysis and the delineation
sampling approaches: destructive sampling, nonde- of conservation units. Two copies of nDNA are found
structive sampling, and noninvasive sampling (Taberlet in the chromosomes of the nucleus of a cell. Nuclear
et al. 1999). In destructive sampling approaches, the DNA has a biparental mode of inheritance as one copy
animal is killed to obtain the genetic sample of blood is inherited from the mother and one copy is inherited
or tissue. This approach was widely used during the from the father. In wildlife genetics, nDNA analyses
1970s and 1980s when analyzing protein data (i.e., are important for individual identification, kinship and
allozymes) but has been replaced by nondestructive parentage studies, gene flow evaluations, and the delin-
and noninvasive genetic sampling for vertebrate ani- eation of conservation and management units (Allen-
mals. Nondestructive sampling involves capturing an dorf et al. 2012). When generating mtDNA or nDNA
animal and obtaining a blood, tissue, feather, mouth data, researchers choose to evaluate particular loci in
swab, or hair sample and then releasing the animal. the genome. These loci may be neutral or nonneutral
Occasionally, nondestructive sampling is conducted (box 6.1). Neutral loci are not affected by natural selec-
without capturing individual animals by using a biopsy tion while nonneutral or adaptive loci affect individual
dart gun on large mammals such as whales, elephants, fitness and are under selection (Hedrick 2009).
and cougars (Hoelzel and Amos 1988; Georgiadis et al. After choosing to collect data for a particular type of
Box 6.1 Glossary of Genetic Terms

allele: A unique genetic variant observed at a hybrid zone: An area of contact between two ge-
particular locus. netically distinct populations where hybridiza-
allele frequency: A measure of the relative fre- tion occurs.
quency of an allele in a population at a particu- inbreeding: Mating between closely related indi-
lar genetic locus. viduals.
bottleneck: A sharp reduction in population size, inbreeding depression: A decrease in fitness due to
leading to a loss of genetic diversity. breeding between closely related individuals
census population size (Nc ): The number of indi- (inbreeding).
viduals in a population, as distinguished from introgression: Gene flow between populations
Ne (see effective population size). whose individuals hybridize. This occurs due to
effective population size (Ne ): The number of backcrossing of hybrids to one or both parent
breeding individuals in an idealized population groups.
that would show the same amount of change locus (loci): A particular location in the genome of
in allele frequencies under random genetic an organism. The term loci is the plural form of
drift or the same amount of inbreeding as the locus.
population under consideration. mitochondrial DNA (mtDNA): A circular DNA mol-
fitness: The relative ability of an individual (or ecule of the mitochondrion, which is haploid
population) to survive, reproduce, and propa- and generally passed on only from mother to
gate genes in an environment. offspring.
gene flow: The movement of alleles between mutation: A change in the DNA sequence of an
populations. This is also referred to as migra- organism. Mutations that occur in the DNA
tion. of germ cells can be passed on to the next
genetic drift: The change in allele frequencies generation.
due to random sampling effects as alleles are neutral locus: A locus where the frequency of al-
passed on from one generation to the next. leles is not affected by selection.
genetic rescue: Increasing genetic diversity of a nonneutral or adaptive locus: A locus where the
population lacking genetic diversity due to frequency of alleles is affected by selection.
drift or inbreeding by adding new individuals nuclear DNA (nDNA): The DNA of the chromo-
from a different source population. somes found in the nucleus of a cell.
genetic structure: Spatial variation in allele identity outbreeding depression: A reduction in fitness
or frequency. of hybrid individuals relative to the parental
genotype: The combination of alleles observed at a types.
particular locus for a specific individual. population viability analysis (PVA): A quantitative
haplotype: An allele or combination of alleles analysis of population dynamics with the goal
passed on from a single parent. of assessing extinction risk.
hybridization: Interbreeding of individuals from positive selection: A process that increases the
genetically distinct populations, subspecies, or prevalence alleles that increases an individual’s
species. fitness.
hybrid swarm: A population of individuals that all purifying selection: A process that decreases the
are hybrids. prevalence of alleles that diminish an indi-
vidual’s fitness.
66 habitats in peril

Box 6.2 Overview of Methods Used in Genetic Analysis

polymerase chain reaction (PCR): The chemical and is the most commonly used approach for
process used to make millions of copies of a evaluating genetic diversity, gene flow, parent-
particular target DNA region or locus using age, and hybridization in wildlife populations
specific primers (Mullis and Faloona 1987). (Selkoe and Toonen 2006).
PCR has revolutionized molecular genetics and amplified fragment length polymorphism (AFLP)
is used in all following methods. analysis: A PCR-based tool for assaying genetic
DNA sequencing: The chemical process used to variation that does not require knowledge of
read the sequence of nucleotides (DNA base the DNA sequence of the target species. The
pairs—A, G, C, T ) at a particular DNA region. amplified DNA fragments are separated by size
Early studies in wildlife genetics were domi- on a polyacrylamide gel and visualized using
nated by a method known as Sanger sequenc- fluorescence or autoradiography (Vos et al.
ing (Sanger et al. 1977), but there are currently 1995).
many new methods categorized as “next gen- single nucleotide polymorphism (SNP) analysis: A
eration sequencing” approaches that parallelize new molecular method that surveys single base
the process and can produce a greater volume pair genetic polymorphisms in many locations
of sequence data in a shorter period of time throughout the genome. This method has not
(Shendure and Ji 2008). yet been used extensively in wildlife genetics,
microsatellite analysis: Also known as short tandem but the use of this method is predicted to ex-
repeats (STRs) or simple sequence repeats pand rapidly in coming years because it surveys
(SSRs). Repeating sequences of two to six a greater proportion of the genome and can
base pairs (e.g., CACACA or GTCGTCGTC). be automated for high throughput analyses
The process of genotyping microsatellite loci (Vignal et al. 2002; Morin et al. 2004).

DNA and locus, a variety of methods are used to obtain crease genetic structure (i.e., differentiation between
genetic data in the form of haplotypes or genotypes. groups). This occurs because spatial distribution and
Fully describing these methods is beyond the scope of quality of habitat are key determinants of population
this chapter, but the most commonly used approaches size and dispersal success, which in turn influence Ne
are summarized in box 6.2. The most important meth- and thus genetic drift. Therefore, rather than simply
odological advance in wildlife and medical genetics describing genetic structure, genetic diversity, or gene
was the development of the polymerase chain reaction flow, researchers increasingly have attempted to deter-
or PCR (Mullis and Faloona 1987). This technique mine how habitat distribution and quality influence
makes it possible to make many copies of particular those characteristics of natural populations. This in-
DNA locus and to use low-quantity DNA sources like terest has contributed to the increasing popularity of
feces, saliva, or bones. We refer readers to the follow- landscape genetic studies for both plants and animals
ing sources for additional information on methodologi- (Manel et al. 2003; Holderegger et al. 2010; Storfer
cal approaches used in wildlife genetics (DeYoung and et al. 2010).
Honeycutt 2005; Beebee and Rowe 2008). Habitat area is often assumed to be the strongest
predictor of population size and thus Ne, particularly
for metapopulations and other systems of fragmented
Impacts of Habitat Loss and
populations (Hanski 1999). This concept can be traced
Fragmentation on Gene Flow, Ne ,
to the theory of island biogeography, which posits that
and Diversity
island area is the most important predictor of species
Habitat loss, fragmentation, and degradation may extinction rate (MacArthur and Wilson 1967). How-
strongly reduce genetic diversity and typically in- ever, this assumption is not always supported (Prugh
 population genetics and wildlife habitat 67

et al. 2008), because habitat quality and configuration “Implications of Changes in Gene Flow and Genetic
(e.g., edge effects or links to other habitat patches) Diversity”). Thus, the impacts of habitat loss, degrada-
may have greater importance. Genetic structure of tion, and fragmentation on small populations may be
fragmented populations is affected by both genetic intensified by loss of fitness related to genetic diversity.
drift and gene flow (Hutchison and Templeton 1999); Other factors may interact with habitat quality and
in systems of small populations, connectivity between fragmentation to influence Ne and thus genetic diver-
patches can have the greatest impact on genetic di- sity. The stability of populations matter: populations
versity within any single population or habitat patch that fluctuate in size will lose more genetic diversity
(e.g., Epps et al. 2006). Therefore, Ne and thus genetic than stable populations (Frankham 1995a). The ef-
diversity may be eroded by (1) habitat loss; (2) habitat fect is worsened if a population declines to a small
degradation, even if habitat area doesn’t decrease; or size for multiple generations (known as a bottle-
(3) habitat fragmentation (fig. 6.1). Although habitat neck), although populations that expand rapidly after
fragmentation is often closely associated with habi- a short-term decline may lose little genetic diversity
tat loss (Fahrig 2003), it can also occur without any (Nei et al. 1975). Thus, strong temporal variation in
change in habitat area for naturally patchy populations habitat quality could reduce Ne. A variety of statistical
if landscape changes influence the ability of organisms tests can be used to deduce demographic events such as
to disperse between patches. Transportation networks population bottlenecks from genetic data (Cornuet and
(roads, railroads, etc.) are likely sources of this type Luikart 1996; Piry et al. 1999; Garza and Williamson
of fragmentation (Balkenhol and Waits 2009; Hold- 2001). Polygynous mating systems where variance in
eregger and Di Giulio 2010). A good example of this reproductive success is high may also result in Ne being
is the effect of fenced highways and canals on desert much smaller than census population size (Nc). If, for
bighorn sheep (Ovis canadensis nelsoni) in California, instance, a single dominant male in a small population
USA. Although those structures are primarily located fathers a large proportion of the offspring, and other
in the arid flats rather than the mountainous habitat males have many fewer offspring, genetic drift and
used by bighorn sheep for foraging and escape terrain, inbreeding will occur more rapidly than expected for
Epps et al. (2005) showed that they contribute to in- species with monogamous mating (Frankham 1995a).
creased genetic structure and reduced genetic diver- Spatial structuring within a population (i.e., nonran-
sity in those small populations. In other cases, such as dom association of groups of individuals) will typically
Kuehn et al.’s (2007) study of roe deer populations in reduce Ne for the population as a whole because drift
Switzerland separated by a fenced highway, roads cause will be accelerated by the comparatively smaller size of
genetic divergence but no detectable decrease in ge- each group. As such, spatial structuring may be cryptic,
netic diversity. especially when movement and dispersal through dif-
Whatever the cause, as Ne declines, genetic drift will ferent habitats are poorly understood, and reductions
increase and genetic diversity will be lost as noted pre- in Ne may be more common than readily apparent, even
viously. Strong genetic drift can even override balanc- for widely distributed species. For instance, Halbert
ing selection (when heterozygous individuals have a fit- et al. (2012) demonstrated that cryptic genetic struc-
ness advantage, such as disease resistance) or positive ture exists within the Yellowstone National Park bison
selection for an advantageous allele. Another problem herd, which is one of only two bison populations that
that arises in small or fragmented populations is in- have persisted in their native habitat since European
creased levels of inbreeding, which will exacerbate loss settlement of North America. They estimated that Ne
of genetic diversity from drift. Inbreeding occurs when of each subpopulation was ~100 and noted that this
close relatives mate and bear offspring, and it results in apparent isolation requires reconsideration of current
decreases in heterozygosity because related individu- management practices, including culling.
als are more likely to have the same allele at any given For a species as a whole, spatial structuring of popu-
locus (Haldane and Waddington 1930). Decrease in fit- lations (i.e., populations divided into subpopulations
ness resulting from inbreeding is referred to as inbreed- with limited dispersal) can help maintain higher ge-
ing depression (see examples in the following section, netic diversity (Templeton et al. 1990). If the number
68 habitats in peril

of subpopulations is large, even if drift causes fixation in different environments, and such patterns are start-
of a single allele (i.e., loss of all other alleles due to ing to be detectable in natural systems (Schoville et al.
drift) in each subpopulation, different alleles will be 2012). For example, a latitudinal gradient in allele fre-
fixed in different subpopulations by random chance, quencies for a gene associated with reproductive tim-
resulting in maintenance of diversity in the system as a ing has been observed in Chinook salmon (Oncorhyncus
whole. However, it is important to recognize that each tshawytscha) (O’Malley et al. 2010). Spatial gradients in
subpopulation may still suffer inbreeding depression or fitness of a particular allele are one of the reasons why
reduced fitness, so fragmentation should not be viewed managers must use caution when seeking to alleviate
as an appropriate strategy for maintaining regional or inbreeding by bringing in individuals from other pop-
species-wide genetic diversity in most cases. ulations. If the source stock for those augmentations
Most research investigating the relationship comes from different habitats, outbreeding depres-
between population genetic structure and habitat sion is possible (Dobzhansky 1948). Outbreeding de-
has focused on neutral genetic diversity. Neutral ge- pression occurs when co-adapted gene complexes are
netic loci offer an excellent means to infer the effects disrupted and can lead to serious problems such as re-
of habitat quality and configuration on demographic producing at the wrong time of year in a particular hab-
history, genetic diversity, and gene flow, as described itat. Outbreeding depression in natural populations has
earlier, and may have future adaptive potential, but in- been observed across many taxa, including birds (Marr
ferring fitness consequences of low genetic diversity et al. 2002), amphibians (Jourdan-Pineau et al. 2012),
or low heterozygosity at neutral loci is difficult (Chap- fish (Roberge et al. 2008), plants (Goto et al. 2011),
man et al. 2009). Although individuals or populations and mammals (Marshall and Spalton 2000). However,
with lower genetic diversity have been shown to have recent literature suggests that inbreeding depression
reduced fitness (e.g., Johnson et al. 2011; Agudo et al. occurs more frequently than outbreeding depression in
2012), effect sizes are often very weak (Chapman et al. natural populations and that concerns about outbreed-
2009), and many studies merely assume that neutral ing are often overstated (Edmands 2007; Frankham
and adaptive genetic diversity are correlated; however, et al. 2011).
evidence is mixed. Studies tend to detect that correla-
tion especially where genetic drift is strong (Radwan
Implications of Changes in Gene Flow and
et al. 2010; e.g., Marsden et al. 2012). Until recently,
Genetic Diversity
few adaptive loci were described well enough to allow
Effects on Population Viability
population-level assessments of adaptive variation, and
many genes with adaptive significance show little varia- Ultimately, changes in gene flow and genetic diversity
tion. Exceptions can be found in research on genetic may affect population viability (Lacy 1997). Although
diversity related to immune-system function and dis- it has long been understood that low genetic diversity
ease resistance (e.g., the microhistocompatability com- has negative consequences, Lande (1988) argued that
plex or MHC genes). Studies of MHC variation show genetic drift and inbreeding are likely to pose signifi-
clear evidence of balancing selection in some systems cant threats only when census population sizes are
(Hedrick 1994), even where neutral genetic diversity is very low and thus populations are already vulnerable.
severely depleted (Aguilar et al. 2004), but in others, However, in naturally fragmented populations where
the effects of genetic drift may overwhelm evidence of local population sizes are small, such effects could ac-
selection (Worley et al. 2006; Alcaide 2010). cumulate rapidly if fragmentation prevents dispersal
New approaches to studying adaptive genetic varia- among habitat patches. Inbreeding has been demon-
tion, in some cases employing “next generation” se- strated to be a serious problem in many captive breed-
quencing techniques (box 6.2), are poised to reveal new ing programs (Ralls et al. 1988) and, increasingly, in
insights into how adaptive genetic variation is affected wild populations (e.g., Frankham 1998; Saccheri et al.
by habitat quality, distribution, and population history 1998). However, because neutral genetic diversity is
(Kirk and Freeland 2011; Miller et al. 2012; Schoville not a perfect index for adaptive genetic diversity, it has
et al. 2012). Selection may favor different genotypes proved difficult to identify thresholds of neutral genetic
 population genetics and wildlife habitat 69

diversity at which negative consequences for fitness is the US Fish and Wildlife Service’s decision to aug-
result. Despite this difficulty, as we discuss in the fol- ment the endangered population of Florida panthers
lowing paragraphs, there is clear evidence of the det- (Puma concolor) with mountain lions from Texas in
rimental consequences of inbreeding and low genetic an attempt to reverse the inbreeding effects (Pimm
diversity in natural populations (Frankham 1995b) and et al. 2006; Johnson et al. 2010). Genetic rescue may
concensus that genetic factors can affect extinction risk also occur naturally as was documented for an inbred
(Jamieson and Allendorf 2012). population of Scandinavian wolves (Canis lupus) that
Many studies have demonstrated links between neu- showed much higher heterozygosity and rapid popula-
tral genetic diversity, MHC diversity, and fitness. Cor- tion growth after a single immigrant arrived and mated
relations between neutral genetic diversity and MHC (Vila et al. 2003). Genetic rescue has been employed
diversity may indicate strong drift or inbreeding (Rad- in other situations (e.g., Hogg et al. 2006; Miller et al.
wan et al. 2010), suggesting that in many cases neutral 2012; Olson et al. 2012; Heber et al. 2013) but must
genetic diversity can be used as an index for adaptive be done cautiously. Such efforts may contribute to out-
genetic diversity. In free-ranging raccoons (Procyon breeding depression (see earlier discussion) if there are
lotor), genetic diversity at neutral microsatellite loci no closely related populations to use as sources, im-
was correlated with reduced endoparasite loads and migrants may kill resident juveniles (e.g., in large car-
prevalence of a single ectoparasite (Ruiz-Lopez et al. nivores where infanticide is common), and the spread
2012). Conversely, Schwensow et al. (2007) found that of disease is an ever-present concern (Tallmon et al.
neither neutral genetic diversity nor adaptive genetic 2004). However, it can be successful even when source
diversity (MHC alleles) correlated with parasite load populations are also inbred or of low genetic diversity
(a potential fitness indicator) in dwarf lemurs (Chei- (Heber et al. 2013). Finally, similar to other transloca-
rogaleus medius). However, they were able to identify tions (e.g., Fischer and Lindenmayer 2000; Letty et al.
particular alleles that were associated with high or low 2007), any attempt to “rescue” a small or inbred popu-
parasite burdens. lation must consider whether the habitat will support
In other cases, even when adaptive genetic diver- a population of adequate size and whether threats to
sity is not assessed, comparative estimates of very low the original population have been adequately addressed
neutral genetic diversity may be correlated with ap- (Hedrick and Fredrickson 2010).
parent inbreeding effects. For instance, Olano-Marin Increasingly, we recognize that genetic diversity is
et al. (2011) found correlations between individual het- affected by habitat area, quality, connectedness, and
erozygosity at neutral loci and reproductive success in other ecological factors. Attempts to maintain popu-
blue tits (Cyanistes caeruleus), and argued that neutral lation viability and genetic diversity must consider how
loci are more sensitive to detecting inbreeding than those processes interact. For instance, the influence of
adaptive loci. Johnson et al. (2011) detected a relation- connectivity on Ne becomes more important as average
ship between inbreeding and reproductive success for population size declines, particularly in metapopula-
Sierra Nevada bighorn sheep (Ovis canadensis sierrae) tions with high rates of population turnover, because
but observed that the effects were too small to influ- of increased rates of genetic drift. A good example of
ence population trajectories. Chapman et al. (2009) this is a study by Cosentino et al. (2012) that evalu-
reviewed many such studies and noted that heterozy- ated forty-one small populations of tiger salamander
gosity–fitness correlations were often weak and that (Ambystoma tigrinum) and demonstrated that coloniza-
there appeared to be a publication bias against studies tion history, connectivity, wetland area, and presence
that failed to find heterozygosity–fitness correlations. of predatory fish all influenced genetic diversity and
To counter the effects of low genetic diversity and divergence. Despite this complexity, because of the
inbreeding, “genetic rescue” may be employed (Tall- primary relationship between Ne and genetic diversity,
mon et al. 2004). This technique requires bringing in a basic understanding of typical population sizes is a
less closely related individuals from other populations first requirement when considering whether genetic
to inject new alleles into a genetically depauperate diversity and gene flow will have a strong influence on
population. One of the best-known examples of this population viability for a given species.
70 habitats in peril

of genes between groups. The spatial location where

Incorporating Genetics in Population
hybridization and introgression occurs is known as
Viability Analysis (PVA)
a hybrid zone (Barton and Hewitt 1985). Hybridiza-
Genetic impacts on extinction risk are rarely modeled tion is a natural evolutionary process which can lead
in population viability analyses (Allendorf and Ry- to positive impacts on biodiversity, such as an increase
man 2002; Beissinger 2002), but attempts to include in genetic diversity and fitness of hybrid individuals
genetics in the PVA of animal populations highlight the (hybrid vigor), adaptive radiations, and the creation of
importance of this variable (Mills and Smouse 1994; new species (Lewontin and Birch 1966; Arnold 1997;
Robert et al. 2004; Haig and Allendorf 2006). Given Seehausen 2004). However, it is also a conservation
the impacts of habitat loss and fragmentation on ge- and management challenge with negative impacts for
netic variation and inbreeding, failure to include this populations including outbreeding depression, loss of
variable in PVA is likely to result in overly optimistic local adaptations, genetic swamping of one parental
projections of population persistence (Allendorf and group, and, in some cases, extinction by hybridization
Ryman 2002). For example, in a simulation study of (Rhymer and Simberloff 1996; Allendorf et al. 2001;
eighteen threatened animal species with initial popu- Wolf et al. 2001).
lation sizes of fifty to one thousand individuals, Brook Allendorf et al. (2001) provided a categorization of
et al. (2002) showed that the average median time to hybridization to help guide conservation and manage-
extinction was increased 25–30% when the effects of ment decisions. Their classification is focused on the
inbreeding depression were included in PVA analyses. cause of the hybridization event (natural or anthropo-
This study also demonstrated a wide variation in im- genic) and the extent of introgression between hybrid-
pacts as species showed little decrease (<5%) in me- izing taxa. They suggest that anthropogenic hybridiza-
dian time to extinction, while others showed extreme tion events that are followed by backcrossing and
impacts with decreases in time to extinction up to 95%. introgression with parental groups are of greatest con-
Most studies that incorporate genetics into PVA cern for conservation and management and should be
generally include inbreeding depression effects only avoided when possible. The three anthropogenic activi-
at the juvenile survival stage of the life cycle, yet it is ties that contribute the most to hybridization are the
known to affect multiple stages (Mills and Smouse introduction of nonnative plants and animals, habitat
1994; Ogrady et al. 2006). When evaluating the im- fragmentation, and habitat modification (Rhymer and
pacts of inbreeding depression across the full life cycle Simberloff 1996; Allendorf et al. 2001). In a review of
of eighteen mammal and twelve bird species, O’Grady hybridization among threatened and endangered ani-
et al. (2006) found that inbreeding depression led to mals, Zamudio and Harrison (2010) found that 75% of
lower population sizes and a 31–41% median reduction hybridization cases could be linked to anthropogenic
in time to extinction. In conclusion, these studies indi- habitat change or introductions of nonnative species.
cate that the probability of survival of threatened spe- In a review of plant and animal hybrid zones, Buggs
cies in fragmented and degraded habitat will be over- (2007) concluded that human-induced habitat changes
estimated if genetic factors are not considered in PVA. were driving factors in eleven of thirty-nine cases of
hybrid zone movement, and climate change could be
shifting hybrid zone location in multiple animal spe-
Impacts of Habitat Loss and
cies. Thus, habitat loss and alteration are serious con-
Fragmentation on Hybridization
cerns for increasing the threat of hybridization among
and Introgression
native taxa.
Another potential genetic impact of human-caused The introduction of nonnative species has led to
habitat alternation is an increase in hybridization and numerous examples of hybridization in invertebrates,
introgression. Hybridization is the mating between mammals, birds, fish, reptiles, and amphibians (re-
individuals from genetically distinct populations, sub- viewed in Rhymer and Simberloff 1996; Zamudio and
species, or species. When hybrids backcross to one or Harrison 2010). A classic example of this is hybridiza-
both parental groups, this causes the introgression tion of introduced mallards (Anas platyrhynchos) with
 population genetics and wildlife habitat 71

local, endemic duck species. Hybridization and intro- tor in driving hybridization among grey wolves and
gression with mallards has been implicated in popu- coyotes (C. latrans) in the Great Lakes region of North
lation declines of the New Zealand grey duck (A. su- America (Lehman et al. 1991; Koblmuller et al. 2009),
perciliosa superciliosa; Rhymer et al. 1994), Hawaiian Algonquin wolves (C. lycaon) and coyotes in Canada
duck (A. wyvilliana; Griffin et al. 1989), Florida mottled (Wilson et al. 2000; Kyle et al. 2006), and red wolves
duck (A. fulvigula fulvigula; Mazourek and Gray 1994), and coyotes in the southeast (Roy et al. 1996; Bohling
and the Australian (Pacific) black duck (A. supercili- and Waits 2011). The impact of habitat conversion on
osa rogersi; Lever 1987). In addition, there is evidence hybridization has also been documented for savannah
that invasiveness can increase as a result of hybridiza- (Loxodonta africana) and forest elephants (L. cyclotis) in
tion. Hybridization between different populations of Africa (Roca et al. 2005).
invaders or with already-established local species may Another major impact of habitat loss and degrada-
promote the establishment of the colonists while si- tion is the decline of population size and density of na-
multaneously allowing the development of new genetic tive species that can lead to Allee (1931, 1938) effects
combinations, which can facilitate the exploitation of that increase the likelihood of mating between native
underused or open niches (Seehausen et al. 2008). and invasive species as native animals have difficulty
This has been a hypothesized mechanism for the suc- finding mates (Courchamp et al. 1999). This has been
cessful and rapid spread of Cuban brown anole (Anolis commonly cited as a factor leading to increases in hy-
sagrei; Kolbe et al. 2004, 2007). bridization among many rare or endangered species
Another major source of hybridization is the inter- (Rhymer and Simberloff 1996; Allendorf et al. 2001;
breeding of domestic or game species with local native Wayne and Vila 2003; Grewal et al. 2004; Muñoz-
species. Examples include domestic dogs (Canis familia- Fuentes et al. 2010; Zamudio and Harrison 2010). One
ris) and wolves (C. lupus) in Europe (Randi 2008), dogs management approach that has been used to address
and Simien jackals (Canis simensis) in Ethiopia (Gotelli this problem for red wolves is selective removal of coy-
et al. 1994), domestic cats (Felis silvestris catus) and otes and hybrid offspring (Miller et al. 2003; Stoskopf
European wild cats (Felis sylvestris sylvestris; Beaumont et al. 2005).
et al. 2001; Randi 2008), domestic cats and the African A final challenge for conservation and management
wildcat (F. libyca; Wiseman et al. 2000), and domestic of hybridization is the unclear legal status of hybrids
ferrets (Mustela furo) and European polecat (M. puto- (Allendorf et al. 2001; Haig and Allendorf 2006; Ell-
rius; Davison et al. 1999). This has also been a major strand et al. 2010). Hybrids are not classified or recog-
concern for native fish populations that hybridize with nized by the International Union for Conservation of
introduced species. For example, extensive introduc- Nature, and only one country (South Africa) has en-
tions of rainbow trout (Oncorhynchus mykiss) and cut- dangered species legislation that specifically mentions
throat trout (O. clarki) in western US watersheds have hybrids. Historically, hybrids were not protected under
resulted in loss of diversity of native species because the US Endangered Species Act, but the policy “that hy-
of massive introgression (Allendorf and Leary 1988; brids between endangered species, subspecies, or popu-
Dowling and Childs 1992; Leary et al. 1993). lations cannot be protected” was withdrawn in 1990 in
Habitat fragmentation and alteration are also ma- response to growing scientific information on the com-
jor factors in the current increase in hybridization and plexities and benefits of hybridization (Allendorf et al.
introgression among wildlife populations. Barriers to 2001). While a new “intercross” policy was drafted by
hybridization are often maintained by habitat differ- the US Fish and Wildlife Service and National Marine
ences, thus land conversion that homogenizes habitats Fisheries Service in 1996, it was never formally ap-
and decreases landscape heterogeneity is believed to proved (Allendorf et al. 2001; Ellstrand et al. 2010).
elevate the probability of hybridization by relaxing Thus, there is no official policy on the conservation
ecological divergent selection among species and / or status or protection of hybrid individuals. This makes
by elevating the costs of mate choice in declining pop- management decisions regarding hybridization chal-
ulations (Allendorf et al. 2001; Hendry et al. 2006). lenging, particularly in cases where purposeful intro-
Habitat homogenization is believed to be a major fac- gression may be necessary to avoid inbreeding depres-
72 habitats in peril

sion (Tallmon et al. 2004) and / or augment population

sizes, such as occurred in the Florida panther (Pimm Box 6.3 Metrics of Genetic Diversity
et al. 2006) and Washington pygmy rabbit (Brachylagus Diversity metrics for DNA sequences:
idahoensis) recovery efforts (USFWS 2012).
• Haplotypic diversity (h): haploid version of
expected heterozygosity (mitochondrial
Using Genetic Methods to Assess and DNA only)
Predict the Impacts of Habitat Alterations • Nucleotide diversity (π): calculated as the
The relationship between habitat, genetic diversity, average number of differences in the nu-
gene flow, and genetic structure may be viewed from cleotides at each position along a sequence
two perspectives. First, because maintaining genetic (mitochondrial or nuclear sequence)
diversity is fundamental to maintaining population • Number of different haplotypes (variants)
viability, habitat loss, fragmentation, and degradation detected (mitochondrial DNA only)
must be avoided if possible. In this context, research- Diversity metrics for SNPs, microsatellites,
ers may seek to describe genetic structure and diver- and other molecular methods used to analyze
sity to identify populations that are isolated or have variation in nuclear DNA (see also review in
very low genetic diversity, and may attempt to quan- Hughes et al. 2008):
tify how habitat configuration and quality influences
those characteristics. Alternatively, because genetic • Expected heterozygosity (He ) is calculated
diversity and structure may reflect habitat loss, frag- based on the observed frequency of alleles,
mentation, or degradation, researchers may use genetic assuming Hardy-Weinberg equilibrium, and
studies primarily as a tool to learn about the influences can be corrected for sample size (Nei 1978)
of habitat on distribution, movement, dispersal, and • Observed heterozygosity (Ho ) is the propor-
demographic history, rather than being concerned with tion of individuals that are heterozygous at
precisely estimating genetic diversity or structure for a particular locus or averaged across a set
their own sake. of loci
• Proportion of polymorphic loci (i.e., loci
containing more than one allele)
Using Neutral and Adaptive Genetic • Allelic richness (Ar ) is the average number
Diversity to Evaluate Habitat of alleles per locus; Ar is quite sensitive to
Methods of assessing genetic diversity (box 6.2) have differences in sample size and therefore
proliferated rapidly because of technological advances should be corrected using subsampling or
in the last several decades, leading to many diversity rarefaction techniques (Leberg 2002)
metrics. Some methods, like allozymes (proteins used
for early studies of genetic variation), have been re-
placed by newer systems assessing DNA sequence or cautiously as an index of diversity across the genome
genotypes at mutation-prone locations (box 6.2). Ge- of a species. However, increasingly, new sequencing
netic diversity is typically measured at the population techniques make it possible to identify adaptive vari-
level (i.e., across individuals). At the individual level, ants and examine diversity at those loci. In some cases,
we typically consider only heterozygosity, which is habitat-related variation in phenotype can be linked
influenced by genetic diversity. Although it is beyond to particular alleles, raising the possibility of under-
the scope of this chapter to fully review statistics for standing fitness consequences of genetic diversity in
estimating genetic diversity at the population level, we different habitats. For instance, Linnen et al. (2013)
briefly summarize some of the more common choices identified multiple mutations that controlled coat
in box 6.3. color in deer mice (Peromyscus maniculatus) and varied
Genetic diversity metrics are often calculated across in nearby populations depending on the color of the
a relatively small number of neutral loci and then used substrate. Mice on light-colored substrates that lacked
 population genetics and wildlife habitat 73

alleles for light coat color were exposed to higher rates best metric varies depending on the study system and
of predation by avian predators, and alleles for light is still widely debated (Jost 2008; Heller and Siegis-
coat color showed evidence of strong selection in those mund 2009; Ryman and Leimar 2009). Many analyses
habitats. now rely on individual-based metrics of genetic differ-
Genetic diversity can reflect habitat quality and thus ence rather than population-based estimates of gene
offers a potential assessment tool, although fewer stud- flow or genetic structure. Such approaches are par-
ies have focused on this relationship compared to the ticularly useful for species that exist in relatively con-
well-known effects of habitat fragmentation on genetic tinuous habitat, where defining populations is prob-
diversity (Pitra et al. 2011). Pitra et al. (2011) found lematic. Individual-based approaches can also be used
that genetic diversity of great bustards (Otis tarda) was to detect more recent landscape changes than many
correlated with habitat quality and population size and population-based approaches (Landguth et al. 2010;
density. Epps et al. (2006) determined that He and al- but see Epps et al. 2013). Common metrics include in-
lelic richness of microsatellite loci in desert bighorn dividual relatedness (reviewed in Csillery et al. 2006)
sheep populations were correlated both with popu- and Rousset’s a (Rousset 2000). In other cases, the spa-
lation connectivity and with population elevation. As tial correlation of alleles is described across study areas
predicted by a previous analysis of population extinc- and areas of discontinuity can be matched up to land-
tions (Epps et al. 2004), higher elevation populations scape features (Manel et al. 2007). Manel et al. (2007)
appeared to have larger Ne and thus maintained more used this approach to determine that brown bears in
genetic diversity over the long term. However, we note Scandinavia, despite being relatively continuously dis-
that if dispersal is density dependent and gene flow is tributed, could be divided into three “populations” with
high, sink habitats (where mortality is high but offset genetic differences accumulating rapidly in the bound-
by immigration) could potentially have higher than ary regions.
expected genetic diversity in some cases because new Early analyses of genetic structure usually focused
individuals are constantly entering those areas. Porlier on large landscapes (even the entire range of a spe-
et al. (2009) examined individual genetic diversity of cies) and evaluated evidence for areas of low or absent
tree swallows (Tachycineta bicolor) across a gradient of gene flow among populations. For instance, Hafner
agricultural intensity and found that more genetically and Sullivan (1995) used allozymes to evaluate genetic
diverse individuals were found in poorer habitats with structure across the range of American pikas (Ochotona
more intensive agriculture. This counterintuitive result princeps) and deduce major phylogeographic divisions,
occurred because more diverse individuals arrived first establish a likely maximum distance over which dis-
and settled in the earliest available habitat, which oc- persal is possible among populations, and evaluate the
curred in areas with intensive agriculture. Thus, any influence of summer drought conditions on genetic
attempt to use genetic diversity as a metric of habitat diversity in at least a qualitative way. In recent years,
quality should carefully consider the characteristics of effects of habitat on genetic structure largely have been
that species and system. addressed through landscape genetic approaches (see
the following section), which allow spatially explicit
tests of how different habitats influence gene flow.
Effects of Habitat on Gene Flow and
Many of these studies are occurring at much finer
Genetic Structure
scales, enabled in part by newer molecular methods
The influence of habitat on gene flow and genetic that target large numbers of highly variable loci and
structure has been of great interest for many years. In- thus give good statistical power for resolving small ge-
terpreting such studies requires a basic understanding netic differences.
of metrics of gene flow and genetic structure, which
we review briefly in box 6.4. There are many metrics of
Landscape Genetics
genetic structure (i.e., differences in allele frequencies
among different groups of individuals) and gene flow The field of landscape genetics is a relatively new disci-
(rates of interbreeding among populations), but the pline that integrates population genetics and landscape
74 habitats in peril

Box 6.4 Estimating Genetic Structure and Gene Flow

Metrics of Genetic Structure of this approach (e.g., geographical information)

The most commonly reported metrics of genetic are implemented in various programs including
structure are FST ( Wright 1943) and its analogues STRUCTURE (Pritchard et al. 2000), GENELAND
such as GST ( Nei 1973) and ϕST. These statistics (Paetkau et al. 2004), and TESS (Chen et al. 2007).
have values ranging from 0, meaning identical
alleles are present at identical frequencies and Gene Flow
thus there is no structure between populations, Gene flow can be estimated from FST as Nm
to 1, meaning no shared alleles and a complete (effective number of migrants × migration rate)
structure. However, FST and GST have a number of using Wright’s island model (Wright 1931), where
problematic properties. For one, the maximum FST = 1 / (1+4Nm) for nuclear loci. However, that
value possible declines with increasing diversity transformation relies on unrealistic assumptions
of the loci considered (Hedrick 2005). A variety of equilibrium between genetic drift and migra-
of alternatives have been proposed that address tion and other simplifications, and so Nm esti-
this problem, including G'ST (Hedrick 2005) and mates should not be interpreted literally (Whit-
Jost’s D ( Jost 2008 ), but consensus about the lock and McCauley 1999). Alternative methods
appropriateness of these statistics remains elusive use coalescent theory (e.g., MIGRATE, Beerli
(e.g., Whitlock 2011). and Felsenstein 2001) or assignment tests (e.g.,
Rannala and Mountain 1997; Wilson and Ran-
Population Assignment Tests nala 2003; Faubet and Gaggiotti 2008) and rely
In this analytical framework, all individuals in on maximum-likelihood or Bayesian approaches.
the study area typically are combined into a These approaches use simulations to explore a
single analysis and sorted into varying numbers wide variety of possible parameter values and
of groups by genetic similarity and the overall determine the solution that best fits observed
performance of each model (i.e., number of patterns in genetic structure. These methods also
groups) evaluated to determine which model is rely on a variety of assumptions and typically
most likely. This framework lends itself well to require extensive exploration of datasets; thus,
descriptive or exploratory analysis rather than their use is not always straightforward, and many
hypothesis testing but does not require a priori researchers still use metrics of genetic structure
definition of populations, although results can be (FST, G'ST ) as an index of gene flow.
influenced by uneven or biased sampling. Variants

ecology with the goal of providing “information about riers and evaluate landscape resistance to gene flow,
the interaction between landscape features and micro- and evaluate current corridors and design future cor-
evolutionary processes, such as gene flow, genetic drift ridors (Holderegger and Wagner 2008; Balkenhol et al.
and selection” (Manel et al. 2003). Landscape genetic 2009a; Segelbacher et al. 2010; Storfer et al. 2010).
approaches are particularly valuable for examining Landscape genetic analyses of adaptive loci can be used
the underlying environmental factors that facilitate or to better understand how environmental factors such
hinder gene flow (Storfer et al. 2007; Holderegger and as temperature or precipitation affect adaptive genetic
Wagner 2008; Storfer et al. 2010) and have multiple variation and evolutionary response to landscape and
conservation and management applications (Segel- environmental change (Holderegger et al. 2006; Ma-
bacher et al. 2010). Landscape genetic approaches can nel et al. 2010). A wide range of statistical approaches
be used to evaluate or predict the impacts of landscape are applied in landscape genetics (Storfer et al. 2007;
change on genetic diversity and gene flow, detect bar- Balkenhol et al. 2009b), and much effort is focused on
 population genetics and wildlife habitat 75

developing and evaluating new methods (Faubet and and gene flow processes, they found that gene flow in
Gaggiotti 2008; Cushman and Landguth 2010; Epper- American marten was primarily driven by elevation
son et al. 2010; Legendre and Fortin 2010; Murphy (Wasserman et al. 2010). This finding allowed them
et al. 2010; Balkenhol and Landguth 2011). A detailed to derive a model of landscape resistance that com-
overview of the specific methods applied in landscape bined with simulation modeling (Landguth et al. 2010)
genetics is beyond the scope of this chapter. However, could be used to predict the effects of predicted climate
we present two case studies to highlight the use of land- change on population connectivity and genetic diver-
scape genetics in studying impacts of environmental sity of the American martens using five potential future
change on wildlife populations. temperature scenarios (Wasserman et al. 2012; Was-
Although the potential for loss of gene flow and ge- serman et al. 2013). They found that even moderate
netic diversity resulting from habitat fragmentation has warming scenarios resulted in very large reductions in
been well understood for many years, those changes population connectivity and genetic diversity. Wasser-
were initially difficult to detect in wild populations. man et al. (2013) were also able to identify important
However, advances in sampling using noninvasive ge- corridors in the current landscape that would remain
netic material and increased abilities of labs to process intact across the climate change scenarios and current
large numbers of samples have made it easier to sample corridors that would be lost as the climate warms.
wildlife populations systematically and over large
landscapes. Epps et al. (2005) used those techniques
Detecting Hybridization
to sample twenty-seven populations of desert bighorn
sheep (Ovis canadensis nelsoni) across a large portion Genetic analysis is the most definite method for de-
of their range in southeastern California and evaluate tecting hybridization among taxonomic groups. In the
how natural and anthropogenic habitat fragmentation 1980s and 1990s, mtDNA data were often used to de-
interacted to shape gene flow and genetic diversity, as tect hybrids (i.e., individuals with the morphology of
previously described. In a follow-up study, Epps et al. one species but the mtDNA of an alternative taxonomic
(2007) tested least-cost models of connectivity in a GIS group). However, this approach is limited in resolu-
by determining which estimates of effective distance tion because of the maternal inheritance of mtDNA
(distance mediated by habitat) were most strongly cor- (hybridization will not be detected when the father is
related with gene flow estimates among populations. from the other taxonomic group) and the need for mor-
In this manner, they determined that gene flow was phological data. More recently, nDNA microsatellite
facilitated by steeply sloped areas between core popu- analysis (and occasionally AFLP analysis; see box 6.2)
lations, identified the maximum effective distance at has been the molecular method of choice for detect-
which gene flow occurred, and then used those models ing hybridization in wildlife populations. When iden-
to evaluate how human-made barriers and population tifying hybrids, it is possible to use diagnostic species-
translocations affected connectivity among different specific alleles (Fitzpatrick and Shaffer 2004; Schwartz
populations of desert bighorn sheep. et al. 2004), but increasingly, hybrids are detected by
Predicting the impact of climate change on wildlife applying complex maximum-likelihood and Bayesian
habitat and populations is one of the most important approaches (box 6.4, Pritchard et al. 2000; Ander-
challenges for future wildlife conservation (Heller and son and Thompson 2002; Corander et al. 2003; Cor-
Zavaleta 2009). In a study of American marten popula- ander and Marttinen 2006; Corrander et al. 2006;
tions of the northern Rocky Mountains in the United Randi 2008) to datasets of 10–100s of microsatellite
States, Wasserman et al. (2010, 2012) have provided loci, AFLPs, or SNPs (box 6.2). Detecting hybridization
one of the best examples of how landscape genetic using genetic approaches is most challenging when the
methods can be applied to predict and evaluate the hybridizing groups are closely related and when back-
impact of climate change on wildlife populations. Us- crossing to parental groups has occurred (Vaha and
ing nDNA microsatellite loci and an individual-based Primmer 2006; Bohling and Waits 2011). However,
causal modeling approach (Cushman et al. 2008) to genetic analyses have added greatly to our ability to
explore relationships between landscape patterns detect and understand hybridization in wildlife popu-
76 habitats in peril

lations, and many of the challenges can be overcome locations. They generated DNA sequence data for ma-
by increasing the number of nDNA loci analyzed per ternally inherited mtDNA, three neutral loci on the
individual. In the following discussion, we review two biparentally inherited X chromosome and one neutral
case studies that illustrate the use of genetic methods locus on the paternally inherited Y chromosome to
to detect the impact of habitat modification on hybrid- evaluate the frequency and directionality of hybridiza-
ization in wildlife populations. tion between forest and savannah elephants. Their re-
One good example of habitat-dependent hybridiza- search indicated that hybridization occurs along forest /
tion occurs between the endangered California tiger savannah ecotones and that forest elephants declined
salamander (Ambystoma californiense) and the non- in regions that lost forest cover during the Holocene
native barred tiger salamander (Ambystoma tigrinum due to introgressive hybridization with savannah el-
mavortium) that is introduced by bait dealers and ephants. Additionally, there is evidence that ongoing
fisherman (Riley et al. 2003). Before the introduction deforestation and landscape conversion may foster ge-
of the barred tiger salamander fifty to sixty years ago, netic replacement of forest elephants, increasing the
the two species had been geographically separated for risk of extinction of this species (Roca et al. 2005).
five million years (Shaffer et al. 2004). Using mtDNA
and nDNA loci, researchers evaluated hybridization of
Conclusions and Future Directions
these species in the Salinas Valley of California where
the landscape has been modified for cattle farming As summarized previously, the environmental changes
and agriculture creating perennial or ephemeral cattle that destroy, degrade, and alter habitats have multiple
ponds, both of which have longer hydroperiods than impacts on wildlife at the genetic level. In addition,
the highly ephemeral vernal pools that were histori- molecular genetic techniques greatly enhance our abil-
cally the natural breeding habitat of A. californiense (Ri- ity to study, understand, and predict the impact of such
ley et al. 2003; Fitzpatrick and Shaffer 2004, 2007a, b). changes on wild populations. As molecular techniques
Overall, they found evidence of hybrid swarms in most and analytical methods continue to rapidly progress,
of the Salinas Valley and demonstrated that hybridiza- we anticipate an increasing role for genetic analysis in
tion and introgression rates were linked to introduction unraveling those relationships and in providing solu-
history and lowest in natural vernal pools, intermedi- tions for conservation and management. For example,
ate in ephemeral cattle ponds, and highest in perennial our increased ability to survey greater numbers of neu-
cattle ponds. They hypothesize that the habitat charac- tral loci will lead to greater precision and accuracy in
teristics of the breeding ponds can promote some form estimates of Ne, gene flow, hybridization, and kin rela-
of environment-dependent reproductive isolation, ei- tionships (Allendorf et al. 2010).
ther through individual mate choice within the breed- Genomic approaches that survey adaptive ge-
ing pond or reduced population size of the introduced netic variation will enable us to better understand
species (Fitzpatrick and Shaffer 2004, 2007b). They the adaptive response to the diversity of natural and
also documented increased fitness of hybrid individuals human-altered habitats and provide better guidance
highlighting the increasing challenge of maintaining on conservation priorities and the appropriate sources
and protecting the genetically pure populations of the for population augmentation, reintroductions, and as-
native endangered California tiger salamander (Fitz- sisted migration (Ouborg et al. 2010; Funk et al. 2012).
patrick and Shaffer 2007a). The use of genomic approaches will also increase our
Studies of African elephants provide another good ability to understand the mechanistic basis of inbreed-
example of natural and human-caused habitat change ing depression and the relationship between genetic
that can lead to increased hybridization. African for- diversity and fitness at the individual and population
est and savannah elephants are distinct species sepa- levels (Allendorf et al. 2010; Ouborg et al. 2010). Ulti-
rated by a hybrid zone (Roca et al. 2001). To evaluate mately, we are interested in better understanding and
historical and current hybridization levels, Roca et al. predicting how environmental change will affect neu-
(2005) collected genetic data from over seven hundred tral and adaptive diversity.
savannah and forest elephants in twenty-one different The rapidly growing field of landscape genetics has
 population genetics and wildlife habitat 77

emerged to help address these questions and will ben- Balkenhol, N., F. Gugerli, S. A. Cushman, L. P. Waits, A. Cou-
efit from the larger neutral and adaptive genetic da- lon, J. W. Arntzen, R. Holderegger, and H. H. Wagner.
tasets generated from new genomics techniques and 2009. “Identifying Future Research Needs in Landscape
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 7 Habitat Fragmentation
and Corridors
K. Shawn Smallwood

S ince its conceptualization, habitat fragmentation

has factored prominently in academic discussions
on threats to biodiversity and species’ conservation.
converted vast additional areas for human use, it is time
to ask whether the theory has advanced sufficiently to
resolve the details that matter. More importantly, it is
Habitat fragmentation has also been incorporated time to examine whether the concepts of habitat frag-
into environmental law; e.g., one of the standards of mentation and corridors have mattered where these
the California Environmental Quality Act is to assess a interrelated concepts needed to be implemented.
project’s potential interference with the movement of My objectives for this chapter are first to compare
fish and wildlife. A principal countermeasure to habi- definitions of habitat fragmentation and corridors and
tat fragmentation—maintaining and improving habitat synthesize these interrelated concepts. Next, I review
connectivity—has also factored into many academic environmental documents to reveal how practitioners
and legal discussions. Natural or constructed corri- perceive these concepts and to what degrees they apply
dors have often been promoted to mitigate the effects them in impact analyses and conservation planning.
of habitat fragmentation by maintaining or improv- Finally, I suggest how these concepts could be more
ing animal movement and gene flow between habitat consistently understood and effectively applied.
patches. Given its prominence in conservation biology
and environmental law, one would expect that the term
Habitat Fragmentation Defined
habitat fragmentation is well understood and based on a
strong theoretical foundation. Wilcox and Murphy (1985) described habitat fragmen-
However, given the widespread confusion over the tation as habitat loss and insularization. They described
meaning of the term habitat (Hall et al. 1997; Guth- the risk of fragmentation as (1) destruction, reduction,
ery and Strickland, this volume), which is a concep- or subdivision of demographic units; (2) loss of po-
tual foundation of habitat fragmentation, it might be tential sources of immigrants; and (3) impedance of
worth examining how well habitat fragmentation and immigration caused by conversion of habitat between
corridors are understood. At its inception, some degree habitat patches. Saunders et al. (1991) described it as
of vagueness around the concept was reasonable. Wil- clearing of natural vegetation resulting in isolated,
cox and Murphy (1985) argued, “That current theory remnant patches of vegetation. Yahner (1996) recom-
is inadequate for resolving many of the details should mended that habitat fragmentation be considered a
not detract from what is obvious and accepted by most process of diminishing size and increasing isolation
ecologists: habitat fragmentation is the most serious of habitat fragments, where the result is habitat loss.
threat to biological diversity and is the primary cause of Villard et al. (1999) described habitat fragmentation
the present extinction crisis.” Nearly three decades af- as a process in which a focal habitat type is partially
ter Wilcox and Murphy (1985), and after humans have or completely removed, resulting in an alternate con-
 habitat fragmentation and corridors 85

figuration and reduced population persistence on the cies to competitors, predators, and parasites that were
landscape. Bender et al. (1998) defined habitat frag- more prevalent along the patch edges (Laurence and
mentation as an event that creates a greater number of Yensen 1991; McCollin 1993; Porneluzi et al. 1993),
habitat patches that are smaller in size than the origi- even leading to pest outbreaks that can further de-
nal contiguous habitat. Francis (this volume) similarly grade or remove the vegetation in habitat fragments
described it as the spatial arrangement and shape of (Roland 1993).
remaining habitat patches remaining after habitat loss. Ultimately, what separates habitat fragmentation
Bender et al. went on to define it as an event that pro- from simple habitat loss is the disproportionate reduc-
duces an even greater population decline than would tion in numerical capacity of the remaining habitat of
occur due to simple habitat loss. the same net area (also see Francis, this volume). In
Karr (1994) described fragmentation as a disrup- other words, habitat reduced to a contiguous area of
tion of the linkages among patches that exchange one hundred ha might support one hundred individuals
ecologically important resources. Wilcox et al. (2002) of a certain species, but one hundred ha of habitat frag-
described it as the loss of contiguity of accessible land- mented by impassable barriers (e.g., irrigation ditches,
scape from the perspective of the organism or some a freeway, residential or commercial development) and
other ecologically important element, where landscape contaminated by physical and biological pollutants
context, the spatial extent of the organisms’ ecologi- might support sixty individuals, or even zero individu-
cal interactions, and its demographic organization are als. Hypothetically, the protection or creation of cor-
critical factors. ridors can maintain or restore some of the numerical
Saunders et al. (1991) argued that habitat fragmen- capacity that would otherwise be lost to fragmentation.
tation not only causes biogeographic changes, i.e., Applied to the previous example, the creation of one or
increasing isolation of habitat patches, but also physi- more movement corridors might restore the capacity
cal changes within the patches due to alterations in of the one hundred ha from sixty individuals to ninety
microclimate. Similarly, Wiens (1997) argued that the individuals. Fragmentation is thought to disproportion-
land conversions causing habitat fragmentation often ately reduce numerical capacity, whereas corridors are
sharpen habitat patch boundaries, alter connectivity, thought to counteract this effect.
and shift the cost-benefit contours on the landscape. Why the disproportionate reduction in fragmented
In other words, land conversions often inject into the habitat? Genetic isolation is often cited, and so are
remaining habitat fragments invasive species (Alberts other processes from island biogeography theory. An-
et al. 1993), light pollution (Rich and Longcore 2006), other factor is the suite of altered physical and biologi-
noise pollution, atmospheric pollution, water pollu- cal conditions caused by greater edge-to-interior ratios
tion (Longcore et al. 1993), soil erosion, and added and by the activities in the emergent nonhabitat ma-
mortality factors such as automobile collisions, line trix (Saunders et al. 1991; Bolger et al. 1997; Wiens
collisions, electrocutions, poaching, and animal dam- 1997). But a third factor is social organization, which is
age control. Furthermore, the net length of boundary a much stronger force often given credit in discussions
increases between habitat fragments and the nonhabi- of habitat. In an extreme example, our one hundred
tat matrix, thereby increasing the number and variety ha of fragmented habitat might consist of one hundred
of ways in which the land conversions can degrade one-ha patches of isolated habitat, each patch capable
the remaining habitat fragments. For example, large of supporting one individual of a particular species
carnivores, due to their greater ranging behavior, will on the basis of resources but incapable of supporting
more often experience conflict with humans when the any individuals beyond a single generation due to in-
habitat fragments they occupy include greater edge-to- sufficient space for breeding pairs, offspring, or larger
interior ratios (Woodroffe and Ginsberg 1998). Loss of demographic units. The spatial extent of habitat must
the large predators can result in mesopredator release, comport with the social organization and associated
thereby putting more predation pressure on other wild- behaviors of the species.
life species residing in the fragments (Zembal 1993). Long-term ecological relationships have estab-
Habitat fragmentation can expose patch-interior spe- lished trajectories of how individuals of each species
86 habitats in peril

interface with their habitats. For example, some spe- confusion because the original definition of the term
cies have developed territorial behaviors to stabilize habitat applies to single species, but habitat is increas-
numerical responses to resource variability and there- ingly used to characterize vegetation complexes asso-
fore cannot tolerate crowding that might initially be ciated with multiple species, i.e., habitat types. When
forced by habitat fragmentation. Many species rou- the discussion refers to wildlife habitat or habitat types,
tinely shift locations of high activity, i.e., high density, the basis of habitat fragmentation is typically a map of
either generationally or multigenerationally. Taylor vegetation cover types that are convenient for mapping
and Taylor (1979) hypothesized that this spatial shift- but probably ill-suited for analysis of habitat fragmenta-
ing of abundance enables species to rest local food re- tion. Measuring species richness in response to habitat
sources or to escape predator or parasite loads. They fragmentation implies the same concept of habitat type
also suggested that dispersed young might naturally as the basis of the analysis. In reality, a map of a species’
congregate in new locations while the natal colonies habitat, i.e., that part of the environment where the
senesce. Increasing barriers to movement and reducing species lives, would rarely match a mapped habitat type
the effective area of habitat patches, whether through intended to apply to multiple species. That is, the non-
land conversions or patch contamination, will act upon habitat matrix derived from such a map probably rarely
the species’ behavior trajectories, which often involve applies to a given species. This source of confusion in
much larger habitat areas than commonly studied by discussions of habitat fragmentation will be discussed
biologists (Smallwood 1999). in the following discussion.
Consider that the mass density of populations of
species of mammalian Carnivora averages nine kg / km2
Corridors Defined
when scaled to the spatial area encompassing the popu-
lation, referred to as the threshold area (Smallwood Beier and Loe (1992) defined the wildlife movement
2001a). This scaling of the spatial extent of a popu- corridor as a linear habitat whose primary function is to
lation to achieve a common mass density suggests an connect habitat patches. Beier and Noss (1998) defined
ecological allometry that is more sensitive to the spatial corridor as linear habitat connecting habitat patches
extent at which the species’ population operates than it of the same type and separated by a dissimilar matrix.
is to density or body mass. Indeed, threshold area was Beier and Noss specifically excluded linear reaches of
proportional to average female brain mass among spe- vegetation that were dissimilar to the habitat patches at
cies of Carnivora, and it was more responsive to female issue; e.g., they excluded riparian corridors that might
brain mass than to body mass (Smallwood 2001a). Spe- stretch between two patches of chaparral. In their
cies of Carnivora aggregate in similar numbers, but meta-analysis of corridor effectiveness, Gilbert-Norton
these numbers spread across larger spatial extents in et al. (2010) defined corridor “as a narrow, linear (or
proportion to increasing average brain mass. I hypothe- near-linear) piece of habitat that connects two larger
size that the habitat areas intervening populations also patches of habitat that are surrounded by a nonhabitat
scale with female brain mass. I also hypothesize that matrix.” They also distinguished between natural and
the habitat areas at which the populations of other ani- constructed corridors. Simberloff and Cox (1987) char-
mal species operate also scale with brain mass. Female acterized what they termed conservation corridors as
brain mass as the axis of similitude would implicate pa- constructed corridors intended to connect habitat re-
rental care and other life-history attributes, along with serves to facilitate immigration and genetic exchange.
basal metabolic rate and sensory perception. Species Forman and Godron (1981) defined another two types
are probably much more sensitive to habitat fragmenta- of corridor as (1) strip corridor: a protected strip
tion than can be measured by extrapolations of average of landscape that is wide enough to include interior
density to the remaining areas of habitat fragments. conditions typical of the interior conditions of habitat
Discussion of habitat fragmentation often centers patches being connected by the strip, and (2) line corri-
on individual species, but also often it encompasses dor: a protected strip of landscape that is too narrow to
multiple species. This dual use of the term causes include interior conditions typical of the interior con-
 habitat fragmentation and corridors 87

ditions of habitat patches being connected by the strip have been thought to be used by animals as convenient
but instead includes edge conditions that are typical of guides to migration, dispersal, home range patrol, and
the edge conditions of the connected habitat patches. other types of long-distance movement. However, biol-
There are many other definitions of corridor in the lit- ogists often assume that this relationship exists, rather
erature, including movement corridor, habitat corridor, than actually measuring it. In fact, animals move across
dispersal corridor, and landscape linkages, so it is clear all types of landscape features and vegetation patterns,
that the corridor concept varies among investigators not just linear features. The capacity for animals and
according to their application. plants to move across the “matrix” landscape might be
Beier and Noss (1998) concluded that corridors just as important as the capacity for movement along
enhance population viability in connected habitat corridors.
patches, but the papers they reviewed generally did Conservation corridors also imply concentrated
not support their conclusion due to pseudoreplication movement of one or more species but only in the con-
and confounding caused by inadequate experimental text of navigating between habitat patches in a chang-
design. In response to criticism of the evidence sup- ing environment. Whether protected or constructed,
porting the claim that habitat corridors improve popu- conservation corridors are intended to enable contin-
lation viability (Simberloff et al. 1992), Beier and Noss ued or improved movement of wildlife between habitat
asserted that an experimental design was unnecessary fragments. However, the effectiveness of conservation
for concluding that corridors are effective. corridors is rarely tested, and many so-called corridors
may themselves be nothing more than remnant frag-
ments of habitat. Furthermore, many conservation cor-
Synthesis of Habitat Fragmentation
ridors have been established to suit the convenience
and Corridors
of the planners. I have rarely seen evidence that ani-
The concepts of habitat fragmentation and corridors mal movement patterns were measured prior to the
are interrelated. These terms are part of the same establishment of conservation corridors. The common
discussion about a particular type of anthropogenic assumption appears to have been that the animals sur-
impact on wildlife species and whether and to what viving habitat fragmentation will use the corridors, per-
degree this type of impact can be moderated by the pro- haps because they have no other choice. Nevertheless,
tection or creation of habitat or movement corridors. the purpose of conservation corridors has been to offset
Habitat fragmentation can certainly happen naturally the adverse impacts of habitat fragmentation. The only
(Morrison et al. 1998), but naturally occurring habi- overlap in the scientific dialogue between naturally oc-
tat fragmentation has been rarely discussed and is of curring corridors and conservation corridors is in the
relatively low interest. Much greater focus has been on protection of the former as the latter as habitat frag-
anthropogenic habitat fragmentation and whether and mentation proceeds.
how to mitigate its adverse effects via corridor protec-
tion or construction. Discussions about corridors have
Habitat Fragmentation and Corridors
been more confusing on the distinction of whether the
in Practice
context is about corridors as naturally occurring land-
scape or habitat features or as a conservation asset that The concepts of habitat fragmentation and corridors
needs to be protected or constructed to counter the ef- have been favorite topics of debate in the scientific lit-
fects of habitat fragmentation. erature, having inspired thousands of papers and books
Corridor implies concentrated movement of one or with one or both of these concepts being addressed.
more species, or disproportionate use of a linear por- Scientists debate definitions related to these concepts,
tion of a landscape. Naturally occurring corridors are and they debate how to measure habitat fragmentation
typically characterized as linear features of the land- and how to identify corridors from within a landscape
scape, such as stream basins and the vegetation that matrix. While these scientific debates have continued
grows along streams. Linear features of the landscape for nearly three decades, other human actions have
88 habitats in peril

transformed large tracts of wildlife habitat for human pede the use of native wildlife nursery site.” This lan-
purposes. It is at the business end where the concepts guage is the closest that CEQA comes to recognizing
of habitat fragmentation and corridors have been in habitat fragmentation as an impact and corridors as
most urgent need of general understanding and imple- the solution. Consultants and resource agency biolo-
mentation. gists often cite this CEQA standard before discussing
It is in environmental reviews, management plans, wildlife movement corridors in the context of project
environmental laws, and, ultimately, actions where the impacts or mitigation. Because environmental reviews
concepts of habitat fragmentation and corridors matter performed in California are supposed to address the
most (Salwasser 1990). The practitioners who need to project’s impacts on wildlife movement, habitat frag-
understand these concepts include environmental con- mentation and corridors should be expected to be dis-
sultants, biologists working for regulatory and natural cussed in environmental review.
resource agencies (i.e., permitting agencies), members I assessed the environmental review documents
of the public who participate with project planning, en- of fifty-seven proposed projects involving 83,413 ha
vironmental lawyers, and decision-makers including (app. 7.A). Wildlife movement corridors were men-
city councils, county supervisors, state commissions, tioned in thirty-four reviews (60%) and analyzed in
and judges. Unless the concepts of habitat fragmen- zero reviews. Habitat fragmentation was mentioned in
tation and corridors are understood clearly by practi- twenty-six reviews (46%) and analyzed in two reviews
tioners, and unless these concepts influence manage- (3.5%). Where the concepts of habitat fragmentation
ment and conservation decisions in meaningful ways, and corridors matter most—where they matter to en-
then, to be frank, the scientific discussion of them has vironmental decision-making—about half of the envi-
been of little value. ronmental reviews made no mention of them.
To ascertain the effectiveness of the concepts of In one of the two reviews that included an analysis of
habitat fragmentation and corridors, I examined envi- habitat fragmentation (app. 7.B, 5), the level of project
ronmental review documents with which I participated impact was calculated in acres and was based on the
as an expert witness over the past twenty years (ap- starting habitat value and ending habitat value. Habitat
pendix 7.A). I examined how the concepts of habitat values were 100% for parcels zoned ≥40 acres, 75% for
fragmentation and corridors were portrayed and im- parcels zoned ≥20 acres, and 25% for parcels zoned
plemented in the face of real proposed projects that, ≥10 acres. However, because >90% of the project area
if approved, could potentially affect wildlife through was composed of parcels zoned ≥40 acres, the project
the process of habitat fragmentation and destruction or area was concluded to have 100% habitat value. In the
protection of corridors. Some of these projects involved other review including a quantitative analysis (app. 7.B,
very small areas, but even a project on a small habitat 36), habitat fragmentation was analyzed as the amount
area can potentially sever habitat linkages, thereby con- of habitat area that will be lost as a percentage of the
tributing to habitat fragmentation. Many of these proj- cumulative impacts in the county.
ects were constructed, some were not, and others are An analysis that is sufficient for the California Envi-
waiting for permits or construction starting dates, but ronmental Quality Act or the National Environmental
the actual outcomes of these proposed projects were Policy Act is not the same type or level of analysis that is
irrelevant to my examination of how the concepts of practiced by research scientists, so even the two crude
habitat fragmentation and corridors were treated in the analyses that I encountered in environmental review
environmental reviews. documents would not have been regarded as scientific
Under the California Environmental Quality Act by the research community. Essentially, there has been
(CEQA), which is the law that I most often work under no scientific analysis performed and no scientific pre-
as an expert witness, a project would have a significant dictions made of the effects of habitat fragmentation
impact on the environment if it were to “interfere sub- or corridor loss or protection resulting from any of the
stantially with the movement of any native resident or environmental reviews with which I have been profes-
migratory fish or wildlife species or with established sionally involved. Consultants and agency biologists
native resident or migratory wildlife corridors, or im- preparing the environmental reviews for about half of
 habitat fragmentation and corridors 89

the projects have expressed familiarity with the terms can extend beyond the boundaries of an area proposed
habitat fragmentation and corridors, but neither knew for development” (app. 7.B, 13). This review went on to
how to perform impact assessments relevant to these describe how fragmentation of a creek could adversely
concepts, or chose not to do so. affect downstream wetlands by altering hydration pe-
riods. Thus, some environmental reviews recognized
the potential for net greater impacts caused by move-
Descriptions of Habitat Fragmentation
ment barriers and habitat fragmentation as compared
Environmental reviews defined or described habitat to simple habitat loss.
fragmentation in various ways, expressing some over-
lap as well as considerable divergence in understand-
Descriptions of Corridors
ing of the concept. One review defined habitat frag-
mentation as habitat loss resulting in displacement of Environmental reviews defined or described corri-
individuals from the developed area (app. 7.B, 1). This dors more often than they defined habitat fragmenta-
review equated habitat fragmentation with habitat loss tion. One review explained that corridors link blocks
characterized by displacement rather than by any over- of habitat, where the habitat in corridors resembles
all population reduction. In another review, this notion habitat preferred by the target species (app. 7.B, 33),
of displacement was rolled into one of crowding within and another defined wildlife corridors as “pathways or
fragmented patches of Swainson’s hawk (Buteo swain- habitat linkages that connect discrete areas of natural
soni) foraging habitat, “increasingly fragmented with open space otherwise separated or fragmented by to-
more intensive uses (agricultural / residential or urban pography and changes in vegetation.” (app. 7.B, 31).
development) on smaller minimum parcel sizes” (app. Corridor definitions sometimes appeared tailored to
7.B, 5). In this version of habitat fragmentation, the af- serve the strategy of the impacts assessment or miti-
fected species suffers the inconvenience of crowding gation plan. Kern County (app. 7.B, 2, 4.3–20) wrote,
but not necessarily any overall numerical reduction. “Wildlife movement corridors, also referred to as dis-
In contrast to the notion of simple displacement, persal corridors or landscape linkages, are generally
or at worst crowding due to habitat fragmentation, an- defined as linear features along which animals can
other review explained that small fragments of habitat travel from one habitat or resource area to another . . .
can only support small populations and are more vul- drainages, ridgelines, and other natural and built linear
nerable to extinction (app. 7.B, 6). Alluding to habitat features and barriers often serve as areas that wildlife
fragmentation, an environmental review document routinely use to access essential natural resources. It is
included the statement, “The ability for wildlife to assumed that wildlife species would use such features
freely move about an area and not become isolated is for movement if they occurred within the survey area.”
considered connectivity and is important to allow dis- This definition implies that wildlife will use the land
persal of a species to maintain and exchange genetic left undeveloped as a corridor, thereby mitigating the
characteristics, forage (food and water), and escape project’s impacts.
from predation” (app. 7.B, 8). Imposing barriers to In the review of a project seeking to mitigate its im-
movement qualified as habitat fragmentation in one pacts by leaving undeveloped land as corridors, PBS&J
review (app. 7.B, 41), and in two other reviews barriers (app. 7.B, 4, 5.4–8) wrote,
were implied: “Fragmentation of open space areas by
urbanization creates ‘islands’ of wildlife habitat that Terms such as habitat corridors, linkages, crossings,
are more or less isolated from each other” (app. 7.B, 4), and travel routes are used to describe physical connec-
and “Habitat fragmentation involves the potential for tions that allow wildlife to move between patches of
dividing sensitive habitat and thereby lessening its suitable habitat in undisturbed landscapes as well as
biological value” (app. 7.B, 10). In another review, the environments fragmented by urban development . . .
public learned that “habitat fragmentation can result Wildlife corridors are usually bounded by urban land
when development occurs within larger regions of areas or other areas unsuitable for wildlife. The corridor
natural habitat. The effects of habitat fragmentation generally contains suitable cover, food, and / or water to
90 habitats in peril

support species and facilitate movement while in the wildlife that provide shelter and sufficient food sup-
corridor. Wildlife corridors link areas of suitable habitat plies to support wildlife species during migration.
that are otherwise separated by areas of non-suitable Movement corridors generally consist of native or
habitat such as rugged terrain, changes in vegetation, or undeveloped matrix habitats (e.g., greenbelts, parks,
human disturbance. Wildlife corridors are essential to or other open spaces) that span contiguous acres of
the regional ecology of a species because they provide unfragmented habitat. Wildlife movement corridors
avenues of genetic exchange and allow animals to ac- are an important element of resident species home
cess alternative territories as dictated by fluctuating ranges.” Pacific Municipal Consultants (app. 7.B, 16,
population densities . . . Wildlife corridors are typically 4.4–20) defined wildlife movement corridors as “tra-
relatively small, linear habitats that connect two or more ditional routes used by wildlife to travel within their
habitat patches that would otherwise be fragmented or home range. Movement corridors typically provide
isolated from one another. wildlife with undisturbed cover and foraging habitat
and are generally composed of several trails in contigu-
This last part of the definition appeared to rationalize ous spans of forested, riparian, riverine, and woodland
leaving only a narrow strip of land as the corridor. habitats. The width of movement corridors varies de-
The review of another large residential develop- pending on the topography. Movement corridors are an
ment—for which the principal mitigation measure essential element of home ranges of a wide variety of
was the protection of land to be left undeveloped wildlife including black bear (Ursus americanus), grey
as movement corridors (app. 7.B, 9)—defined wildlife fox (Urocyon cinereoargenteus), mountain lion (Felis
movement corridors as the “gentlest topography and concolor), and other migratory wildlife.”
more open habitat.” A subsequent review document for According to EGI (app. 7.B, 34), “Wildlife move-
this project identified a wildlife corridor as a drainage ment corridors and habitat linkages are areas that con-
or riparian vegetation in a canyon, which traverses no nect suitable wildlife habitat areas in a region otherwise
other topographic features and will not be surrounded fragmented by rugged terrain, changes in vegetation, or
by development in the future. Similarly, the consul- human disturbance. Corridors are generally local path-
tants who prepared the planning for a multispecies ways connecting short distances usually covering one
habitat conservation plan across a large portion of San or two main types of vegetation communities. Linkages
Diego County (app. 7.B, 33) defined a wildlife corri- are landscape level connections between very large
dor as “a route used by one or more species to move core areas and generally span several thousand feet
between two areas of habitat. A corridor can be defined and cover multiple habitat types. Natural features such
by topographical features such as ridges or valleys, as canyon drainages, ridgelines, or areas with vegeta-
habitat types such as bands of riparian vegetation, ar- tion cover provide corridors and linkages for wildlife
eas of natural open space passing between two man- travel. The habitat connectivity provided by corridors
made constraints, or even game trails used by many and linkages is important in providing access to mates,
generations of animals” (page 6-1). The review defined food, and water, allowing the dispersal of individuals
two types of corridor—local and regional (page 6-1): away from high-density areas, and facilitating the ex-
“Regional corridors connect open space in a region change of genetic traits between populations.”
and allow activities such as dispersal of young, genetic According to ECORP (app. 7.B. 36), corridors are
transfer between subpopulations, and seasonal migra- where surrounding habitat concentrates wildlife move-
tion. Local corridors are routes within a habitat used ment, or which link large areas of undeveloped open
regularly by an animal to commute between resources space. Other reviews defined wildlife corridors as “ar-
such as denning sites, water sources, and hunting or eas that connect suitable wildlife habitat areas within
foraging areas.” a region, especially where species are known to mi-
According to another review (app. 7.B, 10), “Wild- grate” (app. 7.B, 37), and “where resident and migra-
life corridors are areas of habitat used by wildlife for tory animals freely move, generally within or between
seasonal or daily migration.” In another (app. 7.B, 14), preferred habitat areas” (app. 7.B, 30).
“Wildlife corridors are routes frequently utilized by Environmental reviews sometimes identified par-
 habitat fragmentation and corridors 91

ticular parts of the project area as wildlife movement might exist nonscientific reasons for this poor transla-
corridors, which were then claimed to be unaffected by tion of scientific concepts into conservation practice,
the project or proposed to be protected or enhanced as but even so, it should be more difficult to abuse con-
mitigation (app. 7.B, 2, 9, 18, 25, 41). Corridor desig- cepts that are clearly and consistently understood. If
nations were arbitrary in these reviews and may have these concepts are going to matter in the future, if they
been influenced by convenience. More often, environ- are going to conserve wildlife in the face of massive
mental reviews claimed that the project site did not anthropogenic land use changes, then barriers must
occur in an established or major wildlife movement be overcome to effective measurement and consistent
corridor (app. 7.B, 2, 3, 5, 7, 11, 12, 14, 15, 17, 19, 20, understanding.
26, 30, 34, 36, 46). The premise implied by this strategy
was that there exists a registry or inventory of wildlife
Barriers to Measuring Habitat
movement corridors that was consulted and turned up
Fragmentation and Corridors
a negative result. Even if there was an inventory of cor-
The Dependent Variable
ridors, the inventory’s scientific credibility would hinge
on species-specificity and empirical foundation. The effects of habitat fragmentation and corridors
must be measureable if they are going to translate into
meaningful theory or sound conservation planning
Summary of Habitat Fragmentation and
and decision-making. It is not enough for conserva-
Corridors in Practice
tion biologists to proclaim that “we know that habitat
Environmental review documents rarely cited peer- fragmentation is happening and corridors are effective
reviewed papers on the topics of habitat fragmentation at rectifying the effects of fragmentation.” The effects
and corridors. Given their varied characterizations of need to be quantified and relevant hypotheses tested at
habitat fragmentation and corridors, it was apparent appropriate spatial and temporal scales. It is therefore
that practitioners often only vaguely understood these necessary to identify what it is that fragmentation and
concepts. The concept of habitat fragmentation was corridors are thought to be affecting.
the most poorly understood of the two concepts, hav- Species richness and species diversity were early
ing been omitted from discussion in most reviews and favorites as dependent variables responding to habi-
defined erroneously in most reviews that addressed tat fragmentation, due to their frequent use in island
the topic. In some cases, the practitioners may have biogeography theory and the often- claimed likeness
crafted definitions of these terms to minimize esti- between habitat fragments and islands. Species rich-
mates of project impacts or to minimize the cost of ness remains in use, as Gilbert-Norton et al. (2010)
mitigation. Among the environmental documents I used it as an indirect measure in their meta-analysis of
reviewed, none of the alleged existing corridors were corridor effectiveness. However, the comparability of
measured on any variables for corridor function. None terrestrial habitat fragments to islands is questionable
of the corridors proposed for protection as project miti- because the fragments occur in very different ecological
gation were accompanied by commitments to test for contexts than do islands. Furthermore, species richness
corridor functionality following project development. and species diversity are community measures, which,
Instead of testing for corridor functionality, those who by a proper definition of habitat (Hall et al. 1997), qual-
performed environmental reviews of the fifty-seven ify as poor measures of habitat fragmentation. In other
projects addressed herein relied on hopeful specula- words, because habitat is a species-specific concept, its
tion that land designated as corridors would actually fragmentation is best measured for the species to which
function as corridors. the habitat is associated. Species richness and species
Based on my review of environmental assessments diversity might be more suitable dependent variables
on the impacts and mitigation planning of fifty-seven in analyses of landscape or ecosystem fragmentation.
projects, I conclude that the concepts of habitat frag- Population persistence would probably generate
mentation and corridors have too often failed to mat- the greatest agreement among ecologists and wildlife
ter where they most needed to be implemented. There biologists as the ultimate expression of the effects of
92 habitats in peril

fragmentation and corridors, although there might

Measuring Habitat
not be agreement on what qualifies as a population
or what time period is sufficient for measuring persis- The concept of habitat fragmentation has been poorly
tence. Abundance could be measured as a variable that understood by the community in greatest need of
indicates the population and could be expressed as the understanding it. This poor understanding might
number of individuals, breeding pairs, or colonies or be exacerbated by a generally poor understanding of
populations. The health of the population can also be habitat. The habitat concept itself has lacked consis-
measured, such as by genetic variation, productivity, tent definition (Guthery and Strickland, this volume),
and survival, but these variables are difficult to measure leading to ample confusion among practitioners as well
and are probably more proximal expressions of the ef- as theorists. The simplest and perhaps most appropri-
fects of fragmentation and corridors. ate definition of habitat would be that part of the en-
Discussed more often than applied, a common mea- vironment within a species’ geographic range where
sure of corridor effectiveness has been movement of the species actually lives (Morrison and Hall 2002).
individuals within the corridor, or corridor use. This However, Hall et al. (1997) found vague or imprecise
measure goes to a hypothesized principal mechanism characterizations of habitat in 82% of the fifty papers
of population decline caused by habitat fragmentation they reviewed. A popular characterization of habitat
and of population rescue provided by corridors. Organ- has been the classification of vegetation into “habitat
isms must be able to disperse, which habitat fragmen- types,” to which species are then associated based on
tation is hypothesized to hamper and which corridors relative abundance of the species in each type. Even
are hypothesized to facilitate. Indeed, Gilbert-Norton vaguer is the characterization of habitat as all types
et al.’s (2010) meta-analysis of corridor effectiveness of natural vegetation bounded by lands converted for
tested whether movement increased with the existence human use (Bolger et al. 1997). This characterization
of natural or created corridors. However, a dependent of habitat glosses over what specifically it is in the en-
variable measured as the rate of movement between vironment that species are selecting; it overlooks the
habitat patches does not express the capacity of the resources in the environment that each species needs
remaining habitat. The investigators using this depen- for metabolism and reproduction, and which limits the
dent variable only assume that improved movement species’ abundance.
translates into greater habitat capacity. They selected The very fact that species are selecting resources
movement as their dependent variable because move- from the environment indicates that the second law
ment is critical to colonization and gene flow, which of thermodynamics applies to the distribution and
they believed increases population viability and de- abundance of wildlife (Smallwood 1993, 2002). The
creases extinction probability. In reality, the movement primary regulator of the distribution and abundance
that is measured could be of the same individual travel- of any wildlife species is the degree to which individu-
ing back and forth between habitat patches, or it could als can metabolize sufficiently and for long enough to
be of individuals dispersing into an ecological sink. successfully reproduce. The primary resource must be
Whereas the measurement of movement rates along energy for metabolism, but then other resources are
a corridor can be reassuring, it cannot alone express needed to locate and process this energy, including
corridor effectiveness because it is in the connected atmosphere (water), nutrients, ecosystem processes,
habitat patches where the corridor’s effects ultimately and the suite of other biological species in the envi-
matter. ronment. Because energy needed for metabolism is the
Investigators have often discussed or used indica- primary resource that defines a species’ habitat, these
tors of habitat fragmentation, including percentage of energy sources should be the core of any definition of
habitat lost, perimeter-to-area ratio, mean patch size, a species’ habitat, and the portions of the environment
and fractal dimension. The usefulness of these indica- that facilitate the metabolism of these resources should
tors depends on the degree of correspondence between round out the definition. Thus, habitat is defined as the
the indicators and direct measures of fragmentation. suite of environmental elements—soils, plant species,
 habitat fragmentation and corridors 93

and other animal species—that includes the species’ temporal grain and extent. As for the null condition in
limiting resources. Habitat is a vague, indicator-level which the investigator assumes that species’ individu-
representation of the parts of the environment that are als would be randomly spaced, deviations between the
essential to the species’ metabolism and reproduction, null condition and the measured condition would be
and hence to its persistence (Morrison et al. 1998). shorter, and these deviations might vary depending on
The complexities of measuring habitat fragmenta- what is meant by random.
tion increase when considering the more fundamental A uniform pattern is easy to define. A uniform pat-
challenges of measuring habitat selection. After all, tern results when the study units are maximally distant
measuring habitat fragmentation presupposes that the from each other. It is the pattern one would expect if
habitat undergoing fragmentation was accurately char- there were no relationships between organisms and en-
acterized as selected habitat. A fundamental problem vironmental resources. An interesting exception to this
with this presupposition is that the species perform- expectation would be when regular patterns of distribu-
ing the measurements is not the species whose habitat tion result from social rules organized around the spe-
selection is being measured, so we should not assume cies’ long-term experience with the resources, but even
that we perceive the species’ environment in the same in cases of socially driven regularity, deviations from a
manner as the species we are studying. We draw infer- uniform pattern should be evident at larger spatial ex-
ences from patterns in the data, and tend to conclude tents. Not only is meaningful pattern relatively easy to
that a species has selected a portion of the environ- measure as deviations from a null pattern of uniformity,
ment based on the species’ disproportionate number but these deviations can also be interpreted from laws
of individuals or time spent in that selected area. Our of thermodynamics and information theory (Shannon
inferences, however, could be confounded by multiple and Weaver 1949; Kullback 1959; Phipps 1981).
factors, including our measurement of the wrong en- Many might regard a random pattern as also easy to
vironmental variables, the wrong spatiotemporal grain define. A random pattern is the pattern of distribution
or extent, and our counting of the wrong study unit, resulting from lack of influence from the locations of
e.g., individuals versus breeding pairs versus colonies other individuals (Fisher 1950; Taylor 1961). Because
versus populations (discussed in the following section). many observed or simulated patterns of distribution
In some cases, disproportionate use of an area might can be regarded as random, the starting point for mea-
represent an ecological sink (Smallwood 2002). suring meaningful patterns of aggregation is less clear
A second fundamental problem with measuring than it is for uniformity. Along a theoretical continuum
habitat selection is the epistemological context of the from uniform to aggregated patterns, randomness oc-
measurements, such as whether the investigator mea- cupies an unknown and perhaps varying proportion of
sures meaningful pattern as deviation from a uniform this range somewhere between uniformity and com-
or random null pattern (Smallwood 2002). The start- pletely aggregated (see Smallwood 2002, fig. 6.3).
ing point matters for measuring meaningful associa- Furthermore, the definition of a random pattern high-
tions with places on the Earth where environmental lights interactions among individuals, which serve as
resources might be selected. The null condition in the most common study unit in habitat selection stud-
which the investigator assumes that species’ individu- ies. It implies that interactions among individuals re-
als would be uniformly spaced would result in greater sult in the meaningful patterns of aggregation that we
deviations between the null condition and the mea- measure, but in so doing it also obscures the influence
sured condition, so long as the appropriate variable(s) of environmental resources. Of course, we know that
is being measured at the appropriate spatiotemporal individuals interact and that these interactions influ-
grain and extent. If the species’ individuals space them- ence the patterns of distribution that we observe, but
selves out in a regular pattern due to social rules (home randomness as the null condition invokes these inter-
ranges), then the study unit might more appropriately actions as paramount and starts the analysis of habitat
consist of social groups rather than individuals, and selection from a different worldview than starting it
measurements might be needed at much larger spatio- from uniformity as the null condition.
94 habitats in peril

A third fundamental problem with measuring habi- founding due to resource instability in past and even
tat selection—and this problem is related to the sec- relic environments, but measuring the effects of habi-
ond when it comes to detecting meaningful patterns of tat fragmentation is prone to further complications
habitat selection—is the spatiotemporal grain and ex- due to a new type of resource instability in the current
tent of the measurements, whether the measurements environment.
are made during a snapshot of the organisms life (or A fourth fundamental problem with measuring hab-
over a season, a year, multiple years, multiple genera- itat selection is achieving replication and interspersion
tions, or multiple periods of the species’ multiannual of “treatments,” which in the context of habitat selec-
cycle of abundance) and whether the measurements tion studies are the categories of vegetation, landscape,
are made within an animal’s home range, across mul- or specific resources being associated with counts of
tiple home ranges, or across an area capable of support- the species under study. Pseudoreplication results from
ing an entire population or larger demographic unit. In inadequate replication and interspersion of treatments,
other words, the sampling resolution and measurement but these conditions are often more difficult to achieve
sensitivity must be sufficient for testing hypotheses of in mensurative studies (Hurlbert 1984). Testing the ef-
habitat selection (Huston 2002) before any meaning- fects of habitat fragmentation and corridors is likely
ful tests can be performed on the effects of habitat even more prone to pseudoreplication than is testing
fragmentation or corridors. As an example, measuring for habitat selection.
habitat selection immediately after land conversions A fifth fundamental problem is often not knowing
fragmented the habitat could be misleading due to tem- why the animals that were counted were located where
porary crowding of individuals in the remaining habitat they were found by researchers. The aggregations we
fragments (Whitcomb et al. 1981; Lovejoy et al. 1986; encounter can be one of four types (Smallwood 2002):
Scott 1993). resource, demographic, early stage, and constrained.
The capacity of a species to utilize an environment A resource aggregation would likely be a temporary col-
(i.e., its habitat), or to populate its numbers within lection of individuals where a limiting resource has be-
an environment, will be limited by the most limiting come available. A demographic aggregation might be a
resource according to Liebig’s Law of the Minimum. temporary collection of breeding pairs or a gathering of
This limitation might vary across the species’ environ- juveniles or subadults developing social relationships.
ment or through time at given locations. This variation An early-stage aggregation would be the collection of
in where and when Liebig’s Law of the Minimum will individuals corresponding with the level of coloniza-
operate defines the ultimate source of instability in tion at the time of the habitat survey. A constrained ag-
the environment with which a species must contend. gregation would be the type of aggregation that is most
Therefore, each species evolves strategies to contend prone to confounding in the context of fragmented
with the instability in its most limiting resource(s), habitat, because it is a forced aggregation, or even an
and the most effective strategies tend to be behavioral expression of crowding. Either way, the constrained ag-
and social in nature, often resulting in time lags and gregation is likely temporary, so measurement of it can
feedback loops affecting numerical responses. Species yield an overoptimistic interpretation of habitat selec-
of wildlife tend to buffer themselves against instability tion (or corridor use).
in the most limiting resources by developing stubborn Also related to habitat selection and how its mea-
social rules, such as home range tenure (home ranges surement can influence interpretation of the effects of
often remain unchanged even in the face of superabun- habitat fragmentation is the effect of location. Although
dant food resources) and mating hierarchies. They also there has been discussion of the influences of the ex-
buffer themselves against resource instability by devel- tent of habitat loss and degree of isolation on habitat
oping stubborn behaviors such as seasonal migrations, fragmentation, as well as the edge-to-interior ratio and
foraging self-limitations, and foraging by gestalt where the types of activities and species composition in the
resources are expected to be found rather than where areas surrounding habitat fragments, there has been
they are known to be found (Hutto 1990). Therefore, little, if any, discussion of the effect of location. The in-
not only is measuring habitat selection prone to con- fluence of location on corridor effectiveness has often
 habitat fragmentation and corridors 95

been discussed, but its influence on habitat fragmenta- ever, conceptualizations of habitat fragmentation and
tion has been neglected. Certain habitat areas are likely corridors vary widely in the scientific literature and are
strategic to a species for one or more reasons, so their either poorly understood or intentionally abused by
removal would have disproportionate consequences for many practitioners. The science supporting these con-
the species. An example would be a landscape bottle- cepts needs to improve. Appropriate dependent vari-
neck, where animals must be able to negotiate for dis- ables representing responses to habitat fragmentation
persal. Another example would be systematic loss of and corridors need to be measured at appropriate spa-
habitat in a particular landscape context, such as the tial grain and extent and over appropriate time periods.
upland grassland areas surrounding ponds. Many spe- Quantitative tests of habitat fragmentation and corri-
cies forage or breed in ponds but spend the rest of their dor effectiveness should be made to single species that
lives in the surrounding grasslands. Therefore the ef- are sufficiently understood to measure habitat variables
fects of habitat fragmentation might vary by location, rather than using overly simplistic maps of vegetation
all else being equal. cover types that are applied to suites of wildlife species
(e.g., Bolger et al. 1997). Much more care is needed on
the measurement of habitat fragmentation and corridor
Permeable Movement Barriers
effectiveness, and in the discussion of these concepts.
Just as difficult as it might be to characterize habi- Definitions of habitat fragmentation and corridors
tat fragments as places where a species would really need to be more explicit. Scientists need to more effec-
choose to reside is to characterize the matrix areas sur- tively convey their concepts of habitat fragmentation
rounding the fragments as nonhabitat and as barriers and corridors to biologists in natural resource agencies
to movement between fragments. For some species, the and environmental consulting firms, as well as to attor-
matrix might be easily shown to block movement, but neys, political decision-makers, and the public. Where
for others the matrix might serve as a portion of the these concepts really matter is where decisions are be-
species’ habitat and be completely or partially perme- ing made about whether and how to further fragment
able to movement. La Polla and Barrett (1993) tested habitat and how to mitigate the effects of this fragmen-
corridor width and presence on captive-reared meadow tation. These decisions are being made now, and with
vole (Microtus pennsylvanicus) density in grassland frag- the proliferation of habitat conservation plans these de-
ments, but the matrix was simply mowed grass. The cisions increasingly involve very large areas of wildlife
mowed grass may have been completely permeable to habitat. Vague and poorly understood concepts such
vole movement between unmowed fragments. Tests for as habitat fragmentation and corridors can cause more
habitat fragmentation and corridor effectiveness need harm than good in wildlife conservation, as I have seen
to clearly characterize the ability of each species’ ability too many times in the aftermath of the environmental
to move across or reside within the matrix areas. reviews I addressed in this chapter.

ACKNOWLEDGMENTS
Conclusions
I appreciate the review comments on this chapter from
Habitat fragmentation likely qualifies as a major threat A. M. Long and an anonymous reviewer. I also appre-
to the persistence of many plant and animal species, ciate M. L. Morrison for our many discussions of the
and corridors might help to mitigate this threat. How- habitat concept.
APPENDIX 7.A
Summary of environmental reviews with which I have had personal experience and whether the reviews
mentioned or analyzed the potential project impacts on habitat fragmentation or movement corridors.
Citations appear in Appendix 7.B by reference (Ref) number in the right column. MW refers to megawatts
in energy projects.
Mention / analysis
of potential impacts
related to:

Habitat Movement
Source Project Acres fragmentation corridors Ref

BLM 2012a 104 MW Steens Mt Wind Energy Project 8,700 yes / no no / no 1

Kern County 2012 250 MW Beacon Photovoltaic Project 2,301 no / no yes /no 2
Yolo County 2008 Clark-Pacific Concrete Precast Factory 90 no / no yes / no 3
PBS&J 2008 Delta Shores residential / commercial development 782 yes / no yes / no 4
Sacramento LAFCo 2011 Expand City of Elk Grove’s Sphere of Influence to 7,869 yes / yes yes / no 5
accommodate future residential / commercial
development
BLM 2010 370 MW Ivanpah Solar Electric Generating System 4,073 yes / no yes / no 6
City of Lancaster 2012 60 MW Summer Solar and Springtime Solar Projects 216 no / no yes / no 7
HDR 2011 600 MW Mount Signal & Calexico Solar Farm Projects 4,228 no / no yes / no 8
Impact Sciences 2002 Newhall Ranch residential development 12,000 yes / no yes / no 9
Caltrans 2006, 2010 Route 84 Safety Improvement Project (Niles Canyon) 24 yes / no yes / no 10
BLM 2012b 20 MW Ocotillo Sol Project 115 yes / no yes / no 11
Foothill Associates 2006 Regional University development 1,282 yes / no yes / no 12
EDAW 2008 Rio del Oro Specific Plan, housing development 3,828 yes / no yes / no 13
Imperial County 2012 200 MW Solar Gen 2 Array Project 2,009 yes / no yes / no 14
Design, Community & Environ- St. John’s Church expansion, requiring removal of 3 no / no yes / no 15
ment 2010 mature forest canopy
Pacific Municipal Consultants Yuba Highlands Specific Plan residential development 2,902 yes / no yes / no 16
2005
Ecology and Environment 2007a, b 67 MW Windy Flats West Wind Energy Project 616 no / no no / no 17
Antonio Mountain Ranch residential development 523 yes / no yes / no 18
BLM 2012c 150 MW Desert Harvest Solar Project 1,464 yes / no yes / no 19
Ecology and Environment 2012 49.9-MW Hudson Ranch Power II Geothermal Project 100 yes / no yes / no 20
and Simbol Calipatria Plant II
WRA Environmental Consultants J&J Ranch, 24 Adobe Lane residential development 20.5 no / no yes / no 21
2009
Harzoff 1992; Huffman & Associ- Putah Creek Parkway Demonstration Project, adding 10 no / no no / no 22
ates 1992 bike path and fence along centerline of riparian
corridor
UC Davis Long-Range Development Plan no / no no / no 23
Yeast 1998 Sunset SkyRanch Airport expansion 108 no / no no / no 24
Raney Planning & Management, Covell Village residential development 422 no / no yes / no 25
Inc. 2004
Planning Consultants Research West Bluff residential development, the last upland 44 no / no yes / no 26
area bordering the Ballona Wetlands
Thomas Reid Associates 1997 Natomas Basin HCP for residential development on 21,300 yes / no no / no 27
wetlands and fields used for rice cultivation
County of Yolo 2010 365-foot-tall Results Radio Tower with guy wires 1.7 no / no no / no 28
Glazner Environmental Consult- Atwood Apartments 55.6 yes / no no / no 29
ing 1999
City of Sierra Madre 2001; EIP Maranatha High School 63 yes / no yes / no 30
Associates 1999
Grassetti Environmental Consult- Creekside Highlands residential development along 7.1 yes / no yes / no 31
ing 2004; Edelstein 2003 riparian zone of Ward Creek
Appendix 7.A continued

Mention / analysis
of potential impacts
related to:

Habitat Movement
Source Project Acres fragmentation corridors Ref

Humboldt Bay Harbor & US Army Dredging and disposal of 5.6 million cubic yards of — no / no no / no 32
Corps of Engineers 1995 benthic marine sediment in Humboldt Bay Harbor
Ogden 1998 San Diego Multiple Habitat Conservation Program for 121,023 yes / no yes /no 33
incidental take permits of 85 species
EGI 2012 140 MW Campo Verde Solar Project 1,990 no / no yes / no 34
ESA 2012 33 MW Casa Diablo IV Geothermal Development 52.5 no / no yes / no 35
Project
ECORP 2011; Kern County 2012 20 MW FRV Orion Solar Project 260 yes / yes yes / no 36
Chambers Group 2012 30 MW Imperial Valley Solar Company 2 Project 158.5 no / no yes / no 37
Glazner Environmental Consult- Atwood Ranch Unit III Subdivision 55.6 no / no yes / no 38
ing 1999
City of Sacramento 2011; AES 2011 20 MW Sutter Landing Park Solar Photovoltaic Project 104 no / no no / no 39
CEC 2007 660 MW Colusa Generating Station, gas-fired power 31 yes / no no / no 40
plant
City of Petaluma 2013 Residential development 59 yes / no yes / no 41
Kern County & ESA 2012 125 MW Pioneer Green Solar Project 720 no / no no / no 42
Swaim 2011 Oakland Zoo expansion 62 yes / no no / no 43
Contra Costa County 2001 Expansion of Temple B’nai Tikyah, Walnut Creek, CA 0.3 no / no no / no 44
Jones & Stokes Associates 1999 Valley elderberry longhorn beetle HCP for gravel mine 9.9 no / no no / no 45
in Yolo County, CA
City of Anderson 2003 Anderson Marketplace (Walmart) commercial devel- 26.6 no / no no / no 46
opment
Sacramento County 2002 The Promenade commercial development 8 no / no no / no 47
North Fork Associates 2000 Silver Bend Apartments 6 no / no no / no 48
Foothill Associates 2002 Winters Highlands residential development 102.6 no / no no / no 49
EIP Associates 2001 UC Merced NCCP / HCCP 2133 yes / no yes / no 50
URS 2001 UC Merced Long Range Development Plan EIR 2000 yes / no no / no 51
Beck 2003 Table Mountain mining 50 no / no no / no 52
Foothill Associates 2004; City of Callahan Estates residential development 26.4 no / no no / no 53
Winters 2004
CEC 2000 Blythe Energy Project 76 yes / no no / no 54
Kern County 2013 100 MW Rosamond Solar Project 400 yes / no yes / no 55
Arnold 2013 7.5 MW LANDPRO Solar Project 80.6 no / no yes / no 56
ESA 2012, Sage Institute, Inc. 2011 Expansion of Metropolitan Air Park, City of San Diego 331 no / no yes / no 57

APPENDIX 7.B
References for environmental review documents reviewed for treatment of corridors and habitat
fragmentation.
Ref Reference

1 BLM (Bureau of Land Management). 2012. North Steens 230-kV Transmission Line Project Draft Environmental Impact Statement.
DOI-BLM- OR-B060-2010-0035-EIS, Hines Oregon.
2 Kern County. 2012. Draft Environmental Impact Report, SCH# 2012011029, Beacon Photovoltaic Project by Beacon Solar LLC, Con-
ditional Use Permit 11, Map 152, Kern County Planning and Community Development Department, Bakersfield, California.
3 Yolo County. 2008. Clark Precast, LLC’s “Sugarland” Project, Initial Study / Mitigated Negative Declaration, Zone File # 2007-078,
Yolo County Planning & Public Works Department, Woodland, CA.
4 PBS&J. 2008. Delta Shores Draft Environmental Impact Report. City of Sacramento. Sacramento, CA.
continued
Appendix 7.B continued

Ref Reference

5 Sacramento LAFCo (Local Agency Formation Commission). 2011. Draft Environmental Impact Report: City of Elk Grove Proposed
Sphere of Influence Amendment (LAFC # 09-10). State Clearinghouse No. 2010092076. City of Elk Grove, Sacramento County,
California.
6 BLM (Bureau of Land Management). 2010. California Desert Conservation Area Plan Amendment / Final Environmental Impact
Statement for Ivanpah Solar Electric Generating System, FEIS-10-31, Needles, CA.
7 City of Lancaster. 2012. Initial Study for Conditional Use Permits 12-08 and 12-09, Summer Solar and Springtime Solar Projects.
Lancaster, CA.
8 HDR (Engineering, Inc.) 2011. Draft Environmental Impact Report Mount Signal and Calexico Solar Farm Projects, Imperial County,
California. County of Imperial, El Centro, California.
9 Impact Sciences. 2002. Newhall Ranch November 2000 Administrative Draft EIR. Los Angeles County Department of Regional Plan-
ning, Los Angeles, California.
10 Caltrans (California Department of Transportation). 2006. Route 84 Safety Improvement Project Negative Declaration (CEQA).
Dividing Border Between Union City and the City of Fremont in Alameda County, California, 04-ALA-84-KP 19.5 / 21.4, EA 174400.
Caltrans, Oakland, California.
10 Caltrans (California Department of Transportation). 2010. Niles Canyon Safety Improvement Project Draft Environmental Impact
Report / Environmental Assessment. Alameda County, California.
11 BLM (Bureau of Land Management). 2012a. El Centro Office Draft Environmental Impact Statement / Draft CDCA Plan Amendment
Ocotillo Sol Project. US Department of the Interior, BLM / CA / ES-2012-009+1793, DES 20-12, DOI-BLM- CA-D000-2012-0005-
EIS. El Centro, California.
12 Foothill Associates. 2006. Biological Resources Assessment: Regional University site and off-site improvements, Placer County, Cali-
fornia. Prepared for KT Development. County of Placer, Auburn, California.
13 EDAW. 2008. Recirculated Draft Environmental Impact Report / Supplemental Draft Environmental Impact Statement, Rio del Oro
Specific Plan Project, State Clearinghouse #2003122057, City of Rancho Cordova and US Army Corps of Engineers, Sacramento
District.
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27 Thomas Reid Associates. 1997. Natomas Basin Habitat Conservation Plan, Sacramento and Sutter Counties, California. City of Sacra-
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Appendix 7.B continued

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31 Edelstein, D. 2003. Analysis of impacts of Creekside Highlands. App. A in Grassetti Environmental Consulting (2004). Initial Study
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31 Grassetti Environmental Consulting. 2004. Initial Study and Draft Mitigated Negative Declaration, Vesting Tentative Map, Tract
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32 Humboldt Bay Harbor Recreation and Conservation District & US Army Corps of Engineers. 1995. Final Environmental Impact State-
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33 Ogden (Ogden Environmental and Energy Services Co., Inc.) 1998. Biological goals, standards, and guidelines for Multiple Habitat
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34 EGI. 2012. Final Environmental Impact Report for the Campo Verde Solar Project, SCH. No. 2011111049. Prepared for Imperial
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35 ESA (Environmental Science Associates). 2012. Casa Diablo IV Geothermal Development Project, Draft Environmental Impact State-
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36 ECORP. 2011. Biological Evaluation and Impact Analysis Orion Solar Site, Kern County, CA. Report to SunEdison, San Francisco,
California.
36 Kern County. 2012. Draft Environmental Impact Report, SCH# 2012031079, FRV Orion Solar Project (PP12232). Bakersfield, Cali-
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37 Chambers Group, Inc. 2012. Draft Environmental Impact Report, Imperial Valley Solar Company 2 Project, Imperial County, Cali-
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39 AES (Analytical Environmental Services). 2011. Biological Resources Assessment, City Of Sacramento 28th Street Solar Photovoltaic
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39 City of Sacramento. 2011. Community Development Department Solar Photovoltaic Park at 28th Street Landfill Project, Initial Study.
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40 CEC (California Energy Commission). 2007. Preliminary Staff Assessment: Colusa Generating Station, Application For Certification
(06-AFC-9), Colusa County, CEC-700-2007-003-PSA, Sacramento, CA.
41 City of Petaluma. 2013. Davidon Homes Tentative Subdivision Map and Rezoning Project Draft Environmental Impact Report and
Technical Appendices. State Clearinghouse No. 2004072137. City of Petaluma, California.
42 Kern County and ESA (Environmental Science Associates). 2012. Draft Environmental Impact Report, Pioneer Green Solar Project
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field, California.
43 Swaim, K. (Swaim Biological, Inc.). 2011. Oakland Zoo California Project Biological Assessment. Report to East Bay Zoological
Society, Oakland, California.
44 Contra Costa County. 2001. Initial Study for the Expansion of the Congregation Temple B’nai Tikyah, Walnut Creek, CA.
45 Jones & Stokes Associates. 1999. Habitat Conservation Plan for the Valley elderberry longhorn beetle for A. Teichert & Sons, Inc.,
Yolo County, California. Prepared for US Fish and Wildlife Service, Sacramento, California.
46 City of Anderson. 2003. Anderson Marketplace Project Draft Environmental Impact Report. City of Anderson Planning Department,
Anderson, California.
47 Sacramento County. 2002. Revised Draft Environmental Impact Report: The Promenade Community Plan Amendment, Rezone,
Use Permit, Variances, and Special Review of Parking. State Clearinghouse No. 1998042028. Sacramento County Department of
Environmental Review and Assessment, Sacramento, California.
48 North Fork Associates. 2000. Biological Resources Assessment for the ±6-acre Silver Bend Apartment Project, Placer County, Cali-
fornia. Auburn, California.
49 Foothill Associates. 2002. Biological Resources Assessment: Winters Highlands Project Site. City of Winters, California.
50 EIP Associates. 2001. University Community Plan EIR. State Clearinghouse No. 2001021056. County of Merced, California.
51 URS. 2001. UC Merced Long Range Development Plan Draft Environmental Impact Report. Merced County, California.
52 Beck, A. J. 2003. Biological assessment of a portion of North Table Mountain, Central Butte County, California. Unpubl. report by
Eco-Analysts, Chico, California.
53 City of Winters. 2004. Initial Study and Negative Declaration for Callahan Estates. City of Winters, California.
54 CEC (California Energy Commission). 2000. Preliminary Staff Assessment: Blythe Energy Power Plant Project, Application for Certi-
fication 99-AFC-8, Riverside County, Sacramento, CA.
continued
100 habitats in peril

Appendix 7.B continued

Ref Reference

55 Kern County. 2013. Addendum to the Environmental Impact Report for the Rosamond Solar Project, ROSAMOND SOLAR MODI-
FICATION PROJECT (PP13292), SGS Antelope Valley Development, LLC, Specific Plan Amendment No. 18, Map No. 232,
Specific Plan Amendment No. 19, Map 232, Zone Change Case No. 36, Map No. 232, Modification to Conditional Use Permit No.
27, Map No. 232, Streets and Highways—Nonsummary Vacation—Map No. 232. Kern County Planning and Community Develop-
ment Department, Bakersfield, California.
56 Arnold, R. (RCA Associates). 2012a. General biological resources assessment, LANDPRO 7.5 MW Solar Project, APN0466-181-059,
060, 061 & 062, San Bernardino County, California. Unpublished report to Sunlight Partners, LLC, Mesa, Arizona.
57 ESA. 2012. Metropolitan Airpark Project Draft Environmental Impact Report, SCH No. 201007054, Project No. 208889, City of San
Diego Development Services Department, San Diego, California.
Sage Institute, Inc. (SII). 2011 (revised by ESA in 2012). Metropolitan Airpark Project Volume 1, 2011 Biology Survey Report, Project
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 8
Julie L. Lockwood and
The Impact of Invasive
Species on Wildlife
J. Curtis Burkhalter

Habitat

N onnative species are an increasingly prevalent

aspect of wildlife habitat. In today’s world, it is
nearly impossible to walk through a forest, meadow, or
as “invasive” only if they are abundant and widespread
enough to cause negative ecological impact (Lockwood
et al. 2013). A negative impact occurs when the pres-
wetland and not readily encounter several nonnative ence of an invasive species causes a reduction in the
plants and animals. Some of these nonnative species distribution, abundance, or individual performance
are so common that it is difficult to imagine them fail- (survival, reproduction) of one or more native species
ing to have an effect on co-occurring native wildlife. (Parker et al. 1999). Only a small fraction of the spe-
Other nonnative species are so rare and localized in cies that have the opportunity to be invasive (i.e., are
their distribution that they seem innocuous. What is transported out of their native range and released in
a wildlife biologist to make of this range of nonnative a nonnative locale) are known to go on to become in-
species impacts? When should the presence of nonna- vasive (Williamson and Fitter 1996). Blackburn et al.
tive species become a concern in regard to their effects (2011) provide a comprehensive and integrative frame-
on native wildlife populations? The answers to these work for considering the process of species invasion
questions are, of course, quite complex. In this chapter, (fig. 8.1). This framework defines the series of barriers
we tackle this complexity by summarizing currently ac- and filters that a nonnative species must transit before
cepted frameworks for defining nonnative species and it becomes invasive. These barriers are not trivial,
measuring their impacts, and then reviewing a suite of and the proportion of species that transit each com-
mechanisms by which nonnative species can degrade monly hovers at around 10–20% (Williamson and Fit-
the habitat quality of native wildlife. This review is ter 1996), although it can reach as high as 40–50% in
designed to provide a scaffold onto which biologists some situations (Jeschke and Strayer 2006).
can begin to place their own experiences in managing Morrison et al. (2006) define wildlife habitat as “an
nonnative species for the benefit of wildlife, and with area with a combination of resources (like food, cover,
which to more readily access the expansive literature and water) and environmental conditions (tempera-
on biological invasions. ture, precipitation, presence or absence of predators
or competitors) that promotes occupancy by individu-
als of a given species (or population) and allows those
Definitions and Framework
individuals to survive and reproduce.” Given this defi-
The title of this chapter introduces three concepts (in- nition, invasive species will negatively impact wildlife
vasive, impact, and wildlife habitat) that we must de- habitat by altering resources or conditions to the extent
fine before moving to mechanisms of invasive species that survival and reproduction of a targeted species is
impacts. The first two terms are related in the sense affected. This impact can be imposed by (1) the invader
that, by most definitions, nonnative species are labeled maintaining a numerical dominance over native wild-
 the impact of invasive species on wildlife habitat 103

Figure 8.1 The invasion process can be divided into a series of stages. For each stage there are associated barriers that
must be overcome before the species can proceed into the next stage. The number of nonnative species that successfully
transit any one stage is generally thought to be quite small (<20%), thus dictating that most introduced nonnative species
fail to reach an abundance and geographical range size that allow them to negatively impact native wildlife. Adapted from
Blackburn et al. (2011)

life, (2) the invader being so common across space that will not be felt evenly across all locations within that
very few individuals in a wildlife population will fail to ecosystem.
come into contact with the invaders, (3) indirect effects Nonnative species typically will not impact native
mediated through one or more native species, (4) non- ecosystems until they reach some critical range size
native species changing ecosystem properties (e.g., and abundance. This threshold is not well defined for
changes in nutrient cycling, disturbance regimes), or any single invader and certainly varies according to
(5) a high per capita negative effect of a single invader the strength of the per capita effect of the invader on
on a single native individual. the native species (fig. 8.2). However defined, it will
These definitions provide a clear way to link wildlife typically take several years between when a nonnative
habitat and invasive species impacts. However, there is species is newly discovered in a habitat and its attain-
hidden complexity behind these concepts that stems ment of a population size and distribution that allow it
from considering how impacts may vary through time to impact co-occurring native wildlife (fig. 8.2). This
and across space. Not all invasive species will show lag between arrival and becoming invasive may be rela-
impacts on wildlife habitat over the duration of their tively short or it can be quite prolonged (e.g., more than
tenure in an ecosystem, and the impact of the invader one hundred years), with the time dictated mostly by
104 habitats in peril

at low numbers for a long period when in fact it is on its

way to causing serious negative impacts (Crooks 2005).
Additionally, unexpected events in time and space (e.g.,
disturbances) can also have unexpected consequences
in regard to how seemingly benign nonnatives may rap-
idly proliferate into new areas following disturbance
events (Keeley et al. 2003). Equally unwelcome is the
possibility that the impacts of a particularly trouble-
some invader will lessen through time, thus leading a
manager to expend great resources on a problem that
will naturally dissipate (Strayer et al. 2006). In the
future, climate change may also alter whether ecologi-
cally benign nonnative species become invasive (Hell-
Figure 8.2 The impact of an invasive species is dependent
mann et al. 2008).
on its abundance, geographical range size, and per capita
effect. The solid line depicts standard logistic popula- It is relatively easy to extend this complexity in
tion growth in a nonnative population, which includes measuring the impact of invasive species across space.
a period of time when the nonnative is inherently rare. Not all locations within a particular habitat will sustain
The dashed line represents population growth with a equal numbers of the invasive species due to relatively
prolonged lag, whereby abundance stays low for longer fine-grained differences in resources and environmen-
than logistic growth would dictate. Native wildlife will tal conditions. The ecological “rules” that govern how
not suffer reduced survivorship or fecundity (impacts) native species respond to such environmental condi-
until some threshold abundance (depicted here) or range tions will also apply to any co- occurring nonnative
size is achieved. With or without a prolonged lag, newly species. Thus, some locations will provide optimal
established nonnative populations will not reach this
habitat for the invader and some only marginal. For
threshold until some time has passed. The time required
a wildlife manager, a sound knowledge of a nonnative
in any one case is dependent on how long the population
species’ niche requirements will greatly inform where
stays rare, as well as the per capita effects of nonnative on
native individuals. this species will impose strong impacts on co- occurring
wildlife and where it will not. Management decisions
can then proceed toward optimizing scarce human
how fast the nonnative population can grow (Crooks and economic capital to ameliorate invasive species
2005). The mechanisms that produce prolonged lags impacts only where they are likely to occur.
are numerous and include genetic impoverishment, With these definitions and frameworks in mind,
Allee effects, or negative biotic interactions, although we will devote the rest of this chapter to reviewing the
no one mechanism has yet explained the majority of various mechanisms by which an invasive species can
lags (Crooks 2005; Lockwood et al. 2013). In addi- affect wildlife habitat. In providing examples of these
tion to lags, there is the possibility that the impacts of mechanisms, we will illustrate the complexity inherent
invasive species will have an acute stage after which in defining and measuring impact. Our review is orga-
impacts will attenuate (Strayer et al. 2006; fig. 8.2). nized so that we cover the direct impacts of invasive
The attenuation of impacts can occur via behavioral or species on wildlife habitat first, followed by indirect
evolutionary adaptation of the local flora and fauna to mechanisms of impact, and finally synergisms between
the invader or through a drop in the abundance or geo- invasive species impacts and other known wildlife hab-
graphical extent of the invasive population (e.g., over- itat stressors (e.g., fragmentation).
shooting a density limitation and then returning to a
number of individuals that is better matched to local
Direct Impacts
resource conditions). The element of time thus intro-
duces the possibility that managers may dismiss a non- Invasive species can impact wildlife habitat by directly
native species as ecologically benign because it persists altering resource levels or environmental conditions
 the impact of invasive species on wildlife habitat 105

such that they reduce the survival or fecundity of co- quality of that habitat. Watling et al. (2011b) tested for
occurring wildlife. Invasive plants can impose such such an effect on a range of native amphibians that use
changes by (1) altering vegetation composition so that oak-hickory forests in Missouri. They found that the
available cover is lost or degraded in usefulness, (2) re- higher the density of Amur honeysuckle, the fewer am-
placing, or otherwise making inaccessible, plants or phibian species recorded. In addition, the presence of
invertebrates that provide critical food, and (3) chang- honeysuckle tended to support the same suite of spe-
ing the chemistry, salinity, water flow, or other physi- cies across space, whereas native vegetation supported
cal aspects of habitat so that suboptimal physiological a wider variety of species. This change in spatial diver-
conditions are created. Invasive animals can directly sity patterns in native amphibians is perhaps due to the
impact native wildlife either by eating or competing honeysuckle creating relatively homogeneous microcli-
with native wildlife, or by producing toxins that harm mates across space, whereas native vegetation created
native wildlife predators that consume them. patches with differing conditions that support differing
suites of amphibians (Watling et al. 2011b). There was
also a clear shift in the set of amphibian species found
Invasive Plants
across a gradient of honeysuckle density, which Watling
The structure that vegetation produces is a key com- et al. (2011b) attributed to the cooler microclimate pro-
ponent of wildlife habitat, providing suitable microcli- duced by heavy honeysuckle cover.
mates, nesting substrate, and protection from preda- Nonnative plants found in North American grass-
tion, and supporting critical food resources (Price and lands provide an example of more contextual and nu-
Waser 1984; Morrison et al. 2006). Vegetation struc- anced effects on wildlife. Grasslands have a particular
ture also strongly influences the physical processes structure characterized by low abundance of woody
(e.g., fire and flood) and chemistry that characterize vegetation, few or no large trees, and a predominance
wildlife habitat (Brooks et al. 2004; Brown et al. 2006). of perennial grasses (White et al. 2000). Several native
Invasive plants can alter vegetation structure to vary- birds are considered grassland obligates, and thus the
ing degrees and concomitantly alter habitat features for loss of grassland habitat directly results in the decline in
wildlife that depend on these structures. Some inva- these birds’ populations (Johnson 2000). Some of these
sive plants radically change vegetation structure, earn- grassland losses are obvious (e.g., conversion to hous-
ing them the label of ecosystem engineer (Wang et al. ing developments), while others are subtle, involving
2006), while others seem to replicate the structure pro- a switch from native to nonnative grasses, sometimes
duced by their native plant counterparts. In the former with the grassland being dominated by a single nonna-
case, managers should expect large impacts on native tive grass. However, the presence of nonnative grasses
wildlife, while in the latter there may be no or only does not automatically transform a grassland into un-
mild impacts. In table 8.1, we provide an overview of suitable habitat for native birds (e.g., Jones and Bock
literature that addresses this continuum, including ex- 2005; Kennedy et al. 2009). The extent to which inva-
amples of invasive plants that alter cover, change food sion by nonnative grasses will influence native birds
resources, and transform the physical environment. seems to be tied to the degree to which the nonnatives
An example of an invasive ecosystem engineer is the mimic the vegetation structures produced by natives
Amur honeysuckle (Lonicera maackii), which has estab- (Kennedy et al. 2009). In this case, structure is impor-
lished itself within the oak-hickory forests of the east- tant for nest construction, placement so that the nest
ern and midwestern United States. This plant produces is concealed from predators, and the support of inverte-
a dense shrub layer that is absent in the typically open brates that provide high nutrition food items for breed-
understory of uninvaded forests (Collier et al. 2002). ing birds and their young. Whether a manager will need
One of the many effects of this invasive honeysuckle is to shift grassland plant composition away from nonna-
a reduction in soil surface temperature and an increase tives to natives will depend crucially on the breeding
in soil-level humidity (Watling et al. 2011b). For spe- biology of the birds he or she wishes to support and
cies such as amphibians that have tight physiological the degree to which the existing vegetation (native or
requirements, such changes can drastically affect the nonnative) supports those needs (Kennedy et al. 2009).
106 habitats in peril

Table 8.1 An overview of literature that addresses the continuum of impacts of invasive plants on wildlife habitat.

Impact Example

Changes in physical properties

Change in disturbance regime Andropogon gayanus increases fire intensity in Australia (Rossiter et al. 2003)
Change in nutrient regimes Tamarix ramosissima leaf litter decomposition changes dissolved organic matter dynam-
ics in southwestern North America streams (Kennedy and Hobbie 2003)
Change in water quality Lythrum salicaria increases tannin concentrations in freshwater ecosystems (Brown et al.
2006)
Change in nutrient regimes Litter from suite of exotics decomposes and releases nitrogen significantly faster than
natives in eastern US hardwood forests (Ashton et al. 2005)
Change in disturbance regime Extensive review of various studies involving invasives and changes in disturbance
regimes (Mack and D’Antonio 1998)
Replacing, or otherwise making inaccessible, food resources
Changes in food availability Centaurea maculosa reduces native plant diversity and, consequently, food availability for
native birds in Montana (Ortega et al. 2006)
Seasonal changes in food availability Lonicera spp. in eastern US forests provide more food for frugivorous avian species during
winter (McCusker et al. 2010)
Changes in food abundance Noxious weeds in western US rangelands shown to reduce food abundance for wildlife
(DiTomaso 2009)
Decrease in food abundance Spartina alterniflora invasions in China result in lower food abundances, thereby reduc-
ing arthropod abundance and community composition (Wu et al. 2009)
Changes in food web dynamics Elymus athericus, a nonnative grass, invasion into European salt marshes induces a tro-
phic cascade in which the dominant arthropod prey for most fish species shifts within
invaded marshes, thereby disturbing trophic function (Laffaille et al. 2005)
Altering vegetation composition so that cover is lost or degraded
Changes in vegetation composition Invasion of Lonicera spp. and Rosa multiflora results in changes to plant cover and results
in a doubling of avian nest depredation (Borgmann and Rodewald 2004)
Indirect impact of changes in plant composition Invasion of Lonicera maackii changes water chemistry that results in greater levels of
activity and / or increased surfacing behavior, exposing tadpoles to increased predation
risk (Watling et al. 2011a)
Decreases in native plant cover Eragrostis lehmanniana and Cenchrus ciliaris invade native grasslands in Texas and reduce
ground-nesting avian community abundance (Flanders et al. 2006)
Changes in vegetation cover and structure Eragrostis lehmanniana invasion into native grasslands in Arizona results in changes in
vegetation structure and cover, resulting in reductions in small mammal abundance
(Litt and Steidl 2011)
Changes in plant composition Nest predation levels increase early in breeding season for birds utilizing the nonnative
shrub Lonicera maackii (Rodewald et al. 2010)

In terms of physical processes, a common mecha- over larger areas than is common when native plants
nism by which invasive plants alter wildlife habitat is dominated (Brooks et al. 2004). The end result has
by altering natural disturbance regimes such as fire. been the transformation of these grasslands into habi-
Wildlife that live in fire-prone ecosystems typically tats that cannot support a whole suite of native wildlife
have evolved mechanisms to withstand fire events and (Brooks et al. 2004).
perhaps profit from them by taking advantage of the Litt and Steidl (2011), for example, explored the
temporary change in vegetation structure or the in- influence of a fire-tolerant invasive grass, Lehmann
crease in available food resources (DiTomaso 2009; Litt lovegrass (Eragrostis lehmanniana), on the small mam-
and Steidl 2011). Thus, invasive plants that alter the mals native to the semideserts of Arizona. They found
frequency, intensity, areal extent, or timing of fire can that each species responded differently to the structural
create conditions that disfavor native wildlife (Brooks changes that lovegrass imposed (Litt and Steidl 2011).
et al. 2004). In grasslands, the invasion of fire-tolerant Those mammals that did well in areas with dense grass
plants can lead to a grass-fire cycle whereby the habitat cover and high vegetation found lovegrass-dominated
is burned much more often, at higher intensities, and areas just as attractive as habitats that contained mostly
 the impact of invasive species on wildlife habitat 107

native grasses. However, those mammals that preferred vasive predators will consume adults, young, or eggs
areas that are sparsely vegetated decreased in the pres- of native wildlife. They will also have nonconsumptive
ence of lovegrass. Litt and Steidl (2011) attribute some effects whereby their presence adds costs to the energy
of this variation in use of the lovegrass to the strongly budget of native prey via mechanisms such as the need
divergent food resources invaded versus uninvaded for increased vigilance at the expense of foraging or
habitat offered small mammals. These results mirror using habitats that are safe from the invasive preda-
those for grassland birds, providing another example tor but suboptimal for foraging or breeding (Preisser
of how the effects of invasive plants can vary according et al. 2005).
to the requirements of native wildlife. Invasive predators are found in terrestrial island
Litt and Steidl (2011) also measured the response and continental regions, and in many freshwater and
of the native small mammals to fire, again across a gra- marine habitats (Cox and Lima 2006). The list of in-
dient of lovegrass dominance. When these grasslands vasive predators that have profoundly affected native
burned, the higher vegetation biomass and differences prey include some rather obvious examples of species
in plant species composition across a gradient of in- that have the teeth and body size to eat most anything
vaded sites tended to alter the magnitude, persistence, that they come across, a list that includes the brown
and direction of fire effects on small mammals. For tree snake (Boiga irregularis), Burmese python (Python
those mammalian species that preferred dense vege- molurus bivittatus), and red fox (Vulpes vulpes) (Harding
tation cover (i.e., dominated by lovegrass), fire dra- et al. 2002; Wiles et al. 2003; Dove et al. 2011). Others
matically reduced the suitability of a site. For species may be a bit more surprising such as wild boar (Sus
that preferred open, sparse vegetation, fire tended to scrofa), red imported fire ant (Solenopsis invicta), and
improve habitat conditions. ship rat (Rattus rattus). This latter group includes spe-
In addition to changing disturbance regimes, inva- cies that are either not typically considered ravenous
sive species have the ability to affect ecosystem prop- predators (boar) or are very small relative to their prey
erties such as nutrient cycling and water quality. Loss (ants and rats). However, it is not the size of the preda-
et al. (2012) demonstrated that the invasion of non- tor that matters as much as its evolutionary “fit” within
native earthworms into North American hardwood the ecosystem it is invading (Cox and Lima 2006).
forests has caused a multitude of changes across tro- Some of the best-known examples of extinction due
phic levels. Nonnative earthworms increase the rate to invasive species impacts come from situations where
of leaf litter decomposition, resulting in faster rates a nonnative predator establishes in an ecosystem where
of nutrient depletion by plant communities. This in the native prey are naïve to its presence (Cox and Lima
turn resulted in changes in plant communities, which 2006; Lockwood et al. 2013). More formally, these
subsequently precipitated declines in the number of hunting behaviors (and other associated features) are
ground-nesting bird species. An additional example termed predator “archetypes,” and prey naiveté results
comes from tidal wetlands of the United States. Brown from the prey having never evolved, or having lost,
et al. (2006) demonstrated that invasion by purple anti-predator defenses for the archetype of the invader
loosestrife (Lythrum salicaria) increases tannin con- (Cox and Lima 2006; Sih et al. 2010). In the context of
centrations in the invaded wetlands, which in turn has wildlife conservation, a manager should expect large
negative impacts on aquatic food webs. negative impacts of an invasive predator when there
is a mismatch between how that predator pursues its
prey and how the native prey respond to the presence
Animal Invasions
and behavior of this predator. These relationships can
Animal invaders will directly affect co-occurring na- be quite complex in detail, and there is a renewed fo-
tive wildlife by serving as their predators, their prey, or cus in ecology on understanding the mechanisms by
their competitors. The evidence for invasive animal im- which prey respond to their predators and what level of
pacts comes largely from invasive predators, although mismatch is necessary for the prey to drop in numbers
there are certainly recorded cases of impact entailing toward extinction (e.g., Banks and Dickman 2007; Salo
all of these mechanisms (Blackburn et al. 2004). In- et al. 2007). What is relatively well known, however,
108 habitats in peril

is that native prey species are more likely to become toads. Cane toads have been purposefully introduced
extinct if they are already rare or have naturally local- to a wide variety of other locations where sugarcane
ized distributions (Cox and Lima 2006). A particularly was (or is) a principle agricultural product, and there
vexing situation can arise when the invasive predators is growing evidence that in many of these places na-
are able to maintain very high densities even when they tive wildlife populations may be at risk due to ingesting
have consumed a large percentage of the individuals of cane toads (e.g., Wilson et al. 2011).
a native prey species (Courchamp et al. 1999; Roemer Finally, there is the possibility that invasive animals
et al. 2001). Such invasive predators are able to eat the will compete with native wildlife and thus reduce the
prey to extinction without themselves suffering an as- native’s survivorship or fecundity. Evidence for this ef-
sociated population decline because their numbers are fect is limited, as it is within invasions more broadly
subsidized by human food sources or by less behavior- (Gurevitch and Padilla 2004; Lockwood et al. 2013).
ally naïve (native or nonnative) alternative prey (e.g., Competition, unlike predation, involves a more gradu-
hyperpredation, Courchamp et al. 2003). ated reduction in the competing species’ survivorship
If we reverse the role of the invader and make it and reproduction. Thus, even though two species may
the novel prey item that is consumed by the native compete for available resources, the effects of this com-
wildlife predator, there is the potential for the invasive petition on either population’s abundance could be
prey to drive predator population numbers down via relatively small or fluctuate substantially across space
its unique antipredation defenses. Here again it is the and through time, especially as compared to other
mismatch between what antipredator mechanisms a mechanisms reducing abundance such as disease or
prey will demonstrate and what the prey has evolved predation (Davis 2009). Nevertheless, there are a few
to recognize and avoid. There are a variety of antipreda- examples of invasive competitors clearly impacting the
tor defenses that animals will employ (Caro and Girling populations of co-occurring wildlife. Perhaps the best
2005; Pease 2011), but in this context it is the produc- example comes from Australia where the invasive com-
tion of toxins that seem to have the most clear negative mon myna (Acridotheres tristis) aggressively excludes
effects on native predators. Perhaps the best-known ex- native parrots from using secondary tree cavities for
ample is the effect of invasive cane toads (Bufo mariuns) nesting (Pell and Tidemann 1997). A similar competi-
on native wildlife in Australia. Cane toads, which are tive effect has been posited between European starlings
native to South and Central America, exude onto their (Sturnus vulgaris) and secondary cavity-nesting birds in
skin a powerful bufogenin that is highly toxic to their North America, although the evidence for a strong ef-
predators (Phillips et al. 2003). In Australia, where the fect is equivocal (Koenig 2003).
cane toad was introduced to serve as a biocontrol agent
in sugarcane plantations, the native wildlife are naïve
Indirect Impacts
to this toxin, and for many species if an individual in-
gests a single cane toad it will perish (Shine 1995). In Invasive species can impact wildlife habitat through a
locations where the cane toad is abundant, a large per- variety of indirect pathways. Such pathways link two or
centage of native predators will consume the toad with more species via a third species (Morin 2011; fig. 8.3).
obvious effects on the consumed individual’s survival There are few fully worked examples of invasive spe-
probability. This added source of mortality has reduced cies having indirect impacts on native wildlife habitat,
the abundance of some native wildlife populations in no doubt due to the difficulty in fully documenting
Australia (Shine 2010). the complex interactions associated. Those that have
Phillips et al. (2003) suggest that snakes may be par- been documented mostly involve the influence of in-
ticularly susceptible to the ill effects of cane toads since vasive herbivores on island wildlife. Wagner and van
their feeding mechanism nearly ensures that they will Driesche (2010), for example, describe several cases
come into contact with enough of the toxin to repre- where invasive herbivores such as feral pigs, cattle,
sent a lethal dose. The feeding trials and other data col- goats, and sheep consume the host plants of specialist
lected by Phillips et al. (2003) suggest that nearly 30% insect herbivores on islands, sometimes to the point
of terrestrial Australian snakes are threatened by cane where the associated insect became extinct. Similarly,
 The ImpacT of InvasIve specIes on WIldlIfe habITaT 109

Spruce-Fir Forest, an indirect effect on wildlife is given by Rodewald et al.

Southern Appalachia
(2010), in which they find invasive shrubs (Lonicera
spp.) creating habitat in which birds experience re-
duced survival early in the breeding season.
Invasive species can also indirectly have an impact
by altering microclimates on which native wildlife
Introduced Balsam
Woolly Adelgid depend. A particularly compelling example comes
from Siderhurst et al. (2010), who documented how
the loss of eastern and Carolina hemlock trees (Tsuga
canadensis and T. caroliniana, respectively) due to the
hemlock woolly adelgid (Adelges tsugae) affected the
habitat of brook trout (Salvelinus fontinalis). Hemlock
Remnant Spruce-Fir woolly adelgids, natives of East Asia, are invasive forest
Forest
pests that suck the sap of native hemlock trees in North
America. Their effects on North American hemlock
trees are harmful enough that most hemlocks will die
within four to fifteen years after initial adelgid infes-
tation (Orwig 2002). Hemlock is a dominant canopy
tree in many North American forests, especially within
riparian zones, and dictates a suite of forest microcli-
Declines in Canopy matic conditions including the degree of shading and
and Subcanopy Birds
temperatures at ground level (Siderhurst et al. 2010).
Figure 8.3 Invasive species can affect native wildlife via Given the hemlock’s central role in these forests, their
their influence on a third species, thus producing indirect loss from the canopy should precipitate many indirect
impacts. For example, the nonnative balsam woolly adelgid impacts on native wildlife. Siderhurst et al. (2010)
has devastated spruce-fir forests in the southern Appa- investigated whether, and when, the loss of hemlock
lachian mountains of North America. The loss of spruce would increase solar radiation to streams and thus also
and fir trees substantially reduces the potential feeding increase water temperatures. They estimated that the
and breeding habitat of birds that specialize in the use of loss of hemlock trees due to adelgid damage increased
canopy and subcanopy trees. The end result has been a
light levels by over 20% in some streams in western
noted reduction in the abundance of in these birds (see
Virginia, resulting in an increase of 2°C in mean daily
Rabenold et al. 1998).
water temperatures. Such an increase will bring water
temperatures to near the maximum thermal tolerance
Barrios- Garcia and Ballari (2012) review cases where of brook trout (Butryn et al. 2013). Siderhurst et al.
wild boar have negatively impacted native wildlife, in- (2010) provide evidence that such changes may not
cluding causing a reduction in the abundance of native come to fruition over the long term, but their results
Hawaiian honeycreepers via the boar, reducing nectar- provide a clear connection between invasive forest
producing plants in forest understories through its root- insects and the abundance of recreationally impor-
ing behavior. The impact of invasive herbivores can be tant fish.
less direct, but perhaps more widespread, through their Invasive plants can increase their abundance or geo-
sometimes profound influence on vegetation cover graphical range through forming mutualistic relation-
(Rabenold et al. 1998; Courchamp et al. 2003). For ex- ships with native or nonnative animals. When these
ample, Donlan et al. (2007) provide strong evidence invasive plants have impacts on wildlife through any
that grazing by feral goats on the Galapagos Islands led of the mechanisms in table 8.1, there exists an indirect
to a substantive reduction in suitable habitat for the route of impact (fig. 8.3). The most common example
endemic and highly threatened Galapagos rail (Lateral- in the literature is the association between birds and
lus spilonotus). One example of an invasive plant having invasive plants where the bird provides pollination
110 habitats in peril

and seed dispersal services to the invader (e.g., Lafleur studies of facilitation involving nonnative species that
et al. 2007). Aslan and Rejmánek (2010) report that “close the loop” and unequivocally show how the es-
nearly half of all their citizen science observations of tablishment of a mutualism will indirectly negatively
bird-invasive plant interactions involved birds eating affect local wildlife. Thus, for example, although birds
the seeds or fruit of the plants and thus likely serving commonly eat the fleshy fruits of many invasive plants,
as a dispersal mutualist for the invader. Best and Arc- it is not clear the extent to which the increased disper-
ese (2009) show that selective grazing by Canada geese sal of such a plant is serving to decrease the abundance
(Branta canadensis) on a suite of nonnative grasses had of any native wildlife species of concern.
a net positive effect on the grasses themselves. Graz-
ing pressure created growing conditions that favored
Synergisms and Synthesis
the production of nonnative grass stems (i.e., reduced
litter accumulation), and the seeds ingested by graz- Wildlife populations are principally driven toward ex-
ing geese were dispersed into locations suitable for the tinction through the loss, fragmentation, and degra-
grasses to establish (Best and Arcese 2009). Mammals dation of their habitat (Damschen et al. 2006). As we
can also form mutualistic interactions with nonnative reviewed previously, invasive species can heap yet more
plants, thus increasing their dispersal capabilities (Da- negative influences on the survivorship and reproduc-
vis et al. 2010). Examples of mutualistic interactions tion of wildlife populations. The degree to which in-
between mammals and nonnative plants include the vasive species are prominent threatening factors in
much-maligned rat that can also directly produce nega- wildlife declines is the subject of debate (e.g., Gure-
tive impacts on native wildlife (Shiels and Drake 2011). vitch and Padilla 2004 vs. Clavero and Garcia-Berthou
Given the volume with which nonnative species 2005). However, there are many examples of wildlife
are transported and released worldwide, there is the having become extinct where the negative impacts of
distinct possibility that an ecosystem will harbor sev- invasive species are clear and significant (Miller et al.
eral invasive species (Lockwood et al. 2013). In such 1989; Clavero and Garcia-Berthou 2005; Wagner and
situations, there is the possibility that a suite of non- Van Driesche 2010). But to what extent do habitat loss
native species will form mutualistic networks that to- and fragmentation interact with species invasions and
gether impose higher levels of impact than each could lead to the loss of native species? Despite the two forces
achieve individually (Simberloff and Von Holle 1999). playing clear roles in biodiversity loss (Hobbs 2000),
When such networks materialize, the end result can they are very often studied independently (Didham
be a transformation of the native ecosystem, or an “in- et al. 2007; Vilà and Ibanez 2011). In a comprehensive
vasional meltdown,” where the end result is the loss of literature survey by Didham et al. (2007), they found
native species (Simberloff and Von Holle 1999; Sim- that of the nearly 12,000 studies of land-use change
berloff 2006). The evidence for invasion meltdowns (e.g., fragmentation) and over 3,500 studies of species
is limited, although there are several compelling case invasions (published between 2002 and 2007), only 1%
studies (Simberloff 2006). However, no example di- considered both simultaneously and 0.03% explicitly
rectly ties the strong facilitation between two nonna- considered the interaction of the two.
tive species to a decrease in the survivorship or fecun- What we know from studies that do consider both
dity of co-occuring native wildlife (Simberloff 2006). in tandem is that the land uses that surround habitat
Although not a direct example of invasional meltdown, fragments play a large role in the number and types of
the introduction of new parasites / diseases by intro- invasive plants that exist within those fragments (e.g.,
duced animals could have far-reaching effects on na- Hobbs 2000; Lindenmayer and McCarthy 2001). In a
tive biota (Prenter et al. 2004). A gripping example of review of empirical evidence linking landscape context
the devastation that introduced parasites can have on with invasive plants, Vilà and Ibanez (2011) condense
native wildlife is seen in the marked reduction in en- this literature into a few solid generalizations. Well
demic Hawaiian aviafauna caused by the introduction supported is the positive relationship between the de-
of avian malaria (van Riper III et al. 1986). gree of habitat fragmentation and the presence of in-
In fact, from our literature review, there are no vasive plants. This result is driven to a large degree by
 the impact of invasive species on wildlife habitat 111

the increased presence of invasive plants near the edge et al. 2007). Existing evidence provides some indication
of fragments. Additionally, there is a positive relation- of how the two forces will combine to determine the
ship between the percentage of urbanized land around fate of native wildlife, but clearly much more remains
the habitat fragment and the incidence of invasive to be explored. Such explorations must do a better job
plants within that fragment. Road density, frequency of directly tying landscape changes (e.g., fragmenta-
of road use, and the existence of road improvements tion) to the numerical abundance and geographical
near a habitat fragment tend to increase the number range size of the invasive species and documenting
of invasive plants species in the fragment. In short, how such landscape changes influence the per capita
the types and relative abundance of land uses around effect of invasive species on native species (Didham
habitat fragments dictate to a great extent the degree et al. 2007). These three factors (the abundance, range
to which that fragment harbors invasive plants (Vilà size, and per capita effect of the invader) have been
and Ibanez 2011). The degree to which we can gener- suggested to be the key components of measuring im-
alize these specific results to the presence of invasive pact on native species (see previous discussion, Parker
animals is uncertain, although several examples show et al. 1999). In the context of wildlife management and
that the land uses around a habitat fragment will in- conservation, this is the end measure that is vital for us
fluence the abundance and impact of invasive animals to understand in responding to crises in wildlife popu-
within that fragment (e.g., Donovan et al. 1997; Tewks- lations. Making these connections explicit clarifies the
bury et al. 2006). impact of our research and provides detailed guidance
The work of Vilà and Ibanez (2011) also highlights to those tasked with managing habitats for our native
understudied elements of the interplay between inva- wildlife.
sive species and habitat loss or fragmentation. For ex-
ample, it is not clear how often habitat corridors aid
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“Extracts of the Invasive Shrub Lonicera maackii Increase
 RESEARCH AND
PA R T I I I •
CONSERVATION
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 9 Thoughts on Models
and Prediction
Bret A. Collier and
Douglas H. Johnson

W ildlife managers and conservationists face

many challenges, such as convincing the public
that wildlife professionals occasionally do know more
derstanding their status and predicting changes. That
knowledge is gained by studying the habitats they use,
as well as when and how they use them. One of the
about wildlife populations than the average citizen, en- greatest challenges wildlife managers must address is
couraging youth to appreciate some things that are not recognizing how the status and sustainability of wild-
electronic, and persuading elected officials to think be- life populations will change as the Earth’s ecosystems
yond the immediate and consider long-term sustainabil- continue in flux. It is a simple model that predicts com-
ity. These challenges would exist even in a stable world. plete habitat loss will be deleterious to species that are
However, the world is very dynamic. The weather is restricted to that habitat. Assessing responses to other
fickle, the climate changes in long-term cycles, and hu- habitat changes requires more complex models and in-
mans now influence climate in unprecedented manners. vokes greater uncertainty. Numerous models have been
The human population grows at exponential rates, its developed for a variety of objectives and a wide range
impact magnified by increasing per capita consumption of species, but their record for accurate prediction is
of resources. And the world shrinks; a disease that once spotty. As ecologists strive to assess current and future
might have been confined to a small rural village in Asia impacts of environmental change on wildlife resources
now may become a worldwide epidemic in a matter of with only limited knowledge of the system, what is the
weeks. Well-intentioned legislation can effect enormous manager to do? In this chapter, there will be no discus-
but unanticipated changes in land use over millions sion of model creation or application, as a vast array of
of hectares across the nation, and changes in energy literature is available on that topic (Shenk and Franklin
prices can potentially have significant environmental 2001; Williams et al. 2002; Millspaugh and Thompson
consequences. How is a wildlife manager to deal with 2009). Rather, this chapter will discuss issues outside
so many major changes well beyond his or her control? the approach of “collect data, analyze, and explain” that
This chapter will not resolve these difficulties. In- typifies the standard approach to prediction in wildlife
stead, we contemplate how wildlife managers and con- ecology and focus on the interplay among modeling,
servationists might think about future planning and monitoring, and careful thinking.
become more proactive relative to wildlife populations
and their habitats.
Known Unknowns
There are things that we think we know. We think we
Thoughts on Models and Prediction
know how to measure habitat, or at least the aspects of
Knowledge of where and when animals occur, and their habitat that we believe are relevant to an animal. We
success in surviving and reproducing, is crucial to un- think we know what affects survival, or at least how to
118 research and conservation

identify the primary biotic and abiotic factors that af- Consider some wildlife examples. If a manager
fect survival. There are many things that we know are wanted to maintain harvest of a species at some level,
unknowns, and wildlife researchers use models to turn a model for control would be appropriate; the output
known unknowns into known knowns, such that we (desired harvest) is known and the input (harvest regu-
know the extent to which we know. lations) need to be determined; the necessary model
would relate regulations to actual harvest. If wildlife
managers were concerned about how grassland birds
The Role of Modeling
might be affected by the elimination of the Conserva-
The need for accurate predictions of ecological out- tion Reserve Program (CRP) and a return of CRP lands
comes in response to environmental changes has to cultivation, they would find useful a model for the
heightened interest in the application of various mod- prediction of densities of grassland birds in various habi-
els. One useful definition of a model is that an object tats (CRP and cultivated lands); inputs would be areas
A* is a model of an object A if one can use A* to ad- of land in each land-use category before and after the
dress relevant questions about A (Minsky 1968). Thus, curtailment of CRP, and outputs would be numbers of
models are used to reduce the infinite-dimensional grassland birds before and after. A scientist interested
set of ecological components and their interactions in the effects of an ingested chemical on egg produc-
to manageable pieces relevant to the specific topic of tion by a bird species would seek to develop a model
interest (Muller et al. 2011). Components in a model for understanding that relates measurements of egg pro-
are linked, typically via mathematical relationships, duction to the amount of the chemical ingested for a
to describe characteristics of the system in an effort (large) number of birds.
to mimic or predict the behavior of the system being Models have many uses in wildlife-habitat ecology;
modeled. Thus, models in wildlife ecology help us to models can describe functional relationships among
reduce the complexity of biological systems by focus- ecological variables, quantify those relationships, and
ing on only components pertinent to the immediate change as additional data are collected, and models
need (Shenk and Franklin 2001). Note that we are not can be used to predict future scenarios. Models rep-
discussing purely statistical models (e.g., those used in resent a way for us to synthesize knowledge and ex-
linear regression) per se but rather a more overarch- plain a system. Further, models provide scientists with
ing framework focusing on some system. For example, the ability evaluate available knowledge, identify areas
population models help us understand how the pro- where information is lacking, formulate hypotheses,
cesses of births, deaths, immigration, and emigration design research focused on filling knowledge gaps, in-
influence population size, and models of an ecosystem tegrate information from a variety of sources, and serve
can be developed to mimic the dynamic interactions as a means of communicating research results (John-
among its plants, animals, and abiotic components. son 2001).
It is convenient to view a system as receiving input
from some source and responding with an output. A
Designing a Model
model of that system can be constructed for one of
three basic objectives, one each aligned with input, What type of modeling approach is best? Wildlife-habitat
model, and output. Prediction involves determining the models range widely, from models for species distribu-
output from the system based on some prescribed in- tions (Elith et al. 2006), models of fine-scale habitat
put and the assumed model. Understanding the system suitability (US Fish and Wildlife Service 1981), to mod-
is based on inferring relationships between observed els focused on response to hypothetical or predicted
inputs and outputs. Control involves manipulating in- perturbations (Boyce 1992; Hanski 1998; DeAngelis
puts to the system to achieve desired outputs. Clearly, and Mooij 2005). Beck and Suring (2009) identified
however, a model built for understanding a system, if it forty frameworks for modeling wildlife-habitat re-
adequately mimics the relationships in the real system, lationships, which vary in model structure, input re-
will be useful for both prediction and control. quirements, and output variables. Thus, we refer the
 thoughts on models and prediction 119

interested reader to Johnson (2001), who states, “A lation parameters encompassing the range of the data
model has value if it provides better insight, predic- (both temporally and spatially) needed for modeling
tions, or control than would have been available with- most systems of interest. In these cases, approaches
out the model” (Johnson 2001, 113). Relatively few such as simulation can be used to identify the relative
models, however, have been confronted with a truly influence of model elements and identify where future
independent set of data to determine their actual value data collection should occur, given that the posited
for their intended purpose. relationships are accurate. Thus, if a model is simple,
Not surprisingly, a wide variety of recommendations slightly biased, but useful, that is better than a model
for wildlife-habitat modeling have been proffered. Most that is complex, unbiased, but not useful. Models that
of these suggestions have been discussed elsewhere in are slightly biased for most species yet useful for all
significant detail, and all of these suggestions are some- species represent a viable modeling alternative versus
how linked in model development; however, here we process- and species-specific models, as data can be
attempt to distill those discussions down to the follow- acquired and shared across system. Models with a sig-
ing set of general guidelines for model development. nificant number of integrated parts often provide good
First, models must have precise, well-defined ques- predictions when focused within sample periods (pe-
tions that focus on key conservation-relevant outputs, riods in which the range of data was collected) but can
while limiting the number of assumptions used. As trend toward unreliability as time and space increase
models represent a construct of a system that we are (Bunnell 1989; Araujo and Rahbek 2006; Beutel et al.
trying to understand, they should clearly define what 2006). We note that, while rarely discussed, data of-
ecological variables are relevant to the modeling ex- ten used for comparisons to modeled predictions are
ercise (Müller et al. 2011). Models can rarely answer frequently model based (e.g., density / abundance data
why questions, typically focusing on how questions, but corrected for imperfect detection, capture-recapture
often asking what if, as most questions in ecology are estimates of population growth) and are thus uncer-
focused on resolving how a system operates relative to tain. Yet, estimates are regularly used to evaluate mod-
system inputs and our preconceptions about the struc- els, often without addressing the inherent uncertainty
ture of the system. Thus, approaching development in these estimates, which may impact the accuracy of
of any model should be (somewhat obviously) based model evaluations. Thus, there exists a modeling loop,
on a well- defined question that relies on reasonable where model evaluation is dependent on modeled
expectations regarding how the system under study predictions.
works and the question(s) that the researcher wants Models and their metrics do not ever represent truth
to evaluate. in and of themselves. Model output that is purported
Ecological systems are inherently complex, thus we to track the observed fluctuation in a population is no
cannot possibly define all of the interacting compo- more representative of truth than one whose predic-
nents (although given the proliferation of model sizes, tions are uncoupled from observed fluctuations. Mod-
some have tried), nor can we realistically parameterize els don’t give answers; rather, they provide estimates
those components or identify the relationships of those of the magnitude of difference in model predictions
components. Ecological and statistical complexity in based on the structure and variation in a model. Thus,
models should be treated differently in that a model models can and should be evaluated at different levels
could have thousands of interacting values (ecologi- of the system. As most ecological systems are hierarchi-
cally complex), yet relationships between model pa- cal in nature, the level of aggregation at which a model
rameters could be deterministic in nature (statistically is evaluated will impact model veracity and usefulness.
simple). It is important to realize that models are only Model-based aggregated parameters (e.g., population
as good as the data and assumptions on which they are growth rates) are more likely to be robust to moder-
based. All models rely on data and have assumptions ate variation in inputs; intermediate parameters (e.g.,
about data—how it was collected, how it is structured, per-capita recruitment) are more likely to be impacted
and so forth. We often lack adequate estimates of popu- by the vagaries of internal model parameters and the
120 research and conservation

mechanistic relationships between those parameters;

whereas input parameters (annual survival) will be
driven by both the posited statistical model and the
amount of data used in the estimation routine. Thus,
model robustness, in many cases, may rely more on
structure and less on accuracy of input values.
Most relationships among variables in real systems
are nonlinear; however relationships frequently pos-
ited between model parameters are regularly linear.
How can this incompatibility be reconciled, beyond
concluding that all models are wrong and giving up on Figure 9.1 The nonlinear response of a species to an envi-
them? The answer is that most nonlinear relationships ronmental gradient X.
are “close enough” to linear to be useful, as long as in-
put variables do not vary “too much.” Of course, “close
enough” and “too much” are vague terms, the first Doing so will require careful monitoring of actual
depending on the objectives of the modeling, and the events in comparison to predictions.
second providing a cautionary note about extrapolating
beyond the range of the data used to develop the model.
Unknown Unknowns
Space is curved, while Euclidean geometry assumes it is
not. Therefore Euclidean geometry is invalid (wrong), Modeling is useful if something is known about the
but it still is useful as a very close approximation to the system and the relevant variables. For example, X af-
truth. The nonlinearity is so slight at distances usually fects Y, but you don’t have complete knowledge, such as
measured on Earth that it can be ignored, but that is exactly how X affects Y. You know what you don’t know.
not so true with cosmological distances. Nonlineari- In this situation, one can propose and formulate vari-
ties among other relationships may not be so obvious. ous plausible relationships between X and Y and gather
Consider a hypothetical species whose abundance re- data, use extant data, or seek expert opinion about de-
sponds to some habitat feature, denoted by X, as shown tails of the relationship and the amount of uncertainty
in figure 9.1. A reasonable example might be the abun- we should expect in the relationship. What, however,
dance of grasshopper sparrows (Ammodramus savanna- if you have no idea about the form of the relationship
rum) in relation to biomass of live vegetation. A study or, even worse, what the relevant variables are? That is,
conducted in a semiarid region (A in the figure) likely you don’t know what you don’t know—the unknown
would find that the species is more common when unknowns situation. Modeling now is not so straight-
vegetation is thicker. Someone studying grasshopper forward. What can be done?
sparrows in a more mesic region (C in the figure) would Possibly one could look backward in time. Imag-
reach the opposite conclusion, whereas a study in the ine the world, say, fifty years ago, but knowing what is
intermediate region (B in the figure) might detect no known now about what has transpired during that half
effect at all of vegetation biomass on grasshopper spar- century. Is it possible to identify events that we now
row abundance. Extrapolating results from any of the realize have had a huge impact but were unappreciated
three studies to regions with different vegetation would a half-century ago?
be misleading. Viewing figure 9.1 makes this last con- For example, the world population was roughly three
clusion obvious, but no researcher conducting a single billion humans—a lot of people, but the exponential
study would have the benefit of such an opportunity. nature of its growth in developing countries was only
The bottom line is that using models for prediction, beginning to become apparent. Some authorities, nota-
which always involves extrapolation over time, will re- bly Paul Ehrlich and Ann Ehrlich (The Population Bomb)
quire extrapolation beyond the range of data on which identified the patterns and described scenarios of ca-
the models were based, so it is especially important to lamities occurring within a decade or so. Such alarmists
determine if and when the models are no longer useful. were widely ignored, especially when the anticipated
 thoughts on models and prediction 121

catastrophes failed to occur as soon as predicted. None- ing management, regardless of whether or not a priori
theless, the problem of overpopulation continues today. hypotheses are posited.
We suspect that virtually every major event that Obviously, not everything can be monitored. How-
has occurred had been anticipated by someone much ever, we think that decisions about what to monitor
earlier. The difficulty for managers and those mak- should be based on at least the following consid-
ing decisions arises in part because numerous other erations:
events have been predicted but had not actually oc-
curred. Who are the true prophets, who are the false 1. Importance of the resource: This criterion seems
prophets, and how can we tell them apart? Instead of trivial as all resources are important to some-
seeking specific predictions about the future, perhaps thing, but it covers a multitude of attributes. Any
we should be seeking environmental leading indicators, resource that has major influence on a number
rescuing the term from the purely economics realm. of other resources qualifies here—such as krill,
What might useful environmental leading indicators representing one of the lowest trophic levels in
be? Certainly, most organisms respond directly to vari- oceans, or freshwater, a resource regularly over-
ous weather features, so climate should be assessed. looked but on which both humans and wildlife
And humans cause enormous changes to wildlife and directly depend.
their habitats, so human numbers, their geographical 2. Susceptibility: Which resources are deemed most
distribution, consumption patterns, and other behav- susceptible? Certain forests are more susceptible
iors somehow should be monitored. From a food web to fire than others. Some wetlands are more read-
viewpoint, the top predators in a system rely on other ily drained than others. Some grasslands are more
species in the system and might well reflect the condi- subject to encroachment by invasive species than
tion of those species. Because water flows downhill, it others. Habitat specialists likely are more suscep-
should be straightforward to identify locations where tible than generalists because they tolerate a more
conditions reflect conditions throughout the entirety narrow range of conditions and would be more
of the upstream area. likely to be vulnerable when conditions change.
These ideas lead to the notion of monitoring with- 3. Potential rate of change: Prairie wetlands, as one
out specific hypotheses in mind, or what has been dis- trivial example, are much more dynamic than rock
paraged by some as “surveillance monitoring.” Nichols formations. Locations with a slow rate of change—
and colleagues (Yoccoz et al. 2001; Nichols and Wil- such as mature forests, unless significant perturba-
liams 2006) have argued that monitoring is useful only tions occur—likely require less intensive monitor-
when there is a specific hypothesis to test or a specific ing than, say, a coastal wetland.
management program to inform. They criticize surveil- 4. Ease, objectivity, and repeatability: Monitoring is
lance monitoring, which they defined as monitoring defined as information collected over time and
not guided by a priori hypotheses and their correspond- space. Thus, methodological approaches must be
ing models (Nichols and Williams 2006). Certainly, if objective (unbiased sampling, for instance) and
any management action is planned or a hypothesis allow for repeatability of the analytical methods
needs to be tested, one should see what happens after such that accurate results can be obtained. For in-
the action or during the experiment. But many impor- stance, one reason why singing birds are the object
tant events are neither planned nor anticipated. One of so many monitoring programs is because they
can think of the Deepwater Horizon oil spill and its are much more readily amenable to current sam-
consequences to natural resources or the Exxon Valdez, pling and surveying methods than, say, crepuscular
Chernobyl, or the Japanese earthquake and subsequent mammals or karst salamanders.
tsunami. One cannot go to a disaster site and learn
what it was like before the disaster; you need to have
Control over Events
had a monitoring or analogous program in place well
before the event. Thus, while not hypothesis driven, The basic approach used to further understanding of
surveillance monitoring can be a critical for inform- wildlife-habitat relationships is to identify locations
122 research and conservation

where individuals are found, collect descriptive data on ecosystem-level concerns could be elevated or lowered
both the individuals and locations, analyze those data proactively as increased data is made available or as
and develop an appropriate model, make predictions conditions change over time.
either for places other than the study area or for a later
time period, and corroborate those results against the
What’s in the Future?
new data as evidence for a model’s utility. The process
of observation and inference represents the workhorse So, how do wildlife scientists and managers become
of wildlife science, where virtually every research out- more proactive? How should they respond to antici-
put is somehow dependent on past data collected. In- pated or predicted changes in the environment when
duction, or generalizing the specific, is then used to often we can only speculate as to the causal relation-
make predictions of future conditions. This process of ships or appropriate response metrics? Despite the
evidence- (data) based science has served us well, and need, truly proactive wildlife management activities
this reliance on previously gathered data comes with a are rare; wildlife managers are better categorized as
cost, in that wildlife sciences are consistently reactive, first responders than as actual managers, as they have
rather than proactive, which limits the actual ability very little control or even the ability to predict major
we have to manage or manipulate ecological systems. events that will influence wildlife populations at large
Speculating what ecological calamities may occur scales. This is regretful as we spend an inordinate
is often met with significant skepticism and resistance, amount of time trying to use models to predict what is
regardless of the potential benefits of looking ahead going to happen after we respond to an issue; we per-
(McCann et al. 2006). Cynicism about such specula- haps do not spend enough time trying to construct and
tion has several foundations. One is the issue of false drive what should happen. Regardless, in some cases,
positives; many disasters that were anticipated did researchers have had the foresight to synthesize pre-
not occur. A second basis for cynicism is the fact that vious research to identify potential overarching issues
speculation is not evidence based and hence may not based on past patterns, such as the reduction of the
represent strong inference (Platt 1964). A third basis ozone atmospheric layer due to human chemical use
is innate conservatism and inertia; it is easier to con- (Crutzen and Ehhalt 1977) or the plight of grassland
tinue doing what we have been doing rather than to birds in the United States (Droege and Sauer 1994;
change because of something that might—but might Knopf 1994). Consider two examples of the difficulties
not—happen. Nonetheless, the benefit of strategically of proactive responses to management issues: the Con-
identifying and addressing imminent, yet perhaps un- servation Reserve Program and the Deepwater Horizon
recognized, issues could be immeasurable to both cur- oil spill, both of which have had impacts on wildlife and
rent and future management of wildlife and wildlife wildlife habitats.
habitats (Sutherland et al. 2008). Horizon scanning is
the process of identifying how a system may be at risk
The Conservation Reserve Program
in the near or intermediate future relative to a suite
of potential issues (disturbance, succession, climate First, the Conservation Reserve Program (CRP) nicely
variation, etc.) and prioritizing the issues, whether or illustrates how the wildlife community responded
not they currently occur, that deserve increased atten- fairly effectively to a major change in the landscape.
tion through research, monitoring, modeling, or evalu- CRP was a component of the 1985 Food Security Act
ation (Sutherland et al. 2008). Regular application of (also known as the farm bill) that allowed owners of
horizon scanning would allow for identification of is- highly erodible cropland to plant it to grasses and forbs,
sues currently affecting an ecological system but also leave it idle except for necessary maintenance, and re-
allow consideration of future conditions, not yet oc- ceive annual payments for the period of the contract,
curring, that may also impact the system. Regular use usually ten or fifteen years. At that time, the United
by wildlife ecologists would allow for both surveillance States had surpluses of major crops, and the Depart-
and hypothesis-driven monitoring strategies to be ap- ment of Agriculture was seeking a way to reduce those
plied or implemented. Additionally, both species- and surpluses. The primary purpose of the CRP was to
 thoughts on models and prediction 123

achieve that goal and thereby maintain adequately high Although the program’s future is in doubt, as enroll-
prices for commodities. By focusing the program on ment is expected to plummet with the currently high
highly erodible lands, the program also addressed its commodity prices, for decades CRP did provide habi-
secondary objective of reducing soil erosion from wind tat for an abundance of game and nongame wildlife,
and water. A third objective, seemingly almost an af- in part because wildlife scientists were able to identify
terthought, was to restore habitat for fish and wildlife. its promise early, gather supportive information, and
The program was immensely popular with producers; make informed recommendations to policy makers.
the area enrolled in CRP climbed from 1,929,064 acres
in 1986 to 32,522,380 acres in 1990 and remained
The Deepwater Horizon Oil Spill
above 30 million acres until 2012, when it dipped
slightly (http: // www.fsa.usda.gov / Internet / FSA_File A recent large oil spill, the Deepwater Horizon offshore
/ historystate8612.xls, accessed March 6, 2013). drilling rig in the Gulf of Mexico, offers a compelling
Much of the area enrolled in CRP was in the Great example of seriously deficient responses to events with
Plains and Midwest, and the extensive conversion of huge potential consequences to wildlife populations
cropland to perennial herbaceous cover resulted in a and habitats. Although the event resulted from human
massive habitat alteration at landscape levels. Expecta- error and could have been avoided, it would be wildly
tions within the wildlife community were that these optimistic to assume that similar events would never
grasslands would support far more birds and other wild- occur. Here, we focus on the assessment of ecological
life than did the croplands they replaced (reviewed by damage caused by the Deepwater Horizon blowout.
Higgins et al. 1988). Wildlife researchers responded to President Barack Obama formed the National Com-
this landscape change by initiating a number of studies mission on the BP Deepwater Horizon Oil Spill and
to determine how wildlife, especially birds, responded. Offshore Drilling to assess the overall response to the
Studies of waterfowl nesting in CRP fields began in event; the following information was derived from the
1989 in the Dakotas and Minnesota (Kantrud 1993; Commission’s report (National Commission 2011).
Luttschwager et al. 1994), and CRP turned out to be a The Clean Water Act mandates the development
boon for nesting waterfowl, adding about two million of a National Contingency Plan, which prescribes the
ducks per year to the fall flight. Studies on grassland nationwide response structure for oil spills (265–66).
birds were initiated in 1990 in the northern prairies Regional Response Teams, co-chaired by the US Coast
(Johnson and Schwartz 1993) and in several Midwest- Guard and EPA, include representatives from state and
ern states in 1991 (Best et al. 1997). To date, numerous other federal agencies and are tasked with developing
grassland species have benefited from the conversion of regional contingency plans as well as preauthorization
cropland to perennial cover (e.g., Johnson and Igl 1995; protocols for certain response strategies. Area commit-
Johnson 2005). tees, which develop area contingency plans, similarly
Because of the attention focused on wildlife ben- include federal and state representatives but are led by
efits of CRP, wildlife received greater priority in sub- the Coast Guard. The area contingency plans are the
sequent farm bills. Modifications based on the obvious most specific and relied-upon during the response to a
benefits were made in amendments to the farm bill spill. In addition, the industry had developed its own
in 1990 and 1995 to enhance consideration of wild- spill response plans (266). The Commission noted that
life (Heard 2000). In particular, environmental ben- industry plans were sent only to the Minerals Manage-
efits were given consideration equal to soil and water ment Service, where few of them received environ-
conservation. A new practice was adopted by the Farm mental review. BP’s response plan was “embarrassing”
Service Agency specifically to benefit nesting water- (133). It listed as a wildlife expert on whom BP would
fowl; the Duck Nesting Habitat Initiative prioritized rely an individual who had died several years before BP
lands in areas with abundant waterfowl and allowed submitted its plan. Among species of concern in case of
greater areas of upland to be enrolled (http: // www a Gulf oil spill, the plan listed seals and walruses, which
.ducksunlimited .org / conservation / habitat / new- crp never occur in the Gulf.
-practice-emphasizes-ducks, accessed March 21, 2013). Because of poor planning before the spill, the re-
124 research and conservation

sponse to it was chaotic. Disagreement among fed- the marine ecosystem. Such research depends on
eral agencies, between federal and state agencies, and sampling, measurements, and investigations that
between industry and public agencies contributed to can be accomplished only during and right after
this problem, as did political posturing and bureau- the spill” (184).
cratic inertia. Further, independent scientific research
related to the spill was hindered by the lack of timely The bottom line is that while this specific incident
access to the site and delayed funding (174). Specifi- could not have been predicted, it is reasonable to as-
cally relevant to the topic of this chapter is the lack sume that some major oil spill would occur at some
of “pretreatment” data. Regarding marine mammals, time in the future. With that anticipation, it would have
Tim Ragen, executive director of the federal Marine been prudent to first develop a systematic monitoring
Mammal Commission, testified before a House of Rep- program to establish an understanding of the condition
resentatives subcommittee and noted, “Unfortunately, of wildlife (and other ecological components) under
the scientific foundation for evaluating the potential natural (pre-“treatment”) conditions and, second, de-
effects of the Deepwater Horizon spill on many marine velop a plan to rapidly implement additional research
mammals inhabiting the Gulf is weak” (181). The Com- after an incident where effects from the incident are
mission also stated: likely and at comparison sites unaffected by the inci-
dent. Such a plan should have a direct and short chain
• “Unfortunately, comprehensive data on conditions of command so that it can be effectively implemented
before the spill—the natural ‘status quo ante’ from on very short notice.
the shoreline to the deepwater Gulf—were gener-
ally lacking” (174).
Conclusion
• “A typical damage assessment can take years.
Two sets of determinations—one concerning the Unlike many of the so-called “hard sciences,” wildlife
baseline conditions against which damages to each ecology rarely can address major questions with de-
species or habitat will be assessed and another signed experiments. The discipline largely relies on ob-
concerning the quantification of those damages— servational studies with severely limited opportunities
are particularly difficult and consequential in for control, randomization, and replication (Shaffer
terms of the overall results” (183–84). and Johnson 2008). Accordingly, inferring causation
• “Without well-established baseline conditions, is a much more daunting task. Moreover, major eco-
there can be inaccurate quantification of damages logical treatments are rarely applied by scientists. They
or required restoration. Given that the ecologi- may be natural events, such as a hurricane or earth-
cal baseline can vary seasonally, annually, and quake. They may be natural events with effects that
over much longer time scales, it can be difficult were exacerbated by human actions, such as flooding.
to pinpoint the exact condition of an ecosystem They may be purely results of human error, such as oil
prior to a spill. Because long-term historical data spills or wildfires. Or they may be consequences of ac-
are often nonexistent or discontinuous, natural tions taken largely for other purposes, such as the farm
resource trustees are likely to be disadvantaged by bill or reservoir dam building. The question is: “How
a lack of sufficient information to fully character- can we capitalize on such events to learn about the af-
ize the condition of relevant ecosystems prior to fected system so that we may better predict, respond to,
the incident in question” (184). or mitigate similar events in the future?” Three scenar-
• “[F]unding for academic and other scientists in ios are proposed depending on whether events can be
the days and weeks immediately after the spill predicted, are partly predictable, or are unpredictable.
was limited. As a result, the nation lost a fleeting A predictable event is one for which the action as
opportunity to maximize scientific understand- well as its location and timing are known. Such events
ing of how oil spills—particularly in the deep could include the construction or removal of a dam on
ocean—adversely affect individual organisms and a river, restoration of a particular plot of land, or preda-
 thoughts on models and prediction 125

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 10
Michael K. Schwartz,
Manage Habitat,
Monitor Species
Jamie S. Sanderlin, and
William M. Block

M onitoring is the collection of data over time.

We monitor many things: temperatures at local
weather stations, daily changes in sea level along the
with the associated opportunity costs and whether
monitoring data could be collected more efficiently.
Choosing exactly what to monitor in wildlife biol-
coastline, annual prevalence of specific diseases, sun- ogy is no trivial matter. For example, should we moni-
spot cycles, unemployment rates, inflation, commodity tor populations, habitats, or both? If populations are
futures—the list is virtually endless. In wildlife biology, monitored, what is the appropriate state variable(s)
we also conduct a lot of monitoring, most commonly to measure: abundance, density, presence, age- class
measuring state variables that either directly or in- structure, genetic variation or structure, survival, re-
directly relate to the size and areal extent of wildlife production, movement? At what scale and resolution
populations. Most of these activities, both in wildlife should managers monitor these variables? Can we use
biology and more generally, are not strongly targeted surrogate measures for monitoring variables of inter-
at answering specific questions. Rather, they reflect est that are either impossible or extremely expensive
measurements of things we consider to be of general to obtain on their own?
interest, but which, as data streams temporally extend, In this review, we focus on recent thinking concern-
often produce a variety of novel insights. For instance, ing the role of monitoring in wildlife science, choice
no one could have predicted the benefits of long-term, of state variables, and the degree to which monitoring
fixed local weather stations or atmospheric CO2 mea- can and should be designed and targeted to answer spe-
surements in evaluating climate change, a purpose cific questions of management significance. Lastly, we
for which neither data stream was initially intended discuss the paradigm of linking monitoring to risk as-
(Lovett et al. 2007; Harris 2010). sessment, decision analysis, and adaptive management.
The general value of monitoring is not debated:
without a time series of previously collected data we
Why Monitor?
are limited in our ability to evaluate current conditions
or make projections into the future. However, the col- We start this review by asking a simple question: why
lection and maintenance of data across extensive time monitor at all? Although this may seem like a simple
and space that characterize an effective monitoring question to address, it is not. First, monitoring must
program are both expensive and difficult. Maintaining be differentiated from assessments and research. We
both resources and infrastructure to regularly collect define monitoring as the measuring of a parameter of
and archive information with established methods and management interest (e.g., abundance of a species),
quality controls is challenging. Thus, it is reasonable to or a surrogate of this parameter, over time (Schwartz
ask not whether these data are useful (they invariably et al. 2007). The critical part of this definition is the
are) but rather how useful they are when compared temporal element. Without considering time, we are
 manage habitat, monitor species 129

not monitoring but rather are conducting an assess- explanation of the mechanism(s) behind the decline. If
ment. In essence by our definition, when multiple our initial goal was to identify what causes elk declines
assessments are combined it becomes a monitoring in an area, this monitoring effort alone, even if success-
effort. Assessments can be very valuable in setting ful in detecting a decline, will have failed as our effort
management objectives. In fact, an assessment is one didn’t collect data in a framework that could determine
of the first steps for initiating a monitoring program. why the decline occurred (see adaptive management
Assessments can be particularly useful if we ask if a in the following discussion). Thus, monitoring may
state variable of interest is meeting a certain thresh- be necessary to track trends in wildlife populations of
old. For example, instead of asking if estimates of elk interest, but it may not be sufficient for us to have the
(Cervus canadensis) abundance (abundance being the information necessary to change a population trajec-
state variable of interest) are changing over time, we tory (e.g., a decline).
can ask: do we have a minimum of number of elk in a We could argue that all “important” species should
particular area after a given management action? The be monitored, and if we see declines, we could sub-
latter question is much simpler and less expensive to sequently address the cause. However, this is much
accomplish. We may even tie these assessments to trig- like the monitoring approach that the US Forest Ser-
gers to make the situation more powerful. For instance, vice (USFS), one of the world’s largest land owners,
we could state (a priori to the assessment) that if we do adopted from their 1982 planning regulations. Some
not have a minimum number of elk in a particular area, of the biggest problems associated with the monitor-
post a management action, then we will take particu- ing requirements from the 1982 planning regulations
lar and precise follow-up actions. These assessments were that monitoring a large number of species was
often are not representative measures of the state vari- too expensive and thus often not conducted, leading to
able, nor do they need to be. For example, surveys can litigation. Furthermore, these monitoring efforts were
concentrate on those areas and populations most likely seldom followed by any attempt to understand mecha-
to change or where detecting change is of the great- nisms behind the changes that were observed. Lastly,
est interest. The point here is that although monitor- the USFS often relied on monitoring vegetation (habi-
ing is very useful, it can be expensive and may not be tat) as a proxy for presence or abundance of wildlife.
needed in all circumstances. An assessment can help us Overall, this approach was untenable and was replaced
decide if monitoring is needed, provide some baseline with a new planning rule as of 2012 (see the following
data for the monitoring effort, and can help us choose discussion), where monitoring is conducted at broad
which variables may be most precise and powerful for scales, focused on ecological conditions, and used only
monitoring. sparingly on focal species.
Once we decide that we need to accomplish more
than a simple assessment and instead need trend, we
Should Habitat Surrogates Be Used?
are in the realm of monitoring. If we are interested in
how the abundance of elk changes over time, we need Monitoring some state variables can be very expen-
to consider a suite of technical questions including sive. One potential alternative to monitoring the state
those relating to statistical power, such as the length variable of interest directly is monitoring a surrogate.
of time that we need to monitor a species before we There is much debate in the literature on the appropri-
would reasonably expect to see an increase or decline, ateness of using surrogates for monitoring (Carignan
or how we plan to estimate abundance given budget and Villard 2002). However, we need to differentiate
constraints. Yet, before we consider technical details and clearly define what we mean by surrogates. Sur-
of a monitoring effort, we first need to clearly define rogates are meant to indicate a variable of interest but
our study objectives and ensure that the product of are either less expensive or less complicated to mea-
our monitoring will satisfy our original goals. That is, sure. In science, surrogates should have a strong sta-
monitoring abundance of elk over a ten-year time hori- tistical link or causal connection to the state variable
zon and having 80% certainty that the population has of interest (e.g., as one variable increases, the other
declined by greater than 25% still does not provide any does by a predictable amount). Many surrogates are
130 research and conservation

used for monitoring, with different levels of success, and empirical concerns, we recommend that if one
depending on what is being monitored. For example, decides to monitor, it needs to be done on focal spe-
there is evidence that species accumulation indices, an cies, unless adequate data has been collected to deter-
index relating the percentage of an area surveyed to mine the causal or statistical relationship between a
the percentage of the target species represented scaled surrogate and population metric of interest. Thus, we
by random and maximum possible curves, can act as recommend that agencies should manage habitat and
weak indicators of biodiversity (Rodrigues and Brooks monitor species in order to meet wildlife monitoring
2007). In wildlife management, we are often interested mandates.
in surrogate measures for abundance, although there
are other state variables that may be more meaningful
Adaptive Management
than abundance.
Unfortunately, too often the surrogate for abun- Once a decision has been made to monitor a focal spe-
dance that is used in practice is vegetation or habitat cies, there is still the issue that monitoring only pro-
(Hunter et al. 1988). While the idea behind using habi- vides information about trend or trajectory but does
tat or vegetation communities as a coarse filter is very not provide any mechanism behind the trend (Nichols
tempting, and has been used extensively in the conser- and Williams 2006). In recognition of this fact, the
vation planning literature, the justification behind it is study of mechanism behind change in population tra-
extremely weak (Oster et al. 2008; Seddon and Leech jectory has been combined with monitoring to create
2008; Cushman et al. 2010; Lindenmayer and Likens an “adaptive management” framework (Walters 1986;
2011). For example, Cushman et al. (2008, 2010) ex- Johnson et al. 1993; Williams 1996; Block et al. 2001;
amined abundance patterns of birds in the Oregon Yoccoz et al. 2001; Westgate et al. 2013). Adaptive man-
Coast Range and found little support for the idea that agement is the structured, iterative process of making
vegetation measures could predict abundance patterns a management decision when there is considerable
at both a plot and subbasin level. They noted that for uncertainty. The adaptive cycle relies on monitoring
this concept to work, several criteria must be met. The the system after a management action by collecting ad-
first is that habitat is a proxy or surrogate for population ditional data and revising models to reduce uncertainty
abundance. This requires that abundance of a species and ultimately inform future decisions. Ideally, studies
is tightly linked with the current and future environ- are structured as experiments to assess cause and ef-
mental conditions such that changes in the environ- fect; in reality, studies are typically pseudoexperiments
ment are predictably reflected in abundance (Cushman or observational (Block et al. 2001). In essence, sys-
et al. 2008). Second, vegetation maps at multiple scales tematic, monitoring data are collected in a hypothesis
must be reliable as a surrogate for habitat. One prob- framework (Nichols and Williams 2006).
lem is that even if there are high correlations between Resource managers are often quick to assert that
vegetation and habitat, and between habitat and abun- they practice adaptive management. In a general sense,
dance, the compounding of relationships makes the use adaptive management is a structured process to learn
of vegetation maps a poor proxy of abundance. Further- from what was done in the past. The key here is that
more, many wildlife-habitat models that relate species the process is structured with a series of steps and
occurrence to broad vegetation types suffer from large feedback loops that facilitate learning. Moir and Block
rates of commission and omission errors (Block et al. (2001) provided a conceptual model for adaptive man-
1994). This is not to say that there are not cases where agement on public lands. It consisted of a seven-step
there are tight statistical links between vegetation and process with the steps linked by a flow of information
abundance, but this may be the exception, not the rule. and feedback loops to inform whether or not manage-
Thus, unless a clear statistical or causal relation- ment actions were meeting their intent. Information
ship between a surrogate measure such as habitat and provided in feedback loops is largely the result of moni-
species abundance, occurrence, or another state vari- toring, and without monitoring, adaptive management
able is established, one should not assume it exists fails (Biber 2011). These feedback loops can evaluate
(Lindenmayer and Likens 2011). Based on theoretical whether: (1) the monitoring design is effective and the
 manage habitat, monitor species 131

correct state variable(s) are being measured; (2) moni- multispecies metrics (e.g., abundance, distribution,
toring is being done as designed; and (3) whether species richness, changes in local extinction and local
or not the management action is achieving its goal. colonization). At the other end of the spectrum from
Clearly, adaptive management hinges on successful context monitoring is targeted monitoring. This is the
monitoring. The failure to conduct valid monitoring is monitoring of how a particular species is responding
considered the “Achilles’ heel” to adaptive management to a management action or general threat (Holthausen
(Moir and Block 2001). et al. 2005). Targeted monitoring also can be the moni-
While the adaptive management approach can toring of sensitive, threatened, or endangered species
work well for species in which data are relatively easy that may not be suffering from ongoing management
to collect and abundant, such as game species, many actions but are threatened by small population pro-
monitoring efforts are focused on rare, endangered, cesses (Caughley 1994). Context and targeted monitor-
and sensitive species. Threatened, endangered, and ing have very different objectives and costs, and likely
sensitive species often occur in low numbers and can involve collection of different types of data. Therefore,
be secretive and difficult to sample (Neel et al. 2013). the first step in designing any monitoring program will
Data will inherently be limited compared to more be stating this broad objective.
abundant game species. Thus, the adaptive manage- For the remainder of this review, we will largely
ment framework is likely to fail for rare species where focus on targeted monitoring. We do not mean to im-
experiments are unethical and data is sparse, unless ply that large, broad-scale context monitoring efforts
adequate sampling effort, sometimes approaching a have been unsuccessful or are not useful. Clearly, ef-
census, is allotted. forts like the Breeding Bird Survey, which relies on
volunteers across the United States to observe birds
during the breeding season, has led to many useful
Articulating the Monitoring Goal
insights into changes in bird distributions and abun-
One failure of many monitoring programs is the lack of dances (Link and Sauer 1998; Sauer et al. 2003; Sauer
articulation of clear and measurable objectives. Clearly, and Link 2011; Stegen et al. 2013). Whereas we see
one of the advantages of the adaptive management the value in context monitoring and advocate the need
framework is that objectives must be stated if data are to for national-level monitoring programs to accomplish
inform competing management hypotheses of how the this, success requires well-coordinated, well-funded,
system functions (Nichols and Williams 2006). How- broad-scale data collection on many species or assem-
ever, adaptive management also has been criticized blages. It therefore represents the minority of monitor-
for not articulating and including triggers to change ing efforts designed each year worldwide (e.g., Green-
management actions, leading to a lack of accountability wood 2003; Kéry and Schmid 2006; Kéry and Plattner
(Nie and Schultz 2012). Outside of the adaptive man- 2007; Biodiversity Monitoring Switzerland [http: //
agement framework, there are hundreds of ongoing www.biodiversitymonitoring.ch]). Thus, in the subse-
monitoring efforts that occur on public lands that have quent sections, we discuss issues of targeted monitor-
no clearly stated objectives or thresholds and are not ing related to metrics to monitor, sampling area, scale,
tied back into decision-making processes. cost, power, effect size, and duration.
The first monitoring goal that needs to be clearly
stated is the purpose of the monitoring effort. Are the
Metrics to Measure and Monitor
data being collected for general context monitoring
(sensu Holthausen et al. 2005), which is defined as An investigator must first decide what it is that he or
monitoring a broad array of ecosystem components she wants to measure (see table 10.1 adapted from
at multiple scales? Breeding Bird Survey data (e.g., Holthausen et al. 2005). As noted earlier, the ultimate
Boulinier et al. 1998) are an example of these types of piece of information sought is how management affects
data. Inherent in context monitoring is the idea that the population(s) under consideration. The most com-
monitoring will detect emerging threats to wildlife mon metric to monitor is abundance. Yet, abundance
populations and unanticipated changes in species or may not be the most biologically meaningful metric
Table 10.1 Population and community measures that may be employed in a monitoring program. This table is
adapted from Holthausen et al.’s (2005 ) table 4.

Population Measures

Abundance / density Abundance and density are the most commonly used monitoring metrics. Abundance
estimates require capturing, marking, and recapturing individuals multiple times.
Note that individuals do not need to be physically captured (e.g., noninvasive DNA
capture-recapture) to provide an estimate with reasonable variance. Because these
measures are often expensive, indices of abundance are frequently used. Density
is abundance per area, although the estimation of the area being considered is not
always clear. Abundance change over a unit of time (λ, finite rate of increase) also
can be monitored.
Occupancy Occupancy assesses presence of an organism at sample locations. Showing presence
at a site necessitates only proof of the species’ existence. Demonstrating absence is
more complex and requires information on the probability of detecting the organ-
ism, given it is present. These data are often placed in an occupancy framework
where multiple site visits at multiple stations can be used to infer changes in oc-
cupancy, which can be linked to abundance (Royle and Nichols 2003) or monitored
as its own metric (MacKenzie et al. 2006). Occupancy models (MacKenzie et al.
2006) also can be used to detect changes in local colonization and local extinction
of a species.
Vital rates (birth / death; immigration / emigration) Vital rates are age-specific birth and death rates, and emigration and immigration
rates. Monitoring vital rates sometimes provides a better measure of trend than
measures of abundance. Collecting vital rate data allows the use of population
viability analyses, which can estimate probability of persistence and allow modeling
of the impact of management actions on persistence.
Range distribution measures Geographic range is the range of a given species at a particular point in time. Monitor-
ing range over time provides information on if a species is expanding, contracting,
or remaining constant. Groups like the IUCN often use range distribution to moni-
tor change in species status (Mace et al. 2008).
Genetic measures Genetic monitoring has been defined as quantifying temporal changes in population
genetic metrics or other population data generated using molecular markers. In
general, three categories of genetic monitoring are recognized. Category I is simply
the use of diagnostic molecular markers for traditional monitoring such as estimat-
ing changes in population abundance, vital rates, hybridization, or geographic range.
Category II used the population genetic data itself to monitor changes in genetic
variation, effective population size, dispersal, or population substructure. Category
III addresses questions of evolutionary potential, such as change in frequencies of
adaptive genes (see Hansen et al. 2012).

Community Measures

Diversity measures Species diversity measures used in community and ecosystem ecology include species
richness and evenness. Species richness can be based on repeated estimates of the
presence or absence of taxa, whereas species evenness requires abundance data.
With recent advances in analytical techniques (Dorazio et al. 2006; Royle and Kéry
2007; Kéry and Royle 2008), we can estimate species richness and community
dynamics from local colonization and local extinction (e.g., Sauer et al. 2013) while
accounting for detection.
Integrity measures Biological integrity is a comparative approach that examines if conditions (which can
be measured using other approaches mentioned previously) are similar today to
historical times or to other locations that are considered “intact.”
 manage habitat, monitor species 133

to choose (Van Horne 1983) because it only provides could be misleading, if the population includes a large
one piece of the puzzle, and other state variables may number of non-territorial animals (e.g., nonbreeding
be the driving factors for population change. For ex- individuals) not easily sampled using traditional meth-
ample, imagine a stream that has two adult fish in the ods. In a situation where there is high mortality of terri-
early spring that produces between five hundred to torial animals that are immediately replaced by surplus,
one thousand fry in the summer, which are subject to the population of territorial animals may appear stable
heavy mortality. Imagine this dynamic continuing for a while the overall population is declining.
decade. An estimate of abundance in the fall would be We recommend conducting sensitivity analyses to
less useful than a measure of connectivity or effective determine which state variables contribute most to
population size. Even if abundance is the desired goal, population change (e.g., Caswell 1978; Williams et al.
as it is easily articulated and fits well into management 2002). If demographic data are not available, the inves-
models (Fancy et al. 2009), estimates of abundance tigator will need to conduct literature reviews, consult
may not be obtainable, especially across an entire geo- with established experts (useful for Bayesian priors
graphic range. In lieu of collecting data to estimate [Gelman et al. 2004]), or use results of similar studies
population abundance, investigators consider various to establish the basis for using specific state variables.
proxy measures (table 10.1) and often consider these All of these sources can be used in a sensitivity anal-
at small spatial scales, sometimes distributed across a ysis, although some sources will be more valuable than
species geographic range. others. For example, there will be more variation in
It is important here to note that while we did not ad- parameter estimates from a literature review of studies
vocate the use of habitat as a surrogate measure, there in different geographic locations or resulting from dif-
are often many surrogates that can be used for formal ferent study methods compared to parameter estimates
estimates of abundance (Slade and Blair 2000; Mc- from a study conducted in the same area (Mills 2012).
Kelvey and Pearson 2001; Tallmon et al. 2010; Tallmon Overall, we recommend that selecting the variables
et al. 2012; Noon et al. 2012). The key difference is that to measure should be based on strong statistically based
modeling efforts that have tested the use of these surro- information. One must understand the biology and
gates for abundance often find them to be good predic- population dynamics of the species being monitored
tors of abundance and, in some cases, more robust for to make better decisions on exactly what to monitor.
detecting trends than estimates of abundance. In other
words, the causal connection has been studied and
Selection of Sampling Areas
validated. For example, Tallmon et al. (2010) showed
that declines in effective population size can be a more Once the study objective is established, the scale of
sensitive measure of declining populations than moni- resolution chosen by ecologists is perhaps the most
toring estimates of abundance under many conditions. important decision in monitoring because it prede-
McKelvey and Pearson (2001) demonstrated that Mt+1 termines procedures, observations, costs, and results
(otherwise known as the number of unique individuals (Green 1979; Hurlbert 1984). A major step in design-
captured at time t+1) showed lower variance and less ing a monitoring program is to define the target popula-
sensitivity to model violations than formal abundance tion. Defining the target population essentially defines
estimates when sample sizes were small (but character- the area to be sampled. This first step establishes the
istic of most field studies). Similarly, there has been a sampling universe from which samples can be drawn
recent explosion of the use of occupancy estimates as a and the extent to which inferences can be extrapolated.
way to monitor trends in wildlife populations (Gaston Although this seems rather straightforward, the
et al. 2000; MacKenzie and Nichols 2004; MacKen- mobility of wildlife can muddle the inferences drawn
zie et al. 2006; Estrada and Arroyo 2012). from the established area. The basis for establishing the
When monitoring populations, it is beneficial to de- sampling frame includes information on habitat-use
termine which parameters are most sensitive to change patterns, range sizes, movement patterns, and logis-
and focus on those. Typically, focus is on population tics of sampling a large area. Ideally, the bounds of the
abundance or density of breeding individuals, which sampling area would correspond to some natural bar-
134 research and conservation

rier that would keep a population closed, but this is breeding animals should be conducted during the
often not the case. Without such bounds, open forms nonbreeding period, and so on. Within a season, tim-
of estimators will need to be used (Marucco et al. 2011; ing can be critically important because detectability
Wilson et al. 2011; Ivan et al. 2013), or a combination of individuals can change for different activities or dur-
of open and closed estimators, where sampling occurs ing different phenological phases (Bailey et al. 2004;
during a short interval when the population is closed MacKenzie 2005). Male passerine birds, for example,
and over multiple seasons or years when the population are generally more conspicuous during the early part of
is open (i.e., robust design [Pollock 1982; Kendall and the breeding season when they are displaying as part
Nichols 1995; Kendall et al. 1995]). of courtship and territorial defense activities. Detection
To reduce this issue associated with population probabilities for many species will be greater during
closure, there have been developments in using an this period than at other times. Another consideration
approach to estimating density (abundance per area) is that the very population under study can change
instead of abundance, using spatially explicit capture- within a season. For example, age-class structures and
recapture models (e.g., Efford 2004; Royle and Young numbers of individuals change during the course of the
2008; Gardner et al. 2009; Russell et al. 2012). The breeding season as juveniles fledge from nests and be-
distribution of individuals and their movements rela- come a more entrenched part of the population.
tive to a trapping array are explicitly modeled using Population estimates for a species therefore may dif-
this framework. For instance, the use of telemetry data fer depending on the timing of data collection. Once
can be used to evaluate geographic closure (White and the timing decision is made, data must be collected
Shenk 2001; Ivan et al. 2012) and to explicitly model an- during the same time in subsequent years to control
imal movement in spatially explicit capture-recapture for some of the within-season variation. Objectives of a
models (Sollmann et al. 2013). To evaluate geographic monitoring effort also dictate when data should be col-
closure, animals are marked and time spent in the lected. If monitoring is conducted to evaluate popula-
study area is estimated and used to scale abundance tion trends of a species based on a demographic model,
estimates and provide density (Ivan et al. 2013). Simi- sampling should be done at the appropriate time to
larly, genetic data can be used to evaluate violations ensure precise and unbiased estimates of the relevant
of closure assumptions by using approaches such as population parameters. Demographic models typically
spatial autocorrelation among genotypes (Peakall require fecundity and survival data to estimate the
and Smouse 2006). These methods provide indices of finite rate of population increase. Sampling for each
average animal gene flow in an area. Unfortunately, of these parameters may be necessary during distinct
there has been little attention paid to the integration times to ensure accurate estimates for the respective
of genetic approaches to gene flow and abundance es- measures. Timing also is essential in the field of genetic
timation, although this is beginning to change (Luikart monitoring (Goudet et al. 2002; Schwartz et al. 2007).
et al. 2010). For example, different measures of gene flow and ge-
netic variation have been found to be more sensitive to
the timing (e.g., pre- or postdispersal) of sample collec-
When to Collect Data and for How Long?
tion (Goudet et al. 2002; Hammond et al. 2006). This
A key aspect of monitoring is to know when to col- makes intuitive sense as populations should be most
lect data, including the timing of data collection and differentiated if sampling occurs postbreeding, but pre-
the length of time over which data should be collected. dispersal, and most similar after a pulse of dispersal.
Both are influenced by the biology of the organism, the Consistency in timing and methods is essential.
objectives of the study, intrinsic and extrinsic factors Length of the study refers to how long a study should
that influence the parameter(s) to be estimated, and be conducted for reliable and accurate parameter esti-
resources available to conduct the study. mates. A number of factors including study objectives,
Obviously, studies of breeding animals should be field methodology, ecosystem processes, biology of the
conducted during the breeding season, studies of non- species, budget, feasibility, and statistical power issues
 manage habitat, monitor species 135

influence how long a monitoring study should occur. what effect size they can detect with a specific effort—
Monitoring studies also must consider temporal quali- only to find at the end of the monitoring period that
ties of the ecological process or state being measured they never had power to detect change.
(e.g., population cycles, successional patterns includ- There are several computer programs available to
ing their frequency, magnitude, and regularity; Frank- examine interactions between sample size, effect size,
lin 1989). Furthermore, a study should engage in data type I error, and statistical power to detect trends
collection over a sufficiently long period to allow the in abundance, count, or occupancy under different
population(s) under study to be subjected to a reason- sampling schemes (Gerrodette 1993; Gibbs 1995; Bai-
able range of extrinsic environmental conditions. Con- ley et al. 2007). For example, program MONITOR esti-
sider two hypothetical wildlife populations that exhibit mates statistical power of a population-monitoring pro-
cyclic behaviors: one that cycles on average ten times gram when supplied with the number of plots, counts
per fifty years and the other exhibiting a complete cycle per plot and their variation, duration of the monitor-
just once every twenty-five years. The population cycles ing program, interval of monitoring, magnitude and
are the results of various intrinsic and extrinsic factors direction of ongoing population trends, and signifi-
that influence population growth and decline. A moni- cance level associated with trend detection. Similarly,
toring program established to sample both populations program TRENDS conducts a power analysis of linear
over a twenty-five-year period may be adequate to un- regression for monitoring wildlife populations (Gerro-
derstand population trends in the species with frequent dette 1993). TRENDS estimates statistical power of a
cycles but may be misleading for the species with the monitoring program after providing data related to the
long population cycle. A longer timeframe would be rate of change, duration of study, precision of the esti-
needed to monitor the population of species with lower mates, and the type I error rate. Alternatively, power
frequency cycles. can be set to a desired level and program TRENDS esti-
Considering only the frequency of population cycles mates one of the other input parameters. For example,
may be inadequate, as the amplitude or magnitude of if 80% power is desired, one can ask how many years a
population shifts also may influence the length of a monitoring effort must be conducted given a precision
study. Sampling the population of a species exhibiting of the estimate and type I error rate.
greater amplitude in population increases and declines Recently, there has been an increase in reliance
would require a longer period to detect a population on occupancy-based approaches to estimate popula-
trend or effect size than a species exhibiting lower am- tion trends. A new spatially based simulation program
plitude in population shifts. called SPACE has been created to evaluate power for
detecting population trends over time based on species
detections (Ellis et al. 2013). SPACE has been used to
Power to Detect Change
evaluate the power of monitoring trend occupancy for
A major question that arises in any study is: “Am I territorial animals while accounting for natural history
sampling enough to detect a trend, should one exist?” (e.g., territorial overlap, different space use of different
“Enough” can be enough area, enough individuals per life stages), habitat use, and sampling scheme (e.g., size
area, enough time, and so forth. The best way to ad- of the grid; Ellis et al. 2013).
dress this issue is through simulations or calculations We suggest that an analysis of power to detect trend
of statistical power (Zielinski and Stauffer 1996). That is initiated before any monitoring effort is conducted.
is, it is important to ask a question like: “If I am inter- If specific programs do not exist to evaluate statisti-
ested in changes in abundance and I mark one hun- cal power for a given method, consulting papers that
dred individuals per year for ten years, will I have a have conducted simulations on power (and type I er-
95% chance of detecting a 20% decline?” Or ask: ror rates) with that method will be insightful. Overall,
“How many samples do I need to have 80% certainty it would be very unrewarding, if not embarrassing, to
of detecting a 20% decline given a ten-year monitoring come to the end of a long-term monitoring program
program?” Too many monitoring efforts fail to quantify where no trend was found, only to realize that power
136 research and conservation

to detect trend with the given protocol was no greater lation trends of the management indicator species will
than a coin flip. be monitored” (section 219.19 of the 1982 planning
regulations). Interestingly, these management indica-
tor species, which were proxies for other species, were
Resources to Detect Change
often monitored through the use of habitat availability.
All of the components of monitoring that we have This was considered a “proxy-by-proxy” approach and
discussed so far are important but rely on sufficient was deemed legally sound (Inland Empire Public Lands
monetary resources to complete these efforts. Agen- Council v. USFS 1995; Lands Council v. McNair 2008),
cies tasked with developing monitoring programs of- although biologically questionable.
ten work with limited budgets; therefore, cost-efficient Monitoring under the 1982 planning regulations
sampling designs are paramount. Thus, we advocate us- was seen as essential to the success of the multiple-use
ing cost as an additional constraint for study design op- mandate and maintenance of viability clause. Unfor-
timization (e.g., Sanderlin et al. 2009; Sanderin et al. tunately, the regulations relied on focused monitor-
2014). We can determine the power to detect an effect ing of a multitude of management indicator species,
size and the amount of sampling needed for accurate which proved too expensive and difficult to accomplish
parameter estimates, but if there are not enough re- (Schultz 2010; Schultz et al. 2013). Thus, monitoring
sources available to complete a study with these design under the 1982 planning regulations often devolved to
specifications, there is no point in continuing with the assessing the status and trend of the amount of habi-
monitoring effort. tat the species required without strong statistical links
between the habitat and the species. We are currently
in a new era where as of April 2012 the USFS is acting
Case Study: Lessons from USFS
under a new planning rule (which has been in prepara-
Monitoring under the Planning Rule
tion for roughly fifteen years) with extensive public and
We can learn a lot about monitoring programs from scientific involvement. This new direction provides a
the successes and failures of the USFS’s efforts to great opportunity to improve management of wildlife
monitor species under the 1982 “planning rule.” The on public lands (Schultz et al. 2013). Our goals in the
US Forest Service is a United States land management remainder of this section are to review the new rule
agency responsible for facilitating multiple-use activi- as it relates to monitoring and then provide advice
ties on seventy-eight million hectares of land. These as to how to choose species to monitor and points to
activities include the maintenance of healthy wildlife consider.
populations, clean water, recreational opportunities, Monitoring under the 2012 planning rule requires
and timber production. Guidance on how to balance monitoring indicators that relate to the status of eco-
competing demands on the landscape is given in the logical conditions and focal species, which are defined
National Forest Management Act (NFMA) of 1976. as a narrow group of species whose status allows in-
One of the most controversial aspects of the NFMA has ference to the integrity of the larger ecosystem. These
been the tiered planning approach and how planning focal species are to “provide meaningful information
rules accomplish NFMA’s requirement to “provide for regarding the effectiveness of the plan in maintaining
a diversity of plant and animal communities based on or restoring the ecological conditions to maintain the
the suitability and capability of the specific land area diversity of plant and animals communities” (USFS
in order to meet overall multiple-use objectives.” Di- 2012). Interestingly, the new monitoring rule has no
rection on how to achieve NFMA’s mandate to provide special monitoring requirement for “species of con-
for biodiversity was given in promulgated planning servation concern” (e.g., Threatened and Endangered
rules, which until 2012 called for the maintenance of Species), although it does have a viability requirement
viable populations of existing native and desired non- for this category. What remains to be tested in this new
native vertebrate species in a planning area. Part of that planning rule is how individual forests choose focal
maintenance of viable populations included identifying species to monitor and then implement a monitoring
management indicator species and noting that “popu- program.
 manage habitat, monitor species 137

In the following list, we discuss the characteristics 4. Species can be monitored: A species may have a
of a focal species that we think will be effective for low range of variability and be a great reflection
making inference to the larger ecosystem. An ideal fo- of land-use changes, but if it is too difficult or
cal species would have many of the following charac- expensive to monitor then it should not be chosen.
teristics: Some species’ life history traits make them easier
to monitor than others (e.g., small mammals that
1. The species must index something greater than have smaller home ranges, are less vagile, and are
itself: A species that only reflects its own suc- abundant may be preferable to a carnivore that is
cess in an ecosystem will likely be too narrow for secretive, rare, and vagile).
capturing ecological integrity. While we would 5. Background knowledge about national / global
like to have data on all species in a planning area, trend is needed for context: It is important to
common sense and the rule itself dictate that focal tie the local dynamics of a species into a broader
species are only limited subsets of species in the monitoring program. Imagine picking boreal toads
ecosystem. Species such as ecosystem engineers (Bufo boreas) or other amphibians for monitor-
(e.g., beavers [Castor canadensis]) or keystone ing thirty years ago. The population’s numbers
species (e.g., pileated woodpeckers [Dryocopus would have likely declined in a forest, and this
pileatus]) are good examples of animal species that may have been attributed to habitat modifications
index something greater than themselves. Oven- or other management actions. This would be the
birds (Seiurus aurocapillus) are another example, wrong conclusion in the face of global amphibian
as they appear to be sensitive to fragmentation of decline. Knowing the magnitude of global decline
mature forests and may index other species and compared to local declines would have provided
processes of these habitats. relevant context.
2. Low range of natural variability: If a species has
large natural fluctuations that are not linked to
Conclusions, Creative Approaches to
climate or other environmental processes, statisti-
Monitoring, and a Need for Large Scale
cal trend will be difficult to obtain. In this case, the
National Efforts
temporal variation swamps out process variation,
hence obviating the ability to identify trend. Simi- Wildlife monitoring means many things to many dif-
larly, a species known to cycle, like snowshoe hare ferent people. For some it is the broad-scale, long-term
(Lepus americanus), would make a poor choice as efforts to provide background information on spe-
a focal species, as statistically detecting trend will cies, while for others monitoring is very focused on a
be complicated. However, there are times when a specific species for a specific reason (e.g., to detect a
species with a large range of natural variability will change in abundance after a wildfire or other natural
be exquisitely sensitive to environmental change, event), and still for others monitoring is meant to
and this relationship (the signal) will override provide data to better understand mechanisms under-
concerns about the large range of natural variabil- lying ecosystem change (i.e., adaptive management).
ity (the noise). Regardless of the reason to enter a monitoring effort,
3. Species need to be statistically linked to habitat at effective monitoring is a challenging endeavor as it can
the appropriate scale: If the purpose of the focal be prohibitively expensive, take a long time frame to
species is to provide inference to or monitor the achieve results, and may require coordination among
integrity of the ecosystem, then there needs to be multiple groups, especially if the effort is broad in
a strong statistical link to the vegetation. Likewise, scope. This will clearly require creative approaches to
selected focal species must be sensitive to particu- monitoring, which include the development of new
lar land management actions. As many ecologi- field approaches and statistical techniques that can
cal processes are scale dependent, this statistical make monitoring more feasible. We have already seen
evaluation should be conducted at a multiple array many of these developments make monitoring efforts
of scales from plot to landscape levels. that were once inconceivable, such as the monitoring
138 research and conservation

of abundance of wolves (Canis lupus) across the Alps in their objectives. This starts with the statement of
(Marucco et al. 2009), now possible with the advent intent of whether this is a broad-scale, context moni-
of DNA technology. Similarly, other technologies, toring effort or a focal / targeted monitoring effort. The
such as trail cameras, have greatly improved our abil- goal should be stated in context of an overarching ob-
ity to obtain reliable data in an inexpensive manner. jective. For instance, it is not enough to simply claim
Developments in occupancy modeling and abundance that we want the USFS to do focal monitoring of a spe-
estimation also are improving our ability to monitor. cies, but it is essential to understand that the intent
And combining the two approaches—where we use of this focal monitoring may be to measure a broader
occupancy estimation with environmental DNA sam- ecosystem process. After these goals are established,
pling (eDNA), which detects presence / absence of spe- we urge those responsible for monitoring efforts to
cies by sampling the environment (e.g., water or air consider all methodological approaches and conduct
samples)—will certainly reduce costs and allow for rigorous power analyses that include cost constraints
more monitoring data to be collected. to ensure the data collected will ultimately be able to
Another recent trend to improve monitoring is to answer the question posed. We also urge consistency
involve local citizens in monitoring efforts as part of in methods and timing of data collection. The world is
“citizen-science” projects (e.g., Schmeller et al. 2008; filled with too many partial datasets, where someone
Silvertown 2009; Miller-Rushing et al. 2012). This changed methods without proper calibration linking
can reduce costs, allow for more monitoring data to the approaches.
be collected, and foster communications between the Penultimately, we encourage the use of the adaptive
public and governmental agencies. These are unique management framework when appropriate. While this
opportunities to educate the public on research that is approach may not always work, we think that, when
conducted on public lands and give a local community possible, setting monitoring in a framework that helps
a sense of ownership with these public lands. us also understand mechanisms behind trends is essen-
We wish to revisit several critical points that we cov- tial. We hardly have enough fiscal resources available
ered earlier in this chapter, which are essential when now to conduct the initial monitoring, let alone funds
designing more effective and efficient monitoring pro- to go back and try to conduct studies to understand
grams. First is the concept of the use of habitat as a trends once they are detected. Adaptive management
proxy or surrogate measure for wildlife populations. and structured decision making allow us to kill the pro-
Although it is very tempting to use habitat as a proxy verbial “two birds with one stone.” However, we cau-
for abundance of a species, there is very little evidence tion that adaptive management is best accomplished
that this is a defensible position. The burden of proof is when thresholds or triggers are in place to force certain
therefore on the group wishing to use habitat as a proxy conservation actions, should these triggers be tripped.
to statistically show the tight relationship between Lastly, in this chapter we focused on the monitor-
the two variables before using them in a monitoring ing of focal species, which included species that re-
program. This does not mean that we are against sur- flect broader ecosystem processes and species that
rogates or indices in general, but we urge caution in are of special management or conservation concern.
their use without first establishing a causal or statisti- While we believe that most monitoring efforts will be
cal relationship, which is often not established when targeted, this does not mean that we do not see the
monitoring habitat as a surrogate for species’ abun- value in context monitoring approaches. We can only
dance. In fact, there is strong statistical evidence that imagine the valuable data that could be produced if we
under some circumstances indices of abundance are had a national wildlife monitoring program that was
as good as collecting abundance, and some of these on the scale of our Forest Inventory and Analysis pro-
indices and surrogates may reflect change better than gram (cf. Manley et al. 2004, 2005). Perhaps such an
estimated abundance (McKelvey and Pearson 2001; effort would have allowed the detection of emerging
Engemann 2003). threats, such as invasive species, faster than our ad hoc
Our next summary point is that once a decision is detection methods currently in place could, ultimately
made to monitor, we implore groups to be very specific saving billions of dollars. A commitment to large-scale
 manage habitat, monitor species 139

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 11
Kevin S. McKelvey
The Effects of Disturbance
and Succession on
Wildlife Habitat and
Animal Communities

T his chapter discusses the study of disturbance and

succession as they relate to wildlife. As such, the
discussion is confined to those disturbance processes
all affect both the postdisturbance wildlife community
and, more importantly, the trajectory of the postdistur-
bance community. Even for well-studied species, co-
that change the physical attributes of habitat, leading herent understandings of their relationships to distur-
to a postdisturbance trajectory. However, even with bance across time and space are therefore often vague.
this narrowing of the scope of disturbances discussed, Commonly, we look at successional changes in habi-
there remain formidable obstacles prior to any co- tat quality by using spatial samples of different ages as if
herent discussion of disturbance. The first, and most they were a temporal series, which implies spatiotem-
fundamental, is definitional: what constitutes distur- poral constancy in successional dynamics. This assump-
bance and succession, and what is habitat? The con- tion has served wildlife research well, but in the face of
cepts of disturbance, habitat, and succession are highly directional climate change and the nearly continuous
scale- dependent; disturbances at one scale become addition of exotic species, this approach is becoming
part of continuous processes at a larger scale, and ideas increasingly untenable. We need to embrace the idea
associated with succession require assumptions of con- that postdisturbance succession is increasingly unlikely
stancy, which become highly problematic as spatiotem- to produce communities similar to those that the dis-
poral scales increase. Literature on the effects of distur- turbance altered: short-term successional patterns are
bance and succession on wildlife, however, focuses on a likely to be influenced by the large and dynamic pool of
narrow range of spatial scales, primarily occurs within exotic plants and animals and longer-term succession
a narrow temporal window immediately following dis- by directional climate change.
turbance, and seldom includes interactions between Together, these observations indicate a need for
areas within the disturbed patch and the landscape studies of disturbance and succession at larger spatio-
that surrounds it. While these largely descriptive stud- temporal scales. It is, however, important to assess
ies undoubtedly have great local value, more general the feasibility of scaling up studies in time and space.
information about the relationships between organ- Clearly, studies that expand both the spatial and tem-
isms and environments shaped by disturbance and poral dimensions of data collection quadratically in-
succession is remarkably limited. Multiple small-scale crease costs; longer time frames do not fit into current
descriptive studies of the immediate postdisturbance competitive funding structures and contain a variety
environment do not appear to coherently aggregate of negatives such as the potential loss of data. Broad-
into larger understandings of the effects of disturbance scale targeted monitoring, however, can provide a
and succession on wildlife. Context is important: the framework allowing acquisition of data at these scales
conditions at the time of the disturbance, in adjacent and can be designed to produce both immediate and
undisturbed patches, and within the broader landscape longer-term results.
144 research and conservation

and other habitat features). Temporal habitat patterns

Disturbance, Succession, and Questions
are as critical for conservation as are spatial patterns,
of Scale
but much harder to study. In many cases, the events
Bormann and Likens (1979) defined disturbance as dis- that structure landscapes and define species ranges are
ruption of the pattern of the ecosystem, principally by rare—often singular. For example, the genetic popula-
external physical forces. This idea, however, assumes tion structures of many species are strongly associated
that ecosystems function as idealized Newtonian sys- with glacial vicariance that occurred during the Pleis-
tems, in stasis until external energy is applied. But few tocene (see Shafer et al. 2010 for a review). To study
ecosystems exist in equilibrium (Sousa 1984). Further, these rare events, we primarily look to the past to gain
Rykiel (1985) noted that whether disturbance is viewed insight into their frequency, size, and postevent succes-
as changing the state of an ecosystem or being part of sional and evolutionary trajectories. Further, for prac-
that state is entirely a function of scale. At one scale, a tical reasons, we frequently use habitats of different
tree-fall is a state-altering disturbance; at a larger scale, ages as surrogates for the passage of time, making the
it is part of a continuous process that creates and main- tacit assumption that if we were to project one area
tains the state of an old-forest ecosystem. The same forward (or backward), it would be similar enough to a
thing is true for larger disturbances; at one scale, fire in surrogate area that we can infer its future or past state
western US forests represents a significant disturbance by studying that surrogate. I refer to this approach as
to ecological function, radically altering wildlife habi- “trading space for time.”
tats. At a larger scale, fire is a part of the ecosystem— The study of temporal patterns of habitat over time,
many ecosystems are dependent on fire to maintain and specifically the validity of using spatial surrogates to
the presence, patterns, and juxtapositions of plant and infer temporal patterns, is tightly linked to the concept
animal species (Habeck and Mutch 1973; Covington of succession. Here I use this term in a neo-Clemensian
and Moore 1994; Nowacki and Abrams 2008). fashion (Clements 1916; Daubenmire 1952): suc-
Similar-scale dependencies are associated with cession assumes a pattern of orderly and predictable
the effects of disturbance on wildlife habitat. Habitat changes in species presence and abundance that occur
is often thought of with a particular scale in mind: over time after disturbance, leading to a fairly stable
perhaps a forest stand or home-range area evaluated terminal state, or climax. As Gleason (1927) noted, the
across a year or the lifespan of an organism. If succes- idea of succession is very appealing: if understood, it
sional thinking is applied, habitat may be defined as allows us to predict the future and to see into the past
existing within a specific sere (e.g., a species may be without needing any data other than what we collect
considered to be associated with early seral or late suc- in the present. However, successional concepts cannot
cessional forests). However, the habitat requirements be accepted naively. The validity of this concept, and its
for population persistence are, like disturbance, com- resulting popularity (or lack thereof), is directly related
plex and span many scales in both space and time. In to the complexity of the system studied and the scale
space, wildlife habitat spans spatial domains measured at which the system is evaluated. Simple systems have
in meters (e.g., specific resources for denning), to ki- fewer succession pathways and fewer species; hence,
lometers (e.g., sufficient resources to support a local the vegetative trajectories are more predictable. For
viable population), to hundreds of kilometers (e.g., a example, in the western forests of the United States,
mosaic of resources and connectivity sufficient to sup- tree communities are often simple and contain large
port long-term metapopulation [Levins 1969, 1970] areas of intact natural vegetative communities. In
persistence and abundance). Additionally, at all spatial these systems, succession-based classification systems
scales, juxtaposition and spatial patterns of habitats (e.g., habitat types; Pfister and Arno 1980) and con-
are important (e.g., Iverson et al. 1987). Temporally, cepts (e.g., potential natural vegetation [PNV]; Küchler
the definition of habitat is even more complex. Again, 1964) are popular. In highly modified landscapes con-
scales vary from almost instantaneous (e.g., the timing cepts, however, concepts like PNV become abstract as
of ice breakup in the Arctic) to millennial (e.g., the pro- none of the vegetation within a study area may exist
cesses of erosion and deposition that create soils, caves, in its putative potential condition (Zerbe 1998). For
 effects of disturbance and succession on wildlife habitat 145

these reasons, looked at objectively, succession is often process, we discuss and label this process as part of a
a problematic concept, but one that has proved useful separate body of literature. Similarly, we are prone to
in many communities and without which our ability view what we do as being more of a disturbance than
to study systems and build predictive models would be what other organisms do. For example, beavers (Cas-
severely limited. However, regardless of its historical tor canadensis) instigate a variety of disturbances (tree
validity, there is good reason to doubt its relevance as a felling, house and dam construction), oftentimes re-
tool to predict future habitat conditions; the likelihood moving all accessible trees from the area adjacent to
of rapid directional climate change and the increasing their pond (Martell et al. 2006) and thereby creating
presence of exotic species compromise our ability to a shifting pattern of disturbance that can, over many
use past patterns to predict the future. Another con- generations, affect large areas (Naiman et al. 1986).
cept, community assembly theory, which views com- However, studies of beaver activity are seldom framed
munities as being the result of a continuous process of or titled as being disturbance studies.
species invasion and extinction (Lodge 1993), may be By choosing to concentrate our studies on mid-scale
more germane. disturbances, such as fires, and the subsequent changes
in habitat, we have, unfortunately, also chosen events
that occur at temporal scales that are inconvenient—
Disturbance and Its Relationship to
postdisturbance successional trajectories frequently
Wildlife Communities
require many years to evolve. To study these processes,
Defining Disturbance
it is therefore necessary either to set up very long-term
The spatial scaling of habitat, including the habitat studies or trade space for time, with its associated as-
patch structure generated by disturbance, is ultimately sumptions and weaknesses. As noted earlier, trading
defined by the grain at which specific organisms per- space for time requires the assumption of a high de-
ceive the world (Kotliar and Wiens 1990) and opera- gree of similarity and transferability across disturbance
tionally defined at some arbitrary time scale. Thus, events and subsequent recoveries. Given these con-
disturbances also are subject to these same scalings: straints, there are two types of disturbances that occur
an event that constitutes a significant disturbance for at scales and with frequencies that allow both study
a woodland salamander (e.g., Ensetina sp.) may be of and reasonable transferability: anthropogenic changes
little importance to a wolverine (Gulo gulo). Thus, such as logging and land conversion, and fires; a great
when we consider disturbance ecology, we should ide- deal of what we identify as the study of disturbance
ally begin with operational environment of the affected therefore concerns these types of disturbances. Likely
organisms, clearly identify the time frame of interest, for these reasons, and based on practical considerations
and define disturbances accordingly. This, however, is such as the ease of aging trees and hence inferring past
seldom done. When we consider disturbance, we often disturbance events, this body of literature has a strong
have a particular scale in mind, generally an intermedi- bias toward the study of forested systems.
ate scale. When we label events as disturbances, we en- In addition to often being appropriate in scale, an-
vision events such as hurricanes or forest fires and par- thropogenic disturbance is highly researchable because
ticularly anthropogenic activities such as logging, land we seek to control many aspects of our disturbances.
clearing, and infrastructure development: scientific For example, in forestry we apply standardized silvicul-
articles that claim to study disturbance overwhelm- tural treatments: a clear-cut treatment, in addition to
ingly study these types of events. However, this scale is removing all trees, generally will also include removal
not the organism’s scale; rather, it is our scale. In part, of residual debris and soil scarification, leaving a fairly
this is simply a matter of semantics: how we define homogenous postdisturbance environment that is
disturbance rather than what we study. For example, replicated across multiple treatment areas. Treatment
we study the effects of tree-fall and subsequent forest blocks also tend to be of similar size—both econom-
gap dynamics on wildlife; Forsman et al. (2010) found ics and policy dictate this. On Forest Service lands in
eleven studies of gap effects on forest birds. However, the United States, for example, clear- cut treatments
while forest gap formation is an obvious disturbance have for many years been limited in size and spacing
146 research and conservation

to ten to twenty ha blocks interspersed with uncut ar- disinterest—is the norm not just for studies of topical
eas (Franklin and Forman 1987). They also occur at natural disasters but is pervasive across disturbance
relatively fixed rates across time and space because types. As noted by Fontaine and Kennedy (2012), in a
we desire fairly constant flows of products supplied recent meta-analysis of fire effects on small mammals
to a fixed array of mills. Thus, a system of clear-cuts and birds, the meta-analysis was limited to short-term
is a much more regularized disturbance pattern than (four years or less) responses because data “at longer
would occur naturally, and for scientific study, these time scales were too sparse to permit quantitative as-
regularizing factors allow us to achieve a level of rep- sessment” (Fontaine and Kennedy 2012, abstract).
lication that is generally absent in natural events. Of
natural disturbances, fires are probably the most ideal anthropogenic disturbance
for study. In fire-prone areas, fires occur frequently Within the constraints noted by Fontaine and Kennedy
enough to be grouped based on covariates such as in- (2012; taxa limited to small mammals and birds and ef-
tensity, aspect, and pre-fire vegetation. Thus, fires form fects to the period immediately following disturbance),
another major class of disturbance studies. In addition there are still hundreds of published papers document-
to these two heavily studied classes of disturbance, a ing the effects of anthropogenic disturbance on wildlife.
third major group of published studies concerns distur- Luckily, there has been a series of recent meta-analyses
bance events that we perceive as being “natural disas- that provide both syntheses of these papers’ findings
ters”: hurricanes, floods, tsunamis, and so forth. and extensive bibliographic references. Kalies et al.
(2010) performed a meta-analysis of twenty-two papers
on the effects of thinning, prescribed burns, and some
Wildlife Studies Associated
wildfire on birds and rodents in the American South-
with Disturbance
west. Not all species or taxa were equally represented;
Formal studies of the effects of disturbances on wild- birds were most commonly studied. Spatial scales were
life are almost exclusively limited to the immediate small (less than four hundred ha) and temporally close
aftermath of the event. This phenomenon is perhaps to the treatment (twenty years or less posttreatment).
most apparent in the study of natural disasters. For ex- Ground-foraging birds and rodents showed neutral
ample, a Google Scholar search on “Hurricane Hugo” density responses to the treatments, whereas aerial,
revealed a total of thirteen journal publications on tree, and bole-foraging birds had positive or neutral
the hurricane’s effects on animals, of which ten (77%) responses to both small- diameter removal and burn-
were published within the first five years. However, few ing treatments but negative responses to overstory re-
studies are currently underway; I was not able to locate moval and wildfire. Hartway and Mills (2012) looked at
a single journal article with animal responses to Hurri- a variety of anthropogenic treatments to increase nest
cane Hugo as the primary subject published after 2002. success, including prescribed burns (nineteen stud-
This pattern is not limited to the study of wildlife: of ies, fifty-two species). They also evaluated the effects
seventy-two identified journal publications on all sub- of livestock exclusion, prescribed burning, removal of
jects concerning the effects of Hurricane Hugo, sixty- predators, and removal of cowbirds (Molothrus ater).
seven (83%) were published in the first five years, and Of the four treatments, prescribed burning was not the
none were published after 2008. Given that changes most effective but did significantly increase nest suc-
in vegetation wrought by Hugo will affect habitat in cess for most species, though there was a large amount
various ways for hundreds of years, there is no biologi- of variation between species groups. Specifically, preco-
cal rationale for this spate of studies of the exceedingly cial bird species were negatively affected by prescribed
ephemeral habitat conditions associated with its imme- burning. Vanderwel et al. (2007) performed a meta-
diate aftermath coupled with low levels of interest asso- analysis of the effects of partial cutting on forest birds.
ciated with the still rapidly evolving habitat conditions All forty-two studies examined the effects of cutting
that currently exist. Further, this pattern—frequent ten years or less after harvest. Of the thirty-four bird
studies immediately after the disturbance followed by species studied, fourteen declined with increasing har-
 effects of disturbance and succession on wildlife habitat 147

vest intensity and six increased. Not surprisingly, deep erality of the pattern observed by Hossack et al. (2013)
forest birds such as ovenbirds (Seiurus aurocapilla) and is unknown.
brown creepers (Certhia americana) were the species
most sensitive to harvesting.
General Strengths and Weaknesses of
Disturbance-Related Meta-analyses
fire
Similar to anthropogenic disturbances, a number of Throughout these meta-analyses, a general pattern
recent meta-analyses document the effects of fire on emerges: bird response to disturbance is much more
wildlife. Not all of these meta-analyses separate natural heavily studied than is mammal response—confin-
fire from anthropogenic treatments such as thinning ing their scope to the effects of fire alone, Leidolf and
or prescribed burns. For example, Zwolak (2009) per- Bissonette (2009) identify 512 studies on birds—and
formed a meta-analysis of fire clear-cutting and partial mammal study is almost exclusively limited to small
cutting on small mammal populations. Of the fifty-six mammals. The general paucity of habitat disturbance
studies used in the meta-analysis, most (fifty-four) doc- studies for mammals other than small mammals is
umented short-term changes. Only eleven documented likely due to several factors. Perhaps the most critical
longer-term (ten to twenty years after the disturbance) factor is that for larger mammals, home range sizes
effects (Zwolak 2009, supplemental, appendix A). and / or densities are not compatible with abundance
Zwolak found that deer mice (Peromyscus maniculatus) estimation within disturbance classes. Another is that
increased and red-backed voles (Myodes gapperi) de- they are far more difficult to locate and count than are
creased after disturbance, with the effect size increas- small mammals and particularly birds. Even for small
ing with rising disturbance intensity. These patterns re- mammals, there is no approach that provides as effi-
laxed as time since disturbance increased. Fontaine and cient an estimate of occurrence and relative abundance
Kennedy (2012) performed a meta-analysis of forty- as a bird point-count. Thus, specific disturbance studies
one papers, documenting the short-term effects of fire of many mammalian species of interest such as large
and fire surrogates on small mammals and birds. They, carnivores are impeded both by the utilization of large
like Zwolak (2009), found that postdisturbance effects areas that likely exceed areas of disturbance and diffi-
on the abundance of species increased with increasing culties in obtaining sufficient sample sizes. The effects
fire severity. A total of 119 bird species and 17 small of disturbance on other groups such as herptofauna are
mammal species were analyzed by Fontaine and Ken- far less studied.
nedy (2012). While generalizations across this broad of In addition to being limited in types of species ana-
a group are difficult, in many ways their results mirror lyzed, meta-analyses are weakened by the need to re-
those of Vanderwel et al. (2007) and Zwolak (2009): duce studies to groups that can be evaluated based on
the largest positive effect was on deer mice in severe a common metric (effect size) and common covariates
burns whereas forest birds such as ovenbirds and her- such as time since disturbance. While such grouping
mit thrush (Catharus guttatus) were negatively affected. is necessary, all disturbances are not created equal;
Interestingly, brown creepers were positively affected postdisturbance abundances may be correlated with
by moderate intensity disturbances. While the period abundances within the general landscape (e.g., Brotons
immediately following disturbance may exhibit the et al. 2005), a covariate that is generally missing from
most profound shifts in structure and composition with disturbance studies. Similarly, the effects of past distur-
a relaxation toward the predisturbance state over time bances may have altered systems leading to different re-
(e.g., Zwolak 2009), this is not always the case. Hos- sponses to the studied disturbance when viewed across
sack et al. (2013), studying salamander response to fire many systems. Fox (2011), for example, notes that dry,
in Montana, found little change in abundance during disturbance-prone grasslands generically have fewer
the period immediately after wildfire but significant granivores than other grasslands; these systems would
declines seven to twenty-one years after disturbance. therefore be expected to have different responses to any
Because we lack very many long-term studies, the gen- specific fire because they are already adapted to the oc-
148 research and conservation

currence of frequent fires. Assuming that a species can the first five years after the burn. However, unlike other
persist through a disturbance and within the immedi- taxa, some bird species are specifically adapted to these
ate postdisturbance landscape, likely the postdistur- immediate postfire conditions, making their study in
bance vegetative trajectory is more important than the these environments especially useful. Notable among
time since disturbance. Monamy and Fox (2000), for the bark insectivores is the black-backed woodpecker
example, found that recolonization trajectories for an (Picoides arcticus). This species colonizes burned areas
old forest specialist differed between two fires based within one year after a fire, occupies burned areas for
on time since disturbance but that the two trajectories three to five years, and peaks in density around three
were concordant when vegetative trajectory rather years after fire (Caton 1996; Saab et al. 2007). Outside
than time was evaluated. This pattern appears logical of this window, when postfire environments are opti-
and is likely general. If this is the case, then standard mal, black-backed woodpeckers are difficult to locate
approaches to successional studies where time since and are distributed diffusely feeding, opportunistically,
disturbance is used to characterize organism response on recently killed trees (Tremblay et al. 2010, Dudley
are likely produce weaker results than studies that mea- et al. 2012).
sure vegetative conditions directly.
Succession
Bird Studies Where Evaluation Close to the Defining Succession
Disturbance Event is Essential
Succession is an idea that emerged in the late 1800s
Clearly, more common and replicable sorts of dis- and was championed by Clements (1916). He en-
turbance, like fire, will be more likely to engender visioned the process as an orderly series of plant as-
meaningful adaptations. However, even for common semblages from the point of initiation through to the
disturbance types, such as fire in dry environments, im- “climax,” a stable state. Just as we define disturbance
mediate postfire conditions will be relatively rare on as events that we find disturbing given our viewpoint,
the landscape. To take advantage of these conditions, similarly, we tend to consider succession in terms of
population responses must be quick and the organisms processes that scale reasonably to our lifetimes. For
need either to be common enough to be on or adjacent these reasons, the genesis of succession as a concept
to the fire when it occurs or to be mobile enough to began with the observations of the dynamics of forests
find these disturbances quickly so as to take advantage (Clements 1916) and is most commonly still evoked in
of the ephemeral postdisturbance environment. It is this context. A great deal of thinking about this con-
not surprising that, for example, deer mice show large cept has occurred since Clements wrote “Plant Succes-
increases in postfire environments. They are general- sion: An Analysis of the Development of Vegetation”
ists and therefore present in the area, can reproduce in 1916, but the underlying ideas have remained re-
rapidly, and normally favor bare areas over dense grass markably stable. Primarily, the concept has been em-
(Pearson et al. 2001) and should therefore prosper in bellished, incorporating ideas such as “secondary” suc-
postfire environments. Similarly, avian species, being cession, in which initiation contains elements of the
highly mobile, can locate desired environments includ- predisturbance ecosystem, and various “disclimaxes,”
ing recent disturbances and can evolve to take advan- where an area is prevented from evolving into its pre-
tage of these conditions. For these reasons, the study sumed true climax by processes such as frequent dis-
of avifauna in postfire environments is both active turbance or grazing (see Meeker and Merkel 1984 for
and biologically appropriate. Saab and Powell (2005), a discussion). As mentioned previously, succession has
in a review of literature on over two hundred species retained popularity in many fields, particularly where
of birds and their associations with fire, showed that ecosystems are simple and primarily composed of na-
aerial, ground, and bark insectivores clearly favored tive species. Even in these simple systems, however,
recently burned habitats whereas foliage gleaners pre- there is a great deal of variability in these trajectories
ferred unburned habitats. As with other studies, virtu- due to the effects of pathogens and increasingly due to
ally all of the associated papers evaluated effects within novel weather patterns owing to climate change. For
 effects of disturbance and succession on wildlife habitat 149

example, Rehfeldt et al. (2009) showed that histori- phenomena across broad scales of space and time are
cally novel weather patterns, but patterns that are pre- necessarily Gleasonian, whereas fields that study small,
dicted to become increasingly common in the future, local phenomena tend to be Clemensian (Hortal et al.
immediately preceded major aspen (Populus tremuloi- 2012). The effects of rapid directional climate change
des) die-offs in Colorado. on this debate are to shrink the spatiotemporal domain
One critical aspect of successional thinking is that, across which constancy is a reasonable conceit. We are
although the initiation conditions might vary as would, relatively sure that climates will change significantly
to a certain extent, the trajectory toward climax con- over the next several centuries and that climate change
ditions, the climax conditions are invariant. As Horn is already affecting ecosystems, but our understandings
(1975) noted, an invariant steady state is a property of both the details of local climate change and, even
of stochastic systems with constant transition proba- more critically, the effects of these changes on habitat
bilities (Markov chains). Usher (1979) observed that and the wildlife that these habitats support are vague
if succession is described as a Markov process, then (Walther et al. 2002). This uncertainty is obviously in-
transition probabilities are likely not constant and will creased by the continuous entry of additional exotic
be affected by many factors such as population pro- species whose interactions with current ecosystems
cesses. Cadenasso et al. (2002), for example, showed are largely unknown. There is a school of thought that
that levels of mammal herbivory play a key role in believes that these specifics, when taken together, are
early successional trajectories of old fields. With many important enough to render the entire concept of suc-
exotic species entering ecosystems, the probability of cession suspect (Ricklefs 2008).
unforeseen biological feedback obviously increases.
For example, the successional trajectories of western
Wildlife Research on the Effects
North American grasslands to disturbance have been
of Succession
radically altered by the introduction of a plethora of ex-
otic plant species. In western Montana, spotted knap- The overwhelming number of disturbance papers that
weed (Centaurea maculosa) has occupied vast areas to measure immediate postdisturbance responses and
the detriment of both native bunch grasses and native the associated lack of longer-term studies dictates that
forbs (Callaway et al. 1999). In an effort to control the direct study of the site-level effects of succession
knapweed, exotic gall flies were introduced. They failed are sparse to nonexistent. One of the few cases where
to control the knapweed but instead became super- long-term studies on primary successional dynamics
abundant, leading to a variety of secondary trajectories are ongoing is within the area affected by the pyroclas-
involving increased densities of deer mice and associ- tic explosion of Mount Saint Helens (e.g., Spear et al.
ated hantavirus (Pearson and Callaway 2006). These 2012). Because large-scale population-level studies of
grass systems continue to be inundated by new exotics the effects of succession on wildlife are difficult and
(e.g., leafy spurge [Euphorbia esula]) and with climate seldom done, most of our understandings of forest
change, other species, such as yellow star-thistle (Cen- succession on wildlife are associated with habitat use
taurea solstitialis), common in adjacent states, may gain data. For example, we know that some species have
foothold. strong associations with certain habitats and that these
For a system to move in an orderly manner from dis- habitats are related to disturbance. The Florida sand
turbance toward a steady-state climax, both the species skink (Plestiodon reynoldsi), for example, is obligately
involved and the climatic background need to remain associated with Florida scrub habitat (Schrey et al.
constant. This strong requirement for constancy was 2011), a cover type that requires high-intensity fire on
first challenged by Gleason (1927), who argued that a a five-to-eighty-year time scale (Laessle 1958). Know-
plethora of factors lead to unique postdisturbance tra- ing this habitat association, we can infer the relation-
jectories. In the eighty-three years that have elapsed ship between this species and succession. Given this
since Gleason’s paper was published, the use and accep- constraint—that we understand habitat relationships
tance of successional concepts has split along disciplin- and know the disturbance regime associated with a
ary lines: disciplines like phylogeography that evaluate particular vegetative community—there is a vast lit-
150 research and conservation

erature on the effects of succession on wildlife (see reasonable to ask whether it is a necessary approach.
Hunter and Schmiegelow 2011 for a recent review). We could, rather than seek to quantify the specific
Interestingly, a great deal of this work focuses on the effects of specific disturbances, endeavor to design
two ends of the successional spectrum: early succes- studies that would allow comparison and generality
sional conditions associated with the immediate af- between disturbances. In the case of hurricanes on the
termath of disturbances and the study of a variety of eastern coast of the United States, for example, rather
organisms that are dependent on old forests. This may than studying the effects of Hugo, one could ask how
be because organisms of interest are associated with much the immediate postdisturbance conditions re-
these two seres. For ungulates, likely the most heavily lated to Hugo resemble those of other recent strong
studied mammal group, the general understanding is “Cape Verde–type” hurricanes that made landfall on
that early seral environments are preferred due to both the southeastern coast of the United States (e.g., Fran
the encouragement of forage and increased forage nu- [1996]; Floyd [1998]; Isabel [2003]). One paper, Rit-
tritional content (e.g., Hobbs and Spowart 1984), and tenhouse et al. (2010), attempted to place the effects
a great deal of focus is therefore on the generation and of hurricanes in a broader context, including both
maintenance of these conditions. Similarly, because immediate poststorm effects and overall changes in
very old forests are in decline worldwide, dependent species composition for birds. Rittenhouse used the
species often follow this trend and may become endan- North American Breeding Bird Survey (BBS; Link and
gered. Species such as the spotted owl (Strix occiden- Sauer 1998) to infer species abundance patterns for
talis) and red-cockaded woodpecker (Picoides borealis) the period 1967–2005. This period included a number
have become iconic in this regard (Bart and Forsman of hurricane landfalls, notably Hurricane Hugo (Sep-
1992; Ligon et al. 1986). These older forests are not tember 1989). Study areas were chosen based on the
only in decline but often are associated with higher intersection of hurricane tracks and a time series of
levels of biological diversity when compared to earlier annual or biennial satellite imagery. The main effect of
seres (e.g., Díaz et al. 2005). The emphasis on early hurricanes was seen in the year immediately following
seral and old growth forests is also due to a perceived the hurricane, where modest decreases in abundance
overabundance of mid-aged forests owing to a variety of and increases in species richness were similar across
human activities. Under natural disturbance regimes, focal areas. Interestingly, although hurricanes caused
forest age structures frequently follow negative expo- a threefold increase in the extent of disturbed forest
nential or Weibull distributions (Van Wagner 1978), within the focal areas, this factor was associated with
or distributions containing significant areas of both changes in community similarity for only three of the
old forest and areas that have recently been disturbed. thirteen avian groups examined and was not associated
Conversely, human-caused disturbance generally leads with patterns of avian abundance or species richness.
to truncated age distributions and can simultaneously Rittenhouse et al. (2010) attribute the lack of rela-
decrease early seral conditions through activities such tionships between disturbance levels and population
as fire suppression (e.g., Betts et al. 2010). responses to landscape scale heterogeneity; refuges
were present in areas where intact forests were the pre-
disturbance land cover. This understanding is similar
Disturbance Viewed in a Landscape
to that of Brotons et al. (2005), who found that local
Context: Studies of Both Disturbance
abundances in areas exterior to disturbances affected
and Succession
postdisturbance abundances within the disturbed
In the case of black-backed woodpeckers, postfire con- area. Thus, to understand postdisturbance trajectories,
ditions are clearly essential habitat, but for many other evaluating the larger landscape may be essential. While
species these environments are rare and noncritical. Rittenhouse et al. (2010) were able to analyze data at
However that may be, based on this review, it is clear spatial and temporal scales sufficient to infer general-
that targeted studies of disturbance occur almost ex- ized relationships, studying disturbances at this scale
clusively during the immediate aftermath of the event. is difficult and was only possible in this case because
While it is clear that this has been our approach, it is of the presence of the BBS, which is relatively unique
 effects of disturbance and succession on wildlife habitat 151

both in its longevity and in its spatially comprehensive- clear. In some cases, low-severity fire appears to have
ness. However, even though Rittenhouse et al. (2010) little effect (Roberts et al. 2011). Clark et al. (2013)
were able to apply forty years of BBS data, because hur- associated negative effects with fire, but sites they
ricanes occurred sequentially within the time period, studied had been burned and salvage logged, and pre-
statistical statements were limited to a period within fire timber harvest had occurred. Due to sample size
ten years after disturbance. constraints, Clark et al. (2013) could not separate an-
A rare example of a meta-analysis of both distur- thropogenic and natural disturbance factors. Further,
bance and succession is found in Schieck and Song while Franklin et al. (2000) found short-term negative
(2006). They performed a meta-analysis of studies on effects, the heterogeneous fire-generated landscape
boreal forest birds for a period of 0–125 years after of northwestern California featured high reproduc-
disturbance. Interestingly, they found that while bird tive and survival rates (Franklin et al. 2000)—the fire
communities changed after a disturbance, composition mosaic in general appeared to produce excellent owl
changes were largely associated with numbers of oc- habitat. In short, the relationship between spotted owls
currences rather than actual changes in species com- and disturbance is complex and our understandings
position. All species with measurable counts (greater are still very nebulous. Using derived information to
than five) were present in all forest types. The ability of conserve spotted owls into the future is also compli-
organisms to persist in all forest types and conditions cated by the twin factors of climate change and exotic
obviates the need to colonize areas except for after the species introduction. Owl demographics are sensitive
most extreme events. It also suggests that immediate both to regional climate and local weather (Glenn et al.
postfire abundance studies may not provide informa- 2010). Further, fire frequency is anticipated to increase
tive data on the landscape-level responses of popula- in the Pacific Northwest with climate change (Mc-
tions to disturbance and succession. Kenzie et al. 2004). Lastly, the invasion of the barred
owl (Strix varia) is having a profound negative effect
on spotted owl demographics across its range (Dug-
Species Relationships to Disturbance and
ger et al. 2011), fundamentally changing the expected
Succession: The Case of the Spotted Owl
population responses to specific vegetative patterns. In
Given the current approach of conducting largely inde- short, the spotted owl provides a case study in all of
pendent and descriptive studies of postdisturbance or- the complexities highlighted in this paper. The effects
ganism response, it is reasonable to ask how well these of disturbance are complex, depending both on time
studies composite into generalized understandings for since disturbance and the scale at which the question
any organism. The spotted owl, being one of the most is asked. Effects of any specific type of disturbance are
heavily studied species, provides a good example of the difficult to quantify because, at the scale of spotted owl
difficulties associated with determining the relation- home ranges, a variety of disturbance types have often
ships between an organism and disturbance while ap- occurred: fire, logging, and postfire salvage. Lastly, our
plying this methodology. As mentioned previously, the understandings concerning the effects of postdistur-
spotted owl has become an iconic organism represent- bance succession on the quality of spotted owl habitat
ing an old forest obligate (Bart and Forsman 1992), and have been fundamentally altered by the invasion of an
for this reason, the long-term viability of owls in fire- exotic competitor.
prone forests has been questioned (Spies et al. 2006).
However, spotted owls exist in a wide variety of for-
Approaches to the Problem
ests, including some that are highly modified (Thome
et al. 1999). Short-term relationships to anthropogenic It is clear that our traditional approaches, involving
disturbance have been highly studied, with uniformly primarily small-scale, short-duration descriptive stud-
negative relationships (e.g., Forsman et al. 1984; Frank- ies within disturbed areas (and sometimes including
lin et al. 2000), except where the dominant prey are undisturbed control areas), while producing much
woodrats (Neotoma fuscipes; Thome et al. 1999). Fire knowledge of local importance, do not provide a co-
is much less heavily studied, and the effects are less herent approach to increasing our knowledge of the
152 research and conservation

more general effects of disturbance and succession on collect appropriate data to assess biological processes
wildlife. It is equally clear that the traditional approach across broader spatiotemporal domains. Because of the
of trading space for time to study succession is becom- high level of scientific design associated with targeted
ing less and less tenable due to the increasing pool of monitoring, intermediate short-term products can
exotic organisms and the directional nature of climate both increase the immediate utility of the monitoring
change. What is less clear is how to move forward in and provide feedback on its overall efficacy. Secondly,
a manner that leads to more rapid increases in our monitoring efforts should take advantage of new tech-
knowledge of these processes while existing in a world nologies and, specifically, genetic sampling (Schwartz
that is in a rapid state of biological flux. The simple et al. 2007). Not only do these methods often provide
answer is to expand our studies in both space and time. the least expensive approach to sampling species (e.g.,
This would allow the formalization of the context in Kendall and McKelvey 2008), but genetic data are
which disturbance occurs, including the effects of nearly unique in that, if the samples are retained, these
landscape pattern and habitat juxtaposition, and would samples can be reanalyzed at a later date. Even without
avoid invoking constancy assumptions to infer future the formal retention of genetic data, the past collection
states. There are, however, major problems associated of materials that contain genetic data has allowed ret-
with this approach. The first is cost. Increasing the rospective studies of populations. For example, Miller
size and duration of a study quadratically increases the and Waits (2003) were able to retrospectively calculate
costs. Further, funding large, long-term projects is diffi- the effective population size of grizzly bears in Yellow-
cult given the competitive grant paradigm under which stone National Park by analyzing 110 museum speci-
a large proportion of science is funded. Competitive mens collected between 1912 and 1981. Because of this
grants generally have both spending and time limits ability to retrospectively analyze data collected in the
that preclude their direct utility for long-term fund- past, genetically based monitoring is less dependent on
ing; long-term studies are often patched together by current technologies than are most other monitoring
acquiring multiple grants, often from different sources. approaches. With genetic data, new technologies can
Additionally, there are many ways that these programs be applied to extant samples, allowing a fully modern
can fail and many examples of historical failures due analysis of all data regardless of collection time. This
to funding drying up, data loss due to inadequate ar- ability also provides a safeguard against data loss and
chiving, failures to maintain quality control across time allows post-hoc testing quality controls: assays can be
and space, and poor initial designs leading to the collec- repeated both to recover data and to ensure that the
tion of large quantities of relatively useless data. Lastly, original analyses were correct.
there is the strong, but often unacknowledged, power
of serendipity associated with many independent stud-
Conclusions
ies. The ability to capture serendipity is decreased if
scientific resources divert from a large number of There has been little to no attempt in the literature to
small, highly independent studies and commit to a view disturbance in terms of the operational environ-
small number of highly directed, long-term projects. ment of the studied species; we define disturbances
There are, however, approaches that merge the at anthropocentric scales. While this is biologically
fields of science and monitoring and which may prove problematic, it is so ubiquitous that it is impossible to
useful in documenting landscape-level changes asso- write a review article without also adopting this con-
ciated with disturbance and subsequent succession. ceit. However, given this constraint, there is a vast body
Targeted monitoring (Yoccoz et al. 2001; Nichols and of literature associated with both the effects of distur-
Williams 2006) combines principles of scientific de- bance and succession on wildlife. However, due both
sign including clearly articulated goals and specified er- to practicalities and interest levels, the vast majority
ror levels with the large-scale, long-term requirements of specific studies on the effects of disturbance focus
for biological monitoring. Increasing the proportion of on the period immediately postdisturbance; few stud-
monitoring that is specifically targeted could allow a ies extend beyond five years postdisturbance. Further,
reallocation of extant long-term funding into efforts to most studies are small in scope and are not designed to
 effects of disturbance and succession on wildlife habitat 153

test concepts; most generalizations are based on com- sociated wildlife communities were following expecta-
positing these small-scale primary studies through lit- tions as well as when and where systems were changing
erature reviews and meta-analyses, of which there are in unexpected ways.
many. However, as the case of the spotted owl demon-
strates, it is difficult, even with many largely unrelated
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 12
Kathi L. Borgmann and
Wildlife Habitat
Restoration
Courtney J. Conway

A s the preceding chapters point out, many wild-

life species and the habitat they depend on are in
peril. However, opportunities exist to restore habitat
cline in species diversity is greater than at any other
time in human history, and the decline is due primar-
ily to loss or degradation of habitat (Millennium Eco-
for many imperiled wildlife species. But what is wildlife system Assessment 2005). As previous chapters point
habitat restoration? We begin this chapter by defining out, a substantial portion of some species’ geographic
habitat restoration and then provide recommendations range has been degraded due to one or more anthro-
on how to maximize success of future habitat restora- pogenic changes (Francis, this volume; Wiens and
tion efforts for wildlife. Finally, we evaluate whether Van Horne, this volume). Hence, the need to restore
we have been successful in restoring wildlife habitat wildlife habitat within degraded areas to help prevent
and supply recommendations to advance habitat res- further losses in biodiversity is now a worldwide fo-
toration. Successful restoration requires clear and cus (Suding 2011). In fact, numerous restoration proj-
explicit goals that are based on our best understand- ects have been initiated all over the world, including
ing of what the habitat was like prior to the disturb- large-scale salt-marsh restoration in the San Francisco
ing event. Ideally, a restoration project would include: Bay (EDAW et al. 2007), seabird colony restoration
(1) a summary of prerestoration conditions that define worldwide (Jones and Kress 2012), and restoration of
the existing status of wildlife populations and their the Everglades in south Florida (US Army Corps of En-
habitat; (2) a description of habitat features required gineers and South Florida Water Management District
by the focal or indicator species for persistence; (3) an 2006). Restoration has, in fact, become a familiar term
a priori description of measurable, quantitative metrics to many of us, but what is restoration? Restoration is
that define restoration goals and measures of success; “the process of assisting the recovery of an ecosystem
(4) a monitoring plan; (5) postrestoration comparisons that has been degraded, damaged, or destroyed” (So-
of habitat features and wildlife populations with adja- ciety for Ecological Restoration International Science
cent unmodified areas that are similar to the restora- and Policy Working Group 2004). Restoration can en-
tion site; and (6) expert review of the entire restoration compass many different aspects; we will focus more
plan (i.e., the five aforementioned components). narrowly on wildlife habitat restoration. Wildlife habi-
tat restoration describes efforts that focus specifically
on improving the resources necessary to promote oc-
Introduction
cupancy or improve habitat suitability for one or more
More than 50% of the land in six out of fourteen of the wildlife species (Hall et al. 1997; Morrison et al. 1998).
world’s biomes has been converted to anthropogenic We assume that habitat restoration efforts are a sub-
land uses (primarily agriculture) since the 1900s (Mil- set of ecological restoration efforts because they are
lennium Ecosystem Assessment 2005). The rate of de- directed explicitly at improving habitat conditions for
158 research and conservation

one or more focal species. However, because the term projects but did not provide information on any spe-
habitat is inherently species-specific, restoration ef- cific restoration action(s).
forts that seek to improve habitat conditions to pro- While reviewing these restoration studies, we noted
mote occupancy for one species may not necessarily problems that were common to many wildlife habitat
provide suitable habitat for other species. Some have restoration projects. Several habitat restoration efforts
argued that efforts to restore areas to explicitly benefit a (1) failed to consider the habitat requirements of spe-
single (or even a few) species are shortsighted, and that cies (Morrison 1998; Block et al. 2001; Miller and
management actions should instead target entire eco- Hobbs 2007; Maslo et al. 2011), (2) lacked quantitative
logical communities (Hobbs and Norton 1996; Palmer criteria to assess the degree of success of the restoration
et al. 1997; Menninger and Palmer 2006). Although effort (i.e., pre- or postrestoration monitoring) (Block
conservation of entire communities is often the goal et al. 2001), and (3) failed to adequately share the out-
of management effort, many habitat restoration efforts come with practitioners or the public (Block et al. 2001;
will necessarily focus on one or a few species due to leg- Bernhardt et al. 2005; Miller and Hobbs 2007; Suding
islative mandates or political necessity. In cases where 2011). Well-planned habitat restoration efforts can im-
wildlife habitat restoration is targeted at one or more prove habitat suitability for wildlife species, but many
focal species, restoration plans should also be designed habitat restoration efforts were conducted with a very
to not adversely affect other endemic species in the general understanding of the habitat requirements of
community. wildlife species. For example, beach stabilization plans
Our goal in this chapter is not to provide in-depth failed to consider nesting requirements of the federally
summaries of the numerous books and papers that have endangered piping plover (Charadrius melodus; Maslo
been written about habitat restoration (Hobbs and et al. 2011). Failing to consider habitat requirements of
Norton 1996; Block et al. 2001; Morrison 2001, 2002; the focal species or suite of species during restoration
Falk et al. 2006). Instead, our goals are to highlight plans can result in failure to achieve restoration goals at
some common problems shared by many past wildlife the expense of the species. Common among many res-
habitat restoration efforts, describe key components toration studies was the failure to adequately evaluate
of a successful habitat restoration project, and discuss the successes and failures of restoration efforts. Moni-
some steps practitioners can take to improve the effi- toring wildlife before and after restoration is required
ciency and maximize the effectiveness of future habitat to determine if restoration has been successful. Fail-
restoration efforts. To help identify areas where past ing to measure wildlife response to restoration stymies
restoration efforts could be improved, we first con- our ability to improve future restoration projects and
ducted a literature review to assess the effectiveness of leaves many practitioners with little guidance regard-
past wildlife habitat restoration projects. We searched ing how to learn from past habitat restoration efforts.
ISI Web of Knowledge (www.isiwebofknowldege.com) In the following discussion, we describe key compo-
for “habitat restoration” OR “habitat enhancement” nents of a successful habitat restoration project.
and also included an additional search for “restora-
tion AND habitat AND influence, OR impact*, OR
Developing a Restoration Plan
response* AND bird*, OR mammal*.” Our search un-
doubtedly overlooked many published (and even more A common criticism of many restoration projects is the
unpublished) restoration efforts, but we assume that lack of an explicit “desired state” or “desired outcome.”
the results of the search are representative of habi- Oftentimes, the desired outcome of restoration is to re-
tat restoration projects. This search resulted in 1,458 store an area to pre-European conditions (Thorpe and
references, but only 58 of these references contained Stanley 2011). However, sufficient details about the
information suitable for inclusion in our review (i.e., conditions that existed prior to European settlement
summarized restoration activities that were conducted are typically not available for most areas (Thorpe and
to benefit terrestrial wildlife species). The other 1,400 Stanley 2011). Hence, restoration efforts that seek to
papers merely mentioned habitat restoration in the return areas to pre-European conditions have led some
abstract or provided a summary of habitat restoration authors to criticize restoration ecology as being past ori-
 wildlife habitat restoration 159

ented and static (Hobbs and Norton 1996; Davis 2000; location or the relative abundance of species prior to
Choi 2007; Hobbs et al. 2011). Instead of focusing on human-induced changes, but we can use data collected
one point in time (i.e., pre-European settlement) as the within the region (historic or current) to suggest what
desired condition for all restoration efforts, we would species are likely absent from the proposed restoration
likely be better served by an approach that restores the site but present in nearby undisturbed sites. By com-
natural disturbance regime or removes the anthropo- paring historic conditions (conditions that were likely
genic factor(s) that resulted in suboptimal conditions. present prior to anthropogenic changes) with existing
The restoration point then becomes the predisturbance conditions at the proposed restoration site, we can
condition (i.e., the likely condition prior to anthropo- start to develop a detailed, quantitative list of desired
genic changes that created the need for restoration) postrestoration conditions (i.e., goals). For example,
and not some predetermined point in time. Careful if reference sites or information based on conditions
consideration of predisturbance data, where available, present prior to anthropogenic changes suggest that a
combined with conditions at suitable reference sites bird species that relies on willows for breeding declined
can often provide a meaningful benchmark for resto- in abundance, then a restoration action might be to re-
ration. If predisturbance conditions are not known, establish willow densities within a range that existed
restoration can be targeted so that the restored areas prior to anthropogenic changes or that exists currently
are near the middle of the full range of environmental at reference sites that support healthy populations of
variation that existed historically at the site (Hobbs and the species. Data sources that might help define con-
Norton 1996). In either case, restoration efforts should ditions prior to human-induced changes include the
focus less on returning an area to some static state that Breeding Bird Survey, Christmas Bird Counts, natural-
existed prior to disturbance in the future but instead ists’ records, and museum specimens, all of which may
on a longer-term goal of achieving a range of condi- help to identify the suite of species formerly present at
tions that will eventually create a self-sustaining system the proposed restoration site.
without the need for continual human intervention. In addition to examining predisturbance data, prac-
We suggest that the first step in developing a resto- titioners should also sample the existing conditions at
ration plan is to summarize what is known about past the proposed restoration site. Descriptions of existing
conditions at the site and evaluate how those differ conditions are useful for (1) identifying species or habi-
from existing conditions at the proposed site. This will tat components that have been lost, (2) determining
help answer the question: “Why does the site currently the environmental attributes that are most in need
need restoration?” Other useful questions to ask at the of restoration, (3) setting goals and benchmarks, and
outset include: For which suite of species has habitat (4) examining postrestoration changes. For wildlife
suitability or abundance declined? What ecosystem habitat restoration efforts, existing conditions should
functions did the site provide prior to anthropogenic include surveys of the plant and animal community
changes (that it no longer provides)? Documenting within the proposed restoration site. Comparisons to
the range of historic environmental conditions and the former survey data (if such data are available) or with
natural disturbance processes that likely existed prior nearby reference communities will help determine
to anthropogenic changes will help determine if par- the environmental attributes that are most in need of
ticular species have likely been lost from the site and restoration. For example, if no willow flycatchers (Em-
will help practitioners develop the goals of the resto- pidonax traillii) are present at a proposed restoration
ration project. Predisturbance accounts of the site or site, but predisturbance data from the same area sug-
conditions at nearby reference sites that support the gest that willow flycatchers were formerly abundant
suite of endemic species can be used to help guide prior to anthropogenic changes, then some important
the selection of the desired postrestoration condition habitat feature(s) are probably lacking at the proposed
(Morrison 2002; Society for Ecological Restoration In- site. An important point to consider is why willow fly-
ternational Science and Policy Working Group 2004). catchers no longer occur at the proposed site. Compari-
In some situations, practitioners will not have an accu- sons among reference sites, predisturbance conditions,
rate list of the species that once occurred in a particular and existing conditions will provide an indication of
160 research and conservation

what habitat components are suboptimal. Existing con- The key to successful restoration is to bring together as
dition or prerestoration data also provides the founda- much information as is known about the focal species
tion from which to compare restoration success after and the community to create a more informed resto-
restoration efforts have been implemented. With ex- ration plan. Although habitat restoration efforts often
isting condition data, practitioners can document the originate due to legislative or political impetus that
“effect size” achieved by their restoration efforts (i.e., focus on one focal species, we caution against focus-
the extent to which restoration actions changed key ing exclusively on a single species. Habitat restoration
environmental conditions). efforts that focus solely on individual species may have
In addition to examining predisturbance records, unintended consequences for the rest of the commu-
the niche requirements and limitations (e.g., dispersal nity, hence practitioners should consider other species
distance) of a suite of focal species (or species of high- in the community when designing a restoration plan.
est concern) should also be considered (Zedler 2000b;
George and Zack 2001; Scott et al. 2001; Morrison
Setting Goals
2002; Miller and Hobbs 2007). Failure to consider
site-specific and species-specific habitat requirements One of the most important components of a restoration
could result in failure to meet the project’s goals. For project is the identification of well- defined, quantita-
example, the project will likely not meet its goals if the tive goals (Hobbs and Norton 1996; Hobbs and Harris
proposed restoration site is not big enough to support 2001). Well-defined goals allow practitioners to com-
a viable population of one of the focal species. Thus, an municate the importance of the restoration project to
understanding of the niche requirements will be help- policy makers and the public as well as judge the suc-
ful to create an effective restoration plan (Morrison cess of restoration actions. Vague goals (e.g., “return the
2002; Rodewald, this volume). Additional factors to site to historic conditions”) are subjective and provide
consider include landscape context and spatial factors no means by which to judge the success of restoration
that can affect occupancy (George and Zack 2001; Scott actions. Because “success” in restoration ecology has
et al. 2001). For example, species that are sensitive to often been seen as a vague and subjective term (Zedler
fragmentation may not occupy a restored site if the site 2007), we need to include more explicit, quantitative
exists in a heavily fragmented landscape. Dispersal ca- measures with which to judge the outcome of habitat
pabilities and metapopulation structure should also be restoration efforts. Determining the extent of change
considered when designing a restoration plan (Scott for each metric that is considered biologically mean-
et al. 2001; Morrison 2002; Menninger and Palmer ingful to the suite of focal species, and doing so prior
2006). Restoration of physical habitat attributes may to restoration, is one way we can reduce the subjective
not result in the desired outcome (colonization and nature of determining the success of restoration efforts.
persistence of one or more desired species) if disper- The Society of Ecological Restoration suggests that an
sal barriers exit or if regional abundance is insufficient ecosystem is restored when the ecosystem is capable of
to allow for colonization of the newly restored area producing a self-sustaining population of species (So-
(Schrott et al. 2005; Menninger and Palmer 2006). For ciety for Ecological Restoration International Science
example, burning and mowing failed to increase plant and Policy Working Group 2004).
species diversity in prairie remnants in Iowa because Despite the recognized need to include metrics of
the patches were too small to support many of the fo- success in restoration efforts, only two (Klein et al.
cal species, and past land-use practices significantly re- 2007; Maslo et al. 2011) of the fifty-eight projects we
duced the local seed bank, limiting recruitment (Van reviewed provided information on the criteria used to
Dyke et al. 2007). Although all factors that may influ- measure the success of restoration efforts. Because few
ence populations cannot be assessed in every restora- projects provided criteria to assess the success of res-
tion project, identifying key ecological attributes and toration efforts, determining whether the actions im-
associated niche requirements will result in a more proved habitat for any wildlife species was difficult. Of
comprehensive plan that has a higher probability of en- the fifty-eight published papers that summarized habi-
hancing habitat conditions for the suite of focal species. tat restoration efforts, 62% of the studies claimed that
 wildlife habitat restoration 161

restoration actions were successful or partially suc- clude) the desired conditions. Clearly articulated goals
cessful, although no criteria were provided by which based on detailed analysis of the full-range of predis-
to judge the success. We could not determine from the turbance conditions (to the extent available), reference
paper if the restoration actions were successful in 22% conditions, species-specific needs, and societal values
of the projects because not enough information was in- will bolster support from policy makers and the public
cluded to determine if the project met its stated goals. and will make restoration projects more likely to suc-
Many past restoration projects, however, failed to ceed (Hobbs 2007; Miller and Hobbs 2007).
establish goals at all. Of the fifty-eight studies we re-
viewed in the published literature, only 14% described
Monitoring and Evaluating
an explicit goal of the restoration actions. Instead, pub-
lished restoration efforts typically addressed whether The field of restoration ecology has primarily focused
restoration actions (often vaguely defined) affected on community ecology and plant succession (Morrison
wildlife species as a post-hoc assessment. Of the stud- 1998; Young 2000) or on restoring ecosystem function
ies we reviewed, 50% examined the effects of restora- while taking an implicit approach to wildlife that can
tion actions on wildlife species several years after the best be described as “if you build it they will come”
actions took place. Failure to consider the impacts of (Palmer et al. 1997; Zedler 2000b). However, restor-
restoration on wildlife species at the outset can limit ing an area to a former vegetative community may not
our ability to infer success of habitat restoration efforts. always result in the restoration of habitat for endemic
While a restored area may appear visually improved, if wildlife species (Zedler 1996, 2000a; Maslo et al.
wildlife species that were once common (prior to dis- 2011). For example, prescribed fire and / or mechanical
turbance) do not reestablish in the restored area, habi- thinning treatments have been implemented on a large
tat restoration efforts may not have been successful. portion of western coniferous forests to reduce fire se-
Hence, endemic wildlife species provide a benchmark verity and restore forests to presettlement conditions
from which to assess habitat restoration goals. Previous (Moore et al. 1999; Dickson et al. 2009; Kalies et al.
reviews of restoration projects have also highlighted 2012). However, we cannot assume that these restora-
the failure of many restoration efforts to identify ex- tion projects improved habitat conditions for the suite
plicit goals; 20% of river restoration projects failed of endemic species without an assessment of the spe-
to establish restoration goals (Bernhardt et al. 2005). cies present. Monitoring endemic wildlife can provide
Moreover, coordinators of restoration / restoration proj- an effective means to measure the extent to which a
ects could rarely provide information about project ob- restoration project has returned the site to its natural
jectives in a manner that described the desired condi- state. If the goal of a restoration project is to return
tions following restoration (Follstad Shah et al. 2007). a site to its natural condition, we would expect that
Restoration goals should be set in conjunction with the suite of species associated with pine woodlands, for
the development of the desired conditions described example, would be present, and populations would be
previously. Goals should be realistic, reviewed by out- self-sustaining. If one of the goals of a restoration effort
side experts, and guided by the needs of the focal spe- is to improve habitat suitability for endemic wildlife,
cies (Hobbs and Norton 1996; Morrison 2002). When postrestoration monitoring efforts must be included
setting goals, practitioners should take into account the as an integral part of the restoration project to docu-
ecological constraints of the system as well as social ment whether project goals are met. Surprisingly, res-
and financial constraints that are likely to affect the toration projects designed to benefit wildlife often fail
project (Ehrenfeld 2000; Morrison 2002; Miller and to include pre- or postrestoration wildlife monitoring
Hobbs 2007). If, for example, the proposed restoration (Block et al. 2001; Bernhardt et al. 2005). For example,
site is too small to support the desired species, and the past reviews of restoration projects found that only 10%
area surrounding the site is nonhabitat or cannot be of river restoration projects included an assessment of
restored, the ecological constraints of the system will any plant or wildlife species (Bernhardt et al. 2005).
likely prevent the goals from being attained. In most Although many of these river restoration projects were
cases, project goals should equate to (or at least in- not designed as wildlife habitat restoration projects,
162 research and conservation

we suggest that monitoring wildlife species should be different conditions as the control site. However, the
incorporated into the project as another method to effects of restoration actions cannot be accurately as-
judge project success. Lack of pre- and postrestoration certained if prerestoration data are not collected and
monitoring efforts greatly reduces our ability to infer no replication exists. Other restoration projects mea-
whether restoration actions have been successful. Fail- sured conditions before and after restoration but did
ure to incorporate a monitoring plan in conjunction not measure the same conditions at control or refer-
with the restoration plan could also lead to inadequate ence sites. Lack of control sites does not allow one to
or inappropriate monitoring. For example, if the resto- account for more widespread (i.e., regional) changes
ration site is small, attempting to estimate abundance that may be responsible for changes over time at the
of multiple wildlife species and species richness will restoration site. For example, a region-wide drought
likely not provide enough information to assess the ef- may cause the abundance of the focal species to de-
fectiveness of the restoration (Block et al. 2001). Thus, crease at the restoration site and hence hinder one’s
practitioners should determine the most appropriate ability to assess the effectiveness of the restoration
monitoring metrics while developing a monitoring actions if data from reference sites are not available.
plan. The parameters chosen as the focus of a monitor- Although 90% of the fifty- eight publications we re-
ing effort would relate directly back to a project goal. viewed monitored a control site, only 19% of projects
Morrison (2002) provides details regarding what to monitored the restoration and control sites both before
monitor and how to monitor wildlife habitat restora- and after restoration. Other reviews have also identi-
tion projects. Most restoration projects that monitor fied a failure to include monitoring of control sites in
wildlife species often record abundance of the focal most restoration efforts. Less than one-third of river
species after restoration. However, practitioners may restoration projects in the southwestern United States
also want to consider measuring productivity as abun- included monitoring at reference or control sites (Foll-
dance can be a misleading indicator of habitat quality stad Shah et al. 2007). In another recent review of 240
(Van Horne 1983). Of the fifty-eight published proj- studies, only 20% included prerestoration monitoring
ects we reviewed, 81% did not incorporate measures and only 58% compared changes at the restoration site
of productivity into their monitoring plan. Measuring to those at control sites (Jones and Schmitz 2009).
productivity can be costly but doing so helps to ensure Before-after control-impact (BACI) designs are the
that habitat restoration efforts do not merely create a preferred alternative for evaluating the effectiveness of
habitat sink for focal species. restoration efforts. BACI designs, as the name implies,
Many restoration efforts focus on a single site (with- compare metrics before and after restoration at treated
out any spatial replication), which limits the inferences sites and control sites to determine the impact of a
that can be made regarding the effectiveness of the res- treatment (i.e., restoration actions). A BACI design is
toration actions elsewhere. Hence, developing an ap- optimal because this sampling design allows research-
propriate sampling design that adequately assesses the ers to assess preexisting conditions (useful for develop-
wildlife responses to restoration is critical to improve ing desired conditions) and to distinguish natural varia-
our ability to make inferences beyond the boundaries tion from effects caused by restoration actions. A BACI
of any one restoration site. However, some past stud- study without replication (i.e., only one restoration
ies have used suboptimal sampling designs to monitor site) cannot make inferences beyond the specific study
responses to restoration. Of the fifty-eight publications site but is still a great improvement over restoration ef-
we reviewed, 78% did not include any measurements forts that do not include any prerestoration monitoring
of site conditions prior to restoration, which limits the or a control site. Monitoring of restoration effective-
conclusions researchers can make regarding changes ness is viewed by many restoration practitioners as too
in wildlife populations as a result of restoration. Of- cost prohibitive, but monitoring provides the only way
ten these studies measured wildlife response at a site to determine whether the thousands of dollars spent
that was enhanced at some point in the past and mea- on restoration actions were worthwhile. Thus, it is
sured the same metrics at a control or reference site to critical to develop a well-thought-out restoration and
determine if the enhanced site exhibited the same or monitoring plan that carefully balances costs against
 wildlife habitat restoration 163

monitoring needs. In our experiences, restoration proj- cess will improve if practitioners estimate the timing of
ects either seek to monitor too many metrics that are anticipated responses of both vegetation and the suite
not aligned with project goals or the metrics monitored of focal wildlife species to restoration actions and use
are not sufficient. Both scenarios tell us little about the those estimates to determine the duration of postres-
effects of restoration actions. Developing a detailed, toration monitoring efforts.
quantitative restoration plan that undergoes external Restoration projects will be successful if they in-
peer review by experts is key to the successful imple- clude an a priori monitoring plan with detailed descrip-
mentation of restoration activities that will likely have tions of what, how, when, and how long to monitor the
a positive effect on wildlife species (Morrison 2002). project site. Monitoring plans must also be adaptive. An
Restoration projects must also consider how long adaptive design is often a key component to a success-
to continue postrestoration monitoring efforts. Of the ful monitoring plan (Block et al. 2001; Lindenmayer
fifty-eight studies we reviewed, 48% monitored wild- and Likens 2009; Block 2013). Adaptive monitoring
life for two or fewer years after restoration efforts were plans include benchmarks or thresholds that trigger ad-
implemented. However, the full effectiveness of many ditional actions to improve the chances of meeting the
restoration actions may not be realized until years af- project goals. Establishing thresholds or benchmarks
ter the restoration actions are implemented (Jones and will help determine whether postrestoration monitor-
Schmitz 2009). The duration of monitoring should be ing efforts need to be extended and may help avert a
long enough to account for the time it will likely re- crisis if effective changes are implemented when those
quire the plant community to respond to restoration thresholds are not met. When developing benchmarks,
actions or to produce optimal habitat conditions for an important aspect to consider is the natural range of
the focal species. For example, if vegetation height is variability within a community for each metric. Deter-
an important habitat component for one of the focal mining where each metric lies in relation to the natural
species, monitoring abundance of the focal species variability of the metric across the area sampled can
before the vegetation has had a chance to respond to help determine whether restoration efforts are work-
restoration actions will not allow for assessment of ing or whether modifications are needed (Hobbs and
the effectiveness of the restoration actions. A more Norton 1996).
appropriate monitoring plan would be to monitor the
abiotic conditions that affect vegetation height in the
Are We Achieving?
first couple of years following restoration to ensure that
vegetation growth is on schedule and then monitor the In 1937 the Pittman-Robertson Act made funds avail-
suite of focal wildlife species in subsequent years. Criti- able for wildlife restoration. Since that time, the
cal thresholds or triggers should also be considered Pittman-Robertson Act has helped improve 41,962,211
when determining how long to plan postrestoration acres of wildlife habitat, provided improvements to
monitoring activities. For example, waiting five years 604,659 acres of waterfowl impoundments, and funded
after restoration actions to sample salt-marsh vegeta- thousands of research projects (US Fish and Wildlife
tion may be sufficient if the vegetation is not expected Service 2012). But the real question is: have these im-
to reach desired condition until year five. One caveat, provements benefited wildlife? Has habitat restoration
however, is that monitoring at only one point in time been successful?
could prove costly if vegetation does not respond to In our review of habitat restoration for mammals
restoration activities as expected and a small change and birds, 62% reported that restoration was at least
soon after restoration could have fixed the problem. partially successful. Several other studies have also
Thus, detailed monitoring could be based on the time conducted reviews of restoration projects to determine
interval established for specific triggers, but more qual- how successful we have been in restoring habitats and
itative assessment of habitat conditions could occur at ecosystems. Jones and Schmitz (2009) examined 240
shorter intervals to ensure that the restoration project ecosystem recovery studies published from 1910 to
is on target to meet the project’s goals. Whatever the 2008 and found that restoration was completely suc-
duration of postrestoration monitoring chosen, suc- cessful (all restoration goals were met) in 35% of eco-
164 research and conservation

system recovery studies examined and was partially lack the information necessary from which to judge
successful (some, but not all, restoration goals were project success (Follstad Shah et al. 2007).
met) in another 38% of studies. In another review of
restoration projects conducted in a variety of ecologi-
Advancing Habitat Restoration
cal communities across the globe, 44% of eighty-nine
restoration projects resulted in increased measures of The field of restoration has been criticized for lacking
biodiversity (abundance, species richness, or diversity) coherent general principles that allow for the transfer
compared to prerestoration levels but lower relative to of knowledge among practitioners (Hobbs and Norton
reference sites (Rey Benayas et al. 2009). In a third 1996; Suding 2011). One factor that contributes to
review of 621 wetland restoration projects, biological poor knowledge transfer and limits our ability to use
structure remained 26% lower compared to reference past knowledge to create more effective restoration
sites even ten years postrestoration (Moreno-Mateos plans is the lack of publication of successes or failures
et al. 2012). The proportion of restoration projects (Zedler 2000b; Follstad Shah et al. 2007; Suding 2011).
that were deemed successful varied among the three In fact, results of many restoration studies are fre-
reviews of restoration projects; however, determining quently not even reported or are only available in gray
project success is challenging and depends on the met- literature, further limiting knowledge transfer (Folls-
rics used to measure success of the restoration project tad Shah et al. 2007; Suding 2011; Maslo et al. 2012).
and the stated goals of the restoration project. Of the For example, only 25% of river restoration projects in
fifty-eight restoration projects we reviewed, 62% indi- the southwestern United States shared results with
cated that restoration actions improved conditions, but agencies, funders, or program managers (Follstad Shah
this number likely overestimates the percentage of suc- et al. 2007). While increased publication of restoration
cessful restoration projects because unsuccessful res- efforts could help remedy this problem, a more effective
toration efforts are less likely to be published (Zedler solution would be to track restoration projects in a pub-
2007) and practitioners likely have a strong impetus lically accessible database as many restoration projects
to deem their project successful to appease their fund- may not be suitable for publication in peer-reviewed
ing sources. Objective measures of project success is journals (or no one person has the time or responsibil-
also difficult because few restoration efforts developed ity to do so), but they do contain valuable information
metrics to measure their successes or failures (Hobbs that can help improve future restoration efforts. A data-
and Norton 1996) and few determined a priori what base of restoration practices that includes goals, moni-
constitutes project success (Suding 2011). Only 8% toring metrics, progress, and outcomes for every resto-
of river restoration projects based project success on ration project will aid in tracking restoration progress
pre- determined criteria (Follstad Shah et al. 2007). as well as inform restoration practices. In an attempt
Moreover, reporting and dissemination of postrestora- to remedy the lack of accessible data on river restora-
tion monitoring results is rare (Suding 2011), and many tion projects, Bernhardt et al. (2005) synthesized river
restoration projects are not published in academic restoration projects into the National River Restora-
journals and are therefore difficult to summarize effec- tion Science Synthesis (NRRSS) database. The NRRSS
tively. In fact, 75% of river restoration projects did not database is exactly what restoration ecology needs to
disseminate the results of their projects (Follstad Shah move forward. This database contained information
et al. 2007). on more than 30,000 river restoration projects across
Are habitat restoration projects effectively enhanc- the United States, including project costs, goals, moni-
ing habitat suitability enough to increase the probabil- toring metrics, and outcomes (Bernhardt et al. 2005).
ity of occupancy for the suite of focal species? The an- Unfortunately, the National Biological Information In-
swer to this question is project specific, but collectively frastructure (NBII), which housed the database, was
the answer remains largely unknown because compre- terminated, and the data are currently not publically
hensive assessments of restoration success and failures accessible. A similar project called the Global Restora-
are uncommon (Suding 2011), and existing datasets tion Network (http: // www.globalrestorationnetwork
 wildlife habitat restoration 165

.org / ) was started by the Society for Restoration prises” to begin learning from unforeseen challenges in
Ecology, in which users input information on their restoration (Hobbs 2009). Restoration projects could
restoration projects. The database contains valuable also be updated annually in the database, to help track
information on monitoring, evaluation, and restora- pitfalls and successes. A database of habitat restoration
tion actions. However, entering information into the projects could provide numerous benefits; however,
database is voluntary and no entries have been added building a database that would be used by practitioners
since 2009. and keeping it updated has its challenges and will likely
A database that houses information on restoration only be successful if practitioners are required to enter
projects and is used by the restoration community is information into the database as a mandatory compo-
needed (Suding 2011). A restoration database (similar nent of funding or mitigation contracts for restoration
to the global restoration network or the NRRSS) could projects.
provide practitioners with a readily accessible source Wildlife habitat restoration has an increasingly im-
of information regarding the location of other similar portant role to play in our endeavor to conserve species
restoration projects, the restoration actions that were diversity, and the collective success of our future res-
most successful at improving habitat suitability for toration efforts will be maximized if (1) both restora-
one’s suite of focal wildlife species, and the restoration tion plans and monitoring plans are well designed and
actions that had little success. A restoration database summarize what is known about habitat requirements
could hold information about specific restoration ac- for the suite of focal wildlife species; (2) goals and ob-
tions that are often not reported in peer-reviewed jectives are clearly articulated, quantitative (i.e., easily
publications. In the fifty- eight published projects we measured), and linked directly to desired outcomes
reviewed, 21% of the projects did not report the size with performance criteria developed by which to judge
of the treated area, and 57% did not provide detailed project success; (3) restoration and associated monitor-
information on the restoration activities performed. ing activities are planned as part of a BACI design with
The database would also help keep track of the cumu- both spatial and temporal controls; and (4) details and
lative area of different vegetation communities that outcomes of restoration actions are readily accessible
have been part of restoration efforts and help identify in a database to ensure that practitioners learn from
areas or communities that are restoration priorities. past mistakes and build on the successes of past efforts.
Focusing restoration projects in strategic locations The mentality of “if we build it, they will come” gives a
is one mechanism that may help maximize the prob- false sense of confidence in the outcome and needs to
ability of success. The database could also include be replaced with carefully planned restoration projects
restoration projects in the planning phase, which that consider the needs of endemic wildlife species to
might promote more cross-disciplinary collaboration help maintain healthy and diverse ecosystems in our
and external peer review (and hence improvements). ever-changing environment.
Databases such as this can also provide information Note: Any mention of trade, firm, or product names
on restoration impacts at a larger scale as many in- in this chapter is for descriptive purposes only and does
dividual restoration projects are often conducted at a not imply endorsement by the U.S. government.
small scale with little spatial replication (Follstad Shah
et al. 2007). LITERATURE CITED
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restoration is the length of time required to see the K. Barnas, S. Brooks, J. Carr, S. Clayton, C. Dahm,
results of the restoration actions. Thus, the dissemi- J. Follstad-Shah, D. Galat, S. Gloss, P. Goodwin, D. Hart,
nation of project information is often delayed and the B. Hassett, R. Jenkinson, S. Katz, G. M. Kondolf, P. S. Lake,
R. Lave, J. L. Meyer, T. K. O’Donnell, L. Pagano, B. Powell,
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and E. Sudduth. 2005. “Synthesizing U.S. River Restoration
the project is progressing. In an attempt to remedy Efforts.” Science 308:636–37.
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Studies to Evaluate the Success of Ecological Restoration on Lindenmayer, D. B., and G. E. Likens. 2009. “Adaptive Monitor-
Wildlife.” Restoration Ecology 9:293–303. ing: A New Paradigm for Long-Term Research and Monitor-
Choi, Y. D. 2007. “Restoration Ecology to the Future: A Call for ing.” Trends in Ecology & Evolution 24:482–86.
New Paradigm.” Restoration Ecology 15:351–53. Maslo, B., J. Burger, and S. N. Handel. 2012. “Modeling Forag-
Davis, M. A. 2000. “ ‘Restoration’—A Misnomer?” Science ing Behavior of Piping Plovers to Evaluate Habitat Restora-
287:1203. tion Success.” Journal of Wildlife Management 76:181–88.
Dickson, B. G., B. R. Noon, C. H. Flather, S. Jentsch, and W. M. Maslo, B., S. N. Handel, and T. Pover. 2011. “Restoring Beaches
Block. 2009. “Quantifying the Multi-scale Response of for Atlantic Coast Piping Plovers (Charadrius melodus): A
Avifauna to Prescribed Fire Experiments in the Southwest Classification and Regression Tree Analysis of Nest-Site
United States.” Ecological Applications 19:608–21. Selection.” Restoration Ecology 19:194–203.
EDAW, Philip Williams and Associates, H. T. Harvey and As- Menninger, H. L., and M. A. Palmer. 2006. “Restoring Ecologi-
sociates, Brown and Caldwell, and Geomatrix. 2007. “South cal Communities: From Theory to Practice.” In Foundations
Bay Salt Pond Restoration Project: Final Environmental of Restoration Ecology, edited by D. A. Falk, M. A. Palmer,
Impact Statement / Report.” www.southbayrestoration.org. and J. B. Zedler. Island Press, Washington, D.C.
Ehrenfeld, J. G. 2000. “Defining the Limits of Restoration: The Millennium Ecosystem Assessment. 2005. Ecosystems and Hu-
Need for Realistic Goals.” Restoration Ecology 8:2–9. man Well-being: Synthesis. Island Press, Washington, D.C.
Falk, D. A., M. A. Palmer, and J. B. Zedler, eds. 2006. Founda- Miller, J. R., and R. J. Hobbs. 2007. “Habitat Restoration—
tions of Restoration Ecology. Island Press, Washington, D.C. Do We Know What We’re Doing?” Restoration Ecology
Follstad Shah, J. J., C. N. Dahm, S. P. Gloss, and E. S. Bern- 15:382–90.
hardt. 2007. “River and Riparian Restoration in the South- Moore, M. M., W. W. Covington, and P. Z. Fule. 1999. “Refer-
west: Results of the National River Restoration Science ence Conditions and Ecological Restoration: A Southwest-
Synthesis Project.” Restoration Ecology 15:550–62. ern Ponderosa Pine Perspective.” Ecological Applications
George, T. L., and S. Zack. 2001. “Spatial and Temporal Con- 9:1266–77.
sideration in Restoring Habitat for Wildlife.” Restoration Moreno-Mateos, D., M. E. Power, F. A. Comin, and R. Yock-
Ecology 9:272–79. teng. 2012. “Structural and Functional Loss in Restored
Hall, L. S., P. R. Krausman, and M. L. Morrison. 1997. “The Wetland Ecosystems.” Plos Biology 10(1): e1001247.
Habitat Concept and a Plea for Standard Terminology.” Morrison, M. L. 1998. “Letter to the Editor.” Restoration Ecology
Wildlife Society Bulletin 25:173–82. 6:133.
Hobbs, R. J. 2007. “Restoration Ecology: Are We Making an ———. 2001. “Introduction: Concepts of Wildlife and Wildlife
Impact?” Restoration Ecology 15:597–600. Habitat for Ecological Restoration.” Restoration Ecology
———. 2009. “Looking for the Silver Lining: Making the Most 9:251–52.
of Failure.” Restoration Ecology 17:1–3. ———. 2002. Wildlife Restoration: Techniques for Habitat Anal-
Hobbs, R. J., L. M. Hallett, P. R. Ehrlich, and H. A. Mooney. ysis and Animal Monitoring. Island Press, Washington D.C.
2011. “Intervention Ecology: Applying Ecological Science in Morrison, M. L., B. G. Marcot, and R. W. Mannan. 1998.
the Twenty-first Century.” BioScience 61:442–50. Wildlife-Habitat Relationships. University of Wisconsin Press,
Hobbs, R. J., and J. A. Harris. 2001. “Restoration Ecology: Madison.
Repairing the Earth’s Ecosystems in the New Millennium.” Palmer, M. A., R. F. Ambrose, and N. L. Poff. 1997. “Ecological
Restoration Ecology 9:239–46. Theory and Community Restoration Ecology.” Restoration
Hobbs, R. J., and D. A. Norton. 1996. “Towards a Conceptual Ecology 5:291–300.
Framework for Restoration Ecology.” Restoration Ecology Rey Benayas, J. M., A. C. Newton, A. Diaz, and J. M. Bullock.
4:93–110. 2009. “Enhancement of Biodiversity and Ecosystem Ser-
Jones, H. P., and S. W. Kress. 2012. “A Review of the World’s vices by Ecological Restoration: A Meta-analysis.” Science
Active Seabird Restoration Projects.” Journal of Wildlife 325:1121–24.
Management 76:2–9. Schrott, G. R., K. A. With, and A. W. King. 2005. “Demographic
Jones, H. P., and O. J. Schmitz. 2009. “Rapid Recovery of Dam- Limitations of the Ability of Habitat Restoration to Rescue
aged Ecosystems.” Plos One 4(5): e5653. Declining Populations.” Conservation Biology 19:1181–93.
Kalies, E. L., B. G. Dickson, C. L. Hambers, and W. W. Cov- Scott, T. A., W. Wehtje, and M. Wehtje. 2001. “The Need for
ington. 2012. “Community Occupancy Responses of Small Strategic Planning in Passive Restoration of Wildlife Popu-
Mammals to Restoration Treatments in Ponderosa Pine lations.” Restoration Ecology 9:262–71.
Forests, Northern Arizona, USA.” Ecological Applications Society for Ecological Restoration International Science and
22:204–17. Policy Working Group. 2004. “The SER International
Klein, L. R., S. R. Clayton, J. R. Alldredge, and P. Goodwin. Primer on Ecological Restoration.” www.ser.org.
2007. “Long-Term Monitoring and Evaluation of the Lower Suding, K. N. 2011. “Toward an Era of Restoration in Ecology:
Red River Meadow Restoration Project, Idaho, USA.” Resto- Successes, Failures, and Opportunities Ahead.” Annual
ration Ecology 15:223–39. Review of Ecology, Evolution, and Systematics 42:465–87.
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Thorpe, A. S., and A. G. Stanley. 2011. “Determining Appropri- Van Horne, B. 1983. “Density as a Misleading Indicator of Hab-
ate Goals for Restoration of Imperilled Communities and itat Quality.” Journal of Wildlife Management 47:893–901.
Species.” Journal of Applied Ecology 48:275–79. Young, T. P. 2000. “Restoration Ecology and Conservation
US Army Corps of Engineers and South Florida Water Manage- Biology.” Biological Conservation 92:73–83.
ment District. 2006. “Comprehensive Everglades Restora- Zedler, J. B. 1996. “Coastal Mitigation in Southern California:
tion Plan.” www.evergladesplan.org. The Need for a Regional Restoration Strategy.” Ecological
US Fish and Wildlife Service. 2012. “Wildlife Restoration Applications 6:84–93.
Program—Accomplishments.” http: // wsfrprograms.fws.gov ———, ed. 2000a. Handbook for Restoring Tidal Wetlands. CRC
/ subpages / grantprograms / WR / WR_Accomplishments.htm. Press, Boca Raton, Florida.
Van Dyke, F., J. D. Schmeling, S. Starkenburg, S. H. Yoo, ———. 2000b. “Progress in Wetland Restoration Ecology.”
and P. W. Stewart. 2007. “Responses of Plant and Bird Trends in Ecology & Evolution 15:402–7.
Communities to Prescribed Burning in Tallgrass Prairies.” ———. 2007. “Success: An Unclear, Subjective Descriptor of
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 ConClusion Synthesis for Advancing
Michael L. Morrison and
Heather A. Mathewson
Useful Knowledge
of Habitat
Unifying Themes or Many Directions?

A s we noted in the preface, the purpose of this book

is to deliver to a broad audience an understand-
ing of current thoughts on the multitude of impacts
settled on at least a basic set of principles concerning
habitat, this “broad audience” would at least be hear-
ing a somewhat similar message (we say “somewhat”
influencing wildlife and wildlife habitats and provide because we know scientists just prefer to keep their
recommendations for a path forward that will advance options open to disagree at any time). Although habitat
management and conservation. Habitat is certainly is an ecological concept and thus not within a strict
one of the most frequently used terms in animal and framework directly amenable for hypothesis testing per
plant ecology (Mathewson and Morrison, this vol- se, having scientists wandering around using habitat in
ume). Elementary school students are introduced substantially different contexts does not in any way ad-
to the term, and there is unlikely to be a hunter or vance ecological understanding and, ultimately, trans-
nature explorer who does not read and use the term lation into meaningful management options.
frequently. This widespread use of habitat is both a Thus, for this closing chapter we went through each
blessing and a curse. It is a blessing because a broad chapter and looked for common messages on habitat.
swath of people have some notion of what it is, which That is, are these authors converging on a common un-
opens up ample opportunity for education and hence derstanding of habitat and how it should be thought
conservation. Unfortunately, it is also a curse because about and studied? Was there a commonality in the
the scientific community has simply refused to come priority research needs expressed? What is the priority
to any meaningful conclusions on what habitat is and list of research needs? Here, we first briefly review with
how we should study, manipulate, and, ultimately, con- the intent of relating common sections within each
serve it. chapter. We then finish by discussing the implications
Hence, we come back to our aim of writing a book the authors of this book presented for advancing our
accessible to a “broad audience.” In the preface, we de- understanding of animal ecology.
fined this audience as being composed of advanced un-
dergraduate and graduate students in natural resource
Messages
management and conservation (e.g., wildlife, range,
conservation biology, recreation and parks); resource In the subsections that follow, we briefly highlight
managers at local, state, and federal levels (e.g., state common messages we gleaned from across the chap-
wildlife and parks departments, USFWS, BLM, Army ters. We are not attempting to review all topics that
Corps); and private land managers. Although this were raised but rather focus on topics that were raised
sounds like a difficult goal to achieve, we approached as important issues in multiple chapters and thus in
the task knowing that once the scientific community different contexts.
170 wildlife habitat conservation

Definitions Populations and Quality

Guthery and Strickland clearly identified a problem Rodewald discussed the general failure to explicitly
with use of the “habitat term” when they summarized link considerations of habitat with individual and
the many uses of the term in the scientific literature. population-level measures of habitat quality, which can
It seems that perhaps in the majority of cases, even then obscure identification of the highest-quality habi-
the author appears to be unsure of the meaning of the tat. She also noted that habitat studies have a heavy bias
term they used. As has been recommended for de- toward a single life stage or season, usually the breed-
cades, Guthery and Strickland urged authors to clearly ing season.
define their intended meaning in a paper and then Likewise, Guthery and Strickland noted that stud-
use that meaning throughout the paper (we will come ies have documented a relationship between recruit-
back to the issue of definitions in the following dis- ment and adult survival, which provides at least some
cussion). support for spatial or temporal differences in habitat
A theme throughout many of the chapters, raised (subcomponent) quality (for a defined area and time).
early on by Mathewson and Morrison, and Guth- However, they also note that many studies ignore the
ery and Strickland, is the need to clearly elucidate dilemma of density dependence, where in sustaining
the theoretical relationship being assumed between populations, increases in productivity entail decreases
what is being described or predicted (dependent vari- in survival and vice versa. Van Horne and Wiens car-
able) and the parameters chosen for measurement; ried a similar message, clearly identifying that time
that is, the habitat variables. Making this connection lags, legacy effects, or carryover effects also erode the
between animal and environment should force the match between habitat occupancy and habitat quality.
author to explain both the parameters and the spatial The discussions by Mathewson and Morrison, Guth-
and temporal scales relevant to the question(s) being ery and Strickland, Rodewald, and others in this book
asked. identify a clear theme where our studies of habitat
In the “classical” view of habitat, Guthery and quality, while informative and an advancement over
Strickland seek to explain where an organism lives, classical descriptive habitat studies, provide us with an
which includes biotic and abiotic requisites and time. incomplete picture at best—and an incorrect picture
They continue on to note that while this classical ver- in many cases—of what factors drive population per-
sion of habitat is logically a dead-end concept, except sistence through space and time. These issues bring us
for descriptive science, defining a minimal set of fac- back to the topic of the arbitrary nature of most study
tors that would describe an animal’s habitat in the areas (see also McKelvey chapter) and the uncertainty
classical sense is not a trivial activity. Thus, Guthery associated with the portion of the biological population
and Strickland go on to explain, as has been done for we chose to study. The issue of identifying the biologi-
decades (Johnson 1980), that a hierarchical approach cal population of study has been thoroughly discussed
can be adopted where a largely arbitrary area (level 1; (e.g., see Morrison 2012 and review therein) but has
e.g., study area) with (demographically, behaviorally) yet to translate into general application in field stud-
meaningful subdivisions (level 2) can be subdivided ies. The chapters in this book go a step further in em-
(level 3) and perhaps further subdivided (level 4). phasizing both the spatial and temporal components
Guthery and Strickland explain that the hierarchical of habitat quality determination, as well as the related
view of habitat represents sequential parsing; namely, and critical topic of potential density dependence. Van
the study of an entity composed of parts that are in Horne and Wiens continue this theme, noting that it
turn composed of parts and so on. They concluded that has long been known that the density of a species in
this approach provides a natural means of studying how an area can influence the range of habitats occupied.
animals partition time and occurrence in the third- When habitat occupancy varies with density, the char-
level units that we call habitats (or as many call “habitat acteristics of the area at a particular time will be sensi-
types”). tive to the species density at that time, which in turn
 synthesis for advancing useful knowledge of habitat 171

might not say much about the relationships with habi- being made now and involve very large areas of wildlife
tat that influence individual fitness. habitat.
Rodewald’s emphasis on separating individual- ver- A frequent theme throughout this book is the po-
sus population-level concepts of habitat quality is criti- tential and real impacts of fragmentation on animal
cal and relates directly back to the initial designation of populations, ranging from genetic changes to outright
the biological population being studied and the linked extinction. Smallwood focused specifically on frag-
spatial scale of study that such work should entail. mentation and related topics, while most of the other
As reviewed by Morrison (2012), most studies pick a chapter authors intertwined fragmentation into the
convenient study area (e.g., a park, ranch, or portions topic they were emphasizing. For example, biological
thereof) without any stated regard for the biological invasions by both plants and animals are substantially
population being sampled (this is in contrast to stating promoted by fragmentation (Lockwood and Burkhal-
the statistical population, which usually has little or no ter); fragmentation can result in a decrease in effective
meaningful relationship to the biological population). population size, increases in genetic drift and loss of
Clearly, individuals make up the biological population. genetic diversity, while increased isolation of fragments
As noted by Morrison (2012), it is how we sample from leads to decreases in gene flow and loss of genetic di-
within that biological population that holds the key to versity (Waits and Epps); reduction in patch size is a
making our studies meaningful in a broad ecological frequent outcome of human-related development ac-
and, ultimately, management-conservation sense. tivities (Francis); and, of course, fragmentation is a
major issue in the development of habitat restoration
plans (Borgmann and Conway).
Heterogeneity
Smallwood reviewed the fundamental problem we
The issue of habitat fragmentation is woven through- have in often not knowing why the animals we counted
out many chapters (e.g., Smallwood, Van Horne and or observed were located where we found them. The
Wiens, Waits, Francis, Lockwood and Burkhalter) and aggregations we encounter can be temporary based on
certainly qualifies as a major threat to the persistence resource availability, based on social interactions, con-
of many plant and animal species. However, concep- strained because of patch size, and so forth. Clearly,
tualizations of habitat fragmentation and corridors these issues substantially impact (i.e., bias) how we
vary widely in the literature, and the science support- view habitat use.
ing these concepts needs to improve (see Smallwood
chapter). The issues of spatial and temporal scales are
Applications
closely linked with how we view fragmentation. Small-
wood concluded that quantitative tests of fragmenta- Schwartz et al. explained why one must take care in
tion and corridor effectiveness should be conducted understanding the underlying biological distribution
that measure habitat variables rather than using overly of the entity (species, group of species, an index) of
simplistic maps of vegetation cover types; the effective- interest when designing a monitoring plan. It is not
ness of corridors is highly species specific. sufficient to simply “monitor,” because the pattern that
As detailed by Smallwood, definitions of habitat results over time might not reflect the actual changes
fragmentation and corridors need to be more explicit. in the biological entity of interest.
Scientists need to more effectively convey their con- Borgmann and Conway (see also Rodewald chapter)
cepts of habitat fragmentation and corridors to biolo- invoked the “if you build it, they will come” view of
gists in natural resource agencies and environmental restoration and management when discussing the ways
consulting firms, as well as to attorneys, political deci- that population dynamics can be influenced by individ-
sion makers, and the public. These concepts really mat- ual behaviors related to habitat selection, because such
ter where decisions are being made about whether and a view fails to recognize that a diverse suite of environ-
how to further fragment habitat, and how to mitigate mental, biological, and social cues are used by animals
the effects of this fragmentation. These decisions are to select habitats (e.g., conspecific attraction). Thus, in
172 wildlife habitat conservation

cases where cues for settlement are lacking, locations study is done and one must justify publication is an at-
that otherwise would be suitable for settlement would tempt made to justify extrapolation of results outside
not be occupied. This theme runs through many chap- the study area.
ters. For example, Van Horne and Wiens noted that it is This theme also directly links our conceptual un-
frequently assumed that places where a species occurs derstanding of how animals select a place to settle (i.e.,
contain good habitat and places where it is absent do habitat) with study design and thus study area identi-
not. It has long been known, however, that the density fication. Every chapter in this book discussed spatial
of a species in an area can influence the range of habi- (and temporal) scale in one or more contexts, empha-
tats occupied. Issues of source-sink, ecological traps, sizing that the process of habitat selection is widely
and related concepts are additional considerations. viewed as a hierarchical process wherein individuals
use a series of cues to decide ultimately where to be
and when to be there, what to eat, where to nest, and
Themes
so forth; Guthery and Strickland explicitly detailed
We did not begin this project with an a priori expecta- this process. Although this model of habitat selection
tion of what the authors would say, and we certainly did is widely held, it is seldom used as a basis for selec-
not expect that some grand answer would emerge from tion of a sampling area; naturally this issue relates di-
the chapters (or that we could even recognize it). We, rectly to our previous discussion of identification of the
like everyone reading this book, understand to varying biological population. Hence, this issue of the process
degrees of specificity that ecology is typified by interac- of habitat selection is a subpart of our discussion of
tions, and any understanding thereof requires a synthe- identification of the biological population.
sis of a lot of information. We are not sure if a grand A secondary but related theme deals directly with
answer did emerge, but we are confident that a grand habitat quality. Much attention across the chapters
direction did emerge. Although all of the authors were was devoted to explaining how measures of quality, be
provided drafts of each chapter, this was done so they they recruitment or individual survival, must be con-
could cross-reference related topics as they deemed ap- ducted on appropriate spatial (i.e., usually larger than
propriate; there was no attempt for them to all get their we study) and temporal (i.e., usually longer than we
stories straight. In writing this closing chapter, we went study) scales. The topic of habitat quality was discussed
through each chapter and looked for major conclusions throughout the chapters, with issues raised concerning
in hopes of finding a number of themes based on a simi- what metrics to measure (e.g., recruitment, survival),
larity in conclusions; the result was synthesized and the interacting issue of density dependence and its
summarized earlier. It became apparent very quickly influence on quality conclusions, and the interacting
that there was, indeed, a central theme that ran across issues of study area size and duration. The issue of spa-
all of the chapters. As would be expected in the field tial scale relates directly to the aforementioned major
of ecology, the theme is multiparted, with each sub- theme, while the issue of temporal scale also becomes
part inexorably linked; part of the theme is based on a study design conundrum. Ultimately, we want to
biology, while the other part is based on study design. know how populations persist in space and time. We
We can summarize the primary theme as: can summarize the embedded, secondary theme as:
Advancing understanding of animal ecology (and thus Advancing understanding of the biological population
habitat) requires that we identify and study a biological requires quantification of those metrics that explain popu-
population in appropriate space and time. lation persistence.
This primary theme might seem obvious, but after
reading the chapters in this book as well as other re- Terminology
lated literature (much of which was cited across the
Guthery and Strickland recommend that because habi-
chapters), we are confident that most of our studies
tat is a concept and thus intrinsically ambiguous, au-
pay little or (usually) no attention to the biological
thors should explicitly define their intended meaning
population when establishing the study. Only after the
 synthesis for advancing useful knowledge of habitat 173

and then unambiguously apply that meaning through-

Implications
out the paper. They thought this process would achieve
the goal of authors and readers having a common un- The ultimate conservation implications of these
derstanding of meaning. Morrison and Hall (2002) also themes are clear and directly related to how we design
recommended such an approach for defining habitat and manage land with the aim of population persis-
on a paper-by-paper basis. We think, however, that such tence. Specification of a biologically relevant study
a procedure—while certainly a useful fallback strat- area relates directly to planning for reserves and other
egy—is not the optimal way to advance how we view management-conservation areas. As Van Horne and
and study habitat. Defining terms does allow readers to Wiens and McKelvey explained, because of natural suc-
know what is meant, but it does not mean that anything cession, climate change, or processes or threats mov-
about the study (and thus paper) was done in a manner ing across the reserve boundary, conditions within a
that actually adds to our knowledge about how and why management area with small and fixed boundaries will
animals occur where they do. likely change in ways that cannot be managed for the
Thus, we think recommending definitions, while benefit of the species. Because most studies are small
necessary, is far from sufficient. Rather, we think that in scope, the generalizations they produce are based on
the necessary procedure is for each author to explicitly compositing these small-scale primary studies through
develop and justify how they identified the biological literature reviews and meta-analyses. It is difficult,
population, how study design decisions were made however, even with many largely unrelated small-scale
based on that population, and what ramifications studies of specific areas, to generate coherent under-
on results these decisions were likely to express. Al- standings of general relationships between an organ-
though many will complain that we are setting the bar ism and its environment. McKelvey clearly showed that
too high, we are simply asking that authors discuss the because the condition of any patch of land, regardless
biological population to the best of their ability. Journal of size, changes substantially though time, our usual
editors frequently reject manuscripts because of few strategy of creating conservation areas that are fixed in
or small study areas; such manuscripts are often then location is usually doomed to failure; or as Van Horne
published in an appropriate regional journal. All we are and Wiens pointed out, incredibly expensive to main-
asking is that authors be transparent. Morrison (2012) tain in a desired state.
discussed these issues in detail, including consider-
ations of subspecies and especially ecotypes as a basis LITERATURE CITED
for study design. In this book, Rodewald succinctly Johnson, D. H. 1980. “The Comparison of Usage and Avail-
summarizes the steps we need to take to move beyond ability Measurements for Evaluating Resource Preference.”
Ecology 61:65–71.
traditional approaches to studying wildlife-habitat Morrison, M. L. 2012. “The Habitat Sampling and Analysis
relationships. Fortunately, we now have a number of Paradigm has Limited Value in Animal Conservation: A
genetic tools (see Waits chapter) that allow us to more Prequel.” Journal of Wildlife Management 76:438–50.
easily explore issues of population structure and limits. Morrison, M. L., and L. S. Hall. 2002. “Standard Terminol-
There remains a role for what Guthery and Strickland ogy: Toward a Common Language to Advance Ecological
Understanding and Application.” In Predicting Species
termed “classical” habitat studies. However, we think
Occurrences: Issues of Accuracy and Scale, edited by J. M.
that failure to recognize and adopt the issues (themes) Scott, P. J. Heglund, M. L. Morrison, J. B. Haufler, M. G.
raised herein will doom us to slow and halting progress Raphael, W. A. Wall, and F. B. Samson, 43–52. Island Press,
in understanding population persistence. Washington, D.C.
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 Index

abiotic components of habitat, 10, 12, 16, Ambystoma tigrinum mavortium, 76 efforts for, 157, 159; spotted owl relation-
27, 118, 163, 170 American martens, 75 ship to, 151; wildlife avoidance responses
Able, K., 3, 4 American oystercatchers, 55 to, 55, 59
abundance of wildlife: density dependence American pikas, 73 anthropogenic stressors, 51, 53, 58, 59–60
and, 17; effects of restoration projects American redstarts, 23 Antilocapra americana, 23, 55
on, 164; habitat fragmentation and, Ammodramus savannarum, 37, 120 anti-predator defenses, 107, 108
92; habitat occupancy and, 37; habitat amphibians, 22, 68, 70, 105, 137 Aphelocoma californica, 57
quality and, 5, 49–50; human activities Amphispiza bilineata, 20 arbitrary area of interest, 13, 15, 16, 170
and, 59; monitoring of, 128–29, 131–33, amplified fragment length polymorphism Arcese, P., 110
138; postdisturbance, 147, 150; spatial (AFLP) analysis, 66, 75 Archilochus alexandri, 57
shifting of, 86; surrogate measures for, Amur honeysuckle, 105 Army Corps of Engineers, 97, 169
130; vegetation features and, 38. See also Anas fulvigula fulvigula, 71 Artemisiospiza belli, 38
density Anas platyrhynchos, 16, 70 arthropods, 5, 106
acid rain, 35 Anas superciliosa rogersi, 71 artificial night lighting, 25, 47, 53–55, 85;
Acridotheres tristis, 108 Anas superciliosa superciliosa, 71 effects on birds, 54; effects on green
adaptive genetic diversity, 63–64, 68, 69, Anas wyvilliana, 71 frog signaling behavior, 53; effects on
72, 74, 76–77 Andropogon gayanus, 106 sea turtle hatchlings, 25, 53; insects at-
adaptive locus, 64, 65, 68, 74 Andropogon sp., 11 tracted by, 54–55; measuring and quan-
adaptive management, 128, 129, 130–31, animal communities: effects of artificial tifying magnitude of effects of, 58–59;
137, 138 lighting on, 53, 54; effects of disturbance predation risk and, 53–54; strategies for
adaptive monitoring, 163 and succession on, 143–53; effects of management of, 53
Adelges tsugae, 109 habitat loss or degradation on, 56–58; Aslan, C. E., 110
African elephants, 36, 71, 76 effects of human presence on, 55, 56; aspen, 149
African wildcats, 71 monitoring of, 136; niche and, 5, 28; assessment, 129. See also monitoring
Ailuropoda melanoleuca, 36 restoration efforts and, 159–60 assumptions of constancy, 35, 143, 149, 152
Alces alces, 23 animal invasions, 107–8 attractive sinks, 49. See also ecological traps
Algonquin wolves, 71 Anolis sagrei, 71 Australia, 22, 34, 56, 106, 108
Allee effect, 27, 71, 104 anthropogenic activities, 5, 20, 38, 47, 59, Australian (Pacific) black ducks, 71
allele(s), 63; definition of, 65; fixed, in 150; hybridization due to, 70; impact
subpopulations, 68; habitat-related on gene flow and genetic diversity, 75; backcrossing of hybrids to parental groups,
variation in phenotype linked to, 72–73; impact on habit fragmentation and cor- 65, 70, 75
major histocompatibility complex, 69; ridors, 87, 91; recreational, 38, 40, 47, Ballari, S. A., 109
metrics of genetic structure, 74; positive 49, 51, 55–56, 59, 136, 169 barred owls, 37, 38, 151
selection for, 65, 67; purifying selection anthropogenic biomes, 47, 157 barred tiger salamanders, 69, 76
of, 65; spatial correlation of, 73; spatial anthropogenic disturbance, 47, 49, 53–59, barren-ground caribou, 23
gradients in fitness of, 68 90, 145–47, 150; artificial night lighting, Barrett, G. W., 95
allele frequency, 63, 65, 68, 73 53–55; due to recreational activities, Barrios- Garcia, M. N., 109
allelic richness (A r), 72, 73 55–56, 59; habitat loss due to, 49; log- beach mice, 53
Allendorf, F. W., 70 ging, land clearing, land conservation, beach stabilization plans, 158
allozymes, 64, 72, 73 145–46; measuring noise and light and bears, 36, 67, 73, 90, 152
Ambystoma californiense, 76 quantifying magnitude of their effects, beavers, 137, 145
Ambystoma tigrinum, 69 58–59; noise, 47, 51–53, 85; restoration Beck, J. L., 118
176 index

before-after control-impact (BACI) designs, blue tits, 24, 54, 69 Catharus guttatus, 147
125, 162, 165 bobolinks, 25 cats, domestic, interbreeding with native
behaviorally naive prey, 107, 108 bobwhites, 11, 13, 16, 17 species, 71
behavioral processes guiding habitat selec- Boiga irregularis, 107 Cenchrus ciliaris, 106
tion, 19–20, 24–27, 28, 171–72 boreal toads, 137 census population size (Nc), 65, 67, 68–69
Beier, P., 86, 87 Bormann, F. H., 144 Centaurea maculosa, 106, 149
Bender, D. J., 85 bottlenecks, 65, 67, 95 Centaurea solstitialis, 149
Bernhardt, E. S., 164 Brachylagus idahoensis, 72 Certhia americana, 147
Best, R. J., 110 Branta canadensis, 110 cerulean warblers, 22
bighorn sheep, 55, 67, 69, 73, 75 Breeding Bird Survey (BBS), 131, 150–51, Cervus canadensis, 129
biocontrol agent, 108 153, 159 Cervus elaphus, 55
biodiversity: effects of habitat loss and frag- breeding season, 19, 23–24, 106, 109, 131, Chapman, J. R., 69
mentation on, 34, 48, 84, 157; effects of 134, 170 Charadrius melodus, 158
restoration projects on, 157, 164; habitat Brook, B. W., 70 Cheirogaleus medius, 69
management for protection of, 35, 38, brook trout, 109 chestnut-backed chickadees, 36, 37, 38
40, 41; hybridization and, 70; invasive Brotons, L., 150 Chihuahuan Desert, 39
species and loss of, 110; monitoring of, Brown, C. J., 107 Chinook salmon, 68
130, 131, 136 Brown, C. R., 56 Christmas Bird Counts, 159
biological integrity, 132 Brown, M. B., 56 chromosomes, 64, 65, 76
biomes, 4, 10; anthropogenic, 47, 157 brown bears, 73 citizen science projects, 110, 138
biotic components of habitat, 4, 10, 12, 16, brown creepers, 147 Clark, D. A., 151
27, 104, 118, 170 brown-headed cowbirds, 50 classical view of habitat, 10, 16, 17, 170, 173
biparental inheritance, 64, 76 brown tree snakes, 107 Clean Water Act, 123
birds: acoustic cues and habitat selection Budeo swainsoni, 89 clear-cutting, 47, 145–46, 147
by, 50; behavioral influences on habitat Bufo boreas, 137 Clemensian concept of succession, 144,
selection by, 25–26; Breeding Bird bufogenin toxin, 108 148, 149
Survey, 131, 150–51, 153, 159; Christmas Bufo mariuns, 108 Clements, F. E., 148
Bird Counts, 159; conspecific attraction Buggs, R. J. A., 70 Clemmys insculpta, 55
among, 25, 50; definition of habitat Bureau of Land Management (BLM), 96, cliff swallows, 11, 56
for, 11; effect of Conservation Reserve 169 climate change, 4, 117, 128, 139, 153, 173;
Program on, 118, 123; effect of invasive Burmese python, 107 directional, 143, 145, 149, 152; fire
species on, 105–7, 108, 109–10; effects burned habitats, 21, 106, 107, 146–47, 148, frequency and, 151; habitat conserva-
of artificial lighting on, 54; effects of rec- 151, 160. See also fire tion and, 38, 39, 41; hybridization and,
reational activities on, 56; fire effects on, 70; invasive species and, 104; landscape
147; habitat management for, 41; habitat Cadenasso, M. L., 149 genetic studies of, 75; monitoring
quality for, 15, 21; habitat restoration California condors, 36 impacts of, 128, 139; nonnative species
programs for, 157, 158, 159, 163; habitat California Environmental Quality Act and, 104; spotted owl demographics
use across life cycles, 21–24; hybridiza- (CEQA), 84, 88 and, 151; succession and, 143, 145, 148,
tion in, 70; inbreeding depression in, 70; California tiger salamanders, 76 149, 152
Migratory Bird Treaty, 35; monitoring Calvert, A. M., 24 coalescent theory, 74
programs for, 121, 122, 123, 131, 134; Canada geese, 110 Colinis virginanus, 11
natural abode of, 14; nesting success of, cane toads, 108 Collaborative Forest Landscape Restoration
28; niche of, 5; noise effects on, 51–53, Canis familiaris, 71 Program, 40
57–58; outbreeding depression in, 68; Canis latrans, 71 common myna, 108
precocial, 146; studies of disturbance Canis lupus, 55, 69, 71, 138 community assembly theory, 145
effects on, 145, 146–47, 148, 150–51; Canis simensis, 71 community dynamics, 132
temporal dynamics influencing habitats canopy cover, 13, 17, 22, 50, 96, 109 connectivity, 39; demographic, across life
of, 37, 38. See also specific species Canyonlands National Park, 55 cycle, 19, 28; habitat fragmentation and,
bison, 55, 67 capercaillie, 14 48, 84, 85, 89, 90; least-cost models of,
Bison bison, 55 Capreolus capreolus, 23 75; migratory, 24, 27; between patches,
Bissonette, J. A., 147 capture-recapture models, 119, 132, 134 genetic diversity and, 67, 69, 73, 75;
black-backed woodpeckers, 148, 150 Cardinalis cardinalis, 26 population abundance and, 133, 144
black bears, 90 Caretta caretta, 25 conservation and management of habitats,
blackbirds, 54 Carolina hemlock, 109 7, 34–41, 173; adaptive management,
Blackburn, T. M., 102 Carpodacus mexicanus, 57 128, 129, 130–31, 137, 138; approaches
black-capped vireos, 25 carrying capacity, 13, 21, 27 to, 34–35; challenge of a changing world
black-chinned hummingbirds, 57, 58 carrying capacity density, 15 on, 39–40; climate change and, 38, 39,
black-throated blue warblers, 23, 27 carryover effects, 23–24, 27, 28, 37, 170 41; cooperative, 40; corridors for wildlife
black-throated sparrows, 20 Carya sp., 13 movement, 87–95; cost-effective, 38,
Block, W. M., 130 Castor canadensis, 137, 145 41; “natural” habitats, 47; proactive,
 index 177

117, 122; public support for, 36; realities density: capture-recapture models for esti- Dryocopus pileatus, 137
of, 35–39; reserves for, 38–39, 40, mation of, 134; effects of domestic dogs Duck Nesting Habitat Initiative, 123
86, 118, 122–23, 173; spatial variation on, 55–56; fitness density, 6, 14–15, ducks, 16, 70–71, 123
and, 38–39; species-specific, 35, 36; 16, 20; habitat occupancy and, 37, 38, dwarf lemurs, 69
systems-centered, 35; temporal variation 170; monitoring of, 132; Van Horne’s
and, 37–38, 39; variation within species mathematical definition of habitat qual- earthquakes, 121, 124
and their responses to habitat, 36–37; a ity and, 5, 9, 10, 14–15, 20, 37, 170, 172; earthworms, nonnative, 107
way forward for, 40–41; wildlife habitat variable relationship between fitness eastern bluebirds, 52
restoration, 157–65 and, 20; vegetation features and, 38. See Eastern chipmunks, 28
Conservation Reserve Program (CRP), 118, also abundance of wildlife eastern hemlock, 109
122–23 density dependence, 15, 17, 73, 170, 172; eastern towhees, 22
conspecific attraction, 19–20, 25, 50, 171 due to crowding effects, 24, 27; inverse, ecological integrity, 136–37
control, 118; before-after control-impact 20; population demographics and, 24, ecological systems, 39, 47, 58, 59, 119, 122.
designs, 125, 162, 165; over events, 26, 27; seasonal events and, 24 See also ecosystems
121–22 dependent variable(s), 170; in measuring ecological traps, 6, 20, 25, 37, 49, 53, 172
corridors for wildlife movement, 48; as responses to habitat fragmentation, 91, ecology of information, 24, 25
conservation effort, 87; definitions of, 92, 95 ecosites, 4, 6
86–87, 95, 171; descriptions in environ- desert bighorn sheep, 55, 67, 73, 75 ecosystem engineers, 105, 137
mental reviews, 89–91, 96–97; habitat destructive genetic sampling, 64 ecosystem processes, 35, 92, 134, 138
fragmentation and, 84–95; impact on detection probability, 134 ecosystems, 4; artificial lighting effects on,
population viability, 87, 92; invasive Didham, R. K., 110 54; in a changing world, 39; complexity
species and, 111; landscape genetic disclimax, 148 and interacting components of, 119; con-
studies of, 74, 75; land uses surrounding, disease resistance, 67, 68 servation and management of, 35, 39,
111; line and strip corridors, 86–87; dispersal mutualist, 110 40; disturbances of, 144, 153; diversity
measuring effectiveness of, 87, 88, disturbance, 35, 39, 122, 143–53; anthro- of, 16; effects of Deepwater Horizon
91–92, 94–95; naturally occurring, 87; pogenic, 47, 49, 53–59, 90, 145–47, spill on, 124; effects of habitat loss and
permeable movement barriers and, 95; 150; approaches to problem of, 151–52; degradation on, 47, 56, 58, 59; effects
in practice, 87–91 classes of, 145–46; context of, 143; of invasive species on, 103, 105–7, 110;
Cosentino, B. J., 69 definition of, 144, 145; due to beaver fragmentation of, 91; horizon scanning
cost-efficient sampling designs, 136 activity, 145; effects on birds, 145, of issues affecting, 122; modeling of,
cougars, 64 146–47, 148, 150–51; effects on physical 119; models of, 117, 118; monitoring of,
courtship displays, 21, 134 attributes of habitat, 143; fire, 144, 145, 131–32, 134, 136–38; multitrophic, 35,
cowbirds, 50, 146 146, 147; funding studies of, 152; genetic 39, 40; no-analog, 28, 39; noise effects
Cox, J., 86 monitoring of effects of, 152; nonnative on, 58; novel, 153; restoration of (see
coyotes, 71 species and, 103, 104, 106–7; spatial and wildlife habitat restoration); succession
critical habitats, 6, 36 temporal scale dependence of, 143–44, in, 148–49
crowding: density dependence and, 24, 27; 145; species relationships to spotted owl, ecotypes, 16, 27, 36, 173
due to habitat fragmentation, 86, 89, 94 151; strengths and weaknesses of meta- effective population size (Ne), 63–64, 132,
Cuban brown anoles, 71 analyses of, 147–48; succession after, 133, 152, 171; vs. census population
cues for habitat selection, 25 143–45, 148–52; targeted monitoring of size, 67; definition of, 63, 65; effects
Cushman, S. A., 130 effects of, 143, 152, 153; viewed in land- of habitat loss and fragmentation on,
cutthroat trout, 71 scape context, 150–51; wildlife studies 64, 66–68, 171; 50 / 500 rule for, 64;
Cyanistes caeruleus, 24, 54, 69 associated with, 146 genetic diversity and, 63, 69; genetic
diversity: habitat quality and, 16; haplo- drift and, 63, 66, 67; habitat area and,
dam construction, 124, 145 typic, 72; monitoring of, 132; nucleo- 66; inbreeding and, 63–64, 65, 67; as
Daubenmire, R., 3 tide, 72. See also biodiversity; genetic measure of declining populations, 133;
decision analysis, 128 diversity of Yellowstone National Park bison
decision support approaches, 40 DNA: collecting samples of, 64; environ- herd, 67
Deepwater Horizon oil spill, 121, 122, mental (eDNA), 138; mitochondrial effect size: of genetic diversity and fitness,
123–24 (mtDNA), 64, 65, 72, 75, 76; mutations 68; of monitoring efforts, 131, 135, 136;
deer mice, 72, 147, 148, 149 of, 63, 65, 72; nuclear (nDNA), 64, 65, of restoration efforts, 160; in studies of
definitions of habitat, 3–4, 9–14, 36, 102, 72, 76; techniques for analysis of, 66 disturbance, 147
144, 169–70, 172–73; classical, 10, 16, DNA sequencing, 66, 72 Ehrlich, Ann, 120
17, 170, 173; deduced meanings, 12–14; dogs, domestic: interbreeding with native Ehrlich, Paul, 120
in dictionaries and articles, 10–11; hier- species, 71; leash laws and effects in elephants, 36, 64, 71, 76
archical, 9, 16, 17, 170; irrational, 16, 17; protected areas, 55–56 Eliahu, M. N., 10
of practicing biologists, 11–12, 16 Dolichonyx oryzivorus, 25 elk, 55, 129
demographic model, 134 Donlan, C. J., 109 Ellis, E. C., 47
demographics. See population demo- Douglas fir, 36 Elton, C., 5
graphics drought, 5, 37, 39, 73, 162 Elymus athericus, 106
178 index

emigration, 19, 118, 132. See also immi- forest elephants, 76; genetic diversity Forman, R. T. T., 86
gration and, 69, 73; habitat fragmentation and, Forsman, J. T., 154
Empidonax traillii, 159 84, 89, 92, 110, 171; by hybridization, Fox, B. J., 147, 148
endangered species, 25, 50, 69, 76, 150, 70; monitoring for, 131, 132; rate of, 66 fractal dimension, 13, 92
158; hybridization among, 70, 71; legisla- Exxon Valdez oil spill, 121 Francis, C. D., 85
tion pertaining to, 35, 36, 71; monitoring Franklin, A. B., 151
of, 131, 136 Fahrig, L., 48 Fretwell, S. D., 37
Endangered Species Act, 35, 71 farm bills, 122, 123, 124 FST, 74
energy metabolism, 92 Farm Service Agency, 123
Ensetina sp., 145 Farrell, S. L., 50 Galapagos rail, 109
environmental conditions: as component fecundity, 12; impact of invasive species on, gall flies, 149
of habitat, 4, 5, 6, 10, 34, 92–93, 102; 105; as measure of habitat quality, 15, gap effects, 145
demographic effects of changes in, 20–21; per capita, 20, 26, 27; survival garden warblers, 23
27–28; genetic effects of, 74–77; habitat and, 15 Garshelis, D. L., 6, 10
quality and, 15, 16; habitat selection feedback loops, 94; for adaptive manage- geese, 35, 110
and, 19, 25, 26, 171; impact of invasive ment, 130 gene flow, 63–69; definition of, 65; effects
species on, 104–5; landscape genetic Felis concolor, 90 of habitat loss and fragmentation on,
studies of, 74–75; leading indicators Felis libyca, 71 64, 66–68, 75, 171; effects of habitat
of, 121; modeling to predict ecological Felis silvestris catus, 71 on, 73; effects on population viability,
effects of changes in, 117, 118, 120, 121; Felis silvestris sylvestris, 71 68–69; estimation of, 74; and genetic
postdisturbance, 143, 145, 147–48, 149, fenced highways and canals, 67 structure of fragmented populations, 67;
150, 151, 152; potential rate of change feral species, impact on native species, landscape genetics for estimation of, 74;
of, 121; recreation effects on, 55, 59; res- 108–9 monitoring of, 134
toration programs and, 159–60; species ferrets, domestic, 71 GENELAND program, 74
abundance and, 37, 130; species moni- 50 /500 rule, 64 gene mutations, 63, 65, 72
toring and, 137, 138; species persistence fire, 35, 105, 124, 137, 144, 145, 146, 147, gene polymorphisms, 66, 72, 75
and, 6; wildlife responses to changes in, 150; invasive plant effects and, 106–7; genetic analysis methods, 66
49–51; world changes in, 39–40 meta-analyses of effects on wildlife, 146, genetic diversity, 63; adaptive, 63–64,
environmental DNA (eDNA), 138 147–48; prescribed burns, 146–47, 161; 68, 69, 72, 74, 76–77; bottlenecks and,
environmental law, 84, 88. See also legis- spotted owl demographics after, 151 65; connectivity between patches and,
lation fire-tolerant grasses, 106–7 67, 69, 73, 75; effective population size
Environmental Protection Agency (EPA), fishing, 40, 55 and, 63, 69; effects of habitat loss and
123 fitness: correlation with heterozygosity, 68, fragmentation on, 64, 66–68, 70, 72, 75;
environmental reviews, 88–89, 91, 95–97 69; definition of, 65; genetic diversity effects on population viability, 68–69; to
environmental stochasticity, 26, 27 and, 68, 76; habitat fitness potential, evaluate habitat, 72–73; extinction risk
Epps, C. W., 67, 73, 75 6, 21; habitat selection, habitat quality, and, 69, 73; fitness and, 68, 76; habitat
equilibrium: between genetic drift and and, 5, 6, 16, 17; hybridization and, 70, quality and, 73; haplotypic, 72; hybrid-
migration, 74; between habitat selection 76; in ideal free and ideal despotic dis- ization and, 70; landscape genetics for
and habitat quality, 6, 26; Hardy- tributions, 26; inbreeding and, 67 (see estimation of, 74; metrics of, 72–73;
Weinberg, 72; view of nature, 39, 144 also inbreeding depression); individual MHC diversity and, 69; neutral, 68–69.
Eragrostis lehmanniana, 106 vs. population-level measures of, 20–21; See also population genetics
Eschrichtius robustus, 24 niche and, 5–6; surrogate measures of, genetic drift, 63–68, 74, 171; definition
Eucalyptus, 22 14–15 of, 63, 65; effective population size and,
Euphorbia esula, 149 fitness advantage, 67 63, 66, 67; and genetic structure of
European blackbirds, 54 fitness density, 6, 14–15, 16, 20 fragmented populations, 67; population
European polecats, 71 fitness potential, 6, 21 viability and, 68, 69
European starlings, 108 floods, 39, 105, 124, 146 genetic impoverishment, 104
European wild cats, 71 Florida mottled ducks, 71 genetic monitoring, 132, 134, 138, 152
Everglades restoration, 157 Florida panthers, 69, 72 genetic rescue, 65, 69
evolutionary traps, 53 Florida sand skinks, 149 genetic structure: definition of, 65; effects
existing conditions, 13; at proposed restora- Florida scrub habitat, 149 of habitat on, 73; impact of habitat
tion sites, 159–60 flushing responses, 55, 56 loss and fragmentation on, 66–67, 72;
exotic species, 28, 30, 106; disturbance, flying foxes, 22 metrics of, 73, 74
succession, and, 143, 145, 149, 151, 152, focal species for monitoring, 137, 138 genetic terms, 65
153; introduction of, 149, 151; invasion Fontaine, J. B., 146, 147 genotype, 65, 66, 68, 72, 134
of, 39, 106, 151. See also invasive species Food Security Act, 122 geographic closure, 134
expected heterozygosity (He), 72, 73 food webs, 39, 106, 107, 121 geographic range, 92; anthropogenic
extinction: assessing risk for, 65, 69, 70; forest elephants, 71, 76 changes to, 157; monitoring of, 132, 133
community assembly theory and, 145; Forest Inventory and Analysis program, giant pandas, 36
due to invasive species, 107–8, 110; of 138 Gilbert-Norton, L., 86, 91, 92
 index 179

Gleason, H. A., 144, 149 responses to, 47, 49–51; species conser- habitat preference, 5, 6, 9, 25, 38
Gleasonian concept of succession, 144, 149 vation and, 35 habitat quality, 4, 5, 9, 10, 170–72; abun-
Global Restoration Network, 164 habitat deterioration, 9, 13 dance and, 5, 49–50; animal fitness and,
Godron, M., 86 habitat elements, 9, 10, 12, 13, 16 5, 6, 14–15, 16, 17; continuum of, 49;
golden-cheeked warblers, 50 Habitat for Humanity, 3 definitions of, 5, 14–16, 17; definitions
grasshopper sparrows, 37, 120 habitat fragmentation, 9, 14, 34, 48, of: Van Horne’s mathematical defini-
gray whales, 24 84–95; anthropogenic, 87, 91; corridors tion, 5, 9, 10, 14–15, 20, 37, 170, 172;
grazing, 38, 39, 47, 49, 148; by feral goats to mitigate effects of, 84–95; crowd- determination of, 49, 170; evaluation of,
on Galapagos Islands, 109; by geese, ing due to, 86, 89, 94; definitions of, 11; fecundity as measure of, 15, 20–21;
35, 110 84–86, 95, 171; dependent variables fitness density and, 14–15; genetic
great bustards, 73 in measuring responses to, 91, 92, 95; diversity and, 73; habitat occupancy
green frogs, 53 descriptions in environmental reviews, and, 37, 49, 50, 170; habitat selection
grey foxes, 90 89, 96–97; effects on genetic structure, and, 6; individual vs. population-level
grey wolves, 71 66–67; extinction related to, 84, 89, 92, measures of, 19, 20–21, 28, 171; noise
Grinnell, J., 5 110, 171; vs. habitat loss, 48–49, 67, 85; effects on, 51; occupancy and, 37, 49, 50,
grizzly bears, 152 habitat restoration and, 171; impact on 170; population demographics and, 5, 6,
ground squirrels, 37 gene flow, effective population size, and 15, 19, 170–71; relative, 14; spatial and
GST, 74 diversity, 66–68, 70, 72, 75, 171; impact temporal components of, 10, 13, 171, 172
G'ST, 74 on hybridization and introgression, habitat relationship, 6, 7, 10, 13; effects of
Gulo gulo, 23, 145 70–72; invasive species and, 110–11, succession on, 149; variation in space
Guthery, F. S., 17 171; island biogeography theory and, and time, 37, 38. See also wildlife-habitat
Gymnogyps californianus, 36 66, 85, 91; measurement of, 91–95; relationships
permeable movement barriers and, 95; habitat restoration. See wildlife habitat
habitat(s): abiotic components of, 10, population persistence and, 85, 91–92, restoration
12, 16, 27, 118, 163, 170; as arbitrary 93, 95, 171; in practice, 87–89, 91; habitat sampling and analysis paradigm, 7
area of interest, 13, 15, 16, 170; biotic synthesis of corridors and, 87; variations habitat selection, 4–6, 10, 13; behavioral
components of, 4, 10, 12, 16, 27, 104, by location, 94–95 processes guiding, 19–20, 24–27, 28,
118, 170; common messages and themes habitat generalists, 121 171–72; conspecific attraction for,
related to, 169–73; conservation and habitat islands, 47, 89, 91 19–20, 25, 50, 171; evolutionary, 4, 6;
management of, 7, 34–41; critical, 6, 36; habitat linkages, 41, 88, 89, 90 hierarchical process of, 4, 172; ideal
definitions of, 3–4, 9–10, 14, 36, 102, habitat loss, 14, 34, 47–48; community- free and ideal despotic distributions, 26,
144, 169–70, 172–73; demographic con- level interactions and indirect effects 27, 37; problems with measurement of,
sequences of, 4, 5, 6, 19–28; disturbance of, 56–58; due to anthropogenic noise, 93–94; random settlement, 26; settle-
of, 143–53; effects of changing world on, light, and recreation, 51–56; vs. habitat ment decisions and, 24, 50, 57–58; site
39–40; fitness potential of, 6, 21; hierar- fragmentation, 48–49, 67, 85; impact dependence and, 24, 26–27
chical outlook of, 9; hierarchical view of, on gene flow, effective population size, habitat specialists, 121
9, 16, 17, 170; organismal, 4; restoration and diversity, 66–68, 70, 72; impact on habitat subcomponents, 9, 13, 14–15, 17,
of (see wildlife habitat restoration); hybridization and introgression, 70–72; 170
saturated, 37; shifts in use across life invasive species and, 110–11; measuring habitat subunits, 16
cycle, 21–24; spatial patterns of, 3, 4, wildlife responses to, 49–51; species habitat suitability, 3, 4, 19, 26–27, 37, 38;
5, 6, 10, 13, 17, 19–20, 23, 25, 38–39, specificity of, 47 habitat restoration projects for improve-
143–44, 145; species specificity of, 4, 10, habitat management. See conservation and ment of, 158; models of, 16; presence of
12; as surrogate measure of abundance, management of habitats humans and, 55, 56; quality continuum
129–30, 133, 138; temporal patterns of, habitat matching, 28 and, 49
4, 11, 17, 19–20, 37–38, 39, 144 habitat matrix, 9 habitat surrogates, 14–15, 129–30, 133,
habitat availability, 5, 13, 136 habitat occupancy, 4, 10, 27, 102; density 138
habitat avoidance, 9, 10, 55, 59 and, 37, 38, 170; habitat quality and, 37, habitat terminology, 3–7, 169–70, 172–73;
habitat components, 9, 159–60, 163; abi- 49, 50, 170; human activities and, 59; ambiguity of, 3–4; definitions and
otic, 10, 12, 16, 27, 118, 163, 170; biotic, measures of, 59, 132, 133; monitoring of, meanings of habitat, 3–4, 9–14, 36, 102,
4, 10, 12, 16, 27, 104, 118, 170 36, 132, 135, 138; restoration efforts for 144, 169–70; misunderstandings and
habitat conditions, 9, 10, 13–14 promotion of, 157–58, 164 misuse of, 3–4, 6, 9, 10; modifiers of, 3,
habitat conservation plans, 90, 95. See habitat patches, 6, 9, 11, 12, 48; arbitrary 4; niche, 5–6, 10; organism-focused, 4;
also conservation and management of area consisting of, 15; corridors connect- vegetation-based, 3, 4, 6, 12, 86
habitats ing, 48, 84–95; edge-to-interior ratios habitat trajectory, postdisturbance succes-
habitat degradation, 47; community-level of, 85, 94; genetic diversity and con- sional, 143, 144, 145, 148–49, 150, 153
interactions and indirect effects of, nectivity between, 67, 69, 73, 75; habitat habitat types, 3, 4, 9, 12, 16, 86, 170
56–58; due to anthropogenic noise, fragmentation and, 48, 171; invasive habitat use, 4–5; across full life cycle,
light, and recreation, 51–56; impact on species in, 111; Simpson’s diversity of, 21–24; demographic consequences of,
gene flow, effective population size, and 13; spatial heterogeneity among, 25; as 19–28
diversity, 66–68; measuring wildlife synonymous with habitat, 15, 16 Haematopus palliatus, 55
180 index

Hafner, D. J., 73 hybrid zone, 65, 70, 76 to, 85, 110–11, 171; indirect effects of,
Halbert, N. D., 67 Hylocichla mustelina, 22 108–10; negative impact of, 102–4;
Halfwerk, W., 52 hyperpredation, 108 plants, 105–7, 109–10
Hall, L. S., 173 irrational outlook, 16, 17
hantavirus, 149 Ibanez, I., 110–11 ISI Web of Knowledge, 158
haplotype, 65, 66, 72 ideal despotic distribution, 26, 27 island biogeography theory, 66, 85, 91
haplotypic diversity (h), 72 ideal free distribution, 26, 27, 37
Hardy-Weinberg equilibrium, 72 immigration, 19; conservation corridors Janzen, D. H., 38
Harrison, R. G., 70 for, 86; genetic rescue and, 69; habitat Johnson, D. H., 16, 119
Hartway, C., 146 fragmentation and, 84; population Johnson, H. E., 69
Hawaiian ducks, 71 models and, 118, 132; into sink habitats, Johnson, M. D., 15, 21
Helmitheros vermivorum, 22 37, 73 Jones, H. P., 163
hemlock, 109 inbreeding: definition of, 65; effective Jost’s D, 74
hemlock woolly adelgid, 109 population size and, 63–64, 65, 67; juxtaposition, 16, 144, 152
hermit thrush, 147 genetic rescue to counter effects of, 69;
heterogeneity: in demographic conse- hybridization and, 65; low genetic diver- Kalies, E. L., 146
quences of habitat use, 19; disturbances sity and, 68–69; in small or fragmented Karr, J. R., 85
and, 150; habitat fragmentation, cor- populations, 67, 68 karst salamanders, 121
ridors, and, 171; hybridization and, 71; of inbreeding depression, 63, 68, 71–72, 76; Kennedy, P. L., 146, 147
patch qualities, 25; spatiotemporal, 25, across life cycle, 70; definition of, 65, 67; keystone species, 137
27–28, 39 outbreeding depression and, 68; time to Kight, C. R., 52
heterozygosity, 72; correlation with fitness, extinction and, 70 Kirtland’s warblers, 39
68, 69; expected (He), 72, 73; inbreeding income breeders, 24 Klimstra, W. D., 13–14
effects on, 67; observed (Ho), 72 index (indices), 13; of abundance, 132, 138; Kokko, H., 25
hickory, 13, 105 of adaptive genetic diversity, 68–69, 72; Krebs, C. J., 13
hierarchical outlook, 9 of fitness, 17; of gene flow, 74, 134; of krill, 121
hierarchical process of habitat selection, habitat (subcomponent) quality, 15; of Kruger National Park (South Africa), 23
4, 172 habitat suitability, 16; species accumula- Kuehn, R., 67
hierarchical view of habitat, 9, 16, 17, 170 tion, 130; use in monitoring programs,
hiking, 55, 56, 59 138 lag, 103–4
historical range of variation, 39 indirect effects: of anthropogenic noise, lag effects, 23. See also carryover effects
Holmes, R., 23 57–58; of habitat degradation and loss, lags, time, 27, 37, 38, 94, 170
home range, 137; corridors and, 87, 90; 56–58; of invasive species, 108–10 land clearing, 145
effects of disturbance on, 144, 147, 151; induction, 7, 122 land conversion, 34, 84–86, 94, 105, 145,
habitat selection and, 93, 94 inference(s): about habitat, 17, 38; about 157; of cropland to perennial cover, 123;
horizon scanning, 122 habitat fragmentation, 93; about species habitat fragmentation due to, 84–86,
Horn, H. S., 149 relationships with habitat, 36; inductive, 94; hybridization and, 71, 76
Hossack, B. R., 147 7; for modeling and prediction, 122; for Lande, R., 68
house finches, 57 monitoring, 133, 136–37; for restoration land management, 6, 40, 136, 137, 169
house sparrows, 52 programs, 162 landscape, 3; demographic effects of
housing development, 28, 47, 85, 89, 90, infrastructure development, 145 context of, 28; disturbance and suc-
96–97, 105 input / output, 118–20 cession in context of, 150–51; habitat
hummingbirds, 57, 58 insects, 54–55, 108, 109 study areas and, 38; human impact on,
hunting, 40, 55, 169 integrity: biological, 132; ecological, 47; naturally occurring corridors in, 87;
Hurricane Hugo, 146, 150 136–37 response of organisms to alterations in,
hurricanes, 39, 124, 145, 146, 150–51 interbreeding of domestic and native spe- 4; as synonymous with habitat, 13
hybridization, 63; of African forest and sa- cies, 71 landscape bottleneck, 95
vannah elephants, 76; Allee and, 71; an- International Union for Conservation of landscape ecology, 6, 77
thropogenic, 70; of California tiger sala- Nature, 71 landscape elements, 13
manders and barred tiger salamanders, interspersion, 16 landscape genetics, 66, 73–75, 76–77
76; classification of, 70; definition of, 65, introgression, 65, 70–71, 76 La Polla, V. N., 95
70; due to interbreeding of domestic and invasion meltdown, 110 Laterallus spilonotus, 109
native species, 71; fitness and, 70, 76; invasive predators, 105, 107–8 leafy spurge, 149
genetic analysis for detection of, 75–76; invasive species, 24, 35, 41, 102–11; leash laws for dogs, 55–56
impact of habitat loss and fragmentation animals, 107–9; biodiversity loss and, legacy effects, 37, 170
on, 70–72; introgression due to, 65, 110; definition of, 102; direct impacts legislation, 3, 6, 35, 71, 84, 88, 117; Cali-
70–71, 76; legal and conservation status of, 104–8; effects on birds, 105–7, 108, fornia Environmental Quality Act, 84,
of hybrids, 71–72 109–10; extinction due to, 107–8, 110; 88; Clean Water Act, 123; Endangered
hybrid swarm, 65, 76 framework of invasion process, 103; Species Act, 35, 71; farm bills, 122, 123,
hybrid vigor, 70 habitat loss and fragmentation related 124; Food Security Act, 122; Marine
 index 181

Mammal Protection Act, 35; National microhabitat, 9, 10 138, 152; of habitat occupancy, 36, 132,
Environmental Policy Act, 88; National microsatellite analysis, 66, 72, 75 135, 138; habitat surrogates used for,
Forest Management Act, 136; Omnibus Microtus pennsylvanicus, 95 129–30, 133, 138; of impacts of climate
Public Land Management Act, 40; MIGRATE program, 74 change, 128, 139; metrics and measures
Pittman-Robertson Act, 163 migration / migratory species: artificial of programs for, 131–33; of potential
Lehmann lovegrass, 106–7 lighting on corridors for, 54; distur- rate of change, 121; purpose of, 131;
Leidolf, A., 147 bances of corridors for, 88, 90; effective reasons for, 128–29, 137; of restoration
Leopold, A., 40 number of migrants × migration rate projects, 158, 161–63; selection of sam-
Lepus americanus, 137 (Nm), 74; gene flow and, 63; neotropi- pling areas for, 133–34; of species abun-
lesser prairie chickens, 21–22 cal, 10, 23; protection under California dance, 128–29, 131–33, 138; statistical
Liebig’s Law of the Minimum, 94 Environmental Quality Act, 88; seasonal power of, 135–36; surveillance, 121,
life cycles: habitat use across, 4, 19, 21–24, interactions of, 23, 27; variation in 122, 125; of susceptible resources, 121;
28; interbreeding depression across, 70 habitat use and demography across life targeted, 131, 138; targeted: of effects of
life stages: monitoring trend occupancy stages, 22 disturbance, 143, 152, 153; by US Forest
during, 135; population demography, Migratory Bird Treaty, 35 Service, 129, 136, 138; when and how
habitat, and, 19, 20, 21–23, 27, 176 migratory connectivity, 24, 27 long to collect data for, 134–35, 138
lighting. See artificial night lighting Mihoub, J., 25 MONITOR program, 135
Likens, G. E., 144 Miller, C. R., 152 moose, 23
line corridors, 86–87 Mills, L. S., 146 Morrison, M. L., 4, 5, 6, 7, 36, 38, 102, 162,
Linnen, C. R., 72 Minerals Management Service, 123 171, 173
Litt, A. R., 106–7 mitochondrial DNA (mtDNA), 64, 65, 72, mosaic(s), 1, 23, 38, 39; of disturbance
locus (loci) in genome, 64, 65, 68; neutral, 75, 76 effects, 144, 151; landscape, 23, 38;
64, 65, 68, 69, 72, 76; nonneutral or mobility of wildlife, 133. See also corridors shifting-mosaic steady-state concepts of
adaptive, 64, 65, 68, 74; polymorphic, 72 for wildlife movement succession, 39
Loe, S., 86 models and prediction, 117–25; before- moths, 54
loggerhead sea turtles, 25 after control-impact (BACI) designs, motorized recreation and vehicles, 55
logging, 145, 151 125, 162, 165; capture-recapture models, mountain biking, 55
Lonicera maackii, 105, 106 119, 132, 134; Conservation Reserve mountain lions, 69, 90
Lonicera spp., 106 Program, 118, 122–23; for control over Mount Saint Helens, 149
Loss, S. R., 107 events, 121–22; Deepwater Horizon mule deer, 21, 23, 55
low-condition habitat, 9 oil spill, 121, 122, 123–24; designing Murphy, D. D., 84
Loxodonta africana, 36, 71 a model, 118–20; known unknowns Mustela furo, 71
Loxodonta cyclotis, 71 for, 117–18; monitoring actual events Mustela putorius, 71
Lythrum salicaria, 106, 107 compared to, 120; population models, mutations, 63, 65, 72
118; predictable, partly predictable, and mutualistic interactions, 10, 55, 109–10
major aspen, 149 unpredictable events, 124–25; proactive Myodes gapperi, 147
major histocompatibility complex (MHC), responses to future issues, 122–24;
68, 69 role of modeling, 118; simulations, 70, natal imprinting, 25
malaria, avian, 110 74, 75, 119, 135; statistical models, 118; National Biological Information Infrastruc-
mallards, 16, 70–71 unknown unknowns for, 120–21. See ture (NBII), 164
managing habitats. See conservation and also monitoring National Commission on the BP Deep-
management of habitats Moir, W. H., 130 water Horizon Oil Spill and Offshore
Marine Mammal Commission, 124 molecular genetic methods, 63, 66, 72, Drilling, 123
Marine Mammal Protection Act, 35 75–76 National Contingency Plan for oil spills,
Markov chains, 149 Molothrus ater, 50, 146 123
mate choice, 27, 35, 53, 67, 71, 76 Monamy, V., 148 National Environmental Policy Act, 88
McGowan, C. P., 55 monitoring, 128–39; adaptive, 163; adap- National Forest Management Act (NFMA),
McKelvey, K. S., 133, 173 tive management framework for, 128, 136
meadow voles, 95 129, 130–31, 137, 138; articulating goal National Marine Fisheries Service, 71
Melospiza melodia, 28 of, 131, 138; assessment and, 129; of bio- National River Restoration Science Synthe-
Merenlender, A. M., 55–56 diversity, 130, 131, 136; choosing focal sis (NRRSS) database, 164, 165
mesquite, 11, 39 species for, 137, 138; context, 131, 138; natural abode, 10, 12–14, 15, 16
meta-analyses, 173; of corridor effective- creative approaches to, 137–38; defining natural disasters, 121, 146
ness, 86, 91, 92 target population for, 133; definition negative biotic interactions, 104
meta-analyses of disturbance, 152; of, 128; determining what to monitor, Neotoma fuscipes, 151
anthropogenic, 146; fire, 146, 147–48; 121, 128, 131–33; ease, objectivity, and nest density, 21
strengths and weaknesses of, 147–48; repeatability of approaches to, 121; effect nesting: birds’ responses to noise and, 50,
and succession, 151 size of, 131, 135, 136; of environmental 52, 57; effects of invasive species on,
metapopulation theory, 3 leading indicators, 121; expense of, 128, 105, 106, 107, 108; habitat needs for, 9,
mice, 53, 58, 72, 147, 148, 149 129, 132, 136, 137–38; genetic, 132, 134, 10, 11, 16, 21; of pelagic seabirds, 21; of
182 index

nesting (continued) Oulanka National Park, 56 selection, 19–20, 24–27, 171–72;

piping plover, beach stabilization plans, outbreeding depression, 65, 68, 69, 70 related to habitat used across full life
and, 158; structures for, 11; of waterfowl ovenbirds, 22, 137, 147 cycle, 21–24; related to interactions
in Conservation Reserve Program fields, overwintering, 23–24 with biotic and abiotic components of
123; water source for, 11 Ovis canadensis nelsoni, 55, 67, 75 habitat, 27; seasonal interactions and,
nest predation, 28, 50, 57, 106 Ovis canadensis sierrae, 69 23–24; spatiotemporal variation in, 20,
nest success, 17, 20–21, 25, 28, 57, 146 24, 27–28
neutral locus, 64, 65, 68, 69, 72, 76 panthers, 69, 72 population dynamics, 6, 19–20; impact
Newtonian systems, 144 parasites, 69, 85, 86, 110 of behavioral processes guiding habitat
New Zealand grey ducks, 71 parrots, 108 selection on, 24–27, 28, 171–72; impact
niche, 5–6, 10; population demographics Passerculus sandwichensis, 23, 37 of habitat use across life cycle on,
and changes in, 27, 28 Passer domesticus, 52 21–24; individual- vs. population-level
niche conservatism, 28 patches. See habitat patches measures of habitat quality and, 20–21;
Nichols, J. D., 121 Pearson, D. E., 133 population viability analysis of, 65, 70; of
noise, anthropogenic, 47, 51–53, 85; per capita contribution to population species being monitored, 133
effects on birds, 51–53, 57–58; effects growth, 15, 23 population genetics, 35, 63–77; to detect
on reproductive success, 52–53; effects per capita effects of invasive species, hybridization, 75–76; effects of gene
on seed dispersal and pollination, 58; 103–4, 111 flow and genetic diversity on population
indirect effects of, 57–58; measuring per capita fecundity, 20, 26, 27 viability, 68–69; future directions for,
and quantifying magnitude of effects of, per capita recruitment, 20, 119 76–77; genetic monitoring, 132; impacts
58–59; from recreational activities, 51, Peromyscus, 58 of habitat loss and fragmentation on
55; sources of, 51 Peromyscus maniculatus, 72, 147 gene flow, effective population size, and
nondestructive genetic sampling, 64 Peromyscus polionotus, 53 diversity, 66–68; impacts of habitat loss
nonhabitat matrix, 85, 86 pests, 85, 109 and fragmentation on hybridization and
noninvasive genetic sampling, 64 Petrochelidon pyrrhonota, 11, 56 introgression, 70–72; incorporating
nonlinear relationships, 120 phenotype, 28, 54, 72 genetics in population viability analysis,
nonnative species, 102; animals, 107–8; Phillips, B. L., 108 70; landscape genetics, 66, 73–75,
attenuation of impacts of, 104; climate philopatric preferences, 25, 38 76–77; methods and theory of, 63–66;
change and, 104; hybridization due to Phleum pratense, 10–11 metrics of genetic diversity, 72; termi-
introduction of, 70–71; invasive, 102–11 phylogeography, 64, 73, 149 nology for, 65; using genetic methods
(see also invasive species); negative im- physiognomic type, 9, 12, 14, 16 to assess and predict impacts of habitat
pact of, 102–3; plants, 105–7; threshold Picoides arcticus, 148 alterations, 72–76
and time lags for impact on ecosystem, Picoides borealis, 150 population growth rates, 15, 20, 27, 119
103–4 Pidgeon, A. M., 20 population persistence, 7, 49, 172, 173;
nonneutral or adaptive locus, 64, 65, 68, 74 pileated woodpeckers, 137 through disturbance, 144, 148; genetics
Norris, D., 24 piñon pine, 58 and, 63, 70; habitat fragmentation and,
northern bobwhites, 11 Pinus edulis, 58 85, 91–92, 93, 95, 171; habitat quality
Northern cardinals, 26 piping plovers, 158 and, 5, 6, 21, 25, 170; habitat restora-
northern spotted owls, 36, 37 Piplio erythrophthalmus, 22 tion and, 157, 160; population viability
Noss, R. F., 86, 87 Pitra, C., 73 analysis for prediction of, 70, 132; of
Notophthalmus viridescens, 22 Pittman-Robertson Act, 163 reintroduced populations, 25
nuclear DNA (nDNA), 64, 65, 72, 76 planning rule of US Forest Service, 129, population trajectories, 49, 56, 69, 129,
nucleotide diversity (p), 72 136 130; postdisturbance, 144
nutrient cycling, 103, 107 plant invasions, 105–7 population viability, 63; behavioral cues
Plestiodon reynoldsi, 149 for habitat selection and, 25; effect of
oak, 13, 17, 38, 105 Poecile rufescens, 36 corridors on, 87, 92; genetic diversity
Obama, Barack, 123 polar bears, 36 and, 68–69, 72; habitat quality and,
observed heterozygosity (Ho), 72 pollination and seed dispersal, 58, 109–10 20–21; inbreeding depression and, 63;
occupancy. See habitat occupancy polymerase chain reaction (PCR), 66 migration and, 23
Ochotona princeps, 73 polymorphisms, 66, 72, 75 population viability analysis (PVA), 65,
Odocoileus hemionus, 23, 55 population assignment tests, 74 132; incorporating genetics in, 70
O’Grady, J. J., 70 population bottlenecks, 65, 67, 95 Populus tremuloides, 149
oil spills, 123–24, 125 population closure, 134 Porlier, M., 73
Olano-Marin, J., 69 population cycles, 135 positive selection, 65, 67
Omnibus Public Land Management Act, 40 population demographics, 4, 19–28; appar- postdisturbance colonization, 148, 151
Oncorhynchus clarki, 71 ent disconnect between habitat and, 19; postdisturbance environment and trajecto-
Oncorhynchus mykiss, 71 habitat quality and, 5, 6, 15, 19, 170–71; ries, 143, 145, 147–48, 149, 150, 151, 152
Oncorhynchus spp., 36 individual vs. population-level measures postdisturbance succession. See succession,
Oncorhynchus tshawytscha, 68 of habitat quality, 20–21; influence of postdisturbance
Otis tarda, 73 behavioral processes guiding habitat postfire conditions, 148, 150, 151
 index 183

potential natural vegetation (PNV), 144 noise from, 51, 55; nonconsumptive, 55; Safriel, U. N., 10
Powell, H. D. W., 148 wildlife avoidance responses to, 55, 59 sage sparrows, 38
power to detect change, 135–36 recruitment, 172; adult survival and, 17, salamanders, 69, 76, 121, 145, 147
prairie warblers, 22 170; per capita, 20, 119; seed, 58, 160 salmon, 35, 68
precipitation, 4, 23, 37, 39, 74, 102 red-backed voles, 147 salmonids, 36
predation risk, 15, 22, 24; animal invasions red-cockaded woodpeckers, 150 Salvelinus fontinalis, 109
and, 107–8; animal responses to, 59; red foxes, 107 sampling areas, 133–34
artificial night lighting and, 53–54; red imported fire ants, 107 sampling design, 136, 162
defenses against, 108; genetic diversity red-spotted newts, 22 sampling scale, 133
and, 72–73; habitat fragmentation and, red squirrels, 28 Sanger sequencing, 66
85, 89; invasive plants and, 105, 106; red wolves, 71 Saunders, D. A., 84, 85
nest predation, 28, 50, 57, 106; noise Reed, S. E., 55–56 savannah elephants, 71, 76
levels and, 52, 57–58; population demo- Refheldt, G. E., 149 savannah sparrows, 23, 37
graphics and, 28 Regional Response Teams, 123 scale dependencies, 143–44, 145
predator archetypes, 107 reintroduced populations, 25–26, 76 scat surveys, 55
predators, 4, 10, 102; avoidance of, 5; of Rejmánek, M., 110 Schieck, J., 151
bird nests, 50, 57; of bobwhites, 17; reproductive success: artificial night light- Schmitz, O. J., 163
defenses against, 27, 107, 108; food ing and, 54; conspecific attraction and, Schwensow, N., 69
abundance and foraging behavior of, 28; 25; density dependence and, 24; ecologi- scrub-jays, 57, 58
food webs and, 121; habitat fragmenta- cal traps and, 25; genetic drift and, 63; seals, 123
tion and, 85, 86; invasive, 105, 107–8; incubation time and, 55; influence of seasonal interactions, 10, 23–24, 27, 37, 41,
noise effects and, 51, 57, 58; removal carryover effects on, 23–24; mate choice 94, 106, 124
of, 124–25, 146; responses to artificial and, 27, 35, 53, 67, 71, 76; noise effects seasonal monitoring, 134
lighting, 54–55; of rodent nests, 28; role on, 52–53, 57; population growth rates, sea turtles, 25, 53
in habitat selection, 25; vegetation and, 15, 20, 27, 119; random settlement and, seed dispersal, 58, 109–10
35, 37, 40 26; site dependence and, 24; wildlife seed recruitment, 58, 160
prediction of ecological outcomes. See responses to environmental change and, Seiurus aurocapilla, 22, 137, 147
models and prediction 49–50. See also fecundity selection: genetic drift and, 67; genetic
prescribed burns, 146–47, 161 reserve areas, 38–39, 40, 173; conservation variation and, 63; natural, 64; positive,
proactive management, 117, 122 corridors connecting, 86; Conservation 65, 67; purifying, 65
Procyon lotor, 69 Reserve Program, 118, 122–23 sensitivity analysis, 23, 133
pronghorn antelope, 23, 55 residential development, 28, 47, 85, 89, 90, sensory ecology, 50, 53
Prosopis glandulosa, 11, 39 96–97, 105 Setophaga caerulescens, 23
protected areas, 38, 41, 55–56 resiliency, 39, 41 Setophaga chyrsoparia, 50
proxy-by-proxy approach, 136 resources: abundance of, 5; assessing Setophaga discolor, 22
pseudoreplication, 87, 94 availability of, 4; habitat defined by, 3–4, Setophaga kirtlandii, 39
Pseudotsuga menziesii, 36 10; impact of invasive species on, 102; Setophaga ruticulla, 23
Puma concolor, 69 individual vs. population-level measures settlement decisions, 24, 50, 57–58. See
purifying selection, 65 of, 21; monitoring based on importance also habitat selection
purple loosestrife, 107 of, 121; susceptibility of, 121 shade-coffee farms, 23
pyroclastic explosion of Mount Saint restoration ecology, 158, 160, 161, 164. See ship rats, 107
Helens, 149 also wildlife habitat restoration short tandem repeats (STRs), 66
Python molurus bivittatus, 107 Restoration Ecology (journal), 165 Sialia sialis, 52
retrospective population studies, 152 Siderhurst, L. A., 109
Quercus sp., 13 risk assessment, 128 Sierra Nevada bighorn sheep, 69
risk-disturbance hypothesis, 59 Sillett, T., 23
raccoons, 69 Rittenhouse, C. D., 150–51, 153 silvicultural treatments, 28, 145
Ragen, Tim, 124 road ecology, 51, 53, 67, 111 Simberloff, D., 86
rainbow trout, 71 Rodewald, A. D., 109 Simien jackals, 71
rainforests, 22 roe deer, 23, 67 Simons, T. R., 55
Rana clamitans, 53 Rollins, D., 11 simple sequence repeats (SSRs), 66
random settlement, 26 Romesberg, H. C., 7 simulations, 70, 74, 75, 119, 135
Rangifer tarandus, 23 Rosa multiflora, 106 single nucleotide polymorphism (SNP)
Rattus rattus, 107 Roseberry, J. L., 13–14 analysis, 66, 72, 75
recreational activities, 38, 47, 49, 55–56, Rountree, R., 3, 4 singular events, 144
136, 169; animal stress hormone re- Rousset’s a, 73 sink habitats, 6, 23, 37, 49, 73, 92, 93, 162,
sponses to, 55; effects of dogs on wildlife Rykiel, Jr., E. J., 144 172
in protected areas for, 55–56; habitat site dependence, 24, 26–27
management and, 40; influence on evo- Saab, V. A., 148 snakes, 107, 108
lutionary trajectories of populations, 56; Sacramento–San Joaquin Delta, 34 snowmobiling, 55, 59
184 index

snowshoe hares, 137 context, 150–51; wildlife research on urban forests, 26

Society for Restoration Ecology, 160 effects of, 149–50 Urocitellus townsendii, 37
Society of Ecological Restoration, 160 Sullivan, R. M., 73 Urocyon cinereoargenteus, 90
Solenopsis invicta, 107 Suring, L. H., 118 Ursus americanus, 90
Song, S. J., 151 surrogate measures: of abundance, 129–30, Ursus maritumus, 36
song sparrows, 28 133, 138; of fitness, 14–15 usable space, 12, 13–14, 15, 16
source-sink habitats, 6, 23, 37, 172. See also surveillance monitoring, 121, 122, 125 US Coast Guard, 123
sink habitats susceptibility, 121 US Department of Agriculture, 122
SPACE program, 135 Sus scrofa, 107 US Fish and Wildlife Service, 69, 71
Spartina alterniflora, 106 Sutherland, W., 25 US Forest Service (USFS), 138, 145; plan-
spatial factors, 3, 4, 5, 6, 10, 13, 17, 19–20, Swaddle, J. P., 52 ning rule, 129, 136
23, 25, 38–39, 67, 143–44, 172 Swainson’s hawk, 89 Usher, M. B., 149
spatially explicit capture-recapture models, Sylvia borin, 23
134 Vanderwel, M. C., 146, 147
spatiotemporal variation, 172; disturbance, Tachycineta bicolor, 52, 73 van Driesche, R. G., 108
succession and, 143–44, 149, 152; Tallmon, M. K., 133 Van Horne, B., 5, 9–10, 14–15, 16, 20, 37,
habitat fragmentation and, 93, 94, 95; Tamarix ramosissima, 106 170, 172, 173
population demographics and, 20, 24, Tamiasciurus hudsonicus, 28 vegetation: postdisturbance trajectory of,
27–28 Tamias striatus, 28 148; as surrogate for abundance, 130;
species accumulation indices, 130 Taylor, L. R., 86 types of, 4, 5, 6, 9, 12, 16, 17, 36–37, 38,
species-distribution models, 37, 39 Taylor, R. A. J., 86 40, 86, 130
species of conservation concern, 36, 136 temperature, 4, 9, 10, 17, 39, 74, 75, 102, vegetation association, 3, 5, 10
species richness: anthropogenic noise and, 109, 128; soil surface, 105; water, 109 vegetation composition, 16, 39, 105, 106
51, 57; effect of habitat fragmentation temporal factors, 4, 11, 17, 19–20, 37–38, vegetation cover type, 10, 11, 12, 48, 50,
on, 86, 91; effect of habitat restoration 39, 144, 172. See also spatiotemporal 149; changes in, 48, 49, 59; impact of
on, 162, 164; after hurricanes, 150; variation invasive herbivores on, 108–9
monitoring of, 131, 132; recreational territorial animals, 53, 86, 133, 134, 135 vegetation structure, 16, 47; impact of
activities and, 56 TESS program, 74 invasive plants on, 105, 106
spotted knapweed, 149 Tetrao urogallus, 14 Vilà, M., 110–11
spotted owls, 36, 37, 38, 150, 151, 153 threshold area, 86 Villard, M.-A., 84
state-and-transition models, 39 thresholds: for adaptive management, 131, Vireo atricapilla, 25
state variables, 128–29, 130, 131, 133 138; of ecosystem effects of invasive spe- vital rates, 26, 132
stationarity, 39 cies, 103–4; of fitness effects of neutral voles, 95, 147
statistical models, 6, 118, 120 genetic diversity, 68–69; of flushing Vulpes vulpes, 107
statistical power, 135–36 responses to humans, 56; of habitat
statistical trend, 137 quality, 49; for monitoring efforts, 129; Wagner, D. L., 108
Steidl, R. J., 106–7 for postrestoration monitoring, 163; of Waits, L. P., 152
stochastic processes, 4, 19, 26, 27, 149 resiliency, 39, 41 walruses, 123
Storch, I., 14 tidal wetlands, 4, 34, 107 Washington pygmy rabbits, 72
strip corridors, 86 tiger salamanders, 69, 76 Wasserman, T. N., 75
Strix occidentalis, 150 time lags, 27, 37, 38, 94, 170 water, 4, 12; for nesting, 11; quality of, 106,
Strix occidentalis carunia, 36 Title IV of the Omnibus Public Land Man- 107; temperature of, 109
Strix varia, 37, 151 agement Act, 40 water balances, 35
STRUCTURE program, 74 toads, 108, 137 water flow, 35, 41, 105, 121
study design: before-after control-impact, Trani, M. K., 13 watersheds, 10, 12, 71
125, 162, 165; for modeling, 118–20 transferability in time and space, 145 Watling, J., 105
Sturnus vulgaris, 108 transition probabilities, 149 Weibull distribution, 150
succession, postdisturbance, 11, 22, 38, 41, transportation networks, 51, 63, 67. See also western scrub-jays, 57, 58
122, 143–45, 148–52, 173; approaches road ecology wetlands, 96, 102; effect of creek fragmen-
to study of, 152; Clemensian concept of, tree swallows, 52, 73 tation on, 89; impact of invasive species
144, 149; climate change and, 143, 145, TRENDS program, 135 on, 107; quality of, 16; restoration of,
148, 149, 152; definition of, 144, 148; triggers, 129, 131, 138, 163 164; Sacramento–San Joaquin Delta,
disclimaxes and, 148; exotic species and, trout, 6, 71 34; surveillance monitoring of, 121; tiger
143, 145, 149, 151, 152, 153; as Markov Tsuga canadensis, 109 salamander genetic diversity and, 69
process, 149; monitoring patterns of, Tsuga caroliniana, 109 whales, 24, 64
135; scale dependence of, 143; second- tsunamis, 121, 146 Wiens, J. A., 9, 10, 85, 170, 171, 172, 173
ary, 148; shifting-mosaic steady-state Turdus merula, 54 Wilcox, B. A., 84, 85
concepts of, 39; species relationships to: Tympanuchus pallidicinctus, 21–22 wild boar, 107, 109
spotted owl, 151; viewed in landscape type I errors, 135 wildlife-habitat models, 118–19, 130
 index 185

wildlife-habitat relationships, 19–20, 28, Pittman-Robertson Act and funding for, wood turtles, 55
173; modeling of, 118 163; reports and databases of projects worm-eating warblers, 22
wildlife habitat restoration, 157–65; for, 164–65; success of, 163–64; success Wright, S., 63; island model, 74
advancement of, 164–65; before-after of: metrics, 160–61, 164
control-impact designs for, 162; compo- wildlife refuges, 10, 14, 55, 150 X chromosome, 76
nents of projects for, 157; definition of, willow flycatchers, 159
157–58; desired conditions for, 159, 161, win-stay, lose-switch strategies, 25 Yahner, R. H., 84
162, 163; developing a plan for, 158–61; wolverines, 23, 145 Y chromosome, 76
goals of, 157, 161–62; “if you build it, wolves, 55, 69, 71, 138 yellow star-thistle, 149
they will come” view of, 19–20, 161, 171; woodland salamanders, 145 Yellowstone National Park, 67, 152
literature review of projects for, 158; woodpeckers, 137, 148, 150
maximizing efforts for, 165; monitor- woodrats, 151 Zamudio, K. R., 70
ing and evaluation of, 158, 161–63; wood thrushes, 22 Zwolak, R., 147