Download the full version of the ebook at ebookfinal.

com

Wildlife Habitat Conservation Concepts Challenges

and Solutions 1st Edition Michael L. Morrison

https://ebookfinal.com/download/wildlife-habitat-
conservation-concepts-challenges-and-solutions-1st-edition-
michael-l-morrison/

OR CLICK BUTTON

DOWNLOAD EBOOK

Download more ebook instantly today at https://ebookfinal.com

Instant digital products (PDF, ePub, MOBI) available
Download now and explore formats that suit you...

Wildlife Restoration Techniques for Habitat Analysis and

Animal Monitoring 1st Edition Michael L. Morrison

https://ebookfinal.com/download/wildlife-restoration-techniques-for-
habitat-analysis-and-animal-monitoring-1st-edition-michael-l-morrison/

ebookfinal.com

Eels Physiology Habitat and Conservation Physiology

Habitat and Conservation 1st Edition Nakashima Sachiko

https://ebookfinal.com/download/eels-physiology-habitat-and-
conservation-physiology-habitat-and-conservation-1st-edition-
nakashima-sachiko/
ebookfinal.com

Wildlife Habitat Management Concepts and Applications in

Forestry 2nd Edition Brenda C. Mccomb

https://ebookfinal.com/download/wildlife-habitat-management-concepts-
and-applications-in-forestry-2nd-edition-brenda-c-mccomb/

ebookfinal.com

Habitat Management for Conservation Malcolm Ausden

https://ebookfinal.com/download/habitat-management-for-conservation-
malcolm-ausden/

ebookfinal.com
Safe Passages Highways Wildlife and Habitat Connectivity
1st Edition Jon P. Beckmann

https://ebookfinal.com/download/safe-passages-highways-wildlife-and-
habitat-connectivity-1st-edition-jon-p-beckmann/

ebookfinal.com

Free Ranging Dogs and Wildlife Conservation 1st Edition

Gompper

https://ebookfinal.com/download/free-ranging-dogs-and-wildlife-
conservation-1st-edition-gompper/

ebookfinal.com

Wildlife Ecology Conservation and Management Third Edition

Fryxell

https://ebookfinal.com/download/wildlife-ecology-conservation-and-
management-third-edition-fryxell/

ebookfinal.com

Evolution and Innovation in Wildlife Conservation Parks

and Game Ranches to Transfrontier Conservation Areas 1st
Edition Sinek
https://ebookfinal.com/download/evolution-and-innovation-in-wildlife-
conservation-parks-and-game-ranches-to-transfrontier-conservation-
areas-1st-edition-sinek/
ebookfinal.com

Discipline With Dignity New Challenges New Solutions 3rd

Edition Richard L. Curwin

https://ebookfinal.com/download/discipline-with-dignity-new-
challenges-new-solutions-3rd-edition-richard-l-curwin/

ebookfinal.com
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
All rights reserved. Published 2015
Printed in the United States of America on acid-free paper
987654321

Johns Hopkins University Press

2715 North Charles Street
Baltimore, Maryland 21218-4363
www.press.jhu.edu

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

A catalog record for this book is available from the British

Library.

Special discounts are available for bulk purchases of this book.

For more information, please contact Special Sales at 410-516-
6936 or [email protected].

Johns Hopkins University Press uses environmentally

friendly book materials, including recycled text paper that
is composed of at least 30 percent post-consumer waste,
whenever possible.
 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
This page intentionally left blank
 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-
This page intentionally left blank
 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
This page intentionally left blank
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
to viability of populations and, ultimately, species per- Royal Society B: Biological Sciences 365:2245–54.
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.
In a world where land management and conserva- Dennis, R. L. H., T. G. Shreeve, and H. V. Dyck. 2003. “To-
tion are the pinnacle goals for most ecological studies, wards a Functional Resource-Based Concept for Habitat: A
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
Approaches for Classifying Benthic Habitats and Evaluat-
tives (Lindenmayer and Hunter 2010). However, this ing Habitat Quality.” Journal of Environmental Management
should not compromise the uniqueness of components 73:165–81.
within systems. Conversely, we are continuously dis- Elton, C. 1927. Animal Ecology. London: Sidgwick and Jackson.
couraged from conducting local-scale, case studies Garshelis, D. 2000. “Delusions of Habitat Evaluation: Measur-
or baseline-monitoring studies, which when planned ing Use, Selection, and Importance.” In Research Techniques
in Animal Ecology: Controversies and Consequences, edited by
appropriately can be integrated into future research
L. Boitani and T. K. Fuller, 111–64. New York: Columbia
programs (Lindenmayer and Likens 2010). Although University Press.
beyond the scope of this chapter, the foundation of any Grinnell, J. 1917. “The Niche-Relations of the California
habitat study is the initial identification of the relevant Thrasher.” Auk 34:427–33.
8 foundation

Hall, L. S., P. R. Krausman, and M. L. Morrison. 1997. “The ———. 2009. Restoring Wildlife: Ecological Concepts and Practical
Habitat Concept and a Plea for Standard Terminology.” Applications. Washington, D.C.: Island Press.
Wildlife Society Bulletin 25:173–82. ———. 2012. “The Habitat Sampling and Analysis Paradigm
Hutchinson, G. E. 1957. “Concluding Remarks.” Cold Spring Has Limited Value in Animal Conservation: A Prequel.”
Harbor Symposium on Quantitative Biology 22:415–27. Journal of Wildlife Management 76:438–50.
Hutto, R. L. 1985. “Habitat Selection by Nonbreeding, Migra- Morrison, M. L., and L. S. Hall. 2002. “Standard Terminol-
tory Land Birds.” In Habitat Selection in Birds, edited by ogy: Toward a Common Language to Advance Ecological
M. L. Cody, 455–76. San Diego: Academic Press. Understanding and Applications.” In Predicting Species Oc-
Johnson, D. H. 1980. “The Comparison of Usage and Avail- currences: Issues of Scale and Accuracy, edited by J. M. Scott
ability Measurements for Evaluating Resource Preference.” et al., 43–52. Washington, D.C.: Island Press.
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
Concepts for Conservation Biology.” Conservation Biology Can Habitat Preference Models Tell Us? TESTS Using a Vir-
24:1459–68. tual Trout Population.” Ecological Applications 13:1580–94.
Lindenmayer, D. B., and G. E. Likens. 2010. “The Science and Rickleffs, R. E., and G. L. Miller. 2000. Ecology. 4th edition.
Application of Ecological Monitoring.” Biological Conserva- New York: W. H. Freeman and Company.
tion 143:1317–28. Romesberg, H. C. 1981. “Wildlife Science: Gaining Reliable
Mannan, R. W., and R. J. Steidl. 2013. “Habitat.” In Wildlife Knowledge.” Journal of Wildlife Management 45:293–313.
Management and Conservation: Contemporary Principles Rountree, R., and K. Able. 2007. “Spatial and Temporal Habitat
and Practices, edited by P. R. Krausman and J. W. Cain, III, Use Patterns for Salt Marsh Nekton: Implications for Eco-
229–45. Baltimore: Johns Hopkins University Press. logical Functions.” Aquatic Ecology 41:25–45.
Mitchell, S. C. 2005. “How Useful is the Concept of Habitat? A Shreeve, T. G., R. L. H. Dennis, and H. Van Dyck. 2004. “Re-
Critique.” Oikos 110:634–38. sources, Habitats and Metapopulations—Whither Reality?”
Morris, D. 2003. “Toward an Ecological Synthesis: A Case for Oikos 106:404–8.
Habitat Selection.” Oecologia 136:1–13. Van Horne, B. 1983. “Density as a Misleading Indicator of Hab-
Morris, D. W. 2011. “Adaptation and Habitat Selection in the itat Quality.” Journal of Wildlife Management 47:893–901.
Eco-evolutionary Process.” Proceedings of the National Acad- Wiens, J. A. 1984. “Resource Systems, Populations, and Com-
emy of Sciences 278:2401–11. munities.” In A New Ecology: Novel Approaches to Interactive
Morrison, M. L. 2001. “A Proposed Research Emphasis to Over- Systems, edited by P. W. Price, C. N. Slobodchikoff, and
come the Limits of Wildlife-Habitat Relationship Studies.” W. S. Gaud, 397–436. New York: Wiley and Sons.
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

LITERATURE CITED Morrison, M. L., and L. S. Hall. 2002. “Standard Terminol-

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
Grange, W. B. 1949. The Way to Game Abundance. Charles Scrib- Structure, edited by S. S. Bell, E. D. McCoy, and H. R.
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.
Johnson, D. H. 1980. “The Comparison of Usage and Avail- Trani, M. K. 2002. “The Influence of Spatial Scale on Land-
ability Measurements for Evaluating Resource Preference.” scape Pattern Description and Wildlife Habitat Assess-
Ecology 61:65–71. ment.” In Predicting Species Occurrences: Issues of Accuracy
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-
Protein.” Southeastern Naturalist 7:595–606. itat Quality.” Journal of Wildlife Management 47:893–901.
Krebs, C. J. 1972. Ecology. Harper and Row, New York, New Wiens, J. A. 2002. “Predicting Species Occurrences: Progress,
York, USA. Problems, and Prospects.” In Predicting Species Occurrences:
Midgley, A. R. 1937. “Modification of Chemical Composition Issues of Accuracy and Scale, edited by J. M. Scott, P. J. He-
of Pasture Plants by Soils.” Journal of the American Society of glund, M. L. Morrison, J. B Haufler, M. G. Raphael, W. A.
Agronomy 29:498–503. Wall, and F. B. Sampson, 739–49. Island Press, Washington,
Morrison, M. L. 2012. “The Habitat Sampling and Analysis D.C., USA.
Paradigm Has Limited Value in Animal Conservation: A
Prequel.” Journal of Wildlife Management 76:1–13.
 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
Sandercock and L. Scott Mills. Biology 70:1838–54.
Calvert, A. M., S. J. Walde, and P. D. Taylor. 2009.
“Nonbreeding-Season Drivers of Population Dynamics in
LITERATURE CITED Seasonal Migrants: Conservation Parallels across Taxa.”
Anders, A., J. Faaborg, and F. Thompson. 1998. “Postfledging Avian Conservation and Ecology 4:5.
Dispersal, Habitat Use, and Home-Range Size of Juvenile Campomizzi, A. J., J. A. Butcher, S. L. Farrell, A. G. Snelgrove,
Wood Thrushes.” Auk 115:349–58. B. A. Collier, K. J. Gutzwiller, M. L. Morrison, and R. N.
Bakermans, M. H., A. C. Vitz, A. D. Rodewald, and C. G. Ren- Wilkins. 2008. “Conspecific Attraction is a Missing Com-
gifo. 2009. “Migratory Songbird Use of Shade Coffee in the ponent in Wildlife Habitat Modeling.” Journal of Wildlife
Venezuelan Andes with Implications for Conservation of Management 72:331–36.
Cerulean Warbler.” Biological Conservation 142:2476–83. Chalfoun, A. D., and T. E. Martin. 2007. “Assessments of Habi-
Bauchinger, U., T. Van’t Hof, and H. Biebach. 2009. “Food tat Preferences and Quality Depend on Spatial Scale and
Availability during Migratory Stopover Affects Testis Metrics of Fitness.” Journal of Applied Ecology 44:983–92.
Growth and Reproductive Behaviour in a Migratory Pas- Clotfelter, E. D., A. B. Pedersen, J. A. Cranford, N. Ram, E. A.
serine.” Hormones and Behavior 55:425–33. Snajdr, V. Nolan Jr., and E. D. Ketterson. 2007. “Acorn Mast
Bearhop, S., W. Fiedler, R. Furness, S. Votier, S. Waldron, J. Drives Long-Term Dynamics of Rodent and Songbird Popu-
Newton, G. Bowen, P. Berthold, and K. Farnsworth. 2005. lations.” Oecologia 154:493–503.
“Assortative Mating as a Mechanism for Rapid Evolution of Danchin, E., L. Giraldeau, T. Valone, and R. Wagner. 2004.
a Migratory Divide.” Science 310:502–4. “Public Information: From Nosy Neighbors to Cultural
Bearhop, S., G. Hilton, S. Votier, and S. Waldron. 2004. “Stable Evolution.” Science 305:487–91.
Isotope Ratios Indicate that Body Condition in Migrating Davis, J. M., and J. A. Stamps. 2004. “The Effect of Natal
Passerines is Influenced by Winter Habitat.” Proceedings of Experience on Habitat Preferences.” Trends in Ecology and
the Royal Society B-Biological Sciences 271:S215–18. Evolution 19:411–16.
Belisle, M., A. Desrochers, and M. Fortin. 2001. “Influence of Descamps, S., S. Boutin, D. Berteaux, A. G. McAdam, and
Forest Cover on the Movements of Forest Birds: A Homing J. Gaillard. 2008. “Cohort Effects in Red Squirrels: The
Experiment.” Ecology 82:1893–1904. Influence of Density, Food Abundance and Temperature
Berger, J. 2004. “The Last Mile: How to Sustain Long-Distance on Future Survival and Reproductive Success.” Journal of
Migration in Mammals.” Conservation Biology 18:320–31. Animal Ecology 77:305–14.
Bock, C., and Z. Jones. 2004. “Avian Habitat Evaluation: Should Doligez, B., C. Cadet, E. Danchin, and T. Boulinier. 2003.
Counting Birds Count?” Frontiers in Ecology and the Environ- “When to Use Public Information for Breeding Habitat
ment 2:403–10. Selection? The Role of Environmental Predictability and
Bolger, D. T., W. D. Newmark, T. A. Morrison, and D. F. Doak. Density Dependence.” Animal Behaviour 66:973–88.
2008. “The Need for Integrative Approaches to Under- Donovan, T. M., P. W. Jones, E. M. Annand, and F. R. Thomp-
stand and Conserve Migratory Ungulates.” Ecology Letters son III. 1997. “Variation in Local-Scale Edge Effects: Mecha-
11:63–77. nisms and Landscape Context.” Ecology 78:2064–75.
Bollinger, E., P. Bollinger, and T. Gavin. 1990. “Effects of Hay- Doran, P., and R. Holmes. 2005. “Habitat Occupancy Patterns
Cropping on Eastern Populations of the Bobolink.” Wildlife of a Forest Dwelling Songbird: Causes and Consequences.”
Society Bulletin 18:142–50. Canadian Journal of Zoology 83:1297–1305.
Boves, T. J. 2011. “Multiple Responses by Cerulean Warblers Drent, R., and S. Daan. 1980. “The Prudent Parent—Energetic
to Experimental Forest Disturbance in the Appalachian Adjustments in Avian Breeding.” Ardea 68:225–52.
Mountains.” PhD dissertation, University of Tennessee, Dunning, J., D. Stewart, B. Danielson, B. Noon, T. Root, R.
Knoxville, TN. Lamberson, and E. Stevens. 1995. “Spatially Explicit
Boulinier, T., and E. Danchin. 1997. “The Use of Conspecific Population-Models—Current Forms and Future Uses.”
Reproductive Success for Breeding Patch Selection in Ter- Ecological Applications 5:3–11.
restrial Migratory Species.” Evolutionary Ecology 11:505–17. Edelaar, P., A. M. Siepielski, and J. Clobert. 2008. “Match-
Broderick, A., B. Godley, and G. Hays. 2001. “Trophic Status ing Habitat Choice Causes Directed Gene Flow: A
Drives Interannual Variability in Nesting Numbers of Neglected Dimension in Evolution and Ecology.” Evolution
Marine Turtles.” Proceedings of the Royal Society B-Biological 62:2462–72.
Sciences 268:1481–87. Esler, D. 2000. “Applying Metapopulation Theory to Conserva-
Brook, B. W., and C. J. A. Bradshaw. 2006. “Strength of Evi- tion of Migratory Birds.” Conservation Biology 14:366–72.
dence for Density Dependence in Abundance Time Series Festa-Bianchet, M. 1998. “Condition-Dependent Reproductive
of 1198 Species.” Ecology 87:1445–51. Success in Bighorn Ewes.” Ecology Letters 1:91–94.
Bunnell, D. B., S. E. Thomas, and R. A. Stein. 2007. “Prey Flesch, A. D., and R. J. Steidl. 2010. “Importance of Envi-
30 foundation

ronmental and Spatial Gradients on Patterns and Con- 2006. “Assessing Habitat Quality for a Migratory Songbird
sequences of Resource Selection.” Ecological Applications Wintering in Natural and Agricultural Habitats.” Conserva-
20:1021–39. tion Biology 20:1433–44.
Fretwell, S. D., and H. L. J. Lucas. 1969. “On Territorial Behav- Kennedy, J., P. R. Witthames, R. D. M. Nash, and C. J. Fox.
ior and Other Factors Influencing Habitat Distribution in 2008. “Is Fecundity in Plaice (Pleuronectes platessa L.)
Birds. Part 1 Theoretical Development.” Acta Biotheoretica Down-Regulated in Response to Reduced Food Intake dur-
19:16–36. ing Autumn?” Journal of Fish Biology 72:78–92.
Friesen, L., P. Eagles, and R. MacKay. 1995. “Effects of Residen- Kennedy, M., and R. Gray. 1993. “Can Ecological Theory
tial Development on Forest Dwelling Neotropical Migrant Predict the Distribution of Foraging Animals—A Critical
Songbirds.” Conservation Biology 9:1408–14. Analysis of Experiments on the Ideal Free Distribution.”
Fuhlendorf, S. D., and D. M. Engle. 2001. “Restoring Hetero- Oikos 68:158–66.
geneity on Rangelands: Ecosystem Management Based on King, D. I., R. M. Degraaf, M.-L. Smith, and J. P. Buonaccorsi.
Evolutionary Grazing Patterns.” BioScience 51:625–32. 2006. “Habitat Selection and Habitat-Specific Survival of
Gates, J., and L. Gysel. 1978. “Avian Nest Dispersion and Fledg- Fledgling Ovenbirds (Seiurus aurocapilla).” Journal of Zoology
ing Success in Field-Forest Ecotones.” Ecology 59:871–83. 269:414–21.
Greenberg, R., P. Bichier, A. C. Angon, and R. Reitsma. 1997. Kokko, H., and W. Sutherland. 2001. “Ecological Traps in
“Bird Populations in Shade and Sun Coffee Plantations in Changing Environments: Ecological and Evolutionary
Central Guatemala.” Conservation Biology 11:448–59. Consequences of a Behaviourally Mediated Allee Effect.”
Griffin, P. C., and L. S. Mills. 2009. “Sinks without Borders: Evolutionary Ecology Research 3:537–51.
Snowshoe Hare Dynamics in a Complex Landscape.” Oikos Kristan, W. B. 2003. “The Role of Habitat Selection Behavior in
118:1487–98. Population Dynamics: Source-Sink Systems and Ecological
Harrison, X. A., J. D. Blount, R. Inger, D. R. Norris, and S. Bear- Traps.” Oikos 103:457–68.
hop. 2011. “Carry- Over Effects as Drivers of Fitness Differ- ———. 2007. “Expected Effects of Correlated Habitat Vari-
ences in Animals.” Journal of Animal Ecology 80:4–18. ables on Habitat Quality and Bird Distribution.” Condor
Hartley, M., and M. Hunter. 1998. “A Meta-analysis of Forest 109:505–15.
Cover, Edge Effects, and Artificial Nest Predation Rates.” Kristan, W. B., III, M. D. Johnson, and J. T. Rotenberry. 2007.
Conservation Biology 12:465–69. “Choices and Consequences of Habitat Selection for Birds.”
Hobbs, N., and T. Hanley. 1990. “Habitat Evaluation Do Use Condor 109:485–88.
Availability Data Reflect Carrying Capacity.” Journal of Wild- Law, B., and C. Dickman. 1998. “The Use of Habitat Mosaics by
life Management 54:515–22. Terrestrial Vertebrate Fauna: Implications for Conservation
Hobbs, N., and D. Swift. 1985. “Estimates of Habitat and Management.” Biodiversity and Conservation 7:323–33.
Carrying-Capacity Incorporating Explicit Nutritional Con- Leston, L. F. V., and A. D. Rodewald. 2006. “Are Urban Forests
straints.” Journal of Wildlife Management 49:814–22. Ecological Traps for Understory Birds? An Examina-
Hoekman, S. T., L. S. Mills, D. W. Howerter, J. H. Devries, tion Using Northern Cardinals.” Biological Conservation
and I. J. Ball. 2002. “Sensitivity Analyses of the Life Cycle 131:566–74.
of Midcontinent Mallards.” Journal of Wildlife Management Lloyd, J., and T. Martin. 2005. “Reproductive Success of
66:883–900. Chestnut-Collared Longspurs in Native and Exotic Grass-
Holmes, R. T. 2011. “Avian Population and Community Pro- land.” Condor 107:363–74.
cesses in Forest Ecosystems: Long-Term Research in the Magi, M., R. Mand, H. Tamm, E. Sisask, P. Kilgas, and V. Tilgar.
Hubbard Brook Experimental Forest.” Forest Ecology and 2009. “Low Reproductive Success of Great Tits in the
Management 262:20–32. Preferred Habitat: A Role of Food Availability.” Ecoscience
Holmes, R., P. Marra, and T. Sherry. 1996. “Habitat-Specific 16:145–57.
Demography of Breeding Black-Throated Blue Warblers Marra, P. P., K. A. Hobson, and R. T. Holmes. 1998. “Linking
(Dendroica caerulescens): Implications for Population Dy- Winter and Summer Events in a Migratory Bird by Using
namics.” Journal of Animal Ecology 65:183–95. Stable-Carbon Isotopes.” Science 282(5395): 1884–86.
Homyack, J. A. 2010. “Evaluating Habitat Quality of Vertebrates Mattisson, J., H. Andren, J. Persson, and P. Segerstrom. 2011.
Using Conservation Physiology Tools.” Wildlife Research “Influence of Intraguild Interactions on Resource Use by
37:332–42. Wolverines and Eurasian Lynx.” Journal of Mammalogy
Houston, A. I., P. A. Stephens, I. L. Boyd, K. C. Harding, and 92:1321–30.
J. M. McNamara. 2007. “Capital or Income Breeding? A McPeek, M., N. Rodenhouse, R. Holmes, and T. Sherry. 2001.
Theoretical Model of Female Reproductive Strategies.” “A General Model of Site-Dependent Population Regula-
Behavioral Ecology 18:241–50. tion: Population-Level Regulation without Individual-Level
Johnson, M. D. 2007. “Measuring Habitat Quality: A Review.” Interactions.” Oikos 94:417–24.
Condor 109:489–504. Mihoub, J., P. Le Gouar, and F. Sarrazin. 2009. “Breeding
Johnson, M. D., T. W. Sherry, R. T. Holmes, and P. P. Marra. Habitat Selection Behaviors in Heterogeneous Environ-
 demographic consequences of habitat 31

ments: Implications for Modeling Reintroduction.” Oikos “Blue Tits Use Fledgling Quantity and Quality as Public
118:663–74. Information in Breeding Site Choice.” Ecology 88:2373–82.
Mills, L. S. 2012. Conservation of Wildlife Populations: Demogra- Parker, G., and W. Sutherland. 1986. “Ideal Free Distribu-
phy, Genetics, and Management. 2nd edition. John Wiley & tions When Individuals Differ in Competitive Ability:
Sons, Ltd., West Sussex, UK. Phenotype-Limited Ideal Free Models.” Animal Behaviour
Mitchell, G. W., C. G. Guglielmo, N. T. Wheelwright, C. R. 34:1222–42.
Freeman- Gallant, and D. R. Norris. 2011. “Early Life Events Perryman, W., M. Donahue, P. Perkins, and S. Reilly. 2002.
Carry Over to Influence Pre-migratory Condition in a Free- “Gray Whale Calf Production 1994–2000: Are Observed
Living Songbird.” Plos One 6:e28838. Fluctuations Related to Changes in Seasonal Ice Cover?”
Morris, D. 1987. “Tests of Density-Dependent Habitat Selec- Marine Mammal Science 18:121–44.
tion in a Patchy Environment.” Ecological Monographs Persson, J. 2005. “Female Wolverine (Gulo gulo) Reproduction:
57:269–81. Reproductive Costs and Winter Food Availability.” Canadian
———. 1988. “Habitat-Dependent Population Regulation and Journal of Zoology 83:1453–59.
Community Structure.” Evolutionary Ecology 2:253–69. Petit, L., and D. Petit. 2003. “Evaluating the Importance of
———. 2003. “How Can We Apply Theories of Habitat Selec- Human-Modified Lands for Neotropical Bird Conservation.”
tion to Wildlife Conservation and Management?” Wildlife Conservation Biology 17:687–94.
Research 30:303–19. Pettorelli, N., S. Dray, J. Gaillard, D. Chessel, P. Duncan, A.
Morrison, M. L. 2001. “A Proposed Research Emphasis to Over- Illius, N. Guillon, F. Klein, and G. Van Laere. 2003. “Spatial
come the Limits of Wildlife-Habitat Relationship Studies.” Variation in Springtime Food Resources Influences the
Journal of Wildlife Management 65:613–23. Winter Body Mass of Roe Deer Fawns.” Oecologia 137:
———. 2009. Restoring Wildlife: Ecological Concepts and Practical 363–69.
Applications. Island Press, Washington, D.C. Pidgeon, A. M., V. C. Radeloff, and N. E. Mathews. 2006.
———. 2012. “The Habitat Sampling and Analysis Paradigm “Contrasting Measures of Fitness to Classify Habitat Quality
Has Limited Value in Animal Conservation: A Prequel.” for the Black-Throated Sparrow (Amphispiza bilineata).”
Journal of Wildlife Management 76:438–50. Biological Conservation 132:199–210.
Mortelliti, A., G. Amori, and L. Boitani. 2010. “The Role of Piper, W. H. 2011. “Making Habitat Selection More ‘Familiar’:
Habitat Quality in Fragmented Landscapes: A Conceptual A Review.” Behavioral Ecology and Sociobiology 65:1329–51.
Overview and Prospectus for Future Research.” Oecologia Podolsky, R., and S. Kress. 1992. “Attraction of the Endangered
163:535–47. Dark-Rumped Petrel to Recorded Vocalizations in the Gala-
Nagy, L., and R. Holmes. 2005. “To Double-Brood or Not? pagos Islands.” Condor 94:448–53.
Individual Variation in the Reproductive Effort in Black- Pulliam, H. 1988. “Sources, Sinks, and Population Regulation.”
Throated Blue Warblers (Dendroica caerulescens).” Auk American Naturalist 132:652–61.
122:902–14. ———. 2000. “On the Relationship between Niche and Distri-
Norris, D. 2005. “Carry-Over Effects and Habitat Quality in bution.” Ecology Letters 3:349–61.
Migratory Populations.” Oikos 109:178–86. Pulliam, H., and B. Danielson. 1991. “Sources, Sinks, and Hab-
Norris, D. R., and P. P. Marra. 2007. “Seasonal Interactions, itat Selection: A Landscape Perspective on Population-
Habitat Quality, and Population Dynamics in Migratory Dynamics.” American Naturalist 137:S50–66.
Birds.” Condor 109:535–47. Pysek, P., V. Jarosik, and T. Kucera. 2002. “Patterns of Invasion
Norris, D., P. Marra, T. Kyser, T. Sherry, and L. Ratcliffe. in Temperate Nature Reserves.” Biological Conservation
2004. “Tropical Winter Habitat Limits Reproductive Suc- 104:13–24.
cess on the Temperate Breeding Grounds in a Migratory Reed, J., and A. Dobson. 1993. “Behavioral Constraints and
Bird.” Proceedings of the Royal Society B-Biological Sciences Conservation Biology: Conspecific Attraction and Recruit-
271:59–64. ment.” Trends in Ecology & Evolution 8:253–56.
Norris, D. R., and C. M. Taylor. 2006. “Predicting the Conse- Robb, G. N., R. A. McDonald, D. E. Chamberlain, and S. Bear-
quences of Carry-Over Effects for Migratory Populations.” hop. 2008. “Food for Thought: Supplementary Feeding as a
Biology Letters 2:148–51. Driver of Ecological Change in Avian Populations.” Frontiers
Norton, M., S. Hannon, and F. Schmiegelow. 2000. “Fragments in Ecology and the Environment 6:476–84.
Are Not Islands: Patch vs. Landscape Perspectives on Song- Robb, G. N., R. A. McDonald, D. E. Chamberlain, S. J. Reyn-
bird Presence and Abundance in a Harvested Boreal Forest.” olds, T. J. E. Harrison, and S. Bearhop. 2008. “Winter
Ecography 23:209–23. Feeding of Birds Increases Productivity in the Subsequent
Owen-Smith, N., D. Mason, and J. Ogutu. 2005. “Correlates Breeding Season.” Biology Letters 4:220–23.
of Survival Rates for 10 African Ungulate Populations: Rodenhouse, N., T. Sherry, and R. Holmes. 1997. “Site-
Density, Rainfall and Predation.” Journal of Animal Ecology Dependent Regulation of Population Size: A New Synthe-
74:774–88. sis.” Ecology 78:2025–42.
Parejo, D., J. White, J. Clobert, A. Dreiss, and E. Danchin. 2007. Rodenhouse, N., T. Sillett, P. Doran, and R. Holmes. 2003.
32 foundation

“Multiple Density-Dependence Mechanisms Regulate The Ecology and Evolution of Migration, edited by R. Green-
a Migratory Bird Population during the Breeding Sea- berg and P. P. Marra, 426–36. John Hopkins University
son.” Proceedings of the Royal Society B-Biological Sciences Press, Baltimore, MD.
270:2105–10. Sillett, T., R. Holmes, and T. Sherry. 2000. “Impacts of a Global
Rodewald, A. D, and M. H. Bakermans. 2006. “What Is the Climate Cycle on Population Dynamics of a Migratory
Appropriate Paradigm for Riparian Forest Conservation?” Songbird.” Science 288:2040–42.
Biological Conservation 128:193–200. Sillett, T., N. Rodenhouse, and R. Holmes. 2004. “Experimen-
Rodewald, A. D., R. P. Rohr, M. A. Fortuna, and J. Bascompte. tally Reducing Neighbor Density Affects Reproduction and
2014. “Community-Level Demographic Consequences Behavior of a Migratory Songbird.” Ecology 85:2467–77.
of Anthropogenic Disturbance: An Ecological Network Skagen, S. K., and A. A. Y. Adams. 2011. “Potential Misuse
Approach.” Journal of Animal Ecology. doi: 10.1111 / 1365- of Avian Density as a Conservation Metric.” Conservation
2656.12224. Biology 25:48–55.
Rodewald, A. D., and D. P. Shustack. 2008. “Consumer Re- Sloan, S., R. Holmes, and T. Sherry. 1998. “Depredation Rates
source Matching in Urbanizing Landscapes: Are Synan- and Predators at Artificial Bird Nests in an Unfragmented
thropic Species Over-Matching?” Ecology 89:515–21. Northern Hardwoods Forest.” Journal of Wildlife Manage-
Rosenberg, K., J. Lowe, and A. Dhondt. 1999. “Effects of For- ment 62:529–39.
est Fragmentation on Breeding Tanagers: A Continental Studds, C. E., T. K. Kyser, and P. P. Marra. 2008. “Natal Disper-
Perspective.” Conservation Biology 13:568–83. sal Driven by Environmental Conditions Interacting across
Runge, J. P., M. C. Runge, and J. D. Nichols. 2006. “The Role the Annual Cycle of a Migratory Songbird.” Proceedings
of Local Populations within a Landscape Context: Defining of the National Academy of Sciences of the United States of
and Classifying Sources and Sinks.” American Naturalist America 105:2929–33.
167:925–38. Studds, C. E., and P. P. Marra. 2005. “Nonbreeding Habitat Oc-
Runge, M. C., and P. P. Marra. 2005. “Modeling Seasonal Inter- cupancy and Population Processes: An Upgrade Experiment
actions in the Population Dynamics of Migratory Birds.” In with a Migratory Bird.” Ecology 86:2380–85.
Birds of Two Worlds: The Ecology and Evolution of Migration, Sutherland, W., and K. Norris. 2002. “Behavioural Models of
edited by R. Greenberg and P. P. Marra, 375–98. John Population Growth Rates: Implications for Conservation
Hopkins University Press, Baltimore, MD. and Prediction.” Philosophical Transactions of the Royal Soci-
Saether, B., T. Ringsby, and E. Roskaft. 1996. “Life History ety of London Series B-Biological Sciences 357:1273–84.
Variation, Population Processes and Priorities in Species Tozer, D. C., D. M. Burke, E. Nol, and K. A. Elliott. 2012.
Conservation: Towards a Reunion of Research Paradigms.” “Managing Ecological Traps: Logging and Sapsucker
Oikos 77:217–26. Nest Predation by Bears.” Journal of Wildlife Management
Saether, B., J. Tufto, S. Engen, K. Jerstad, O. Rostad, and 76:887–98.
J. Skatan. 2000. “Population Dynamical Consequences of Van Horne, B. 1983. “Density as a Misleading Indicator of Hab-
Climate Change for a Small Temperate Songbird.” Science itat Quality.” Journal of Wildlife Management 47:893–901.
287:854–56. Vitz, A. C., and A. D. Rodewald. 2006. “Can Regenerating
Schlaepfer, M., M. Runge, and P. Sherman. 2002. “Ecologi- Clearcuts Benefit Mature-Forest Songbirds? An Exami-
cal and Evolutionary Traps.” Trends in Ecology & Evolution nation of Post-breeding Ecology.” Biological Conservation
17:474–80. 127:477–86.
Schmidt, K. A. 2004. “Site-fidelity in Temporally Correlated ———. 2007. “Vegetative and Fruit Resources as Determinants
Environments Enhances Population Persistence.” Ecology of Habitat Use by Mature-Forest Birds during the Post-
Letters 7:176–84. breeding Period.” Auk 124:494–507.
Schmidt, K. A., S. R. X. Dall, and J. A. van Gils. 2010. “The ———. 2011. “Influence of Condition and Habitat Use on Sur-
Ecology of Information: An Overview on the Ecologi- vival of Post-fledging Songbirds.” Condor 113:400–11.
cal Significance of Making Informed Decisions.” Oikos Wagner, R. H., and E. Danchin. 2010. “A Taxonomy of Biologi-
119:304–16. cal Information.” Oikos 119:203–9.
Sherry, T., and R. Holmes. 1996. “Winter Habitat Quality, Ward, M. P., and S. Schlossberg. 2004. “Conspecific Attraction
Population Limitation, and Conservation of Neotropical and the Conservation of Territorial Songbirds.” Conservation
Nearctic Migrant Birds.” Ecology 77:36–48. Biology 18:519–25.
Sillett, T. S., and R. T. Holmes. 2002. “Variation in Survivor- Webster, M. S., P. P. Marra, S. M. Haig, S. Bensch, and R. T.
ship of a Migratory Songbird throughout its Annual Cycle.” Holmes. 2002. “Links Between Worlds: Unraveling Migra-
Journal of Animal Ecology 71:296–308. tory Connectivity.” Trends in Ecology and Evolution 17:76–83.
———. 2005. “Long-Term Demographic Trends, Limiting Fac- Werner, G., and J. Gilliam 1984. “The Ontogenetic Niche and
tors, and the Strength of Density Dependence in a Breeding Species Interactions in Size Structured Populations.” Annual
Population of Migratory Songbird.” In Birds of Two Worlds: Review of Ecology and Systematics 15:393–425.
 demographic consequences of habitat 33

———. 1989a. The Ecology of Bird Communities. Vol I. Foun- Wilbur, H., and J. Collins. 1973. “Ecological Aspects of Amphib-
dations and Patterns. Cambridge University Press, Cam- ian Metamorphosis.” Science 182:1305–14.
bridge UK. Witherington, B. E. 1997. “The Problem of Photopollution for
———. 1989b. “Spatial Scaling in Ecology.” Functional Ecology Sea Turtles and Other Nocturnal Animals.” In Behavioral
3:385–97. Approaches to Conservation in the Wild, edited by J. R. Clem-
Wiens, J., and C. Graham. 2005. “Niche Conservatism: Inte- mons and R. Buchholz, 303–28. Cambridge University
grating Evolution, Ecology, and Conservation Biology.” An- Press, Cambridge UK.
nual Review of Ecology Evolution and Systematics 36:519–39. Zanette, L. Y., A. F. White, M. C. Allen, and M. Clinchy. 2011.
Wilbur, H. 1980. “Complex Life-Cycles.” Annual Review of Ecol- “Perceived Predation Risk Reduces the Number of Offspring
ogy and Systematics 11:67–93. Songbirds Produce Per Year.” Science 334:1398–1401.
 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-
Exploring the Variety of Random
Documents with Different Content
The Project Gutenberg eBook of The Sabbath,
the Crystal Palace, and the People
 This ebook is for the use of anyone anywhere in the United
States and most other parts of the world at no cost and with
almost no restrictions whatsoever. You may copy it, give it away
or re-use it under the terms of the Project Gutenberg License
included with this ebook or online at www.gutenberg.org. If you
are not located in the United States, you will have to check the
laws of the country where you are located before using this
eBook.

Title: The Sabbath, the Crystal Palace, and the People

Author: James Baldwin Brown

Release date: May 23, 2020 [eBook #62202]

Language: English

Credits: Transcribed from the [1853?] Arthur Hall, Virtue, & Co.
edition by David Price. using scans from the British
Library

*** START OF THE PROJECT GUTENBERG EBOOK THE SABBATH,

THE CRYSTAL PALACE, AND THE PEOPLE ***
Transcribed from the [1853?] Arthur Hall, Virtue, & Co. edition by
David Price, email [email protected] using scans from the British
Library.

THE
SABBATH, THE CRYSTAL
PALACE,
AND THE PEOPLE.

“GO YE, AND LEARN WHAT THAT MEANETH, I WILL

HAVE MERCY AND NOT SACRIFICE.”

BY
JAMES BALDWIN BROWN, A.B.,
MINISTER OF CLAYLANDS CHAPEL, LONDON.

LONDON:
PUBLISHED FOR THE AUTHOR, BY
ARTHUR HALL, VIRTUE, & CO., PATERNOSTER ROW.
MDCCCLIII.

Price Sixpence.
 THE SABBATH, THE CRYSTAL
PALACE, AND THE PEOPLE.

The relation between the Church of Christ and human society has
long been ill-defined and unsettled. The Church has to present to
society, in its struggles and sufferings, an aspect in which the
kindliness of human sympathy and interest is blent with the severity
of truth; and this is always difficult. Between worldly compliances
on the one hand, and bigoted formalities on the other, it is hard to
strike the mean. Between the two extremes the Church is prone to
alternate. This question,—“The opening of the Crystal Palace during
a portion of the Lord’s Day,” demands the statement of the feeling
and thought of the Church upon this subject at the present time.
The spirit of a party is quite as significant as its acts and
expressions, for that spirit is a living fountain, out of which other
acts and feelings will flow forth; and as the utterance of the mind of
the Church upon this great question will probably determine the
character of its relation to, and influence on society for some years
to come, we should watch most carefully, not over our words and
deeds only, but over the spirit in which we address ourselves to this
discussion. We must be prepared either to reform or re-affirm our
first principles, as to the relation between the Church and the human
world—for this is, emphatically, a question of first principles; it has
been dealt with too much in detail; we must look to the foundations
if we would settle it aright. Nor is it a matter of merely casual and
momentary importance to which party we attach ourselves, and
what cry we raise. The party will do more work on us personally
than we shall do for the party. It is possible (it has been so before,
it may be so again) that we may be taking for ourselves and for the
Church many backward steps, by joining ourselves unthinkingly to
those who, whether right or wrong, certainly are most loud and
dogmatic in their tone. It is possible, that by calmly taking our stand
on a principle which has but few supporters, we may find ourselves,
though we appear to stand alone on earth, in holy fellowship with
the clear-eyed watchers of all earth’s transactions, who bend over
heaven’s blue cope to regard us, and with the God of truth and
love. Therefore let us watch and pray while we thoughtfully
consider this question, for it is a solemn matter, and affects the weal
of the Church and the world, and our own with them, far more
deeply than at first appears.
Before entering on the argument, I may be allowed to state, in a few
words, the reasons which have induced me to place my thoughts
upon this subject before the public. At the last meeting of the
Congregational Union, I took occasion to state my objection to a
proposed petition from the meeting against the opening of the
Crystal Palace on the Lord’s day. I was, at the time, wholly
unsupported, but have since found reason to believe that there was
a large amount of hearty and intelligent sympathy with my
objections which did not express itself. I wrote to the editor of the
British Banner, stating the fact, and developing more fully my views.
Since the publication of my letter, I have been subjected, in the
pages of religious papers, to misconstructions and
misrepresentations, especially from anonymous correspondents.
This was, of course, to be expected; but hardly, perhaps, the
singular want of comprehension, both of my views and of their own,
which some of the letter-writers displayed. Having an intense dislike
to a newspaper warfare, I felt it due to myself and many of my
brethren to state calmly my views, and to advocate them to the best
of my ability. Whence this address to Christian people, which I offer
with sincerity and earnestness, trusting to their Christian charity and
candour to give a fair consideration to the principles and conclusions
which it sets forth. I have thrown my argument into the form of an
address, which, for many reasons, I prefer.
The question to be considered in the following pages is this:—“Is it
wise and right for us, as Christians, to offer any opposition to the
opening of the Crystal Palace during a portion of the Lord’s day?” I
trust that all into whose hands this may come, are of one mind as to
the value and importance of the Sabbath. But there are those who
take the negative on this question, who are as loyal to the Sabbath,
as honestly desirous to have it better and more extensively
observed, as the straitest of those who have set themselves forth as
its special defenders. In making this clear, it will be needful to
examine the views of the great parties who have expressed
themselves against interference in this matter, but on grounds with
which I cannot accord.
There are many who take no part in the protest of, at any rate, a
large portion of the religious public, simply on the ground that an
appeal to Government, on any subject involving religious principles
and considerations, is undesirable in itself, and dangerous, inasmuch
as it may be made the precedent for future interference, in some
more serious form. Gallio is their model ruler, a man “who cared for
none of these things.” They hold that the State should have no
thought and no voice on such matters. That such a movement as
that of the Crystal Palace Company may safely be left to stand or fall
by its own merits. If it be good, they are sure it will come to
something; if bad, they have a happy faith that it will come to
nought. No doubt, such theories have a strong intrenchment in the
order of a ruling Providence. Somehow, good things do live, bad
things do die, notwithstanding man’s efforts to the contrary; but still,
bad things are long a-dying, and God expects us all—rulers, too,
according to the measure of their rule—to help to end them, and get
them decently buried out of sight. Had there been a petition to
Government to open all the beer-shops in the kingdom all day and
all night on the Sabbath, and the Government said yea, should we
dare to sit calmly by, and trust that the evil would cure itself? Is it
not the business of Government to help to protect, by governmental
effort and action, the whole community from the effects of the worst
passions and most degrading views of the community? Though, no
doubt, the highest condition of a State is that in which a healthy,
moral, public feeling renders such interference needless. We have
hardly yet arrived at this Utopia of politics, and this excessive
jealousy of Government, in the present condition of England,
obstructs many useful measures. The objection to Government
expressing itself on this subject seems to be a radically unsound
one. The question must be discussed on quite other principles, if a
healthy settlement of it is to be made.
There is a second party, which refuses to join in the protest through
indifference to the Sabbath, or latitudinarian views of its nature and
claims. To them, the Sabbath appears to be simply a human
institution; a thing invented by priests for priestly purposes; an
enslavement of man’s free spirit; a formalism which mars the pure
essence of devotion, makes it a thing of times and seasons, and
desecrates every other day that it may consecrate one. “Every day
is a Sabbath,” according to this doctrine; to attach holiness to any
particular day, is to rob all the rest of the holiness which belongs to
them. To tell a man that he can worship best in sanctuaries in
Sabbath seasons, is to fetter his right and liberty of worship at all
times and places; and, in short, the Sabbath is regarded as the very
key-stone of that arch of formalism on which the Church rears the
superstructure of her power. Such is the latitudinarian view of the
Sabbath; and, of course, those who hold it, rejoice in the prospect of
the opening of the Crystal Palace on that day. In order to discern
the falseness of their views, we must glance at the true idea of the
Christian Sabbath.
Of all the popular cants of the day, perhaps that is at once the most
pretentious, and the most heartless, which asserts a necessary
antagonism between form and spirit, soul and body, the spiritual and
the material, and sets itself up as the special champion of spirit and
the spiritual, by maintaining that all forms and organizations, all
times and seasons, all modes and habitudes, are systematic
conspiracies against the liberties and rights of men. This is the
latest resurrection of the old ascetic spirit, and must end—as all
attempts to emancipate ourselves from the conditions of life and
development which God has implied in the constitution of our being
and of all things—have ended, in blank immorality and shameless
denial of all moral law. This is the great danger of our times. It is
not formalism that we have chiefly to fear. There is more peril of
our casting off all form and order, than of our being mastered and
bound by any one. The Sabbath is of God. He who causeth the
outgoing of the morning and evening to rejoice, causeth the
outgoing of the Sabbath morning to rejoice over the human world.
He who gives to the weary body the refreshment of nightly slumber,
gives to the weary mind and spirit the rest of the Sabbath day. We
may steal the hours from slumber, but the wrong will in the end
avenge itself; and it is at our peril, and to our certain detriment as
men, societies, or nations, that we steal its offices from the Sabbath.
The moon, the fairest and the benignest minister that attends our
earth, marks out our weeks for us. She chimes with notes of silvery
clearness the sevens, while the sun intones the units on the bells of
time. In full tune with living nature we keep our Sabbaths. We
enter into the universal harmony when we consecrate our seventh
day. This surface analogy rests on the deepest principles; and if, to
any, this orderly procession of the Sabbaths seem a formalism, a
yoke of bondage, then the day, the night, the periodic mealtime, the
Christmas festival, the birthday greeting, must be to them a torment
and an insult. The arrangements of all things must be, to such, a
maddening discord. Even the primitive simplicities of barbaric
existence, if they could recur to it, would not emancipate them. But
such an experience might, perhaps, convince them, that these
consecrations are the records of a vital progress—that these
seasons, cut off from the lump of untrained, untrimmed, unformed
existence, are the string courses of the masonry of that living temple
of society, which mark the lines of its emergence from the dark
ocean of primitive barbaric chaos and night.
A tendency to think lightly of what has been consecrated by the
religious feeling of the pious for long generations, is by no means a
beautiful or commendable thing. We may be sure that some deep
reason underlies the disposition of pious minds with one consent,
through successive ages, to fall into a certain form or mode of action
and thought. God has other ways of giving to the world
commandments than by speaking or writing them. It is as distinctly
a command of God to us to work by day and sleep by night, as if it
had been written on the tables of stone; and any organized attempt
to violate it exacts the penalties of a broken command. And God
has given to all men an implicit direction to observe a Sabbath, in
the instinct of its need which he has implanted in the constitution of
humanity. This is witnessed even among the most degraded
peoples, who have wandered farthest from the first condition, and
most injured the original constitution, by means of the miserable
substitutes wherewith they are fain to supply its place. But the
question might fairly be asked here—“Is this all the obligation of the
Sabbath day? Is it simply, that as sleep is good for the body every
night, so it is good for man that, once in seven days, he should rest
from his labours? If this be all, where is the religious obligation of
the Sabbath?” No, this is not all. When we speak of a man resting
from his labours, we speak of a man’s rest, and not of a brute’s. A
brute rests from toil that he may renew the tension of his muscles; a
man, that he may renew the tension of his soul. A man’s work is not
the work of his muscles only, nor is a man’s life the life of a labourer,
of a student, of a priest, but all these in unison. And when God
said, “Thou shalt rest on the seventh day from all thy work,” he said
it to man—to a being endowed with a spirit capable of “looking
before and after,” capable of looking up to Him, whose natural and
joyful employment, on his rest-days, should be to refresh his higher
powers, by special direction of them to their appropriate objects, lest
he should become embruted by his needful daily toil. We cannot
separate, even in thought, when we look at it in relation to God,
between the rest of the Sabbath and spiritual activity and devotion.
The fact, that it is a man for whom God provides the Sabbath,
seems to indicate the kind of use which is to be made of it. The
man who deliberately holds himself back from a high exercise, of
which God has made him capable, and prefers to make his Sabbath
more like a brute’s than a man’s, commits, in relation to it, that most
heavy of transgressions, “a coming short of the glory of God.” There
is a sin which no written law can touch—an inward sin, a sin of life,
which consists in living below the idea which God has implanted; a
sin which may co-exist with the most perfect legal righteousness,
and which can only be denominated “a coming short of the glory of
God.” Human rulers may separate in their legislation between the
rest and the religious purpose of the Sabbath; they may say, “The
first we will care for, it is within our province; the second we can
take no cognizance of by formal statutes;” but the Divine Ruler can
make no such separation. To rest from toil, and to restrain the
thoughts and the heart from going out through His creation and up
to Him, is to commit that sin against Him which the Apostle specifies
as the fundamental transgression—a coming short of His glory. You
see that it is a sin which may be committed in churches and chapels,
as well as in parlours, in Parks, in Crystal Palaces, and on sea
beaches. I fear, the Sabbath is not to any of us what it ought to be,
in the measure in which it ought to be—a season of inward renewing
of strength for daily labour—a height to which God affords us leisure
and strength to climb, that we may look beyond the stars which
watch our daily travail, to Him.
Thus far we have hardly glanced at the Sabbath as a positive
institution of the Lawgiver and Ruler of this world. The view of the
Sabbath which has already been presented, seems to underlie all
Divine legislation (and there is such) upon the subject. This is the
foundation on which it rests, towards the realization of this it works.
We miss much of the meaning of the old Jewish legislation, by not
going deep down beneath the positive commandment, and studying,
as we are able—nay, if friends of Christ, bound to do—the necessity
out of which the law arises, the feature of the original Divine
constitution which it is intended to illustrate and guard. This method
will bring us into true tune with the commandment, our observance
will then be spiritual, that of a friend, a child, not a slave. Every law
must have its reason in the nature of things, must be intended to
direct attention to, vindicate, or restore, some reality which is in
danger of being disregarded. It is needful to see what is behind the
law, in order to understand it truly. God saw that there was in man
a fearful tendency to deny the God that made him, by withholding
himself from the higher exercises, the higher life, of which God
made him capable—getting rid of spiritual burden and care, and
making his life as much like a brute’s as it could possibly become.
He saw, moreover, that the struggle between the higher element and
the lower, the spirit and the flesh, which had broken loose from the
spirit’s control, would be a long and sharp one, in each human soul,
and in the world at large; and, tenderly compassionating his afflicted
and distracted prodigal children, He came, even in the very hour of
their apostacy, to help the higher nature in its conflicts, and finally
give to it the victory by allying it for ever with himself. His purpose
of mercy had a methodic development—first the germ, then the
blade, then the ear, and after that the full corn in the ear. He chose
Abraham as the man through whom He would enter into relations
with his descendants, the Jewish people, constituting them the
people through whom He would enter into relations with the whole
human world. He gave to the Jews a law, the purpose of which was
to bring out the original features of the constitution of man, and his
primal relations to God and to all things. That which had been lost
in the fall was re-established by the Jewish economy, and every
ordinance of God, which man, the sinner, had trampled upon and
spurned, was brought forth again, and sealed afresh, in the sight of
all the people, and, through history, in the sight of all men, with the
seal of the Almighty and the All-wise. The Jews, in this solemn
covenant, were the representative people, through whom God was
addressing the whole human race. It was, above all things needful,
to bring out the idea that there was power to support the original
constitution of things, though man had been suffered to attempt, for
a time, at any rate, to violate it. God needed to bear most solemn
witness before men, that no violation of it could be successful; that
it was girded with the living splendour of His righteousness, and
sustained by the terrible resources of His power. This, man had
forgotten. “Am I my brother’s keeper?” said one, in wanton
contempt of the law of that relationship of brotherhood which had
been established by God. “These be thy gods,” said others, as they
danced around their golden calf, and, under the very shadow of
Sinai, made light of the living Lord. Men were growing wanton in
the unbounded license to sin, to break every Divine command
without immediate and palpable penalty. God had to tell this people,
and through them the world, with a terrible simplicity and sternness
which even they could understand, that an awful sanction attended
the original unwritten laws which He had established, and that the
penalty of a systematic breach of them must be death. Let us take
the legislation on the Sabbath as a specimen.
The law is written thus, Exodus xx. 8, 9, 10, 11:—“Remember the
Sabbath day, to keep it holy. Six days shalt thou labour, and do all
thy work; but the seventh day is the Sabbath of the Lord thy God: in
it thou shalt not do any work, thou, nor thy son, nor thy daughter,
thy man-servant, nor thy maid-servant, nor thy cattle, nor thy
stranger that is within thy gates: for in six days the Lord made
heaven and earth, the sea, and all that in them is, and rested the
seventh day: wherefore the Lord blessed the Sabbath day, and
hallowed it.” The comment on it is given in Exodus xxxi. 12, 13, 14,
15, 16, 17:—“And the Lord spake unto Moses, saying, Speak thou
also unto the children of Israel, saying, Verily, my Sabbaths ye shall
keep: for it is a sign between me and you throughout your
generations; that ye may know that I am the Lord that doth sanctify
you. Ye shall keep the Sabbath, therefore; for it is holy unto you:
every one that defileth it shall surely be put to death: for whosoever
doeth any work therein, that soul shall be cut off from among his
people. Six days may work be done; but in the seventh is the
Sabbath of rest, holy to the Lord: whosoever doeth any work in the
Sabbath day, he shall surely be put to death. Wherefore the
children of Israel shall keep the Sabbath, to observe the Sabbath
throughout their generations, for a perpetual covenant. It is a sign
between me and the children of Israel for ever: for in six days the
Lord made heaven and earth, and on the seventh day he rested, and
was refreshed.” It has been said that this is a too tremendous
sanction to a mere formal institution. We have seen that this is not
a mere formal institution, a day appointed arbitrarily by the
Lawgiver, as for wise reasons our lawgivers appoint days and
seasons upon earth. It is a part of the original constitution, a beam
through the gloom of the first sentence, “In the sweat of thy brow
shalt thou eat bread;” a station for man the wayfarer to rest and
refresh himself, as he ploughs and digs, and fights and forces his
way through the jungle of earth to God. God, in this Sabbath
institution, is bringing out a fundamental and primal law, and He
gives to it the very highest sanction, a witness to all fundamental
laws that the breach of them is death. That God did not regard it as
a mere formal institution is most evident, in that He himself protests
against the mere form of it, when it was offered to him as a sacrifice
widowed of the congenital soul. Isaiah i. 13–15. “Bring no more
vain oblations; incense is an abomination unto me; the new moons
and Sabbaths, the calling of assemblies, I cannot away with; it is
iniquity, even the solemn meeting. Your new moons and your
appointed feasts my soul hateth: they are a trouble unto me; I am
weary to bear them. And when ye spread forth your hands, I will
hide mine eyes from you: yea, when ye make many prayers, I will
not hear: your hands are full of blood.” The Sabbath of the Jews
was a perennial monitor, pointing backwards to the earliest time.
“Remember the Sabbath day,” remember the fountain of present
and familiar liberties and joys. The time came when the alphabet of
morals had been conned and thoroughly mastered, when the whole
world was to be put into possession of treasure which had been
accumulating in the hands of the Jews for ages. It is a fine
illustration of the manner in which Judaism in its living germs, which
had never been suffered to perish, expanded into Christianity, to
trace the transition from the Jewish Sabbath to the Sabbath of the
Christian church. It is more true, perhaps, to speak of growth than
transition.
It seems very strange to some that a feature of Judaism so strongly
marked, has so little formal recognition and reinforcement in the
New Testament. What is the foundation of the Christian Sabbath?
On what ground of reason or of law does it rest? Of law, none
whatever. To the Pharisees, the men who sat in Moses’ seat, Christ
appeared to be a Sabbath breaker. Whatever His estimation of it,
His conduct with regard to it was such, that it could at any rate be
said, with a colour of truth, “This man is not of God, because he
keepeth not the Sabbath day.” On every occasion on which there is
a question on the subject, Christ appears in strong antagonism to
the Churchmen of his time; and on one occasion he defends himself
by an appeal to an irregularity of David, which he justified on the
ground of necessity; a mode of defence which must have greatly
surprised them, which surprise he takes no pains to mitigate. And
yet, mark you, he never allowed that he violated the spirit of the
Sabbath institution; nay, he charged the rigid Sabbatarians with the
violation, giving them a new lesson to learn—“Go ye, and learn what
that meaneth, I will have mercy and not sacrifice.” For the rest, we
have to gather the Sabbath law out of the New Testament in the
following fashion:—
On the first day of the week, the Saviour arose from the dead, and
laid the foundation of a new creation. On the first day of the week,
Jesus appeared to his disciples, (John xx. 19 and 26.) On the first
day of the week, the Apostles were baptized with tongues of fire.
On the first day of the week, the disciples came together to break
bread at Troas, (Acts xx. 7.) On this day Paul thinks a work of
charity specially graceful and appropriate, (1 Cor. xvi. 2.) There is
little doubt, but no decisive evidence, that the assemblies of
Christians alluded to “passim” in the Epistles, are to be referred to
this day; and John, in Patmos, “was in the Spirit on the Lord’s day.”
As far as the New Testament is concerned, we believe that this is
all. Some, then, might say that it is simply by implication and
inference, and not by distinct commandment, that the Church keeps
a Sabbath. The Jewish Sabbath, with all its awful sanctions,
vanishes, and no one can quote any law of the New Testament
which sets up anything in its place. No one can show by what
authority the day was altered, nor tell how far the authority, which
changed the day, could transfer the sanctions and penalties from the
old day to the new. In mere word, this is true enough. It is untrue
enough, at heart. How does the Sabbath come down to us? As a
day set apart, through all Christian ages, by the deliberate
consecration of the whole Catholic church from apostolic times. We
must not rail too much at tradition. The Lord’s day is certainly more
a tradition than a commandment. And what, in the Christian church,
is the office of the Spirit in relation to Divine things, and, among
them, Divine laws? Is it not the office of the Spirit to lead men to
see from within the wisdom and goodness of the commandment,
and lovingly to adopt it; showing to the soul of man the wisdom of
the Divine constitution in everything, and securing, not a formal, but
a loving adhesion to it; abandoning entirely the system of formal
prohibitions and penalties, as inconsistent with the method of grace
and of love? Does the absence of a Divine testimony against theft
or adultery, in Christ’s epitome of the commandments, legalize theft
or adultery? And does the absence of a renewed formal
announcement of the original Divine provision for man—the
Sabbath-day—mean that God ceases to care for it, that He annuls
the first constitution, and leaves man to work on till death like a
brute? On the contrary, it means that the Spirit of God, and not the
law, has taken charge of the institution, and that He will make its
power felt and its worth acknowledged, wherever He bears witness
within human souls. But it is a very terrible sin to take it out of His
charge, and make it law again, by presenting it in any other way
than He presented it to the early Christian world. Has not the Lord’s
day come down to us just as we should have anticipated—through
the Spirit? Was there not a deep reason beneath the old institution,
Exodus xxxi. 17—“It is a sign between me and the children of Israel
for ever: for in six days the Lord made heaven and earth, and on the
seventh day He rested, and was refreshed.” Is there not as deep a
reason underneath the new? Is it not the Lord’s day? The Jewish
day commemorated a creation complete; the Christian Sabbath, a
creation begun. The one a last day—the finishing of a work—looking
backwards to the original condition and constitution; the other a first
day, with the week days, the ages, the eternities before it,
commemorating a new condition and constitution, or the old one
transfigured, to bear fruit through a future eternity. For myself, I
would handle as reverently the Lord’s day which the Spirit has
delivered to us through the Church, as the Jew his Sabbath,
protected by the penalties of the law. Only, as Christians, we are
bound to go further back than the Jew could go, dared go, even
beneath and behind the commandment, and enter into the counsels
of Him who gave to us the Sabbath day. We are to regard the
purpose and spirit of the Sabbath. A man who goes to church twice
and worships, and thinks that therein he has fulfilled the law of the
Sabbath, and brands him as a law-breaker ipso facto who has never
been to church at all, fearfully misunderstands the matter. The thing
is good; its goodness is now its argument; and that observance
which is not in full tune with its goodness, may be worth much
before men, but is worth nothing before God. A father who has
children at home, in early years compels them by law to join the
family circle, and observe the laws and orders of the house. When
they are full grown, he tries to make the home attractive, and leaves
it to the sense of duty, love, and pleasure, to work conformity. This
Christian view of the Sabbath is the very opposite of that which
makes every day a Sabbath, and dishonours the one. The
endeavour to realize every day a Sabbath feeling, if genuine and
constant, will add fresh value to the Sabbath of refreshment and
repose; while a disposition to slight the Lord’s day, on the pretext of
keeping all days holy, will end in exhausting life of its spiritual
element, and destroying the very soul of man’s work.
Having laid down these principles with regard to the Christian
Sabbath, in refutation of the rationalistic argument, let us proceed to
the practical application of them to the matter in hand. The case is
just this. An institution is about to be formed and an exhibition
opened, which must be regarded as the legitimate fruit of the forty
years’ peace which that stern warrior conquered for us, whom we
have just attended with befitting splendour to his burial, whose
mourners were a million and a half of men. The Crystal Palace of
last year was its direct result—the World’s Great Show—the Jubilee
of Commerce, celebrated with festal pomp in London—the heart of
the body commercial—whence also had gone forth, forty years ago,
the head and the hands which had torn the troubler of the world’s
peace from his throne, and inaugurated the new era. England
gained the victory, and at the end of a long generation, unparalleled
for activity and enterprise in the history of man, she called the world
to her capital to see and to taste the fruits. Her grey old warrior was
spared to see it. The peaceful pageant of commerce passed by him
and did him homage, and then he was taken to his rest. It is surely
remarkable, that Wellington was spared to see the crown of his
labours—the fruit of generations of peaceful activities, which will be
marked in history by his name—and then was removed before the
event was consummated which may cost the world another struggle,
and the blood of her bravest men. Such was the Crystal Palace of
1851.
The Crystal Palace of 1853 will have another and profounder
character. It is an attempt to consecrate these fruits of the world’s
strivings to the education and elevation of the great masses, whom
commerce has too long used up, and then flung aside to perish. It
is a deliberate effort to gather all the ripest fruit of the world’s most
earnest strivings, most glorious victories, and present them, in order
to minister to the development of the men, women, and children of
the present generation. It is no mere exhibition, except in the sense
in which the whole world is an exhibition. It is the grandest
conception of a hitherto mechanical and money-loving generation,
and has its root far deeper than in a desire for 5 per cent. returns.
It grows up through the force of a conviction, which is now wrought
into the mind of the community, that the intellect and wealth which
commerce has developed, owe a ministry to the people of the land;
and that while the merchant princes can pillage the Continent,
Egypt, Palestine, India, and China, of their treasures, to minister to
their own vanity, amusement, or instruction, the united strength of
the intellect and wealth of the country should build a Palace, far
transcending all private palaces, for the great people whose industry
has made our England the queen of the kingdoms of the earth. It is
emphatically a People’s Palace, and the organization of it on this
gigantic scale, by men of shrewd understanding, is certainly a sign
that the tide of public feeling has turned towards higher, more
intellectual, more elevating pursuits and recreations, than it affected
some fifteen years ago. The fact that the keenest speculators are
now ransacking our world for the treasures of art, science, and the
early history of our race, wherewith to adorn this Palace, is a proof
that the very class which has been most prone to renounce all the
higher attributes of humanity, and to make its life like that of the
brutes that perish, is beginning to resume the exercise of those
higher attributes, and to waken to a sense of what a man’s life
includes. It is possible that many may regard this view of the New
Exhibition as overstrained. Many expect that people will go to the
Crystal Palace to see the sight, and, when they have seen it, will go
to the public-houses and finish there. Such a view is both shallow
and faithless. No doubt, those who like to think so will find plenty
which will appear to sustain their views. “The people” is a vague
term. It needs patient and intelligent observation, not on the
outskirts, but in the centre of a great popular movement, to discern
its character. The experience of the last few years does not confirm
such anticipations. The working classes, who visit the Museum in
Great Russell Street, the Zoological Gardens on Monday, Hampton
Court, and Kew, add nothing to the disorder and drunkenness of the
metropolis. It is wonderfully rare, even in the more distant
exhibitions and places of intelligent enjoyment which have been
mentioned, to meet with disorder. Certainly, the drunkenness and
disorder of London have been greatly diminished by the opening of
places of resort for the working classes where mind as well as body
may be fed. Every thing in the past justifies the extension of the
experiment, and on the grandest scale. To the poor man, these
things are not so much exhibitions as they are to us who can more
frequent them. They partake of the dignity of events, stir up the
fountains of manhood, perhaps long stagnant, and make him feel
life’s meaning and life’s worth, and thus they expand his soul. Now,
it is worth while seriously to consider—Is this different from the
object which God proposed in the institution of the Sabbath day? It
stops far short of that object; but, as far as it goes, does it not travel
in the same direction, and aim at the lifting man up from the brutish
condition into which a too slavish daily toil would plunge him, to feel
how much a man is better than a brute? God, in the Sabbath
institution, seeks to make him feel that man is so much better than a
brute, that he can talk to God as a Friend, and love Him as a Father.
No human effort can teach man that;—no Sabbath observance, even
of the strictest sort, can make man observe the Lord’s day, as the
Lord counts observance; but when men, in the mass, are gone so far
from the Sabbath that they systematically prostitute and pollute it,
does God frown on human institutions, which work, however slowly
and dimly, towards the realization of the benefit? Does He not say
of all such, “Forbid them not, for he that is not against us is on our
side?” If through the unconscious influence of Christianity, which
has leavened even our speculating fraternities, “the earth is helping”
the Kingdom; if the back-water of the mill-wheel of the Church has
come round, and is adding its strength to the current; let us look on
it lovingly and hopefully, as a state of things to be fostered and led
onward to what is better, in no wise to be resisted and banned. And
if men want to see this thing which is to elevate and educate them
on the Lord’s day—the great mass of the people being notoriously
averse to the Sabbath of the Church—we should not say, roughly
and fiercely, “You shall not,” but recognize it, as far as it goes, as a
sign of progress, hoping that, by all the humanizing influences which
are brought to bear on them, we may regain a hold on them, and
lead them on to a true appreciation of the Christian idea of the
Sabbath. The gist of the matter seems to lie here:—at present, we
have but slight hold on the working classes; they care not for our
Sabbaths, and find no pleasure in our worship. Is it right, on the
ground that it would be a formal breach of that Sabbath which is
now really and flagrantly violated, to refuse our consent to a
movement which will secure for them some portion, at any rate, of
benefit on the Lord’s day?
With many, much of the argument turns on this word “formal”
breach. It is worth while to consider more fully its meaning, and to
inquire into the real notions of Sabbath observance which prevail in
the Christian church. We are all agreed, that the highest kind of
good was contemplated by God in the institution of the Sabbath; we
are not all agreed that, failing the highest good, a lesser good which
is within our reach may lawfully be secured. Many good people
reason thus:—“The Sabbath is God’s day. It is the portion of man’s
time which He has cut off and consecrated to himself; the mere time
is His, as well as the spirit of the worshipper; to recognize any
secular employment of such time, is to recognize a robbery of God.”
“Remember thou keep holy the Sabbath day.”
To this view of Sabbath observance it may be objected—1. It
recognizes a distinction in kind between the common day and the
sacred day, which has no warrant in the New Testament Scripture.
Is not “work” a command, as well as “worship?” And is not every
commandment holy, and obedience holy? Is it only on the first day
that we are to be fervent in spirit? Are we not to be serving the
Lord by diligence in business, with fervent spirit, every day in the
week, in our places of business, and the haunts of men? The idea
that the Sabbath is a kind of tribute paid to God, to allow us to go
free to work on week-days—a sort of composition—is foreign both to
the spirit and the letter of the teaching of the New Testament. The
Lord’s day is to be a day of refreshing, of renewing of soul, that
every day may be a day of Divine service, and have more, not less,
of the sacred element infused into it. The idea of a tribute of one
day out of seven, false as it is, rules very much of the feeling of the
religious public on this question, and perhaps in a measure affects
all our hearts. But a second objection, and a plainer one, to this
view of the law of Sabbath observance, is this:—there is hardly a
Christian to be found who professes practically to carry it out. How
much of our Sabbath is literally spent in devotion, how much in
cheerful converse with the family circle, how much in meals, how
much in criticism of the sermon, not always of a highly spiritual cast,
how much in passing references to the leading topics of the day?
Let us be honest and single-minded, and answer from our
consciences these questions. If we were to try our Sabbath
observance by that law which did consecrate the time in its unity
and completeness, which of us could stand? Do we regard this as a
sin? By no means; because we feel that Christian Sabbath
observance is in the spirit, and not in the letter; in the direction of
our thoughts and desires without hindrance from the work-day world
to God; in the feeling of rest—of Sabbath calm—of holy peace and
joy—which has possessed us, and made our Sabbath more a thing
to be measured by the Spirit’s instruments, and registered in the
Spirit’s record, than by the clock. And if a man has no homage to
render, no spiritual good to gain, because he desires none, what
does he gain, what does the Church gain, what does the world gain,
if he be compelled in some sort to recognize it, by being debarred
from a pursuit which would at any other time be beneficial to him;
provided always that his liberty be no detriment to those who have
better ways of spending the day? Bring him to church to hear the
Gospel! By all means. I would that all places of pleasure were shut
up on Sabbaths, because men felt that they had better work on
hand than to visit them. But if the man says, “I will not enter your
churches—I hate them;”—we ask, where is the gain to him, to any
one, in saying, “There are the ale-houses, there are the tea-gardens
—you are a sinner for going, but there they are; there is, moreover,
a train specially provided by Government for you to travel; but this
Crystal Palace, so grand and beautiful, we can and will debar you
from. Whatever good it has to offer you, you shall not get it on
Sabbath days!” It seems very foolish to confess that all attempts to
make the masses devote the hours of the Sabbath to devotion are
futile—that no serious limit can be set successfully to the efforts of
private speculators to tempt men to demoralizing Sabbath
desecration—and yet, when a scheme is set on foot which aims at
remedying in some sort the mischief, by elevating instead of
degrading men, because it is so good and so complete as to be of
national importance, its progress is barred at once.
This is the true secret of much serious opposition. The national
character of this act is felt by many to be the chief objection to it.
Now let us understand what the word “national” implies. There is,
no doubt, a very true sense in which a Government represents a
nation. But it may represent truly or falsely; if falsely, does God
regard it as the representative of the national mind and will? If the
nation is bent on not keeping the Sabbath in the highest manner,
would the dictum of the Government against all other ways of
spending it, constitute a national Sabbath observance? All that we
could gain in that case would be an appearance—an appearance
how awfully contradicted in Clerkenwell, Rag-fair, Lambeth, Chelsea,
Greenwich, and in every place of dense population or public resort.
And does God care for this appearance? Is it a cloak that hides any
thing from his eye? Think you, that it seems to Him a thing for the
sake of which it is worth while to sacrifice one instrument which may
help man out of the pit of brutal degradation which the Sabbaths of
this metropolis disclose to us, yawning in the very heart of the
wisest, the greatest, the richest, the most godly city in the world?
Many remember fondly what Sabbaths once were, and willingly shut
their eyes to the change. In spite of the decent appearance of our
streets—and God forbid that we should ever lose it!—the reality is
too decisively the other way, for us to hope that a Government Act
can give to us a character before man or before God. The national
thought and feeling utters itself every year more loudly. It will not
help us, while the fire is raging, to batten down the hatches, and
step the deck as trimly as if the cry of alarm had never been heard.
We may shut the Crystal Palace and be no nearer a national Sabbath
keeping, nay, farther from it,—as plagues pent up in the kilns of
their own corruption but taint the air more widely, and cover with
their black shadow a broader surface of the land.
Some fear that the act of the Legislature will add a sanction to
Sabbath breaking by which many will be emboldened. Alas! the
balance is not so tremulous that the weight of Government in either
scale will make it kick the beam. It is to be feared that many will
frequent the Crystal Palace on Sunday, who otherwise would be in
the House of God. This is, no doubt, a very serious matter, but a
simply preventive legislation will not remedy the moral mischief out
of which the evil springs. For such, no system of safeguards can be
successful. Men are beyond the reach of protection, who would use
the term “national sanction” as the cloak of sin.
Thus much on the opposition arising from views of the nature of
Sabbath observance and its claims. We must now pass on to notice
a class of objections founded on the nature of the Exhibition itself,
and its probable influence on the heart, mind, and manners of men.
Many expect that men will get more harm from the accessories than
good from the thing itself. Here, again, much is to be said on either
side. There will, doubtless, be beer-houses, tea-gardens, skittle-
grounds, and all the apparatus of demoralization (though the
licensing magistrates may do much to prevent it); but may we not
fairly expect that large numbers will spend, on their travelling and
admission, money which, if they stayed at home, would be drunk or
played away; that many will take their families with them, which is
always an elevating thing to the poor; that some, at any rate, who
go for pleasure, may find deeper thoughts awakened, and turn with
disgust from grosser amusements which, in other states of mind,
would delight them; and that, on the whole, an immense amount of
vice and sensuality will be spared, though, alas, there will be enough
to waken sorrow in all good men. There will just be a battle
between the interest which the Crystal Palace will awaken, and
meaner, grosser things. Will the baser triumph, when both are fairly
brought to bear on men? There is enough in the history of public
exhibitions, during the last twenty years, to rebuke our faithlessness
and teach us hope. My whole argument rests on the fact of existing
and increasing neglect of worship and church ordinances on the
Sabbath day. For how much of the existing disaffection the Church
is responsible, God only knoweth; but, certainly, obstruction and
prevention come from us with singularly bad grace. Attraction is our
one great power. What we can attract and win, we keep. What we
constrain we cannot attach, and our chains are but ropes of sand.
One fruitful source of Sabbath desecration is the unnatural condition
in which men and women are compelled to live, in the heart of our
great cities. Every thing around them blights the gentlest and most
gracious thoughts and feelings of our nature. They live from hand
to mouth—they snatch at each day’s existence—they have no rest,
no sense of possession in the present, no hope for the future. They
are out of reach of true rest on the Sabbath. It is mere mockery to
talk about it. Keep them at home, and the Sabbath cannot be a
delight, except when an enthusiastic spirit can wholly emancipate
itself from circumstances. God’s ordinance seems to them a
delusion and a snare. Picture the miserable houses, the foul air, the
dirty, damp, mouldy walls, the reeking smoke, the pestilential
exhalations from the open sewer, the cries of drunkenness, and the
curses of blasphemy, amidst which we expect half a million of men
in our metropolis to pass the Sabbath. No wonder that they fly from
it, fly to the ale-house, and drown there their disgust and despair.
They are hardly within reach of our Christian instructions,—alas, for
the seed sown in such stony ground! The circumstances of their life
expose it to fearful peril; a broad change must pass over the moral
and mental condition of these people before, as a class, they can be
expected to receive gladly the Gospel and bring forth its fruits. Let
them get out to something they will care for—something that will
teach them—God’s clear air and sunshine, the violet odour, balmy to
them as the breath of Paradise, the bird song, the breeze among the
boughs, the fresh clean meadows, the sparkling wreathing river, and
they are at once within reach of better and holier thoughts. Or if no
thoughts come to them, for impressions shape themselves into
thoughts but slowly in minds inept, yet a genial refreshing dew has
passed over their spirits—they feel that the city life, with its squalor
and misery, is man’s work not God’s,—and at last, though the
thought may be long in ripening, they may come to think that it may
be true after all, as the Bible says, that in God the poor man has a
defender and keeper, and a remedy for all the ills which sin and
selfishness have entailed upon the world.
Those who see much of this class will be deeply convinced that we
have no means of reaching them at present as a class, though the
direct and earnest attention of the Church, in all its branches, to
their condition and needs, is a most hopeful symptom; but looking at
this great class, and their relation to society—so benighted, so
withered in soul, as to resist sternly the Divinest influences—I
confess it seems to me a terrible responsibility to keep them away,
on any day, from anything which would do them even a little good.
It is but little that we can hope, and that little will be slowly
realized. With sorrow we open the way for them—sorrow, that they
will not choose a more excellent way. We believe in God’s high
purpose in the institution of the Sabbath day, and fling wide the
gates of our sanctuaries. But if men pass scornfully or scowlingly,
let us at least be thankful, tearfully thankful, if they are not passing
by our doors to dens of vice and crime. Let us not tell them, If you
do not come here, the Lord does not care where you go. Let us tell
them that He follows them everywhere with His care,—that He has
spread abroad the expanse of nature for them,—that He has given
art, science, commerce, and history to man;—it may chance that
many, hearing this, may desire to know more of Him, and learn from
himself what He means by a Sabbath day.
It may be said, and with justice, that the majority of the frequenters
of the Crystal Palace will hardly be of the class which has been
described. Not the poorest, but the class above the poorest—the
well-paid artizans, the shopmen and shopwomen, the mercantile
clerks and the kindred classes will furnish at any rate a large
proportion of the visitors to this Exhibition. It is worth one’s while to
consider thoughtfully whether we are prepared to apply the same
rigid rule, as regards the measure of time to be devoted to public
services on the Lord’s day, to the poor workman and shopman
confined to the hot dense air of the factory or shop during all the
disposable hours of the week, and living probably in a home but
sparely visited by the light and air of heaven,—between which home
and the sanctuary he ought, according to the present theory, to
divide his time on the Sabbath,—and to the rich man who, in the
afternoon or evening, can walk in his own garden, pluck the fruit of
his own vine and fig-tree, ventilate his lungs in the purest and
balmiest air, and, being refreshed in body, can go down to God’s
house in comfort and refresh his soul. We must beware, lest we
make the lot of the poor more bitter by the yoke of our law of
ordinances which are in themselves beautiful and benignant, lest he
take the name of his God in vain. Many hold forth a rest day in the
week as the remedy. This simply means, in most cases, the sacrifice
of four or five shillings a week. Six days’ labour can hardly supply
the needs of a poor man’s family, especially its higher needs. The
loss of some shillings is certainly the loss of some books, some
schooling, some innocent amusements for his children, and is, not
seldom, the loss of bread.
The last, and in the estimation of a large number of religious men,
the most serious aspect of the question which I will refer to is this:
—“The opening of the Crystal Palace,” it is said, “is but the door to
the opening of the national institutions, the theatres, and, finally, the
factories and shops on the Sabbath.”
But there is evidently a limiting principle at work, recognized by the
public, and expressed by the Government. The limitation of the
hours, the clause against the sale of spirituous liquors, and the close
of that portion of the building which would involve actual handiwork,
are recognized by the public at once as right and seemly. There is a
public feeling in England which will care for these things, which are
supposed to be imperilled; a public feeling which, during the next
years, it will be the office of the Church to nurture and unfold. If
that public feeling fail us, then, unquestionably, our condition will be
most serious; and the question must be argued, how far a
Government may maintain, in the face of public feeling, a system of
preventions and prohibitions? That it has the right, up to a certain
point, most people feel; that there is a point beyond which it is most
unsafe, all are agreed. But that point will never be reached in
England, while the Church is faithful to her country and to her Lord.
If that point ever be reached, on us will be the sin. Ministers of the
Church of Christ! there has been too long a schism in the army. The
shepherds have been battling for precedence and prerogative, and
the sheep have strayed. The call to us now is for work; work—not
by platform crudities, vanities, and falsities; not by protests,
preventions, petitions, and bills of spiritual rights;—but by earnest,
manful, godly, spiritual effort to make the Gospel known and felt as
the power of God unto salvation. The power of the Gospel is not felt
as it once was. Admirable sermons are preached, and with
admirable emphasis and theatric art, but power does not go forth
from them. Men look at us, hear us, admire us, but our prehensile
power is gone. We do not lay hold on men. We have dealt too
much in a religion of exclusions and negations. We need to take
hold on men, and say,—“We have good news—good news of God.”
The cry, “Good news! light! bread! life!” should be heard more loudly
from our pulpits. We should not lack hearers, if we could make
them feel that we had good news to tell. Were our Master among
us at this crisis, He would not work by protests and prohibitions.
Wherever the people were, there would He be, vehement against
organized hypocritic wickedness, but patient, gentle, merciful to the
souls gone astray in their darkness and misery. Oh, that our hearts
could catch that tone of touching human tenderness wherewith He
would address this weary and burdened generation, “Come unto me,
all ye that are weary and heavy laden, and I will give you rest.” But
He is with us alway, and, by ways that we little discern and
sympathize with, He may be leading this generation to himself. Let
us be humble-hearted and full of charity; let us work, work harder
and more lovingly, with more oneness of heart and voice, to make
men feel that it is God’s good news to them which we have in
charge, and then will our Sabbaths return to us, fresh and pure,
beautiful and blest, as that George Herbert wrote of.

“SUNDAY.
“O day most calm, most bright,
The fruit of this, the next world’s bud,
Th’ indorsement of supreme delight,
Writ by a friend, and with his blood;
The couch of time; care’s balm and bay:
The week were dark, but for thy light:
Thy torch doth show the way.