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Conservation Biology
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Conservation Biology
Foundations, Concepts, Applications
Conservation Biology
Foundations, Concepts, Applications
Second Edition
Fred Van Dyke
Wheaton College
Illustrators and Permissions Assistants
Michael J. Bigelow
Lauren Anderson
Editorial Assistant
Jo Ebihara
Springer Science and Business Media
ISBN 978-1-4020-6890-4 e-ISBN 978-1-4020-6891-1
Library of Congress Control Number: 2007942036
© 2008 Springer Science + Business Media B.V.
No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered
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Printed on acid-free paper
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Fred Van Dyke
Wheaton College, Illinois
USA
Cover Photograph Description
Wiwik is a citizen of Indonesia, currently completing her
BA degree in psychology. Her professors think she is an
excellent student, but Wiwik is featured on the cover of
this textbook for another reason. She works as a caregiver
to orphaned orangutans at the Nyaru Menteng Orangutan
Rehabilitation Center in Indonesia’s Kalimantan province.
The orangutan (Pongo pygmaeus) is listed by the World
Conservation Union (IUCN) as an endangered species. The
greatest threat to current populations, now confined to the
island of Borneo, is tropical deforestation. But orangutan
populations are also threatened in another way. The demand
for infant orangutans in the pet trade, and the large sums of
money that an infant orangutan can bring on the black market, motivate many people to shoot nursing mother orangutans and sell the infant. Although illegal, many consider the
potential gain worth the risk. If caught, the hunter may be
fined or sent to jail, but that solves only part of the problem.
The infant and now orphaned orangutan has no skills or
experience needed to live alone in the forest. Released back
to the wild in this state, it would perish.
Because orangutans rely so extensively on learning
rather than instinct, they spend 7–8 years with their mothers before becoming independent. To rehabilitate orphaned
orangutans, workers like Wiwik educate them in an intensive “curriculum” of foraging techniques, nest building,
arboreal locomotion, predator avoidance, and other skills
needed to survive in the wild. In our cover photo, Wiwik
is showing two 5- to 6-year-old female orangutans, Sirius
(next to Wiwik on her left) and Betty (back to the camera)
how to open a termite mound and extract the termites from
it for food.
Wiwik’s efforts are informed by careful scientific
studies of orangutan food habits, behavior, social relations, and habitat requirements, but the problem she is
attempting to solve, the problem of the conservation
of the orangutan, is not, strictly speaking, a scientific
question. It is a complex problem that requires an
understanding not only conservation science, but of
economics, law, ethics, and national and international
politics. That is why I chose to make my initial invitation to readers with this picture. It is a picture of the
complexity of conservation biology: an international,
multi-disciplinary effort in which experiment and
management must be combined, and are often blurred,
in projects fraught with uncertainty, but not bereft of
hope. As this text goes to press, organizations like
the Orangutan Conservancy, which administers Nyaru
Menteng, are working with the Indonesian government
and national and international conservation organizations to make the first successful transitions of these
orphaned orangutans into independent, free-living
individuals who can again become part of wild populations. It is an effort in which a dedicated worker like
Wiwik may have as much to contribute as the world’s
most famous conservation scientist. And, if there is to
be hope for success, it is an effort that will need both.
It is also an effort that will need a new generation of
conservation scientists, managers, policy makers, and
activists if it is to continue. I offer this book as an
invitation to the reader, whatever your background and
wherever in the world you may be, to learn about this
effort, and to join it.
v
Foreword
I am really delighted that a second expanded edition
of this textbook is being printed. The first edition has
been useful enough and personally it has been invaluable for my teaching in Latin America. This edition
has more international examples and so will appeal to
an even wider audience. This is a significant update
and revision of a teaching resource of major importance
to the teaching of conservation biology.
Conservation is becoming a more urgent discipline
day by day as the gravity of the environmental crisis
deepens. The reality of climate change is upon us and
we know that it is seriously affecting biodiversity in
many parts of the world. It is therefore essential to train
students based on the large amount of good conservation
science that is now available. The author of this text has
made a thorough study of this literature and analysed it
succinctly and accurately. This text covers both the historic and the most recent developments in conservation.
It treats biodiversity as a whole ranging from the species
and habitat diversity to DNA and genetic diversity.
The environmental crisis is so great that science alone
will not resolve it. It has become a moral and ethical and
even a religious crisis. One of the aspects I most like about
this text is that as well as treating the practical side of conservation it discusses the philosophical and ethical aspects. It
also presents a balanced view of policy and the economics
of the subject. If the political and the ethical issues are not
addressed we will not win the battle to make the necessary changes that will conserve a major proportion of the
world’s biodiversity. However, there is also much in this
text for the practical field based conservationist.
Fred Van Dyke has done an excellent job of presenting
an ever expanding field in an accessible way. The abundant
illustrations and the tables are a great help to make the data
readily understandable. I urge all students who take up this
text to consider all these aspects carefully to prepare you for
an exciting career that is vital for the future of our planet.
Ghillean Prance
Director of the Royal Botanic Gardens, Kew, 1988–1999
vii
Preface
In the United States, there is a saying among career politicians. “You’re not really an incumbent until you win your
first re-election.” So in writing textbooks, one is not at
all confident of having made much of a contribution after
only one edition. The first edition is very educational to the
author, but many of the best lessons begin to sink in only
after the text is in press, at which point they are too late to
benefit the reader.
This second edition of Conservation Biology:
Foundations, Concepts, Applications reflects some of
what I learned from the experience of writing the first edition, both what to do and what not to do, what to leave in
and what to leave out. Of course, one not only learns from
writing, but from having written. Once a book is released
into the world, it takes on a life of its own, and an author
learns as much from the responses of his readers as from
the research he invested in writing. This second edition is
not only informed by new developments in the discipline
of conservation biology, but also by the experience of
those who have taught the discipline using my text, and I
am grateful for their feedback and candor.
Some things are unchanged from the first edition, not
only in the book but in the convictions that have motivated me to repeat the effort. I believe, more strongly
than ever, that conservation biology should be taught as a
unity of thought and practice expressed through a coherent foundation of concepts, theories, facts, and values, not
as a loose assemblage of impressive disciplinary expertise. A unified textbook of conservation biology does not
attempt to present every subject that conservation biologists have studied, but instead defines the context and
relationships of controlling ideas, problems, and applications of the discipline. Critical facts and case histories are
important, but they are meaningless without context. We
do not remember facts that we memorized years, or even
days, ago simply for the purpose of passing a test. Rather,
we remember information that skilled teachers imbued
with meaning, that inspired and enlightened us, and that
led to an understanding of our own discipline, and our
place and purpose in it. I have also learned from my colleagues who specialize in the educational process itself
(my valued friends in the “Education” department), that
effective learners who display high, long-term retention
rates of what they have studied learn highly organized
information. Ineffective learners with low retention rates
learn disorganized information. Therefore, as in the first
edition, I have given considerable attention to the organization of ideas in every chapter, and among chapters, so
that the organization of ideas is itself part of the way that
students will understand the concepts presented. It is the
connections of these ideas to one another that matter as
much as the ideas themselves.
Guided by these convictions, I have organized this second
edition around the same fundamental questions that guided
the first edition, questions that I believe give meaning to
diverse research and management efforts in all aspects of
conservation biology. First, how did conservation biology
become a distinct discipline, and what keeps conservation biology from being absorbed into related disciplines?
Second, what are the fundamental intellectual, conceptual,
and practical problems that conservation biologists must
address and solve? Third, what is the role of conservation
biology in achieving “success” in conservation in ways that
affect all dimensions of the human experience?
So What’s New in the Second Edition?
Although these foundational convictions remain unchanged,
practical experience with the first edition, my own and my
colleagues who used it, has led me to change some specifics. The first edition was encumbered with a strong North
American bias, especially in its explanation of the history
of conservation and conservation law. Without presuming
to have eliminated that bias entirely, I have been intentional
in this second edition in presenting the history of conservation, and its legal empowerment and support, in the context of a global conservation community and perspective.
ix
North American examples and experiences have a place in
understanding that perspective, but they will no longer be
offered as unreflectively or as exclusively as I did in my
first effort.
Although the first edition enjoyed course adoptions in
over 150 institutions worldwide, even those professors
most enthusiastic about the text almost unanimously
expressed discontent with the fifth chapter, “The Historic
and Foundational Paradigms of Conservation Biology.”
I have come to agree with that judgment. The paradigm
chapter was repetitive, cumbersome, and not well connected to the rest of the text, so I have removed it. Some
consideration of historic conservation science paradigms
remains important, but these are now dealt with either in
the explanation of the history of conservation and conservation biology or in chapters that address more specific
topics in genetics, populations, habitats, landscape ecology, or ecosystem management. I have also eliminated
the chapter on Restoration Ecology, not because this
subject is unimportant, but because restoration manifests
itself differently in genetics than it does in landscapes,
and so restoration is dealt in these more specific contexts
in individual chapters.
As there are elements that have been removed, there
are things that have been added. The chapter on conservation genetics and the chapter on the conservation of
populations, each formerly presented as a single chapter,
have been split into two chapters for each topic. For both
genetics and populations, the initial chapter explores the
theory underpinning an understanding of their applications
in conservation, and a second chapter then gives more
detailed examination to specific applications and case histories that make use of such theory. Of course, these kinds
of boundaries are invariably leaky, and the reader will find
these distinctions expressed more in emphasis than in
absolute, uncompromising divisions. In every chapter, old
or new, I was blessed with the reviews of highly qualified
scientists on specific topics, and, with their help and guidance, I have added examples of the most recent theoretical
developments, experiments, and field studies on those topics in each chapter.
Perhaps the most important change for users of the first
edition will be the new chapter five, which takes up a topic
that was ignored in my first edition. That is the subject
of global climate change, its current and projected effects
on world biodiversity, and the conservation strategies currently being developed to mitigate those effects. Writing
an entirely new chapter is harder than intellectually refurbishing material you have written before, but there was
no question that a chapter on climate change had to be
included. Readers will judge the quality of the effort, but
the intent is simple: to make climate-mediated conservation strategies part of “ordinary science” and discussion in
conservation biology. Given the current state of things, I
could not do otherwise.
Finally, I take the same risks I took in the first edition
in speaking directly to students and prospective future
practitioners of conservation biology in the final chapter
(Chapter 14). Conservation biology is not practiced by
textbooks. It is practiced by conservation biologists. I
believe if you want someone to do something, the right
and courteous thing to do is to ask them to do it, explain
why, and show them the means and resources they will
need to succeed. In this second edition as in the first, I
have decided to make that appeal directly to my student
readers in answering the question: how does one become a
conservation biologist?
The Differences in This Book
and the Difference It Makes
Like the first edition, this second edition strives to be
genuinely interdisciplinary in its approach. Although an
understanding of biological facts and concepts is essential and given pride of place, the text takes seriously the
contributions of law, political science, economics, ethics,
sociology, and other disciplines to the modern conservation effort. Additionally, my book does not conceal issues
of ambiguity and uncertainty in conservation science, or
issues of controversy in conservation ethics and policy.
I do not believe that we should shield students from the
inherently controversial, and often contentious, nature
of the scientific effort, nor should we try to protect them
from the messy uncertainties that inevitably arise when
we attempt to translate research results into management
decisions. Many years of working with US state and
federal conservation agencies, combined with the shared
experiences of other conservation professionals from
around the world whom I respect, have convinced me
that such uncertainty is best acknowledged quickly and
forthrightly, lest it lead to recommendations that are more
precise than accurate, and produce professionals that are
more arrogant than useful. Thus, this second edition, like
the first, not only explains, but also critiques the foundational and current practices, techniques, and concepts of
conservation biology. My purpose in taking this approach
is not to create a spirit of negativism or confusion, but
to provoke current and future conservation biologists to
examine their foundational premises carefully and make
continued efforts to improve all aspects of conservation
practice.
I close this preface by noting one other thing that is
the same in the first and second editions, but something
that only the feedback from the first edition has made me
realize that I should explicitly state. Compared to other
texts in a variety of disciplines, including conservation
biology, my book will often use more direct quotes from
primary sources, take more time to explain or dissect an
individual case history or theory, or give greater attention
x Preface
to examining the sources or implications of ideas. There
is a method to this madness. With my own students, my
goal is to teach them ways to think about, understand, and,
eventually, critique even the best examples of scientific
investigation. I do this because I do not want them to be
students forever. I need help. I need their help, and I want
them, in time, to become my helpers and, eventually, my
colleagues. I cannot accomplish this transformation simply
by summarizing the final conclusions of innumerable studies and presenting that distillation as a “list” of things to
know, however efficient and time-saving such an approach
might be. I must instead attempt to get them to re-create
the intellectual process that led to the formation of the conclusion. That takes more time, and requires an emphasis
on processing information rather than simply presenting
it. For those who believe that the greater the density of
information and summary conclusion in a textbook the
better, I can only say, with respectful candor, that my book
is not the one you should use for that approach. For those
who share my commitment to helping students re-create an
intellectual process of discovery, insight, and application
that, with practice, eventually leads them to form original
intellectual creations and applications of their own, I hope
you will find this book a useful tool, and I welcome your
feedback on how to do it better. I invite all users of this
book, whether instructors, students, or inquisitive readers
who want to know more about conservation biology, to
join me in the adventure of this effort.
Preface xi
Acknowledgements
I am grateful to the reviewers who gave me the benefit of
their time, knowledge, experience and expertise to enhance
my understanding of all of the topics covered in this text,
and to point out my errors of fact and interpretation. I am
just as grateful for their insights about teaching, letting
me know, from their experience, what works in a classroom and what does not. They have saved me as much
embarrassment, and users of the book as much frustration,
as they could. Errors and misjudgments that remain are
entirely my responsibility. My illustration and permission
assistants, Michael J. Bigelow and Lauren Anderson, displayed unflagging patience and fortitude in working with
a difficult boss, and amazing creativity in finding ways to
transform some of my abstract concepts into tables, graphs
and figures that captured concepts elegantly and visually
for the students who will use this book. Their pictures
often really were worth a thousand words. At Wheaton
College, Biology Department Chair Rod Scott, Dean
of Natural and Social Sciences Dorothy Chappell, and
Wheaton College Provost Stanton Jones were exceedingly
gracious and generous in providing me with time, funds,
and, at critical moments, much needed encouragement
to stick with the work and not give up. I am particularly
grateful to Drs. Jones, Chappell and Scott for approving
and providing funds, sometimes on very short notice, for
this work from the Aldeen Grant program of Wheaton
College, and to the college alumni and donors who made
such funds possible to support this effort. My editorial
assistant Jo Ebihara checked every reference, constructed
the index, and developed the glossary with skill and speed,
two traits that can be dangerous to combine in most people, but were superbly manifested in Jo. As with my first
edition, I continued to be blessed with the world’s most
capable secretary, Teresa Cerchio Brown, who copied,
assembled, and arranged electronic and print material until
it had cluttered every part of her computer and office, but
never complained and never failed to get everything right.
Everyone associated with my new publisher, Springer
Science and Business Media, was most helpful and considerate. My editor, Catherine Cotton, worked diligently
and patiently to develop our original book proposal and
guide it through the process of corporate acceptance and
approval. Ria Kanters, Publishing Assistant in Forestry,
Ecology, and Conservation in charge of actually assembling my material into a book, displayed the same levels
of diligence and patience (perhaps even more patience) in
enduring some of my missed deadlines and late-arriving
material and assembling it into the present text. Finally
I am most grateful to my wife, Linda, who must have
thought during some parts of the writing process that her
husband had died or run off, since he was rarely seen at
home. The end of a thing is better than the beginning, and
the end of this book, for now, allows me to look forward to
beginning to re-enter something like a normal life.
Fred Van Dyke
Wheaton College
27 September 2007
xiii
About The Author
Fred Van Dyke is a professor of biology at Wheaton College
(Illinois). He has previously served on the faculties of
Northwestern College (Iowa) and the Au Sable Institute for
Environmental Studies, as a wildlife biologist for the Montana
Department of Fish, Wildlife and Parks, as a scientific and
professional consultant to the US National Park Service,
the US Forest Service, the Pew Charitable Trust, and to
various private environmental and conservation consultants.
He is the author of numerous publications on animal home
range and habitat use, management and conservation of
animal populations, management of successional processes to
conserve habitat, and conservation values and ethics.
xv
Contents
Cover Photograph Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
Foreword. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii
About The Author . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xv
Second and First Edition Reviewers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxix
1 The History and Distinctions of Conservation Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1. Perspectives and Questions for an Inquiry into Conservation Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.2. The Origins of Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.2.1. Conservation in Historical Context. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.2.2. Cultural Foundations of Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.2.3. Conservation as Expression of Privilege. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.2.4. Conservation as Right Relationship with Nature – The Arcadian Vision. . . . . . . . . . . . . . . . . . . . . . . . . 6
1.2.5. Conservation as Knowledge – The Invitation to Study and Appreciate Nature . . . . . . . . . . . . . . . . . . . . 7
1.2.6. Conservation to Save Species – Origins of the First Conservation Organizations . . . . . . . . . . . . . . . . . . 8
1.2.7. Conservation as Preservation of Landscape – The Washburn Expedition Goes to Yellowstone . . . . . . . 9
1.3. Intellectual Foundations and History of Conservation in the United States . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
1.3.1. Conservation as Moral Mission – John Muir and Theodore Roosevelt . . . . . . . . . . . . . . . . . . . . . . . . . . 11
1.3.2. “Scientifi c Conservation” Through Sustained Yield – Moral Mission Gives Way to
Utilitarian Purpose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
1.3.2.1. The Federal Government Empowers Conservation as Science and Democratic Ideal . . . . . . . . 12
1.3.2.2. German Infl uences in Conservation – Forest Monocultures and Maximum Yields. . . . . . . . . . . . . 14
1.3.2.3. The Rise of the Resource Conservation Ethic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
1.3.2.4. Aldo Leopold and the Formation of the “Wilderness Ideal” in Conservation . . . . . . . . . . . . . . 16
1.4. The Emergence of Global Conservation – Shared Interests Lead to Cooperation . . . . . . . . . . . . . . . . . . . . . . . 17
1.4.1. Multilateral Treaties – The Beginnings of International Conservation Efforts. . . . . . . . . . . . . . . . . . . . . 17
1.4.1.1. Conservation Driven by Shared Commercial Interests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
1.4.1.2. International Protection of Migratory Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
1.4.2. Forums for International Conservation – The United Nations and the
International Union for the Conservation of Nature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
1.5. Conservation in the Developing World: New Expressions of Resource Management,
National Parks and Nature Preserves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
1.6. Return to Start: What is the Place of Conservation Biology in the World
Conservation Effort? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
1.6.1. The Emergence of Conservation Biology from the Applied Sciences . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
1.6.2. Conceptually Distinctive Characteristics of Conservation Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
1.7. Synthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
xvii
xviii Contents
2 Values and Ethics in Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
2.1. What Does Science Have to Do with Value?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
2.1.1. Avoiding the Absurd – Being Self-Aware of Values in Conservation Decisions . . . . . . . . . . . . . . . . . . . . 30
2.1.2. Recognizing Management Actions as Value Judgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
2.1.3. Values and Ethics – Defi nitions and Initial Assessments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
2.2. The Problem of Categories: How Do We Classify Different Kinds
of Conservation Values?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
2.2.1. An Overview of Value Categories. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
2.2.2. Instrumental Values. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
2.2.2.1. General Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
2.2.2.2. Determining Attitudes with Sociological Surveys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
2.2.2.3. Tools of Economic Valuation: Cost–Benefi t Analysis
and Contingency Valuation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
2.2.2.4. Contingent Valuation Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
2.2.2.5. Criticisms of Contingent Valuation Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
2.3. The Problem of Moral Value: Assigning Intrinsic Values in Conservation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2.3.1. Where Does Intrinsic Value Reside?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2.3.2. Ecocentrism as a Basis for the Intrinsic Value . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
2.3.3. Intrinsic Value in the Judeo-Christian Tradition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
2.3.4. Other Western Religious Traditions – Islam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
2.3.5. Eastern Religious Traditions and Conservation – Hinduism and Buddhism . . . . . . . . . . . . . . . . . . . . . . 44
2.3.5.1. Hinduism. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
2.3.5.2. Buddhism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
2.3.6. Practical Implications – Faith-Based Organizations in Conservation. . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
2.3.6.1. “Goal Rational” Versus “Value Rational” Conservation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
2.3.6.2. Jewish and Christian FBOs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
2.3.6.3. FBOs in Islam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
2.3.6.4. Conservation Activism in Hinduism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
2.3.6.5. Conservation FBOs in Buddhism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
2.3.6.6. Future Roles and Contributions of FBOs in Global Conservation . . . . . . . . . . . . . . . . . . . . . . . 48
2.4. The Problem of Practice: Do Conservation Values Require Conservation Virtues? . . . . . . . . . . . . . . . . . . . . . . 49
2.4.1. The Problem of Plastic Trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
2.4.2. From Values to Virtues: Virtue-Based Ethics in Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
2.4.3. What are Appropriate Conservation Virtues? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
2.5. Orphaned Orangutans: Ethical Applications in Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
2.6. Synthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
3 The Legal Foundations of Conservation Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
3.1. Conservation Law and Policy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
3.1.1. Context and Defi nition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
3.1.2. Historical Origins of Conservation Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
3.2. Environmental and Conservation Law in Individual Nations: Modern Examples
from the United States, South Africa, and Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
3.2.1. General Considerations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
3.2.2. Common Characteristics of Effective National Conservation Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
3.2.3. The US National Environmental Policy Act (NEPA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
3.2.3.1. NEPA’s History and Content. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
3.2.3.2. NEPA and US Federal Lands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
3.2.3.3. Preparation of an Environmental Impact Statement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
3.2.3.4. Shortcomings of the National Environmental Policy Act. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
3.2.4. The US Endangered Species Act . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
3.2.4.1. Historical Origins and Content . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
3.2.4.2. The Endangered Species Act and Landowner Confl icts:
The Case of the Red-Cockaded Woodpecker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Contents xix
3.2.4.3. San Bruno Mountain and the Evolution of Habitat Conservation Planning . . . . . . . . . . . . . . . . 67
3.2.4.4. Criticisms of the Endangered Species Act . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
3.2.5. Water as an Inalienable Reserve – South Africa and Australia Establish Radical
Categories for Conservation Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
3.3. International Conservation Law: Concept and Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
3.3.1. General Considerations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
3.3.2. A Forum for Cooperation and Legal Foundation – The United Nations
and Its Environmental Programs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
3.3.2.1. Background and Context. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
3.3.2.2. Stockholm: The Beginnings of Modern International Conservation Law . . . . . . . . . . . . . . . . . 71
3.3.2.3. Protection of Endangered Species: The Convention on International
Trade in Endangered Species of Wild Fauna and Flora (CITES) . . . . . . . . . . . . . . . . . . . . . . . . 72
3.3.2.4. Rio 1992 – Combining Conservation and Economics
in International Agreements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
3.4. The Process: Creating and Enforcing International Conservation Law. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
3.5. The Problem of Interdependence: How Does One Nation Promote Global
Conservation without Negative Effects on Other Nations? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
3.5.1. The Nature of International Legal Interdependence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
3.5.2. Case History I: Tuna and Dolphins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
3.5.3. Case History II: Shrimp and Sea Turtles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
3.5.4. Outcomes and Future Prospects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
3.6. Synthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
4 Biodiversity: Concept, Measurement, and Challenge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
4.1. Biodiversity and Conservation Biology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
4.2. The Problem of Concept and Quantity: How Do We Know What Biodiversity is and
How Do We Measure it? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
4.2.1. A Conceptual Defi nition of Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
4.2.2. Biodiversity and the Defi nition of Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
4.2.3. Contemporary Issues of the Species Concept . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
4.2.4. Implications of the Species Concept in Conservation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
4.2.5. Measuring Biodiversity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
4.2.5.1. What Biodiversity Measurements Tell Us. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
4.2.5.2. Alpha Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
4.2.5.3. Beta Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
4.2.5.4. Gamma Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
4.2.6. Application and Integration of Diversity Measures to Address Issues
in Conservation: A Case Study from Eastern Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
4.2.7. Problems of Diversity Indices and Alternative Measures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
4.3. The Problem of Process and Pattern: What Explains Variation in Local Biodiversity? . . . . . . . . . . . . . . . . . . . 95
4.3.1. Niche Assembly Theories of Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
4.3.2. The Unifi ed Neutral Theory of Biodiversity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
4.4. The Problem of Dispersion: Where is Biodiversity Located? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
4.4.1. Global Patterns of Biodiversity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
4.4.2. Biodiversity Indices: Can We Find “Hotspots” with Incomplete Information? . . . . . . . . . . . . . . . . . . . . 100
4.5. The Problem of Quantity: How Much Biodiversity is There?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
4.5.1. General Considerations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
4.5.2. Biodiversity and Rarity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
4.5.2.1. The Problem of Rarity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
4.5.2.2. Habitat Generalists Versus Habitat Specialists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
4.5.2.3. Large Populations Versus Small Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
4.5.2.4. Widespread Distribution Versus Restricted Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
4.5.3. The Problem of Endemism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
4.5.3.1. Endemism in the Extreme – A Case History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
xx Contents
4.5.3.2. Endemism, Biodiversity, and Rarity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
4.5.3.3. Endemism and Island Species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
4.6. The Problem of Application: How Do We Manage Biodiversity? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
4.6.1. The Problem with “Hotspots” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
4.6.2. Identifying Areas of Conservation Value Using Remotely Sensed Data . . . . . . . . . . . . . . . . . . . . . . . . . 107
4.6.3. Tracking Biodiversity Using Indicator Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
4.6.3.1. Biodiversity Indicators: Using “Surrogate” Species as Biodiversity Indices . . . . . . . . . . . . . . . 107
4.6.3.2. Taxon-Based Biodiversity Indicators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
4.6.3.3. Structure- and Function-Based Biodiversity Indicators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
4.6.3.4. Bison as an Example of a Function-Based Keystone Species . . . . . . . . . . . . . . . . . . . . . . . . . . 110
4.6.3.5. Ecological Redundancy and Function-Based Biodiversity Indicators . . . . . . . . . . . . . . . . . . . . 110
4.7. The Problem of Conservation: How Do We Identify and Prioritize Areas to Preserve Biodiversity? . . . . . . . . 111
4.7.1. Current Global Prioritization Strategies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
4.7.2. Management Approaches to Biodiversity at Landscape Levels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
4.7.2.1. Gathering Appropriate Background Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
4.7.2.2. Maintaining Ecological and Evolutionary Processes Promoting Biodiversity . . . . . . . . . . . . . . 113
4.7.2.3. Regional Biodiversity Management – Defi ning Functional Conservation Areas . . . . . . . . . . . . 113
4.7.3. Building Biodiversity Conservation into Existing Management Plans –
The New South Wales Environmental Services Scheme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
4.8. Synthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
5 Biodiversity Conservation and Climate Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
5.1. Climate and Climate Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
5.1.1. Why Does Climate Change Threaten Biodiversity? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
5.1.2. What Is “Climate” and What Is “Climate Change”? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
5.1.3. Should Contemporary Global Warming Be Called “Climate Change?” . . . . . . . . . . . . . . . . . . . . . . . . . 124
5.1.4. The Implications of Rapidly Rising CO2
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
5.1.5. Why We Call It “Climate” Change – Non-temperature Variations
in Climate in a Warming World . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
5.2. The Global Fingerprint of Climate Change on Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
5.2.1. Extinction Patterns in Edith’s Checkerspot Butterfl y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
5.2.2. Finding the Global Fingerprint of Climate Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
5.2.3. Can Climate Change Cause Extinction of Local Populations?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
5.2.3.1. Climate Change and Pikas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
5.2.3.2. Climate Change and Desert Bighorn Sheep . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
5.3. Climate Change in Ecosystems – Species Loss and System Degradation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
5.3.1. Climate Change at Ecosystem Levels: Biome Boundaries and Elevational Shifts. . . . . . . . . . . . . . . . . . 131
5.3.2. Life Zone Changes in Tropical Forests. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
5.3.3. Elevational Shifts in Tropical Cloud Forests: The Case of the Golden Toad . . . . . . . . . . . . . . . . . . . . . . 132
5.4. Climate-Mediated Mechanisms of Ecosystem Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
5.4.1. Climate Infl uences on a Keystone Species: The Case of the Whitebark Pine . . . . . . . . . . . . . . . . . . . . . 133
5.4.2. Climate Infl uences on Ecosystem Processes: Invasive Species in a Warmer World. . . . . . . . . . . . . . . . . 136
5.4.3. Climate Infl uences on Ecosystem Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
5.4.3.1. The Future of Coral in Warmer Oceans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
5.4.3.2. Loss of Polar Sea Ice: Implications for Polar Biodiversity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
5.5. Conservation Planning and Climate Change: Creating Climate-Integrated
Conservation Strategies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
5.5.1. The Bioclimate Envelope: Modeling Climate Effects on Individual Species. . . . . . . . . . . . . . . . . . . . . . 140
5.5.2. Climate Change-Integrated Strategies for Conservation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
5.5.3. Modeling Efforts to Predict Future Responses to Ongoing Climate Change . . . . . . . . . . . . . . . . . . . . . . 145
5.5.4. Errors of Application: Use and Misuse of Endangerment Criteria
to Model Climate Change Effects on Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
5.6. Policy Initiatives for Climate Change and Conservation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
5.7. Synthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150