Conservation biology: evolution in action

Conservatio n

Biology

EVOLUTION IN ACTION

Edited by

SCOT T P CARROL L CHARLE S W FO X

CONSERVATIO N BIOLOG Y

Evolutio n i n Actio n

Edited by

Scott P. Carroll

Charles W . Fox

DAI HOC TEfXl N60T8N

TRUNG TAM HOC LIEU

OXFOR D

UNIVERSITY PRESS

2008

OXTOR D

UNIVERSITY PRESS

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Library of Congress Cataloging-in-Publication Data

Conservation biology: evolution in action / edited by Scott P. Carroll

and Charles W. Fox.

p. cm.

Includes bibliographical references and index.

ISBN: 978-0-19-530679-8; 978-0-19-530678-1 (pbk.)

1. Conservation biology. I. Carroll, Scott P. II. Fox, Charles W.

QH75.C6615 2008

576—dc22 2007044664

98765432 1

Printed in the United States of America

on acid-free paper

Prefac e

At no time in the nearly four billion years since

the origin of life on earth has our planet seen

such tremendous environmental change. Even the

major mass extinctions in prehuman earth his￾tory (Raup & Sepkoski, 1982) are mere blips

in comparison with the current biodiversity crisis.

Human actions and impacts such as the elimina￾tion, fragmentation, and conversion of habitats;

mass poisoning; overharvesting; species introduc￾tions; and climate change dramatically alter the

local and global carrying capacities of other species.

But they also do more. By modifying the chal￾lenges organisms face, and the resources they have

to address those challenges, we are altering the con￾ditions under which behavioral and physiological

traits are expressed and in which ecological interac￾tions occur. These changes affect the selective envi￾ronments encountered by organisms, influencing

evolutionary dynamics, which in turn feed back to

affect ecological dynamics. Conservation problems

are thus eco-evolutionary in nature, rather than just

ecological, demographic, or genetic (Kinnison &

Hairston, 2007). This ecoevolutionary nature of

responses to environmental change is the focus of

this book.

There is thus a clear and present need to develop

practical approaches to managing our biodiver￾sity problems that consider a role of evolution

occurring during the time frame of the conser￾vation program. Evolutionary theory is the pre￾dictive core of the biological sciences, and it

provides the foundation for designing new and

integrative strategies. Central to our perspective is

the discovery that a great many organisms, from

microbes to trees, are rapidly evolving in response to

their changing environments. As risks and resources

change in form, distribution, and abundance, they

create new niches, affect competition, add or sub￾tract enemies, and generally recast the landscape for

surviving taxa. Selection is now operating in new

directions and at new intensities, and the degree

to which populations respond adaptively can deter￾mine their capacity to persist. Moreover, adaptive

evolution, emerging from the demographic and

genetic chaos suffered by "refugee taxa," may prove

to be of foremost importance in altering the form

and structure of species, interspecific interactions,

and communities in the coming years, decades, and

millennia.

If heretofore unanticipated, widespread evolu￾tion is itself a major component of global change.

Static models—those that treat the ecological play￾ers as passive bystanders in the ecological play—

are now obsolete. Understanding and managing

ongoing adaptation to global change requires new

perspectives to accommodate, exploit, and man￾age evolutionary processes of conservation con￾cern, which include population structuring and the

pace and extent of gene flow, the maintenance

and expression of phenotypic polymorphisms and

plasticity, niche specialization versus generalism,

costs versus benefits of harvesting and of genetic

vi Preface

engineering, and diversity management decisions

above and below the species level. If we succeed

in protecting species and biota demographically,

a chief outcome may be to provide raw mate￾rial for both targeted and unmanaged evolution.

Accordingly, the principal challenge of evolution￾ary conservation biology is to predict and then

manage evolutionary dynamics, and make con￾servation (and preservation) plans that maximize

evolutionary potential—for example, by protecting

communities that have unprecedented assemblages

of juxtaposed, and rapidly evolving, remnant taxa.

It is our hope that the authors of this volume provide

insights that ultimately contribute to the success of

such efforts.

This volume is intended to introduce, explore,

and elaborate evolutionary approaches to conserva￾tion biology. The volume is divided into five parts,

each of which is preceded by a brief introduction

and commentary. The chapters in Part I, "Popula￾tion Structure and Genetics of Threatened Taxa,"

present the history and general concepts of con￾servation genetics, and examine the interaction of

genetic and demographic factors. Part II, "Con￾serving Biodiversity within and among Species,"

focuses on evolutionary processes, their relationship

to biodiversity at different taxonomic levels, and

how they influence practical conservation issues,

including the reintroduction of threatened taxa and

the loss of distinctive populations to hybridization.

The chapters in Part III, "Evolutionary Responses

to Environmental Change," examine both genetic

and phenotypic modes of adaptation to the stresses

and opportunities associated with global change

phenomena. Part IV, "Conservation of the Co￾evolving Web of Life," examines the evolutionary

and co-evolutionary causes and consequences of

changing interspecific dynamics, including species

invasions, extinctions, and host parasite dynamics.

The fifth and concluding portion of the volume,

"Evolutionary Management," presents evolution￾ary analyses of three critically important areas:

reserve design, management of transgene flow into

the wilds, and the sustainable harvest of wild

populations.

All chapters in this book were reviewed by peers.

usually two or three scientists with expertise

i n

the topics covered by the chapter. These reviewer

offered insightful commentary on the chapters an

have made this a much better volume. ^ e thank

Karina Acevedo-Whitehouse, Paul Agapow. Fred

Allendorf, Suzanne Alonzo, Mike Angilletta. Tris￾tan Armstrong, Leslie Blancas, Janette Boughman,

Juan Bouzat, Linda Broadhurst, Jeremy Burdon,

Mar Cabeza, Christina Caruso, Denis Couvet,

Richard Cowling, George Gilchrist, John Kelly,

Holly Kindsvater, Mike Kinnison, Mike Loeb, Arne

Mooers, Patrick Nosil, Stephen O'Brien, Julian

Olden, Otso Ovaskainen, William Perry, David

Reed, Gerald Rehfeldt, Kevin Rice, Kim Scrib￾ner, Mike Singer, David Tallmon, John Thomp￾son, Peter Thrall, Andrew Weeks, Alastair Wilson,

and a few reviewers who asked to remain anony￾mous, for their constructive comments on individual

chapters. We especially thank Mike Loeb for copy￾editing chapters and compiling the final version of

the book.

Last, and most important, we thank the authors

for their dedication to this project. The success of

this volume, and its influence on the conservation

community, ultimately depends on the quality of

the chapters and thus on the hard work, creativity,

and insight of the contributing authors. Thanks to

all of you!

Scott P. Carroll

Charles W. Fox

REFERENCES

Kinnison, M. T., & N. G. Hairston Jr. 2007.

Eco-evolutionary conservation biology:

Contemporary evolution and the dynamics of

persistence. Funct Ecol. 21: 444-454

Raup, D., & J. Sepkoski. 1982. Mass extinctions

in the marine fossil record. Science

215:1501-1503.

Content s

List of Contributors

Part I Population Structure and Genetics of Threatened Taxa

Introduction

Charles W. Fox, Scott P. Carroll

1 The History, Purview, and Future of Conservation Genetics

John C. Avise

2 Effects of Population Size on Population Viability: From Mutation to

Environmental Catastrophes

David H. Reed

3 Demographics versus Genetics in Conservation Biology

Barry W. Brook

4 Metapopulation Structure and the Conservation Consequences

of Population Fragmentation

Julianno B. M. Sambatti, Eli Stahl, Susan Harrison

5 The Influence of Breeding Systems and Mating Systems on Conservation

Genetics and Conservation Decisions

Michele R. Dudash, Courtney J. Murren

Part II Conserving Biodiversity within and among Species

Introduction

Fred W. Allendorf

6 The Importance of Conserving Evolutionary Processes

Thomas B. Smith, Gregory F. Grether

7 Phylogenetic Diversity and Conservation

Daniel P. Faith

Contents

8 Genetic Considerations of Introduction Efforts

Philippine Vergeer, N. Joop Ouborg, Andrew P. Hendry

9 Hybridization, Introgression, and the Evolutionary Management

of Threatened Species

Judith M. Rhymer

Part III Evolutionary Responses to Environmental Change

Introduction

George W. Gilchrist, Donna G. Folk

10 Evolution in Response to Climate Change

Julie R. Etterson

11 Evolutionary Dynamics of Adaptation to Environmental Stress

George W. Gilchrist, Donna G. Folk

12 Managing Phenotypic Variability with Genetic and

Environmental Heterogeneity: Adaptation as a First Principle

of Conservation Practice

Scott P. Carroll, Jason V. Watters

13 Genetic Diversity, Adaptive Potential, and Population

Viability in Changing Environments

Elizabeth Grace Boulding

Part IV Conservation of the Coevolving Web of Life

Introduction

John N. Thompson

14 The Geographic Mosaic of Coevolution and Its Conservation

Significance

Craig W. Benkman, Thomas L. Parchman, Adam M. Siepielski

15 The Next Communities: Evolution and Integration of

Invasive Species

Scorr P. Carroll, Charles W. Fox

16 Ecosystem Recovery: Lessons from the Past

Geerat J. Vermel]

17 Host-Pathogen Evolution, Biodiversity, and Disease

Risks for Natural Populations

Sonia Altizer, Amy B. Pedersen

Part V Evolutionary Management

Introduction

Michael T Kinnison