Tài liệu Mountain Goats Ecology, Behavior, and Conservation of an Alpine Ungulate doc

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Mountain Goats

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Mountain Goats

Ecology, Behavior, and Conservation

of an Alpine Ungulate

Marco Festa-Bianchet

Steeve D. Côté

Washington • Covelo • London

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Copyright © 2008 Island Press

All rights reserved under International and Pan-American Copyright Conventions. No part

of this book may be reproduced in any form or by any means without permission in writing

from the publisher: Island Press, 1718 Connecticut Avenue NW, Suite 300, Washington, DC

20009, USA.

Island Press is a trademark of The Center for Resource Economics.

Library of Congress Cataloging-in-Publication Data

Festa-Bianchet, Marco.

Mountain goats : ecology, behavior, and conservation of an alpine ungulate /

Marco Festa-Bianchet, Steeve Côté.

p. cm.

Includes bibliographical references and index.

ISBN 978-1-59726-170-8 (hardcover : alk. paper) --

ISBN 978-1-59726-171-5 (pbk. : alk. paper)

1. Mountain goat. I. Côté, Steeve D. II. Title.

QL737.U53F48 2007

639.97'96475--dc22

2007025958

Printed on recycled, acid-free paper

Manufactured in the United States of America

10 9 8 7 6 5 4 3 2 1

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À Wendy et Mélanie,

qui ont toujours accepté et respecté nos longs séjours

sur la montagne . . .

E per Alberto,

a cui sarebbe piaciuto vedere le capre.

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Contents

Acknowledgments xi

1 Ecological Questions, Conservation Challenges, and

Long-Term Research 1

2 The Study Area and the Goat Population 17

3 Caw Ridge Study Methods and Limitations 36

4 Home Ranges, Forage Availability, and Habitat Use 48

5 Social Organization 71

6 Body and Horn Growth 91

7 Individual Variability in Yearly and Lifetime Reproductive

Success of Females 118

8 Female Reproductive Strategy 137

9 Survival and Dispersal 157

10 Density-Dependence and the Question of

Population Regulation 177

11 Female Reproductive Strategy and Ungulate

Population Dynamics 191

12 Management and Conservation of Mountain Goats 204

13 Long-Term Monitoring of Marked Individuals and Advances in

Ecology and Conservation 219

Appendix 231

Literature Cited 235

Index 257

ix

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Acknowledgments

Over sixteen years, many people helped us continue our research on

mountain goats, and several funding agencies and organizations sup￾ported us financially or helped us logistically. First and foremost, we wish

to thank the many collaborators, students, and assistants who helped us

collect data in the field. These people endured snowstorms in July, me￾chanical breakdowns, questionable food, pesky marmots, and the pres￾ence of grizzly bears to observe, capture, or otherwise study mountain

goats. We hope all have fond memories of Caw Ridge. In alphabetical or￾der, they are: Chantal Beaudoin, Félix Boulanger, Étienne Cardinal,

Guillaume Côté, Étienne Drouin, Donald Dubé, Catherine Gagnon,

Yanick Gendreau, Sandra Hamel, Dave Hildebrand, Mélina Houle, Paul

Jones, Sandro Lovari, Fanie Pelletier, Alberto Peracino, Sabrina Popp,

Sébastien Rioux, Giorgia Romeo, John Russell, Ken Seidel, Geneviève

Simard, Martin Urquhart, Lucie Vallières, Vanessa Viera, and Sébastien

Wendenbaum. We wish to single out Martin Urquhart, who ensured that

our first critical years of fieldwork were successful, Yanick Gendreau,

who contributed much effort and enthusiasm to collecting data from

1998 to 2002, and Sandra Hamel, whose passion for mountain goats and

for Caw Ridge defies description, and who was the main force behind this

research from 2002 through 2006.

The real work in ecological research is done by graduate students, and

the Caw Ridge study was no exception. We are thankful to have had so

many enthusiastic and resourceful graduate students, the secret weapon of

this research program. Many have gone on to successful careers in wildlife

conservation or research, and we are proud of their achievements. In

xi

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chronological order, they are François Fournier, Martine Haviernick,

Alejandro Gonzalez-Voyer, Yanick Gendreau, Sandra Hamel, and Julien

Mainguy.

The logistics of the Caw Ridge study are complex, as is to be expected

of any research program in remote areas with difficult access and unreli￾able communication lines. We thank all those that made our life easier in

the field by providing logistical support, helped us in the laboratory, or

generally got us out of trouble: Bill Allen, Janet Ficht, Steven Cross,

Mike Ewald, Dave Hobson, Dave McKenna, Bertrand Mercier, Kirby

Smith, and Shane Ramstead. Institutional logistical support was provided

by the Alberta Forest Service, Grande Cache Correctional Center (we

are very grateful for the construction of our traps and the field cabin),

Renewable Resources Consultants, and Smoky River Coal, Ltd.

As we wrote various drafts of this book, we received many wise and

constructive comments from several colleagues, including Tim Coulson,

Jean-Michel Gaillard, Sandra Hamel, Wendy King, Julien Mainguy,

Fanie Pelletier, Cliff Rice, Kathreen Ruckstuhl, and Kirby Smith. We

thank Dave Coltman, Tim Coulson, Jean-Michel Gaillard, Jon Jorgen￾son, and Kirby Smith for ideas and discussions that helped shape our

thinking about mountain ungulates and their conservation.

We acknowledge the pivotal role played by Kirby Smith in the Caw

Ridge Mountain goat study. Kirby first suggested that we choose Caw

Ridge as a study area, set up the initial capture operation, helped us de￾fine the goals of the study, and provided logistic support on innumerable

occasions. His unswerving dedication to wildlife conservation and to the

study of mountain goats was among the main assets of this research pro￾gram. His good humor, hospitality, and knowledge were always much

appreciated.

We were able to carry on this long-term study because we were finan￾cially supported by agencies that recognize the importance of funda￾mental research on wild animals. Our study was generously supported

by the Natural Sciences and Engineering Research Council of Canada

(NSERC, which provided operating and equipment grants to us and

scholarships to some of our students), the Fonds Québécois de la Re -

cherche sur la Nature et les Technologies, Alberta Fish and Wildlife, Al￾berta Conservation Association (ACA), Alberta Recreation, Parks and

Wildlife Foundation, Alberta Wildlife Enhancement Fund, Challenge

Grants in Biodiversity (ACA), International Order of Rocky Mountain

Goats, Rocky Mountain Goat Foundation, the Université de Sher￾brooke, and Université Laval. We thank Alberta Fish and Wildlife and

the British Columbia Ministry of the Environment for supporting the

publication of this book.

xii Acknowledgments

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chapter 1

Ecological Questions, Conservation

Challenges, and Long-Term Research

The conservation of biodiversity and the management of wildlife require

an understanding of the basic ecology of animals. That deceptively sim￾ple statement conceals a difficult problem, because understanding the

“basic ecology” of species demands years of research. The processes that

affect population dynamics of large mammals often develop over many

years and cannot be understood without long-term monitoring. Impor￾tant events (such as forest fires, extreme weather, or the appearance of

new predators, competitors, or diseases) may have drastic long-term ef￾fects on population dynamics but they may be too rare to be detected, let

alone quantified, by a few years of research. In addition, the many factors

affecting a species’ abundance seldom act in isolation. Instead, interac￾tions between factors are commonplace: for example, body mass may af￾fect survival only at high population density (Festa-Bianchet et al. 1997),

and the impacts of a harsh winter may vary substantially according to a

population’s age structure (Coulson et al. 2000). Similar complex rela￾tionships affect the consequences of different harvest levels, which can be

radically different according to the sex–age composition of the popula￾tion and of harvested animals.

Consequently, an in-depth understanding of ungulate ecology re￾quires data collected over many years and can best be served by long￾term studies that seek to answer fundamental questions: What affects

population size? What factors determine age- and sex-specific mortality?

How do individuals differ in their ability to contribute to population re￾cruitment, and why do those differences exist?

1

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A long-term approach to the study of the ecology and conservation of

large herbivores is particularly appropriate because of their longevity and

complex population structure. An individual can experience varying lev￾els of environmental conditions over its lifetime. Consequently, the re￾productive strategy of large herbivores likely evolved in response to the

range of environmental conditions that an individual may encounter over

its lifetime. The complex population structure of large herbivores, some￾times involving a dozen or more coexisting cohorts, means that the pop￾ulation present today is the result of processes and events that took place

over the previous decade and will affect population dynamics over the

next one. It is therefore essential that management programs to conserve

large herbivores, including those that involve some harvest, be mindful

of the differences among individuals. In addition, the consequences of

conservation actions (or of harvest strategies) can persist over many

years. Because of the importance of differences among individuals and of

processes occurring over several years, biological knowledge useful for

conservation of large herbivores can therefore be best obtained by long￾term monitoring of known individuals within a population.

Public finances typically sustain fundamental ecological studies. Our

mountain goat study is no exception. In addition to producing novel in￾formation, ecological studies have an obligation to clearly communicate

the applied implications of their results. The conservation of biodiversity

requires long-term research, and long-term research should make a con￾tribution to conservation. We will use our sixteen years of research on

mountain goats to show how some aspects of the biology of this species

play a fundamental role in its conservation. We will do so by examining

the adaptations of mountain goats to their alpine environment, and by

underlining differences and similarities between mountain goats and

other large herbivores, in particular other mountain-dwelling ungulates.

Why Study Mountain Goats?

Mountain goats provide research challenges and opportunities from both

a fundamental and an applied viewpoint. There is much concern for the

conservation of this species, which appears highly sensitive to both har￾vest and disturbance. In addition, its alpine habitat is very sensitive to hu￾man activities and is likely at risk from the effects of climate change.

Four factors combined to provide the stimulus to study mountain

goats at Caw Ridge. First, an unexplained and rapid decline in mountain

goat numbers in Alberta led to the complete closure of goat hunting in

the province in 1987 (Smith 1988b). Combined with the lack of informa￾2 Mountain Goats

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