Wildlife Ecology, Conservation, and Management

Wildlife Ecology, Conservation, and Management

Wildlife Ecology, Conservation, and Management

489 Pages ·2005·2.7 MB ·English

Wildlife Ecology, Conservation, and Management

WECA01 08/17/2005 04:34PM Page i


Wildlife Ecology, Conservation, and Management WECA01 08/17/2005 04:34PM Page ii


To our colleagues Graeme Caughley, Peter Yodzis, and James N.M.


Smith who have influenced both our approach to wildlife biology


and the writing of this book. WECA01 08/17/2005 04:34PM Page iii


Wildlife Ecology, Conservation,


and Management


Second Edition


Anthony R.E. Sinclair PhD, FRS


Biodiversity Research Centre, University of British Columbia,


Vancouver, Canada


John M. Fryxell PhD


Department of Integrative Biology, University of Guelph, Guelph,


Canada


Graeme Caughley PhD


CSIRO, Canberra, Australia WECA01 08/17/2005 04:34PM Page iv


© 2006 by Anthony R.E. Sinclair and John M. Fryxell


© 1994 by Blackwell Science


BLACKWELL PUBLISHING


350 Main Street, Malden, MA 02148-5020, USA


9600 Garsington Road, Oxford OX4 2DQ, UK


550 Swanston Street, Carlton, Victoria 3053, Australia


The right of Anthony R.E. Sinclair and John M. Fryxell to be identified as the Authors of this


Work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988.


All rights reserved. No part of this publication may be reproduced, stored in a retrieval system,


or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or


otherwise, except as permitted by the UK Copyright, Designs, and Patents Act 1988, without the


prior permission of the publisher.


First edition published 1994


Second edition published 2006 by Blackwell Publishing Ltd


1 2006


Library of Congress Cataloging-in-Publication Data


Sinclair, A.R.E. (Anthony Ronald Entrican)


Wildlife ecology, conservation, and management / Anthony R.E.


Sinclair, John M. Fryxell, Graeme Caughley. – 2nd ed.


p. cm.


Rev. ed. of: Wildlife ecology and management / Graeme Caughley,


Anthony R.E. Sinclair.


Includes bibliographical references and index.


ISBN-13: 978-1-4051-0737-2 (pbk. : alk. paper)


ISBN-10: 1-4051-0737-5 (pbk. : alk. paper)


1. Wildlife management. 2. Wildlife conservation. 3. Animal ecology.


I. Fryxell, John M., 1954– II. Caughley, Graeme. III. Caughley, Graeme.


=


Wildlife ecology and management. IV. Title. 20


SK355.C38 2005


639.9–dc22


2005007229


A catalogue record for this title is available from the British Library.


Set in 9.5/12pt Berkeley


by Graphicraft Limited, Hong Kong


Printed and bound in India


by Replika Press


The publisher’s policy is to use permanent paper from mills that operate a sustainable forestry


policy, and which hasbeen manufactured from pulp processed using acid-free and elementary


chlorine-free practices. Furthermore, the publisher ensures that the text paper and cover board


used have met acceptable environmental accreditation standards.


For further information on


Blackwell Publishing, visit our website:


www.blackwellpublishing.com WECA01 08/17/2005 04:34PM Page v


Contents


Preface xi


1 Introduction: goals and decisions 1


1.1 How to use this book 1


1.2 What is wildlife conservation and management? 2


1.3 Goals of management 3


1.4 Hierarchies of decision 5


1.5 Policy goals 7


1.6 Feasible options 8


1.7 Summary 8


Part 1 Wildlife ecology 9


2 Biomes 11


2.1 Introduction 11


2.2 Forest biomes 12


2.3 Woodland biomes 14


2.4 Shrublands 14


2.5 Grassland biomes 15


2.6 Semi-desert scrub 17


2.7 Deserts 17


2.8 Marine biomes 17


2.9 Summary 18


3 Animals as individuals 19


3.1 Introduction 19


3.2 Adaptation 19


3.3 The theory of natural selection 19


3.4 Examples of adaptation 21


3.5 The effects of history 23


3.6 The abiotic environment 27


3.7 Genetic characteristics of individuals 27


3.8 Applied aspects 33


3.9 Summary 35


v WECA01 08/17/2005 04:34PM Page vi


vi CONTENTS


4 Food and nutrition 36


4.1 Introduction 36


4.2 Constituents of food 36


4.3 Variation in food supply 40


4.4 Measurement of food supply 42


4.5 Basal metabolic rate and food requirement 46


4.6 Morphology of herbivore digestion 49


4.7 Food passage rate and food requirement 51


4.8 Body size and diet selection 52


4.9 Indices of body condition 53


4.10 Summary 59


5 The ecology of behavior 60


5.1 Introduction 60


5.2 Diet selection 60


5.3 Optimal patch or habitat use 66


5.4 Risk-sensitive habitat use 69


5.5 Quantifying habitat preference using resource


selection functions 70


5.6 Social behavior and foraging 72


5.7 Summary 77


6 Population growth 78


6.1 Introduction 78


6.2 Rate of increase 78


6.3 Fecundity rate 82


6.4 Mortality rate 82


6.5 Direct estimation of life-table parameters 84


6.6 Indirect estimation of life-table parameters 85


6.7 Relationship between parameters 87


6.8 Geometric or exponential population growth 88


6.9 Summary 89


7 Dispersal, dispersion, and distribution 90


7.1 Introduction 90


7.2 Dispersal 90


7.3 Dispersion 92


7.4 Distribution 93


7.5 Distribution, abundance, and range collapse 98


7.6 Species reintroductions or invasions 99


7.7 Dispersal and the sustainability of metapopulations 104


7.8 Summary 108


8 Population regulation, fluctuation, and competition within


species 109


8.1 Introduction 109


8.2 Stability of populations 109


8.3 The theory of population limitation and regulation 111 WECA01 08/17/2005 04:34PM Page vii


CONTENTS vii


8.4 Evidence for regulation 116


8.5 Applications of regulation 120


8.6 Logistic model of population regulation 121


8.7 Stability, cycles, and chaos 125


8.8 Intraspecific competition 131


8.9 Interactions of food, predators, and disease 134


8.10 Summary 134


9 Competition and facilitation between species 135


9.1 Introduction 135


9.2 Theoretical aspects of interspecific competition 136


9.3 Experimental demonstrations of competition 138


9.4 The concept of the niche 143


9.5 The competitive exclusion principle 146


9.6 Resource partitioning and habitat selection 146


9.7 Competition in variable environments 153


9.8 Apparent competition 153


9.9 Facilitation 154


9.10 Applied aspects of competition 159


9.11 Summary 162


10 Predation 163


10.1 Introduction 163


10.2 Predation and management 163


10.3 Definitions 163


10.4 The effect of predators on prey density 164


10.5 The behavior of predators 165


10.6 Numerical response of predators to prey density 169


10.7 The total response 170


10.8 Behavior of the prey 176


10.9 Summary 178


11 Parasites and pathogens 179


11.1 Introduction and definitions 179


11.2 Effects of parasites 179


11.3 The basic parameters of epidemiology 180


11.4 Determinants of spread 183


11.5 Endemic pathogens 184


11.6 Endemic pathogens: synergistic interactions with food


and predators 184


11.7 Epizootic diseases 186


11.8 Emerging infectious diseases of wildlife 187


11.9 Parasites and the regulation of host populations 188


11.10 Parasites and host communities 190


11.11 Parasites and conservation 191


11.12 Parasites and control of pests 194


11.13 Summary 195 WECA01 08/17/2005 04:34PM Page viii


viii CONTENTS


12 Consumer–resource dynamics 196


12.1 Introduction 196


12.2 Quality and quantity of a resource 196


12.3 Kinds of resources 196


12.4 Consumer–resource dynamics: general theory 197


12.5 Kangaroos and their food plants in semi-arid


Australian savannas 200


12.6 Wolf–moose–woody plant dynamics in the boreal forest 207


12.7 Other population cycles 212


12.8 Summary 215


Part 2 Wildlife conservation and management 217


13 Counting animals 219


13.1 Introduction 219


13.2 Estimates 219


13.3 Total counts 219


13.4 Sampled counts: the logic 221


13.5 Sampled counts: methods and arithmetic 226


13.6 Indirect estimates of population size 235


13.7 Indices 241


13.8 Summary 243


14 Age and stage structure 244


14.1 Age-specific population models 244


14.2 Stage-specific models 247


14.3 Sensitivity and elasticity of matrix models 248


14.4 Short-term changes in structured populations 251


14.5 Summary 252


15 Model evaluation and adaptive management 253


15.1 Introduction 253


15.2 Fitting models to data and estimation of parameters 254


15.3 Measuring the likelihood of models in light of


the observed data 256


15.4 Evaluating the likelihood of alternative models using AIC 258


15.5 Adaptive management 264


15.6 Summary 267


16 Experimental management 268


16.1 Introduction 268


16.2 Differentiating success from failure 268


16.3 Technical judgments can be tested 269


16.4 The nature of the evidence 272


16.5 Experimental and survey design 274


16.6 Some standard analyses 279


16.7 Summary 287 WECA01 08/17/2005 04:34PM Page ix


CONTENTS ix


17 Conservation in theory 289


17.1 Introduction 289


17.2 Demographic problems contributing to risk of extinction 289


17.3 Genetic problems contributing to risk of extinction 291


17.4 Effective population size (genetic) 297


17.5 Effective population size (demographic) 298


17.6 How small is too small? 299


17.7 Population viability analysis 300


17.8 Extinction caused by environmental change 305


17.9 Summary 310


18 Conservation in practice 312


18.1 Introduction 312


18.2 How populations go extinct 312


18.3 How to prevent extinction 321


18.4 Rescue and recovery of near extinctions 323


18.5 Conservation in national parks and reserves 324


18.6 Community conservation outside national parks and reserves 332


18.7 International conservation 332


18.8 Summary 334


19 Wildlife harvesting 335


19.1 Introduction 335


19.2 Fixed quota harvesting strategy 335


19.3 Fixed proportion harvesting strategy 341


19.4 Fixed escapement harvesting strategy 344


19.5 Harvesting in practice: recreational 346


19.6 Harvesting in practice: commercial 346


19.7 Age- or sex-biased harvesting 347


19.8 Bioeconomics 347


19.9 Game cropping and the discount rate 352


19.10 Summary 353


20 Wildlife control 355


20.1 Introduction 355


20.2 Definitions 355


20.3 Effects of control 356


20.4 Objectives of control 356


20.5 Determining whether control is appropriate 357


20.6 Methods of control 358


20.7 Summary 364


21 Ecosystem management and conservation 365


21.1 Introduction 365


21.2 Definitions 365


21.3 Gradients of communities 366


21.4 Niches 366


21.5 Food webs and intertrophic interactions 366


WECA01 08/17/2005 04:34PM Page i


Wildlife Ecology, Conservation, and Management WECA01 08/17/2005 04:34PM Page ii


To our colleagues Graeme Caughley, Peter Yodzis, and James N.M.


Smith who have influenced both our approach to wildlife biology


and the writing of this book. WECA01 08/17/2005 04:34PM Page iii


Wildlife Ecology, Conservation,


and Management


Second Edition


Anthony R.E. Sinclair PhD, FRS


Biodiversity Research Centre, University of British Columbia,


Vancouver, Canada


John M. Fryxell PhD


Department of Integrative Biology, University of Guelph, Guelph,


Canada


Graeme Caughley PhD


CSIRO, Canberra, Australia WECA01 08/17/2005 04:34PM Page iv


© 2006 by Anthony R.E. Sinclair and John M. Fryxell


© 1994 by Blackwell Science


BLACKWELL PUBLISHING


350 Main Street, Malden, MA 02148-5020, USA


9600 Garsington Road, Oxford OX4 2DQ, UK


550 Swanston Street, Carlton, Victoria 3053, Australia


The right of Anthony R.E. Sinclair and John M. Fryxell to be identified as the Authors of this


Work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988.


All rights reserved. No part of this publication may be reproduced, stored in a retrieval system,


or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or


otherwise, except as permitted by the UK Copyright, Designs, and Patents Act 1988, without the


prior permission of the publisher.


First edition published 1994


Second edition published 2006 by Blackwell Publishing Ltd


1 2006


Library of Congress Cataloging-in-Publication Data


Sinclair, A.R.E. (Anthony Ronald Entrican)


Wildlife ecology, conservation, and management / Anthony R.E.


Sinclair, John M. Fryxell, Graeme Caughley. – 2nd ed.


p. cm.


Rev. ed. of: Wildlife ecology and management / Graeme Caughley,


Anthony R.E. Sinclair.


Includes bibliographical references and index.


ISBN-13: 978-1-4051-0737-2 (pbk. : alk. paper)


ISBN-10: 1-4051-0737-5 (pbk. : alk. paper)


1. Wildlife management. 2. Wildlife conservation. 3. Animal ecology.


I. Fryxell, John M., 1954– II. Caughley, Graeme. III. Caughley, Graeme.


=


Wildlife ecology and management. IV. Title. 20


SK355.C38 2005


639.9–dc22


2005007229


A catalogue record for this title is available from the British Library.


Set in 9.5/12pt Berkeley


by Graphicraft Limited, Hong Kong


Printed and bound in India


by Replika Press


The publisher’s policy is to use permanent paper from mills that operate a sustainable forestry


policy, and which hasbeen manufactured from pulp processed using acid-free and elementary


chlorine-free practices. Furthermore, the publisher ensures that the text paper and cover board


used have met acceptable environmental accreditation standards.


For further information on


Blackwell Publishing, visit our website:


www.blackwellpublishing.com WECA01 08/17/2005 04:34PM Page v


Contents


Preface xi


1 Introduction: goals and decisions 1


1.1 How to use this book 1


1.2 What is wildlife conservation and management? 2


1.3 Goals of management 3


1.4 Hierarchies of decision 5


1.5 Policy goals 7


1.6 Feasible options 8


1.7 Summary 8


Part 1 Wildlife ecology 9


2 Biomes 11


2.1 Introduction 11


2.2 Forest biomes 12


2.3 Woodland biomes 14


2.4 Shrublands 14


2.5 Grassland biomes 15


2.6 Semi-desert scrub 17


2.7 Deserts 17


2.8 Marine biomes 17


2.9 Summary 18


3 Animals as individuals 19


3.1 Introduction 19


3.2 Adaptation 19


3.3 The theory of natural selection 19


3.4 Examples of adaptation 21


3.5 The effects of history 23


3.6 The abiotic environment 27


3.7 Genetic characteristics of individuals 27


3.8 Applied aspects 33


3.9 Summary 35


v WECA01 08/17/2005 04:34PM Page vi


vi CONTENTS


4 Food and nutrition 36


4.1 Introduction 36


4.2 Constituents of food 36


4.3 Variation in food supply 40


4.4 Measurement of food supply 42


4.5 Basal metabolic rate and food requirement 46


4.6 Morphology of herbivore digestion 49


4.7 Food passage rate and food requirement 51


4.8 Body size and diet selection 52


4.9 Indices of body condition 53


4.10 Summary 59


5 The ecology of behavior 60


5.1 Introduction 60


5.2 Diet selection 60


5.3 Optimal patch or habitat use 66


5.4 Risk-sensitive habitat use 69


5.5 Quantifying habitat preference using resource


selection functions 70


5.6 Social behavior and foraging 72


5.7 Summary 77


6 Population growth 78


6.1 Introduction 78


6.2 Rate of increase 78


6.3 Fecundity rate 82


6.4 Mortality rate 82


6.5 Direct estimation of life-table parameters 84


6.6 Indirect estimation of life-table parameters 85


6.7 Relationship between parameters 87


6.8 Geometric or exponential population growth 88


6.9 Summary 89


7 Dispersal, dispersion, and distribution 90


7.1 Introduction 90


7.2 Dispersal 90


7.3 Dispersion 92


7.4 Distribution 93


7.5 Distribution, abundance, and range collapse 98


7.6 Species reintroductions or invasions 99


7.7 Dispersal and the sustainability of metapopulations 104


7.8 Summary 108


8 Population regulation, fluctuation, and competition within


species 109


8.1 Introduction 109


8.2 Stability of populations 109


8.3 The theory of population limitation and regulation 111 WECA01 08/17/2005 04:34PM Page vii


CONTENTS vii


8.4 Evidence for regulation 116


8.5 Applications of regulation 120


8.6 Logistic model of population regulation 121


8.7 Stability, cycles, and chaos 125


8.8 Intraspecific competition 131


8.9 Interactions of food, predators, and disease 134


8.10 Summary 134


9 Competition and facilitation between species 135


9.1 Introduction 135


9.2 Theoretical aspects of interspecific competition 136


9.3 Experimental demonstrations of competition 138


9.4 The concept of the niche 143


9.5 The competitive exclusion principle 146


9.6 Resource partitioning and habitat selection 146


9.7 Competition in variable environments 153


9.8 Apparent competition 153


9.9 Facilitation 154


9.10 Applied aspects of competition 159


9.11 Summary 162


10 Predation 163


10.1 Introduction 163


10.2 Predation and management 163


10.3 Definitions 163


10.4 The effect of predators on prey density 164


10.5 The behavior of predators 165


10.6 Numerical response of predators to prey density 169


10.7 The total response 170


10.8 Behavior of the prey 176


10.9 Summary 178


11 Parasites and pathogens 179


11.1 Introduction and definitions 179


11.2 Effects of parasites 179


11.3 The basic parameters of epidemiology 180


11.4 Determinants of spread 183


11.5 Endemic pathogens 184


11.6 Endemic pathogens: synergistic interactions with food


and predators 184


11.7 Epizootic diseases 186


11.8 Emerging infectious diseases of wildlife 187


11.9 Parasites and the regulation of host populations 188


11.10 Parasites and host communities 190


11.11 Parasites and conservation 191


11.12 Parasites and control of pests 194


11.13 Summary 195 WECA01 08/17/2005 04:34PM Page viii


viii CONTENTS


12 Consumer–resource dynamics 196


12.1 Introduction 196


12.2 Quality and quantity of a resource 196


12.3 Kinds of resources 196


12.4 Consumer–resource dynamics: general theory 197


12.5 Kangaroos and their food plants in semi-arid


Australian savannas 200


12.6 Wolf–moose–woody plant dynamics in the boreal forest 207


12.7 Other population cycles 212


12.8 Summary 215


Part 2 Wildlife conservation and management 217


13 Counting animals 219


13.1 Introduction 219


13.2 Estimates 219


13.3 Total counts 219


13.4 Sampled counts: the logic 221


13.5 Sampled counts: methods and arithmetic 226


13.6 Indirect estimates of population size 235


13.7 Indices 241


13.8 Summary 243


14 Age and stage structure 244


14.1 Age-specific population models 244


14.2 Stage-specific models 247


14.3 Sensitivity and elasticity of matrix models 248


14.4 Short-term changes in structured populations 251


14.5 Summary 252


15 Model evaluation and adaptive management 253


15.1 Introduction 253


15.2 Fitting models to data and estimation of parameters 254


15.3 Measuring the likelihood of models in light of


the observed data 256


15.4 Evaluating the likelihood of alternative models using AIC 258


15.5 Adaptive management 264


15.6 Summary 267


16 Experimental management 268


16.1 Introduction 268


16.2 Differentiating success from failure 268


16.3 Technical judgments can be tested 269


16.4 The nature of the evidence 272


16.5 Experimental and survey design 274


16.6 Some standard analyses 279


16.7 Summary 287 WECA01 08/17/2005 04:34PM Page ix


CONTENTS ix


17 Conservation in theory 289


17.1 Introduction 289


17.2 Demographic problems contributing to risk of extinction 289


17.3 Genetic problems contributing to risk of extinction 291


17.4 Effective population size (genetic) 297


17.5 Effective population size (demographic) 298


17.6 How small is too small? 299


17.7 Population viability analysis 300


17.8 Extinction caused by environmental change 305


17.9 Summary 310


18 Conservation in practice 312


18.1 Introduction 312


18.2 How populations go extinct 312


18.3 How to prevent extinction 321


18.4 Rescue and recovery of near extinctions 323


18.5 Conservation in national parks and reserves 324


18.6 Community conservation outside national parks and reserves 332


18.7 International conservation 332


18.8 Summary 334


19 Wildlife harvesting 335


19.1 Introduction 335


19.2 Fixed quota harvesting strategy 335


19.3 Fixed proportion harvesting strategy 341


19.4 Fixed escapement harvesting strategy 344


19.5 Harvesting in practice: recreational 346


19.6 Harvesting in practice: commercial 346


19.7 Age- or sex-biased harvesting 347


19.8 Bioeconomics 347


19.9 Game cropping and the discount rate 352


19.10 Summary 353


20 Wildlife control 355


20.1 Introduction 355


20.2 Definitions 355


20.3 Effects of control 356


20.4 Objectives of control 356


20.5 Determining whether control is appropriate 357


20.6 Methods of control 358


20.7 Summary 364


21 Ecosystem management and conservation 365


21.1 Introduction 365


21.2 Definitions 365


21.3 Gradients of communities 366


21.4 Niches 366


21.5 Food webs and intertrophic interactions 366


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