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To understand modern principles of sustainable management andthe conservation of wildlife species requires intimate knowledgeabout demography, animal behavior, and ecosystem dynamics. Withemphasis on practical application and quantitative skilldevelopment, this book weaves together these disparate elements ina single coherent textbook for senior undergraduate and graduate students.It reviews analytical techniques, explaining the mathematical andstatistical principles behind them, and shows how these can be usedto formulate realistic objectives within an ecological framework. This third edition is comprehensive and up-to-date, andincludes: * Brand new chapters that disseminate rapidly developing topics inthe field: habitat use and selection; habitat fragmentation,movement, and corridors; population viability. analysis, theconsequences of climate change; and evolutionary responses todisturbance * A thorough updating of all chapters to present important areas ofwildlife research and management with recent developments andexamples. * A new online study aid - a wide variety of downloadablecomputer programs in the freeware packages R and Mathcad, availablethrough a companion website. Worked examples enable readers topractice calculations explained in the text and to develop a solidunderstanding of key statistical procedures and population modelscommonly used in wildlife ecology and management. The first half of the book provides a solid background in keyecological concepts. The second half uses these concepts to developa deeper understanding of the principles underlying wildlifemanagement and conservation. Global examples of real-lifemanagement situations provide a broad perspective on theinternational problems of conservation, and detailed case historiesdemonstrate concepts and quantitative analyses. This third editionis also valuable to professional wildlife managers, park rangers,biological resource managers, and those working in ecotourism.
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Cover
Dedication
Title Page
Copyright
Preface
About the companion website
Chapter 1: Introduction: goals and decisions
1.1 How to use this book
1.2 What is wildlife conservation and management?
1.3 Goals of management
1.4 Hierarchies of decision
1.5 Policy goals
1.6 Feasible options
1.7 Summary
Part 1: Wildlife ecology
Chapter 2: Food and nutrition
2.1 Introduction
2.2 Constituents of food
2.3 Variation in food supply
2.4 Measurement of food supply
2.5 Basal metabolic rate and food requirement
2.6 Morphology of herbivore digestion
2.7 Food passage rate and food requirement
2.8 Body size and diet selection
2.9 Indices of body condition
2.10 Summary
Chapter 3: Home range and habitat use
3.1 Introduction
3.2 Estimating home range size and utilization frequency
3.3 Estimating habitat availability and use
3.4 Selective habitat use
3.5 Using resource selection functions to predict population response
3.6 Sources of variation in habitat use
3.7 Movement within the home range
3.8 Movement among home ranges
3.9 Summary
Chapter 4: Dispersal, dispersion, and distribution
4.1 Introduction
4.2 Dispersal
4.3 Dispersion
4.4 Distribution
4.5 Distribution, abundance, and range collapse
4.6 Species reintroductions or invasions
4.7 Summary
Chapter 5: Population growth and regulation
5.1 Introduction
5.2 Rate of increase
5.3 Geometric or exponential population growth
5.4 Stability of populations
5.5 The theory of population limitation and regulation
5.6 Evidence for regulation
5.7 Applications of regulation
5.8 Logistic model of population regulation
5.9 Stability, cycles, and chaos
5.10 Intraspecific competition
5.11 Interactions of food, predators, and disease
5.12 Summary
Chapter 6: Competition and facilitation between species
6.1 Introduction
6.2 Theoretical aspects of interspecific competition
6.3 Experimental demonstrations of competition
6.4 The concept of the niche
6.5 The competitive exclusion principle
6.6 Resource partitioning and habitat selection
6.7 Competition in variable environments
6.8 Apparent competition
6.9 Facilitation
6.10 Applied aspects of competition
6.11 Summary
Chapter 7: Predation
7.1 Introduction
7.2 Predation and management
7.3 Definitions
7.4 The effect of predators on prey density
7.5 The behavior of predators
7.6 Numerical response of predators to prey density
7.7 The total response
7.8 Behavior of the prey
7.9 Summary
Chapter 8: Parasites and pathogens
8.1 Introduction and definitions
8.2 Effects of parasites
8.3 The basic parameters of epidemiology
8.4 Determinants of spread
8.5 Endemic pathogens
8.6 Endemic pathogens: synergistic interactions with food and predators
8.7 Epizootic diseases
8.8 Emerging infectious diseases of wildlife
8.9 Parasites and the regulation of host populations
8.10 Parasites and host communities
8.11 Parasites and conservation
8.12 Parasites and control of pests
8.13 Summary
Chapter 9: Consumer–resource dynamics
9.1 Introduction
9.2 Quality and quantity of a resource
9.3 Kinds of resource
9.4 Consumer–resource dynamics: general theory
9.5 Kangaroos and their food plants in semi-arid Australian savannas
9.6 Wolf–moose–woody plant dynamics in the boreal forest
9.7 Other population cycles
9.8 Summary
Chapter 10: The ecology of behavior
10.1 Introduction
10.2 Diet selection
10.3 Optimal patch or habitat use
10.4 Risk-sensitive habitat use
10.5 Social behavior and foraging
10.6 Summary
Chapter 11: Climate change and wildlife
11.1 Introduction
11.2 Evidence for climate change
11.3 Wildlife responses to climate change
11.4 Mechanisms of response to climate change
11.5 Complex ecosystem responses to climate change
11.6 Summary
Part 2: Wildlife conservation and management
Chapter 12: Counting animals
12.1 Introduction
12.2 Total counts
12.3 Sampled counts: the logic
12.4 Sampled counts: methods and arithmetic
12.5 Indirect estimates of population size
12.6 Indices
12.7 Harvest-based population estimates
12.8 Summary
Chapter 13: Age and stage structure
13.1 Introduction
13.2 Demographic rates
13.3 Direct estimation of life table parameters
13.4 Indirect estimation of life table parameters
13.5 Relationships among parameters
13.6 Age-specific population models
13.7 Elasticity of matrix models
13.8 Stage-specific models
13.9 Elasticity of the loggerhead turtle model
13.10 Short-term changes in structured populations
13.11 Environmental stochasticity and age-structured populations
13.12 Summary
Chapter 14: Experimental management
14.1 Introduction
14.2 Differentiating success from failure
14.3 Technical judgments can be tested
14.4 The nature of the evidence
14.5 Experimental and survey design
14.6 Some standard analyses
14.7 Summary
Chapter 15: Model evaluation and adaptive management
15.1 Introduction
15.2 Fitting models to data and estimation of parameters
15.3 Measuring the likelihood of the observed data
15.4 Evaluating the likelihood of alternate models using AIC
15.5 Adaptive management
15.6 Summary
Chapter 16: Population viability analysis
16.1 Introduction
16.2 Environmental stochasticity
16.3 PVA based on the exponential growth model
16.4 PVA based on the diffusion model
16.5 PVA based on logistic growth
16.6 Demographic stochasticity
16.7 Estimating both environmental and demographic stochasticity
16.8 PVA based on demographic and environmental stochasticity
16.9 Strengths and weaknesses of PVA
16.10 Extinction caused by environmental change
16.11 Extinction threat due to introduction of exotic predators or competitors
16.12 Extinction threat due to unsustainable harvesting
16.13 Extinction threat due to habitat loss
16.14 Summary
Chapter 17: Conservation in practice
17.1 Introduction
17.2 How populations go extinct
17.3 How to prevent extinction
17.4 Rescue and recovery of near-extinctions
17.5 Conservation in National Parks and reserves
17.6 Community conservation outside National Parks and reserves
17.7 International conservation
17.8 Summary
Chapter 18: Wildlife harvesting
18.1 Introduction
18.2 Fixed-quota harvesting strategy
18.3 Fixed-proportion harvesting strategy
18.4 Harvesting in practice: dynamic variation in quotas or effort
18.5 No-harvest reserves
18.6 Age- or sex-biased harvesting
18.7 Commercial harvesting
18.8 Bioeconomics
18.9 Game cropping and the discount rate
18.10 Summary
Chapter 19: Wildlife control
19.1 Introduction
19.2 Definitions
19.3 Effects of control
19.4 Objectives of control
19.5 Determining whether control is appropriate
19.6 Methods of control
19.7 Summary
Chapter 20: Evolution and conservation genetics
20.1 Introduction
20.2 Maintenance of genetic variation
20.3 Natural selection
20.4 Natural selection and life history tradeoffs
20.5 Natural selection due to hunting
20.6 Natural selection due to fishing
20.7 Selection due to environmental change
20.8 Ecological dynamics due to evolutionary changes
20.9 Heterozygosity
20.10 Genetic drift and mutation
20.11 Inbreeding depression
20.12 How much genetic variation is needed?
20.13 Effective population size
20.14 Effect of sex ratio
20.15 How small is too small?
20.16 Summary
Chapter 21: Habitat loss and metapopulation dynamics
21.1 Introduction
21.2 Habitat loss and fragmentation
21.3 Ecological effects of habitat loss
21.4 Metapopulation dynamics
21.5 Territorial metapopulations
21.6 Mainland–island metapopulations
21.7 Source–sink metapopulations
21.8 Metacommunity dynamics of competitors
21.9 Metacommunity dynamics of predators and prey
21.10 Corridors
21.11 Summary
Chapter 22: Ecosystem management and conservation
22.1 Introduction
22.2 Definitions
22.3 Gradients of communities
22.4 Niches
22.5 Food webs and intertrophic interactions
22.6 Community features and management consequences
22.7 Multiple states
22.8 Regulation of top-down and bottom-up processes
22.9 Ecosystem consequences of bottom-up processes
22.10 Ecosystem disturbance and heterogeneity
22.11 Ecosystem management at multiple scales
22.12 Biodiversity
22.13 Island biogeography and dynamic processes of diversity
22.14 Ecosystem function
22.15 Summary
Appendices
Glossary
References
Index
End User License Agreement
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Cover
Table of Contents
Preface
Part 1: Wildlife ecology
Chapter 1: Introduction: goals and decisions
Figure 2.1
Figure 2.2
Figure 2.3
Figure 2.4
Figure 2.5
Figure 2.6
Figure 2.7
Figure 2.8
Figure 2.9
Figure 2.10
Figure 2.11
Figure 2.12
Figure 3.1
Figure 3.2
Figure 3.3
Figure 3.4
Figure 3.5
Figure 3.6
Figure 3.7
Figure 3.8
Figure 4.1
Figure 4.2
Figure 4.3
Figure 4.4
Figure 4.5
Figure 4.6
Figure 4.7
Figure 4.8
Figure 5.1
Figure 5.2
Figure 5.3
Figure 5.4
Figure 5.5
Figure 5.6
Figure 5.7
Figure 5.8
Figure 5.9
Figure 5.10
Figure 5.11
Figure 5.12
Figure 5.13
Figure 5.14
Figure 5.15
Figure 5.16
Figure 5.17
Figure 5.18
Figure 5.19
Figure 5.20
Figure 5.21
Figure 5.22
Figure 5.23
Figure 6.1
Figure 6.2
Figure 6.3
Figure 6.4
Figure 6.5
Figure 6.6
Figure 6.7
Figure 6.8
Figure 6.9
Figure 6.10
Figure 6.11
Figure 6.12
Figure 6.13
Figure 6.14
Figure 6.15
Figure 6.16
Figure 6.17
Figure 6.18
Figure 6.19
Figure 7.1
Figure 7.2
Figure 7.3
Figure 7.4
Figure 7.5
Figure 7.6
Figure 7.7
Figure 7.8
Figure 7.9
Figure 7.10
Figure 7.11
Figure 8.1
Figure 8.2
Figure 8.3
Figure 8.4
Figure 8.5
Figure 8.6
Figure 9.1
Figure 9.2
Figure 9.3
Figure 9.4
Figure 9.5
Figure 9.6
Figure 9.7
Figure 9.8
Figure 9.9
Figure 9.10
Figure 9.11
Figure 9.12
Figure 9.13
Figure 9.14
Figure 10.1
Figure 10.2
Figure 10.3
Figure 10.4
Figure 10.5
Figure 10.6
Figure 10.7
Figure 10.8
Figure 10.9
Figure 10.10
Figure 10.11
Figure 11.1
Figure 11.2
Figure 11.3
Figure 11.4
Figure 11.5
Figure 12.1
Figure 12.2
Figure 13.1
Figure 13.2
Figure 13.3
Figure 13.4
Figure 13.5
Figure 13.6
Figure 13.7
Figure 14.1
Figure 14.2
Figure 14.3
Figure 14.4
Figure 14.5
Figure 15.1
Figure 15.2
Figure 15.3
Figure 15.4
Figure 15.5
Figure 16.1
Figure 16.2
Figure 16.3
Figure 16.4
Figure 16.5
Figure 16.6
Figure 16.7
Figure 16.8
Figure 16.9
Figure 16.10
Figure 16.11
Figure 16.12
Figure 17.1
Figure 18.1
Figure 18.2
Figure 18.3
Figure 18.4
Figure 18.5
Figure 18.6
Figure 18.7
Figure 18.8
Figure 18.9
Figure 18.10
Figure 18.11
Figure 18.12
Figure 18.13
Figure 18.14
Figure 19.1
Figure 20.1
Figure 20.2
Figure 20.3
Figure 20.4
Figure 20.5
Figure 20.6
Figure 20.7
Figure 20.8
Figure 21.1
Figure 21.2
Figure 21.3
Figure 21.4
Figure 21.5
Figure 22.1
Figure 22.2
Figure 22.3
Figure 22.4
Table 1.1
Table 1.2
Table 1.3
Table 2.1
Table 5.1
Table 6.1
Table 6.2
Table 6.3
Table 7.1
Table 7.2
Table 8.1
Table 9.1
Table 12.1
Table 12.2
Table 12.3
Table 12.4
Table 13.1
Table 13.2
Table 13.3
Table 13.4
Table 14.1
Table 14.2
Table 14.3
Table 15.1
Table 15.2
Table 16.1
Table 16.2
Table 17.1
Table 17.2
Table 19.1
Table 19.2
Table 19.3
Table 20.1
To our colleagues Graeme Caughley, Jamie Smith, and Peter Yodzis, who have influenced both our approach to wildlife biology and the writing of this book.
Third Edition
John M. Fryxell PhD
Department of Integrative Biology, University of Guelph, Guelph, Canada
Anthony R. E. Sinclair PhD
Biodiversity Research Centre, University of British Columbia, Vancouver, Canada
The late Graeme Caughley PhD
CSIRO, Canberra, Australia
This edition first published 2014 © 2014 by John Wiley & Sons Ltd
First edition © 1994 by Blackwell Science; Second edition © 2006 by Anthony R. E. Sinclair and John M. Fryxell.
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Library of Congress Cataloging-in-Publication Data
Fryxell, John M., 1954–
Wildlife ecology, conservation, and management.—Third edition / John M. Fryxell, Anthony R.E. Sinclair, Graeme Caughley.
pages cm
Includes index.
ISBN 978-1-118-29106-1 (cloth)—ISBN 978-1-118-29107-8 (pbk.) 1. Wildlife management. 2. Wildlife conservation. 3. Animal ecology. I. Sinclair, A. R. E. (Anthony Ronald Entrican) II. Caughley, Graeme. III. Title.
SK355.C38 2014
639.9—dc23
2014013806
A catalogue record for this book is available from the British Library.
Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books.
Cover image © Luke Vander Vennen, used by permission.
Modern principles of sustainable management and conservation of wildlife species require a clear understanding of demography, animal behavior, and ecosystem dynamics. Our book weaves together these disparate elements in a single coherent text intended for senior undergraduate and graduate students. The first half provides a solid background in key ecological concepts such as demography, population growth and regulation, competition within and among species, and predator–prey interactions. The second half uses these key ecological concepts to develop a deeper understanding of the principles underlying wildlife management and conservation, including population viability assessment, sustainable harvesting, landscape planning, and ecosystem management.
New quantitative methods, developed over the last 10 years, are now so fundamental to management that we have included them at the most basic levels. Several chapters of the book will be useful to practicing wildlife managers. For example, we have included modern approaches to estimating animal abundance and habitat selectivity, the use of age- and stage-structured data in demography studies, and the use of models as efficient methods for making conservation and management decisions. As a study aid, we have included a wide variety of downloadable computer programs in R and Mathcad on an accompanying website. These are intended to help readers develop a solid understanding of key statistical procedures and population models commonly used in wildlife ecology and management.
In this edition we have arranged the sequence of chapters to reflect the progression from individuals to populations, communities, and ecosystems. Four new chapters have been added to cover rapidly developing topics: effects of climate change on wildlife, the evolutionary response by wildlife populations to rapidly changing conditions, home range use and habitat selection as a consequence of patterns of individual movement, and the importance of corridor use and metapopulation dynamics for wildlife populations living in the highly fragmented landscapes that increasingly characterize the modern world.
Anne Gunn and David Grice were invaluable in bringing together the first edition of this book after Graeme Caughley fell ill. Fleur Sheard prepared the line drawings for that edition. Since then we have continued to benefit from the helpful contributions of a number of people, including Tal Avgar, Andrew McAdam, Cort Griswold, David Grice, Sue Briggs, Andrea Byrom, Steve Cork, Charles Krebs, Graham Nugent, John Parkes, Roger Pech, Laura Prugh, Wendy Ruscoe, Dolph Schluter, Julian Seddon, Grant Singleton, David Spratt, Eric Spurr, Vernon Thomas, and Bruce Warburton. We also thank the Natural Sciences and Engineering Research Council of Canada for continuing support over the years.
Our close friend and colleague, Graeme Caughley, died in 1994. We have retained the substance and spirit of his scholarship, expanding the fields where advances have occurred since the first edition. For this new edition we are indebted to Sue Pennant and Anne Sinclair, who are always willing (if not necessarily eager) to provide a fresh set of eyes for proofreading of the new material.
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