Wildlife Ecology, Conservation, and Management

To understand modern principles of sustainable management and the conservation of wildlife species requires intimate knowledge about demography, animal behavior, and ecosystem dynamics. With emphasis on practical application and quantitative skill development, this book weaves together these disparate elements in a single
coherent textbook for senior undergraduate and graduate students. It reviews analytical techniques, explaining the mathematical and statistical principles behind them, and shows how these can be used to formulate
realistic objectives within an ecological framework. 

This third edition is comprehensive and up-to-date, and includes:

• 
  Brand new chapters that disseminate rapidly developing topics in the field: habitat use and selection; habitat fragmentation, movement, and corridors; population viability. analysis, the consequences of climate change; and evolutionary responses to disturbance
• 
  A thorough updating of all chapters to present important areas of wildlife research and management with recent developments and examples.
• 
  A new online study aid – a wide variety of downloadable computer programs in the freeware packages R and Mathcad, available through a companion website. Worked examples enable readers to practice calculations explained in the text and to develop a solid understanding of key statistical procedures and population models commonly used in wildlife ecology and management.

Professor John Fryxell currently teaches in the Department of Integrative Biology at the University of Guelph, Canada, where he has worked closely with a number of university and government scientists to develop sustainable conservation strategies for elk, woodland caribou, wolves, and marten. Previous to this he worked at the University of British Columbia and as Wildlife Consultant for the Provincial Government of Newfoundland and Labrador. His research has focused on the role of behavior in population and community dynamics of large mammals. He has a continuing interest in African wildlife, including long-term studies on the demography and spatial ecology of large herbivores and their predators in Serengeti National Park. 

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 6

1.5 Policy goals 7

1.6 Feasible options 7

1.7 Summary 8

Part 1 Wildlife ecology 9

2 Food and nutrition 11

2.1 Introduction 11

2.2 Constituents of food 11

2.3 Variation in food supply 14

2.4 Measurement of food supply 17

2.5 Basal metabolic rate and food requirement 20

2.6 Morphology of herbivore digestion 23

2.7 Food passage rate and food requirement 26

2.8 Body size and diet selection 27

2.9 Indices of body condition 28

2.10 Summary 33

3 Home range and habitat use 35

3.1 Introduction 35

3.2 Estimating home range size and utilization frequency 36

3.3 Estimating habitat availability and use 38

3.4 Selective habitat use 40

3.5 Using resource selection functions to predict population response 42

3.6 Sources of variation in habitat use 42

3.7 Movement within the home range 45

3.8 Movement among home ranges 48

3.9 Summary 51

4 Dispersal, dispersion, and distribution 53

4.1 Introduction 53

4.2 Dispersal 53

4.3 Dispersion 55

4.4 Distribution 56

4.5 Distribution, abundance, and range collapse 61

4.6 Species reintroductions or invasions 62

4.7 Summary 67

5 Population growth and regulation 69

5.1 Introduction 69

5.2 Rate of increase 69

5.3 Geometric or exponential population growth 73

5.4 Stability of populations 73

5.5 The theory of population limitation and regulation 76

5.6 Evidence for regulation 81

5.7 Applications of regulation 85

5.8 Logistic model of population regulation 86

5.9 Stability, cycles, and chaos 88

5.10 Intraspecific competition 90

5.11 Interactions of food, predators, and disease 93

5.12 Summary 93

6 Competition and facilitation between species 95

6.1 Introduction 95

6.2 Theoretical aspects of interspecific competition 96

6.3 Experimental demonstrations of competition 98

6.4 The concept of the niche 103

6.5 The competitive exclusion principle 106

6.6 Resource partitioning and habitat selection 106

6.7 Competition in variable environments 113

6.8 Apparent competition 113

6.9 Facilitation 114

6.10 Applied aspects of competition 119

6.11 Summary 122

7 Predation 123

7.1 Introduction 123

7.2 Predation and management 123

7.3 Definitions 123

7.4 The effect of predators on prey density 124

7.5 The behavior of predators 125

7.6 Numerical response of predators to prey density 129

7.7 The total response 130

7.8 Behavior of the prey 136

7.9 Summary 138

8 Parasites and pathogens 139

8.1 Introduction and definitions 139

8.2 Effects of parasites 139

8.3 The basic parameters of epidemiology 140

8.4 Determinants of spread 143

8.5 Endemic pathogens 144

8.6 Endemic pathogens: synergistic interactions with food and predators 144

8.7 Epizootic diseases 146

8.8 Emerging infectious diseases of wildlife 147

8.9 Parasites and the regulation of host populations 150

8.10 Parasites and host communities 151

8.11 Parasites and conservation 152

8.12 Parasites and control of pests 155

8.13 Summary 156

9 Consumer–resource dynamics 157

9.1 Introduction 157

9.2 Quality and quantity of a resource 157

9.3 Kinds of resource 157

9.4 Consumer–resource dynamics: general theory 158

9.5 Kangaroos and their food plants in semi-arid Australian savannas 161

9.6 Wolf–moose–woody plant dynamics in the boreal forest 167

9.7 Other population cycles 172

9.8 Summary 175

10 The ecology of behavior 177

10.1 Introduction 17710.2 Diet selection 177

10.3 Optimal patch or habitat use 183

10.4 Risk-sensitive habitat use 186

10.5 Social behavior and foraging 187

10.6 Summary 190

11 Climate change and wildlife 191

11.1 Introduction 191

11.2 Evidence for climate change 191

11.3 Wildlife responses to climate change 192

11.4 Mechanisms of response to climate change 196

11.5 Complex ecosystem responses to climate change 199

11.6 Summary 201

Part 2 Wildlife conservation and management 203

12 Counting animals 205

12.1 Introduction 205

12.2 Total counts 205

12.3 Sampled counts: the logic 207

12.4 Sampled counts: methods and arithmetic 212

12.5 Indirect estimates of population size 220

12.6 Indices 227

12.7 Harvest-based population estimates 228

12.8 Summary 231

13 Age and stage structure 233

13.1 Introduction 233

13.2 Demographic rates 233

13.3 Direct estimation of life table parameters 235

13.4 Indirect estimation of life table parameters 236

13.5 Relationships among parameters 238

13.6 Age-specific population models 239

13.7 Elasticity of matrix models 242

13.8 Stage-specific models 243

13.9 Elasticity of the loggerhead turtle model 245

13.10 Short-term changes in structured populations 246

13.11 Environmental stochasticity and age-structured populations 246

13.12 Summary 249

14 Experimental management 251

14.1 Introduction 251

14.2 Differentiating success from failure 251

14.3 Technical judgments can be tested 252

14.4 The nature of the evidence 255

14.5 Experimental and survey design 257

14.6 Some standard analyses 262

14.7 Summary 271

15 Model evaluation and adaptive management 273

15.1 Introduction 273

15.2 Fitting models to data and estimation of parameters 274

15.3 Measuring the likelihood of the observed data 276

15.4 Evaluating the likelihood of alternate models using AIC 278

15.5 Adaptive management 281

15.6 Summary 284

16 Population viability analysis 285

16.1 Introduction 285

16.2 Environmental stochasticity 285

16.3 PVA based on the exponential growth model 286

16.4 PVA based on the diffusion model 287

16.5 PVA based on logistic growth 290

16.6 Demographic stochasticity 291

16.7 Estimating both environmental and demographic stochasticity 294

16.8 PVA based on demographic and environmental stochasticity 296

16.9 Strengths and weaknesses of PVA 296

16.10 Extinction caused by environmental change 298

16.11 Extinction threat due to introduction of exotic predators or competitors 298

16.12 Extinction threat due to unsustainable harvesting 300

16.13 Extinction threat due to habitat loss 302

16.14 Summary 302

17 Conservation in practice 305

17.1 Introduction 305

17.2 How populations go extinct 305

17.3 How to prevent extinction 315

17.4 Rescue and recovery of near-extinctions 316

17.5 Conservation in National Parks and reserves 317

17.6 Community conservation outside National Parks and reserves 322

17.7 International conservation 323

17.8 Summary 324

18 Wildlife harvesting 325

18.1 Introduction 325

18.2 Fixed-quota harvesting strategy 325

18.3 Fixed-proportion harvesting strategy 329

18.4 Harvesting in practice: dynamic variation in quotas or effort 332

18.5 No-harvest reserves 334

18.6 Age- or sex-biased harvesting 335

18.7 Commercial harvesting 340

18.8 Bioeconomics 340

18.9 Game cropping and the discount rate 344

18.10 Summary 346

19 Wildlife control 347

19.1 Introduction 347

19.2 Definitions 347

19.3 Effects of control 348

19.4 Objectives of control 348

19.5 Determining whether control is appropriate 349

19.6 Methods of control 350

19.7 Summary 356

20 Evolution and conservation genetics 357

20.1 Introduction 357

20.2 Maintenance of genetic variation 358

20.3 Natural selection 359

20.4 Natural selection and life history tradeoffs 361

20.5 Natural selection due to hunting 363

20.6 Natural selection due to fishing 365

20.7 Selection due to environmental change 367

20.8 Ecological dynamics due to evolutionary changes 372

20.9 Heterozygosity 374

20.10 Genetic drift and mutation 375

20.11 Inbreeding depression 376

20.12 How much genetic variation is needed? 377

20.13 Effective population size 378

20.14 Effect of sex ratio 379

20.15 How small is too small? 380

20.16 Summary 380

21 Habitat loss and metapopulation dynamics 381

21.1 Introduction 381

21.2 Habitat loss and fragmentation 381

21.3 Ecological effects of habitat loss 384

21.4 Metapopulation dynamics 386

21.5 Territorial metapopulations 389

21.6 Mainland–island metapopulations 390

21.7 Source–sink metapopulations 391

21.8 Metacommunity dynamics of competitors 392

21.9 Metacommunity dynamics of predators and prey 393

21.10 Corridors 394

21.11 Summary 398

22 Ecosystem management and conservation 399

22.1 Introduction 399

22.2 Definitions 400

22.3 Gradients of communities 400

22.4 Niches 400

22.5 Food webs and intertrophic interactions 400

22.6 Community features and management consequences 402

22.7 Multiple states 404

22.8 Regulation of top-down and bottom-up processes 405

22.9 Ecosystem consequences of bottom-up processes 407

22.10 Ecosystem disturbance and heterogeneity 408

22.11 Ecosystem management at multiple scales 410

22.12 Biodiversity 411

22.13 Island biogeography and dynamic processes of diversity 413

22.14 Ecosystem function 415

22.15 Summary 417

Appendices 419

Glossary 423

References 435

Index 489

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