Introduction to Population Ecology

Introduction to Population Ecology, 2nd Edition is a comprehensive textbook covering all aspects of population ecology.  It uses a wide variety of field and laboratory examples, botanical to zoological, from the tropics to the tundra, to illustrate the fundamental laws of population ecology. Controversies in population ecology are brought fully up to date in this edition, with many brand new and revised examples and data.

Each chapter provides an overview of how population theory has developed, followed by descriptions of laboratory and field studies that have been inspired by the theory. Topics explored include single-species population growth and self-limitation, life histories, metapopulations and a wide range of interspecific interactions including competition, mutualism, parasite-host, predator-prey and plant-herbivore. An additional final chapter, new for the second edition, considers multi-trophic and other complex interactions among species. 

Throughout the book, the mathematics involved is explained with a step-by-step approach, and graphs and other
visual aids are used to present a clear illustration of how the models work. Such features make this an accessible introduction to population ecology; essential reading for undergraduate and graduate students taking courses in population ecology, applied ecology, conservation ecology, and conservation biology, including those with little mathematical experience.

Larry Rockwood is Professor of Biology and Environmental Science and Policy, and Chairman of the Department of Biology at George Mason University in Fairfax, Virginia, USA.  He earned his B.S. degree in Biopsychology and his Ph.D. degree in Biology at the University of Chicago.  His early research was conducted in Costa Rica where he studied foraging patterns in leaf-cutting ants.  More recently he has collaborated on a variety of projects from human-coyote conflicts to aspects of avian ecology and plant ecology.  He has been teaching introductory ecology, population ecology and tropical ecology for almost 40 years. In 2014 Dr Rockwood was presented with the David J. King Award in recognition of “outstanding contributions to enhancing teaching and learning” by George Mason University.

Preface ix

Acknowledgments xi

About the companion website xiii

Part 1 Single species populations 1

1 Density independent growth 5

1.1 Introduction 5

1.2 Fundamentals of population growth 8

1.3 Types of models 10

1.4 Density independent versus density dependent growth 12

1.5 Discrete or "geometric" growth in populations with non-overlapping generations 12

1.6 Exponential growth in populations with overlapping generations 16

1.7 Examples of exponential growth 18

1.8 Applications to human populations 19

1.9 The finite rate of increase (λ) and the intrinsic rate of increase (τ) 23

1.10 Stochastic models of population growth and population viability analysis 25

1.11 Conclusions 30

References 30

2 Density dependent growth and intraspecific competition 33

2.1 Introduction 33

2.2 Density dependence in populations with discrete generations 37

2.3 Density dependence in populations with overlapping generations 42

2.4 Nonlinear density dependence of birth and death rates and the Allee effect 46

2.5 Time lags and limit cycles 51

2.6 Chaos and behavior of the discrete logistic model 53

2.7 Adding stochasticity to density dependent models 54

2.8 Laboratory and field data 55

2.9 Behavioral aspects of intraspecific competition 60

2.10 Summary 64

References 64

3 Population regulation 69

3.1 Introduction 69

3.2 What is population regulation? 70

3.3 Combining density-dependent and density-independent factors 71

3.4 Tests of density dependence 73

3.5 Summary 77

References 78

4 Populations with age structures 81

4.1 Introduction 81

4.2 Survivorship 83

4.3 Fertility 90

4.4 Mortality curves 94

4.5 Expectation of life 96

4.6 Net reproductive rate, generation time, and the intrinsic rate of increase 97

4.7 Age structure and the stable age distribution 99

4.8 Projecting population growth in age-structured populations 99

4.9 The Leslie or population projection matrix 102

4.10 A second version of the Leslie matrix 103

4.11 The Lefkovitch modification of the Leslie matrix 104

4.12 Dominant latent roots and the characteristic equation 105

4.13 Reproductive value 107

4.14 Conclusions: sensitivity analysis 109

References 112

5 Metapopulation ecology 115

5.1 Introduction 115

5.2 Metapopulations and spatial ecology 116

5.3 MacArthur and Wilson and the equilibrium theory 120

5.4 The Levins or classical metapopulation 124

5.5 Lande's extension of the Levins model 125

5.6 Extinction in metapopulations 127

5.7 Metapopulation dynamics of two local populations 127

5.8 Source-sink metapopulations and the rescue effect 129

5.9 Nonequilibrium and patchy metapopulations 130

5.10 Spatially realistic models 130

5.11 Assumptions and evidence for the existence of metapopulations in nature 135

5.12 Summary 138

References 139

6 Life history strategies 145

6.1 Introduction 145

6.2 Power laws 149

6.3 The metabolic theory of ecology 152

6.4 Cole and Lewontin 154

6.5 The theory of τ- and κ-selection versus fast and slow life histories 159

6.6 Cost of reproduction and allocation of energy 162

6.7 Clutch size 163

6.8 Latitudinal gradients in clutch size 164

6.9 The effects of predation and disease on life history characteristics 165

6.10 Bet-hedging 166

6.11 The Grime general model for three evolutionary strategies in plants 166

6.12 Summary 168

References 168

Part 2 Interspecific interactions among populations 173

7 Interspecific competition 177

7.1 Introduction 177

7.2 Interspecific competition: early experiments and the competitive exclusion principle 178

7.3 The Lotka–Volterra competition equations 180

7.4 Laboratory experiments and competition 186

7.5 Resource-based competition theory 187

7.6 Spatial competition and the competition-colonization trade-off 194

7.7 Evidence for competition from nature 196

7.8 Indirect evidence for competition and 'natural experiments' 198

7.9 Summary 205

References 205

8 Mutualism 209

8.1 Introduction 209

8.2 Ant–plant mutualisms 210

8.3 Modeling mutualism 215

8.4 Summary: the costs of mutualism 217

References 217

9 Host–parasite interactions 221

9.1 Introduction 221

9.2 Factors affecting microparasite population biology 223

9.3 Modeling host–microparasite interactions 224

9.4 Dynamics of the disease 226

9.5 Immunization 229

9.6 Endangered metapopulations and disease 230

9.7 Social parasites 232

9.8 Summary 235

References 235

10 Predator/prey interactions 239

10.1 Introduction 239

10.2 The Lotka-Volterra equations 248

10.3 Early tests of the Lotka–Volterra models 250

10.4 Functional responses 252

10.5 Adding prey density dependence and the type II and III functional responses to the Lotka-Volterra equations 256

10.6 The graphical analyses of Rosenzweig and MacArthur 258

10.7 Use of a half saturation constant in predator/prey interactions 262

10.8 Parasitoid/host interactions and the Nicholson–Bailey models 264

10.9 Section summary 267

10.10 Field studies 268

10.11 The dangers of a predatory lifestyle 277

10.12 Escape from predation 277

10.13 Summary 281

References 282

11 Plant–herbivore interactions 287

11.1 Introduction 287

11.2 Classes of chemical defenses 289

11.3 Constitutive versus Induced Defense 294

11.4 Plant communication 296

11.5 Novel defenses/herbivore responses 296

11.6 Detoxification of plant compounds by herbivores 297

11.7 Plant apparency and chemical defense 298

11.8 Soil fertility and chemical defense 299

11.9 Modeling plant–herbivore population dynamics 299

11.10 Summary: the complexities of herbivore–plant interactions 303

References 306

12 Multi-trophic interactions 311
Jonathan Witt

12.1 Introduction 311

12.2 Trophic cascades 312

12.3 Trophic cascades and antropogenic change 317

12.4 Intraguild predation 319

12.5 Intraguild predation and prey suppression 321

12.6 Intraguild predation and mesopredator release 322

12.7 Cannibalism 323

References 326

Appendix 1: Problem sets 333

Appendix 2: Matrix algebra: the basics 337

Appendix 3: List of mathematical symbols used in this book 343

Index 351

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