BIOLOGY Ecology, Genetics, and Evolution

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1 Met a population BIOLOGY Ecology, Genetics, and Evolution Edited by Ilkka Hanski Department of Ecology and Systematics University of Helsinki FIN Helsinki, Finland Michael L Gilpin Department of Biology University of California, San Diego La Jolla, California ACADEMIC PRESS San Diego London Boston New York Sydney Tokyo Toronto Hi

2 Contributors Preface xv xi P A R T CONCEPTUAL FOUNDATIONS 1 The Metapopulation Approach, Its History, Conceptual Domain, and Application to Conservation Ilkka Hanski and Daniel Simberloff I. Introduction 5 II. Brief History of Metapopulation Studies 7 III. Conceptual Domain and Metapopulation Approaches 9 IV. Metapopulations and Conservation Biology 16 V. Conclusions 26

3 vi 2 Empirical Evidence for Metapopulation Dynamics Susan Harrison and Andrew D. Taylor I. Introduction 27 II. Single-Species Metapopulations 30 III. Multispecies Metapopulations 35 IV. Discussion 39 3 Metapopulation Dynamics and Landscape Ecology John A. Wiens I. Introduction 43 II. Approaches to Patchiness 44 III. What Is Landscape Ecology? 45 IV. How Is Landscape Ecology Relevant to Metapopulation Dynamics? 48 V. Metapopulations, Landscapes, and Conservation 58 VI. Conclusions 60 P A R T METAPOPULATION THEORY 4 Metapopulation Dynamics: From Concepts and Observations to Predictive Models Ilkka Hanski I. Introduction 69 II. An Example of Classical Metapopulation Dynamics with Rampant Population Turnover 70 III. Classical Metapopulation Dynamics: The Levins Model 73 IV. Minimum Viable Metapopulation Size 76 V. Predictive Models of Metapopulation Dynamics 78 VI. Nonequilibrium Metapopulations 85 VII. Four Conservation Messages 88 VIII. Concluding Remarks 91 5 Structured Metapopulation Models Mats Gyllenberg, Ilkka Hanski, and Alan Hastings I. Why Use Structured Models? 93 II. Modeling Structured Metapopulations 96

4 vii III. Steady States and Metapopulation Extinction 100 IV. Examples 101 V. Discussion 107 VI. The Mathematical Formalism Two-Species Metapopulation Models Sean Nee, Robert M. May, and Michael P. Hassell I. Introduction 123 II. Two-Species Patch Models 127 III. Spatially Explicit Metapopulations 137 IV. Conclusion From Metapopulation Dynamics to Community Structure: Some Consequences of Spatial Heterogeneity Robert D. Holt I. Introduction 149 II. Effects of Landscape Heterogeneity on Local Community Composition 150 III. Metapopulation Dynamics of Food Chains 155 IV. Apparent Competition in Metacommunities 161 V. Conclusions Genetic Effective Size of a Metapopulation Philip W. Hedrkk and Michael E. Gilpin I. An Example 166 II. Methods 167 III. Results 171 IV. Conclusions The Evolution of Metapopulations H. H. Barton and Michael C. Whitlock I. Introduction 183 II. Neutral Variation in Metapopulations 185 III. Selection in Metapopulations 189 IV. Adaptation to Local Conditions 193 V. Speciation and the "Shifting Balance" 200 VI. Discussion 209

5 viii P A R T» METAPOPULATION PROCESSES 10 Extinction Models for Local Populations Patrick Foley I. Introduction 215 II. Local Populations Are Vulnerable to Extinction due to Demographic and Environmental Stochasticity 217 III. Environmental Stochasticity 220 IV. Models with Demographic and Environmental Stochasticity 228 V. Robustness of the Model 232 VI. Catastrophies and Genetic Impoverishment 235 VII. Implications for Metapopulation Dynamics 237 VIII. Conclusion Studying Transfer Processes in Metapopulations: Emigration, Migration, and Colonization Rolf A. Ims and Nigel G. Yoccoz I. Introduction 247 II. Transfer of Individuals in Metapopulations: Defining the Components 248 III. Indirect, Global Approaches to Parameter Estimation 251 IV. Direct, Patch-Specific Approaches 254 V. Experimental Approaches to Studying Transfer Rates in Metapopulations 262 VI. Conclusion Migration within Metapopulations: The Impact upon Local Population Dynamics Peter B. Stacey, Veronica A. Johnson, and Mark L Taper I. Introduction 267 II. The Effect of Stochastic Variation in Migration Rate on Metapopulation Dynamics 271 III. Empirical Evidence 279 IV. Conclusions 289

6 ix 13 Evolution of Migration Rate and Other Traits: The Metapopulation Effect Isabelle Olivieri and Pierre-Henri Gouyon I. Introduction 293 II. How Does a Metapopulation Persist in a Landscape? An Example of a Model with Local Disturbances and Successional Processes 295 III. Within-Population Selection versus Colonization Selection: An Insight into the Metapopulation Effect 298 IV. Does Selection Adjust Migration Rate at the Metapopulation Level? 301 V. The Metapopulation Effect: Other Traits 311 VI. Influence of Migration on the Evolution of Life-History Traits 320 VII. Conclusion 321' 14 Spatial Processes in Host-Parasite Genetics Steven A. Frank I. Introduction 325 II. Dimensionality and Colonization Extinction Dynamics 326 III. Introduction to the Examples 332 IV. Plant-Pathogen Interactions 333 V. Cytoplasmic Male Sterility 339 VI. Other Systems of High Dimension 347 VII. Theories and Tests 350 P A R T 15 Butterfly Metapopulations Chris D. Thomas and Ilkka Hanski I. Introduction 359 II. Consequences of Landscape Structure on Population Structure 360 III. Population Turnover in Butterflies 364 IV. Theoretical Predictions Tested 368

7 V. Adding Realism to the Metapopulation Concept 373 VI. Conclusions Tritrophic Metapopulation Dynamics: A Case Study of Ragwort, the Cinnabar Moth, and the Parasitoid Cotesia populous Ed van der Meijden and Catharina A. M. van der Veen-van Wijk I. Introduction 387 II. Materials and Methods 389 III. Population Dynamics of Ragwort 391 IV. Mechanisms of Persistence in the Ragwort 395 V. Population Dynamics of the Cinnabar Moth and the Parasitoid Cotesia popularis 396 VI. Mechanisms of Persistence in the Cinnabar Moth 401 VII. Discussion, Spatially Correlated Dynamics in a Pika Metapopulation Andrew T. Smith and Michael E. Gilpin I. Introduction 407 II. Relevant Pika Natural History 409 III. The Bodie Site 411 IV. Methods 412 V. Results 415 VI. Discussion A Case Study of Genetic Structure in a Plant Metapopulation Barbara E. Giles and Jerome Goudet I. Introduction 429 II. Theory 430 III. The Silene Metapopulation on Islands in the Baltic 434 IV. Patterns of Genetic Structure on the Islands 439 V. Other Genetic Metapopulation Studies 449 Appendix: Information About the Studied Islands 453 Bibliography 455 Index 505

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