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G, Balance of Nature? Stuart L. Pimm Technisehe Unive^Hat Darmstadt FAC:IH;:RE;CH 10 — BIOL The University of Chicago Press Chicago and London — R ! [;• ! i o t h e k Schn;os::^:;nr;:ra6e 10 D-6 4 2 8 7 D a r m s t a d t Inv.-Nr. Contents Preface / xi - 1. WHY "THE BALANCE OF NATURE"? . 1 What Ecology Is About and What Ecologists Study / 1 The Problems of Scaling Up / 3 What Is the "Balance of Nature"? / 4 Beginnings: Elton and MacArthur / 6 The Last of the Old Meetings / 8 A Theoretical Basis / 9 Heresies / 12 Are Theoretical and Empirical Studies Incompatible? / 13 How This Book Is Organized / 15 2. RESILIENCE , 18 What Is It and How Do We Measure It? / 18 Resilience and the Life-History Characteristics of Individual Species / 21 Food-Web Effects on Resilience / 26 Ecosystem Effects on Resilience / 30 Summary and Conclusions / 32 Appendix / 33 A Note On Temporal Variability 35 3. TEMPORAL VARIABILITY AND THE INDIVIDUAL SPECIES 39 Introduction / 39 Body Size and Resistance / 40 Longevity / 43 Resilience and Temporal Variability: Models / 44 Resilience and Temporal Variability: Empirical Studies / 47 Other Factors / 53 Summary and Conclusions / 54 Appendix: The Measurement of Population Variability / 54 vi CONTENTS 4. THE EFFECTS OF FOOD-WEB STRUCTURE ' 59 The Ideas of MacArthur and Watt / 59 ^ Temporal Variability and the Diversity of Prey / 63 Temporal Variability and the Diversity of Predators / 71 Abundance: The Relative Contributions of Predator and Prey Diversity / 78 Summary and Conclusions / 80 5. THE VARIABILITY OF THE ENVIRONMENT , 83 Environmental Variability Across Space / 83 Environmental Change Over Time / 87 Summary and Conclusions / 97 6. NONLINEAR DYNAMICS, STRANGE ATTRACTORS, AND CHAOS 99 The Complexity of Population Change / 99 Where to Look for the Beast / 101 What to Expect: A Bestiary / 107 Meanwhile, in the Real World . . . / 115 Summary and Conclusions / 131 7. EXTINCTIONS 135 From Population Variability to Community Persistence / 135 The Effect of Population Size on Extinction / 135 The Effects of Individual Longevity and Population Resilience / 144 Temporal Variability in Density / 149 The Effects of Spatial and Demographic Structure / 151 Changes in Reproductive Rate at Low Density / 156 Feedback Cycles / 162 , Summary and Conclusions / 163^ A Note on Persistence 169 8. SPECIES DIFFERENCES AND COMMUNITY STRUCTURE AS EXPLANATIONS OF WHY INTRODUCTIONS FAIL The Effects of Individual Adaptations / 173 The Role of Population Parameters / 175 Community Structure and Species Assembly / 181 Summary / 187 173 Contents J ^ vii 9. PATTERNS IN SPECIES COMPOSITION — 189 Alternatives / 189 "Competition and the Structure of Ecological Communities" / 191 Rejecting Competition as an Explanation / 193 The Debate Continues / 195 Patterns as Hypotheses: A Catalog / 198 Experiments in Persistence: Introduced Species / 207 Persistence and Predation / 217 Summary / 220 10. FOOD-WEB STRUCTURE AND COMMUNITY PERSISTENCE 222 Does Pattern Indicate Persistence? / 222 Food-Chain Lengths / 223 Feeding on More than One Trophic Level / 225 Compartments / 227 Predator-Prey Ratios"/ 229 J Niche Geometry / 232 Cohen's Cascade Model / 242 Summary / 243 - 11. COMMUNITY ASSEMBLY; OR, WHY ARE THERE SO MANY KINDS OF COMMUNITIES? 245 Assembly and Persistence / 245 Models of Community Assembly / 245 Implications: Possible Topologies for Community Assembly / 249 Multispecies Simulations / 256 Empirical Studies / 258 Summary and Conclusions / 268 ^ A Note on Resistance ^ 271 12. SMALL-SCALE EXPERIMENTAL REMOVALS OF SPECIES Do Species Removals Have Effects? / 277 Four Hypotheses as to which Species can be Removed / 281 Looking Upward: Resistance to Changes in Lower Trophic Levels / 289 Summary / 292 277 viii CONTENTS 13. FOOD WEBS AND RESISTANCE 294 v A Food-Web Theory / 294 Indirect Effects in Removal Experiments / 301 Uncertainty in Removal Experiments / 303 The Time Scale of Removal Experiments / 306 Summary / 312 x 14. CHANGES IN TOTAL DENSITY AND SPECIES COMPOSITION 315 Resistance at the Community Level / 315 A Theory for Changes in, Species Composition / 315 A Theory for Changes in Total Density / 321 Testing the Theories / 328 Summary / 333 v 15. T H E CONSEQUENCES OF INTRODUCTIONS AND EXTINCTIONS i 334 i Resistance on a Large Scale / 334 Do Extinctions and Introductions Cause Further Changes in Species Composition? / 335 Theories of Resistance: Vacant Niches / 344 The Food-Web Theory of Resistance: Its Relevance To Species Introductions and Secondary Extinctions / 346 The Role of History / 353 A Comment about Habitat Islands / 355 Summary / 356 1 j j ! ; j i 16. MULTISPECIES M O D E L S AND T H E I R LIMITATIONS 357 Recipes: Getting Community Dynamics from Food-Web Diagrams / 361 17. CONCLUSIONS AND CAVEATS ^ Resilience/371 , Persistence / 378 Variability / 372 ~ Resistance / 382 Extinction / 376 The Difficulties: The Inevitable Approaches to Long-Term, Large-Scale, and Hence Multispecies Studies / 387 Literature Cited / 391 Species Index / 419 Subject Index / 426 j i ! 371 '< > I ; ] I