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Contents

Preface x Acknowledgements xiii 1 Natural Selection, Ecology and Behaviour 1 Watching and wondering 1 Natural selection 5 Genes and behaviour 6 Selfish individuals or group advantage? 11 Phenotypic plasticity: climate change and breeding times 18 Behaviour, ecology and evolution 21 Summary 22 Further reading 22 Topics for discussion 23 2 Testing Hypotheses in Behavioural Ecology 24 The comparative approach 25 Breeding behaviour of gulls in relation to predation risk 26 Social organization of weaver birds 28 Social organization in African ungulates 30 Limitations of early comparative studies 31 Comparative approach to primate ecology and behaviour 33 Using phylogenies in comparative analysis 37 The comparative approach reviewed 45 Experimental studies of adaptation 46 Summary 49 Further reading 50 Topics for discussion 51 3 Economic Decisions and the Individual 52 The economics of carrying a load 52 The economics of prey choice 59 Sampling and information 62 The risk of starvation 63 Environmental variability, body reserves and food storing 65 Food storing birds: from behavioural ecology to neuroscience 66 The evolution of cognition 71 Feeding and danger: a trade-off 73 Social learning 75 Optimality models and behaviour: an overview 79 Summary 81 Further reading 82 Topics for discussion 82 4 Predators versus Prey: Evolutionary Arms Races 83 Red Queen evolution 83 Predators versus cryptic prey 86 Enhancing camouflage 92 Warning colouration: aposematism 95 Mimicry 100 Trade-offs in prey defences 103 Cuckoos versus hosts 105 Summary 113 Further reading 114 Topics for discussion 114 5 Competing for Resources 116 The Hawk-Dove game 116 Competition by exploitation: the ideal free distribution 119 Competition by resource defence: the despotic distribution 123 The ideal free distribution with unequal competitors 123 The economics of resource defence 126 Producers and scroungers 130 Alternative mating strategies and tactics 131 ESS thinking 142 Animal personalities 143 Summary 144 Further reading 145 Topics for discussion 146 6 Living in Groups 147 How grouping can reduce predation 148 How grouping can improve foraging 159 Evolution of group living: shoaling in guppies 163 Group size and skew 164 Group decision making 169 Summary 177 Further reading 177 Topics for discussion 178 7 Sexual Selection, Sperm Competition and Sexual Conflict 179 Males and females 180 Parental investment and sexual competition 182 Why do females invest more in offspring care than do males? 184 Evidence for sexual selection 186 Why are females choosy? 189 Genetic benefits from female choice: two hypotheses 194 Testing the hypotheses for genetic benefits 196 Sexual selection in females and male choice 201 Sex differences in competition 204 Sperm competition 205 Constraints on mate choice and extra-pair matings 208 Sexual conflict 209 Sexual conflict: who wins? 216 Chase-away sexual selection 218 Summary 220 Further reading 221 Topics for discussion 221 8 Parental Care and Family Conflicts 223 Evolution of parental care 223 Parental investment: a parent's optimum 227 Varying care in relation to costs and benefits 229 Sexual conflict 232 Sibling rivalry and parent-offspring conflict: theory 238 Sibling rivalry: evidence 240 Parent-offspring conflict: evidence 243 Brood parasites 249 Summary 252 Further reading 252 Topics for discussion 253 9 Mating Systems 254 Mating systems with no male parental care 254 Mating systems with male parental care 264 A hierarchical approach to mating system diversity 279 Summary 280 Further reading 281 Topics for discussion 281 10 Sex Allocation 282 Fisher's theory of equal investment 285 Sex allocation when relatives interact 286 Sex allocation in variable environments 296 Selfish sex ratio distorters 304 Summary 305 Further reading 305 Topics for discussion 306 11 Social Behaviours: Altruism to Spite 307 Kin selection and inclusive fitness 308 Hamilton's rule 313 How do individuals recognize kin? 318 Kin selection doesn't need kin discrimination 322 Selfish restraint and kin selection 325 Spite 327 Summary 331 Further reading 332 Topics for discussion 333 12 Cooperation 334 What is cooperation? 334 Free riding and the problem of cooperation 336 Solving the problem of cooperation 337 Kin selection 339 Hidden benefits 341 By-product benefit 341 Reciprocity 345 Enforcement 350 A case study - the Seychelles Warbler 354 Manipulation 356 Summary 358 Further reading 358 Topics for discussion 359 13 Altruism and Conflict in the Social Insects 360 The social insects 360 The life cycle and natural history of a social insect 364 The economics of eusociality 366 The pathway to eusociality 366 The haplodiploidy hypothesis 367 The monogamy hypothesis 371 The ecological benefits of cooperation 375 Conflict within insect societies 379 Conflict over the sex ratio in the social hymenoptera 379 Worker policing in the social hymenoptera 386 Superorganisms 389 Comparison of vertebrates with insects 390 Summary 392 Further reading 392 Topics for discussion 393 14 Communication and Signals 394 The types of communication 395 The problem of signal reliability 396 Indices 397 Handicaps 405 Common interest 411 Human language 416 Dishonest signals 417 Summary 421 Further reading 422 Topics for discussion 423 15 Conclusion 424 How plausible are our main premises? 424 Causal and functional explanations 436 A final comment 438 Summary 441 Further reading 441 References 442 Index 489

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