It is 14 years since the last edition and the science or sciences of animal behaviour have progressed enormously. Topics which justified only a brief mention in an introductory text then, for example sperm competition as a factor in mating systems, have prospered to require textbooks of their own. How then to approach a new edition which cannot be allowed to become significantly larger.
It is our conviction that an introduction to the whole field, or at least a substantial part of it, remains as important as ever. Discussing new areas of research is best done from a firm basis of the basic concepts and for us these are still embodied in Niko Tinbergen's 1963 'Four Questions for Ethology' - function, evolution, causation and development. So the plan of our book remains essentially that of the other editions. There has been some extensive rewriting and we have tried to give good coverage to those areas where there have been important advances, notably in the evolution of behaviour and its development. We continue to give extensive references so that readers can easily get into the literature of areas that catch their interest. There is a great deal of new literature to be explored but we have never cited new work unless it really adds something. Often concepts are best illustrated by some of the now classical papers.
This edition benefits enormously from the encouragement we have had from Cambridge University Press to illustrate more widely and, in particular, to use colour photographs. Animals behaving provide stunning images and we are delighted with many of the new figures here. It remains to acknowledge the great help and support we have had from Martin Griffiths - it was he who first persuaded us to undertake a new edition. His team at CUP have helped us at every stage and it has been a pleasure to work with them.

Preface page ix

1 Introduction 1

Questions about animal behaviour 5

The escaping cockroach 7

The courtship of the sage grouse 11

Units of the nervous system 16

Reflexes and more complex behaviour 20

Diversity and unity in the study of behaviour 30

Summary 31

2 The development of behaviour 33

Young animals grow up 34

Instinct and learning in their biological setting 40

The characteristics of instinct and learning 45

Genetics and behaviour 49

Development and changes to the nervous system 57

Hormones and early development 62

Early experience and the diversity of parental behaviour 68

Play 78

Imprinting 84

Bird song development 94

Conclusions 106

Summary 106

3 Stimuli and communication 109

What stimuli are and how they act 111

Diverse sensory capacities 114

The problem of pattern recognition 126

Sign stimuli (key features) 128

'Supernormal' stimuli 130

Neuroethological basis of sign stimuli 135

Other solutions for pattern recognition: generalized feature detection 139

Communication 144

What is communication? 145

Animal signals as effective stimuli 150

Honesty and deception in animal signalling 159

The honeybee dance 164

The calls of vervet monkeys 173

Summary 177

4 Decision-making and motivation 179

Decision-making on different time scales 182

Decision-making and 'motivation' 193

Measuring motivation 193

Is motivation specific or general? 199

Goals as decision points 202

Homeostasis and negative feedback 205

Competition between motivations 209

Inhibition/disinhibition 211

Decision-making with incomplete information: the role of signals 213

Conflict and 'abnormal' behaviour 217

The physiology of decision-making 220

Hormones and sequences of behaviour 227

Conflict and physiological stress 232

Decision-making, motivation and animal welfare 233

Conclusions 236

Summary 237

5 Learning and memory 239

Learning as part of adaptation 239

Sensitization and habituation 242

Associative learning 246

Specialized types of learning ability 257

What do animals actually learn? 262

Are there higher forms of learning in animals? 264

The comparative study of learning 268

Social learning and culture 272

The nature of animal minds 276

The nature of memory 294

Summary 298

6 Evolution 301

The adaptiveness of behaviour 302

Genes and behavioural evolution 310

Kin selection and inclusive fitness 316

Evolutionarily stable strategies 328

Sex and sexual selection 336

Species isolation and species selection 345

Tinbergen's fourth question: the phylogeny of behaviour 348

Summary 353

7 Social organization 355

The individual in the crowd 355

Advantages of grouping 358

Diverse social groups 364

Eusociality: division into castes 364

Territory in the social organization of vertebrates 369

Mating systems and social organization 373

Dominance in social systems 378

Diverse mammalian social behaviour 381

Primate social organization 388

Summary 403

References 405

Figure credits 435

Index 442

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