Table of Contents

List of Drawings at Part Openings ix Preface xi PART I Leks and Their Taxonomic Occurrence 1 1 What Are Leks? 3 1.1 Introduction 3 1.2 Definition of the Lek 4 1.3 Level of Aggregation: Exploded Leks 7 1.4 Aggregation on Landmarks 11 1.5 Resource Availability: Resource-based Leks 13 1.6 Handling Time 18 1.7 Stability of Arenas and Territories 19 1.8 Sexual Selection and Mating System 20 1.9 Summary 21 2 A Taxonomic Overview 23 2.1 Introduction 23 2.2 Basic Prerequisites for Lekking 24 2.3 Arthropods Other Than Insects 25 2.4 Insects 25 2.5 Fish 28 2.6 Amphibians 30 2.7 Salamanders and Newts 32 2.8 Reptiles 33 2.9 Birds 34 2.10 Ungulates and Other Mammals 39 2.11 Summary 40 PART II Sexual Selection 49 3 Determinants of Male Mating Success 51 3.1 Introduction 51 3.2 Mating Skew 52 3.3 Traits Beneficial for Males 54 3.4 Male-Male Competition or Female Choice? 78 3.5 Costs of Sexually Selected Traits 82 3.6 Lek Organization 84 3.7 Summary 90 4 Female Mating Adaptations 92 4.1 Introduction 92 4.2 Reasons for Female Preference 94 4.3 Costs of Female Choice 108 4.4 Female Mating Strategies 110 4.5 Summary 120 5 Black Grouse: A Case Study 122 5.1 Introduction 122 5.2 Methods of Observation 122 5.3 Correlates of Mating Success 124 5.4 Experiments 131 5.5 Synthesis 135 6 Comparative Studies 137 6.1 Introduction 137 6.2 Different Comparative Approaches 139 6.3 Social Selection 146 6.4 Summary 147 PART III Lek Evolution 149 7 A Review of Hypotheses 151 7.1 Introduction 151 7.2 Predation Risk 151 7.3 Information Sharing 156 7.4 Passive Attraction 156 7.5 Habitat Limitation 157 7.6 The Hotspot Model 159 7.7 Increased per Capita Mating Success on Larger Leks 163 7.8 The Black Hole Model 169 7.9 The Hotshot Model 170 7.10 Summary 173 8 Intraspecific Variation 175 8.1 Introduction 175 8.2 Intraspecific Variability 176 8.3 Intra-Individual Variability and Alternative Reproductive Behaviors 178 8.4 Summary 183 9 Game Theory Models of Leks 184 9.1 Introduction 184 9.2 Ideal Free Theory 184 9.3 Summary 195 PART IV Conclusions 199 10 Concluding Remarks and Prospects for Future Studies 201 10.1 General Conclusions 201 10.2 Future Prospects 203 10.3 A Final Word 207 References 209 Author Index 239 Subject and Species Index 246

Reviews

Leks are among the greatest wonders of the natural world. These aggregations of displaying male birds, mammals, and (by some definitions) insects, apparently existing solely as mating arenas where females come, mate with one or more highly popular males, and leave having obtained nothing more than sperm to fertilize their offspring, have captivated naturalists for centuries. Hoglund and Alatalo, experts in avian behavior and sexual selection, attempt to answer these questions and to place lekking systems in a broad context of sexual selection theory. Science