Duncan J. Irschick, Ph.D. is an Assistant Professor in the Department of Biology at the University of Massachusetts at Amherst. His research interests are in the interface among organism design, function, and ecology. Broadly, he is interested in the evolution of complex functional systems in all its facets. His research integrates micro evolutionary and macro evolutionary approaches, and?applies both experimental and descriptive approaches to understand the causes of, and ultimately the consequences of, this diversity. Mark Briffa, Ph.D. is a Lecturer in Marine Biology at the University of Plymouth in the School of Biological Sciences. His main research focuses on contest behavior, where animals compete directly against one another over ownership of a limited resource. He is particularly interested in how animals use communication to resolve conflicts of interest — this often involves 'repeated signals' that are?performed in a structured way. Jeffrey Podos, Ph.D. is an Associate Professor at the University of Massachusetts in the Biology Department. His research addresses the interface of animal behavior, organism biology, and evolutionary biology. His work focuses on vocal behavior and evolution in vertebrates, especially songbirds.

“Though it covers a variety of topics (e.g., deception, biomechanics, condition) important for understanding this research area, the book is definitely a resource for specialists.  It will be useful only to those interested in the most current research on particular aspects of animal communication . . . Summing Up: Recommended. Graduate students, researchers”.  (Choice, 1 January 2015)

• Contributors ix1 INTRODUCTION 1Duncan J. Irschick, Mark Briffa, and Jeffrey PodosReferences 72 EARLY LIFE-HISTORY EFFECTS, OXIDATIVE STRESS, AND THE EVOLUTION AND EXPRESSION OF ANIMAL SIGNALS 11Nick J. Royle, Josephine M. Orledge, and Jonathan D. BlountIntroduction 11Signaling 12Early Life-History Effects and Resource Allocation Trade-Offs 13Oxidative Stress as a Mediator of Resource Allocation Trade-Offs 15Signals Expressed During Development 20Signals Expressed During Adulthood 25Competition-Dependent Sexual Signals 32Conclusions 34Acknowledgments 36References 363 A PERFORMANCE-BASED APPROACH TO STUDYING COSTS OF RELIABLE SIGNALS 47Jerry F. Husak, Justin P. Henningsen, Bieke Vanhooydonck, and Duncan J. IrschickIntroduction 47Receiver-Independent Costs 51Receiver-Dependent Costs 55Compensatory Traits 59Conclusions 63Acknowledgments 64References 654 COGNITIVELY DRIVEN CO-OPTION AND THE EVOLUTION OF COMPLEX SEXUAL DISPLAYS IN BOWERBIRDS 75Gerald Borgia and Jason KeagyIntroduction 75Cognition, Co-Option, and Complex Display 78Delayed Male Maturity, Male–Male Courtship, and Display TraitAcquisition 81Female Signaling to Affect Male Display Intensity: An Innovation that Improves Courtship Success 82Mate Searching and Flexibility in Adaptive Decision-Making 83Female Uncertainty and Flexibility in Active Mate Assessment 84Long-Term Age-Related Improvement in Decoration Display: Symmetrical Decoration Displays on Older Males’ Bowers 84Anticipation of Male Routes during Courtship: Paths on Display Courts of Spotted Bowerbirds 86Some Other Possible Cognitive Display-Related Behaviors of Bowerbirds 87Construction of Successive Scenes for Females Visiting the Bower 88Cognitive Aspects of Bower Building: Age-Related Improvement in Construction and Novel Techniques for Maintaining Symmetry 90Cognitive Flexibility and Innovation in Display 93Decoration Stealing: An Innovation for Display Trait Acquisition 94Cooperating with Relatives for Display: An Innovation to ReduceSexual Competition 95Vocal Mimicry: Learning and Innovation in Use of Co-Opted Displays 96Co-Option Mechanism 98Cognition in Display Trait Acquisition 100References 1015 INTEGRATING FUNCTIONAL AND EVOLUTIONARY APPROACHES TO THE STUDY OF COLOR-BASED ANIMAL SIGNALS 111Darrell J. Kemp and Gregory F. GretherIntroduction 111Color Signal Production in More Detail 115Signals, Honesty, and Condition-Dependence 116Coloration as an Honest Advertisement 117Trinidadian Guppies (Poecilia Reticulata) 118Pierid Butterflies (Subfamily Coliadinae) 122Birds 127Discussion/Conclusion/Future Work 129Acknowledgments 131References 1316 AGONISTIC SIGNALS: INTEGRATING ANALYSIS OF FUNCTIONS AND MECHANISMS 141Mark BriffaAnimal Contests and the Evolution of Agonistic Signals 141Empirical Approaches to Testing Theory: “Physiological Costs,” “Stamina,” and “Performance” 154Energy Status and Agonistic Signals 156Whole Body Performance and Agonistic Signals 159Conclusions 164References 1677 ACOUSTIC SIGNAL EVOLUTION: BIOMECHANICS, SIZE, AND PERFORMANCE 175Jeffrey Podos and S.N. PatekIntroduction 175Biomechanics 178Body Size 183Performance 187Concluding Remarks 194Acknowledgments 195References 1958 DISHONEST SIGNALING DURING AGGRESSIVE INTERACTIONS: THEORY AND EMPIRICAL EVIDENCE 205Robbie S. Wilson and Michael J. Angilletta Jr.Introduction 205The Evolution of Signaling 206The Theory of Dishonesty 208Dishonest Signaling in Aggressive Interactions between Conspecifics 209Conclusions 223References 2239 FUNCTIONAL APPROACH TO CONDITION 229Dustin J. Wilgers and Eileen A. HebetsIntroduction 229Practical Approaches to Condition 230Condition and Animal Performance 235Condition and Mate Choice 239Summary 241References 242Index 253

Duncan J. Irschick, Ph.D. is an Assistant Professor in the Department of Biology at the University of Massachusetts at Amherst. His research interests are in the interface among organism design, function, and ecology. Broadly, he is interested in the evolution of complex functional systems in all its facets. His research integrates micro evolutionary and macro evolutionary approaches, and?applies both experimental and descriptive approaches to understand the causes of, and ultimately the consequences of, this diversity. Mark Briffa, Ph.D. is a Lecturer in Marine Biology at the University of Plymouth in the School of Biological Sciences. His main research focuses on contest behavior, where animals compete directly against one another over ownership of a limited resource. He is particularly interested in how animals use communication to resolve conflicts of interest — this often involves 'repeated signals' that are?performed in a structured way. Jeffrey Podos, Ph.D. is an Associate Professor at the University of Massachusetts in the Biology Department. His research addresses the interface of animal behavior, organism biology, and evolutionary biology. His work focuses on vocal behavior and evolution in vertebrates, especially songbirds.

“Though it covers a variety of topics (e.g., deception, biomechanics, condition) important for understanding this research area, the book is definitely a resource for specialists.  It will be useful only to those interested in the most current research on particular aspects of animal communication . . . Summing Up: Recommended. Graduate students, researchers”.  (Choice, 1 January 2015)

• Contributors ix1 INTRODUCTION 1Duncan J. Irschick, Mark Briffa, and Jeffrey PodosReferences 72 EARLY LIFE-HISTORY EFFECTS, OXIDATIVE STRESS, AND THE EVOLUTION AND EXPRESSION OF ANIMAL SIGNALS 11Nick J. Royle, Josephine M. Orledge, and Jonathan D. BlountIntroduction 11Signaling 12Early Life-History Effects and Resource Allocation Trade-Offs 13Oxidative Stress as a Mediator of Resource Allocation Trade-Offs 15Signals Expressed During Development 20Signals Expressed During Adulthood 25Competition-Dependent Sexual Signals 32Conclusions 34Acknowledgments 36References 363 A PERFORMANCE-BASED APPROACH TO STUDYING COSTS OF RELIABLE SIGNALS 47Jerry F. Husak, Justin P. Henningsen, Bieke Vanhooydonck, and Duncan J. IrschickIntroduction 47Receiver-Independent Costs 51Receiver-Dependent Costs 55Compensatory Traits 59Conclusions 63Acknowledgments 64References 654 COGNITIVELY DRIVEN CO-OPTION AND THE EVOLUTION OF COMPLEX SEXUAL DISPLAYS IN BOWERBIRDS 75Gerald Borgia and Jason KeagyIntroduction 75Cognition, Co-Option, and Complex Display 78Delayed Male Maturity, Male–Male Courtship, and Display TraitAcquisition 81Female Signaling to Affect Male Display Intensity: An Innovation that Improves Courtship Success 82Mate Searching and Flexibility in Adaptive Decision-Making 83Female Uncertainty and Flexibility in Active Mate Assessment 84Long-Term Age-Related Improvement in Decoration Display: Symmetrical Decoration Displays on Older Males’ Bowers 84Anticipation of Male Routes during Courtship: Paths on Display Courts of Spotted Bowerbirds 86Some Other Possible Cognitive Display-Related Behaviors of Bowerbirds 87Construction of Successive Scenes for Females Visiting the Bower 88Cognitive Aspects of Bower Building: Age-Related Improvement in Construction and Novel Techniques for Maintaining Symmetry 90Cognitive Flexibility and Innovation in Display 93Decoration Stealing: An Innovation for Display Trait Acquisition 94Cooperating with Relatives for Display: An Innovation to ReduceSexual Competition 95Vocal Mimicry: Learning and Innovation in Use of Co-Opted Displays 96Co-Option Mechanism 98Cognition in Display Trait Acquisition 100References 1015 INTEGRATING FUNCTIONAL AND EVOLUTIONARY APPROACHES TO THE STUDY OF COLOR-BASED ANIMAL SIGNALS 111Darrell J. Kemp and Gregory F. GretherIntroduction 111Color Signal Production in More Detail 115Signals, Honesty, and Condition-Dependence 116Coloration as an Honest Advertisement 117Trinidadian Guppies (Poecilia Reticulata) 118Pierid Butterflies (Subfamily Coliadinae) 122Birds 127Discussion/Conclusion/Future Work 129Acknowledgments 131References 1316 AGONISTIC SIGNALS: INTEGRATING ANALYSIS OF FUNCTIONS AND MECHANISMS 141Mark BriffaAnimal Contests and the Evolution of Agonistic Signals 141Empirical Approaches to Testing Theory: “Physiological Costs,” “Stamina,” and “Performance” 154Energy Status and Agonistic Signals 156Whole Body Performance and Agonistic Signals 159Conclusions 164References 1677 ACOUSTIC SIGNAL EVOLUTION: BIOMECHANICS, SIZE, AND PERFORMANCE 175Jeffrey Podos and S.N. PatekIntroduction 175Biomechanics 178Body Size 183Performance 187Concluding Remarks 194Acknowledgments 195References 1958 DISHONEST SIGNALING DURING AGGRESSIVE INTERACTIONS: THEORY AND EMPIRICAL EVIDENCE 205Robbie S. Wilson and Michael J. Angilletta Jr.Introduction 205The Evolution of Signaling 206The Theory of Dishonesty 208Dishonest Signaling in Aggressive Interactions between Conspecifics 209Conclusions 223References 2239 FUNCTIONAL APPROACH TO CONDITION 229Dustin J. Wilgers and Eileen A. HebetsIntroduction 229Practical Approaches to Condition 230Condition and Animal Performance 235Condition and Mate Choice 239Summary 241References 242Index 253

Duncan J. Irschick, Ph.D. is an Assistant Professor in the Department of Biology at the University of Massachusetts at Amherst. His research interests are in the interface among organism design, function, and ecology. Broadly, he is interested in the evolution of complex functional systems in all its facets. His research integrates micro evolutionary and macro evolutionary approaches, and?applies both experimental and descriptive approaches to understand the causes of, and ultimately the consequences of, this diversity. Mark Briffa, Ph.D. is a Lecturer in Marine Biology at the University of Plymouth in the School of Biological Sciences. His main research focuses on contest behavior, where animals compete directly against one another over ownership of a limited resource. He is particularly interested in how animals use communication to resolve conflicts of interest — this often involves 'repeated signals' that are?performed in a structured way. Jeffrey Podos, Ph.D. is an Associate Professor at the University of Massachusetts in the Biology Department. His research addresses the interface of animal behavior, organism biology, and evolutionary biology. His work focuses on vocal behavior and evolution in vertebrates, especially songbirds.

“Though it covers a variety of topics (e.g., deception, biomechanics, condition) important for understanding this research area, the book is definitely a resource for specialists.  It will be useful only to those interested in the most current research on particular aspects of animal communication . . . Summing Up: Recommended. Graduate students, researchers”.  (Choice, 1 January 2015)

• Contributors ix1 INTRODUCTION 1Duncan J. Irschick, Mark Briffa, and Jeffrey PodosReferences 72 EARLY LIFE-HISTORY EFFECTS, OXIDATIVE STRESS, AND THE EVOLUTION AND EXPRESSION OF ANIMAL SIGNALS 11Nick J. Royle, Josephine M. Orledge, and Jonathan D. BlountIntroduction 11Signaling 12Early Life-History Effects and Resource Allocation Trade-Offs 13Oxidative Stress as a Mediator of Resource Allocation Trade-Offs 15Signals Expressed During Development 20Signals Expressed During Adulthood 25Competition-Dependent Sexual Signals 32Conclusions 34Acknowledgments 36References 363 A PERFORMANCE-BASED APPROACH TO STUDYING COSTS OF RELIABLE SIGNALS 47Jerry F. Husak, Justin P. Henningsen, Bieke Vanhooydonck, and Duncan J. IrschickIntroduction 47Receiver-Independent Costs 51Receiver-Dependent Costs 55Compensatory Traits 59Conclusions 63Acknowledgments 64References 654 COGNITIVELY DRIVEN CO-OPTION AND THE EVOLUTION OF COMPLEX SEXUAL DISPLAYS IN BOWERBIRDS 75Gerald Borgia and Jason KeagyIntroduction 75Cognition, Co-Option, and Complex Display 78Delayed Male Maturity, Male–Male Courtship, and Display TraitAcquisition 81Female Signaling to Affect Male Display Intensity: An Innovation that Improves Courtship Success 82Mate Searching and Flexibility in Adaptive Decision-Making 83Female Uncertainty and Flexibility in Active Mate Assessment 84Long-Term Age-Related Improvement in Decoration Display: Symmetrical Decoration Displays on Older Males’ Bowers 84Anticipation of Male Routes during Courtship: Paths on Display Courts of Spotted Bowerbirds 86Some Other Possible Cognitive Display-Related Behaviors of Bowerbirds 87Construction of Successive Scenes for Females Visiting the Bower 88Cognitive Aspects of Bower Building: Age-Related Improvement in Construction and Novel Techniques for Maintaining Symmetry 90Cognitive Flexibility and Innovation in Display 93Decoration Stealing: An Innovation for Display Trait Acquisition 94Cooperating with Relatives for Display: An Innovation to ReduceSexual Competition 95Vocal Mimicry: Learning and Innovation in Use of Co-Opted Displays 96Co-Option Mechanism 98Cognition in Display Trait Acquisition 100References 1015 INTEGRATING FUNCTIONAL AND EVOLUTIONARY APPROACHES TO THE STUDY OF COLOR-BASED ANIMAL SIGNALS 111Darrell J. Kemp and Gregory F. GretherIntroduction 111Color Signal Production in More Detail 115Signals, Honesty, and Condition-Dependence 116Coloration as an Honest Advertisement 117Trinidadian Guppies (Poecilia Reticulata) 118Pierid Butterflies (Subfamily Coliadinae) 122Birds 127Discussion/Conclusion/Future Work 129Acknowledgments 131References 1316 AGONISTIC SIGNALS: INTEGRATING ANALYSIS OF FUNCTIONS AND MECHANISMS 141Mark BriffaAnimal Contests and the Evolution of Agonistic Signals 141Empirical Approaches to Testing Theory: “Physiological Costs,” “Stamina,” and “Performance” 154Energy Status and Agonistic Signals 156Whole Body Performance and Agonistic Signals 159Conclusions 164References 1677 ACOUSTIC SIGNAL EVOLUTION: BIOMECHANICS, SIZE, AND PERFORMANCE 175Jeffrey Podos and S.N. PatekIntroduction 175Biomechanics 178Body Size 183Performance 187Concluding Remarks 194Acknowledgments 195References 1958 DISHONEST SIGNALING DURING AGGRESSIVE INTERACTIONS: THEORY AND EMPIRICAL EVIDENCE 205Robbie S. Wilson and Michael J. Angilletta Jr.Introduction 205The Evolution of Signaling 206The Theory of Dishonesty 208Dishonest Signaling in Aggressive Interactions between Conspecifics 209Conclusions 223References 2239 FUNCTIONAL APPROACH TO CONDITION 229Dustin J. Wilgers and Eileen A. HebetsIntroduction 229Practical Approaches to Condition 230Condition and Animal Performance 235Condition and Mate Choice 239Summary 241References 242Index 253