Peter R. Grant and B. Rosemary Grant are professors emeriti at Princeton University. In recognition of their decades of work studying the ecology, behavior, genetics, and evolution of Darwin's finches, they were awarded the 2005 Balzan Prize and the 2009 Kyoto Prize.

• List of Illustrations xi List of Tables xv Preface xvii CHAPTER ONE: The Biodiversity Problem and Darwin's Finches 1 Biodiversity 1 The Choice of Organisms 2 Darwin's Finches 3 Diversity of Darwin's Finch Species 5 Species and Populations 8 Overview of the Book 11 CHAPTER TWO: Origins and History 13 Introduction 13 Phylogeny 14 Ancestors 16 The Time of Arrival 16 Colonization 17 The Ecological Theater 18 A Change of Scenery 21 The Evolutionary Play 22 Recent History 22 Summary 25 CHAPTER THREE: Modes of Speciation 26 The Formation of New Species 26 Two Groups from One 27 Divergence in Allopatry 28 Coexistence in Sympatry 29 Sympatric Speciation 30 Parapatric Speciation 31 Testing the Models 33 Summary 33 CHAPTER FOUR: Colonization of an Island 35 Speciation: The Initial Split 35 Establishment of a New Population 35 Founder Effects: Expectations from Theory 36 A Colonization Event 38 Inbreeding 39 Recurrent Immigration 39 An Alternative Phenology of Founder Effects 42 Conclusion 42 Species Elsewhere 44 Summary 45 CHAPTER FIVE: Natural Selection, Adaptation, and Evolution 46 Adaptation 46 Beak Sizes and Diets 47 Adaptive Evolution When the Environment Changes 50 Natural Selection 52 Evolution 54 Oscillating Directional Selection 55 Extrapolating from Short to Long Term 58 The Sources of Variation 58 How Beaks Are Formed 59 Depth and Width 59 Length 62 Summary 63 CHAPTER SIX: Ecological Interactions 65 Introduction 65 Competition 66 Patterns of Coexistence 66 Diets Inferred from Beaks 67 Interpreting the Patterns 67 Character Displacement and Release 68 Character Displacement Observed 68 The Competitive Role of G. Magnirostris 69 Selection under Contrasting Conditions 73 Evolution of Character Displacement 73 Summary 75 CHAPTER SEVEN: Reproductive Isolation 76 Pre-mating Barrier to Interbreeding 76 Factors Involved in the Discrimination between Species 76 Beaks 77 Song 77 Learning 79 Song Differences between Species 80 Song Divergence in Allopatry 81 Adaptation to Habitat 83 Change of Songs as a Consequence of Morphological Divergence 84 The Role of Chance 86 Simulating Secondary Contact 88 Summary 91 CHAPTER EIGHT: Hybridization 92 Introduction 92 Hybridization 92 Why Hybridization Occurs 93 When Hybridization Does Not Occur 96 Hybrid Fitness 97 Introgression on Daphne Major 100 Introgression in the Archipelago 103 Reinforcement 103 Reproductive Character Displacement 105 Evolutionary Significance of Introgression 106 Summary 107 CHAPTER NINE: Species and Speciation 108 Introduction 108 From Process to Product:What Is a Species? 109 A Working Definition 110 How Many Species of Darwin's Finches? 111 Certhidea olivacea: One Species or Two? 112 Geospiza difficilis: One Species or Three? 113 From Product Back to Process 114 Fission and Fusion 116 Summary 119 CHAPTER TEN: Reconstructing the Radiation of Darwin's Finches 120 Introduction 120 The Shape of the Radiation 121 Speciation and Extinction 123 Speciation 125 Extinction 126 Implications for Phylogeny 127 Adaptive Landscape 128 A Pattern of Ecological Segregation 133 Specialization 134 The Buildup of Complex Communities 134 Summary 135 CHAPTER ELEVEN: Facilitators of Adaptive Radiation 137 Introduction 137 Environmental Opportunity 138 Geographical Suitability 139 Ecological Opportunity 140 High Diversification Potential 142 Behavioral Flexibility 142 Introgressive Hybridization 145 Hybridization and Animal Breeding 146 Environmental Conditions Conducive to Introgression 146 Finches versus Mockingbirds 148 Summary 150 CHAPTER TWELVE: The Life History of Adaptive Radiations 152 Introduction 152 The First Stage of Adaptive Radiation 153 The Second Stage of Adaptive Radiation 154 Haldane's Rule 157 The Third Stage of Adaptive Radiation 158 Synthesis 160 Summary 162 CHAPTER THIRTEEN: Summary of the Darwin's Finch Radiation 163 What Happened and Why 163 What Is Missing? 165 Epilogue 166 Glossary 168 References 175 Author Index 201 Subject Index 210

"One of the most compelling documentations of the operation of natural selection. In this book, the Grants aim to capture the key insights provided by Darwin's finches into mechanisms of adaptation and speciation generally. They succeed in making a complex topic accessible without losing the excitement inherent in tackling a difficult problem. There is enough depth to stimulate serious students of evolutionary biology, enough explanation for general readers and an approachable style that will please both."--Roger Butlin, Times Higher Education "This Princeton-based couple presents their own accessible summary of their life's research in How and Why Species Multiply. The authors explain the scientific hypotheses involved...with admirable clarity...yet the book's real strength is not theory but data. The Grants' account makes exciting and lucid reading. Among those who should take note are doubters of 'old-fashioned' research methods who marvel at the prospects of genomics in the lab and wonder what use bird bands have in modern science."--Hanna Kokko, Science "Distilled into 200 pages, this is the life's work of two of evolutionary biology's greatest advocates, Peter and Rosemary Grant. In this book they meld insights from geography, behaviour, ecology and genetics to paint a complex but compelling picture of the evolutionary process. [A] must-have primer for any biology student."--Henry Nicholls, New Scientist "The authors' assertion that 'speciation is a process and not an event,' comes across clearly in this concise and accessible tale of 3 million years of finch evolution."--Science News "The book illustrates how laboratory work, particularly in developmental biology and molecular genetics, can be combined effectively with observations and experimental work in the field."--J. S. Schwartz, emeritus, CUNY College of Staten Island, for CHOICE "This book presents a succinct and most readable summary of one of the most important contemporary field experiments in evolution and adaptive radiation. It should be basic reading for any biologist."--Ghillean Prance, Biologist "[T]he book is authoritative, well prepared and edited ... and attractive. The Grants have provide and excellent third part for the Darwin's finch trilogy, and this volume should serve admirably as a summary of the knowledge that they have accumulated."--A. Townsend Peterson, Quarterly Review of Biology "How and Why Species Multiply is so impressive and such a stimulating read for two primary reasons, the first being the data presented throughout the work. Rarely do we have such detailed data on any natural system and the book draws great strength from this. The second reason is the commanding role given to ecological interactions in explaining the evolutionary dynamics of Darwin's finches."--Utku Perktas, Ibis "The book is valuable as a condensed version of the huge amount of fine work the authors have done on the finches. It should be accessible to scientists and informed lay audiences alike. The theory and ecological aspects are very compelling."--Robert M. Zink, Bioscience

Peter R. Grant and B. Rosemary Grant are professors emeriti at Princeton University. In recognition of their decades of work studying the ecology, behavior, genetics, and evolution of Darwin's finches, they were awarded the 2005 Balzan Prize and the 2009 Kyoto Prize.

• List of Illustrations xi List of Tables xv Preface xvii CHAPTER ONE: The Biodiversity Problem and Darwin's Finches 1 Biodiversity 1 The Choice of Organisms 2 Darwin's Finches 3 Diversity of Darwin's Finch Species 5 Species and Populations 8 Overview of the Book 11 CHAPTER TWO: Origins and History 13 Introduction 13 Phylogeny 14 Ancestors 16 The Time of Arrival 16 Colonization 17 The Ecological Theater 18 A Change of Scenery 21 The Evolutionary Play 22 Recent History 22 Summary 25 CHAPTER THREE: Modes of Speciation 26 The Formation of New Species 26 Two Groups from One 27 Divergence in Allopatry 28 Coexistence in Sympatry 29 Sympatric Speciation 30 Parapatric Speciation 31 Testing the Models 33 Summary 33 CHAPTER FOUR: Colonization of an Island 35 Speciation: The Initial Split 35 Establishment of a New Population 35 Founder Effects: Expectations from Theory 36 A Colonization Event 38 Inbreeding 39 Recurrent Immigration 39 An Alternative Phenology of Founder Effects 42 Conclusion 42 Species Elsewhere 44 Summary 45 CHAPTER FIVE: Natural Selection, Adaptation, and Evolution 46 Adaptation 46 Beak Sizes and Diets 47 Adaptive Evolution When the Environment Changes 50 Natural Selection 52 Evolution 54 Oscillating Directional Selection 55 Extrapolating from Short to Long Term 58 The Sources of Variation 58 How Beaks Are Formed 59 Depth and Width 59 Length 62 Summary 63 CHAPTER SIX: Ecological Interactions 65 Introduction 65 Competition 66 Patterns of Coexistence 66 Diets Inferred from Beaks 67 Interpreting the Patterns 67 Character Displacement and Release 68 Character Displacement Observed 68 The Competitive Role of G. Magnirostris 69 Selection under Contrasting Conditions 73 Evolution of Character Displacement 73 Summary 75 CHAPTER SEVEN: Reproductive Isolation 76 Pre-mating Barrier to Interbreeding 76 Factors Involved in the Discrimination between Species 76 Beaks 77 Song 77 Learning 79 Song Differences between Species 80 Song Divergence in Allopatry 81 Adaptation to Habitat 83 Change of Songs as a Consequence of Morphological Divergence 84 The Role of Chance 86 Simulating Secondary Contact 88 Summary 91 CHAPTER EIGHT: Hybridization 92 Introduction 92 Hybridization 92 Why Hybridization Occurs 93 When Hybridization Does Not Occur 96 Hybrid Fitness 97 Introgression on Daphne Major 100 Introgression in the Archipelago 103 Reinforcement 103 Reproductive Character Displacement 105 Evolutionary Significance of Introgression 106 Summary 107 CHAPTER NINE: Species and Speciation 108 Introduction 108 From Process to Product:What Is a Species? 109 A Working Definition 110 How Many Species of Darwin's Finches? 111 Certhidea olivacea: One Species or Two? 112 Geospiza difficilis: One Species or Three? 113 From Product Back to Process 114 Fission and Fusion 116 Summary 119 CHAPTER TEN: Reconstructing the Radiation of Darwin's Finches 120 Introduction 120 The Shape of the Radiation 121 Speciation and Extinction 123 Speciation 125 Extinction 126 Implications for Phylogeny 127 Adaptive Landscape 128 A Pattern of Ecological Segregation 133 Specialization 134 The Buildup of Complex Communities 134 Summary 135 CHAPTER ELEVEN: Facilitators of Adaptive Radiation 137 Introduction 137 Environmental Opportunity 138 Geographical Suitability 139 Ecological Opportunity 140 High Diversification Potential 142 Behavioral Flexibility 142 Introgressive Hybridization 145 Hybridization and Animal Breeding 146 Environmental Conditions Conducive to Introgression 146 Finches versus Mockingbirds 148 Summary 150 CHAPTER TWELVE: The Life History of Adaptive Radiations 152 Introduction 152 The First Stage of Adaptive Radiation 153 The Second Stage of Adaptive Radiation 154 Haldane's Rule 157 The Third Stage of Adaptive Radiation 158 Synthesis 160 Summary 162 CHAPTER THIRTEEN: Summary of the Darwin's Finch Radiation 163 What Happened and Why 163 What Is Missing? 165 Epilogue 166 Glossary 168 References 175 Author Index 201 Subject Index 210

"One of the most compelling documentations of the operation of natural selection. In this book, the Grants aim to capture the key insights provided by Darwin's finches into mechanisms of adaptation and speciation generally. They succeed in making a complex topic accessible without losing the excitement inherent in tackling a difficult problem. There is enough depth to stimulate serious students of evolutionary biology, enough explanation for general readers and an approachable style that will please both."--Roger Butlin, Times Higher Education "This Princeton-based couple presents their own accessible summary of their life's research in How and Why Species Multiply. The authors explain the scientific hypotheses involved...with admirable clarity...yet the book's real strength is not theory but data. The Grants' account makes exciting and lucid reading. Among those who should take note are doubters of 'old-fashioned' research methods who marvel at the prospects of genomics in the lab and wonder what use bird bands have in modern science."--Hanna Kokko, Science "Distilled into 200 pages, this is the life's work of two of evolutionary biology's greatest advocates, Peter and Rosemary Grant. In this book they meld insights from geography, behaviour, ecology and genetics to paint a complex but compelling picture of the evolutionary process. [A] must-have primer for any biology student."--Henry Nicholls, New Scientist "The authors' assertion that 'speciation is a process and not an event,' comes across clearly in this concise and accessible tale of 3 million years of finch evolution."--Science News "The book illustrates how laboratory work, particularly in developmental biology and molecular genetics, can be combined effectively with observations and experimental work in the field."--J. S. Schwartz, emeritus, CUNY College of Staten Island, for CHOICE "This book presents a succinct and most readable summary of one of the most important contemporary field experiments in evolution and adaptive radiation. It should be basic reading for any biologist."--Ghillean Prance, Biologist "[T]he book is authoritative, well prepared and edited ... and attractive. The Grants have provide and excellent third part for the Darwin's finch trilogy, and this volume should serve admirably as a summary of the knowledge that they have accumulated."--A. Townsend Peterson, Quarterly Review of Biology "How and Why Species Multiply is so impressive and such a stimulating read for two primary reasons, the first being the data presented throughout the work. Rarely do we have such detailed data on any natural system and the book draws great strength from this. The second reason is the commanding role given to ecological interactions in explaining the evolutionary dynamics of Darwin's finches."--Utku Perktas, Ibis "The book is valuable as a condensed version of the huge amount of fine work the authors have done on the finches. It should be accessible to scientists and informed lay audiences alike. The theory and ecological aspects are very compelling."--Robert M. Zink, Bioscience

Peter R. Grant and B. Rosemary Grant are professors emeriti at Princeton University. In recognition of their decades of work studying the ecology, behavior, genetics, and evolution of Darwin's finches, they were awarded the 2005 Balzan Prize and the 2009 Kyoto Prize.

• List of Illustrations xi List of Tables xv Preface xvii CHAPTER ONE: The Biodiversity Problem and Darwin's Finches 1 Biodiversity 1 The Choice of Organisms 2 Darwin's Finches 3 Diversity of Darwin's Finch Species 5 Species and Populations 8 Overview of the Book 11 CHAPTER TWO: Origins and History 13 Introduction 13 Phylogeny 14 Ancestors 16 The Time of Arrival 16 Colonization 17 The Ecological Theater 18 A Change of Scenery 21 The Evolutionary Play 22 Recent History 22 Summary 25 CHAPTER THREE: Modes of Speciation 26 The Formation of New Species 26 Two Groups from One 27 Divergence in Allopatry 28 Coexistence in Sympatry 29 Sympatric Speciation 30 Parapatric Speciation 31 Testing the Models 33 Summary 33 CHAPTER FOUR: Colonization of an Island 35 Speciation: The Initial Split 35 Establishment of a New Population 35 Founder Effects: Expectations from Theory 36 A Colonization Event 38 Inbreeding 39 Recurrent Immigration 39 An Alternative Phenology of Founder Effects 42 Conclusion 42 Species Elsewhere 44 Summary 45 CHAPTER FIVE: Natural Selection, Adaptation, and Evolution 46 Adaptation 46 Beak Sizes and Diets 47 Adaptive Evolution When the Environment Changes 50 Natural Selection 52 Evolution 54 Oscillating Directional Selection 55 Extrapolating from Short to Long Term 58 The Sources of Variation 58 How Beaks Are Formed 59 Depth and Width 59 Length 62 Summary 63 CHAPTER SIX: Ecological Interactions 65 Introduction 65 Competition 66 Patterns of Coexistence 66 Diets Inferred from Beaks 67 Interpreting the Patterns 67 Character Displacement and Release 68 Character Displacement Observed 68 The Competitive Role of G. Magnirostris 69 Selection under Contrasting Conditions 73 Evolution of Character Displacement 73 Summary 75 CHAPTER SEVEN: Reproductive Isolation 76 Pre-mating Barrier to Interbreeding 76 Factors Involved in the Discrimination between Species 76 Beaks 77 Song 77 Learning 79 Song Differences between Species 80 Song Divergence in Allopatry 81 Adaptation to Habitat 83 Change of Songs as a Consequence of Morphological Divergence 84 The Role of Chance 86 Simulating Secondary Contact 88 Summary 91 CHAPTER EIGHT: Hybridization 92 Introduction 92 Hybridization 92 Why Hybridization Occurs 93 When Hybridization Does Not Occur 96 Hybrid Fitness 97 Introgression on Daphne Major 100 Introgression in the Archipelago 103 Reinforcement 103 Reproductive Character Displacement 105 Evolutionary Significance of Introgression 106 Summary 107 CHAPTER NINE: Species and Speciation 108 Introduction 108 From Process to Product:What Is a Species? 109 A Working Definition 110 How Many Species of Darwin's Finches? 111 Certhidea olivacea: One Species or Two? 112 Geospiza difficilis: One Species or Three? 113 From Product Back to Process 114 Fission and Fusion 116 Summary 119 CHAPTER TEN: Reconstructing the Radiation of Darwin's Finches 120 Introduction 120 The Shape of the Radiation 121 Speciation and Extinction 123 Speciation 125 Extinction 126 Implications for Phylogeny 127 Adaptive Landscape 128 A Pattern of Ecological Segregation 133 Specialization 134 The Buildup of Complex Communities 134 Summary 135 CHAPTER ELEVEN: Facilitators of Adaptive Radiation 137 Introduction 137 Environmental Opportunity 138 Geographical Suitability 139 Ecological Opportunity 140 High Diversification Potential 142 Behavioral Flexibility 142 Introgressive Hybridization 145 Hybridization and Animal Breeding 146 Environmental Conditions Conducive to Introgression 146 Finches versus Mockingbirds 148 Summary 150 CHAPTER TWELVE: The Life History of Adaptive Radiations 152 Introduction 152 The First Stage of Adaptive Radiation 153 The Second Stage of Adaptive Radiation 154 Haldane's Rule 157 The Third Stage of Adaptive Radiation 158 Synthesis 160 Summary 162 CHAPTER THIRTEEN: Summary of the Darwin's Finch Radiation 163 What Happened and Why 163 What Is Missing? 165 Epilogue 166 Glossary 168 References 175 Author Index 201 Subject Index 210

"One of the most compelling documentations of the operation of natural selection. In this book, the Grants aim to capture the key insights provided by Darwin's finches into mechanisms of adaptation and speciation generally. They succeed in making a complex topic accessible without losing the excitement inherent in tackling a difficult problem. There is enough depth to stimulate serious students of evolutionary biology, enough explanation for general readers and an approachable style that will please both."--Roger Butlin, Times Higher Education "This Princeton-based couple presents their own accessible summary of their life's research in How and Why Species Multiply. The authors explain the scientific hypotheses involved...with admirable clarity...yet the book's real strength is not theory but data. The Grants' account makes exciting and lucid reading. Among those who should take note are doubters of 'old-fashioned' research methods who marvel at the prospects of genomics in the lab and wonder what use bird bands have in modern science."--Hanna Kokko, Science "Distilled into 200 pages, this is the life's work of two of evolutionary biology's greatest advocates, Peter and Rosemary Grant. In this book they meld insights from geography, behaviour, ecology and genetics to paint a complex but compelling picture of the evolutionary process. [A] must-have primer for any biology student."--Henry Nicholls, New Scientist "The authors' assertion that 'speciation is a process and not an event,' comes across clearly in this concise and accessible tale of 3 million years of finch evolution."--Science News "The book illustrates how laboratory work, particularly in developmental biology and molecular genetics, can be combined effectively with observations and experimental work in the field."--J. S. Schwartz, emeritus, CUNY College of Staten Island, for CHOICE "This book presents a succinct and most readable summary of one of the most important contemporary field experiments in evolution and adaptive radiation. It should be basic reading for any biologist."--Ghillean Prance, Biologist "[T]he book is authoritative, well prepared and edited ... and attractive. The Grants have provide and excellent third part for the Darwin's finch trilogy, and this volume should serve admirably as a summary of the knowledge that they have accumulated."--A. Townsend Peterson, Quarterly Review of Biology "How and Why Species Multiply is so impressive and such a stimulating read for two primary reasons, the first being the data presented throughout the work. Rarely do we have such detailed data on any natural system and the book draws great strength from this. The second reason is the commanding role given to ecological interactions in explaining the evolutionary dynamics of Darwin's finches."--Utku Perktas, Ibis "The book is valuable as a condensed version of the huge amount of fine work the authors have done on the finches. It should be accessible to scientists and informed lay audiences alike. The theory and ecological aspects are very compelling."--Robert M. Zink, Bioscience