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Charles Darwin's experiences in the Galápagos Islands in 1835 helped to guide his thoughts toward a revolutionary theory: that species were not fixed but diversified from their ancestors over many generations, and that the driving mechanism of evolutionary change was natural selection.
In this concise, accessible book, Peter and Rosemary Grant explain what we have learned about the origin and evolution of new species through the study of the finches made famous by him: Darwin's finches. Drawing upon their unique observations of finch evolution over a thirty-four-year period, the Grants trace the evolutionary history of fourteen different species from a shared ancestor three million years ago. They show how repeated cycles of speciation involved adaptive change through natural selection on beak size and shape, and divergence in songs. They explain other factors that drive finch evolution, including geographical isolation, which has kept the Galápagos relatively free of competitors and predators; an increase in the number of islands and climate change over the last three million years, which enhanced opportunities for speciation; and flexibility in the early learning of feeding skills, which helped species to exploit new food resources.
Throughout, the Grants show how the laboratory tools of developmental biology and molecular genetics can be combined with observations and experiments on birds in the field to gain deeper insights into why the world is so biologically rich and diverse.
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
Peter R. Grant is the Class of 1877 Professor of Zoology at Princeton University. His books include Ecology and Evolution of Darwin's Finches (Princeton).
B. Rosemary Grant is a senior research scholar in the Department of Ecology and Evolutionary Biology at Princeton University. She is the coauthor, with Peter R. Grant, of Evolutionary Dynamics of a Natural Population: The Large Cactus Finch of the Galapagos.
"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
"I really enjoyed this book. It is a splendid introduction both to the finches and to radiation on islands. The authors' statement that 'Speciation is a process and not an event' should become one of the most famous quotes in evolutionary biology."
– John A. Endler, University of California, Santa Barbara
"A brilliant synthesis. The authors have written a concise summary of current understanding of one of the classic case studies of evolutionary diversification, Darwin's finches of the Galápagos. I can think of no parallel to this work. This book will be an inspiration to students. The Grants' love of the subject and the research comes through clearly."
– Jonathan B. Losos, Harvard University
"This is a book that summarizes decades of research on Darwin's finches and integrates it into a very accessible synthesis. What really distinguishes the book, of course, is the authority of the authors, who have lived with these birds for many years and have unparalleled familiarity with them. Readers will benefit enormously from the scholarship in this book."
– David B. Wake, University of California, Berkeley
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