A reprint of a classical work in the Princeton Legacy Library. Originally published in 1999.
After his famous visit to the Galápagos Islands, Darwin speculated that "one might fancy that, from an original paucity of birds in this archipelago, one species had been taken and modified for different ends." Ecology and Evolution of Darwin's Finches is the classic account of how much we have since learned about the evolution of these remarkable birds. Based upon over a decade's research, Grant shows how interspecific competition and natural selection act strongly enough on contemporary populations to produce observable and measurable evolutionary change. In this new edition, Grant outlines new discoveries made in the thirteen years since the book's publication. Ecology and Evolution of Darwin's Finches is an extraordinary account of evolution in action.
"There is poetic justice in science: this classic modern study of natural selection in operation, by Peter Grant, has been carried out with Darwin's own Galápagos finches, the birds that will forever be associated with his concept of natural selection [...] If you somehow fell into the minority who didn't read the first edition, here is your chance to reform!"
– Jared Diamond, author of the Pulitzer Prize-winning Guns, Germs and Steel
"[This] is the best long-term, fine-scale study of speciation ever conducted [...] The science Peter Grant presents here is more interesting and important than ever."
– From the foreword by Jonathan Weiner
Preface to the 1999 Edition xv
ONE Introduction 3
Charles Darwin, 6 After Darwin, 9. The first synthesis, 9. Evolutionary inference, 10. Plan of the book, 12.
TWO Characteristics of the Islands 19
Origins and ages, 19. Distribution and sizes, 21. Climate, 21. Plants, 2Z Vegetation, 28. Changes in the past, 29. Changes in recent times, 30. Cocos Island, 31 Summary, 31.
THREE General Characteristics and Distributions of Finches 45
The main groups, 45. Genera, 51. Species, 51. Subspecies, 60. Distributions, 60. Patterns among the islands, 62. Extinctions, 64. Other land birds, 64.
FOUR Patterns of Morphological Variation 77
Introduction, 77. The major simple patterns, 77 The minor simple patterns, 79. Correlations between traits, 79. Size, 80. Allometry, 82. Shape, 82. Multivariate shape variation, 89. Geographical variation in size, 92. Summary, 95.
FIVE Growth and Development 100
Introduction, 100. Variation in egg size, 100. Absolute growth, 102. Relative growth, 106. Summary, 111.
SIX Beak Sizes, Beak Shapes, and Diets 113
Introduction, 113. Feeding mechanics, 113. Feeding types, 116. Ecological significance of beak differences between species, 117. Dietary differences between species, 118. Dietary differences between populations of the same species, 128. Dietary differences among individuals in a variable population, 132. Summary, 138.
SEVEN The Importance of Food to Finch Populations 147
Introduction, 147. Plant phenology in the and zone, 147 Finch phenology, 148. Finch populations in relation to food supply, 152. Extreme conditions, 152. Food limitation of population sizes, 154. The frequency of food limitation, 168. Other factors limiting finch populations, 171. Interspecific competition for food, 173. Summary, 173.
EIGHT Population Variation and Natural Selection 175
Introduction, 175. Relative variation, 175. Theoretical background, 177. Field studies, 180. Genetic variation, 180. Natural selection, 183. Sexual selection, 192. Countervailing selection, 193. A summary of selection pressures, 195. Sexual dimorphism, 196. Genetic drift, 197. Enhancement of genetic variation, 199, Variation in relation to abundance, 20Z Other species, 208. Summary, 219.
NINE Species-Recognition and Mate Choice 222
Introduction, 222. The possible cues used in species-recognition, 222. Morphological cues, 224. Song, 251. Song and bill morphology as species cues, 241. Imprinting, 242. The learning of heterotypic song, 244. Misimprinting, 246 Beyond species-recognition: mate choice, 249. Summary, 251.
TEN Evolution and Speciation 253
Evolution, 253. Origins, 253 The number of species, 256 The pattern of speciation, 257 The time framework 260. Allopatric speciation, 263. Alternative models of speciation, 273. Parapatric speciation, 274. Sympatric speciation, 275. Alternatives to gradual genetic change, 280. Conclusions and summary, 283.
ELEVEN Ecological Interactions during Speciation 285
Introduction, 285. Ecological isolation, 285. Causes of initial differentiation, 286. An alternative view, 288. Differentiation entirely in allopatry, 289. The food supply hypothesis, 291. Lack's evidence for competition, 294. Tests of the competition hypothesis, 300. Different explanations reconciled, 310. Conclusions and summary, 312.
TWELVE Competition and Finch Communities 314
Introduction, 314. Combinations of species, 315. Structure determined by competition, 317 Minimum differences between coexisting species, 321. Greater than minimum differences, 323. A digression on methods of analysis, and on bias, 328 Predictive models, 331. The classical case of character release, 340. Conclusions and summary, 346
THIRTEEN The Evolution of Reproductive Isolation 348
Introduction, 348. Experimental tests, 348. Implications of the experimental results, 350. Reinforcement? 353. Absence of speciesfrom islands, 354. Summary, 355.
FOURTEEN Adaptation: Body Size, Plumage Coloration, and Other Traits 357
Introduction, 357 Historical survey, 358. Body size, 359. Plumage, 364. Otherfeatures, 371. Summary, 373.
FIFTEEN Reconstruction of Phylogeny 375
Introduction, 375. Reconstructing the process of morphological divergence, 375. Comparison with contemporary selection, 379. Further evolution, 380. Ontogeny, 381. Phylogeny. 383. Summary, 387.
SIXTEEN Recapitulation and Generalization 389
Introduction, 389. Patterns and processes among modem finches, 389. Evolution, 397 Generalizations, 401.
APPENDIX: Spanish and English Names of the Major Galapagos Islands 413
Author Index 471
Subject Index 476
Color Plates Following page 16
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