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SPECIATION
JERRY A. COYNE
University of Chicago
H. ALLEN ORR
University of Rochester
Sinauer Associates, Inc. • Publishers
Sunderland, Massachusetts U.S.A.
© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured
or disseminated in any form without express written permission from the publisher.
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Contents
Introduction 1
1 Species: Reality and Concepts 9
The Reality of Species 10
Species Concepts 25
Sexually reproducing eukaryotic taxa 12
The biological species concept (BSC) 26
Groups with little or no sexual
reproduction 17
Advantages of the BSC 38
Conclusions 25
Other species concepts 48
Problems with the BSC 39
Why Are There Species? 48
2 Studying Speciation 55
The Problem of Speciation 57
Identifying and Measuring Reproductive
Isolation 61
Absolute strength of isolating barriers 62
Relative strength of isolating barriers 63
Prezygotic versus postzygotic
isolation 65
Which isolating barriers caused
speciation? 69
Comparative Studies of Isolating
Barriers 72
How fast does reproductive isolation
appear? 72
Which traits promote the evolution of
reproductive isolation? 81
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CONTENTS
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3 Allopatric and Parapatric Speciation 83
Allopatric Speciation 85
Theory 112
Vicariant speciation 86
Experimental evidence 117
Peripatric speciation 105
Evidence from nature 118
Parapatric Speciation 111
Conclusions 123
4 Sympatric Speciation 125
Theory 127
Disruptive sexual selection 128
Disruptive natural selection 130
Conclusions 136
Experimental Evidence 138
Evidence from Nature 141
Evidence from habitat “islands” 143
Evidence from host races and hostspecific species 157
Allochronic (temporal) isolation in
sympatry 166
Comparative studies of the
biogeography of speciation 168
Conclusions 175
5 Ecological Isolation 179
Habitat Isolation 182
Detecting and measuring habitat
isolation 184
The problem of allopatry 185
Examples of habitat isolation 186
Relative importance of pollinator
isolation 197
The evolution of pollinator isolation 198
The genetics of pollinator isolation 201
Temporal (Allochronic) Isolation 202
Relative importance of habitat
isolation 188
Detecting and measuring temporal
isolation 203
The evolution of habitat isolation 188
Examples of temporal isolation 204
The genetics of habitat isolation 191
Relative importance of temporal
isolation 205
Pollinator (Floral) Isolation 193
Detecting and measuring pollinator
isolation 194
Examples of pollinator isolation 195
The evolution of temporal isolation 206
The genetics of temporal isolation 210
Conclusions 210
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CONTENTS
6 Behavioral and Nonecological Isolation 211
Mating System “Isolation” 211
Behavioral Isolation 213
Relative importance of mechanical
isolation 229
Detecting and measuring behavioral
isolation 213
The evolution of mechanical
isolation 230
Examples of behavioral isolation 214
The genetics of mechanical isolation 231
Relative importance of behavioral
isolation 215
Gametic (Postmating, Prezygotic)
Isolation 232
The evolution of behavioral isolation 216
Examples of gametic isolation 233
The genetics of behavioral isolation 223
Relative importance of gametic
isolation 238
Mechanical Isolation 227
Examples of mechanical isolation 228
The evolution of gametic isolation 241
Conclusions 245
7 Postzygotic Isolation 247
Extrinsic Postzygotic Isolation 249
Chromosomal speciation: data 259
Intrinsic Postzygotic Isolation 253
Genic incompatibilities 267
The Frequency of Various Forms of
Postzygotic Isolation 255
The evolution of genic incompatibilities:
the Dobzhansky–Muller model 269
The Evolution of Extrinsic versus
Intrinsic Postzygotic Isolation 255
Mathematical models of genic
speciation 272
Genetic Modes of Intrinsic Postzygotic
Isolation 256
Wolbachia and cytoplasmic
incompatibility 276
Chromosomal speciation: theory 256
Conclusions 280
8 The Genetics of Postzygotic Isolation 283
Haldane’s Rule 284
The phenomenon 284
The causes of Haldane’s rule 286
Conclusions 298
The Genetic Basis of Postzygotic
Isolation 299
How many genes cause postzygotic
isolation? 299
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CONTENTS
Complexity of hybrid
incompatibilities 307
Developmental basis of postzygotic
isolation 309
Probability of hybrid
incompatibilities 308
Are duplicate genes important? 312
Where are the genes causing postzygotic
isolation? 308
ix
Which genes cause postzygotic
isolation? 313
9 Polyploidy and Hybrid Speciation 321
Polyploidy 321
Recombinational Speciation 337
Classification 322
What is recombinational speciation? 337
Pathways to polyploidy 324
Theory 338
Incidence 326
The data: frequency and artificial
hybrids 342
Frequency of auto- versus
allopolyploidy 328
Ecology and persistence 330
Why is polyploidy rarer in animals than
in plants? 333
The data: natural recombinational
speciation 344
The data meet the theory 350
10 Reinforcement 353
The Data 354
Selection experiments 355
The revival of reinforcement 372
Alternative Explanations 375
Evidence from nature: case studies 357
Publication bias 375
Evidence from nature: comparative
studies 362
Differential fusion 376
Reinforcement of postzygotic
isolation 365
Ecological character displacement 377
The Theory 366
Early enthusiasm 366
Direct ecological effects 377
Runaway sexual selection 378
Sympatric speciation 378
Distinguishing the Alternatives 379
Objections to reinforcement 369
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11 Selection versus Drift 383
Speciation by Selection 383
Theoretical Criticisms 394
Natural selection 385
Recent Peak Shift Models 396
Sexual selection 386
The Data 398
Mathematical theories of selection-based
speciation 387
Speciation by Drift 387
Evidence from the laboratory 398
Evidence from nature 401
Conclusions 410
Peak shift models 388
12 Speciation and Macroevolution 411
Rates of Speciation 411
Factors Affecting Speciation Rates 429
What is a speciation rate? 412
Tests for the effects of key factors 431
Theory and speciation rates 413
Calculating speciation intervals 416
Distinguishing speciation from
extinction 435
Extreme rates of speciation 425
The data 436
What is the effect of biogeography? 427
Conclusions 441
Conclusions 428
Species Selection 442
Appendix: A Catalogue and Critique of Species Concepts 447
Genotypic Cluster Species Concept 447
Evolutionary Species Concept 456
Recognition Species Concept 451
Ecological Species Concept 457
Cohesion Species Concept 452
Phylogenetic Species Concepts 459
References 473
Author Index 523
Subject Index 533
© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured
or disseminated in any form without express written permission from the publisher.