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Over the last two decades, the study of speciation has expanded from a modest backwater of evolutionary biology into a large and vigorous discipline. Thus, the literature on speciation, as well as the number of researchers and students working in this area, has grown explosively. Despite these developments, there has been no book-length treatment of speciation in many years. As a result, both the seasoned scholar and the newcomer to evolutionary biology had no ready guide to the recent literature on speciation—a body of work that is enormous, scattered, and increasingly technical. Although several excellent symposium volumes have recently appeared, these collections do not provide a unified, critical, and up-to-date overview of the field. Speciation is designed to fill this gap.

Aimed at professional biologists, graduate students, and advanced undergraduates, Speciation covers both plants and animals (the first book on this subject to do so), and deals with all relevant areas of research, including biogeography, field work, systematics, theory, and genetic and molecular studies. It gives special emphasis to topics that are either controversial or the subject of active research, including sympatric speciation, reinforcement, the role of hybridization in speciation, the search for genes causing reproductive isolation, and mounting evidence for the role of natural and sexual selection in the origin of species. The authors do not hesitate to take stands on these and other controversial issues. This critical and scholarly book will be invaluable to researchers in evolutionary biology and is also ideal for a graduate-level course on speciation.

Jerry A. Coyne is Professor in the Department of Ecology and Evolution at the University of Chicago. He earned his Ph.D. (Biology) at Harvard University, followed by an NIH Postdoctoral Fellowship in the Department of Genetics at the University of California, Davis. He has taught undergraduate and graduate courses spanning a wide range of topics, including evolutionary biology, speciation, genetic analysis, social issues and scientific knowledge, and scientific speaking and writing. Dr. Coyne was awarded a Guggenheim fellowship in 1989. He has served as Vice President of the Society for the Study of Evolution (1996) and as Associate Editor of Evolution (1985-1988; 1994-2000) and The American Naturalist (1990-1993). His work is widely published, not only in scientific journals, but in such mainstream venues as The New York Times, the Times Literary Supplement, and The New Republic. His research interests include population and evolutionary genetics, speciation, ecological and quantitative genetics, chromosome evolution, and sperm competition.

H. Allen Orr is Professor in the Department of Biology at the University of Rochester, where he has taught courses in evolution, quantitative and population genetics, evolutionary genetics, and speciation. He completed his Ph.D. in Ecology and Evolution at the University of Chicago and undertook postdoctoral study at the University of California, Davis. Dr. Orr was awarded both the Young Investigator Prize (American Society of Naturalists, 1992) and the Dobzhansky Prize (Society for the Study of Evolution, 1993). Other honors include the David and Lucile Packard Fellowship in Science and Engineering (1995-2000) and a Guggenheim fellowship (2000-2001). Dr. Orr has served on the editorial boards of Evolution (1998-2000) and Genetical Research (1996-present), authored or coauthored numerous articles in scientific journals, and been a frequent contributor of book reviews and critical essays to such publications as The New York Review of Books, The New Yorker, and Boston Review. His research interests include population genetics, the genetics of speciation in Drosophila, and the genetics of adaptation.

Introduction
1. Species: Reality and Concepts
The Reality of Species
Sexually reproducing eukaryotic taxa
Groups with little or no sexual reproduction
Conclusions
Species Concepts
The biological species concept (BSC)
Advantages of the BSC
Problems with the BSC
Other species concepts
Why Are There Species?
2. Studying Speciation
The Problem of Speciation
Identifying and Measuring Reproductive Isolation
Absolute strength of isolating barriers
Relative strength of isolating barriers
Prezygotic versus postzygotic isolation
Which isolating barriers caused speciation?
Comparative Studies of Isolating Barriers
How fast does reproductive isolation appear?
Which traits promote the evolution of reproductive isolation?
3. Allopatric and Parapatric Speciation
Allopatric Speciation
Vicariant speciation
Peripatric speciation
Parapatric Speciation
Theory
Experimental evidence
Evidence from nature
Conclusions
4. Sympatric Speciation
Theory
Disruptive sexual selection
Disruptive natural selection
Conclusions
Experimental Evidence
Evidence from Nature
Evidence from habitat "islands"
Evidence from host races and host-specific species
Allochronic (temporal) isolation in sympatry
Comparative studies of the biogeography of speciation
Conclusions
5. Ecological Isolation
Habitat Isolation
Detecting and measuring habitat isolation
The problem of allopatry
Examples of habitat isolation
Relative importance of habitat isolation
The evolution of habitat isolation
The genetics of habitat isolation
Pollinator (Floral) Isolation
Detecting and measuring pollinator isolation
Examples of pollinator isolation
Relative importance of pollinator isolation
The evolution of pollinator isolation
The genetics of pollinator isolation
Temporal (Allochronic) Isolation
Detecting and measuring temporal isolation
Examples of temporal isolation
Relative importance of temporal isolation
The evolution of temporal isolation
The genetics of temporal isolation
Conclusions
6. Behavioral and Nonecological Isolation
Mating System "Isolation"
Behavioral Isolation
Detecting and measuring behavioral isolation
Examples of behavioral isolation
Relative importance of behavioral isolation
The evolution of behavioral isolation
The genetics of behavioral isolation
Mechanical Isolation
Examples of mechanical isolation
Relative importance of mechanical isolation
The evolution of mechanical isolation
The genetics of mechanical isolation
Gametic (Postmating, Prezygotic) Isolation
Examples of gametic isolation
Relative importance of gametic isolation
The evolution of gametic isolation
Conclusions
7. Postzygotic Isolation
Extrinsic Postzygotic Isolation
Intrinsic Postzygotic Isolation
The Frequency of Various Forms of Postzygotic Isolation
The Evolution of Extrinsic versus Intrinsic Postzygotic Isolation
Genetic Modes of Intrinsic Postzygotic Isolation
Chromosomal speciation: theory
Chromosomal speciation: data
Genic incompatibilities
The evolution of genic incompatibilities: the Dobzhansky-Muller model
Mathematical models of genic speciation
Wolbachia and cytoplasmic incompatibility
Conclusions
8. The Genetics of Postzygotic Isolation
Haldane's Rule
The phenomenon
The causes of Haldane's rule
Conclusions
The Genetic Basis of Postzygotic Isolation
How many genes cause postzygotic isolation?
Complexity of hybrid incompatibilities
Probability of hybrid incompatibilities
Where are the genes causing postzygotic isolation?
Developmental basis of postzygotic isolation
Are duplicate genes important?
Which genes cause postzygotic isolation?
9. Polyploidy and Hybrid Speciation
Polyploidy
Classification
Pathways to polyploidy
Incidence
Frequency of auto- versus allopolyploidy
Ecology and persistence
Why is polyploidy rarer in animals than in plants?
Recombinational Speciation
What is recombinational speciation?
Theory
The data: frequency and artificial hybrids
The data: natural recombinational speciation
The data meet the theory
10. Reinforcement
The Data
Selection experiments
Evidence from nature: case studies
Evidence from nature: comparative studies
Reinforcement of postzygotic isolation
The Theory
Early enthusiasm
Objections to reinforcement
The revival of reinforcement
Alternative Explanations
Publication bias
Differential fusion
Direct ecological effects
Ecological character displacement
Runaway sexual selection
Sympatric speciation
Distinguishing the Alternatives
11. Selection versus Drift
Speciation by Selection
Natural selection
Sexual selection
Mathematical theories of selection-based speciation
Speciation by Drift
Peak shift models
Theoretical Criticisms
Recent Peak Shift Models
The Data
Evidence from the laboratory
Evidence from nature
Conclusions
12. Speciation and Macroevolution
Rates of Speciation
What is a speciation rate?
Theory and speciation rates
Calculating speciation intervals
Extreme rates of speciation
What is the effect of biogeography?
Conclusions
Factors Affecting Speciation Rates
Tests for the effects of key factors
Distinguishing speciation from extinction
The data
Conclusions
Species Selection
Appendix. A Catalogue and Critique of Species Concepts
Genotypic Cluster Species Concept
Recognition Species Concept
Cohesion Species Concept
Evolutionary Species Concept
Ecological Species Concept
Phylogenetic Species Concepts
References
Author Index
Subject Index
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