Douglas J. FutuymaState University of New York at Stony Brook

EvolutionDas Original mit

bersetzungshilfen

Chapter 19 (Evolution of Genes and Genomes)by Scott V. Edwards, Harvard University

Chapter 20 (Evolution and Development)by John R. True, State University of New York at Stony Brook

Ubersetzung von Andreas Held

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Evolutionsbiologie 1

Der Stammbaum des Lebens: Klassifikation und Phylogenie 17

Evolutionsmuster 43

Evolution in den Fossilbelegen 67

Eine Geschichte des Lebens auf der Erde 91

Die Geographie der Evolution 117

Die Evolution der Biodiversitat 139

Die Entstehung der genetischen Variability 161

Variability 189

Genetische Drift: Evolution nach dem Zufallsprinzip 225

Natiirliche Selektion und Anpassung 247

Die genetische Theorie der naturlichen Selektion 269

Die Evolution phanotypischer Merkmale 297

Konflikt und Kooperation 325

Arten 353

Artbildung 379

Fitness: Fortpflanzungserfolg 405

Coevolution: Die Evolution von Wechselbeziehungenzwischen Arten 429

Die Evolution von Genen und Genomen 449

Evolution und Entwicklung 473

Makroevolution: Evolution oberhalb der Artebene 501

Evoiutionswissenschaft, Kreationismus und Gesellschaft 523

Contents

Evolutionary Biology 1What Is Evolution? 2Before Darwin 4Charles Darwin 6Darwin's Evolutionary Theory 7Evolutionary Theories after Darwin 8The Evolutionary Synthesis 9

Fundamental principles of evolution 9

Evolutionary Biology since the Synthesis 11Philosophical Issues 12Ethics, Religion, and Evolution 12Evolution as Fact and Theory 13

2 The Tree of Life:Classification andPhytogeny 17

3Patterns of Evolution 43Evolutionary History and Classification 45Inferring the History of Character Evolution 46Some Patterns of Evolutionary Change Inferred from

Systematics 48Most features of organisms have been modified from

pre-existing features 48Homoplasy is common 51Rates of character evolution differ 54Evolution is often gradual 55Change in form is often correlated with change in function 55Similarity between species changes throughout ontogeny 56Development underlies some common patterns of morpho-

logical evolution 56

Phylogenetic Analysis Documents Evolutionary Trends 61Many Clades Display Adaptive Radiation 62

Classification 19Inferring Phylogenetic History 22

Similarity and common ancestry 22Complications in inferring phylogeny 23The method of maximum parsimony 25An example of phylogenetic analysis 28Evaluating phylogenetic hypotheses 29

Molecular Clocks 32Gene Trees 34Difficulties in Phylogenetic Analysis 35Hybridization and Horizontal Gene Transfer 39

4 Evolution in the FossilRecord 67

Some Geological Fundamentals 68Rock formation 68Plate tectonics 68Geological time 69The geological time scale 69

The Fossil Record 71Evolutionary changes within species 71Origins of higher taxa 71

The Hominin Fossil Record 79Phylogeny and the Fossil Record 83

CONTENTS v i i

Evolutionary Trends 83Punctuated Equilibria 84Rates of Evolution 86

5 A History of Lifeon Earth 91

Before Life Began 92The Emergence of Life 92Precambrian Life 94

Prokaryotes 94Eukaryotes 95Proterozoic life 96

Paleozoic Life: The Cambrian Explosion 97Paleozoic Life: Ordovician to Devonian 99

Marine life 99Terrestrial life 101

Paleozoic Life: Carboniferous and Permian 102Terrestrial life 102Aquatic life 103

Mesozoic Life 103Marine life 103Terrestrial plants and arthropods 105Vertebrates 107

The Cenozoic Era 109Aquatic life 109Terrestrial life 109The adaptive radiation of mammals 111Pleistocene events 112

6 The Geographyof Evolution 117

Biogeographic Evidence for Evolution 118Major Patterns of Distribution 119Historical Factors Affecting Geographic Distributions 121Testing Hypotheses in Historical Biogeography 123

Examples of historical biogeographic analyses 125The composition of regional biotas 128

Phylogeography 129Ecological Approaches to Biogeography 132

The theory of island biogeography 134Structure and diversity in ecological communities 134Community convergence 134

Effects of History on Contemporary Diversity Patterns136

7The Evolution ofBiodiversity 139

Estimating Changes in Taxonomic Diversity 140Estimates of diversity 140Rates 141

Taxonomic Diversity through the Phanerozoic 143Rates of origination and extinction 144Causes of extinction 146Declining extinction rates 146Mass extinctions 148Origination and diversification 151The role of environmental change 156

The Future of Biodiversity 157

8 The Origin of GeneticVariation 161

Genes and Genomes 162Gene Mutations 165

Kinds of mutations 166Examples of mutations 169Rates of mutation 171Phenotypic effects of mutations 174Effects of mutations on fitness 176The limits of mutation 178

Mutation as a Random Process 178Recombination and Variation 179Alterations of the Karyotype 181

Polyploidy 181Chromosome rearrangements 182

9Variation 189Distinguishing Sources of Phenotypic Variation 190Fundamental Principles of Genetic Variation in

Populations 192Frequencies of alleles and genotypes: The Hardy-Weinberg

principle 193An example: The human MN locus 194The significance of the Hardy-Weinberg principle: Factors in

evolution 196Frequencies of alleles, genotypes, and phenotypes 197Inbreeding 197

VIII CONTENTS

Genetic Variation in Natural Populations 199Polymorphism 199Genetic variation in viability 199Inbreeding depression 201Genetic variation in proteins 202Variation at the DNA level 204Multiple loci and the effects of linkage 205Variation in quantitative traits 207

Variation among Populations 212Patterns of geographic variation 212Adaptive geographic variation 216Gene flow 216Allele frequency differences among populations 217Geographic variation among humans 219

7 / j Genetic Drift: EvolutionJL \J at Random 225

The Theory of Genetic Drift 226Genetic drift as sampling error 226Coalescence 227Random fluctuations in allele frequencies 229

Evolution by Genetic Drift 231Effective population size 231Founder effects 232Genetic drift in real populations 232

The Neutral Theory of Molecular Evolution 235Principles of the neutral theory 236Variation within and among species 238Do comparisons among species support the neutral

theory? 239

Gene Flow and Genetic Drift 241Gene trees and population history 241The origin of modern Homo sapiens revisited 243

11Natural Selectionand Adaptation 247

Adaptations in Action: Some Examples 248The Nature of Natural Selection 250

Design and mechanism 250Definitions of natural selection 251Natural selection and chance 251Selection of and selection for 252

Experimental Studies of Natural Selection 252Bacterial populations 252Inversion polymorphism in Drosophila 253Male reproductive success 254Population size in flour beetles 255Selfish genetic elements 256

Levels of Selection 257Selection of organisms and groups 257Species selection 258

The Nature of Adaptations 260Definitions of adaptation 260Recognizing adaptations 261

What Not to Expect of Natural Selection andAdaptation 264

The necessity of adaptation 264Perfection 264Progress 264Harmony and the balance of nature 265Morality and ethics 265

12The Genetical Theory ofNatural Selection 269

Fitness 270Modes of selection 270Defining fitness 271Components of fitness 272

Models of Selection 273Directional selection 273Deleterious alleles in natural populations 278

Polymorphism Maintained by Balancing Selection 280Heterozygote advantage 280Antagonistic and varying selection 282Frequency-dependent selection 283

Multiple Outcomes of Evolutionary Change 286Positive frequency-dependent selection 286Heterozygote disadvantage 286Adaptive landscapes 287Interaction of selection and genetic drift 287

Molecular Signatures of Natural Selection 288Theoretical expectations 288Examples 290

The Strength of Natural Selection 293

CONTENTS i x

13Evolution ofPhenotypic Traits 297

Evolution Observed 298Components of Phenotypic Variation 299How Polygenic are Polygenic Characters? 301Linkage Disequilibrium 303Evolution of Quantitative Characters 304

Genetic variance in natural populations 304Response to selection 305Responses to artificial selection 306

Selection in Natural Populations 308Measuring natural selection on quantitative characters 308Examples of selection on quantitative characters 309

A Neutral Model of the Evolution of QuantitativeCharacters 311

What Maintains Genetic Variation in QuantitativeCharacters? 312

Correlated Evolution of Quantitative Traits 312Correlated selection 312Genetic correlation 313Examples of genetic correlation 314How genetic correlation affects evolution 314

Can Genetics Predict Long-Term Evolution? 316Norms of Reaction 317

Phenotypic plasticity 317Canalization 319Evolution of variability 319

14Conflict andCooperation 325

Direct benefits of mate choice 333Sensory bias 333Indirect benefits of mate choice 333Antagonistic coevolution 337

Social Interactions and the Evolution ofCooperation 339

Theories of cooperation and altruism 339Interactions among related individuals 341

A Genetic Battleground: The Nuclear Family 343Mating systems and parental care 343Infanticide, abortion, and siblicide 345Parent-offspring conflict 345

Genetic Conflicts 346Parasitism, mutualism, and the evolution of individuals 348

15Species 353

A Framework for Conflict and Cooperation 326Levels of organization 326Inclusive fitness and kin selection 326Frequency-dependent selection on interactions 327Evolutionarily stable strategies 327

Sexual Selection 329The concept of sexual selection 329Contests between males and between sperm 330Sexual selection by mate choice 332

What Are Species? 354Phylogenetic species concepts 355The biological species concept 355Domain and application of the biological species concept 357When species concepts conflict 358

Barriers to Gene Flow 359Premating barriers 359Postmating, prezygotic barriers 362Postzygotic barriers 362

How Species Are Diagnosed 363Differences among Species 364The Genetic Basis of Reproductive Barriers 366

Genes affecting reproductive isolation 366Functions of genes that cause reproductive isolation 368Chromosome differences and postzygotic isolation 369Cytoplasmic incompatibility 370The significance of genetic studies of reproductive

isolation 371

Molecular Divergence among Species 372Hybridization 373

Primary and secondary hybrid zones 373Genetic dynamics in a hybrid zone 374The fate of hybrid zones 375

CONTENTS

16Speciation 379Modes of Speciation 380Allopatric Speciation 381

Evidence for allopatric speciation 381Mechanisms of vicariant allopatric speciation 383Ecological selection and speciation 384Sexual selection and speciation 386Reinforcement of reproductive isolation 387Peripatric speciation 389

Alternatives to Allopatric Speciation 392Parapatric speciation 392Sympatric speciation 393

Polyploidy and Recombinational Speciation 396Polyploidy 396Recombinational speciation 398

How Fast Is Speciation? 399Consequences of Speciation 401

17How to Be Fit:Reproductive Success 405

Individual Selection and Group Selection 406Life History Evolution 407

Life history traits as components of fitness 407Trade-offs 408

The Theory of Life History Evolution 411Life span and senescence 411Age schedules of reproduction 412Number and size of offspring 413The evolution of the rate of increase 414Male reproductive success 415

Modes of Reproduction 416The evolution of mutation rates 417Sexual and asexual reproduction 417The problem with sex 418Hypotheses for the advantage of sex and recombination 419

Sex Ratios, Sex Allocation, and Sex Determination 422The evolution of sex ratios 422Sex allocation, hermaphroditism, and dioecy 424

Inbreeding and Outcrossing 424Advantages of inbreeding and outcrossing 425

18Coevolution: EvolvingInteractions amongSpecies 429

The Nature of Coevolution 430Phylogenetic Aspects of Species Associations 431Coevolution of Enemies and Victims 432

Models of enemy-victim coevolution 434Examples of predator-prey coevolution 435Infectious disease and the evolution of parasite virulence 437

Mutualisms 439The Evolution of Competitive Interactions 441

Community patterns 443Multispecies interactions 444

19Evolution of Genesand Genomes 449

Evolution of Genes and Proteins 451Adaptive evolution and neutrality 452Sequence evolution under purifying and positive selection 453Adaptive molecular evolution in primates 454Adaptive evolution across the genome 456

Genome Diversity and Evolution 456Diversity of genome structure 456

Viral and microbial genomes: The smallest genomes 457The C-value paradox 458Repetitive sequences and transposable elements 459

The Origin of New Genes 461Lateral gene transfer 461Exon shuffling 462Gene chimerism and processed pseudogenes 463Motif multiplication and exon loss 464Gene duplication and gene families 465

Phylogenetic and Adaptive Diversification in GeneFamilies 468

Gene conversion 468Phylogenetic patterns following gene duplication 469Selective fates of recently duplicated loci 469Rates of gene duplication 470

CONTENTS x i

20 Evolution andDevelopment 473

Hox Genes and the Dawn of Modern EDB 474Types of Evidence in Contemporary EDB 478The Evolving Concept of Homology 479Evolutionarily Conserved Developmental Pathways 480The Evolution of Gene Regulation: The Keystone of

Developmental Evolution 484Modularity in morphological evolution 485Co-option and the evolution of novel characters 486The developmental genetics of heterochrony 488The evolution of allometry 489

Developmental Constraints and MorphologicalEvolution 490

The Developmental Genetic Basis of Short-TermMorphological Evolution 493

The Molecular Genetic Basis of Gene RegulatoryEvolution 494

Toward the EDB of Homo sapiens 496

21Macroevolution:Evolution above theSpecies Level 501

Rates of Evolution 502Rates of character evolution 502Punctuated equilibrium, revisited 502Stasis 504

Gradualism and Saltation 506Phylogenetic Conservatism and Change 508

Stabilizing selection 508Limitations on variation 509

The Evolution of Novelty 510Accounting for incipient and novel features 510Complex characteristics 512

Trends and Progress 513Trends: Kinds and causes 513Examples of trends 514Are there major trends in the history of life? 515The question of progress 518

Evolutionary Science,Creationism, andSociety 523

Creationists and Other Skeptics 524Science, Belief, and Education 525The Evidence for Evolution 528

The fossil record 528Phylogenetic and comparative studies 528Genes and genomes 529Biogeography 529

Failures of the argument from design 529Evolution and its mechanisms, observed 531

Refuting Creationist Arguments 532On arguing for evolution 537

Why Should We Teach Evolution? 537Health and medicine 538Agriculture and natural resources 540Environment and conservation 541Understanding nature and humanity 541

Glossar mit deutschenErlauterungen 545

Literature Cited 559

Index 585