The Structure of Evolutionary Theory Read online

  STEPHEN JAY GOULD

  The Structure Of

  Evolutionary Theory

  ______________________________

  THE BELKNAP PRESS OF

  HARVARD UNIVERSITY PRESS

  CAMBRIDGE, MASSACHUSETTS

  AND LONDON, ENGLAND

  [Page v]

  Copyright © 2002 by the President and Fellows of Harvard College

  All rights reserved

  Printed in the United States of America

  Library of Congress Cataloging-in-Publication Data

  Gould, Stephen Jay.

  The structure of evolutionary theory / Stephen Jay Gould.

  p. cm.

  Includes bibliographical references (p. )

  ISBN 0-674-00613-5 (alk. paper)

  1. Evolution (Biology) 2. Punctuated equilibrium (Evolution) I. Title.

  QH366.2.G663 2002

  576.8—dc21 2001043556

  Sixth printing, 2002

  [Page vi]

  _________________________________________

  For Niles Eldredge and Elisabeth Vrba

  May we always be the Three Musketeers

  Prevailing with panache

  From our manic and scrappy inception at Dijon

  To our nonsatanic and happy reception at Doomsday

  All For One and One For All

  [Page vii]

  Contents

  Chapter 1:

  Defining and Revising the Structure of Evolutionary Theory 1

  Part I, Chapters 2-7

  The History of Darwinian Logic and Debate 91

  Segue to Part II 585

  Part II, Chapters 8-12

  Towards a Revised and Expanded Evolutionary Theory 593

  Bibliography 1344

  Illustration Credits 1388

  Index 1393

  About This P2P ePub Edition

  [Page ix]

  Expanded Contents

  Chapter 1: Defining and Revising the Structure of Evolutionary Theory 1

  • Theories Need Both Essences and Histories 1

  • The Structure of Evolutionary Theory: Revising the Three Central Features of Darwinian Logic 12

  • Apologia Pro Vita Sua 24

  A Time to Keep 24

  A Personal Odyssey 33

  • Epitomes for a Long Development 48

  Levels of Potential Originality 48

  An Abstract of One Long Argument 53

  Part I: The History of Darwinian Logic and Debate

  Chapter 2: The Essence of Darwinism and the Basis of Modern Orthodoxy: An Exegesis of the Origin of Species 93

  • A Revolution in the Small 93

  • Darwin as a Historical Methodologist 97

  One Long Argument 97

  The Problem of History 99

  A Fourfold Continuum of Methods for the Inference of History 103

  • Darwin as a Philosophical Revolutionary 116

  The Causes of Nature's Harmony 116

  Darwin and William Paley 116

  Darwin and Adam Smith 121

  The First Theme: The Organism as the Agent of Selection 125

  [Page x]

  The Second Theme: Natural Selection as a Creative Force 137

  The Requirements for Variation 141

  Copious 141

  Small 143

  Undirected 144

  Gradualism 146

  The Adaptationist Program 155

  The Third Theme: The Uniformitarian Need to Extrapolate: Environment as Enabler of Change 159

  • Judgments of Importance 163

  Chapter 3: Seeds of Hierarchy 170

  • Lamarck and the Birth of Modern Evolutionism in Two-Factor Theories 170

  The Myths of Lamarck 170

  Lamarck as a Source 174

  Lamarck's Two-Factor Theory: Sources for the Two Parts 175

  The First Set: Environment and Adaptation 176

  The Second Set: Progress and Taxonomy 179

  Distinctness of the Two Sets 181

  Lamarck's Two-Factor Theory: The Hierarchy of Progress and Deviation 175

  Antinomies of the Two-Factor Theory 189

  • An Interlude on Darwin's Reaction 192

  • No Allmacht without Hierarchy: Weissman on Germinal Selection 197

  The Allmacht of Selection 197

  Weismann's Argument on Lamarck and the Allmacht of Selection 201

  The Problem of Degeneration and Weismann's Impetus for Germinal Selection 203

  Some Antecedents to Hierarchy in German Evolutionary Thought 208

  Haeckel's Descriptive Hierarchy in Levels of Organization 208

  Roux's Theory of Intracorporeal Struggle 210

  Germinal Selection as a Helpmate to Personal Selection 214

  Germinal Selection as a Full Theory of Hierarchy 219

  • Hints of Hierarchy in Supraorganismal Selection: Darwin on the Principle of Divergence 224

  Divergence and the Completion of Darwin's System 224

  The Genesis of Divergence 232

  [Page xi]

  Divergence as a Consequence of Natural Selection 234

  The Failure of Darwin's Argument and the Need for Species Selection 236

  The Calculus of Individual Success 238

  The Causes of Trends 240

  Species Selection Based on Propensity for Extinction 246

  Postscript: Solution to the Problem of the “Delicate Arrangement” 248

  • Coda 249

  Chapter 4: Internalism and Laws of Form: Pre-Darwinian Alternatives to Functionalism 251

  • Prologue: Darwin's Fateful Decision 251

  • Two Ways to Glorify God in Nature 260

  William Paley and British Functionalism: Praising God in the Details of Design 262

  Louis Agassiz and Continental Formalism: Praising God in the Grandeur of Taxonomic Order 271

  An Epilog on the Dichotomy 278

  • Unity of Plan as the Strongest Version of Formalism: The Pre-Darwinian Debate 281

  Mehr Licht on Goethe's Leaf 281

  Geoffroy and Cuvier 291

  Cuvier and Conditions of Existence 291

  Geoffroy's Formalist Vision 298

  The Debate of 1830: Foreplay and Aftermath 304

  Richard Owen and English Formalism: The Archetype of Vertebrates 312

  No Formalism Please, We're British 312

  The Vertebrate Archetype: Constraint and Nonadaptation 316

  Owen and Darwin 326

  • Darwin's Strong but Limited Interest in Structural Constraint 330

  Darwin's Debt to Both Poles of the Dichotomy 330

  Darwin on Correlation of Parts 332

  The “Quite Subordinate Position” of Constraint to Selection 339

  Chapter 5: The Fruitful Facets of Galton's Polyhedron: Channels and Saltations in Post-Darwinian Formalism 342

  • Galton's Polyhedron 342

  [Page xii]

  • Orthogenesis as a Theory of Channels and One-Way Streets: the Marginalization of Darwinism 351

  Misconceptions and Relative Frequencies 351

  Theodor Eimer and the Ohnmacht of Selection 355

  Alpheus Hyatt: An Orthogenetic Hard Line from the World of Mollusks 365

  C.O. Whitman: An Orthogenetic Dove in Darwin's World of Pigeons 383

  • Saltation as a Theory of Internal Impetus: A Second Formalist Strategy for Pushing Darwinism to a Causal Periphery 396

  William Bateson: The Documentation of Inherent Discontinuity 396

  Hugo de Vries: A Most Reluctant Non-Darwinian 415

  Dousing the Great Party of 1909 415

  The (Not So Contradictory) Sources of the Mutation Theory 418

&n
bsp; The Mutation Theory: Origin and Central Tenets 425

  Darwinism and the Mutation Theory 439

  Confusing Rhetoric and the Personal Factor 439

  The Logic of Darwinism and Its Different Place in de Vries' System 443

  De Vries on Macroevolution 446

  Richard Goldschmidt's Appropriate Role as a Formalist Embodiment of All that Pure Darwinism Must Oppose 451

  Chapter 6: Pattern and Progress on the Geological Stage 467

  • Darwin and the Fruits of Biotic Competition 467

  A Geological License for Progress 467

  The Predominance of Biotic Competition and Its Sequelae 470

  • Uniformity on the Geological Stage 479

  Lyell's Victory in Fact and Rhetoric 479

  Catastrophism as Good Science: Cuvier's Essay 484

  Darwin's Geological Need and Kelvin's Odious Spectre 492

  A Question of Time (Too Little Geology) 496

  A Question of Direction (Too Much Geology) 497

  Chapter 7: The Modern Synthesis as a Limited Consensus 503

  • Why Synthesis? 503

  • Synthesis as Restriction 505

  The Initial Goal of Rejecting Old Alternatives 505

  [Page xiii]

  R. A. Fisher and the Darwinian Core 508

  J. B. S. Haldane and the Initial Pluralism of the Synthesis 514

  J. S. Huxley: Pluralism of the Type 516

  • Synthesis as Hardening 518

  The Later Goal of Exalting Selection's Power 518

  Increasing Emphasis on Selection and Adaptation between the First (1937) and Last (1951) Edition of Dobzhansky's Genetics and the Origin of Species 524

  The Shift in G. G. Simpson's Explanation of “Quantum Evolution” from Drift and Nonadaptation (1944) to the Embodiment of Strict Adaptation (1953) 528

  Mayr at the Inception (1942) and Codification (1963): Shifting from the “Genetic Consistency” to the “Adaptationist” Paradigm 531

  Why Hardening? 541

  • Hardening on the Other Two Legs of the Darwinian Tripod 543

  Levels of Selection 544

  Extrapolation into Geological Time 556

  • From Overstressed Doubt to Overextended Certainty 566

  A Tale of Two Centennials 566

  All Quiet on the Textbook Front 576

  Adaptation and Natural Selection 577

  Reduction and Trivialization of Macroevolution 579

  Segue to Part II 585

  Part II: Towards a Revised and Expanded Evolutionary Theory

  Chapter 8: Species as Individuals in the Hierarchical Theory of Selection 595

  • The Evolutionary Definition of Individuality 595

  An Individualistic Prolegomenon 595

  The Meaning of Individuality and the Expansion of the Darwinian Research Program 597

  Criteria for Vernacular Individuality 602

  Criteria for Evolutionary Individuality 608

  • The Evolutionary Definition of Selective Agency and the Fallacy of Selfish Genes 613

  [Page xiv]

  A Fruitful Error of Logic 613

  Hierarchical vs. Genie Selectionism 614

  The Distinction of Replicators and Interactors as a Framework for Discussion 615

  Faithful Replication as the Central Criterion for the Gene-Centered View of Evolution 616

  Sieves, Plurifiers, and the Nature of Selection: The Rejection of Replication as a Criterion of Agency 619

  Interaction as the Proper Criterion for Identifying Units of Selection 622

  The Internal Incoherence of Gene Selectionism 625

  Bookkeeping and Causality: The Fundamental Error of

  Gene Selectionism 632

  Gambits of Reform and Retreat by Gene Selectionists 637

  • Logical and Empirical Foundations for the Theory of Hierarchical Selection 644

  Logical Validation and Empirical Challenges 644

  R. A. Fisher and the Compelling Logic of Species Selection 644

  The Classical Arguments against Efficacy of Higher-Level Selection 646

  Overcoming These Classical Arguments, in Practice for Interdemic Selection, but in Principle for Species Selection 648

  Emergence and the Proper Criterion for Species Selection 652

  Differential Proliferation or Downward Effect? 652

  Shall Emergent Characters or Emergent Fitnesses Define the Operation of Species Selection? 656

  Hierarchy and the Sixfold Way 673

  A Literary Prologue for the Two Major Properties of Hierarchies 673

  Redressing the Tyranny of the Organism: Comments on Characteristic Features and Differences among Six Primary Levels 681

  The Gene-Individual 683

  Motoo Kimura and the “Neutral Theory of Molecular Evolution” 684

  True Genie Selection 689

  The Cell-Individual 695

  The Organism-Individual 700

  The Deme-Individual 701

  The Species-Individual 703

  [Page xv]

  Species as Individuals 703

  Species as Interactors 704

  Species Selection as Potent 709

  The Clade-Individual 712

  • The Grand Analogy: A Speciational Basis for Macroevolution 714

  Presentation of the Chart for Macroevolutionary Distinctiveness 714

  The Particulars of Macroevolutionary Explanation 716

  The Structural Basis 716

  Criteria for Individuality 720

  Contrasting Modalities of Change: The Basic Categories 721

  Ontogenetic Drive: The Analogy of Lamarckism and Anagenesis 722

  Reproductive Drive: Directional Speciation as an Important and Irreducible Macroevolutionary Mode Separate from Species Selection 724

  Species Selection, Wright's Rule, and the Power of Interaction with Directional Speciation 731

  Species-level Drifts as More Powerful than the Analogous Phenomena in Microevolution 735

  The Scaling of External and Internal Environments 738

  Summary Comments on the Strengths of Species Selection and its Interaction with Other Macroevolutionary Causes of Change 741

  Chapter 9: Punctuated Equilibrium and the Validation of Macroevolutionary Theory 745

  • What Every Paleontologist Knows 745

  An Introductory Example 745

  Testimonials to Common Knowledge 749

  Darwinian Solutions and Paradoxes 755

  The Paradox of Insulation from Disproof 758

  The Paradox of Stymied Practice 761

  • The Primary Claims of Punctuated Equilibrium 765

  Data and Definitions 765

  Microevolutionary Links 774

  Macroevolutionary Implications 781

  Tempo and the Significance of Stasis 782

  Mode and the Speciational Foundation of Macroevolution 783

  [Page xvi]

  • The Scientific Debate on Punctuated Equilibrium: Critiques and Responses 784

  Critiques Based on the Definability of Paleontological Species 784

  Empirical Affirmation 784

  Reasons for a Potential Systematic Underestimation of Biospecies by Paleospecies 789

  Reasons for a Potential Systematic Overestimation of Biospecies by Paleospecies 792

  Reasons Why an Observed Punctuational Pattern Might Not Represent Speciation 793

  Critiques Based on Denying Events of Speciation as the Primary Locus of Change 796

  Critiques Based on Supposed Failures of Empirical Results to Affirm Predictions of Punctuated Equilibrium 802

  Claims for Empirical Refutation by Cases 802

  Phenotypes 802

  Genotypes 810

  Empirical Tests of Conformity with Models 812

  • Sources of Data for Testing Punctuated Equilibrium 822

  Preamble 822

  The Equilibrium in Punctuated Equilibrium: Quantitatively Documented Patterns of Stasis in Unbranched Segments of Lineages 824

&nbs
p; The Punctuations of Punctuated Equilibrium: Tempo and Mode in the Origin of Paleospecies 839

  The Inference of Cladogenesis by the Criterion of Ancestral Survival 840

  The “Dissection” of Punctuations to Infer Both Existence and Modality 850

  Time 851

  Geography 852

  Morphometric Mode 852

  Proper and Adequate Tests of Relative Frequencies: The Strong Empirical Validation of Punctuated Equilibrium 854

  The Indispensability of Data on Relative Frequencies 854

  Relative Frequencies for Higher Taxa in Entire Biotas 856

  Relative Frequencies for Entire Clades 866

  Causal Clues from Differential Patterns of Relative Frequencies 870

  [Page xvii]

  • The Broader Implications of Punctuated Equilibrium for Evolutionary Theory and General Notions of Change 874

  What Changes May Punctuated Equilibrium Instigate in Our Views about Evolutionary Mechanisms and the History of Life? 874

  The Explanation and Broader Meaning of Stasis 874

  Frequency 875

  Generality 876

  Causality 877

  Punctuation, the Origin of New Macroevolutionary Individuals, and Resulting Implications for Evolutionary Theory 885

  Trends 886

  The Speciational Reformulation of Macroevolution 893

  Life Itself 897

  General Rules 901

  Particular Cases 905

  Horses as the Exemplar of “Life's Little Joke” 905

  Rethinking Human Evolution 908

  Ecological and Higher-Level Extensions 916