The structure of evolutionnary theory

STEPHEN JAY GOULD The Structure Of Evolutionary Theory THE BELKNAP PRESS OF HARVARD UNIVERSITY PRESS CAMBRIDGE, MASSACHUSETTS AND LONDON, ENGLAND 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) Evolution (Biology) Punctuated equilibrium (Evolution) I Title QH366.2.G663 2002 576.8—dc21 Sixth printing, 2002 2001043556 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 Contents Chapter 1: Defining and Revising the Structure of Evolutionary Theory 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 vii Expanded Contents Chapter 1: Defining and Revising the Structure of Evolutionary Theory • Theories Need Both Essences and Histories • The Structure of Evolutionary Theory: Revising the Three Central Features of Darwinian Logic 12 • Apologia Pro Vita Sua A Time to Keep A Personal Odyssey 24 24 33 • Epitomes for a Long Development Levels of Potential Originality An Abstract of One Long Argument 48 48 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 One Long Argument The Problem of History A Fourfold Continuum of Methods for the Inference of History 97 97 99 103 • Darwin as a Philosophical Revolutionary The Causes of Nature's Harmony Darwin and William Paley Darwin and Adam Smith The First Theme: The Organism as the Agent of Selection 116 116 116 121 125 ix x Contents The Second Theme: Natural Selection as a Creative Force The Requirements for Variation Copious Small Undirected Gradualism The Adaptationist Program The Third Theme: The Uniformitarian Need to Extrapolate: Environment as Enabler of Change • Judgments of Importance Chapter 3: Seeds of Hierarchy • Lamarck and the Birth of Modern Evolutionism in Two-Factor Theories The Myths of Lamarck Lamarck as a Source Lamarck's Two-Factor Theory: Sources for the Two Parts The First Set: Environment and Adaptation The Second Set: Progress and Taxonomy Distinctness of the Two Sets Lamarck's Two-Factor Theory: The Hierarchy of Progress and Deviation Antinomies of the Two-Factor Theory 137 141 141 143 144 146 155 159 163 170 170 170 174 175 176 179 181 175 189 • An Interlude on Darwin's Reaction 192 • No Allmacht without Hierarchy: Weissman on Germinal Selection The Allmacht of Selection Weismann's Argument on Lamarck and the Allmacht of Selection The Problem of Degeneration and Weismann's Impetus for Germinal Selection Some Antecedents to Hierarchy in German Evolutionary Thought Haeckel's Descriptive Hierarchy in Levels of Organization Roux's Theory of Intracorporeal Struggle Germinal Selection as a Helpmate to Personal Selection Germinal Selection as a Full Theory of Hierarchy 197 197 201 • Hints of Hierarchy in Supraorganismal Selection: Darwin on the Principle of Divergence Divergence and the Completion of Darwin's System The Genesis of Divergence 203 208 208 210 214 219 224 224 232 Contents Divergence as a Consequence of Natural Selection The Failure of Darwin's Argument and the Need for Species Selection The Calculus of Individual Success The Causes of Trends Species Selection Based on Propensity for Extinction Postscript: Solution to the Problem of the "Delicate Arrangement" • Coda Chapter 4: Internalism and Laws of Form: Pre-Darwinian Alternatives to Functionalism 234 236 238 240 246 248 249 251 • Prologue: Darwin's Fateful Decision 251 • Two Ways to Glorify God in Nature William Paley and British Functionalism: Praising God in the Details of Design Louis Agassiz and Continental Formalism: Praising God in the Grandeur of Taxonomic Order An Epilog on the Dichotomy 260 • Unity of Plan as the Strongest Version of Formalism: The Pre-Darwinian Debate Mehr Licht on Goethe's Leaf Geoffroy and Cuvier Cuvier and Conditions of Existence Geoffroy's Formalist Vision The Debate of 1830: Foreplay and Aftermath Richard Owen and English Formalism: The Archetype of Vertebrates No Formalism Please, We're British The Vertebrate Archetype: Constraint and Nonadaptation Owen and Darwin 262 271 278 281 281 291 291 298 304 312 312 316 326 • Darwin's Strong but Limited Interest in Structural Constraint Darwin's Debt to Both Poles of the Dichotomy Darwin on Correlation of Parts The "Quite Subordinate Position" of Constraint to Selection 330 330 332 339 Chapter 5: The Fruitful Facets of Galton's Polyhedron: Channels and Saltations in Post-Darwinian Formalism 342 • Galton's Polyhedron 342 xi xii Contents • Orthogenesis as a Theory of Channels and One-Way Streets: the Marginalization of Darwinism Misconceptions and Relative Frequencies Theodor Eimer and the Ohnmacht of Selection Alpheus Hyatt: An Orthogenetic Hard Line from the World of Mollusks CO Whitman: An Orthogenetic Dove in Darwin's World of Pigeons • Saltation as a Theory of Internal Impetus: A Second Formalist Strategy for Pushing Darwinism to a Causal Periphery William Bateson: The Documentation of Inherent Discontinuity Hugo de Vries: A Most Reluctant Non-Darwinian Dousing the Great Party of 1909 The (Not So Contradictory) Sources of the Mutation Theory The Mutation Theory: Origin and Central Tenets Darwinism and the Mutation Theory Confusing Rhetoric and the Personal Factor The Logic of Darwinism and Its Different Place in de Vries' System De Vries on Macroevolution Richard Goldschmidt's Appropriate Role as a Formalist Embodiment of All that Pure Darwinism Must Oppose 351 351 355 365 383 396 396 415 415 418 425 439 439 443 446 451 Chapter 6: Pattern and Progress on the Geological Stage 467 • Darwin and the Fruits of Biotic Competition A Geological License for Progress The Predominance of Biotic Competition and Its Sequelae 467 467 470 • Uniformity on the Geological Stage Lyell's Victory in Fact and Rhetoric Catastrophism as Good Science: Cuvier's Essay Darwin's Geological Need and Kelvin's Odious Spectre A Question of Time (Too Little Geology) A Question of Direction (Too Much Geology) 479 479 484 492 496 497 Chapter 7: The Modern Synthesis as a Limited Consensus 503 • Why Synthesis? 503 • Synthesis as Restriction The Initial Goal of Rejecting Old Alternatives 505 505 Contents R A Fisher and the Darwinian Core J B S Haldane and the Initial Pluralism of the Synthesis J S Huxley: Pluralism of the Type 508 514 516 • Synthesis as Hardening The Later Goal of Exalting Selection's Power Increasing Emphasis on Selection and Adaptation between the First (1937) and Last (1951) Edition of Dobzhansky's Genetics and the Origin of Species The Shift in G G Simpson's Explanation of "Quantum Evolution" from Drift and Nonadaptation (1944) to the Embodiment of Strict Adaptation (1953) Mayr at the Inception (1942) and Codification (1963): Shifting from the "Genetic Consistency" to the "Adaptationist" Paradigm Why Hardening? 518 518 531 541 • Hardening on the Other Two Legs of the Darwinian Tripod Levels of Selection Extrapolation into Geological Time 543 544 556 • From Overstressed Doubt to Overextended Certainty A Tale of Two Centennials All Quiet on the Textbook Front Adaptation and Natural Selection Reduction and Trivialization of Macroevolution 566 566 576 577 579 Segue to Part II 585 524 528 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 xiii xiv Contents A Fruitful Error of Logic Hierarchical vs Genie Selectionism The Distinction of Replicators and Interactors as a Framework for Discussion Faithful Replication as the Central Criterion for the GeneCentered View of Evolution Sieves, Plurifiers, and the Nature of Selection: The Rejection of Replication as a Criterion of Agency Interaction as the Proper Criterion for Identifying Units of Selection The Internal Incoherence of Gene Selectionism Bookkeeping and Causality: The Fundamental Error of Gene Selectionism Gambits of Reform and Retreat by Gene Selectionists • Logical and Empirical Foundations for the Theory of Hierarchical Selection Logical Validation and Empirical Challenges R A Fisher and the Compelling Logic of Species Selection The Classical Arguments against Efficacy of Higher-Level Selection Overcoming These Classical Arguments, in Practice for Interdemic Selection, but in Principle for Species Selection Emergence and the Proper Criterion for Species Selection Differential Proliferation or Downward Effect? 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Specter of Catastrophic Mass Extinction: Darwin to Chicxulub The Paradox of the First Tier: Towards a General Theory of Tiers of Time • An Epilog on Theory and History in Creating the Grandeur of