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
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 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