Unofficial SJG Archive

The Unofficial Stephen Jay Gould Archive

Unofficial SJG Archive

  The Structure of Evolutionary Theory (2002)

The world's most revered and eloquent interpreter of evolutionary ideas offers here a work of explanatory force unprecedented in our time—a landmark publication, both for its historical sweep and for its scientific vision.

With characteristic attention to detail, Stephen Jay Gould first describes the content and discusses the history and origins of the three core commitments of classical Darwinism: that natural selection works on organisms, not genes or species; that it is almost exclusively the mechanism of adaptive evolutionary change; and that these changes are incremental, not drastic. Next, he examines the three critiques that currently challenge this classic Darwinian edifice: that selection operates on multiple levels, from the gene to the group; that evolution proceeds by a variety of mechanisms, not just natural selection; and that causes operating at broader scales, including catastrophes, have figured prominently in the course of evolution.

Then, in a stunning tour de force that will likely stimulate discussion and debate for decades, Gould proposes his own system for integrating these classical commitments and contemporary critiques into a new structure of evolutionary thought.

In 2001 the Library of Congress named Stephen Jay Gould one of America’s eighty-three Living Legends—people who embody the “quintessentially American ideal of individual creativity, conviction, dedication, and exuberance.” Each of these qualities finds full expression in this peerless work, the likes of which the scientific world has not seen—and may not see again—for well over a century.

STEPHEN JAY GOULD is the Alexander Agassiz Professor of Zoology at Harvard and the Vincent Astor visiting Professor of Biology at New York University. A MacArthur Prize Fellow, he has recieved innumerable honors and awards and has written many books, including Ontogeny and Phylogeny and Time's Arrow, Time's Cycle (both from Harvard). He lives in Cambridge, Massachusetts, and New York City.

    Michael Ruse

I urge you to get this work and (if not to read it straight through) to dip into it judiciously, for there is much of great worth here, and as a guide to the way that today’s evolutionists think it is surely destined to be a classic.

Review of Structure of Evolutionary Theory Isis 94 (2003): 397.


  • Theories Need Both Essences and Histories1
  • The Structure of Evolutionary Theory: Revising the Three Central Features of Darwinian Logic12
  • Apologia Pro Vita Sua24
  •         A Time to Keep24
  •         A Personal Odyssey33
  • Epitomes for a Long Development48
  •         Levels of Potential Originality48
  •         An Abstract of One Long Argument53
  • A Revolution in the Small93
  • Darwin as a Historical Methodologist97
  •         One Long Argument97
  •         The Problem of History99
  •         A Fourfold Continuum of Methods for the Inference of History103
  • Darwin as a Philosophical Revolutionary116
  •      The Causes of Nature's Harmony116
  •           Darwin and William Paley116
  •           Darwin and Adam Smith116
  •      The First Theme: The Organism as the Agent of Selection125
  •      The Second Theme: Natural Selection as a Creative Force137
  •           The Requirements for Variation141
  •             Copious141
  •             Small141
  •             Undirected141
  •           Gradualism146
  •           The Adaptationist Program155
  •      The Third Theme: The Uniformitarian Need to Extrapolate: Environment as Enabler of Change159
  • Judgments of Importance163
  • Lamarck and the Birth of Modern Evolutionism in Two-Factor Theories170
  •      The Myths of Lamarck170
  •      Lamarck as a Source174
  •      Lamarck's Two-Factor Theory: Sources for the Two Parts175
  •             The First Set: Environment and Adaptation176
  •             The Second Set: Progress and Taxonomy179
  •             Distinctness of the Two Sets181
  •      Lamarck's Two-Factor Theory: The Hierarchy of Progress and Deviation175
  •      Antinomies of the Two-Factor Theory189
  • An Interlude on Darwin's Reaction192
  • No Allmacht without Hierarchy: Weissman on Germinal Selection197
  •      The Allmacht of Selection197
  •      Weismann's Argument on Lamarck and the Allmacht of Selection201
  •      The Problem of Degeneration and Weismann's Impetus for Germinal Selection203
  •      Some Antecedents to Hierarchy in German Evolutionary Thought203
  •             Haeckel's Descriptive Hierarchy in Levels of Organization208
  •             Roux's Theory of Intracorporeal Struggle210
  •      Germinal Selection as a Helpmate to Personal Selection214
  •      Germinal Selection as a Full Theory of Hierarchy214
  • Hints of Hierarchy in Supraorganismal Selection: Darwin on the Principle of Divergence224
  •      Divergence and the Completion of Darwin's System224
  •      The Genesis of Divergence232
  •      Divergence as a Consequence of Natural Selection234
  •      The Failure of Darwin's Argument and the Need for Species Selection236
  •             The Calculus of Individual Success238
  •             The Causes of Trends240
  •             Species Selection Based on Propensity for Extinction246
  •      Postscript: Solution to the Problem of the "Delicate Arrangement"248
  • Coda249
  • Prologue: Darwin's Fateful Decision251
  • Two Ways to Glorify God in Nature260
  •             William Paley and British Functionalism: Praising God in the Details of Design262
  •             Louis Agassiz and Continental Formalism: Praising God in the Grandeur of Taxonomic Order271
  •             An Epilog on the Dichotomy278
  • Unity of Plan as the Strongest Version of Formalism: The Pre-Darwinian Debate281
  •      Mehr Licht on Goethe's Leaf281
  •      Geoffroy and Cuvier291
  •             Cuvier and Conditions of Existence291
  •             Geoffroy's Formalist Vision298
  •             The Debate of 1830: Foreplay and Aftermath304
  •      Richard Owen and English Formalism: The Archetype of Vertebrates312
  •             No Formalism Please, We're British312
  •             The Vertebrate Archetype: Constraint and Nonadaptation316
  •             Owen and Darwin326
  • Darwin's Strong but Limited Interest in Structural Constraint330
  •      Darwin's Debt to Both Poles of the Dichotomy330
  •      Darwin on Correlation of Parts332
  •      The "Quite Subordinate Position" of Constraint to Selection339
  • Galton's Polyhedron330
  • Orthogenesis as a Theory of Channels and One-Way Streets: the Marginalization of Darwinism351
  •      Misconceptions and Relative Frequencies351
  •      Theodor Eimer and the Ohnmacht of Selection355
  •      Alpheus Hyatt: An Orthogenetic Hard Line from the World of Mollusks365
  •      CO. Whitman: An Orthogenetic Dove in Darwin's World of Pigeons383
  • Saltation as a Theory of Internal Impetus: A Second Formalist Strategy for Pushing Darwinism to a Causal Periphery396
  •      William Bateson: The Documentation of Inherent Discontinuity396
  •      Hugo de Vries: A Most Reluctant Non-Darwinian415
  •             Dousing the Great Party of 1909415
  •             The (Not So Contradictory) Sources of the Mutation Theory418
  •             The Mutation Theory: Origin and Central Tenets425
  •             Darwinism and the Mutation Theory439
  •                Confusing Rhetoric and the Personal Factor439
  •                The Logic of Darwinism and Its Different Place in de Vries' System443
  •             De Vries on Macroevolution446
  •      Richard Goldschmidt's Appropriate Role as a Formalist Embodiment of All that Pure Darwinism Must Oppose451
  • Darwin and the Fruits of Biotic Competition467
  •      A Geological License for Progress467
  •      The Predominance of Biotic Competition and Its Sequelae470
  • Uniformity on the Geological Stage479
  •      Lyell's Victory in Fact and Rhetoric479
  •      Catastrophism as Good Science: Cuvier's Essay484
  •      Darwin's Geological Need and Kelvin's Odious Spectre492
  •             A Question of Time (Too Little Geology)496
  •             A Question of Direction (Too Much Geology)497
  • Why Synthesis?503
  • Synthesis as Restriction505
  •      The Initial Goal of Rejecting Old Alternatives505
  •      R. A. Fisher and the Darwinian Core508
  •      J. B. S. Haldane and the Initial Pluralism of the Synthesis514
  •      J. S. Huxley: Pluralism of the Type516
  • Synthesis as Hardening518
  •      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 Species524
  •      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" Paradigm531
  •      Why Hardening?541
  • Hardening on the Other Two Legs of the Darwinian Tripod543
  •      Levels of Selection544
  •      Extrapolation into Geological Time556
  • From Overstressed Doubt to Overextended Certainty566
  •      A Tale of Two Centennials566
  •      All Quiet on the Textbook Front566
  •             Adaptation and Natural Selection577
  •             Reduction and Trivialization of Macroevolution579
  • The Evolutionary Definition of Individuality595
  •      An Individualistic Prolegomenon595
  •      The Meaning of Individuality and the Expansion of the Darwinian Research Program597
  •             Criteria for Vernacular Individuality602
  •             Criteria for Evolutionary Individuality608
  • The Evolutionary Definition of Selective Agency and the Fallacy of Selfish Genes613
  •      A Fruitful Error of Logic613
  •      Hierarchical vs. Genie Selectionism614
  •             The Distinction of Replicators and Interactors as a Framework for Discussion615
  •             Faithful Replication as the Central Criterion for the Gene-Centered View of Evolution616
  •             Sieves, Plurifiers, and the Nature of Selection: The Rejection of Replication as a Criterion of Agency619
  •             Interaction as the Proper Criterion for Identifying Units of Selection622
  •             The Internal Incoherence of Gene Selectionism625
  •             Bookkeeping and Causality: The Fundamental Error of Gene Selectionism632
  •             Gambits of Reform and Retreat by Gene Selectionists637

D e f i n i n g   a n d   R e v i s i n g

T h e  S t r u c t u r e  o f  E v o l u t i o n a r y  T h e o r y 

Theories Need Both Essences and Histories

In a famous passage added to later editions of the Origin of Species, Charles Darwin (1872, p. 134) generalized his opening statement on the apparent absurdity of evolving a complex eye through a long series of gradual steps by reminding his readers that they should always treat "obvious" truths with skepticism. In so doing, Darwin also challenged the celebrated definition of science as "organized common sense," as championed by his dear friend Thomas Henry Huxley. Darwin wrote:

"When it was first said that the sun stood still and world turned round, the common sense of mankind declared the doctrine false; but the old saying of Vox populi, vox Dei [the voice of the people is the voice of God], as every philosopher knows, cannot be trusted in science."

Despite his firm residence within England's higher social classes, Darwin took a fully egalitarian approach towards sources of expertise, knowing full well that the most dependable data on behavior and breeding of domesticated and cultivated organisms would be obtained from active farmers and husbandmen, not from lords of their manors or authors of theoretical treatises. As Ghiselin (1969) so cogently stated, Darwin maintained an uncompromisingly "aristocratic" set of values towards judgment of his work—that is, he cared not a whit for the outpourings of vox populi, but fretted endlessly and fearfully about the opinions of a very few key people blessed with the rare mix of intelligence, zeal, and attentive practice that we call expertise (a democratic human property, respecting only the requisite mental skills and emotional toughness, and bearing no intrinsic correlation to class, profession or any other fortuity of social circumstance).

Darwin ranked Hugh Falconer, the Scottish surgeon, paleontologist, and Indian tea grower, within this most discriminating of all his social groups, a panel that included Hooker, Huxley and Lyell as the most prominent members. Thus, when Falconer wrote his important 1863 paper on American fossil elephants (see Chapter 9, pages 745-749, for full discussion of this incident), Darwin flooded himself with anticipatory fear, but then rejoiced in his critic's generally favorable reception of evolution, as embodied in the closing sentence of Falconer's key section: "Darwin has, beyond all his cotemporaries [sic], given an impulse to the philosophical investigation of the most backward and obscure branch of the Biological Sciences of his day; he has laid the foundations of a great edifice; but he need not be surprised if, in the progress of erection, the superstructure is altered by his successors, like the Duomo of Milan, from the roman to a different style of architecture."

In a letter to Falconer on October 1, 1862 (in F. Darwin, 1903, volume 1, p. 206), Darwin explicitly addressed this passage in Falconer's text. (Darwin had received an advance copy of the manuscript, along with Falconer's request for review and criticism—hence Darwin's reply, in 1862, to a text not printed until the following year): "To return to your concluding sentence: far from being surprised, I look at it as absolutely certain that very much in the Origin will be proved rubbish; but I expect and hope that the framework will stand."

The statement that God (or the Devil, in some versions) dwells in the details must rank among the most widely cited intellectual witticisms of our time. As with many clever epigrams that spark the reaction “I wish I'd said that!”, attribution of authorship tends to drift towards appropriate famous sources. (Virtually any nifty evolutionary saying eventually migrates to T. H. Huxley, just as vernacular commentary about modern America moves towards Mr. Berra.) The apostle of modernism in architecture, Ludwig Mies van der Rohe, may, or may not, have said that "God dwells in the details," but the plethora of tiny and subtle choices that distinguish the elegance of his great buildings from the utter drabness of superficially similar glass boxes throughout the world surely validates his candidacy for an optimal linkage of word and deed.

Architecture may assert a more concrete claim, but nothing beats the extraordinary subtlety of language as a medium for expressing the importance of apparently trivial details. The architectural metaphors of Milan's cathedral, used by both Falconer and Darwin, may strike us as effectively identical at first read. Falconer says that the foundations will persist as Darwin's legacy, but that the superstructure will probably be reconstructed in a quite different style. Darwin responds by acknowledging Falconer's conjecture that the theory of natural selection will undergo substantial change; indeed, in his characteristically diffident way, Darwin even professes himself "absolutely certain" that much of the Origin's content will be exposed as "rubbish." But he then states not only a hope, but also an expectation, that the "framework" will stand.

We might easily read this correspondence too casually as a polite dialogue between friends, airing a few unimportant disagreements amidst a commitment to mutual support. But I think that this exchange between Falconer and Darwin includes a far more "edgy" quality beneath its diplomacy. Consider the different predictions that flow from the disparate metaphors chosen by each author for the Duomo of Milan—Falconer's "foundation" vs. Darwin's "framework." After all, a foundation is an invisible system of support, sunk into the ground, and intended as protection against sinking or toppling of the overlying public structure. A framework, on the other hand, defines the basic form and outline of the public structure itself. Thus, the two men conjure up very different pictures in their crystal balls. Falconer expects that the underlying evolutionary principle of descent with modification will persist as a factual foundation for forthcoming theories devised to explain the genealogical tree of life. Darwin counters that the theory of natural selection will persist as a basic explanation of evolution, -even though many details, and even some subsidiary generalities, cited within the Origin will later be rejected as false, or even illogical.

I stress this distinction, so verbally and disarmingly trivial at a first and superficial skim through Falconer's and Darwin's words, but so incisive and portentous as contrasting predictions about the history of evolutionary theory, because my own position—closer to Falconer than to Darwin, but in accord with Darwin on one key point—led me to write this book, while also supplying the organizing principle for the "one long argument" of its entirety. I do believe that the Darwinian framework, and not just the foundation, persists in the emerging structure of a more adequate evolutionary theory. But I also hold, with Falconer, that substantial changes, introduced during the last half of the 20th century, have built a structure so expanded beyond the original Darwinian core, and so enlarged by new principles of macroevolutionary explanation, that the full exposition, while remaining within the domain of Darwinian logic, must be construed as basically different from the canonical theory of natural selection, rather than simply extended.

Book Details
ISBN: 0-674-00613-5 | 1464 pages
Published: March 2002,
Publisher: Belknap, Harvard University Press
Cover design: Marianne Perlak
Cover photo: Snail (Mollusca) on Grass at Sunset
by Jan Tove Johansson

Back cover:

Book Reviews
  • Anonymous (2002). The grand view: A master of evolution takes a last long look at Darwin. The Economist (Dec. 5th): 82.
  • Ayala, Francisco (2005). On Stephen Jay Gould's monumental masterpiece. Theology and Science 3 (March): 97-118.
  • Barash, David P. (2002). Grappling with the Ghost of Gould. Human Nature 2 (9 July): 283-292.
  • Borrello, Mark E. (2004). Radicals and revolution. Stud. Hist. Phil. Biol. & Biomed. Sci. 35 (1): 209–216.
  • Brown, Andrew (2002). Adventures in Evolution. The Guardian.
  • DiMichele, William (2003). A season with Steve Gould. American Journal of Science 303 (March): 259-261.
  • Doughty, Howard A. (2005). The Structure of Evolutionary Theory. College Quarterly 8 (Winter):.
  • Dormann, Carsten (2002). Stephen Jay Gould's Structure Of Evolutionary Theory. Dept. of Biometry, Univ. of Freiburg.
  • Erwin, Douglas H. (2004). One very long argument. Biology and Philosophy 19 (1): 17-28.
  • Flannery, Tim (2002). A new Darwinism? New York Review of Books 49 (May 23): 52–54.
  • Futuyma, Douglas J. (2002). Stephen Jay Gould a la recherche du temps. Science 296 (April 26): 661-663.
  • Ghiselin, Michael T. (2002). An autobiographical anatomy. Hist. Philos. Life. Sci. 24 (2): 285-291.
  • Grantham, Todd (2004). Constraints and spandrels in Gould's Structure of Evolutionary Theory. Biology and Philosophy 19: 29-43.
  • Hengeveld, Rob (2003). Book Review: The Structure of Evolutionary Theory. Acta Biotheoretica 51 (1): 67-72.
  • Hull, David L. (2002). A Career in the Glare of Public Acclaim. BioScience 52 (Sept.): 837-841.
  • Jablonski, David (2002). A more modern synthesis. American Scientist 90 (July–August): 368-371.
  • Korthof, Gert (2004). Stephen Jay Gould as a critic of orthodox Neo-Darwinism.
  • McGarr, Paul (2003). Revolutions in evolution. International Socialism Journal 100 (Autumn): 81-112.
  • McShea, Daniel (2004). A revised Darwinism. Biology and Philosophy 19 (1): 45-53.
  • Michael, Michaelis (2003). S.J. Gould's Last Words. Metascience 12 (2): 214-216.
  • Monastersky, Richard (2002). Revising the book of life. The Chronicle, March 15, A14.
  • Orr, H. Allen (2002). The descent of Gould. The New Yorker (Sept. 30): 132.
  • Perlman, David (2002). A Darwinian leap. San Francisco Chronicle April 14, RV-1.
  • Padian, Kevin (2002). A magisterial valedictory. The Lancet 340 (Sept. 28): 1027.
  • Padian, Kevin (2003). Evolution's past and future. Comptes Rendus Palevol 2 (6): 335–352.
  • Quammen, David (2003). The man who knew too much. Harper's Magazine (June): 73–80.
  • Radick, Gregory (2003). The Exemplary Kuhnian: Gould's Structure Revisited. Hist. Stud. in the Nat. Sciences 42 (Apr.): 143-157.
  • Raymo, Chet (2002). Gould's last book is fitting epitaph. Boston Globe (May 28): C6.
  • Ridley, Mark (2002). Stephen Jay Gould wants an evolution revolution. New York Times, March 17, sect. 7, col. 1, p. 11.
  • Ruse, Michael (2003). The Structure of Evolutionary Theory. Isis 94 (2): 397.
  • Shermer, Michael (2002). Grand design. Washington Post April 14, BW04.
  • Stearns, Stephen (2002). Less would have been more. Evolution 56 (11): 2339-45.
  • Sterelny, Kim (2003). Last will and testament. Philosophy of Science 70 (2): 255-263.
  • Topper, David (2003). Structure of Evolutionary Theory (review). Leonardo 36 (April): 157-159.
  • Turner, John R.G. (2002). Toe-breaker or epoch-maker? The Spectator June 29.
  • Wake, David B. (2002). A few words about evolution. Nature 416 (April 25): 787-788.
  • Wolpoff, Milford H. (2003). What Is Evolution? Evolutionary Anthropology 12 (1): 53–55.
  • Zimmerman, William F. (2003). Stephen Jay Gould's final view of evolution. Quarterly Review of Biology 78 (4): 454-459.