The Structure of Evolutionary Theory

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by Stephen Jay Gould


  So if this second ad hominem argument won't even wash for a presumably naive press, how can colleagues regard the attention of a far more sophisti­cated professional community as nothing but a spinoff from our hype and rhetoric?

  When this charge has been laid against me, cited evidence almost always rests upon two supposed claims (and their canonical quotations) expressed in my putatively most radical paper of 1980, entitled: “Is a new and general the­ory of evolution emerging?” I wrote this paper for the 5th anniversary of Paleobiology, as a companion piece to a longer analysis of biological research in our profession: “The promise of paleobiology as a nomothetic, evolution­ary discipline” (see Gould, 1980a and b).

  The received legend about this paper — I really do wonder how many col­leagues have ever based their comments on reading this article with any care, or even at all — holds that I wrote a propagandistic screed featuring two out­rageously exaggerated claims: first, the impending death of the Modern Syn­thesis; and second, the identification of punctuated equilibrium as the exter­minating angel (or devil). I do not, in fact and in retrospect (but not in understatement), regard this 1980 paper as among the strongest, in the sense of most cogent or successful, that I have ever written — but neither do I reread it with any shame today. Some of my predictions have fared poorly, and I would now reject them — scarcely surprising for a paper that tried to summa­rize all major theoretical revisions then under discussion among evolutionists. [Page 1003] For example, I then read the literature on speciation as beginning to favor sympatric alternatives to allopatric orthodoxies at substantial relative fre­quency, and I predicted that views on this subject would change substantially, particularly towards favoring mechanisms that would be regarded as rapid even in microevolutionary time. I now believe that I was wrong in this pre­diction.

  But the relatively short section devoted to punctuated equilibrium (Gould, 1980b, pp. 125-126) presents this subject in a standard and unsurprising manner, and I would not change any major statement in this part of the paper. (My reassessment away from high relative frequency for rapid speciation in microevolutionary time, and back to the peripatric orthodoxy of our original views, represents a rethinking of another section of this 1980 paper, and does not speak to the validity of punctuated equilibrium. As I have emphasized throughout this chapter, punctuated equilibrium was formulated as the ex­pected macroevolutionary expression of conventional allopatric speciation — so a return to this conventional model can scarcely threaten the theory's va­lidity!)

  The supposed general death of the Synthesis. Given the fu­ror provoked, I would probably tone down — but not change in content — the quotation that has come to haunt me in continual miscitation and misunder­standing by critics: “I have been reluctant to admit it — since beguiling is often forever — but if Mayr's characterization of the synthetic theory is accurate, then that theory, as a general proposition, is effectively dead, despite its per­sistence as textbook orthodoxy” (Gould, 1980b, p. 120). (I guess I should have written the blander and more conventional “due for a major reassess­ment” or “now subject to critical scrutiny and revision,” rather than “effec­tively dead.” But, as the great Persian poet said, “the moving finger writes, and having writ...” and neither my evident piety nor obvious wit can call back the line — nor would tears serve as a good emulsifier for washing out anything I ever wrote!)

  Yes, the rhetoric was too strong (if only because I should have anticipated the emotional reaction that would then preclude careful reading of what I ac­tually said). But I will defend the content of the quotation as just and accu­rate. First of all, I do not claim that the synthetic theory of evolution is wrong, or headed for complete oblivion on the ashheap of history; rather, I contend that the synthesis can no longer assert full sufficiency to explain evolution at all scales (remember that my paper was published in a paleobiological journal dedicated to studies of macroevolution). Two statements in the quotation should make this limitation clear. First of all, I advanced this opinion only with respect to a particular, but (I thought) quite authorita­tive, definition of the synthesis: “if Mayr's characterization of the synthetic theory is accurate.” Moreover, I had quoted Mayr's definition just two para­graphs earlier. The definition begins Mayr's chapter on “species and transspecific evolution” from his 1963 classic — the definition that paleobiologists would accept as most applicable to their concerns. Mayr wrote (as I explicitly quoted): “The proponents of the synthetic theory maintain that all evolution [Page 1004] is due to the accumulation of small genetic changes, guided by natural selec­tion, and that transspecific evolution is nothing but an extrapolation and magnification of the events that take place within populations and species.”

  Second, I talked about the theory being dead “as a general proposition,” not dead period. In the full context of my commentary on Mayr's definition, and my qualification about death as a full generality, what is wrong with my statement? I did not proclaim the death of Darwinism, or even of the strictest form of the Modern Synthesis. I stated, for an audience interested in macro-evolutionary theory, that Mayr's definition (not the extreme statement of a marginal figure, but an explicit characterization by the world's greatest expert in his most famous book) — with its two restrictive claims for (1) “all evolu­tion” due to natural selection of small genetic changes, and (2) transspecific evolution as “nothing but” the extrapolation of microevolutionary events — must be firmly rejected if macroevolutionary theory merits any independent status, or features any phenomenology requiring causal explanation in its own domain. If we embrace Mayr's definition, then the synthesis is “effec­tively dead” “as a general proposition” — that is, as a theory capable of providing a full and exclusive explanation of macroevolutionary phenomena. Wouldn't most evolutionary biologists agree with my statement today?

  Nonetheless, I was reviled in many quarters, and in prose far more intem­perate and personal than anything I ever wrote, for proclaiming the death of Darwinism, and the forthcoming enshrinement of my own theory as a re­placement (see, for example, A. Huxley, 1982; Thompson, 1983; Cain, 1988; Vogel, 1983; Ayala, 1982; Stebbins and Ayala, 1981a and b; Mayr, 1982a; and Grant, 1983, under the title: “The synthetic theory strikes back”).

  Many reasons underlie this error, and I do accept some responsibility for my flavorful prose (but not for any lack of clarity in intended meaning, or for any statement stronger than Mayr's dismissive words about my own profes­sion of macroevolution). One common reason, perhaps the most prominent of all, arises from careless scholarship and cannot be laid at my doorstep. I provided the full quotation that offended so many colleagues, along with Mayr's accompanying words, so necessary to grasp the definition that I used. But my statement is usually quoted in deceptively abridged form, leading to a false reading clearly opposite to what I intended. I usually find my words cited in the following abridgment: “The synthetic theory ... is effectively dead, despite its persistence as textbook orthodoxy.” Much commentary has been based upon this truncated and distorted version, not on my actual words. Fill in those three dots before you fire.

  Homo unius libri. An old and anonymous Latin proverb states: cave ab homine unius libri — beware the man of one book. I do appreciate the at­tention that punctuated equilibrium has received, and, as a fallible mortal, I am not adverse to the recognition that this debate has brought me. But as a curious and general consequence of extensive publicity for a single achieve­ment, the totality of one's work then tends to be read as a long and unitary commentary upon this singular idea or accomplishment. The Latin motto should therefore be read from both ends: we should be wary of a person who [Page 1005] has only one good idea, but we should also not automatically assimilate an entire life by synecdoche to the single aspect we know best. Leonardo's war machines bear little relationship to the Mona Lisa; Newton's chronology of ancient kingdoms never mentions gravity or the inverse square law; and Mickey Mantle was also the best drag bunter and faste
st runner in baseball.

  Perhaps I should be flattered by the implied importance thus accorded to punctuated equilibrium, but I do maintain interests, some just as consuming, and some (I hope) just as replete with implications for evolutionary theory. Critics generally complete their misunderstanding of my 1980 paper by first imagining that I proclaimed the total overthrow of Darwinism, and then sup­posing that I intended punctuated equilibrium as both the agent of destruc­tion and the replacement. But punctuated equilibrium does not occupy a ma­jor, or even a prominent, place in my 1980 paper.

  This article tried to present a general account of propositions within the Modern Synthesis that, in my judgment, might require extensive revision or enlargement, especially from the domain of macroevolution. I did speak ex­tensively — often quite critically — about the reviled work of Richard Goldschmidt, particularly about aspects of his thought that might merit a rehear­ing. This material has often been confused with punctuated equilibrium by people who miss the crucial issue of scaling, and therefore regard all state­ments about rapidity at any level as necessarily unitary, and necessarily flow­ing from punctuated equilibrium. In fact, as the long treatment in Chapter 5 of this book should make clear, my interest in Goldschmidt resides in issues bearing little relationship with punctuated equilibrium, but invested instead in developmental questions that prompted my first book, Ontogeny and Phylogeny (Gould, 1977b). The two subjects, after all, are quite separate, and rooted in different scales of rapidity — hopeful monsters in genuine saltation, and punctuated equilibrium in macroevolutionary punctuation (produced by ordinary allopatric speciation). I do strive to avoid the label of homo unius libri. I have even written a book about baseball, and another about calendrics and the new millennium.

  The section on punctuated equilibrium in my 1980 paper is both short in extent, and little different in content from my treatment of the subject else­where. I began with the usual definition: “Our model of 'punctuated equi­libria' holds that evolution is concentrated in events of speciation and that successful speciation is an infrequent event punctuating the stasis of large populations that do not alter in fundamental ways during the millions of years that they endure” (p. 125). I then made my usual linkage to ordinary allopatric speciation, not to any novel or controversial mechanism of microevolution. Moreover, I emphasized the scaling error that so often leads people to confuse punctuated equilibrium with saltationism:

  Speciation, the basis of macroevolution, is a process of branching. And this branching, under any current model of speciation — conventional allopatry to chromosomal saltation — is so rapid in geological translation (thousands of years at most compared with millions for the duration of [Page 1006] most fossil species) that its results should generally lie on a bedding plane, not through the thick sedimentary sequence of a long hillslope ... It [gradualism] represents, first of all, an incorrect translation of conven­tional allopatry. Allopatric speciation seems so slow and gradual in eco­logical time that most paleontologists never recognized it as a challenge to the style of gradualism — steady change over millions of years — pro­mulgated by custom as a model for the history of life (p. 125).

  Finally, I stressed that the radical implications of punctuated equilibrium lay in proposed explanations for such macroevolutionary phenomena as cladal trends, not in any proposal for altered mechanisms of microevolution: “Evo­lutionary trends therefore represent a third level superposed upon speciation and change within demes . . . Since trends 'use' species as their raw material, they represent a process at a higher level than speciation itself. They reflect a sorting out of speciation events . . . What we call 'anagenesis,' and often at­tempt to delineate as a separate phyletic process leading to 'progress,' is just accumulated cladogenesis filtered through the directing force of species selec­tion.”

  In tres partes divisa est: the 'urban legend' of punctu­ated equilibrium's threefold history. The opponents of punctu­ated equilibrium have constructed a fictional history of the theory, primarily (I suppose) as a largely unconscious expression of their hope for its minor im­portance, and their jealousy towards its authors. This history even features a definite sequence of stages, constructed to match a classic theme of Western sagas: the growth, exposure and mortification of hubris (try Macbeth as a prototype, but he dies before reaching the final stage of penance; so try Faust instead, who lusts for the world and ends up finding satisfaction in draining a swamp). This supposed threefold history of punctuated equilibrium also ranks about as close to pure fiction as any recent commentary by scientists has ever generated.

  In stage one, the story goes, we were properly modest, obedient to the theoretical hegemony of the Modern Synthesis, and merely trying to bring paleon­tology into the fold. But the prospect of worldly fame beguiled us, so we broke our ties of fealty and tried, in stage two, to usurp power by painting punctuated equilibrium as a revolutionary doctrine that would dethrone the Synthesis, resurrect the memory of the exiled martyr (Richard Goldschmidt), and reign over a reconstructed realm of theory. But we were too big for our breeches, and the old guard still retained some life. They fought back mightily and effectively, exposing our bombast and emptiness. We began to hedge, re­treat, and apologize, and have been doing so ever since in an effort to regain grace and, chastened in stage three, to sit again, in heaven or Valhalla, with the evolutionary elite.

  Such farfetched fiction suffers most of all from an internal construction that precludes exposure and falsification among true believers, whatever the evidence. Purveyors of this myth even name the three stages, thus solidifying the false taxonomy. Dawkins (1986), for example, speaks of the “grandiloquent [Page 1007] era ... of middle-period punctuationism [which] gave abundant aid and comfort to creationists and other enemies of scientific truth.” In the other major strategy of insulation from refutation, supporters of this “urban leg­end” about the modest origin, bombastic rise, and spectacular fall of punctu­ated equilibrium forge a tale that allows them to read any potential disconfirmation as an event within the fiction itself. (Old style gradualism pursued exactly the same strategy in reading contrary data as marks of imperfect evidence within the accepted theory — and thus could not be refuted from within. I am struck by the eerie similarity between the structure of the old theory and the historical gloss invented by opponents of a proposed replace­ment.)

  In particular, and most offensive to me, the urban legend rests on the false belief that radical, “middle-period” punctuated equilibrium became a saltational theory wedded to Goldschmidt's hopeful monsters as a mechanism. I have labored to refute this nonsensical charge from the day I first heard it. But my efforts are doomed within the self-affirming structure of the urban leg­end. We all know, for so the legend proclaims, that I once took the Goldschmidtian plunge. So if I ever deny the link, I can only be retreating from an embarrassing error. And if I continue to deny the link with force and gusto, well, then I am only backtracking even harder (into stage 3) and apologizing (or obfuscating) all the more. How about the obvious (and accurate) alterna­tive: that we never made the Goldschmidtian link; that this common error embodies a false construction; and that our efforts at correction have always represented an honorable attempt to relieve the confusion of others.

  But the urban legend remains too simplistically neat, and too resonant with a favorite theme of Western sagas, to permit refutation by mere evidence. So Dennett (1995, pp. 283-284) writes: “There was no mention in the first pa­per of any radical theory of speciation or mutation. But later, about 1980, Gould decided that punctuated equilibrium was a revolutionary idea after all ... [But] it was too revolutionary, and it was hooted down with the same sort of ferocity the establishment reserves for heretics like Elaine Morgan. Gould backpedaled hard, offering repeated denials that he has ever meant anything so outrageous.” And Halstead (1985, p. 318) wrote of me (with equal pov­erty in both logic and grammar): “He seems to be setting up a face-saving for­mula to enable him to retreat from his earlier
aggressive saltationism, having had a bit of a thrashing, his current tack is to suggest that perhaps we should keep the door open in case he can find some evidence to support his pet theo­ries so let us be 'pluralist.'”

  I do not, of course, claim that our views about punctuated equilibrium have never changed through the years of debate (only a dull and uninteresting theory could remain so static in the face of such wide discussion). Nor do I maintain a position that would be even sillier — namely, that we made no im­portant errors requiring corrections to the theory. Of course we made mis­takes, and of course we have tried to amend them. But I look upon the history of punctuated equilibrium (from my partisan vantage point of course) as a fairly standard development for successful theories in science. We did, indeed, [Page 1008] begin modestly and expand outward thereafter. (In this sense, punctuated equilibrium has grown in theoretical scope, primarily as macroevolutionary theory developed and became better integrated with the rest of evolutionary thought — and largely through articulation of the hierarchical model, as dis­cussed in the previous chapter).

  We started small as a consequence of our ignorance and lack of perspective, not from modesty of basic temperament. As stated before, we simply didn't recognize, at first, the interesting implications of punctuated equilibrium for macroevolutionary theory — primarily gained in treating species as Darwinian individuals for the explanation of trends, and in exploring the extent and causes of stasis. With the help of S. M. Stanley, E. S. Vrba and other col­leagues, we developed these implications over the years, and the theory grew accordingly. But we never proposed a radical theory for punctuations (ordi­nary speciation scaled into geological time), and we never linked punctu­ations to microevolutionary saltationism.

 

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