The Structure of Evolutionary Theory

(Michael S) #1

Punctuated Equilibrium and the Validation of Macroevolutionary Theory 10 09


claim. All attempts collapse upon close examination. Dennett, for example, who
insists (1997, p. 64) that "for a while he [Gould] had presented punctuated
equilibrium as a revolutionary 'saltationist' alternative to standard neo-Darwinism,"
documents his supposed best case by assuring readers (1995, p. 285) that "for a while,
Gould was proposing that the first step in the establishment of any new species was a
doozy—a non-Darwinian saltation." Dennett directly follows this claim with his
putative proof, yet another quotation from my 1980 paper, which he renders as
follows: "Speciation is not always an extension of gradual, adaptive allelic
substitution to greater effect, but may represent, as Goldschmidt argued, a different
style of genetic change—rapid reorganization of the genome, perhaps non-adaptive"
(Gould, 1980b, p. 119).
I regard Dennett's case as pitiful, but the urban legend can offer no better. First
of all, this quotation doesn't even refer to punctuated equilibrium, but comes from a
section of my 1980 paper on the microevolutionary mechanics of speciation.
Secondly, Dennett obviously misreads my statement in a backwards manner. I am
trying to carve out a small theoretical space for a style of microevolutionary rapidity
at low relative frequency—as clearly stated in my phrase "not always an extension of
gradual..." But Dennett states that I am proposing this mechanism as a general
replacement for gradual microevolutionary change in all cases of speciation—"the
first step in the establishment of any new species" in his words. But my chosen
phrase—"not always"— clearly means "most of the time," and cannot be read as
"never." In short, I made a plea for pluralism, and Dennett charges me with
usurpation. Then, when I try to explain, I am accused of beating a retreat to save face.
When placed in such a double bind, one can only smile and remember Schiller's
famous dictum: Mit Dummheit kdmpfen die Gotter selbst vergebens.
Finally, the claim that we equated punctuated equilibrium with saltation makes
no sense within the logical structure of our theory—so, unless we are fools, how
could we ever have asserted such a proposition? Our theory holds, as a defining
statement, that ordinary allopatric speciation, unfolding gradually at
microevolutionary scales, translates to punctuation in geological time.
Microevolutionary saltation also scales as punctuation—so the distinction between
saltation and standard allopatry becomes irrelevant for punctuated equilibrium, since
both yield the same favored result!
Moreover, the chronology of debate proves that we did not issue disclaimers on
this subject only to cover our asses as we retreated from exaggerations of our
supposed second phase, because we have been asserting this clarification from the
very beginning—that is, from the first paper we ever wrote to comment upon
published reactions to punctuated equilibrium. Our first response appeared in 1977,
long before we issued the supposed clarion call of our false revolution in 1980. We
wrote (Gould and Eldredge, 1977, p. 121), under the heading "Invalid claims of
gradualism made at the wrong scale": "The model of punctuated equilibria does not
maintain that nothing occurs gradually at any level of evolution. It is a theory about
speciation and its deployment in the fossil record. It claims that an important pattern,
continuous

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