Punctuated Equilibrium and the Validation of Macroevolutionary Theory 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