The Structure of Evolutionary Theory

(Michael S) #1

Punctuated Equilibrium and the Validation of Macroevolutionary Theory 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 fastest 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
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