1020 THE STRUCTURE OF EVOLUTIONARY THEORY
But a close analysis of Dawkins's and Dennett's arguments exposes the
parochiality of their judgment. They regard punctuated equilibrium as trivial because
our theory doesn't speak to the restricted subset of evolutionary questions that, for
them, defines an exclusive domain of interest for the entire subject. These men
virtually equate evolution with the origin of intricately adaptive organic design—
"organized adaptive complexity," or O.A.C. in Dawkins's terminology. They then
dismiss punctuated equilibrium on the narrow criterion: "if it doesn't explain the
focus of my interests, then it must be trivial." Dawkins (1984, p. 684), for example,
properly notes the implications of punctuated equilibrium for validation of higher-
level selection, but then writes: "Species-level selection can't explain the evolution of
adaptations: eyes, ears, knee joints, spider webs, behavior patterns, everything, in
short, that many of us want a theory of evolution to explain. Species selection may
happen, but it doesn't seem to do anything much." "Everything"? Does nothing else
but adaptive organismal design excite Dawkins's fancy in the entire and maximally
various realm of evolutionary biology and the history of life—the "endless forms
most beautiful and most wonderful" of Darwin's closing words (1859, p. 490).
But the truly curious aspect of both Dawkins's and Dennett's charge lies in their
subsequent recognition, and fair discussion, of the important theoretical implication
of punctuated equilibrium: the establishment of species as Darwinian individuals, and
the consequent validation of species sorting and selection as a prominent process in a
hierarchical theory of Darwinian evolution. In 1984, Dawkins acknowledged that this
aspect of punctuated equilibrium "does, in a sense, move outside the neo-Darwinian
synthesis, narrowly interpreted. This is about whether a form of natural selection
operates at the level of entire lineages, as well as at the level of individual
reproduction stressed by Darwin and neo-Darwinism." In his 1986 book, Dawkins
then devotes a substantial part of the chapter following his rejection of punctuated
equilibrium to an evaluation of species selection. But he finishes his exploration by
reimmersion in the same parochial trap of denying importance because the
phenomenon doesn't explain his exclusive interest in adaptive organismal design: "To
conclude the discussion of species selection, it could account for the pattern of
species existing in the world at any particular time. It follows that it could also
account for changing patterns of species as geological ages give way to later ages,
that is, for changing patterns in the fossil record. But it is not a significant force in the
evolution of the complex machinery of life ... As I have put it before, species
selection may occur but it doesn't seem to do anything much!" (Dawkins, 1986, pp.
268 - 269). But doesn't "the pattern of species existing in the world at any particular
time" and "changing patterns in the fossil record" represent something of
evolutionary importance?
At the end of his long riff against punctuated equilibrium, Dennett also pauses
for breath and catches a glimmer of the concept that seems important and
theoretically intriguing to many students of macroevolution (Dennett, 1995, pp. 297-
298):