774 THE STRUCTURE OF EVOLUTIONARY THEORY
and rates required to impart such dominance to species-individuals in the course of
macroevolution. For now, and for empirically minded paleontologists, the study of
relative frequencies in entire faunas, rather than the extraction of apparently
idealized cases, should be pursued as a primary strategy of research.
Critics have sometimes stated that punctuated equilibrium rests upon
declaration rather than documentation. (Maynard Smith once compared the theory
to "Aunt Jobisca's" maxim about ancient verities "known" by folk wisdom a
priori.) We do indeed assert that working paleontologists know the fact of
dominant stasis in their bones—but this claim represents a fair consensus about the
history of a field, and does underscore a paradox of non-concordance between deep
practical knowledge and imposed theoretical expectation. We have never tried to
argue that such a "professional feeling" constitutes documentation for punctuated
equilibrium. As with all scientific theories, punctuated equilibrium will live or die
by concrete and quantifiable evidence. As with any good hypothesis, punctuated
equilibrium becomes operational when workable definitions can be provided for
key claims and expectations—in this case, for stasis, punctuation, and relative
frequency. Contrary to the impression of some critics who have not followed the
primary literature of paleobiology during the last 25 years, punctuated equilibrium
has proven its fruitfulness and operational worth by being tested—and usually
confirmed, but sometimes confuted—in a voluminous literature of richly
documented cases (see pp. 822-874).
Microevolutionary Links
Eldredge and I coined the term-punctuated equilibrium in a paper first presented
(Gould and Eldredge, 1971) at a symposium entitled "Models in Paleobiology" at
the 1971 Annual Meeting of the Geological Society of America. T. J. M. Schopf,
the organizer of the symposium, conceived the enterprise as a tutorial in modern
evolutionary theory for professional invertebrate paleontologists. By accidents of
history, invertebrate paleontologists generally receive their advanced academic
degrees from geology departments, not from biology. Fossils became primary tools
for stratigraphic correlation long before the development of evolutionary theory,
and even before all scientists had accepted them as remains of ancient organisms!
Given traditions of narrowness in postgraduate education—particularly in Europe,
where students often attend no formal courses at all, and certainly no courses for
credit, outside the department that will grant their degree—most paleontologists,
before the present generation, did not receive any explicit training in evolutionary
biology, and could not articulate the basic concepts of population genetics or
theories of speciation. In paleontological usage, "evolution" designated little more
than the inferred pathway of phylogeny. This "little learning" often became the
"dangerous thing" of Alexander Pope's classic couplet, as paleontologists derived
their understanding of evolution from memories of old textbooks, or from shared
impressions amounting to little more than the blind leading the blind. This situation
has now changed dramatically—