Gould began his higher education at
Antioch College, graduating with a double major in geology and philosophy in 1963. During this time, he also studied at the
University of Leeds in the United Kingdom. After completing graduate work at
Columbia University in 1967 under the guidance of
Norman Newell, he was immediately hired by
Harvard University, where he worked until the end of his life (1967–2002). In 1973, Harvard promoted him to professor of geology and
curator of
invertebrate paleontology at the institution's
Museum of Comparative Zoology. In 1983, he was awarded a fellowship at the
American Association for the Advancement of Science, where he later served as president (1999–2001). The AAAS news release cited his "numerous contributions to both scientific progress and the public understanding of science". He also served as president of the
Paleontological Society (1985–1986) and of the
Society for the Study of Evolution (1990–1991).)
Punctuated equilibrium stability followed by episodic bursts of evolutionary change via rapid cladogenesis. It is contrasted (below) to
phyletic gradualism, a more gradual, continuous model of evolution. Early in his career, Gould and his colleague
Niles Eldredge developed the theory of
punctuated equilibrium, which describes the rate of speciation in the fossil record as occurring relatively rapidly, which then alternates to a longer period of evolutionary stability. According to Gould, punctuated equilibrium revised a key pillar "in the central logic of
Darwinian theory". it merely modified
neo-Darwinism in a manner that was fully compatible with what had been known before. Other biologists emphasize the theoretical novelty of punctuated equilibrium, and argued that evolutionary stasis had been "unexpected by most evolutionary biologists" and "had a major impact on paleontology and evolutionary biology". Comparisons were made to
George Gaylord Simpson's work in
Tempo and Mode in Evolution (1941), in which he also illustrated relatively sudden changes along evolutionary lines. Simpson describes the paleontological record as being characterized by predominantly gradual change (which he termed horotely), although he also documented examples of slow (bradytely), and rapid (tachytely) rates of evolution. Punctuated equilibrium and phyletic gradualism are not mutually exclusive (as Simpson's work demonstrates), and examples of each have been documented in different lineages. The debate between these two models is often misunderstood by non-scientists, and according to
Richard Dawkins has been oversold by the media. Some critics jokingly referred to the theory of punctuated equilibrium as "evolution by jerks", which prompted Gould to describe
phyletic gradualism as "evolution by creeps".
Evolutionary developmental biology Gould made significant contributions to
evolutionary developmental biology, especially in his work
Ontogeny and Phylogeny.
Selectionism and sociobiology Gould was a champion of
biological constraints, internal limitations upon developmental pathways, as well as other non-selectionist forces in evolution. Rather than direct
adaptations, he considered many higher functions of the
human brain to be the unintended
side consequence of
natural selection. Gould believed this feature of human mentality undermines an
essential premise of human
sociobiology and
evolutionary psychology.
Against Sociobiology In 1975, Gould's Harvard colleague
E. O. Wilson introduced his analysis of animal behavior (including human behavior) based on a sociobiological framework that suggested that many social behaviors have a strong evolutionary basis. In response, Gould,
Richard Lewontin, and
others from the Boston area wrote the subsequently well-referenced letter to
The New York Review of Books entitled, "Against 'Sociobiology'". This
open letter criticized Wilson's notion of a "deterministic view of human society and human action". But Gould did not rule out sociobiological explanations for many aspects of animal behavior, and later wrote: "Sociobiologists have broadened their range of selective stories by invoking concepts of
inclusive fitness and
kin selection to solve (successfully I think) the vexatious problem of
altruism—previously the greatest stumbling block to a Darwinian theory of social behavior... Here sociobiology has had and will continue to have success. And here I wish it well. For it represents an extension of basic Darwinism to a realm where it should apply."
Spandrels and the Panglossian paradigm in Fulnek,
Czech Republic With Richard Lewontin, Gould wrote an influential 1979 paper entitled, "
The Spandrels of San Marco and the Panglossian Paradigm", which introduced the architectural term "
spandrel" into evolutionary biology. In architecture, a spandrel is a triangular space which exists over the
haunches of an arch. Spandrels—more often called
pendentives in this context—are found particularly in classical architecture, especially Byzantine and Renaissance churches. When visiting
Venice in 1978, Gould noted that the spandrels of the
San Marco cathedral, while quite beautiful, were not spaces planned by the architect. Rather the spaces arise as "necessary architectural byproducts of mounting a dome on rounded arches." Gould and Lewontin thus defined "
spandrels" in the evolutionary biology context to mean any biological feature of an organism that arises as a necessary side consequence of other features, which is not directly selected for by natural selection. Proposed examples include the "masculinized genitalia in female
hyenas, exaptive use of an
umbilicus as a brooding chamber by snails, the shoulder hump of the giant
Irish deer, and several key features of human mentality". In
Voltaire's
Candide,
Dr. Pangloss is portrayed as a clueless scholar who, despite the evidence, insists that "all is for the best in this best of all possible worlds". Gould and Lewontin asserted that it is
Panglossian for evolutionary biologists to view all traits as atomized things that had been naturally selected for, and criticised biologists for not granting theoretical space to other causes, such as phyletic and developmental
constraints. The relative frequency of spandrels, so defined, versus adaptive features in nature, remains a controversial topic in
evolutionary biology. An illustrative example of Gould's approach can be found in
Elisabeth Lloyd's case study suggesting that the female orgasm is a by-product of shared developmental pathways. Gould also wrote on this topic in his essay "Male Nipples and Clitoral Ripples," prompted by Lloyd's earlier work. Gould was criticized by philosopher
Daniel Dennett for using the term spandrel instead of pendentive, a spandrel that curves across a right angle to support a dome. Robert Mark, a professor of civil engineering at Princeton, offered his expertise in the pages of
American Scientist, noting that these definitions are often misunderstood in
architectural theory. Mark concluded, "Gould and Lewontin's misapplication of the term spandrel for pendentive perhaps implies a wider latitude of design choice than they intended for their analogy. But Dennett's critique of the architectural basis of the analogy goes even further astray because he slights the technical rationale of the architectural elements in question." In a review of
Full House,
Richard Dawkins approved of Gould's general argument, but suggested that he saw evidence of a "tendency for lineages to improve cumulatively their adaptive fit to their particular way of life, by increasing the numbers of features which combine together in adaptive complexes. ... By this definition, adaptive evolution is not just incidentally progressive, it is deeply, dyed-in-the-wool, indispensably progressive."
Cultural evolution Gould's arguments against progress in evolutionary biology did not extend towards a notion of progress in general or notions of
cultural evolution. In
Full House, Gould compares two notions of progress against one another. While the first concept of progress, evolutionary progress, is argued to be invalid for a number of biological considerations, Gould permits that evolution may operate in human cultural evolution through a
Lamarckian mechanism. Gould goes on to argue that the disappearance of the 0.400
batting average in baseball is paradoxically due to the inclusion of better players in the league, rather than players becoming worse over time. In his view such a process is likely reflective in a number of cultural phenomena including sports, the visual arts, and music where, unlike in biological systems, the realm of aesthetic possibilities is constrained by a "right wall" of human limits and aesthetic preferences. Gould later goes on to state that his arguments for biological evolution should not be applied to cultural change lest they be employed by, "so-called 'political correctness' as a doctrine that celebrates all indigenous practice, and therefore permits no distinctions, judgements, or analyses." Around this time cladistics rapidly became the dominant method of classification in evolutionary biology. Inexpensive but increasingly powerful personal computers made it possible to process large quantities of data about organisms and their characteristics. Around the same time the development of effective
polymerase chain reaction techniques made it possible to apply cladistic methods of analysis to biochemical and genetic features as well.
Technical work on land snails ,
the Bahamas Most of Gould's empirical research pertained to
land snails. He focused his early work on the
Bermudian genus
Poecilozonites, while his later work concentrated on the
West Indian genus
Cerion. According to Gould "
Cerion is the land snail of maximal diversity in form throughout the entire world. There are 600 described species of this single genus. In fact, they're not really species, they all interbreed, but the names exist to express a real phenomenon which is this incredible morphological diversity. Some are shaped like golf balls, some are shaped like pencils. ... Now my main subject is the evolution of form, and the problem of how it is that you can get this diversity amid so little genetic difference, so far as we can tell, is a very interesting one. And if we could solve this we'd learn something general about the evolution of form." Given
Cerion extensive geographic diversity, Gould later lamented that if
Christopher Columbus had only catalogued a single
Cerion it would have ended the scholarly debate about which island Columbus had first set foot on in America.
Influence Gould is one of the most frequently cited scientists in the field of evolutionary theory. His 1979 "spandrels" paper has been cited more than 5,000 times. In
Paleobiology—the flagship journal of his own speciality—only
Charles Darwin and
George Gaylord Simpson have been cited more often. Gould was also a considerably respected historian of science. Historian
Ronald Numbers has been quoted as saying: "I can't say much about Gould's strengths as a scientist, but for a long time I've regarded him as the second most influential historian of science (next to
Thomas Kuhn)." Gould's undergraduate course, Science B-16: History of the Earth and Life, was taught in a Harvard Science Center lecture hall with a 250-seat capacity. Science B-16 was so oversubscribed that an annual lottery was held to see which students would be allowed to enroll in the course. If a student was denied course enrollment three times, then their fourth entry into the lottery provided them with a guaranteed seat in the class.
The Structure of Evolutionary Theory Shortly before his death, Gould published
The Structure of Evolutionary Theory (2002), a long treatise recapitulating his version of modern evolutionary theory. In an interview for the Dutch TV series
Of Beauty and Consolation Gould remarked, "In a couple of years I will be able to gather in one volume my view of how evolution works. It is to me a great consolation because it represents the putting together of a lifetime of thinking into one source. That book will never be particularly widely read. It's going to be far too long, and it's only for a few thousand professionals—very different from my popular science writings—but it is of greater consolation to me because it is a chance to put into one place a whole way of thinking about evolution that I've struggled with all my life."
As a public figure Gould became widely known through his popular essays on
evolution in the
Natural History magazine. His essays were published in a series entitled
This View of Life (a phrase from the concluding paragraph of
Charles Darwin's
Origin of Species) from January 1974 to January 2001, amounting to a continuous publication of 300 essays. During production, the only phrase Gould objected to was a line in the script that introduced him as the "world's most brilliant paleontologist". In 2002, the show paid tribute to Gould after his death, dedicating the
season 13 finale to his memory. Gould had died two days before the episode aired. He devoted considerable time to fighting against
creationism,
creation science, and
intelligent design. Most notably, Gould provided expert testimony against the equal-time creationism law in
McLean v. Arkansas. Gould later developed the term "non-overlapping magisteria" (NOMA) to describe how, in his view, science and religion should not comment on each other's realm. Gould went on to develop this idea in some detail, particularly in the books
Rocks of Ages (1999) and ''The Hedgehog, the Fox, and the Magister's Pox
(2003). In a 1982 essay for Natural History'' Gould wrote: Gould also spoke out against creationist misuse of his work and theory, especially with respect to how his theory of punctuated equilibrium relates to the presence of transitional fossils or forms: An
anti-evolution petition drafted by the
Discovery Institute inspired the
National Center for Science Education to create a pro-evolution counterpart called "
Project Steve," which is named in Gould's honor. Since 2013, Gould has been listed on the Advisory Council of the
National Center for Science Education. In 2011 the executive council of the
Committee for Skeptical Inquiry (CSI) selected Gould for inclusion in CSI's "Pantheon of Skeptics" created to remember the legacy of deceased CSI fellows and their contributions to the cause of scientific skepticism. Gould also became a noted public face of science, often appearing on television. In 1984 Gould received his own
NOVA special on
PBS. Other appearances included interviews on
CNN's
Crossfire and
Talkback Live,
NBC's
The Today Show, and regular appearances on PBS's
Charlie Rose show. Gould was also a guest in all seven episodes of the Dutch talk series
A Glorious Accident, in which he appeared with his close friend
Oliver Sacks. In 1999, Gould became the first recipient of the
GSA Public Service Award. Gould was featured prominently as a guest in
Ken Burns's
PBS documentary
Baseball, as well as
PBS's Evolution series. Gould was also on the Board of Advisers to the influential
Children's Television Workshop television show
3-2-1 Contact, where he made frequent guest appearances. == The "Darwin Wars" ==