by Greg Mayer
David Raup, one of the leading figures in the return of paleontology to the “high table” of evolutionary biology in the late 20th century, died this past Thursday, July 9, at the age of 82. Raup attended the University of Chicago as an undergraduate, got his doctorate at Harvard, and was associated most prominently first with the University of Rochester, and then again the University of Chicago, from which he retired in 1995.
Beginning in the 1970s, paleontology was rejuvenated by a renewed interest in what the fossil record shows about both the broad scale patterns of changes in biodiversity through time, and the details of how particular lineages change through time. Raup was one of the most influential figures in this recrudescence, along with his colleagues Stephen Jay Gould, Tom Schopf, and Jack Sepkoski. The great British geneticist John Maynard Smith, said in 1984 of this flowering of paleontology, “The paleontologists have been too long missing from the high table. Welcome back.”
Raup’s most distinguished contributions came in two areas, both marked by a sophisticated, quantitative, approach. In the first, he made great strides in the area of theoretical morphology, developing mathematical descriptions of the possible shapes of mollusk shells, and then asking which parts of the morphological space defined by these equations are occupied, which are not occupied, and why. In a popular exposition based on Raup’s work, Richard Dawkins called this morphological space “The Museum of All Shells”. The mathematical description of shells that don’t exist (i.e. those that are in the parts of the morphological space not occupied) might seem odd or unnecessary, but understanding the possibilities of morphological transformation is key to understanding what it is that constrains, and what it is that enables or directs, evolutionary change. As A.S. Eddington put it, “We need scarcely add that the contemplation in natural science of a wider domain than the actual leads to a far better understanding of the actual.”
In the second, and more extensive, area of his distinguished contributions, Raup looked at levels of diversity, origination, and extinction through time in order to describe the pattern of these events and to model processes that could account for them. He was particularly interested in the relative influences of random versus deterministic factors in explaining the broad patterns in the history of diversification and extinction, notably detecting a periodicity in the history of mass extinctions.
In addition to his intellectual contributions, Raup had a significant effect on the institutional development of the field. In 1975, he was one of the founding members of the editorial board of Paleobiology, a journal dedicated to advancing, and a marker of, paleontology’s growth and renewed influence in the broader discipline of evolutionary biology. He had two papers in the inaugural issue, one coauthored with Schopf, Gould, and Daniel Simberloff. His other major contribution to the institutional development of the discipline was the publication, with Steven M. Stanley, of the influential textbook, Principles of Paleontology (1971; second edition 1978). Unlike previous paleontological textbooks, Principles had nary a key for identifying fossils or a compilation of taxa and their geological distributions: it was about the principles: systematics, biostratigraphy, paleoecology, evolution, and biogeography. On my own bookshelf, it sits inches away from an earlier influential text– Moore, Lalicker, and Fischer’s Invertebrate Fossils— which has a lonely chapter on principles, followed by 22 chapters and 700 pages of dense taxonomic and morphological detail. Raup stood out from traditional paleontologists, even among his fellow young Turks, for doing little or no descriptive systematic and stratigraphic work– even Gould had a long (and little-known) parallel publishing career on the systematic and zoogeographic nitty gritty of West Indian land snails of the genus Cerion— and his textbook reflects this.
The University of Chicago remained a hotbed of palebiology after Raup’s retirement, and his influence there is still strongly felt, with luminaries such as Dave Jablonski and Raup’s former student Mike Foote carrying on the tradition; Mike, with Arnold Miller of the University of Cincinnati, has brought out a third edition (2006) of Raup’s textbook.
h/t Bob Richards
Dawkins, R. 1996. Climbing Mount Improbable. W.W. Norton, New York.
Foote, M. and A.I. Miller. 2006. Principles of Paleontology. 3rd edition. W.H. Freeman, New York.
Maynard Smith, J. 1984. Palaeontology at the high table. Nature 309:401-402 pdf
Moore, R.C., C.G. Lalicker, and A.G. Fischer. 1952. Invertebrate Fossils. McGraw-Hill, New York.
Raup, D.M. 1966. Geometric analysis of shell coiling: general problems. Journal of Paleontology 40:1178-1190. pdf
Raup, D.M. and J.J. Sepkoski. 1984. Periodicity of extinctions in the geologic past. Proceedings of the National Academy of Sciences USA 81:801-805. pdf
Raup, D.M. and S.M. Stanley. 1971. Principles of Paleontology. W.H. Freeman, San Francisco. (2nd edition, 1978).