David M. Raup, 1933-2015

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.

David M. Raup, 1933-2015

David M. Raup (1933-2015) in 1981.

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.”

Raup's (1966) morphological space-- the

Raup’s (1966) morphological space– “The Museum of All Shells”.

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.

A 26 million year periodicity of mass extinction (still debated) from Raup and Sepkoski (1980).

A 26 million year periodicity of mass extinction (still debated) from Raup and Sepkoski (1984).

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.

Principles of Paleontology (2nd edition, 1978).

Principles of Paleontology (2nd edition, 1978).

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).


  1. Frank
    Posted July 16, 2015 at 5:03 pm | Permalink

    I had to use that Moore et al. book in a Paleozoology class in 1973. Of course it was daunting to a freshman biology major, and not terribly interesting. When I started grad school, I recall that it was indeed interesting to see new hypotheses in paleobiology inform evolutionary biology, even if the statistical support for the periodicity argument was not that clear.

  2. Mark Sturtevant
    Posted July 16, 2015 at 5:24 pm | Permalink

    A very interesting post. I used to teach about the morphospace of molluscs, using a modified form of that figure.

  3. Posted July 16, 2015 at 5:24 pm | Permalink

    He died July 9th, not June 9th.

  4. Diane G.
    Posted July 16, 2015 at 5:48 pm | Permalink

    Thank you, very helpful post about someone I’d heard of but wasn’t sure quite why.

  5. Posted July 16, 2015 at 7:09 pm | Permalink

    Really nice post. I’d like to also point out that in addition to being a brilliant scientist–maybe the most brilliant and original paleontologist since G. G. Simpson–Dave was a really great guy. Perhaps not the most outgoing, but a droll sense of humor and very generous with other scientists and students. For such a decorated scientist–Schuchert Award, Paleontological Society Medal, National Academy of Sciences–he had virtually no ego. A rare combination.

    82 years is a pretty good run, but this was still sudden and very sad for many of us. He’ll be missed but, I suspect, not soon forgotten.

    By the way, his two popular books are great reads. The Nemesis Affair is probably the best book about the extinction impact debates in the 1980s, and Extinction: Bad Genes or Bad Luck? is a really thought-provoking meditation on the role of mass extinction in evolution. Well worth reading still!

    • Posted July 16, 2015 at 7:35 pm | Permalink

      Sorry to hear this news – thanks for posting it. David Raup was brilliant and creative. For many of us, Raup’s book on Extinction was his most influential popular work.
      Before 1980, the idea that a meteor killed the dinosaurs was greeted with skepticism. Actually, “horror and disbelief” according to David Raup, who added, “It was like suggesting that the dinosaurs had been shot by little green men from a spaceship.”
      Raup was 60 years old when “Extinction:Bad Genes or Bad Luck” was released in 1992. He retired shortly after. But he stood in favour of the notion of mass extinction via asteroid/comet impacts. This made him one of the first renowned palaeontologists to agree with the theory that space bombardment plays a role in extinctions.
      If anyone should like a brief recap of Raup’s creative-thinking side (including his endorsement of Nemesis, the Sun’s presumed sister star)I wrote a bit about him a few months ago: http://mountainmystery.com/2014/10/03/the-bad-luck-of-extinction/

    • Posted July 16, 2015 at 10:04 pm | Permalink

      Thanks for these thoughts on Raup as a person. Although our paths crossed once or twice, I never knew him personally, and thus could not add anything in this regard.

      Readers interested in learning more about the influence of Raup and the other “young Turks” (as I called them above) in changing the course of paleontology will want to read David’s Rereading the Fossil Record (2012, U. Chi. Press); the last chapter is entitled “Paleontology at the High Table?” (David is, btw, the son of Jack Sepkoski.)


    • Dominic
      Posted July 17, 2015 at 3:28 am | Permalink

      I read his two popular books, Extinction: Bad Genes or Bad Luck? (1992) & The Nemesis Affair: A Story of the Death of Dinosaurs and the Ways of Science (1999). I honestly cannot recall the latter one now but the former was very interesting – not a long book.

      Great History of Science post.

  6. bacopa
    Posted July 16, 2015 at 7:26 pm | Permalink

    Raup was at least somewhat in support of the idea that our sun had a brown dwarf companion star with a long elliptical orbit. While this idea is clearly outside of his area of expertise, it is not a completely unfounded hypothesis. He covered the subject in the books The Nemesis Affair and Extinction: Bad Genes or Bad Luck?.

    The brown dwarf supposedly enters the solar system every 26 million years and knocks asteroids loose. While that kind of orbital period seems a little long to me, I am no expert on orbital dynamics. I presume that Raup consulted with physicists before publishing to determine it was possible, but he might not have.

    Oh well, a little minor crankery can be forgiven. And there are really no negative consequences to the Nemesis hypothesis, unlike Pauling’s vitamin C stuff.

    • Torbjörn Larsson, OM
      Posted July 17, 2015 at 4:50 am | Permalink

      It would be interesting to study the consequences of near or real pathological science. Of course, the jury on the consequences of the much more massive religious magic speculations is also out.

      Hopefully you are right, and the only negative consequence is a slightly increased mortality in needlessly stressed skeptics. =D

    • Posted July 17, 2015 at 4:13 pm | Permalink

      Actually, Dave was a big believer in Pauling’s regimen! He was a habitual chainsmoker (although he quit several years before he died) and enjoyed a martini or three most evenings, and claimed that vitamin C kept him healthy.

      Normally, I’d consider someone like that a crank. But Dave enjoyed taking unpopular positions just to see where they might lead–and you never quite knew when he was being tongue-in-cheek.

  7. Allen
    Posted July 17, 2015 at 4:29 am | Permalink

    I have the 2nd Ed. text at home. It was a constant companion in my B.Sc. days.

  8. Torbjörn Larsson, OM
    Posted July 17, 2015 at 4:47 am | Permalink

    Thank you, very interesting! Picking up an interest in a science area you have not studied often means skipping on the basics at first, and I had no context on Raup.

    As I have commented elsewhere on WEIT, I know him mostly for his and Sepkoski’s awful paper looking for patterns in mass extinctions.

    I wrote this back in 2010 as a response to a blog comment, when I was fresh interested in astrobiology and WISE started to turn up a null result on Nemesis (dwarf star gravitationally bound to our system):


    I don’t see any solid evidence for in Nemesis and/or periodic mass extinctions:

    1) There is no periodicity in the data, either modern or the original Raup-Sepkoski, as shown by simple autocorrelation:

    “Quantitatively, extinction rates in the Fossil Record 2 family data (3) and Sepkoski’s family and genus data (1, 2) are not correlated with themselves at any time lag (49), which is a necessary condition for periodicity to hold. That said, analyses of origination rates in all three datasets (49, 50) suggest short-term autocorrelation. However, the current dataset shows no autocorrelation in either kind of rate (Fig. S1), and a standard spectral analysis (Fig. S2) also suggests purely random variation through the time series (i.e., white noise).” [Alroy, PNAS, 2008.]

    I’m not aware of any criticism to this result.

    2) This is an old contentious area with no consensus.

    3) I have looked at Raup and Sepkoski 1986 paper, which I assume contains much the same analysis as the 1984. It is terrible.

    Among other things they introduce Fourier-techniques, but does not test against null data by other filters than their own peak detector _which admit any sampled data local maximum in the noise_. Noise in, noise out.

    In any case, no periodicity means no Nemesis as envisioned.

    [ http://www.universetoday.com/69136/wise-mission-completes-all-sky-infrared-survey/#comment-82860 ; edited and much pruned of irrelevancies re Raup]


    Trying to remember the criticism now is of course as so often coming back with just a vaguely familiar feeling. I can say this though, as a physicist by training and publication, I am convinced of the utility and soundness of autocorrelation methods, and skeptic of applying filters needlessly and without null hypotheses.

    [Coincidentally, I am not so fond of Schopf either because of a similar pattern search overreach re early fossils, heavily criticized by the late Brasier. (2015 seems a bad year for biology.) But of course there is no consensus there either.]

    So that was my only impression of him up to now. Glad to see the impressive width of his efforts, and I can better understand the interest in mass extinction results in him and from him.

    • Posted July 17, 2015 at 4:23 pm | Permalink

      Seriously, Torbjörn? You’re gonna post that bile on a thread dedicated to the memory of a recently-deceased person who many people admired and respected (and, additionally, where the son of one of the authors of that “terrible” paper has just posted)?

      Oh, wait–I forgot that every thread is really all about you and how smart you are. Stay classy.

      (By the way, both Sepkoski and Raup acknowledged that there were problems with some of the analysis of periodicity. Periodicity was, however, merely a side-note to their discovery of the signal for the “Big 5” mass extinctions in the past, which was a much more important result and, perhaps, a more appropriate note to remember the passing of Dave with.)

  9. Posted July 17, 2015 at 11:49 am | Permalink

    Nice to see the idea of a state space (the shell-space) coming up again.

    What are the prerequisites for the 2006 textbook?

    • Posted July 17, 2015 at 6:25 pm | Permalink

      I’m not sure– I don’t have the 2006 edition. Intro paleontology courses suffer from the fact that the students generally come from two very different backgrounds– geology or biology– and thus a course or textbook must decide how much of each of those important background subjects to include. My own undergraduate paleontology course (in which we used Moore, Lalicker, & Fischer), taught by Peter Bretsky (one of the less prominent ‘young Turks’), began with a recap of invertebrate zoology (which I already well knew), because almost all the students were geology majors, and thus I had to learn about stratigraphy, sedimentation, etc. on my own.

      You could ask Mike Foote, and tell him I sent you.


  10. Raskos
    Posted July 17, 2015 at 2:15 pm | Permalink

    Was sorry to see this. My forays into morphospace have been restricted to the empirical rather than the theoretical, but I still remember being gobsmacked by the elegance and simplicity of Raup’s shell space when I was in graduate school.

One Trackback/Pingback

  1. […] Why Evolution is True: David M. Raup, 1933–2015 […]

%d bloggers like this: