Post by Greg Mayer: More thoughts on E.O. Wilson

January 8, 2022 • 1:20 pm

by Greg Mayer

As WEIT readers well know by now, E.O. Wilson died last month at the age of 92. Jerry knew Ed better than I did, but my scientific interests were actually much closer to his than were Jerry’s. I was greatly influenced by Ed and Robert MacArthur‘s theory of island biogeography. The Harvard Gazette just republished an interview with Ed from 2014, and the interviewer asked what he thought his most important contribution was. Ed said there were several, but his work with MacArthur is the first thing he mentioned.

My well-worn copy of MacArthur and Wilson, purchased in 1977, when I was an undergraduate.

I had learned about their ideas–especially the idea that the number of species on an island was the result of a dynamic equilibrium between ongoing colonization and extinction–in my undergraduate classes. I got a copy of their book and began reading it between my sophomore and junior years of college. When I went off to graduate school to work on island lizards with Ernest Williams, I expected that I would wind up applying and exemplifying the principles of the theory.

The title page of my copy, signed by Ed, with an added sketch of an ant; inscribed on 6 October 2007, at the “Island Biogeography at 40” symposium held at the Museum of Comparative Zoology (MCZ). I had brought my copy with me; Ed and I chatted, and he graciously signed it, adding the ant unbidden. The symposium resulted in a book edited by Jon Losos and Bob Ricklefs (reference below).

Ed’s work in island biogeography included theoretical formulation (though the mathematics was primarily MacArthur’s), analysis of faunal lists and distribution patterns, and, very ambitiously, experimental manipulation of species numbers on mangrove islets in Florida. One of his contributions to island biogeography that has gone largely unremarked, but which I regard as of real significance, are his studies of Cenozoic fossil ant faunas in amber from the Dominican Republic. He found that the ancient amber ant fauna was more characteristic of a continental fauna, rather than that of an oceanic island. I was very impressed by this work; at the time it was, and remains to this day, among the strongest evidence that the Greater Antilles are old continental islands– islands once attached to or closely adjacent to the mainland, from which their fauna derives.  (Several of his papers on the amber ants are open access, and are linked below.)

Ed gave a seminar about his amber fossil work while I was a grad student, and we talked afterwards, and he suggested we continue the discussion. Knowing how busy Ed was, I went to make an appointment to see him with his secretary, rather than rely on us passing in the hallway to set a time (Ed was not often in the hallway). His secretary was very suspicious about who I was and what I wanted, and said I would need to write a letter requesting a meeting. The demand seemed to me beyond the pale, considering that I was a grad student in the MCZ and that Ed wanted the meeting. I never wrote the letter, and that conversation never continued.

A few years later, in 1988, James D. ‘Skip’ Lazell and I wrote that the discovery of bolitoglossine salamanders (a mainland group, with little capacity to cross a salt water barrier) in the Dominican amber would be powerful evidence from the vertebrate fossil record for an old continental origin of the Greater Antillean fauna (Wilson’s work having provided invertebrate evidence). In 2015 George Poinar and Dave Wake reported exactly that– a well-preserved bolitoglossine salamander from the Dominican amber!

Many people (including Ed) have commented on Ed’s rather sour relationship with Dick Lewontin. What is sometimes overlooked in these discussions is that both of them were involved with a group known as the “Marlboro Circle”. This group, which sought to conceptually unify ecology and evolutionary biology, included, among others, Ed’s key collaborator, Robert MacArthur, and also Richard Levins. (Levins was a Marxist who made important contributions to island biogeography, collaborated on publications with MacArthur and Lewontin, and visited Wilson’s experimental mangrove islands.)

I find the activities of this group fascinating– in many ways they set the agenda for ecology and evolutionary biology for the second half of the 20th century. It is one of the most important episodes in the biology of that century–that the group was marked by tragedy (MacArthur’s death at 42) and a falling out among its members lends pathos as well. The episode has, unfortunately, been little studied or remarked upon by historians of science. My last communication with Ed (in 2020) was about some of the sources for the data that went into the origination of his and MacArthur’s formulation of their theory.

I mentioned above that I had expected my studies to apply and exemplify the theory of island biogeography, but as I learned more about reptile distribution and did my own studies in the West Indies, I found that species number on islands was not the result of a dynamic equilibrium between ongoing colonization and extinction. Rather medium (10^4 to 10^5 ka, especially ice age changes in sea level) and long term (10^6 to 10^7 ka; the origin and diversification of major lineages) geological and evolutionary events were much more important in shaping the fauna.

Nonetheless, MacArthur and Wilson, like Darwin in the Origin and Mendel in his seminal paper, had limned many pathways for further progress, and despite failing to find evidence of their equilibrium process, I had, and continue to have, the greatest respect for their work. For my thesis defense, I planned to bring two past members of the Marlboro Circle– Dick Lewontin, my advisor, and Ed Wilson, original island biogeographer– back together, at least briefly.

My thesis defense talk was entitled “A theory of island biogeography, with especial reference to the amphibians and reptiles of the West Indies”. (This was a deliberate mashup of the title of Ed’s book and the title of a monograph from 1914 by Thomas Barbour, an early curator of herpetology at the MCZ, “A contribution to the zoogeography of the West Indies, with especial reference to amphibians and reptiles“.) I gave the talk at Dick’s weekly Population Biology Seminar on the 3rd floor of the MCZ Labs, one floor below Ed’s lab.

I tried to make sure that Ed would attend, despite its location, by placing a copy of my seminar announcement in his mailbox. He did come, and sat at the middle of one side of the great table that sat in the middle of Dick’s lab (Dick was further back in the room). I argued in my talk that the equilibrium theory was a special case of a more general theory, and that the equilibrium theory per se didn’t apply very well to West Indian amphibians and reptiles.

After my talk, among the questions were one or two from Ed. He defended the applicability of his and MacArthur’s theory to broader situations than the ones where it fit best, and, indeed, I concur that they had anticipated modifications, expansions, and refinements that would improve it– that’s why I had said there was a more general theory of which theirs could be a special case. After the questions, Ed left.

Although much of Ed’s public reputation rests, rightly, on Ed’s advocacy for biodiversity, and on the controversy over sociobiology (which accounts for essentially all the negative bits), in remembering Wilson we should not lose sight of his other accomplishments.

Losos, J.B. and R.E. Ricklefs, eds. 2010. The Theory of Island Biogeography Revisited. Princeton University Press, Princeton, N.J. publisher

MacArthur, R.H. and E.O. Wilson. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton, New Jersey. publisher

Wilson, E.O. 1985. Ants of the Dominican amber (Hymenoptera: Formicidae). 1. Two new myrmicine genera and an aberrant Pheidole. Psyche 92:1-9. pdf

Wilson, E.O. 1985. Ants of the Dominican amber (Hymenoptera: Formicidae). 2. The first fossil army ants. Psyche 92:11-16. pdf

Wilson, E.O. 1985. Ants of the Dominican amber (Hymenoptera: Formicidae). 3. The subfamily Dolichoderinae. Psyche 92:17-37. pdf

The number of species on islands

July 27, 2020 • 1:00 pm

by Greg Mayer

[The following is a trivial, and speculative, discussion about a small part of an important paper in the history of ecology and evolutionary biology.]

There are fewer species of any given group of plants or animals on an island than on an equivalent area of the mainland; and the larger the island, the more species there will be. These two general rules of natural history have been known since at least the 19th century, and are known under the rubrics that island biota are depauperate, and the species-area relationship, respectively. There have been many, not always mutually exclusive, explanations for these phenomena, and in the early 1960s they were on the minds of at least several biologists.

The most important resulting paper was Robert MacArthur and Ed Wilson‘s classic “An equilibrium theory of insular zoogeography.” This paper, together with the expansion of the ideas contained within it into a book-length monograph, The Theory of Island Biogeography (1967), were extremely influential in setting out the questions to be asked, and how to go about answering them, across many areas of ecology and evolutionary biology, not just the phenomena of island life.

MacArthur and Wilson (1963).

In the paper, MacArthur and Wilson proposed (among other things) that the number of species on an island resulted from a dynamic equilibrium between ongoing immigration and extinction of species living on the island, and that the relation of these demographic processes to various physical properties of islands led to the species-area relationship. They illustrated this relationship in two figures showing the relationship between the area of an island or island group and the number of species of land and freshwater birds occurring on that island or island group. One figure was for islands in and near the Sunda group, the other for the Moluccas, Melanesia, and Polynesia. Both figures are interesting, but the second one concerns us today.

Figure 2 of MacArthur and Wilson (1963).

For reasons relating to some research projects I’m pursuing during the pandemic, I was led to look closely at this figure, including the sources of the data as indicated in the legend of the figure: three papers by Ernst Mayr, and a book by James C. Greenway. I have a copy of Greenway’s book (both editions, actually), and Mayr’s 1943 paper is readily available online to anyone from SORA (a wonderful free repository containing vast swaths of the ornithological literature).

Neither of the two earlier papers are readily available, but the University of Wisconsin, Madison, Library scanned a copy of the earliest, and sent me a pdf (kudos to the staff of the Library there for working very hard during the pandemic to keep the scholarly literature available); and I happened to have a reprint of the other, a paper from the modestly obscure Proceedings of the Sixth Pacific Science Congress. Mayr published two papers in the proceedings, and I have reprints of both; the second is the one cited by MacArthur and Wilson.

Mayr (1940a).


Mayr (1940b).

The provenance of my copies is of interest. By looking at the stamps and annotations, you can see that the copies were originally in the library of the Bird Department of the Museum of Comparative Zoology, which stamped them, and added the author and date notation at upper left. (Departments of major museums usually maintain libraries physically separate from the main library of the institution.) Reprints were the principal way in which scientific literature circulated in the 1940s, and it is most likely that the Bird Department obtained the copies shortly after publication, not when Mayr moved to the MCZ in 1953. As duplicates (and Mayr’s arrival may be why the Department had extra copies), they then passed into the possession of my friend and colleague Bob O’Hara; we were graduate students together in the 1980s, and he worked closely with Mayr. At some point, also in the 1980s, Bob gave them to me, and I penciled my name and a correction to the date of publication on them. (See Note on the date of publication below.)

So now we come to the matter at hand. I was trying to track down the actual numbers that went into MacArthur and Wilson’s Figure 2, and some of the data came from Mayr (1940b), the second of the proceedings papers. It’s a 20 page paper, with no table of species numbers by islands, so I was reading it carefully to find what numerical data I could. It’s all on page 202– species numbers for 11 islands or island groups that were included in Figure 2. And a little later, while copying the numbers on to a data sheet, I noticed a pencil mark next to the paragraph with the data. (It is the only mark in the paper, aside from those on the first page.)

A pencil mark highlighting species number data in Mayr (1940b).

The mark was not mine; and it would not be Bob O’Hara’s, who would not have been interested in the particular questions addressed by these data. So who would have been using the MCZ Bird Department Library’s copy prior to the 1980s to highlight data included in MacArthur and Wilson (1963)? It occurred to me, could it have been Ed Wilson himself?

Wilson was at the MCZ with Mayr, and would have had access to this copy. In their paper, he and MacArthur thank Mayr (and a few others) for “material aid and advice during the course of the study.” There is thus no question that they consulted Mayr, and used data from a number of Mayr’s papers and books (in Figure 1, as well as Figure 2). If they looked up the data themselves (as opposed to Mayr directly telling them the species numbers), then this particular copy, now in my possession, is a likely copy for them to have used, and Wilson is the likely person to have made the mark highlighting the data that was used. This then, is my speculative (and trivial) suggestion: that Wilson used this copy in the preparation of his and Robert Mac Arthur’s classic and influential paper.

MacArthur and Wilson’s (1963) Figure 2 plots data for 26 islands or island groups. In the cited references, I can find data for only 21 of them; I do not know where they got the data for Tonga, Kei, Tanimbar, Buru, and Ceram. Jurgen Haffer (2007:163), Mayr’s biographer (and himself an accomplished contributor to ornithological science) records the following interesting tidbit:

Mayr (pers. comm.) had copious data on island sizes, distances from mainlands or other islands, number of species, etc. When he tried to determine relations among all these figures he got into mathematical problems and turned this material over to a graduate student with mathematical abilities. However this student got sidetracked into other problems and this material was never exploited.

This account was recalled to Haffer by Mayr decades later, and what data was compiled, when it was compiled, and who the student was, isn’t known.

Haffer (2007: 170) also states that in the 1933 and 1940b papers Mayr “clearly discussed what became later known as the equilibrium theory of insular biogeography”, a claim that has been echoed by other authors. This is not quite correct. Mayr discussed a number of relevant factors contributing to the characteristics of island faunas in 1940b, and the paper is well worth reading and studying today. But he did not formulate in any clear way, even verbally, MacArthur and Wilson’s later theory.  (My very limited German does not allow me to properly assess the 1933 paper, but Mayr’s two 1940 papers in English seem to parallel closely the earlier paper in German.)

For example, Mayr (1940b:215) does note that on a smaller island a species will have a smaller “effective breeding population” and thus be vulnerable to extinction; this is a striking formulation, obviously influenced by theoretical population genetics (likely gotten from Theodosius Dobzhansky). But in discussing extinction on New Caledonia he is clearly discussing evolutionary events stretching over much of the Tertiary (i.e. tens of millions of years), and not the turnover in ecological time of insular populations contemplated by MacArthur and Wilson’s equilibrium theory. The latter has been demonstrated to occur over annual and decadal scales on, for example, smaller islands in the British Isles.

While not excluding the possibility of a MacArthur and Wilson style ecological equilibrium (because he does not consider the situation), Mayr is clearly discussing the origin and persistence (or not) of endemic forms (species, genera, even families) over evolutionary time. Mayr himself (quoted in Haffer, 2007: 163) offers a much more nearly accurate appreciation of his views, stating that his own ideas embraced the “basic thesis” of MacArthur and Wilson; he did not “clearly discuss” the equilibrium theory.

Note on the date of publication. The Sixth Pacific Science Congress was held in 1939, but the Proceedings were published later, from 1940-1943.  Greenway (1958), who knew Mayr well, cited them as 1941, and this may have been the source of my handwritten correction, but I cannot now recall. Haffer’s (2007) definitive list, based on Mayr’s own lists, cites them as 1940.

Greenway, J.C. 1958. Extinct and Vanishing Birds of the World. American Committee for International Wild Life Protection, New York.

Haffer, J. 2007. Ornithology, Evolution, and Philosophy: The Life and Science of Ernst Mayr 1904–2005. Springer, Berlin.

MacArthur, R.H. and E.O. Wilson. 1963. An equilibrium theory of insular zoogeography. Evolution 17:373-387.  pdf

MacArthur, R.H. and E.O. Wilson. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton, N.J.

Mayr, E. 1933. Die Vogelwelt Polynesiens. Mitteilungen aus dem Zoologischen Museum in Berlin 19:306-323.

Mayr, E. 1940a. Borders and subdivisions of the Polynesian Region as based on our knowledge of the distribution of birds. Proceedings of the Sixth Pacific Science Congress 4:191-195.

Mayr, E. 1940b. The origin and history of the bird fauna of Polynesia. Proceedings of the Sixth Pacific Science Congress 4:197-216. (This paper was reprinted in Mayr (1976) but Mayr sometimes updated papers in that collection, and for my purposes I needed to see the original.)

Mayr, E. 1976. Evolution and the Diversity of Life. Harvard University Press, Cambridge, Mass.

Mayr, E. 1943. The zoogeographic position of the Hawaiian Islands. Condor 45:45-48. pdf