The “peppered moth paradigm” (Grant 1999 ) was never without problems. As early as 1937, E. B. Ford argued that melanics had higher viability but had been kept in check by visual predators prior to the Industrial Revolution (Ford 1937 ). The absence of negative selection is not the same as the presence of positive selection. In a 1971 Festschrift in honor of Ford's 70th birthday, David Lees cautioned that for both Biston betularia and Phigalia pedaria, “at least in some cases, factors other than air pollution are responsible for their melanic polymorphisms” (p. 210–211). There was a lag time of about a decade before criticisms of Kettlewell's Biston experiments began to appear. When Michael Majerus published Melanism: Evolution in Action in 1998, the storm broke (Majerus 1999 ). Readers of Evolution will know that some reviewers of this book overreacted to its critique of Kettlewell, declaring that the whole Biston story should be thrown out. Bruce Grant then published his closely argued overview and reappraisal in this journal (Grant 1999 ). Predictably, creationists seized on the controversy to yell “Fraud!” and demand the removal of the story from textbooks. For them, industrial melanism was the best—if not the only—case of natural selection documented in the wild, so if Kettlewell was discredited the impact on evolution would be catastrophic. Of Moths and Men raises the stakes yet again.

Judith Hooper is a freelance journalist. Her previous book, The Three-Pound Universe (Hooper and Teresi 1986 ) was a popular exposition of neuropsychology (the “universe” in question is the human brain). It featured interviews and personality portraits of “quirky, brilliant scientists,” as Diane Ackerman put it in the New York Times Book Review. Although it had plenty of pop topics—dream interpretation, near-death experiences, altered states of consciousness—it was in fact fairly serious-minded and a “good read.” Of Moths and Men follows the same formula. I was prepared for a piece of hack sensationalism, especially after reading the publisher's promotional materials (“chronicles one of the most famous experiments—and one of the greatest feuds—in the history of twentieth-century evolutionary biology,” “portrays Kettlewell as a deluded scientist who distorted facts and suppressed evidence he didn't like,” “Tyrannized by his mentor—the powerful E. B. Ford  a Darwinian zealot determined to crush all enemies in his path—Kettlewell ended his life a suicide,” “A story of hubris and heartbreak ”). Amazingly, the book is much better than that. Unlike a potboiler, it's fully footnoted.

Judith Hooper did her homework. The list of people she interviewed or consulted is very impressive, ranging from Michael Majerus, Miriam Rothschild, Laurence Cook, Bryan Clarke, James Cadbury, Kauri Mikkola, Bruce Grant, and other working scientists, to historian of genetics Will Provine, to staff members and laboratory neighbors at Oxford during the peppered-moth years. Of course, merely talking to someone does not guarantee that one asks the right questions or understands or interprets the answers correctly. But, as a sometime journalist myself, I am favorably impressed. She also consulted Kettlewell family members and the appropriate archival resources. Some early reviews of the book have taken her to task for scientific errors. I found remarkably few—11 in 359 pages of text, none of them important. Actually, three of the 11 are errors of personal identification. On p. 286 Paul Kammerer is identified as British (he was Viennese) and on p. 308 I am called a population geneticist, which may surprise my colleagues. The third example merely underrates Vladimir Nabokov as an evolutionist (p. 313; Hooper is unfamiliar with his scientific work, and perhaps has not read Ada). One specific “error” flagged by two reviewers is not an error at all. Hooper calls the North American peppered moth Biston cognataria, a separate species from the Old World B. betularia. That was, in fact, the usage until recently—and, as usual, the decision about conspecificity when wholly allopatric entities are at issue involves a judgment call, though the biological evidence (from Cyril Clarke, David West, and Bruce Grant as well as Kettlewell) certainly argues strongly for conspecificity. Moreover, the matter is only relevant with reference to the interpretation of Kettlewell's (1965) “evolution of dominance” paper, which was never convincing but which Hooper does not criticize in detail anyway. I am told privately that Hooper has gotten a lot of things about specific people and events wrong. Alas, reviewers, like witnesses on the stand, are not free to repeat hearsay. One specific point: Sir Cyril Clarke may have been “butterfingered,” but it was due to a specific medical condition, and he did have technicians besides his wife (p. 273); the reader can decide if this sort of thing is a serious enough slip to impeach Hooper's credibility.

The book may not truly be “a story of hubris and heartbreak,” but it does revolve around the personalities of E. B. Ford and H. B. D. Kettlewell. An odder couple of collaborators is hard to imagine. I seem to fit the general pattern to the extent that I really liked Kettlewell and enjoyed his company, while Ford always gave me the creeps. Hooper retells some of the many apocryphal anecdotes about Ford. (She somehow misses the claim that he once hired a hot-air balloon in order to look for moths spiraling upward toward the full moon. Moth collectors agree broadly that light traps attract very little when the moon is full.) She has fewer Kettlewellian anecdotes (she should have interviewed me!). David Lees (quoted on pp. 210–211) sums up the two men very well! “Ford was an ascetic, an aesthete and a misogynist, but a real intellectual. Kettlewell was a larger-than-life bon vivant with a liking for female company, shooting and fishing, but no intellectual.” All of this was evident to me during my brief stint at Parks Road. I have often wondered what Bernard and Hazel talked about when he came home from having to deal with “Henry” (as Ford was known).

I have also wondered why Kettlewell's early work had so many problems with experimental design. Surely, I assumed, his protocols had been vetted by Ford—if not designed by him!—and Ford would not have been so careless. Besides, R. A. Fisher was still alive then. Why were the experiments so naïve? Hooper would have us believe Kettlewell was largely on his own—that Ford wanted results but was not very engaged with how he got them: a rather surprising notion.

This leads directly to the imputation of fraud. Kettlewell's initial mark-release-recapture experiment was not going well at the end of June 1953, when he received a message from Ford encouraging him to do better. The recapture rates improved immediately and never relapsed. To her credit, Hooper researched the weather records to see if anything changed that might have altered the moths' behavior; she found nothing. That's it: that's the “smoking gun.” Hooper thinks Kettlewell was under tremendous pressure to please Ford, and obliged. No unbiased jury would be convinced. (Ford, if he were still with us, would surely ask her about the phase of the moon.)

Kettlewell's most persistent critic, the American Ted Sargent, advances a complex model of industrial melanism whereby environmental factors modulate gene expression. It is plausible but highly unparsimonious, and therefore the burden of persuasion is on Sargent. He shows little interest in testing it empirically himself. Hooper tries half-heartedly to make Sargent the “hero” of the book—but he comes across as a classic square peg in a round hole, a man with as many personal “issues” as the Oxonians, and no more sympathetic. The matter is not helped by a sappy story involving Sargent and a supposed “mockingbird death song” (p. 257) which is intended to show us what a keen observer he is.

Grant has pointed out that even if we throw out Kettlewell's work—just for the sake of argument—there is abundant evidence from independent sources to support not only the reality of industrial melanism but also the sufficiency of natural selection to explain it. This is so even if the actual agent(s) of selection remain totally unknown—a great irony, since the goal of the Oxford school was to make explicit the ecological processes responsible for differential survival and reproduction.

There is a striking parallel here with the case of Cyril Burt, the controversial, pioneering hereditarian of IQ. His estimate of +0.77 as the IQ correlation of identical twins reared apart was discredited amid accusations of fraud (Gillie 1976 ; Gould 1981 ), only to be obtained again by independent and untainted investigators (+0.78 in a Minnesota study; Bouchard et al. 1990 ). No matter how hard the creationists may wish to sweep the industrial melanism story away, it's here to stay; it transcends the merely personal. (Melanism itself is apparently not here to stay; it is in free fall on both sides of the Atlantic.)

If Kettlewell had little or no input from Ford and Fisher, his mistakes are that much more understandable. But no one else was doing this kind of thing, and was there anyone around in 1953 who might have done it better? As an experiment, read a few issues of Evolution or Ecology or American Naturalist from around 1950. A surprising amount of what you read will be downright embarrassing by today's standards. But it was “cutting edge” then. To damn Kettlewell 50 years after the fact is of a piece with using contemporary notions of “political correctness” to damn Shakespeare as an anti-Semite or banish Mark Twain from the curriculum because of the n-word.

Bernard Kettlewell did die a suicide, but because of chronic pain, not any guilt or angst over his supposed scientific sins. Here's the Bernard I remember: I had been to his home at Steeple Barton. He took me to the station to catch the train back to Oxford. It was a dank, dreary, drizzly day. I was suitably dressed, but Bernard, in his trademark Bermuda shorts, was clearly freezing. I told him to go home; I was perfectly capable of taking a train unaided. “My good man!” he blustered, “What kind of host would I be if I abandoned my guest at the station!” The train by now was quite late. A self-propelled maintenance car hove into view on the tracks, carrying a couple of laborers with buckets and shovels. Bernard leaned over the tracks and yelled “Yoo-hoo! British Rail! Are you the 5:43?”

Bouchard, T. J., D. T. Lykken, M. McGue, N. L. Segal, and A. Tellegen. 1990. Sources of human psychological differences: the Minnesota study of twins reared apart. Science. 250:223–228. [PubMed Citation]

Ford, E. B. 1937. Problems of heredity in the Lepidoptera. Biol. Rev. 12:461–503.

Gillie, O. 1976. Crucial data faked by eminent psychologist. Sunday Times (London), Oct. 24, pp. 1–2.

Gould, S. J. 1981. The mismeasure of man. W. W. Norton, New York.

Grant, B. 1999. Fine-tuning the peppered moth paradigm. Evolution. 53:980–984.

Hooper, J., and D. Teresi. 1986. The three-pound universe. Macmillan, New York.

Kettlewell, H. B. D. 1965. Insect survival and selection for pattern. Science. 148:1290–1295.

Lees, D. 1971. The distribution of melanism in the Pale Brindled Beauty moth, Phigalia pedaria, in Great Britain. Pp. 152–174 in R. Creed, ed. Ecological genetics and evolution. Essays in honour of E. B. Ford. Blackwell, Oxford, U.K.

Majerus, M. 1998. Melanism: evolution in action. Oxford Univ. Press, Oxford, U.K.

¹Of Moths and Men: An Evolutionary Tale. The Untold Story of Science and the Peppered Moth. Judith Hooper. 2002. W. W. Norton, New York and London. xx + 377 pp. HB $26.95, ISBN 0-393-05121-8.

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