One day in the early spring of 1993, Richard Palmer received a paper by a Danish ornithologist, Anders Pape Möller. Palmer, an associate editor at Evolution, was impressed by the paper, but he was troubled by one of Möller's key statistics.
Although he had met Möller only once, Palmer was familiar with his work. Both were fascinated by the promise of fluctuating asymmetry, the subject of the paper in question. "If you measure the right and left sides of the body very precisely, they're never exact mirror images," explains Palmer, an evolutionary biologist at the University of Alberta in Edmonton. "Those differences are random, and what they tell you is the inability of the right side of the body to produce an exact mirror of the left."
Möller's paper claimed that asymmetry in the tail feathers of the barn swallow was passed from fathers to their sons; in other words, it was heritable. But Palmer pointed out in a three-page letter to Möller that the statistical significance of his findings hinged upon a single outlying data point, and therefore "it would be more prudent to present the data, indicate the sensitivity of the statistical result to a single point, and conclude that it is not possible to say much about the heritability of asymmetry with the present data." Instead of addressing Palmer's concerns and those of the two reviewers, however, Palmer says he felt that Möller was just trying to make those concerns go away. Möller ultimately softened the language of the paper and Palmer accepted it for publication, although he says, "I was left with the sense that it was more important for him to get the paper published than to be correct."
Palmer wasn't alone. Evolutionary biologist Bob Montgomerie of Queens College says it's no secret that Möller bickers with editors and referees. As a frequent reviewer of Möller's papers, Montgomerie found himself endlessly pointing out mistakes, but "the stuff was getting published anyway."
Meanwhile, a handful of Möller's colleagues had begun distancing themselves from him. His collegial relationship with evolutionary biologist Andrew Pomiankowski of University College London deteriorated after a dispute over one of their papers. Adrian Thomas, an ornithologist at the University of Oxford, stopped replying to Möller's E-mails regarding a proposed collaboration. Rumors began circulating about the ecologist, including one back-of-the-envelope calculation that retraced his putative bicycle route at his field research site using the sampling methodology described in concurrent studies. The velocities required an athlete of Olympic caliber.
These suspicions would move into the pages of journals, and eventually into a full-fledged investigation that cast serious doubt on one of Möller's papers. In 2005, Möller's bird-banding permit was revoked, effectively ending his 34-year study of barn swallows. "I've slept badly for five years, now," says Möller via the phone from his lab at the Pierre and Marie Curie University in Paris. "I don't think I have done anything wrong." He says his students have been harassed, his collaborators have been discouraged from working with him, and his family has suffered. His friend, Tim Mousseau, a biologist at the University of South Carolina, has seen firsthand how the investigations have affected him. "I think it was very hurtful for somebody who has dedicated their entire life to the pursuit of knowledge," he says, adding: "The only recognition he wants is for his science."
A FARMER'S BOY
Möller was born in the town of N?rresundby on the day after Christmas in 1953. N?rresundby lies in the peatlands of Denmark's sparsely populated Jutland peninsula and is the site of Lindholm H?je, a major Viking burial ground dating back more than a thousand years. While his ancestors took to the sea, Möller took to the land: "I was a farmer's boy."
During his youth, he tended to his father's cows, sheep, and chickens, and, when he had the chance, he watched birds. In the fall of 1969, 15-year-old Möller visited Thorkil Duch, an electrician and an amateur naturalist in the area, who advised him to keep a notebook of his observations. Duch also taught him to capture birds and wrap identifying bands around their legs so that Möller could keep track not just of species but also individuals. Möller returned home and started banding the barn swallow, a slight, nimble bird that would launch his scientific career.
Four years and untold notebooks later, Möller published his first scientific article on barn swallows in a Danish bird journal. He continued to publish throughout high school but was advised not to pursue a career in biology. "I was told there were so few positions that it would never pay off," he says. He went into biology anyway, and was accepted to a doctoral program at the University of Arhus. There, he quickly distinguished himself as a skilled ornithologist and a diligent worker. He wrote modest papers, focusing on mundane but telling details on the lives of common birds: when crows forage, how magpies die, and where blackbirds lay their eggs.
"I was left with the sense that it was more important for him to get the paper published than to be correct."- Richard Palmer
Shortly after receiving his doctorate in 1985, he was publishing 20 to 30 papers a year in international journals: Behavioral Ecology and Sociobiology, Animal Behavior, Evolution, and Oikos, and in 2002 was selected as an ISI Highly Cited researcher in the field of ecology and the environment. He has now published nearly 600 papers. Dolph Schluter, another former editor of Evolution, says, "He's not just prolific. He's good. He's drawn comparisons [and] pointed to relationships that people will be digging through for years."
Part of Möller's success stemmed from his ability to forge productive collaborations. A list of his coauthors is a who's who in the field of behavioral ecology, and he was as likely to collaborate with a top scientist as with a provincial one.
When asked about his tremendous output, Möller laughs nervously and attributes it to his life on the farm: "You had to work hard to earn your dinner."
A FIELD TAKES OFF
Palmer, who published a review of asymmetry in Science in 2004, describes the early pioneers in the field with reverence: Lee Van Valen was "brilliant" and Kenneth Mather wrote "wonderful" papers. Articles on the topic had been trickling in since the 1940s, but the field really took off in the early 1990s thanks to Möller. "Without a doubt," Palmer says, "you can trace the spectacular popularity in this whole subject area to one paper Möller wrote on barn swallows."
Möller had previously shown that longer tails exhibited greater symmetry than shorter tails, a finding which led him to postulate that symmetry could be an indicator of "good genes." Möller's talent, Pomiankowski says, "is taking theoretical ideas and seeing ways they can be tested with data." So Möller promptly modified the length and asymmetry of the birds' tail feathers and found that females preferred the most symmetrical males (see sidebar). A paper, "Female Swallow Preference for Symmetrical Male Sexual Ornaments," was published in Nature in 1992 and was immediately touted by media outlets around the world: symmetry equals attractiveness.
Scientists were skeptical. "The results were too amazing to believe at face value, which was partly what made us look so closely at the paper," says evolutionary biologist Gerald Wilkinson at the University of Maryland, who criticized the study in a published note to Nature. He and ornithologist Gerald Borgia had noticed inconsistencies with error bars on graphs and doubted the paper's conclusions. Möller published a response to their criticisms, but as Wilkinson recalls, "The only way we could reconcile what he said is if his figures had been in error, if they had been crafted improperly."
In 1993, despite the doubts, evolutionary geneticist Therese Markow invited Palmer and Möller to a conference she organized at the Mission Palms Hotel in Tempe, Ariz. During the conference, Möller first suggested that asymmetry was heritable. This idea is a precondition for his "good genes" theory of sexual selection to apply to his barn swallows: If symmetric tails were not heritable, then they could not have evolved under sexual selection. "A rule of thumb is that everything is heritable," says Möller. "Some things have high heritability and some have a low heritability. This is one of the traits that has a low heritability, but it's very interesting."
Möller mentioned several important studies that demonstrated heritability, but the other attendants insisted that there were none. (Palmer agrees that some evidence exists for the heritability of asymmetry, but he says that it is one of the "squishier" connections.) Möller and Randy Thornhill, who was also at the meeting, set out to prove them wrong by performing a meta-analysis of the relationship between asymmetry and heritability. Thornhill, a professor at the University of New Mexico, Albuquerque, and coauthor of the controversial book Natural History of Rape, had been accused of sloppy science in the past. Palmer says he puts the two "in the same basket."
"I've slept badly for five years, now. I don't think I have done anything wrong."- Anders Möller
Palmer rejected the manuscript at Evolution after receiving two "vitriolic" reviews that raised serious questions about its quality. Möller and Thornhill stood by their conclusions, and eventually the paper landed at a less prominent journal, Journal of Evolutionary Biology. The editor there sensed the brewing controversy and, in an unorthodox move, invited seven commentaries to be published alongside the original article in 1997.
The overall tone of these responses ranged from accusations of sloppiness to hyperbole to outright dishonesty. One set of authors suggested that Möller and Thornhill had a hidden agenda in analyzing their data: supporting their "good genes" model of sexual selection. Pomiankowski, who wrote a gentler response to the paper, says, "I was privy to earlier versions of his analysis, and the numbers kept on changing." In their reply, Möller and Thornhill deny a hidden agenda, adding that "there is a real danger to a scientific field when established workers in the field view their colleagues as competitors and use innuendos and direct claims of malpractice to try to get an edge." If Möller and Thornhill really thought they were fooling anyone, they were only fooling themselves.
THE FINAL STRAW
Then, in 1998, Möller published his 33rd paper in the Danish ecological journal Oikos, describing a relationship between asymmetry in oak leaves and damage caused by plant-eating insects. A year later, Oikos editor-in-chief Nils Malmer received an E-mail from Jorgen Rab?l, a former professor in Möller's lab at the University of Copenhagen, who suggested that the data had been fabricated. Möller was shocked. "I had saved all these bloody leaves from these trees," he recalls. "I thought perhaps there was something wrong with these measurements." He went back to his crackling leaf samples and remeasured them. He soon realized that the new data failed to support the conclusions in the Oikos paper. He felt humiliated and did what he and Malmer agreed was the only honorable response: He published a retraction.
That could have been the end of it. But to Möller's dismay, Rab?l brought the case before the Danish Committee on Scientific Dishonesty in 2001. Rab?l presented the committee with files he had obtained from Möller's technician, and the committee then requested Möller's own data files. Möller delayed for months, insisting that the raw data had been stolen along with his laptop in 1996. Instead, he sent the committee a transformed data set that served as the basis for the paper's three tables. The committee noted inconsistencies in even these files and ruled in 2003: "Neither the raw data kept at the University of Copenhagen nor the data forwarded by the defendant could have generated the results that emerged from the article."
Möller insists that the investigation did not prove his guilt but was instead a character assassination. Indeed, Rab?l had been fired after Möller complained of his lack of productivity, and Möller maintains that the accusations were part of Rab?l's revenge. Möller notes that a second investigation, conducted by his home institution, the National Center for Scientific Research (CNRS) in Paris, did not find him guilty of intentionally committing fraud. But even that verdict states that the committee was "lacking the material evidence necessary to establish innocence."
Möller still publishes at a healthy pace, although he says his manuscripts are rejected twice as frequently as before the investigation. "He's under the microscope," says former Evolution editor Schluter at the University of British Columbia. Yet a look at his recent papers shows that while he is keen on citing his own work, he rarely cites opposing views, perhaps hoping, as Palmer remarks, that they'll just "go away."
Perhaps in response, scientists remain critical and even unkind to Möller. In 2000, Palmer published an unusual essay in the newsletter for the International Society of Behavioral Ecology. It was a fable concerning the fictitious Traumweber brothers, Andy and Randy, expert tailors in the "remote kingdom of Gl?cklichtal, nestled high in the European Alps." Palmer wrote that the maestro of Gl?cklichtal's symphony noticed that audiences "seemed pleased with performances conducted in the Traumweber tuxedo, but dissatisfied when he performed in his imported tuxedo." After careful investigation, "Andy Traumweber discovered the imported jacket was less precisely made, most particularly in the tails: one was distinctly longer than the other." The title of the piece, "The Emperor's Codpiece," came from its final coup:
According to a palace informer, the Emperor was particularly anxious about his imperial private parts, which he felt were so asymmetrical that they deviated too far from the norm. Fortunately, the Traumweber brothers were able to allay his fears with a profound revelation: In certain very special cases, increased expression of a predictable asymmetry actually signals increased fitness, and one of those cases is testicles (Möller 1994), at least if men are like birds. That's why they subsequently fashioned the Emperor's codpiece to enhance his already conspicuous asymmetry.
The president of the society, Nick Davies, issued an ambivalent apology in the subsequent newsletter.
"Results were too amazing to believe at face value, which was partly what made us look so closely at the paper."- Gerald Wilkinson
In a devastating book review of Asymmetry, DevelopmentalStability, and Evolution, evolutionary geneticist David Houle at Florida State University, wrote that Möller and his coauthor John Swaddle at the College of William and Mary "repeat the original conclusions of Möller and Thornhill's (1997) meta-analysis of the heritability of asymmetry down to the wildly inflated estimate of average heritability. Although they do address some of the criticisms of others, these are, in effect, dismissed as technical points that do not affect the overall conclusions."
In closing his review, Houle widens his scope to include the gullible souls who jumped aboard the fluctuating asymmetry bandwagon in the 1990s as well as all scientists who succumb too easily to the enthusiasm accompanying new ideas. "We have little choice," writes Houle, "but to seek inspiration from gurus of the newest ideas; sometimes they turn out to be partially right. However, we should never believe them without a struggle. If an idea seems too good to be true, it is probably not true."
These days, Möller's most vocal defender seems to be Mousseau. "I like Rich [Palmer] a lot," Mousseau says, "He's a friend of mine, but he's quite emotional and somewhat irrational in his stance: he just doesn't like Möller." Palmer privately wrote Mousseau and cautioned him not to be so cavalier in defending his colleague. Mousseau, in turn, wrote letters to both Nature and Science with more than 20 coauthors, defended Möller on discussion boards, and started a petition to give Möller back his bird-banding permit.
Pomiankowski says Möller is in the "limbo land" in which many scientists investigated for fraud find themselves. "I find it an unsatisfactory situation to be in, but that's where we are. I would much prefer that he was properly absolved for what happened or found properly guilty." He'd rather know the truth now. "It's very hard to understand what motivates another person," says Pomiankowski. "You can concoct an explanation about why things go wrong, but who knows?"