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    A Fluctuat

    Although he had met Moller only once,Palmer was amiwith his work. Both were ascinated y the promiseof fluctuaaslrnmetry the subjectof the paper n question. If you meathe right and left sidesofthe body very precisely,hey're neexactmirror images," xplainsPalmer,an evolutionarybioloat the Universityof Alberta in Edmonton. "Thosedifferencerandom,and what they tell you s the inability of the right sidthe body o producean exactmirror of the left."Moller'spaperclaimed hat as).rnmetryn the tail feathethe barn swallow waspassedrom fathers o their sons; n otwords, t was heritable. But Palmerpointed out in a three-pletter to Msller that the statistical significanceof his findinhingedupon a singleoutlyingdatapoint, and herefore it wobe more prudent to present he data, ndicate the sensitivitthe statistical result to a singlepoint, and conclude hat it ispossibleo saymuch about he heritability of asyrnmetrywithpresentdata." nsteadof addressingPalmer's oncernsand thof the two reviewers,however,Palmersayshe felt that Moller wjust trying to make those concernsgo away.Msller ultimatsoftened he languageofthe paper and Palmer accepted t

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    A FLUCTUATINGEALITY

    publication, although he says, I was left with the sense hat itwas more important for him to get the paper published han tobe correct."Palmerwasn't alone. Evolutionarybiologist Bob Montgom-erie of QueensCollegesays t's no secret hat Msller bickers witheditors and referees.As a frequent reviewer of Moller's papers,Montgomerie ound himself endlessly ointing out mistakes, ut

    "the stuffwas getting published anyway."Meanwhile, a handful of Msller's colleagueshad begun dis-tancing themselves rom him. His collegial relationship with evo-lutionary biologist Andrew Pomiankowski of University CollegeLondon deteriorated after a dispute over one oftheir papers.Adrian Thomas, an ornithologist at the University of Oford,stopped replying to Moller's E-mails regarding a proposedcollab-oration. Rumors began circulating aboutthe ecologist, ncludingone back-of-the-envelope calculation that retraced his putativebicycle route at his field researchsite using the sampling meth-odolory described n concurrent studies. The velocities requiredan athlete of Olympic caliber.These suspicions would move into the pagesof journals,and eventually into a full-fledged investigation that cast seriousdoubt on one of Msller's papers. n2OO5, Msller's bird-bandingpermit was revoked,effectively ending his 3,tr-yearstudy of barnswallows."I've slept badly for five years,now," saysMsller viathe phone ftom his lab at the Pierre and Marie Curie Universityin Paris. I dont think I havedone anythingwrong."He sayshisstudentshavebeen harassed,his collaborators havebeen dis-couraged rom working with him, and his family has suffered.His friend, Tim Mousseau,a biologist at the University of SouthCarolina,hasseen irsthandhow the investigationshaveaffectedhim. "I think it was very hurbful for somebodywho has dedicatedtheir entire ife to thepursuit ofknowledgei he says, dding: "Theonly recognitionhe wants s for his science.'A FARMER'SOYMoller was born in the town of Norresundby on the day afterChristmas in 1953. Norresundby lies in the peatlandsof Den-mark's sparselypopulatedJutland peninsula and is the site ofLindholm Haje, a majorViking burial ground dating back morethan a housandyears.While his ancestorsook to the sea,Mollertook to the land: "I was a farmer'sboy."TH E SCIENTIST anuarv00 7

    , , IWASLEFT ITHTHESENSEHATWASMOREMPORTANTOR IMTGET HEPAPERUBLISHEDHAN BECORRE-RICHARD ALM

    During his youth, he tended to his father'scows,sheep,chickens,and,when he had the chance,he watchedbirds. Infall of 1969, 5-year-old Moller visited Thorkil Duch, an elecian and an amateurnaturalist in the area,who advisedhimkeep a notebook of his observations.Duch alsotaught himcapture birds andwrap identiffing bandsaround their legsso Msller could keep rack not just of species ut also ndividuMsller returned home and started banding the barn swalloslight, nimble bird that would launch his scientific career.

    Four years and untold notebooks later, Moller publishedfirst scientific article on barn swallows in a Danish bird journHe continued to publish throughout high school but was advnot to pursue a caxeer n biolory. "I was told there were so positions that it would never pay off" he says. He went ibiology anyway, and was accepted o a doctoral program at University of Arhus. There, he quickly distinguished himsela skilled ornithologist and a diligent worker. He wrote modpapers, focusing on mundane but telling details on the livecommon birds: when crows forage, how magpiesdie, and whblackbirds lay their eggs.

    Shortly a"fter eceiving his doctorate in 1985, he waspubliing 20 to 30 papersayear in internationaljournals: BehaoiEcolog and Sociobiology,Anirnal Behaoior, Eaolution, Oikos,and n 2oo2 wasselected san ISI Highly Cited researin the field of ecologyand the environment. He has now plished nearly 600 papers.Dolph Schluter,another ormer edof Enolution, says, He'snot just prolific. He'sgood.He's dracomparisons and] pointed to relationships hat peoplewilldigging through for years;"

    Part of Msller's success temmed rom his ability to forgepductivecollaborations.A list of his coauthors s a who's whothe field of behavioralecolog'y,nd he wasas ikely to collabowith a top scientist aswith a provincialone.

    When askedabouthis tremendousoutput, Moller laughsnvously and attributes it to his life on the farm: 'You had to whard to earnyour dinner."A FIELD AKES FFPalmer,who publisheda reviewof as5,'rnmelryn Sciencet 2odescribeshe earlypioneers n the field with reverence:Lee VValen was 'brilliant" and Kenneth Mather wrote "wonder

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    papers.Articles on the topichadbeen rickling in since he 94,Os,but the field really took offin the early 199os hanks to Moller.'Without a doubt," Palmersays, you can trace the spectacularpopularity in this whole subject axea o one paper Moller wroteon barn swallows."Msller had previously shown that longer tails exhibitedgreater symmetry than shorter tails, a finding which led him topostulate hat symmetry could be an indicator of "goodgenes."Moller's talent, Pomiankowski says, is taking theoretical deasand seeingways hey can be tested with datal So Moller promptlymodified he engthand asymmetryof the birds'tail feathersandfound that femalespreferred the most s)rynmetricalmales(seesidebar).A paper, "FemaleSwallow Preference or SymmetricalMale SexualOrnaments,"was publishedinNature in 1992 andwas mmediately outedby media outlets around he world: sym-metry equals attractiveness.

    Scientistswere skeptical. The results were too amazing obelieveat facevalue,which was partly what made us look socloselyat the paper,"saysevolutionarybiologist GeraldWilkinsonat the University of Maryland, who criticized the study in a pub-lished note to Nature. He and ornithologist Gerald Borgia hadnoticed inconsistencies with error bars on graphs a.nddoubtedthepaper's onclusions.Moller publisheda responseo their criti-cisms,but asWilkinson recalls, The onlywaywe could reconcilewhat he said is if his figures had been in error, if they had beencrafted improperly."

    HERITABTESYMMETRY?In 1993, despite the doubts, evolutionary geneticist ThereseMarkow invited Palmer and Moller to a conferenceshe organizedat the Mission Palms Hotel in Tempe,Ariz. During the conference,Msller first suggestedhat as1'rnmetry asheritable.This idea sa preconditionfor his "goodgenes" heory ofsexual selection oapply to his barn swallows: If symmetric tails were not heritable,

    Thornhill, aprofessorat the University ofNew Mexico, Albuquque,and coauthor ofthe controversialbook Notural HistorRape,hadbeenaccused f sloppysciencen the past.Palmerhe puts the two "in the same basket."Palmer rejected the manuscript at Eoolution aft,er ecing two "vitriolic" reviews that raised seriousquestionsabits qualrty.Moller and Thornhill stood by their conclusions,eventually he paper anded at a essprominentjotrnal, JouofEoolutionary Biologg. The editor there sensed he brewcontroversyand, in an unorthodox move, nvited sevencommtaries o be publishedalongsidehe original article n 1997.The overall tone of these responses anged rom acctions ofsloppiness o hyperbole o outright dishonesty.Oneof authors suggested hat Msller and Thornhill had a hidagenda n analyzing heir data: supporting their "good gemodel of sexualselection.Pomiankowski,who wrote a genresponse o the paper,says, I was privy to earlier versionhis analysis,and the numbers kept on changing." n their reMsller and Thornhill deny a hidden agenda,adding that "this a real danger to a scientific field when establishedworkin the field view their colleaguesascompetitors and use nendosand direct claims of malpractice o try to get an edgeMsller and Thornhill really thought they were fooling anythey were only fooling themselves.THEFINALSTRAWThen, in 1998,Moller publishedhis 33rd paper in the Daecologicalourna) Oikos,describing a relationship between a.metry in oak eavesand damagecaused y plant-eating nseA year late4 Oikos editor-in-chief Nils Malmer receiveE-mail from JorgenRabol, a former professor n Moller's athe University of Copenhagen, ho suggestedhat the databeen abricated.Msller was shocked. I had savedall thesebloleaves rom these rees," e recalls. I thought perhaps here

    , , I 'VESLEPTADLYOR IVE EARS,OW. DON'TTHINKHAVE ONEANYTHING RONG."then they could not have evolvedunder sexualselection. A. ule ofthumb is that everything s heritable,"saysMoller. "Some hingshavehigh heritability and somehavea ow heritability. This isoneof the traits that has a low heritability, but it's very interesting."

    Msller mentioned several mportant studies that demon-strated heritability, but the other attendants insisted that therewere none. Palmeragreeshat someevidence xists or the heri-tability of asymmetry but he says hat it is one of the "squishier"connections.)Msller and Randy Thornhill, who was also at themeeting, set out to prove them wrong by performing a meta-analysisof the relationshipbetweenasyrnmetryand heritability.

    -ANDFRSMALLsomething wrong with these measurements."He went backtocrackling eaf samplesand remeasured hem. He soonrealthat the new data failed to support the conclusions inthe Opaper. He felt humiliated and did what he and Malmer agwas he only honorable esponse:He publisheda retraction

    ,, Vi*it tfur *rvw"th*-sqiqrttisi,**r * ,I:lr n irt*er* ctivc.::'t' . l!ru*ii*e *l &itsller'l v*rk and re*r.lace*l*ti*cs rry, i:r igin;r lr j*curne*tat i**andc*senrentesy.

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    That could havebeen the end of it. But to Moller's dismay,Rabolbrought he case efore he DanishCommitteeon ScientificDishonestyn 2oo1. Rabolpresentedhe committeewith files hehad obtained rom Msller's technician,and the committee henrequested Moller's own data files. Msller delayed or months,insisting that the raw data had been stolen alongwith his laptopin f996. Instead,he sent the committeea transformed datasetthat servedas he basis or the paper's hree ables.The commit-tee noted inconsistenciesn even hese iles and ruled in 2OO3:"Neither the raw data kept at the University of Copenhagen orthe data forwarded by the defendantcould havegenerated heresults that emerged rom the article."Msller insists that the investigationdid not prove his guiltbut was instead a character assassination. ndeed. Rabol hadbeen fired after Moller complained of his lack of productivity,and Msller maintains that the accusationswerepart of Rabol'srevenge.Msller notes hat a second nvestigation,conductedbyhis home nstitution, the National Center or ScientificResearch(CNRS) n Paris,did not find him guilty of intentionallyeommit-ting fraud. But even that verdict statesthat the committee was"lacking the material evidencenecessaryo establish innocence."

    THEAFTERMATHMoller still publishes at a healthy pace, although he says hismanuscriptsare rejected wice as requentlyas before he inves-tigation. "Het under the microscope,"says ormer Eztolutioneditor Schluter at the University of British Columbia. Yet a lookat his recent papersshows hat while he is keen on citing his ownwork, he rarely cites opposingviews,perhapshoping, as Palmerremarks, hat they'll ust "go away."Perhapsn response,cientists emaincriticalandevenunkindto Moller. In 2OOO,Palmerpublished an unusual essayn thenewsletter or the nternational Societgof Behazsioral cology.IIwasa fable concerning the fictitious Traumweber brothers, Andyand Randy, expert tailors in the'remote kingdom of GliicHichtal,nestledhigh in the EuropeanAlps."Palmer wrote that the maestroof Gliicklichtal's symphony noticed that audiences"seemedpleasedwith performances onductedn the taumweber tuxedo,but dissatisfiedwhen heperformed n his mported tuxedo."Aftercareful nvestigation, "AndyTraumweberdiscovered he importedjacketwas esspreciselymade,mostparticularly in the tails: onewas distinctly longer than the other."The title of the piece,"TheEmperor'sCodpiece,"ame rom its final coup:According to a palace nformer, the Emperor was particu-

    larly anxiousabout his imperial private parts, which he feltwere so asymmetrical that they deviated too far from thenorm. Fortunately, the Traumweberbrothers were able toallay his fears with a profound revelation: In certain veryspecialcases,ncreasedexpression fa predictableasym-metry actually signals increased itness, and one of thosecases s testicles Moller 1994,), t least f men are ike birds.That'swhythey subsequentlyfashionedhe Emperor's od-piece o enhancehis alreadyconspicuousas]rynmetry.

    "THERESULTS ERETOBELIEVE FACESOCLOSELYTHE

    TOOAMAZINGMADEUSLOOKPAPER.''-GERALD 4//LK/NS

    The presidentof the society,Nick Davies, ssuedan amblent apology n the subsequent ewsletter.In a devastating book review of Asgmmetry, DerselopmeStability, and,Eoolution, evolutionary geneticist David HouleFlorida StateUniversity, wrote that Moller andhis coauthor JoSwaddleat the Collegeof William and Mary "repeat he origconclusions f Msller and Thornhill's (rgg7)meta-analysis fheritability of asymmetrydown to the wildly inflated estimataverage eritability. Although they do addresssomeof the ccismsof others, heseare, n efiect,dismissedas echnicalpothat do not affect the overall conclusions."

    In closing his review, Houle widenshis scope o includegullible souls who jumped aboard the fluctuating asymmbandwagon n the 1990s as well as all scientistswho succutoo easily o the enthusiasmaccompanying ew ideas. Wehlittle choicei writes Houle, "but to seek nspiration from guruthe newest deas;sometimes hey turn out to be partially rigHowever,we should never believe them without a struggle. Ifidea seemsoo good o be true, it is probablynot true."Thesedays,Moller's most vocal defenderseems o be Moseau. I like Rich [Palmer] a loti Mousseausays, He'sa frieof mine, but he'squite emotional and somewhat rrationahis stance: he just doesn't ike Moller." Palmerprivately wrMousseauand cautionedhim not to be so cavalier n defendhis colleague.Mousseau, n turn, wrote letters to both Natand,Sciencewith more than 20 coauthors, defended Msllerdiscussionboards,and started a petition to giveMoller backbird-bandingpermit.

    Pomiankowski saysMoller is in the "limbo land" in whmany scientists nvestigated or fraud find themselves.I finan unsatisfactory situation to be in, but that's where we arwould much prefer that he was properlyabsolvedor what hpenedor found properly guilty." He d rather know the truth n"Itt very hard to understand what motivatesanotherperson,"sPomiankowski.'You an concoctanexplanationaboutwhythingowrong, but who knows?" sHavea comment?Emailus at mai l@the-sci entistom

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    ;: l he nAndersPapeMollerpublished is nowinfamous 99 2Nature aperon barnswallit wasa leapof faith o testwhether emalmightchoosemalesbased n asymmetrybetweenhe eftan d rightsides f the bod

    controversyof

    Such symmetry ad beenknown o correlate ith stress nd othenvironmentalactors, ut not o inateselect iori.

    Moller s a strongproponent f the "goodgenes"modelofsxualSelect ion,2.3hichholds hat cert iain haracterist ics,orexample,he tail eathers f a peacock, ignal he presence fgenes hat wil l be beneficialo offspring, nd hereforemakeanindiv idualmoreat t ract iveo potent ia lmates.Mol lerbegan o thihat peihapsmoresubt le ignals f a mate's ual i ty , uchas aisymmetry, lsomattered.a e used o measure nly he engthoone sideof the ail,bu t he says, Therewas oneyear ha t the swlowscameback romAfr icaand herewere ots of indiv idualshwereasymmetric." o esi his heory,he capturedwild malesanvaried he asymmetry etween he left and rightoutermost ai lfeathers y 20 mm. Sureenough,maleswith moresymmetric afeathers eceivedmorevisits rom females,matedearlier, nd hgreater eproductive uccess.l

    As AndrewBalmford ndAdr ianThomas ndicatedn heirreply o Mollei 'spape4snvoking exual elect ionwas not necesary o explainwh y birdshave ymmetr icalai ls :Lopsided i rdswouldhave rouble ly ing.GeraldBorgia ndGeraldWilk insonfound ha i Msller 'sstandard rrorswere es s han one-tenth fthoseMollerhad reported reviously. oreoveq he differenceamong reatmentswereso great,Borgia nd Wilkinson uspectthat maleswereassigned onrandomlyo treatments-6 ichardPalmer aspqinted ut that Mol ler 'smanipulat ionsf asymmetry were en imesas greatas he natural ar iat ion f 2-3 mm.However,Msller 'sbarnswallow tudieshaveneverbeen nvestgated or scientif ic ishonestyR rerencrs1. A.P Msller, "Female swallow preference for slmnetrical male sexual

    omainents;' Natzre, 357 1238 4O, 7992.2. A.P. Moller, "Female choice for male sexual omaments in the monosamous

    swallowl' Nature, 332i64O-42, -t988.

    MALESWITH MCRESYMMETRICAILFEATHERSECEIVMORE ISITSROM EMALIS,ATED ARLIER,NDHA: GREA;IEREPRODUCTIVEUCCE3. A.P. Moilea 'r9iability of costs of male tail ornaments in a swallow,"Mafzre,339:Ig2-5,1989.a. A.P. Moller, "Fiuctuating asFnmetry in male sexualornaments may reliablyrevealmalequalityl'Animal Behauiour,4,Otitgi-7, LggO.5. A. Balmford,A Thomm, "Swallowingornamental asymmetwi'Nature,359:487, 992.6. G Borgia,G Wilkinsirn,"swillowing omamental as).rnmetryiNdture,

    359t4a7-8,1992.