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AN ORNITHOLOGIST FROM
GOES
TO PARIAH
SUPERSTAR
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F
 A Fluctuating

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l\tt-,.iicri ,:f 1i;it.;.j4;t;jilt:, l;:ji.'r,: lr;,, r-i,:';itllij
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By BrendanBorrell

Although he had met Moller only once,Palmer was familiar

with his work. Both werefascinatedby the promiseof fluctuating
aslrnmetry the subjectof the paper in question."If you measure
the right and left sidesofthe body very precisely,they're never
exactmirror images,"explainsPalmer,an evolutionarybiologist
at the University of Alberta in Edmonton. "Thosedifferencesare
random, and what they tell you is the inability of the right side of
the body to produce an exactmirror of the left."
Moller's paper claimed that as).rnmetryin the tail feathersof
the barn swallow was passedfrom fathers to their sons;in other
words, it was heritable. But Palmer pointed out in a three-page
letter to Msller that the statistical significanceof his findings
hinged upon a singleoutlying data point, and therefore"it would
be more prudent to present the data, indicate the sensitivity of
the statistical result to a singlepoint, and concludethat it is not
possibleto saymuch aboutthe heritability of asyrnmetrywith the
presentdata."Insteadof addressingPalmer'sconcernsand those
of the two reviewers,however,Palmer sayshe felt that Moller was
just trying to make those concernsgo away.Msller ultimately
softened the languageofthe paper and Palmer acceptedit for

26 T H E S C IE NT IS T Ja n u a rv2QOT
A FLUCTUATING
REALITY

,,IWASLEFT
WITHTHESENSETHATIT
WASMOREIMPORTANT FORHIMTO
GETTHEPAPERPUBLISHED
THANTO
BECORRECT.''
-RICHARDPALMER

publication, although he says,"I was left with the sensethat it During his youth, he tended to his father's cows,sheep,and
was more important for him to get the paper published than to chickens,and, when he had the chance,he watchedbirds. In the
be correct." fall of 1969, t5-year-old Moller visited Thorkil Duch, an electri-
Palmer wasn't alone. Evolutionary biologist Bob Montgom- cian and an amateur naturalist in the area,who advisedhim to
erie of QueensCollegesaysit's no secret that Msller bickers with keep a notebook of his observations.Duch also taught him to
editors and referees.As a frequent reviewer of Moller's papers, capture birds and wrap identiffing bands around their legs sothat
Montgomerie found himself endlesslypointing out mistakes,but Msller could keep track not just of speciesbut also individuals.
"the stuffwas getting published anyway." Msller returned home and started banding the barn swallow, a
Meanwhile, a handful of Msller's colleagueshad begun dis- slight, nimble bird that would launch his scientific career.
tancing themselvesfrom him. His collegial relationship with evo- Four years and untold notebooks later, Moller published his
lutionary biologist Andrew Pomiankowski of University College first scientific article on barn swallows in a Danish bird journal.
London deteriorated after a dispute over one oftheir papers. He continued to publish throughout high school but was advised
Adrian Thomas, an ornithologist at the University of Oford, not to pursue a caxeerin biolory. "I was told there were so few
stopped replying to Moller's E-mails regarding a proposed collab- positions that it would never pay off" he says. He went into
oration. Rumors began circulating aboutthe ecologist, including biology anyway, and was accepted to a doctoral program at the
one back-of-the-envelope calculation that retraced his putative University of Arhus. There, he quickly distinguished himself as
bicycle route at his field research site using the sampling meth- a skilled ornithologist and a diligent worker. He wrote modest
odolory described in concurrent studies. The velocities required papers, focusing on mundane but telling details on the lives of
an athlete of Olympic caliber. common birds: when crows forage, how magpies die, and where
These suspicions would move into the pages of journals, blackbirds lay their eggs.
and eventually into a full-fledged investigation that cast serious Shortly a"fterreceiving his doctorate in 1985,he was publish-
doubt on one of Msller's papers.In2OO5, Msller's bird-banding ing 20 to 30 papersayear in internationaljournals: Behaoioral
permit was revoked, effectively ending his 3,tr-yearstudy of barn Ecolog and Sociobiology,Anirnal Behaoior, Eaolution, and
swallows. "I've slept badly for five years, now," saysMsller via Oikos,and in 2oo2 was selectedasan ISI Highly Cited researcher
the phone ftom his lab at the Pierre and Marie Curie University in the field of ecologyand the environment. He has now pub-
in Paris."I dont think I have done anything wrong." He sayshis lished nearly 600 papers.Dolph Schluter,another former editor
students have been harassed,his collaborators have been dis- of Enolution, says,"He's not just prolific. He's good. He's drawn
couragedfrom working with him, and his family has suffered. comparisons[and] pointed to relationshipsthat people will be
His friend, Tim Mousseau,a biologist at the University of South digging through for years;"
Carolina,has seenfirsthand how the investigationshaveaffected Part of Msller's successstemmedfrom his ability to forge pro-
him. "I think it was very hurbful for somebodywho has dedicated ductive collaborations.A list of his coauthors is a who's who in
their entire life to the pursuit ofknowledgei he says,adding: "The the field of behavioralecolog'y,and he was aslikely to collaborate
only recognition he wants is for his science.' with a top scientist aswith a provincial one.
When askedabouthis tremendousoutput, Moller laughsner-
vously and attributes it to his life on the farm: 'You had to work
A FARMER'S BOY
hard to earn your dinner."
Moller was born in the town of Norresundby on the day after
Christmas in 1953. Norresundby lies in the peatlands of Den-
A FIELD TAKESOFF
mark's sparselypopulated Jutland peninsula and is the site of
Lindholm Haje, a major Viking burial ground dating back more Palmer,who publisheda reviewof as5,'rnmelry in Scienceit 2oo4,
than a thousandyears.While his ancestorstook to the sea,Moller describesthe early pioneersin the field with reverence:Lee Van
took to the land: "I was a farmer'sboy." Valen was 'brilliant" and Kenneth Mather wrote "wonderful"

TH E S C I E N T I S TJ a n u a r 2v 0 0 7
papers.Articles on the topic had beentrickling in sincethe l94,Os,
but the field really took offin the early 199osthanks to Moller.
'Without a doubt," Palmer says,"you can trace the spectacular
popularity in this whole subject axeato one paper Moller wrote
on barn swallows."
Msller 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."
Moller's talent, Pomiankowski says,"is taking theoretical ideas
and seeingwaysthey can be testedwith datal So Moller promptly
modified the length and asymmetryof the birds'tail feathersand
found that femalespreferred the most s)rynmetricalmales (see
sidebar). A paper, "Female Swallow Preferencefor Symmetrical
Male SexualOrnaments,"was publishedinNature in 1992 and
was immediatelytouted by media outlets around the world: sym-
metry equals attractiveness.
Scientistswere skeptical."The results were too amazing to
believe at face value, which was partly what made us look so
closelyat the paper,"saysevolutionarybiologist GeraldWilkinson
at the University of Maryland, who criticized the study in a pub-
lished note to Nature. He and ornithologist Gerald Borgia had
noticed inconsistencies with error bars on graphs a.nddoubted
the paper'sconclusions.Moller publisheda responseto their criti-
cisms,but asWilkinson recalls,"The onlywaywe could reconcile
what he said is if his figures had been in error, if they had been
crafted improperly."
ASYMMETRY?
HERITABTE
In 1993, despite the doubts, evolutionary geneticist Therese
Markow invited Palmer and Moller to a conferenceshe organized
at the Mission PalmsHotel in Tempe,Ariz. During the conference,
Msller first suggestedthat as1'rnmetrywas heritable.This idea is
a precondition for his "good genes"theory ofsexual selectionto
apply to his barn swallows: If symmetric tails were not heritable,
,,I'VE BADLY
SLEPT FORFIVEYEARS,
NOW.I DON'TTHINK
I HAVEDONE
ANYTHINWRONG."
G
then they could not haveevolvedunder sexualselection.iA.rule of
thumb is that everythingis heritable,"saysMoller. "Somethings
havehigh heritability and somehave a low heritability. This is one
of the traits that has a low heritability, but it's very interesting."
Msller mentioned several important studies that demon-
strated heritability, but the other attendants insisted that there
were none. (Palmeragreesthat someevidenceexistsfor the heri-
tability of asymmetry but he saysthat it is one of the "squishier"
connections.)Msller and Randy Thornhill, who was also at the
meeting, set out to prove them wrong by performing a meta-
analysisof the relationship betweenasyrnmetryand heritability.
Thornhill, a professorat the University ofNew Mexico, Albuquer-
que, and coauthor ofthe controversialbook Notural History of
Rape,hadbeenaccusedof sloppysciencein the past.Palmersays
he puts the two "in the samebasket."
Palmer rejected the manuscript at Eoolution aft,erreceiv-
ing two "vitriolic" reviews that raised serious questions about
its qualrty. Moller and Thornhill stood by their conclusions, and
eventuallythe paperlanded at a lessprominentjotrnal, Journal
ofEoolutionary Biologg. The editor there sensedthe brewing
controversy and, in an unorthodox move, invited sevencommen-
taries to be published alongsidethe original article in 1997.
The overall tone of these responsesranged from accusa-
tions ofsloppiness to hyperbole to outright dishonesty.One set
of authors suggestedthat Msller and Thornhill had a hidden
agendain analyzing their data: supporting their "good genes"
model of sexual selection.Pomiankowski,who wrote a gentler
responseto the paper, says,"I was privy to earlier versions of
his analysis,and the numbers kept on changing."In their reply,
Msller and Thornhill deny a hidden agenda,adding that "there
is a real danger to a scientific field when establishedworkers
in the field view their colleaguesas competitors and use innu-
endos and direct claims of malpractice to try to get an edge."If
Msller and Thornhill really thought they were fooling anyone,
they were only fooling themselves.
THEFINALSTRAW
Then, in 1998, Moller published his 33rd paper in the Danish
ecologicaljourna) Oikos, describing a relationship between a.sym-
metry in oak leavesand damagecausedby plant-eating insects.
A year late4 Oikos editor-in-chief Nils Malmer received an
E-mail from Jorgen Rabol, a former professorin Moller's lab at
the University of Copenhagen,who suggestedthat the data had
been fabricated. Msller was shocked."I had savedall thesebloody
leavesfrom thesetrees,"he recalls."I thought perhapsthere was
-ANDFRSMALLER
something wrong with these measurements."He went back to his
crackling leaf samplesand remeasuredthem. He soon realized
that the new data failed to support the conclusions inthe Oikos
paper. He felt humiliated and did what he and Malmer agreed
was the only honorableresponse:He published a retraction.
,, I Vi*it tfur *rvw"th*-sqiqrttisi,**r* f,I:lr*n irt*er*ctivc
.::'t' . l!ru*ii*e *l &itsller'l :v*rk and ire*r.lace*l*ti*cs rryitll
, i:rigin;rlrj*curne*tati**andc*senrentesy.
lanuary2OOTTHE SCIENTIST 29
 That could have been the end of it. But to Moller's dismay,
Rabolbrought the casebeforethe Danish Committeeon Scientific
Dishonestyin 2oo1. Rabol presentedthe committeewith files he
had obtained from Msller's technician, and the committee then
requested Moller's own data files. Msller delayed for months,
insisting that the raw data had been stolen along with his laptop
in f996. Instead, he sent the committee a transformed data set "THERESULTS
WERETOOAMAZING
that servedasthe basisfor the paper'sthree tables.The commit-
tee noted inconsistenciesin even these files and ruled in 2OO3: TOBELIEVE FACE
"Neither the raw data kept at the University of Copenhagennor MADEUSLOOK
the data forwarded by the defendant could have generatedthe SOCLOSELYTHEPAPER.''
results that emerged from the article." -GERALD\4//LK/NSO/V
Msller insists that the investigation did not prove his guilt
but was instead a character assassination.Indeed. Rabol had
been fired after Moller complained of his lack of productivity,
and Msller maintains that the accusationswere part of Rabol's
revenge.Msller notesthat a secondinvestigation,conductedby
his home institution, the National Centerfor ScientificResearch The president of the society,Nick Davies,issuedan ambiva-
(CNRS)in Paris,did not find him guilty of intentionally eommit- lent apologyin the subsequentnewsletter.
ting fraud. But even that verdict states that the committee was In a devastating book review of Asgmmetry, Derselopmental
"lacking the material evidencenecessaryto establish innocence." Stability, and,Eoolution, evolutionary geneticist David Houle at
Florida State University, wrote that Moller and his coauthor John
Swaddleat the Collegeof William and Mary "repeatthe original
THEAFTERMATH
conclusionsof Msller and Thornhill's (rgg7) meta-analysisofthe
Moller still publishes at a healthy pace, although he says his heritability of asymmetrydown to the wildly inflated estimateof
manuscriptsare rejectedtwice as frequently as beforethe inves- averageheritability. Although they do addresssome of the criti-
tigation. "Het under the microscope,"says former Eztolution cismsof others,theseare, in efiect,dismissedastechnical points
editor Schluter at the University of British Columbia. Yet a look that do not affect the overall conclusions."
at his recent papers showsthat while he is keen on citing his own In closing his review, Houle widens his scopeto include the
work, he rarely cites opposingviews,perhapshoping, as Palmer gullible souls who jumped aboard the fluctuating asymmetry
remarks,that they'll just "go away." bandwagon in the 1990s as well as all scientistswho succumb
Perhapsin response,scientistsremain critical and evenunkind too easilyto the enthusiasmaccompanyingnew ideas."We have
to Moller. In 2OOO,Palmer published an unusual essayin the little choicei writes Houle, "but to seekinspiration from gurus of
newsletterfor the International Societgof Behazsioral Ecology.II the newestideas; sometimesthey turn out to be partially right.
was a fable concerning the fictitious Traumweber brothers, Andy However, we should never believe them without a struggle. If an
and Randy, expert tailors in the'remote kingdom of GliicHichtal, idea seemstoo goodto be true, it is probably not true."
nestledhigh in the EuropeanAlps." Palmer wrote that the maestro Thesedays,Moller's most vocal defenderseemsto be Mous-
of Gliicklichtal's symphony noticed that audiences "seemed seau."I like Rich [Palmer] a loti Mousseausays,"He's a friend
pleasedwith performancesconductedin the taumweber tuxedo, of mine, but he's quite emotional and somewhat irrational in
but dissatisfiedwhen he performedin his imported tuxedo."After his stance: he just doesn't like Moller." Palmer privately wrote
careful investigation, "Andy Traumweber discoveredthe imported Mousseauand cautionedhim not to be so cavalierin defending
jacket was lesspreciselymade,most particularly in the tails: one his colleague.Mousseau,in turn, wrote letters to both Nature
was distinctly longer than the other." The title of the piece, "The and,Sciencewith more than 20 coauthors, defended Msller on
Emperor'sCodpiece,"camefrom its final coup: discussionboards,and started a petition to give Moller back his
According to a palace informer, the Emperor was particu- bird-banding permit.
larly anxious about his imperial private parts, which he felt Pomiankowski saysMoller is in the "limbo land" in which
were so asymmetrical that they deviated too far from the many scientistsinvestigatedfor fraud find themselves."I find it
norm. Fortunately, the Traumweber brothers were able to an unsatisfactory situation to be in, but that's where we are. I
allay his fears with a profound revelation: In certain very would much prefer that he was properly absolvedfor what hap-
specialcases,increasedexpressionofa predictable asym- pened or found properly guilty." He d rather know the truth now.
metry actually signals increased fitness, and one of those "Itt very hard to understand what motivates another person,"says
casesis testicles(Moller 1994,),at least if men are like birds. Pomiankowski.'Youcan concoctan explanationaboutwhythings
That'swhythey subsequentlyfashioned the Emperor'scod- go wrong, but who knows?" s
piece to enhancehis already conspicuousas]rynmetry. H av e a c om m ent? Em ai lus at m ai l @ the- s c i enti sctom

January
2007 TH E S C IE N TIS T 31
 ;:l hen AndersPapeMollerpublishedhis now-
infamous1992 Naturepaperon barnswallows,l
it was a leapof faithto test whetherfemales
might choosemalesbasedon asymmetry
betweenthe left and right sidesof the body.
Suchasymmetryhad beenknownto correlatewith stressand other
environmental factors,but not to inateselectiori.
Moller is a strongproponentof the "goodgenes"model of
s€xualSelection,2.3 which holdsthat certiaincharacteristics, for
example,the tail feathersof a peacock,signalthe presenceof
genesthat will be beneficialto offspring,and thereforemakean
l o r e a t t r a c t i v et o p o t e n t i a lm a t e s .M o l l e r b e g a nto th i n k
i n d i v i d u am
i h a t p e i h a p sm o r e s u b t l es i g n a l so f a m a t e 'sq u a l i t y ,s ucha s ta i l
symmetry,also mattered.a He usedto measureonly the lengthof

of one sideof the tail, but he says,"Therewas one yearthat the swal-
l o w s c a m e b a c kf r o m A f r i c aa n d t h e r ew e r e l o t s o f i n di vi d u a l sth a t
were asymmetric."To tesi his theory,he capturedwild malesand
variedthe asymmetrybetweenthe left and right outermosttail
feathersby 20 mm. Sureenough,maleswith more symmetrictail
feathersreceivedmore visitsfrom females,mated earlier,and had
greaterieproductivesuccess.l
A s A n d r e w B a l m f o r da n d A d r i a nT h o m a si n d i c a t edi n th e i r

controversy replyto Mollei's pape4sinvokingsexualselectionwas not neces-

s a r yt o e x p l a i nw h y b i r d sh a v es y m m e t r i c atla i l s :L o p si d e db i r d s
w o u l d h a v et r o u b l ef l y i n g .G e r a l dB o r g i aa n d G e r a l dWi l ki n so n
foundthai Msller'sstandarderrorswere lessthan one-tenthof
those Moller had reportedpreviously.Moreoveqthe differences
amongtreatmentswere so great,Borgiaand Wilkinsonsuspected
that maleswere assignednonrandomlyto treatments-6 Richard
P a l m e rh a s p q i n t e do u t t h a t M o l l e r 'sm a n i p u l a t i o n os f a sym m e -
t r y w e r et e n t i m e s a s g r e a ta s t h e n a t u r a lv a r i a t i o no f 2- 3 m m .
However,Msller's barn swallowstudieshaveneverbeeninvesti-
gatedfor scientificdishonesty

Rtrerencrs
1. 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 MCRESYMMETRIC
TAILFEATHERS
RECEIVFD
MOREVISITS
FROMFEMALIS,
MATED EARLIER,
ANDHAD
: GREA;IER
REPRODUCTIVE
SUCCESS,

3. 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 sexual ornaments may reliably
revealmale qualityl'Animal Behauiour,4,Otitgi-7, LggO.
5. A. Balmford,A Thomm, "Swallowingornamental asymmetwi'Nature,
359:487,1992.
6. G Borgia,G Wilkinsirn, "swillowing omamental as).rnmetryiNdture,
359t4a7-8,1992.