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Sexual Selection, Physical Attractiveness, and Facial Neoteny: Cross-cultural Evidence and Implications [and Comments and Reply]
Doug Jones C. Loring Brace William Jankowiak Kevin N. Laland Lisa E. Musselman
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Recommended Citation
Musselman, L. E., Langlois, J. H., & Roggman, L. A. (1996). Comment on: Sexual selection, physical attractiveness, and facial neoteny: Cross-cultural evidence and implications, by Doug Jones. Current Anthropology, 37, 739-740.

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Authors

Doug Jones, C. Loring Brace, William Jankowiak, Kevin N. Laland, Lisa E. Musselman, Judith H. Langlois, Lori A. Roggman, Daniel Pérusse, Barbara Schweder, and Donald Symons

This article is available at DigitalCommons@USU: http://digitalcommons.usu.edu/fchd_facpub/602

Sexual Selection, Physical Attractiveness, and Facial Neoteny: Cross-cultural Evidence and Implications [and Comments and Reply] Author(s): Doug Jones, C. Loring Brace, William Jankowiak, Kevin N. Laland, Lisa E. Musselman, Judith H. Langlois, Lori A. Roggman, Daniel Pérusse, Barbara Schweder, Donald Symons Reviewed work(s): Source: Current Anthropology, Vol. 36, No. 5 (Dec., 1995), pp. 723-748 Published by: The University of Chicago Press on behalf of Wenner-Gren Foundation for Anthropological Research Stable URL: http://www.jstor.org/stable/2744016 . Accessed: 09/12/2011 18:16
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CURRENT ANTHROPOLOGY Volume 36, Number 5, December I995 ? I995 byThe Wenner-Gren Foundation Anthropological for Research. rights All reserved ooII-3204/95/36o5-0004$3.oo

Malinowski (i96i[i929]), Berndt (i95i), Weiner (I976), Gregor (i985), Boone (i986), Euba (i986), Munn (i986), i. I thank KimHill oftheUniversity New Mexicofor of assistance Grinker (i990), and Jankowiak (I993). However, existing in in collecting some of the data reported this paperand Davic researchstill suffers fromseveral limitations.First,the Buss,ConradKottak, John and MitanioftheUniversity Michi. workofsocial of is psychologists heavilyempiricaland degan,CarlosAlberto Carosoand MariaHilda ParaisooftheFedera] of and BjarneForsterwald the Puerta scriptive,with little in the way of theorythat would of University Bahia (Brazil), features attractive or Barra Mission(Paraguay) helpwithdifferent for phasesofthestudy explain why people findparticular This research was supported NSF Doctoral Dissertation by Re. even why theyexperiencephysicalattraction all. Secat searchImprovement GrantBNS-goo6394 and by grants from the ond, culturalanthropologists have rarely made research of and University Michigan's Department Anthropology Evolu. of on standardsof attractiveness and theirconsequences a The tionandHumanBehavior Program. Department Anthropol. of the literature access to library computeJ majorobjectiveoffieldwork; ethnographic and ogyat CornellUniversity provided facilities. our relationship appearanceinfluence[s] with others." More recently, therehas been a revivalof interestin the topic of attractivenessand an explosion of social psychological research on the subject, demonstrating significant agreementacross ratersin judgmentsof attractivenessand significant social consequences of atDOUG JONES is Visiting Scholarin Anthropology CornellUni- tractiveness. at This literature has been surveyedat book N.Y. I4853, U.S.A. [dmjs@comell.edu]). Bornin versity (Ithaca, length by Patzer (i985), Hatfield and Sprecher(i986), i959, he was educated at Princeton University (B.A., i98i) and Bull and Rumsey (i988), and Jackson (i992). There is the University of Michigan (M.A., i989; Ph.D., I994). He has materialon standardsof been conducting research standards physical on of attractiveness also considerableethnographic in theUnitedStates,Paraguay, Brazil,and Russia since i989. He attractivenessin non-Westernsocieties, for example,

ers,males seem to be moreconcemedthanfemales withtheatof tractiveness potential sexualpartners, becausehuperhaps in mans showfarmoreage-related variance femalethanin male The selection male attraction markers for to fecundity. resulting offemaleyouthmaylead incidentally attraction females to to cues in an exaggerated This paper displaying age-related form. cross-cultural evidencethatmales in fivepopulations reports U.S. Americans, Russians, Ache,and Hiwi) showan (Brazilians, attraction females to withneotenous facialproportions combi(a nationoflargeeyes,small noses,and fulllips) evenafter female for. studiesshowthatfemale modage is controlled Two further els have neotenous relative U.S. unto cephalofacial proportions and of transformed dergraduates thatdrawings facesartificially to make themmoreor less neotenous perceived correare as moreor less attractive. These results several spondingly suggest the further lines ofinvestigation, between including relationship facialand bodilycues and the consequences attraction neoof to evolution. tenyformorphological

The first publicationbyDarwin to discuss human evolution at length bears the double title "The Descent of Man, and Selection in Relation to Sex" (i98I[i87i]) and consists of two works back to back. The first work discusses human evolution and argues "that man is the modifieddescendantof some pre-existing form"(p. 9). The second presents the topic of sexual selection-a formof naturalselection resulting from"the advantage which certainindividualshave overotherindividualsof the same sex and species, in exclusive relationto reproduction" (p. 256). Darwin yoked human evolution and sexual selection together a single volume because he in believed that sexual selection had played a major role both in the descent of humans fromearlierformsand in the differentiation human races. of Physicalattractiveness its relation thetheory sexualseand to of In the half-century after Darwin publishedthis work, lectiondeserve renewed attention from cultural biological and ana numberofauthorstook up the topic ofphysicalattracThis paperfocuseson an anomaly thropologists. associated with attractiveness-in species,in contrast manyoth- tiveness across cultures. Many, such as Westermarck our physical to

Sexual Selection, Physical Attractiveness, and Facial Neoteny

Five Populations" of (HumanNature4:27i-96) and author the forthcoming Physical Attractiveness theTheory Sexual Seand of lection:Resultsfrom Five Populations University of (AnnArbor: MuseumofAnthropology, press).The present Michigan in paper was submitted XI 94 and accepted2o I 95; thefinalversion I4 reached Editor's the office II 95. 24

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Cross-cultural Evidenceand Implications

byDoug Jones

years ago, Berscheid and Walster (I974:i58)

to relatethe development standards physicalattracof of tiveness in humans to the theoryof sexual selection. Subsequently, however, the social sciences grew indivorcedfromevolutionary creasingly theoryand from the study of physical variation (Degler i99i). Almost 2o summarized (i92i) and Ellis (i926), followed Darwin's lead in trying

the consequences of this divorceforthe studyof physical attractiveness: "Most social scientistshave shown a studied professional disinterest in . . . how our physical

recordsfew if any attemptsto quantifyagreementbe723

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to the "new physical anthropology" the I940S and 'sos, of tween individuals about standardsof attractiveness, consequences of being and the neglect of sexual selection in the earlymodern assess the social and life-history or by perceived as attractiveor unattractive, to compare synthesiswas inherited biological anthropology. Unrelativelyfew serious quantitativestudies standards of attractivenessacross societies. Thus the til recently, in paid attentionto the possible studyof attractiveness the social sciences is underde- in physical anthropology veloped in importantrespects-undertheorizedin psy- role of sexual selection in human evolution. Hulse's and underresearched (i967) studyofskin-color variationin modernJapan chology and both undertheorized suggested that sexual selection mightinfluencethe evoluin culturalanthropology. too, the topic of physical tion of this trait,and a numberof studies of assortative In biological anthropology causes and matingforphysicaltraits(Spuhleri968) suggested and its possible evolutionary attractiveness how This neglect matingpatternsmightinfluencegenotypefrequencies. neglected. consequenceshas been relatively of is partofa widerneglectofthe theory sexual selection However, Hulse's work inspiredlittle commentor folbiologybetween the I930S low-up, and the literatureon assortativemating was in the field of evolutionary and the I970S-a period that West-Eberhard observed. (i983:i56) largelysilent about the causes of the patterns Era" of sexual selection theory(see Thus in spite of Darwin's argumentthat sexual seleccalls "the Forgotten also Cronin i99i). In the I930S and '40s, evolutionary tion played a centralrole in human evolution,serious biologists formulatedthe "modern synthesis"-a syn- quantitativestudiesofadaptationin physicalanthropolon thesis of Darwin's theoryof evolutionand the new sci- ogy have focused overwhelmingly adaptationto the Only recently, ence of genetics.The pioneersof the modernsynthesis physical environment. with the rise of adaptationto ecologi- human behavioral ecology,have anthropologists had theirhands fullinvestigating begun to They were less concernedwith the evo- trying bringthe moderntheory sexual selectionto cal constraints. of lution of social behaviorand had littleuse forDarwin's the studyofhuman evolution(Chagnonand Irons I979, Betzig,Turke,and Borgerhoff Mulder i988). But to date, theoryof sexual selection. Why did the theoryof sexual selection take so long most such studies have focusedon sexual selectionand to win acceptance in evolutionarybiology?Sexual be- mate choice more in relation to behaviorthan in relahavior and social behaviorin generalpose special prob- tion to morphology. In conclusion,the studyof physicalattractiveness lems for evolutionarytheory. These problems result in of fromthe fact that in social evolution the fitnessof a the contextofthe theory sexual selection,which was trait commonly depends on its frequency(Maynard givena centralplace byDarwin,Westermarck, Ellis, and Smith i982). How well a particularshape of tail serves has not been a major topic foranthropological inquiry It will generallynot depend on the in the last half-century. is not that investigation a given bird in flight has shown attractiveness be ill-defined inconsequento in or frequency the population of tails of various shapesBy thatis, its fitness frequency-independent. contrast, tial; both social psychological studies and the ethnois of the attractiveness that tail to membersof the other graphicliteraturesuggestat least moderateagreement of within cultures and at sex will depend on the preferences the other sex, in judgmentsof attractiveness which will oftendepend in turn on the frequenciesof least a moderatelyimportantrole forattractiveness in various shapes of tail-that is, its fitnessis frequency- social and especially sexual interactions. Rather, particular theoretical have keptboth cultural dependent. presuppositions fromgivingthese topics selection presentsa numberof and biological anthropologists Frequency-dependent selec- theirfull attention.Cultural anthropologists have been seeming paradoxes. Under frequency-dependent tion there is no guarantee that natural selection will reluctantto deal with the more "biological" side ofhufavorgenetic variantsthat maximize mean population man behavior-reluctant to considerhuman behavioras for fitness.Instead,populationsmay attainan evolutionary theproductnot just ofstrivings individualgainssuch and social status but of adaptations equilibriumin which no individualcan gain byadopting as materialcomfort a different even thoughall would be betteroff forgeneticreproduction. have strategy, Biologicalanthropologists if all acted differently. they may enter an endless oftenheld a view of adaptation that could not readily Or "arms race" in which each triesto get ahead of the oth- accommodatefrequency-dependent selection,including gains. Fre- the possibilitythat one individual's reproductive ers withoutanyone's enjoyingany long-term sucselection may favor traits that in- cess mightcome at the expense of others. quency-dependent The theoryof sexual selection has advanced so farin success ofindividualsbut reduce crease thereproductive of the viabilityofgroupsand lower theproductivity eco- recentyears that it may be time forrenewedattention systems.It may favorthe evolutionofwaste and extrav- to the relationshipbetween sexual selection and stanin agance, ratherthan efficiency, sexual and othersig- dards of physical attractivenessin our species. The nals. It may result in coevolutionary positive feedback theoryof sexual selection does not imply that people in traits.It was only begin- always maximize inclusive fitness choosinga matecycles that amplifyarbitrary and in humans as in of ningin the I96os,with the development sociobiology adaptations are often imperfect, thattheproblemsassoci- otheranimals estheticstandards and evolutionary the game theory, may be partly prodselectionwere addressed uct of nonadaptive"sensorybiases." Nor does it imply ated withfrequency-dependent that standardsof beauty are completely"hard-wired"; in a sustained fashion. The modernsynthesisof the I930S and '40s inspired in humans as in otheranimals the development stanof

JONES

Sexual Selection and Facial Neoteny | 725
(Daly and Wilson i983:279-3i2). Human

dards of attractivenessis likely to involve a range of to mechanisms, from innate templates to imprinting imitationand otherformsof social learning.It does imply,however,thathuman beings,like otheranimals,are likely to have genetic adaptations for assessing the "mate value" of potential mates and that studying attractivenesswithout considering these adaptations would be like tryingto understandthe eye without it treating as an organof vision.

Sexual Selection and Physical Attractiveness: A Human Anomaly
Sexual selection occurs when some organismsgain an at edge in matingand fertilization the expense ofothers of theirsex. Sexual selection may take the formof contestswithinone sex over chances to mate with and fertilize membersof the othersex or courtshipof one sex by the other.2 In many animal species, male reproductive success is moredependenton matingsuccess thanis femalereproductivesuccess, so sexual selectioncommonlyacts with greaterintensity on males than on females (Trivers

of males more than females show a syndrome traitsas- i992:65-67). This sex difference not limitedto Westernsociety. is sociated with intense sexual selection.This "sexual se37 lection syndrome"includes behavioraltraits:males are Buss (i 989) reviewssurveydata from populationsammore likely than females to resortto violence against ples from33 countriesand findsthat in everysample part- males are more concernedthan femaleswith the physisexual rivals and to forcecopulations on resisting of ners; males cpmmonly expend more time and energy cal attractiveness a potential mate. The average sex is risks than femalesin courtship;males difference more pronouncedamong the non-Western and take greater will generally courtand attempt copulationwitha wider populationsin his sample. While Buss's studyincludes holds apparently thenwill females.The sexual selection no tribalpopulations,the same pattern rangeofpartners forthese as well. Fordand Beach (I 95I:94), summarizing traits: males comsyndromealso includes life-history monly take longerthan females to attain sexual matu- evidence from nearly 2oo cultures, conclude that alvariation ritybecause of the sexual competitionthat they face though there is a great deal of cross-cultural "in most societies the frommaturemales; males commonlyhave highermor- in standards of attractiveness, talityrates than females as a resultof intrasexualcom- physicalbeautyofthe femalereceivesmoreexplicitconpetition; males commonly senesce more rapidlythan siderationthan does the handsomenessofthe male. The ratesreducethe selec- attractivenessof the man usually depends predomifemalesbecause highermortality thanupon his tion pressureforlongevity. Finally,the sexual selection nantlyupon his skills and prowessrather physical appearance." Gregersen(i983) reportssimilar traits:males are more includes morphological syndrome in likely than females to display anatomical specializa- findings a more recentreviewofnearly300 societies, and nonurbanized. tions for intra- and intersexual aggression,including mostlynon-Western In otherwords,human beingsseem to be an exception horns,antlers,enlargedcanine teeth,and body sizes in excess of the ecological optimum; males commonly to the general rule among animals that male attracThe show greater development of sexual advertisements, tiveness mattersmore than female attractiveness. bothtactile (complexgenitalia)and visual (elaborateand importance attached to female (as opposed to male) in physical attractiveness our species stands in need of colored adornments). brightly an explanation. Among humans, considerableanatomical and behavioral evidence suggeststhat males have been subject to sexual selectionthanfemales,althoughthe dif- An stronger Fecundity, Age,
--- 1r , +lfe nn"r%h"er

and i992, I97I, WilliamsI975, Clutton-Brock Parker Andersson The resultis thatin manyspecies, I994).

males are largerthan females.Human males attainsexat ual maturity a later age than human femalesand seIn nesce more rapidly. all societieswith appreciablelevels of violent conflict,male-male aggressionis more Violentcompecommonthanfemale-female aggression. tition is more common among human males than among females,and male sexual coercion of femalesis is farmore common than the reverse.Polygyny much And in most respects, more common than polyandry. human females seem to be more selective than human males in theirchoice of sexual partners. In one respect,however, human beings reversethe between more and less sexusual patternof differences ually selected sexes-men are more concerned than of women with the physical attractiveness a potential has sexual partner.This sex difference been foundrepeatedly in studies by social psychologists.A recent meta-analysis (FeingoldI 990) ofthe social psychological in of on literature sex differences effects physicalattractiveness on romantic attractionshows consistentand strong sex differences, including content analyses of "lonely-heartsadvertisements,"studies of attractiveness and reporteddating success, reportsof interpersonal attractionfollowingdyadic interaction, and surdesiredin a mate (see also Jackson veysofcharacteristics

mammals

han"r

Anomaly Explained? and Neoteny

third form sexual selection, of sexual coercionof one sex by the other.

2. Smuts and Smuts (I993) argue that it is useful to distinguish a

believe that human behavior is Many anthropologists in fromthat of other so radicallydifferent its ontogeny organismsthat the theoryof sexual selection is not ap-

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plicable to human physical attraction. Polhemus (i988:8) probably expresses the attitude of a whole of school ofanthropology "the body" concerning huthe man irrelevanceof the theoryof sexual selection when he writes: A male baboon has a fixedidea of what a desirable femalebaboon should look like.... The same general principleis true of any animal that reproduces by sexual selection. But thereis an important difference between baboons and ourselves.For otheranimals the physicalideal is ioo% instinctively determined.Thus all baboons of a particular species pursue the same ideal.... For humans,on the other hand, ideals of beautyare learned.... In a worldwide and historicalframework, thereis no such thingas naturalhuman beauty. Ifthisview ofthe difference betweenhuman and nonhuman psychologywere correct,the anomaly of female in attractiveness our species mightbe merelyone more consequence of our havingfreedourselvesfromthe instinctiveconstraints that hobble the lives of otheranimals. This view, however,is doublywrong. First,learningoftenplays a large role in the acquisition ofstandards attractiveness of amongnonhumananimals. An immense literaturedemonstrates that early experienceinfluenceslater mate choice via imprinting (Immelman I972). Imitation,too, plays a role in mate choice among nonhuman animals, and social transmission of matingpreferences can even resultin "fads" in mate choice that change fromone breedingseason to the next (Pruett-Jones i992). in Second, physical attraction humans cannot be entirelya productof enculturation.This is shown most of dramatically the experiments Langloiset al. (i987). by In these experiments, infantsbetween the ages of two and three months were exposed to picturesof women ratedattractive and unattractive adult raters;infants by spent more time looking at faces ratedattractive. This held even across racial/culturalboundaries: for European-American infants looking at faces of AfricanAmerican women rated by African-American men and for African-American infants exposed to EuropeanAmericanfaces ratedby European-American men. Thus students of physical attractiveness are asking fortrouble if they start out assuming that nonhuman animals are creaturesof instinctand humans constructions of culture. A betterstartingpoint regarding the role of learningin behavioris suggestedby several decades of researchin comparativepsychology:as a general rule,organisms have relatively "hard-wired" canor alized responses to stimuli that have had relatively fitnessconsequences over evolutionary time unvarying and relativelyflexiblelearnedresponsesto stimuli that have been associated sometimes with positive fitness consequences and sometimes with negative. In other words,giventhatlearningentails costs,in termsoftrial and error, organismsare expectedto adapt to selectively invariantsin theirenvironments with correimportant spondingbehavioral,cognitive,or motivationalinvariances (Seligman I970, Johnston 1982).

How can we apply this principleto the anomaly of in female attractiveness our species? Let us definethe mate value of a potential sexual partner, as the exA, pectedreproductive success from matingwithA divided by some baseline expected reproductive success. The baseline expectedreproductive success mightbe the expected reproductive success frommatingat randomor from mating with an individual of maximum mate value. As a generalrule we expect that human beings, and otheranimals,are likelyto have bothrelatively canalized, "hard-wired"responses to visual stimuli that have been consistently associated with highmate value the throughout evolutionary historyof the species and flexiblelearnedresponsesto stimulithathave relatively been associated sometimes with high mate value and sometimeswith low. In otherwords,standards physiof cal attractiveness likelyto have both species-typical are and population-specific components,and variation in these componentsmay be predictablegiven knowledge of human biology and local circumstances (Symons I979). For example, since fat stores may be selectively advantageousin environments subject to episodic food in and disadvantageous environments shortage requiring considerablephysicalmovement,one mightexpectthat estheticresponsesto fatnesswould varybetweenpopulations dependingon social learningand on individual assessments of the consequences of being fat or thin, ratherthan developingin a uniform fashionwithinthe human species. one mightexpect human beings to have By contrast, a relatively invariant, species-typical emotional response to signs of aging, because age has a relatively invariant association withfecundity thuswithmate and value. In a classic article Henry (i96i) reviewsdata on rates in a wide range of "naturalage-specific fertility fertility" (noncontracepting) populations. The levels of in averfertility these populationsrangefroma lifetime age of 6 to i i children per marriedfemale, but the shapes of the curves of fertility versus age are remarkably similar across all populations. For all populations, female fertility rates at age 30-34 are around 85% of ratesat age 20-24, with further declines to around35 % for women aged 40-44 and o% forwomen aged 50-54. More recentwork suggeststhat the curve of naturalfediffers cundity(potentialreproduction) somewhatfrom the curve of natural fertility be(actual reproduction) cause the latteris influencedby such variables as age of spouse and frequencyof intercourse(James I979, Menken,Trussell,and Larsen i986). Studiesthatcontrol [orthe lattervariablessuggestthatthe decline in female between 2o and 35 is less pronouncedthan Fecundity the decline in female natural fertility-butthe overall similar. shapes of the two curves are fairly The shape ofthe curveoffecundity versusage is very different males. Goldman and Montgomery for (i989), data fromseveral traditional reviewing societies,report declines to about 90% formen between 45 and Eertility men, and to about 8o% formen 50, relativeto younger for 55,aftercontrolling age ofwifeand durationof Dver marriage. Fecundityversus age curvesthus have two important

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Sexual Selection and Facial Neoteny 1727

in characteristics that may help to explain the anomalyof intermediate juvenility.(It was judged to be 23 years female attractiveness:the curves (i) are relativelyin- old.) In otherwords, the level of juvenilitythat maxidoes not maximize pervariantin shape across populationsand (2) show an ear- mizes perceived vulnerability lier and more pronounceddecline in fertility among fe- ceived sexiness. Kenrickand co-workers (KenrickI994) males than among males. Given the general rule that show that forteenage males the ideal sexual partner is organismscommonlyhave invariantresponsesto stim- older than they are-again, more consistentwith the thatmales are concerned uli that have had relatively invariant fitness conse- hypothesis withcues to female quences over evolutionary time, the firstcharacteristic fecunditythan with the hypothesisthat males prefer more easily dominatedfemales.Thus current suggeststhathuman beingsare likelyto have relatively younger, invariantestheticresponses to signs of aging.The sec- evidence suggests that female attractivenesscannot ond characteristicsuggests that these responses are simply be equated with powerlessnessand that somein likely to be stronger males' evaluations of females thing more than changes in perceivedvulnerability is involved in age-related changes in physical attracthan in females' evaluations of males. This does not add up to a completetheory physical tiveness.However,nothingin evolutionary of theory rules of attractiveness, course, or even a complete theoryof out the possibilitythat markersof female submissiveage-related changesin physicalattractiveness. Fecundity ness may be attractiveto men, and the topic certainly is onlyone componentofmate value. Othercomponents deservesmore research. include the abilityand willingnessto provisionoffspring There may be room forargumentabout why attracand heritableviabilityor attractiveness ("good genes"), tivenesschangeswith age, but,in spiteofa considerable devotedto the claim thathuman sexualityand and these componentsofmate value may also varywith literature age, while sensorybias will ensure that attractiveness standardsof physical attractiveness culturallyconare theredoes not seem to be any evidence from does not trackmate value perfectly. Nevertheless,age- structed, related changes in fecundity likely to be a particu- any society that seriously challenges the proposition are larly importantcomponent of age-relatedchanges in thatphysicalattractiveness perceivedto decline from is physical attractiveness, especially in females,both be- youngadulthoodto old age, especiallyforfemales."The of cause these changes have been relatively invariantover correlation femaleage and sexual attractiveness so is obvious that ethnographers the history the species and because othercomponents intuitively of apparently take of mate value such as provisioning abilityand inclina- it forgranted-as they do the bipedalismof the people tion may be more readilyassessable on the basis of be- theystudy-and the significance femaleage tends to of be mentionedonly in passing,in discussions of somehavior than on the basis of physicalappearance. There is one alternative explanation for male at- thingelse" (SymonsI979:i88). Symonscitespassingrefof tractionto youthfulfeaturesin females that requires erences to the effects aging on female attractiveness in ethnographies the Kgatla,pre-revolutionary a more extended treatment. Gowaty (I992:23I-40) of China, the Yanomamo, and the Tiwi. Additionalreferences writes: can be foundin ethnographies TrobriandIslanders(Maliof There should be strongselection on males to control nowski I987 [i929], WeinerI976) and Gawa (Munn females' reproduction directcoerciveconthrough I986) ofMelanesia, Mende (Boone i986) of SierraLeone, trolof females.... Evolutionary whether thinkers, and Mehinaku ofAmazonia (Gregori985), to name just informed feministideas or not, are not surprised by a few. A number of social psychological studies (refactsof patriarchal by one of the overwhelming culviewed in Jackson i992) have documented such agetures,namely that men ... seek to constrainand related declines in physical attractiveness and demoncontrolthe reproductive capacities of women.... Justratedthe expectedsex differences well. as venilizationdecreases the threatsome men may feel Let us summarizethe argument to this point.Huup when confronted with women; many men are comman beingsare anomalous among sexuallyselectedspefortable aroundwomen whom theycan clearlydomicies in the importanceattached to female (relativeto nate and are profoundly uncomfortable around male) appearance in mate choice. Human beings are women whom theycannot so clearlydominate.The anomalous in anotherrespect as well: female fertility hypothesisthatfemininity signals abilityto be domicommonlydeclines to zero long beforethe end of the nated through juvenilizationis an alternative but life to, span. As a resultofmenopause thereis considerably not necessarilymutuallyexclusive of,otherevolumore age-related variance in fecundity among adult fetionaryhypothesesthat posit thatfemininity sigmales than among adult males in our species. The secvalue and reproductive nals, sometimes deceptively, ond anomaly may explain the first: the importance fertility. attachedto female attractiveness our species may rein seem to be at odds withthishypothesis. flect the operation of adaptations for assessing ageSeveralfindings and McArthur a Berry subjectswith a se- related changes in fecundity, component of female (i986) presented ries of outline profiledrawingsrepresenting individuals mate value. Whetherforthis reason or another,social evidence providesoverrangingfromjuvenile to adult and collected ratingsof psychologicaland ethnographic for support theproposition thathumanbeings perceived social characteristicsof each drawing.The whelming was the have relativelyinvariantestheticresponsesto signs of drawingrated weakest and least threatening agingand thatthese responsesoperatemore strongly in most juvenile-looking. (Subjects judged this drawingto males' evaluations of femalesthan vice versa. The drawingrated sexiest was representa 4-year-old.)

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Thus far we have been exclusively concernedwith changes in attractivenesswith age ratherthan differbetweenindividualsofthe same ences in attractiveness mechanismsdo not operage. However,ifage-detecting accuracy,thenadaptationsforchoosing ate with perfect to a mate of a particularage may lead incidentally nonadaptivebiases in the choice ofmates fromamongindividuals who fall within a particularage-class. In other varieswithage,individwords,giventhatattractiveness uals may be more or less attractivethan othersof the same age in part because they have facial proportions or associatedwithyounger olderages. Because theretention oftraitsfromearlystagesofthe lifecycle into later stages,relativeto ancestorsor to othermembersof the population, is known as neoteny ("holding on to given youth"),the propositionabove may be rephrased: varies with age, neotenymay be a that attractiveness component of facial attractiveness.This proposition may hold with particularforceforfemale facial attracof tiveness: a by-product the human male's attraction fecundity may be an attraction to markersof youthful of markers youthto an exagto adult femalespresenting geratedor "supernormal"degree. with the anomaly of female attractiveness Beginning thatneoteny in our species,we are led to the hypothesis The may be a componentoffemalefacialattractiveness. of remainder this paperwill be givenoverto testingand this hypothesis. elaborating

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4

k=O.1

Cephalofacial Age Markers changesin physicalattracTo considerhow age-related estheticresponses tiveness mightresult fromdiffering to differing facial proportions, is necessaryto review it how the sizes and shapes of facial featureschangewith age. It is convenient to divide these changes into changes in hard tissue (bone) and changesin softtissue (cartilageand connectivetissue). Hard tissue. In human beings as in othermammals, the neurocranium-the portionofthe skull housingthe brainbut also includingthe contiguousorbitalregiongrows rapidly early in development,while the facial skeleton proper-including the nasal and masticatory complexes-attains its maximum rate of growthonly later (Enlow iggo). As a result,juvenile mammals present a characteristic "cute" appearance,with relatively and reducedsnouts. large eyes, high foreheads, skeletonassoChangesin the shape ofthe craniofacial ciated with aging are closely approximated a simple by mathematical transformation called cardioidal strain (Mark, Shaw, and Pittengeri988). Cardioidal strainof degreek maps each point {x, y} onto {x', y'} = {x (i k y/r), (i - k y/r)}where r2 = x2 + y2. A shape y subject to positive cardioidal strain (k > o) shows a downward and outward expansion in featureslocated towardthe bottomand a downwardand inwardcontraction in featureslocated towardthe top. Negative cardioidal strainpresentsthe opposite changes. Pictures of the faces ofchildrenor youngadults subjectedto a posi.

FIG. i. The effects negative (top row) and positive of on (bottomrow) cardioidal transformations a 5 x 5 grid and on -afemale face.

are tive cardioidaltransformation perceivedas olderand less cute; pictures subjected to a negative transformation are perceivedas youngerand cuter.Full facial and of and dogs profile drawings theheads ofbirds, monkeys, and even front and side drawingsofVolkswagenBeetles can be made to appear more or less "mature" or "cute" by subjecting them to positive or negative cardioidalstrain.Figurei illustrates effects positiveand negthe of ative cardioidal strain on a square grid and on a face. The transformed gridswere producedwith the Mathematica a2.2 softwarepackage, while the transformed faceswere redrawn from originalfacewith the assisthe tance of polar coordinategraphpaper. Attractionto "cute" proportions may be unlearned: even at 4 monthsof age infantsorientpreferentially toward pictures of infant rather than adult facesinfantile facialproporalthoughit is not knownwhether tions per se are the relevantcue (McCall and Kennedy I have cited research(Langlois et al. i987) showi980). as ing thatinfants youngas 2 monthsofage orientpreftowardattractive ratherthan unattrattive feerentially male faces. If, as I will argue,female attractiveness is

JONES

Sexual Selection and Facial Neoteny | 7e29

partly a matter of cephalofacial neoteny,then infant for preferences attractivefemale faces may be part of a more general attractionto faces or facelike stimuli low cardioidalstrain.3 manifesting Softtissue. Skeletal growthslows down (butdoes not of stop [Behrents i985]) with the attainment adulthood. However,otherchangesin facialproportions resultfrom the growthof cartilage and the atrophyof connective tissue. These affect relativesizes ofeyes,noses, ears, the and lips. "Beginningat age 25, the eyebrowssteadily rim descendfroma positionwell above the supraorbital to a point farbelow it; saggingof the lateral aspect of the eyebrowsmake the eyes seem smaller" (Larrabee steadily throughoutadulthood: ears get bigger, and with innoses get longer,wider, and more protrusive creasingage. Withthe loss ofconnectivetissue,the vermilion or red zone of the lips gets thinner (Enlow I990,

and Makielski I993:I4).

tissues grow Cartilaginous

As a resultofchangesin hardand softtissue with age, it is possible to estimate ages of adults using information about the relativesizes of eyes, noses, and lips.

Larrabee MakielskiI993, SusanneI977). and

Neoteny and Attractiveness

A youthfulor neotenous face is one that combines a high ratio of neurocranialto lower-facial featureswith a small nose and ears and full lips. The appendixgives a summaryof a numberof studies ofneotenyand facial attractiveness. all Virtually ofthemfindthatneotenous as facialproportions, defined above,contribute female to Resultsformales are equivocal. One limattractiveness. itationofthese studiesis thattheyare confined Westto ernsocieties or societies strongly influenced Western by ideals of physical attractiveness. For example, Wagatsuma (i968) shows thatcontactwithEuropeansand U.S. Americans has had a significant influenceon Japanese standardsof attractiveness over the past hundredyears. Because the line of argumentpresentedabove suggests to thatattraction femalecephalofacialneotenyis a good candidate fora human universal,it is important esto is tablishwhethersuch attraction characteristic nonof Westernizedsocieties as well. This paper reports results from an ongoing crosscultural study of criteriaand consequences of physical STUDY I: AGE PREDICTORS AS ATTRACTIVENESS attractiveness one ofwhose aims is to investigate possi- PREDICTORS ACROSS CULTURES ble universals of attractiveness collectingdata from by of dataon standards I989 and i992 I collected as wide a rangeof populationsas practical.Populations Between in studied to date include two relativelyisolated indige- physicalattractiveness fourpopulations:U.S. Amerinous South Americangroups,the Ache (or Guayaki) of cans, Brazilians,Russians, and Ache Indians. Kim Hill, at currently the Universityof New Mexico, assisted with data collectionamongthe Ache and collectedaddifor tional data among the Hiwi. The researchmaterialremaybe a releaser in 3. Neotenousfacialproportions children of parental behavior (Lorenz I943). McCabe (I988) reviews studies ported on in this paper includes facial photographs to and individualsin threepopulationsand interview a willingness nurture thatsubjectsreport greater showing data and to with a highratioof neurocranial lower-facial ratingsof attractiveness facial photographs children protect of fromfive She also cites researchconductedat severallocalities features. derivefromundergradpopulations.Facial photographs undercourtprotection showingthat abused children (ages 3-6) dimensions-that uates at the University of Michigan in Ann Arbor, to havelowerratiosofneurocranial lower-facial groups. studentsat the FederalUniversity Bahia in Salvador, schoolcontrol of nursery is, areless cute-than age-matched

eastern Paraguay and the Hiwi (or Cuiva) of southern Venezuela, as well as three Westernized societiesBrazil,the United States,and Russia. Each of the Westernized populations is the subject of a large body of ethnographicliterature;below I provide a very brief summary of some facts relevant to physical attractivenessin the two indigenousSouth Americanpopulations. The Ache and the Hiwi were first peacefullycontactedby outsiders the I96os and I970S; up to that in time theyhad lived as hunter-gatherers. Membersofthe two populations know what outsiders look like, but most have little contact with outsiderson a day-to-day basis. Both groups maintain a stronglyethnocentric standardof physical attractiveness. For the Ache, Kim Hill (personalcommunication)writes,"The Ache have commentedon how ugly Europeansare parfrequently because oftheirlong noses (theycalled us pyta ticularly puku-long nose-behind our backs) and because they are so hairy." These responses are similar to those reportedby Wagatsuma (i968) forthe firstgenerationof Japanese exposedto contactswithWesterners. Although the Ache and the Hiwi have had little or no contact with Asians or Asian-Americans, theyare curiousabout of photographs East Asian faces, generallyattractedto them, and aware of the similarity between these faces and theirown. A previous study (Jonesand Hill I993) showed much stronger agreementin ratingsof attractivenessamong the threeWesternsocieties in the sample (mean correlationsin ratingsof attractiveness = r and Ache and Hiwi. The .64) than between Westerners lattercorrelations were still significantly positive (r = . I 8), however, a suggesting universalas well as a culturespecificcomponentto standardsof attractiveness. Below I presentthreestudies addressingthe topic of neotenyand femalefacial attractiveness. Data formales are included forpurposesof comparison.Since previous studiesofneotenyand facial attractiveness show a positive relationshipforfemales and an equivocal relationship formales, I use one-tailedstatisticaltests forfemales and two-tailed tests for males. The firststudy includes data fromall five populations. The othertwo studies are more preliminary, further but is fieldwork planned to test the artificialstimuli of the thirdstudy in a wider rangeof populations.

730

CURRENT

ANTHROPOLOGY

Volume 36, Number S, December I995 and Jones(n.d.)providefurther discussion ofproceduresinvolvedin taking, and rating, measuring photographs. For each photograph each populationsample I calin culated thefollowing measuresoffacialproportions: eye width(EW = mean of D [left left endocanthion, exocanthion] and D [rightendocanthion,rightexocanthion]), nose height (NH = D [glabella, subnasale]), and lip height (LH = D [labiale superius, labiale inferius]), where D (a,b) is the Euclidean distance between photographiclandmarksa and b, and landmarknames follow definitionsin Farkas (I98I). For the analysis below I have dividedeach measure by face height(FH = D [glafor bella, gonion])to correct differences sizes offaces, in producingthree indices of facial proportions:relative eye width (EW/FH),relativenose height(NH/FH), and relativelip height(LH/FH). These indices were selected to measure the relative sizes of the threemajor facial below that features-eyes, nose, and lips. I demonstrate they do vary as expected with age-eye width and lip heightdecrease, and nose height increases. Two other featuresthat mighthave been included,ear heightand cheek width,are omittedfromthis analysis: ear height was difficult measure fromphotographs, to and cheek width did not show a consistentrelationwith age. Table i presentssummarystatisticsforthese three indices of facial proportions and correlationsbetween these indices (each subject to log transformation) and for ratingsof physical attractiveness different combinations ofphotographic subjectsand raters.The table provides some support the hypothesized for berelationship tween neotenous facial proportions and attractiveness, especiallyforrelativeeye width.However,simplypiling and reporting up a list of measures of facial proportions their correlationswith attractiveness ratingsprovides only a weak test of the neoteny hypothesis.A better fromdifferent cues, is test, combininginformation age presentedbelow. This analysis proceeds in two stages, the first resultingin several equations that can be used to calculate the predictedage of each photographic subject on the basis of relative sizes of facial featuresand the second using these predictedages to producean index of neoteny. Age-predictor equations. I begin by using stepwise multivariateregressionto produce equations that predict age as a functionof relativesizes of facial features. Forthis stageI use onlyphotographs Ache, since Ache of span a much widerrangeofages than eitherofthe other two photographic sample populations. Using the stepwise multiple linear regression routinefromSystat s.o with bothp to enterand p to discard set at .I 5 produces the followingequation:
(I993)

data were col tographs and questionnaire/interview lected froma second sample of University Michigai of frommiddle-and lower-classresident undergraduates, of Salvador,Brazil,from nativesofanotherAche village from studentsat the Russian StateUniversity the Hu of manities in Moscow, and froma Hiwi settlement. U.S. Americanphotographic subjects and raterswere recruitedin introductory anthropology and psycholog, courses and by flyersposted on campus. For Braziliai and Ache photographicsubjects, photographicequip mentwas set up in public places, and interested individ uals were invited to participate. Brazilian, Russian were recruited goingfrom Ache, and Hiwi raters by doo, to door and by approachingpotential ratersin publi4 places. U.S. photographic subjects and raterswere largelyo European ancestry;attractiveness ratingsof and fron and African-Americans omittedii Asian-Americans are this analysis because a restricted sample providesa bet ter test of hypotheses. Brazilian subjects and raterc identified themselvesas beingofmixedancestry largely mostlyAfricanand European with some Indian ances try.(The issues raised forthe studyof physical attrac tivenessby Brazil's combinationofrace mixtureand ra cial stratification cannot be treatedat any lengthher( but are discussed in Jones[n.d.].) Russian raterswere largelyofRussian nationality with some othernational ities of the former Soviet Union presentas well. Mear Brazilian,U.S. American,and Ache femalesand 24, 2I and 32 forBrazilian, U.S. American,and Ache males age ranges were I7-34, i8-25, I4-5I, I9-32, I8-30 and i6-6o.

brazli, and natives ot two Actlevillages. Katingsot phO

were 23, ages of photographic subjects

20,

and 29 fo:

Facial photographs were taken indoorsin the Unite( States and outdoors in Brazil and Paraguayat a fixec distance in a standardposition. Ratingsof photograph! were collected by having ratersrank subsamples of a photographic sample population.For Brazilianand U.S Americanphotographic samples, new subsampleswere drawn at random for each rater.Because Ache photo. graphicsubjectsspanneda widerrangeofages,individuals fromthe Ache photographic sample were assignec to fixed subsamples with others of similar age. Each raterratedphotographs membersof the opposite sex of drawn froma single photographicsample population; ratings of all three photographicsample populations were collected in each of fivepopulationsof raters(ex. U.S. Americans).The attractiveness ceptforHiwi rating relativeto a given popula. ratingof a given photograph tion of ratersis the mean attractiveness ratingof thai across ratersfromthatpopulation,with age photograph of photographicsubject partialled out. For Ache, re. ported correlationsbetween attractiveness ratingsanc relevantvariablesare means of correlations withinsubsamples (more exactly,means calculated using Fisher's and z-transformation its inverse [Sokal and Rolf I969: An Apple Scanner connected with a Macin. 520-23]). tosh II was used to measure{x, y} coordinatesof a num. ber of facial photographiclandmarks. Jones and Hill

Predicted Age i

=

-

141

log[EW/FH] log[LH/FH]- i28.

-62

(I;

females)

The variance accounted for(R2) is .23. Equation i predictsages ofAche femalesas a function of relative eye width and relative lip height; relative nose height drops out of the regression.However, an

JO N E S

Sexual Selection and Facial Neoteny | 73 I

TABLE

and RatingsofAttractiveness Correlationsbetween Measures of Facial Proportions
Males Rating Females FemalesRatingMales EW/FH
-.09

I

of Photographs
Brazilians

Ratedby
Brazilians (i 9, I I) U.S. Americans (i2, 20) Russians (iI, I4) Ache (iI, I3) Hiwi (4, 4) Mean

EW/FH
.39
.I9

NH/FH
.0I
-.I4

LH/FH
.24 + -.07 .12

NH/FH
-.I9 -*4I -.43+

LH/FH
.I7 .42+

N

= 49, 20

-.35
-.IO

.32

.I3

S.D.

.43 -* .28* .2IO
.OI4 .23

.29* -.I9 .589
.030

.23 .2I .I25
.020 .I7 -.04 .03

-.07 .03 .I85 .oi8
.00 -.03 .07 -.II

-.I5 -47* .585
.033

.27
.IO

53* .I35 .027 .04
-.I5 .05 -.0I

U.S. Americans N = 5I, 35

Brazilians (2o, 23) U.S. Americans (ii, Russians(I2, I4) Ache (2o, 2i) Hiwi (o, o)
Mean

-.26+
.04 -.20

i8)

n.a.
.225 .oi8

.30* .2 I .25 +

.i8 .02
.09 -.04

n.a.
.58I
.024 .00 .I9

.IO

n.a.
.I1I

.13

n.a.
.OI3 .2 .22 .I3 .2I

S.D.

.202

n.a.
.574
.025

n.a.
.ii8
.022 .I9 .I8 .I4 .14

.022
.IO

Ache N = 4I,

36

Brazilians (I7, I6) U.S. Americans (i2, Ache (i5, I5) Hiwi (7, 4)
Mean
Russians (I2, I2)

-.I2 i5) -.09

-. .26 -*99+ -.43 - .I7

.o8
-.03 .00

-.22
.I5

.3 I - .32 .599
.035 -.02

S.D.

.I9 .I89
.OI 3

.49 .i28
.023

.3I+
.I77 .OI3

-.II .607
.032

.0I II8
.029

in Numbers parentheses are NOTE: EW/FH, eye width/face height; NH/FH,nose height/face height; LH/FH,lip height/face height. of for numbers raters femaleand male subjects.
+p < .I *p < -O5

Indices of neoteny and facial attractiveness. neoA equation can be produced with nearly the same pretraitsorpresents dictivepoweras Equation i byleavingrelativelip height tenousfaceis one thatretainsyouthful of markers youthin an exaggerated form relativeto othThe resulting equation is: out of the regression. ers of the same age. Equations I to 3 can be used to = PredictedAge2 - io8 log[EW/FH] (2;females) produce indices of neoteny: I definefacial neotenyas the difference between the actual age of a face and the + I39 log[NH/FH] - I7 age predictedby one of the equations above. In other words, The variance accounted for(R2) is .17. In otherwords,ages ofAche femalescan be predicted Neoteny = Age - Predicted i i Age (4; females) as a functionof relativeeye widthin combinationwith eitherrelativelip heightor relativenose height. Neoteny2 = Age - Predicted Age 2 (5; females) of For Ache males, stepwise multiple regression age on relativeeye width,relativenose height,and relative Neoteny3 = Age - Predicted Age 3 (6; males) lip heightproducesanotherequation: Thus a face with unusually large eyes, small nose, and (3 males) full lips in relation to face heightwill have a low predictedage accordingto Equations I to 3 and a highindex -6i log[LH/FH] - I36. of neotenyaccordingto Equations 4 to 6. Table 2 shows correlationsbetween indices of neoThe variance accounted for(R2) is .55. comHowever, when relative lip height is excluded from teny and ratings of attractivenessfor different When corsubjectsand raters. two variablesdo not yield binationsofphotographic the the regression, remaining equation. Thus we are left relation coefficientsfor two differentsamples are predictor anothersignificant similar,the samples can be treatedas a sinequations forfemales sufficiently with two alternative age-predictor coefficients pooled (Sokal gle sample and the correlation and one formales.

Predicted 3 = Age

-

I46

log[EW/FH]

732

CURRENT

ANTHROPOLOGY

Volume

36, Number

S, December

I995

TABLE

2

Correlations

between

Indices

of Neoteny

and Ratings of Attractiveness

Males Rating Females of Photographs
Brazilians

Females Rating
Males

Ratedby
Brazilians
U.S.
(I9, II)

NeotenyI
37* .o6
.23 + .25*
*

Neoteny2
.24* .04 .II .25

Neoteny3
-.23 - .I7 -*05 -.02

N

= 49, 20

Americans (I2, Russians(ii, I4) Ache (II, I3) Hiwi (4, 4)
Mean S.D.

2o)

.38** yrs. 7.5 yrs.
I.o .34* i8)
.20+

45

- i.6 yrs. 5.5 yrs.
.36**

- I.3 yrs. 7.I yrs. 03
-.I3

*35

N = 5I, 35

U.S. Americans

Brazilians (2o,
U.S.

23)
(ii,

Ache

Americans Russians (I 2, I4)
(2o, 2I)

Hiwi (o, o) Mean
S.D.

-.2 yrs. 7.8 yrs. .36*
.I3+ .23 +
*

n.a.

.23 + .25 +

.28*

.26* -.I yrs. 4.5 yrs.
30* *

.o8

n.a.

.i8

n.a. - i.6 yrs.
7.7 yrs.
- .IO -.I5 .02 -.05

-.07

Pooled samples (Brazil + U.S.) N = Ioo, 55

Brazilians (39, 44) U.S. Americans (23, 38) Russians (23, 28) Ache (3I, 34)

.i6+ .I8*
.32**

Hiwi

n.a.
-.IO

32 **

n.a.
-.I3

n.a.
.00 .I4

Ache N = 4I, 36

Brazilians (I 7, I 6) U.S. Americans (i2,
Ache (i5,
Hiwi

Russians(1I2,
I5)

i5)

-.05
-.I9 .07 *38*

-.i6
-.29 .12

I12)

.0I
.2I -

Mean S.D.

(7, 4)

.2 yrs. 9.2 yrs.

-.2 yrs.

.24 +

9.6 yrs. n.a. n.a.

.4 yrs. 9.3 yrs.

.07

N

Pooled samples (All)
= I4I,
9I

Brazilians (56, 5o) U.S. Americans (35, 53)
Ache (46, 49) Hiwi (7, 5)

Russians(35,

40)

n.a. n.a. n.a.
.25** .3I** I24**

-.07 -.07

n.a.
.26**

.02
.03
.23*

NOTE:

+p < .I

in and male subjects. Numbers parentheses numbers raters female are of for

*p < .05 * p < .0I

and Rolf I969:5 20-23); table 2 shows pooled data where for pooled correlations The rows reporting appropriate. female subjects-for Brazilians, U.S. Americans,Russians, and Ache ratingBrazilian and U.S. photographs and forAche and Hiwi ratingall three sets of photographs-provide consistentsupportforthe proposition that neoteny is a component of female facial attractiveness.4Across five populations of raters,and across

two indices of neoteny,females are perceived as more attractiveto the extentthat their predictedages, as calculated fromtheirfacial proportions, less than their are actual ages.

tiveness connection Westerners for Achefemales probably rating is an largely artifact responses eye shape.Achehaveepicanthic of to and Ache witha highrelative width(log[EW/FH]) folds, eye also have relatively narrow eyes.Westemraters show an aversion all to narrow eyes-correlations betweenlog (EW/EH) and ratings of = facialattractiveness - .3 I, - .33, and - .44 for 4. The rangeof correlation coefficients such that,forWestern female is Brazilian, and raters(Brazilians, and The U.S. Americans, Russians), is not legiti- U.S. American, Russianraters. apparent it Westemaversion withwide eyesdisappears mateto pool coefficients all three for female photographic subject to Achefemales wheneyeshapeis confor. for samples.While Westemratersare attracted neotenousfacial trolled In addition, all populations raters, muchwider to of the proportions whenrating U.S. American Brazilian and women, they rangeof ages amongAche photographic subjectsmakes it more to seemto showa mildaversion neotenous to facial possiblecorrelates attractiveness of when difficult uncover proportions other than of ratingAche women. This slightreversal the neoteny/attrac-age.

JONES

Sexual Selection and Facial Neoteny | 733

Table 3 also includes data on ages predictedfrom Equations I to 3 above. Female models have extremely A second test of the neoteny hypothesisincorporate( neotenous faces: predictedages based on facial propormeasurements of two new samples of facial photo tions are 6.8 years and 7.4 years. This does not mean graphs,of U.S. female and male models, on the theor thatthe facialproportions models matchthose ofreal of features attractive of facesmigh 7-year-olds, thatthe distinguishing since the ages predictedby these equations stand out especially clearly in comparisons betweei are based on naive linear extrapolationsof age-related these samples and Universityof Michigan undergradu changes in adult facial proportions. It does argue ates. The two model samples consist of (i) photographl strongly, however,that female models' faces represent of ten femalemodels displayedon the coversof Cosmo a "supernormalstimulus," presentingin exaggerated politan and Glamour magazinesbetween i989 and I99 formthe featuresthat distinguishyoung women from (five fromeach magazine) and (2) photographsof tel old ones. in male models displayedin advertisements Esquire an! magazines on fil GQ. Both samples were selected from STRAIN AND FACIAL For the female mode STUDY 3: CARDIOIDAL at the Ann ArborPublic Library. samples,I beganwith the most recentissue ofeach mag ATTRACTIVENESS successivelyearlierissues un A thirdtest of the neotenyhypothesisemployeddrawazine and workedthrough til I foundfivesuitable facesfromeach magazine covei ings offaces made more and less neotenousby differing Models not facingdirectlytoward the camera, model degrees of cardioidal strain. From the Universityof and withtheirmouthsopen,celebrities, non-Caucasian Michigan photographicsample I selected nine photowere excluded. Since Esquire and GQ normallyfeaturi graphs: three of females given high ratingsof attraccelebrities rather than models on their covers, mall tiveness by University of Michigan undergraduates, models were selected fromconsecutive advertisement threeof females given averageratings,and threeof fea in these magazines, subject to the same restrictions males given low ratings (high-, medium-, and lowan( attractiveness right females.I used calipersto measurefaceheight, I subgroups). made a line drawingof each lefteye width,nose height, and lip heightand calculate( faceand producedtwo new drawings each ofthe origfor relativeeye width (EW/FH),relativenose height(NH, inal nine by subjecting each originaldrawing negative to FH), and relativelip height(LH/FH).The mean and stan and positive cardioidalstrain.For each trial,a raterwas darddeviationofthese measurements and the compara presentedwith one originaland two transformed drawfor of ble figures my sample ofUniversity Michiganun ingsofa single face(k = - o.i, O, O.i) andaskedto rank dergraduate females are presented in table 3. Fo the threeversionsin orderofattractiveness. Raterswere betweenmodels and undergradu drawnfrom females,all differences studentsin an intermediate-level anthropolates are in the expected,neotenous direction-model ogy class; each set of drawingswas rated at least four have largerelativeeye width,small relativenose height times. The entireprocedurewas repeatedusing photoand large relativelip height.T-tests(conductedon log graphsof nine males, divided as beforeinto high-,metransformed variables) show that all mean difference dium-,and low-attractiveness subgroups. between femalemodels and studentsare significant. For both males and females,faces subjected to positive cardioidal strain (k = O.i) were rated consistently TABLE 3 than the originalfaces (fig.2). The results less attractive and PredictedAges of Studentsanc are highlysignificant < .oi, binomial test) forboth Facial Proportions (p Models of Both Sexes sexes. Results fornegativecardioidalstrain(k = - O.i) are more complicated. For females, average attractiveness ratingswere higherforneotenous faces than Students Models fororiginalfaces. For males, attractiveness were ratings lowerforneotenousfacesthanfororiginal faces.Results S.D. S.D. Mean Mean were marginallysignificant and nonsignificant, respectively(p = .o6, p = .ii, binomial test).Since male and female trendsare in the opposite direction,the differFemales ence between the trendsforthe two sexes is strongly .OI* EW/FH .0I9 .23 .24 significant5 NH/FH .024 .49 .58 .034* .022 .2 LH/FH I.7 OI55** For both sexes thereseemed to be an interaction beSTUDY 2: STUDENTS AND MODELS

Males EW/FH
NH/FH

Predicted Age i (yrs.) Predicted Age 2 (yrs.)

20.2

20.2 .20

4.5
.OI3 .025 .022

8.o

7.4

6.8
.20

3.I*

3.8*
.OI4

*

LH/FH

.57
.I2 22.8

.47
.I2 23.I

.025**

Predicted Age 3 (yrs.)
*p < .05 **p < .0I

8.3

.oi8 5.8

that 5. The null hypothesis thiscase is thatdf(theprobability in a neotenousfemaleface,k = O.I, will be ratedmoreattractive (theprobability a that thantheoriginal face,k = o) is equal to dm moreattractive theoriginal than neotenous malefacewill be rated of face).But forall possiblevalues ofdf= di,,theprobability getresults bothforfemales and formales-that is, tingthe observed for the productof the probability males and the probability for females-is less than.oi.

734

CURRENT

ANTHROPOLOGY

Volume 36, Number S, December I995 tive temales are apparently neotenousenoughthatmating theirfaces more neotenous via negative cardioidal straindoes not make them more attractive (althoughit does not make them less attractive either), while unattractivefemales are rated more attractivewhen their faces are made more neotenous. Attractivemales are neotenous enough alreadythatmakingtheir apparently facesmoreneotenousvia negativecardioidalstrainactually lowerstheirattractiveness, while moderately attracmales are apparently tive and unattractive not so neotenous that making theirfaces more neotenous makes them less attractive(but it does not make them more attractiveeither).Thus there is a limit to how farincreasing neoteny results in increasing attractiveness, and that limit is reached sooner formales than forfemales.

2.5

2 i~ .5-I -0.1

,
0

[
0.1

Females

3 -2.5~~--

attractiveness subgroup hi X Di

-'

2

,/

med. lo Summaryand Implications in In humans, mate value declines with age beginning early adulthood. It declines more quickly for females thanformales. Consequently,we expecthuman beings, especially males, to have adaptationsforassessing agerelated changes in mate value. Regardlessof whether age-related changesin physicalattractiveness (especially forfemales)resultfromsuch adaptationsor fromother facesare attractive causes, it is possible thatparticularly those that presentage-related cues in an exaggerated or form.Largeeyes in relationto face height, supernormal small noses, and full lips are markersof youth.The rein sults presented this papersuggestthatneotenousfeatures are indeed criteriaof female attractiveness even when age is controlledfor.Specifically, i. Women whose facial proportions make them look thantheiractual age (as measuredbyregression younger equations predicting age as a functionof facial proportions) are perceived as more attractiveby male raters fromfivepopulations (but see n. 4). 2. A sample of U.S. female models has significantly than a sample ofU.S. moreneotenousfacialproportions and a strikingly female undergraduates low predicted to age, about 7 years,according regression equationspredictingage as a functionof facial proportions. 3. Cardioidal strain,a mathematical transformation shown by earlierresearchto providea good model for changesin facial proportions duringthe course ofmaturationand to affect perceivedages of faces,also has the an effecton female facial attractivenessaccordingto is U.S. raters.The effect nonlinear,suggesting thatneotenyis a componentof attractiveness only up to a certain point. in 4. Results formale attractiveness the above studies are weak and/orinconsistent. What are some possible future directions research for on neoteny and physical attractiveness and in the anthropologyof physical attractivenessmore generally? Researchto date providesstrong for support the hypothesis thatneotenyis a componentoffemalefacial attractiveness in Western societies. By contrast,the results

-~~~ 1 -0.1 Males 0 0.1

FIG. 2. The effects negative and positive cardioidal of on of transformations the perceived attractiveness faces of both sexes (includingfaces previouslyrated medium, and unattractive). attractive, of tween attractiveness subgroupand the effect negative cardioidalstrain.For the threeunattractive females,i I out of I 2 ratersfoundneotenous faces more attractive than original faces, while for medium and attractive foundneotenousfacesmore facesjust I 3 out of24 raters attractive.For the three attractive males, II out of I2 ratersfound neotenous faces less attractivethan nonneotenous faces, while for medium and unattractive faces just i 6 out of 3I ratersfoundneotenousfaces less In attractive. otherwords,negativecardioidalstrainreof sultedin a markedincreasein theattractiveness unattractivefemale faces and a marked decrease in the atof male faces but littlechangein tractiveness attractive of the attractiveness otherfaces. In summary, resultsfromdrawingsof faces subjected to cardioidal transformations support the hypothesis thatneotenyis a componentoffacial attractiveness, at least forfemales,but theyalso suggestthattherelationis ship between neoteny and attractiveness nonlinear. For all attractiveness subgroupsand both sexes, making facesless neotenousvia positivecardioidalstrainmakes them less attractive. But highlyand moderately attrac-

k cardioidal strain,

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Sexual Selection and Facial Neoteny 1735

ratio,anothercanforthe two non-Westernized societies presentedin this (I993a,b) notes that the waist-to-hip paper are only a beginning;much more researchwill didate fora universalcomponentof femalephysicalatvaries not only with age but, even among have to be carriedout in a wide rangeof such societies tractiveness, associwith a varietyof researchinstruments beforewe can women of the same age, with endocrineprofiles The theoretical state with any confidencewhetherthe neoteny/attrac- ated withfecundity. case thatneotenous tiveness connection is a human universal. Assuming cephalofacialfeaturesprovide such information not is that this connection continues to be supportedby fur- very strong,because such features,unlike secondary such as enlarged therresearch,many additionaltopics will remainto be sexual characteristics breasts, buttocks, addressed.These include the evolutionary unbases ofmale hips, and thighs,do not appear suddenlyat puberty attraction femalecephalofacialneotenyand thepossi- der the controlof female sex hormones.However, the to ble consequences of such attraction human morpho- topic clearly warrants investigation by reproductive for ecologistsand medical anthropologists. logical evolution. i. Evolutionary causes of attraction to neoteny2. Evolutionaryconsequences of attraction to neofurther puzzles. While the connectionbetweenneoteny teny.Darwin unitedhis discussionsofhuman evolution and female facial attractivenessproposed here may and sexual selection in a single volume because he partlyexplain the anomaly of female attractiveness, thoughtthat the latterhad played a major role in the it The resultsof this paper suggestthat it is time raises otherpuzzles. An attraction markersof youth former. to as may be adaptiveformen insofar it leads themto find for renewed attention to the possible connection beyounger women more sexually attractivethan older tween sexual selection and morphologicalevolutionin women, but it will not be adaptive if it leads men to our species. The evolution of modern Homo sapiens findjuveniles more attractive thanyoungwomen. Why, over the past ioo,ooo yearshas been markedby a trend findmarkersofyouthattrac- toward increasing craniofacial neoteny, including rethen,do males apparently tive even among females at the age of maximumfertil- duced prognathism, increased brachycephaly, and genity?Part of the answer may be that mate choice in the eral gracilizationin a numberof populations (Weidenreal world involves attentionto more than just facial reich I945, Newman i962, Brace and Mahler I97I, cues. If real-world mate choice involves both attention FrayerI98I). Biological anthropologists have generally to markers of cephalofacial neoteny to discriminate invoked natural selection for ecological adaptation or to youngadults fromold adults and attention secondary nonadaptiveforcessuch as pleiotropyor biased mutasexual characteristics fromthe rest of the body to dis- tion to explain these trends.The analysis in this paper criminateadults fromjuveniles, then attractionto su- suggeststhatsexual selectionmay also be involved.Sexin neotenous facial featuresis less puzzling. ual selection forneotenous features femalesis likely pernormally This line of argumentsuggeststhat the neotenous fe- to have become a particularly forcein human powerful male faces generatedin Study 3 will be perceived as evolutiononce increasesin lifeexpectancy had resulted more attractive, relative to nonneotenous faces, if in a larger fraction the adult femalepopulationliving of attachedto drawingsofunequivocallyadult bodies. An- past the age of menopause, thus increasing the ageotherpartofthe answercould be thatfemalemate value related variance in adult fecundity. Parallel trendstoof may be highestsomewhat beforethe age of maximum ward neoteny in males could be a by-product such fertility. Symons(I979:I89-90) arguesthatmales are selection in femalesor a directproductof sexual selecmost physically to attracted femalesofmaximumrepro- tion on males. Whether sexual selectionfor neoongoing ductive value, ratherthan maximum fecundity. Repro- tenous features(or other physical traits)can be meaductivevalue is a measureofexpectedlifetime reproduc- sured in living populations is a topic for future tion, and an individual choosing a partner with a investigation. The conventionalwisdom in the social sciences has long-term relationshipin mind should be adapted to take into account a mate's future reproductive potential been that evolutionarytheory,includingthe theoryof A as well as her current in fecundity.6 testableimplication sexual selection,is more or less irrelevant explaining is ofthis line ofargument thatthe attractiveness neo- human social behavior.The successes ofsuch new fields of tenous features(in comparisonwith markersof sexual as evolutionary and human behavioralecolpsychology will vary dependingon whetherindividuals ogy,as well as improvedunderstanding the physical of maturity) are consideringlong-term short-term of or relationships. underpinnings human behavior,are likely to forcea in Finally,neotenous facial proportions females might reassessmentofthis conventionalwisdom.The studyof is provide informationabout levels of ovarian function physical attractiveness a particularly promisingtest above and beyond the information the they provideabout case forinvestigating relationshipbetween biology age (Johnstonand Franklin I993). By analogy, Singh and culture.While this paper has emphasizedthe "biowiththe modern logical" side ofphysicalattractiveness, of theory sexual selectionas a starting point,thistheory 6. Strictly speaking, reproductive value is nota perfect measure of will undoubtedlyhave to be expanded and revised to long-term matevalue because a personcommitted a long-term allow forthe unique importance social learning our to of in relationship die orbecomeinfertile may before endofa mate's the the fateofthe hypothereproductive career. Whenthis possibility takeninto account, species (Laland I994). Whatever is matevalue becomesa function bothofmate'sage andofego'sage. sis that neoteny is a universal of female facial attracThis topicis treated moredetailin Jones in (n.d.). tiveness, this paper will achieve one of its aims if it

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encouragesboth biological and culturalanthropologists smaller chins and jaws than typical female faces. Feto explore a topic that has lain fortoo long in the no- males' ideal male faces were similar to typical male faces. man's-landbetween the two subdisciplines.

Appendix: Research on Neoteny and Female Facial Attractiveness and Apatow (I983-84) McArthur Population. U.S. Americans. Research methods. Researchersused a police Identikitto produce"average" male and femalefacesand faces and combinationsof "baby-faced" mature with different features:large and small eyes, low and high vertical small and largenoses and ears,all placementoffeatures, and all threemature featurestogether, threebaby-faced featurestogether. Results. For both sexes, faces with large eyes were rated more attractivethan average faces, which were rated more attractivethan faces with small eyes. The for was stronger femalesthanformales. For other effect (i) traits, forfemalesonly,averagefaceswereratedmore features,which attractivethan faces with baby-faced were rated more attractivethan faces with maturefeatures; (2) formales only,averagefaces were ratedmore which were than faces with maturefeatures, attractive fearated more attractivethan faces with baby-faced tures. Fauss (I986) Population. Germans. Research methods. Researcherasked subjectsof both sexes to constructthe face of an ideal memberof the opposite sex using a police Identi-kit. Results. Ideal males and females had wide mouths with full lips and a gracile lower jaw and chin. Ideal females also had gracile noses and high-archedeyebrows.

and Franklin(I993) Johnston Population. U.S. Americans. Research methods. Researchersused a "geneticalgorithm" to allow subjects to generateattractivefemale faces by a process analogous to artificialselection. A computerprogramgenerateda small population of female faces froma set of randombinarystrings("genotypes") which specified shapes and positions of facial features. Subjects assigned attractivenessratings to faces. A new generationof faces was producedby "seto lecting" genotypesin proportion theirattractiveness and adding small random "mutations" to the binary process continueduntil each The trial-and-error strings. face. subject had "evolved" the most attractive smallereyefaceshad significantly Results.Attractive fuller chin lengths, smaller lower-facialproportions, mouths than averageundergraduates. lips, and narrower Populations. English,Japanese. threefaResearch methods. Researchersconstructed cial images using graphics software: (i) a composite of (blend)of digitizedphotographs 6o Caucasian English females, (2) a composite of the most attractivei 5 females (as rated by English raters),and (3) an attracthe tiveness "caricature" that exaggerated featuresdisthe second composite fromthe first.They tinguishing threeimagesbased used the same techniquesto generate of femalesas ratedby Japanese on photographs Japanese males. Results.Images wererated3, 2, i (mostto least attractive) by Japaneseand Caucasian ratersratingboth own group and across groups. Attractivefaces had higher cheekbones, a thinnerjaw, largereyes relative to the size of the face, and shorterverticaldistances between jaw and mouth and between mouth and nose.

May,and Yoshikawa Perrett, (i994)

Cunningham(I986) U.S. American Populations. U.S. Americans (raters), (photographs). and international Research methods. Researcher collected measurements of relative size of facial featuresin a sample of of photographs 50 females.The sample included23 phoof tographs U.S. Americancollege studentsand 27 photographs of Miss Universe contestants. Photographs were rated by a sample of U.S. American undergrad- C. LORING BRACE of Museum ofAnthropology, University Michigan, uates. were associated Ann Arbor,Mich. 48109, U.S.A. 25 III 95 ratings Results. Higherattractiveness with larger and more widely separated eyes, wider It is all verywell to show that males may rate females cheeks, and smallernoses. cheekbones,narrower on various scales of "attractiveness"and that neotenousness may be one of them, but in orderforthis to Riedl (I990) on have any cumulative effect the appearanceof future Population. Austrians. that this is Research methods. Subjects were instructedto pro- generationsit also has to be demonstrated reproduction(Ryan facial image using a computer somehow related to differential duce the most attractive program which allowed them to manipulate sizes, I995). StephenJayGould has arguedthataspects ofhuman neoteny emerged because people find iA "cute" shapes, and positions of facial features. Results. Males' ideal femalefaces had largereves and (I977:35 O), and Joneshas expended considerableeffort

Comments

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Joneshas provideda concise, lucid, and ultimatelyaesthetically pleasingresponseto one ofhuman evolution's most intriguing questions: Why is the human male so interestedin a potential partner'srelativephysical atIt tractiveness? should come as no surpriseto anyone who has been partiallyconscious since the onset of pubertythat men around the globe have long been inof to reproduce" (Dobzhansky I972:77). And,in thepast, tenselypreoccupiedwith the relative attractiveness women. Althoughthe existenceof the this was true whethermales thoughttheywere "cute" youthful-looking the pattemofgazingis seldom contested, reasex-linked or not. if Evidently, all femaleshave the same opportunity sons forits persistenceaccount forsome of the liveliest to reproduce, male choice is not going to have any effect and, at times, most emotionallychargeddebates in the on the characteristics futurefemale form.Jonesis historyof social science. of Cultural anthropologistsare more comfortableexquite rightthathuman craniofacial form has undergone increasing gracilization over the past ioo,ooo years plaining the originsof men's keen interestin the nufroman unresolved (Brace, and Hunt i99I), buthe has not made a ances offemalebeautyas stemming Smith, tradicase thathuman choice had anything do with it. Fur- Oedipus complex,the persistenceof a patriarchal to worldorderprothereare some veryimportant thermore, aspects ofthat tion,or the demands of a postcapitalist gracilizationwhich cannot be detectedby readyvisual moting a consumptionethic that, in turn,encourages discrimination eithermales or females.The decrease the sexual objectificationof the female, but not the by in bone density and the thinningof the bones of the male, body.By failingto studythe phenomenonfroma culturalexplanationsareprone perspective, cranialvault cannot be visually discerned, and it would comparative withthe social facbe a most extraordinary person who would go to the to confusethephenomenon'sorigins embelextent of assessing the relative degreeof incisor shov- torsthatshape its culturalexpressionor stylistic eling and lingual-tubercle developmentor third-molar lishment.From the perspectiveof Foucault and his adand for is, agenesis and the relative reductionof the hypoconulid mirers, example,everysocial interaction first men's preference about power, and therefore when consideringthe attractiveness a potentialsex- foremost, of ual partner. That leaves "the neotenyhypothesis"right for youthfulnessis less, about aesthetics than about where it was some years ago, when it was judged to domination and control. It is not clear, however, if arises froma for be "largely,if not totally,a bankruptconcept" whose men's preference female youthfulness persistence was due mainly to "anthropocentrism" will to dominate as much as froma general abilityto relativefecundity, withan a (Shea I989:97)-in this case one could call it male chau- objectify potentialpartner's of unintended consequence beingthe transformation the vinist anthropocentrism. The emergence of "modern" human formover the femaleinto an object of aestheticcontemplation. is It is importantto rememberthat youthfulness an past ioo,ooo years and more is a consequence ofreducresourceoffemalepower,which oftenresults tions from Middle Pleistocene levels of robustness important which I have treatedin considerabledetail elsewhere in the heighteningof men's apprehensionand sexual and (Brace, Smith, Hunt i99i; BraceI995, n.d.).I sug- anxiety.While doing researchin the People's Republic struckbytheparadoxofmen's gested that those reductionswere the effects muta- ofChina I was repeatedly of for becominginfearing tions that were not weeded out when selection forthe yearning while simultaneously maintenance of the formerly necessary levels of ro- volved with a beautifulwoman. Althoughit was underbustness was relaxed. The mutationsthat produce the stood that beautifulwomen (whichin the Chinese contrendobservedare not "biased" but just the most likely textmeant women in theirearlytwenties)were harder minimal kind of change that can occur. This is the to control,manage, or mold, when I broachedthe topic at acknowledged, least mechanism that I labeled the probablemutationeffect of the ideal mate men invariably beauty to is (Brace I995). What it produces,in effect, evolutionby in the realm of fantasy,that they preferred When selectionis reducedor suspended,every- submissiveness.Chinese men are not alone in beingapentropy. been maintainedsimplytends prehensive of female beauty. Cautionary tales from thingthat had formerly to run down. In actual mechanisticterms,what we see around the globe repeatedlywarn men of the hidden is not really "neoteny," or the selective retentionof dangersand potentiallydire consequences of becoming The but youthful form, the increasingfailureof the develop- involved with a prettywoman who is a stranger. the archetype and, to a lesserextent, fatale mental process to producethe formerly necessaryadult femme-fatale This is the most likelyresultof the most status is a panhuman theme, suggestingthat cultures configuration. wam men and women to avoid becoming in likely mutationsoccurring the relaxationor absence everywhere ofselection-which is the minimumworking definition overlyfixedon what is most desiredin the oppositesex. For men it is physical beauty,whereas forwomen it is of the probablemutationeffect.

in demonstrating males evaluate femaleappearance that to that effect. However, not only does this ignorethe role of female choice in reproductive behaviorthatwas such an important partofDarwin's argument thefirst in place (Mayr I972: 90-9 1)-a perspective which has seen a recentresurgence interest(e.g., StrierI992, Cronin of I993)-but it does not take into account the fact that, in the available human examples, it is the males who ''are characterized an appreciably by highervariance in their reproductivebehavior" while "women are uniformly exposed to the risk of pregnancy and rarelyfail

University Nevada, of DepartmentofAnthropology, Las Vegas,Nev. 89154, U.S.A. 2 V 95

WILLIAM

JANKOWIAK

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accessing a man's relative social standing.The cross- are likely to have . . . species-typical. . . components, tales revealsthat and variationin these componentsmay be predictable culturalpervasivenessoffemme-fatale beauty,which is always associated with youthfulness, given knowledge of human biology and local circumthis prediction is is seen as conferring greaterpower on its possessor. In stances." I wonderjust how powerful is this context,youthfulness seldom taken as evidence likely to be. We can only speculate about such local circumstances, don't know whetherthe matingprefwe of submissivnessand thus powerlessness. I findJones'sexperimentaldesign an ingenious way erences have been consistentover time, and we can be will to bracket,forpurposesof analysis,the possible effects less certainthanwith otherspecies thatpreferences success forthe trait of social learningwhile focusingon the impact of sex- translateinto greaterreproductive At predicin linked differences the perceptionof relativephysical carrier. best,we are able to make non-specific attractiveness. would be interesting know what ef- tions such as "one mightexpect human beings to have It to fect sex orientationand thus potential erotic interest a . . . species-typicalemotional response to signs of the evolutionwould have on his researchfindings. sex orientation aging." While I do not wish to denigrate If and inwere controlledfor,lesbians and straight women might ary approach,which I considerboth important I be expected to differ their appreciationof relative sightful, believe it is essential thatwe be conscious of in in youthfulness females.I further suspect that lesbians the dangersinherentin this strategy. more than straight women will findrelativematurity We would do well to ask ourselvesa numberofqueshave we simplyfocusedon the traitswhich aesthetically pleasing. Because straightwomen often tions: First, come to value what men desirein the oppositesex, they support our hypothesis,ignoringthose that do not? will also find youthfulwomen more attractive.Con- Clearly,thereare many age-indicative traits.Would we men gays have got the same answers if we had focused on, say, versely,I predictthat comparedwith straight will findrelatively men aesthetically youthful more at- head shape, skin quality,and hair color? Secondly,have tractive.If my speculations-and that is all theyarewe consideredalternative explanationsforthe findings? turnout to be accurate,theneroticpreference with Cosmides and mayoper- Here I findmyselfin disagreement ate as the trigger heightened dampenedinterest for or in Tooby (I987), who writethat"learningis not an altemathe culturalobjectification a potentialsex partner. of tive hypothesis" to an evolutionaryexplanation. In a his By grounding analysis in an evolutionary frame- trivialsense this statementmust be true,since learning work, Jonesprovides a convincingexplanationforthe must itselfhave evolved and its operationmust be conhuman male's proclivity focus on youthfulness an strainedand informed processes operatingat other to as by importantaspect of female attractiveness. His request levels and on othertime scales. However,this does not that cultural anthropologists could be learned study the psychological negatethe factthata matingpreference processes of sexual attractionharks back to the disci- or unlearned,and if learned it could be independent of pline's historical mission to study the particularand or influenced by the social environment.Nor, since universalaspects ofhuman experience.It remainsto be there is empirical and theoreticalevidence that social seen how many anthropologists will followhis lead and leamingmay allow maladaptivetraitsto spread(Cavallienter the troubledwaters of documentingthe parame- Sforza and Feldman i98I, Boyd and Richerson I985, tersof our sex-linkedhuman nature. Durham i99I), should we assume thatthereis an adaptive explanationforall human behaviour.I believe that studiessuch as Jones'sare invaluable,precross-cultural KEVIN N. LALAND to ciselybecause it is plausible thatattraction facialneofad. Sub-DepartmentofAnimal Behaviour,Madingley, tenyor any othertraitis a society-specific Thirdly, are we just tellingstories?Human evolutionary Cambridge CB3 8AA, England. 23 Iv 95 history of began not in the Pleistocene but with the beginning Jonesillustratesthe value of an evolutionary perspec- life. This means that there is no shortageof hominid, tive. I agree that the relationship models whichwe could betweenhuman mat- ape, primate, social carnivore or and at- use as the basis foran adaptivestory. ing preferences sexual selectiondemandsfurther Evolutionary argutention,and cross-cultural comparisonsare particularly ments are so easy to constructthat empirical support I of insightful. appreciatethe rigor Jones's analysis,espe- should demand more than a cursoryreview of circumhandedout to cially since human sociobiologyand evolutionary psy- stantialevidence or a quick questionnaire chologyhave oftenstrayedtowardmethodologicallax- undergraduates. Jones'sstudy is to be welcomed both ity. I am a little suspicious of his use of one-tailed for the modesty of its claims and for the rigorof its statisticsforfemales (cf.two-tailedformales), particu- analysis. are a larlysince the findings used to justify hypothesis Finally,we should be aware thathuman sexual selecbut his conclusions are tion may operateby means, and at rates,atypicalof aniconcerninga sex difference, robust.His studytouches on important mal populations. Classically, researchersinterestedin probablyfairly the points regarding natureand complexitiesof human human sexual selectionhave treatedculturalinfluences sexual selection, which I address below. First,I raise on mating preferences a confounding as factorwhich some concernsabout the evolutionary of psychological ap- obscuresunderstanding how sexual selectionhas opa mine erated.In contrast, recenttheoretical proach. analysislof writesthat"standardsofphysicalattractiveness has demonstrated that matingpreferences not have do Jones

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Sexual Selection and Facial Neoteny | 739

to be innate to generatesexual selection (Laland I994) cuses on the facial gestaltas the basis forperceptions of Ratherthan obscuringsexual selection,learnedand so attractiveness. This view is consistentwith the literacially transmittedpreferences(for instance, for body ture demonstrating that faces are perceivedas wholes shape, hair color, or footsize) may themselvesgenerat( (e.g., Hosie, Ellis, and Haig I988, Morton and Johnson sexual selection, increasing the frequencyof the pre- I99I, Purcelland Stewart I988, RhodesI986, Sergent trait.Since, in comparisonwith genetictransmis I984, Young, Hellawell, and Hay I987). Furthermore, ferred sion, social transmissiontypicallyresults in a morc this approach is more parsimoniousthan feature-based of rapiddiffusion a preference through population,cul- approaches; both Jonesand Cunninghamfinddifferent a sexual selectionmaybe unusuallyfast, patternsof resultsformale and femalefaces,while we turally generated and the alleles underlying favored traitsmay be selected posit,and find, same pattern resultsforbothmale the of in to high frequency just a handfulof generations. and femalefaces. This analysis suggeststhat (i) thereshould be local, Accordingto Jones'stheory, faces with extremefeasociety-specific correlations between favoredtraitsand turesrepresenting neotenywill be perceivedas more atmating preferences;(2) sexual selection may account tractivethan otherfaces because of selection pressures. for cross-cultural variation in traitsunderlying attrac- As he points out, directionalselectionfavorstraitsthat tiveness;and (3) recentselectionmayhave modified any are extremein theirdimensions(largeantlers,big tails, favored the predilections throughout Pleistocene.These etc.). However, another formof selection-stabilizing theoretical renforce importance empirical selection-is more prevalentthan directionalselection the of findings studies such as Jones'swhich explorematingpreference and favorstraitsthat are the average of values in the patternsacross societies. But in focusingon those as- population i982, Dobzhansky I970). Thus,one (Barash which are universal, could just as easily predictthat selection would favor pects ofhuman matingpreferences we should not neglect the factthat otheraspects show faces with average configurations ratherthan extreme considerablecross-cultural variability(Ford and Beacb features. I95 I, Rosenblatt I 974). Males in all societies may yearn 2. The relationship neotenyto attractiveness. of Jones foran attractive mate,but in some societies "attractive" claims, as does Cunningham,that neotenyis an essenmeans small feet, protruding buttocks, or pendulous tial component of facial attractiveness females bein breasts.How can we account forsuch local preferences? cause it signals fecundity.However, we have shown And could theyexplain cross-cultural variationin ana- that,althoughneotenymay be a componentof attractomical or personality traits? tiveness,it is not essential to it. Empirically, neoteny if is fundamental attractiveness, to judgmentsof neoteny and attractiveness must be significantly highlycorand related. However, we (Langlois, Roggman, and MusLISA E. MUSSELMAN, H. LANGLOIS, JUDITH AND Departmentof Psychology,University Texas, of Austin, Tex. 78712, U.S.A. 9 v 95 treatiseon the importance Jones's paperis an interesting of physical attractiveness sexual selection,but sevfor eral points raised in it are in need of further consideration. i. The nature of attractiveness. Jonesclaims that atin is tractiveness "undertheorized psychology." fact, In thereare two current theoretical on perspectives the natureof facial attractiveness thathe does not consider. One perspective,consistent with Jones's approach, centerson the importanceof facial features defining for attractiveness.Cunningham and his colleagues (Cunningham I986, Cunningham,Barbee, and Pike I990) facesare those thatpossess a consuggestthatattractive stellation of mature, neotenous, and expressivefacial Their approachinvolvesmeasuring sizes of features. the individualfacialfeatures and correlating these particular with overallfacial attractiveness measurements ratings. a We have offered different theoreticalapproach in which we defineattractivefaces as those whose facial are closest to the average population configurations configuration (Langlois and Roggman 1990, Langlois,
LORI A. ROGGMAN

fromthat of Cunningham and Jonesin that it is not to Cunningham, "neotenous features" include eye concernedwith particularfacial features;instead,it fo- height,eye width,nose length,nose tip width,nostril

Roggman, and Musselman I994). This approach differs I986,

that judgmentsof attractiveness and age are unrelated in samples of college-agefemalefaces,indicating that a neotenous appearanceis not requiredforattractiveness. while ofcourseJones correct pointing Furthermore, is in out that old faces are perceivedas less attractivethan young faces, it is also certainlypossible to think of faces that are far fromattractive.Even young-looking infants, who are certainlyall neotenous,show the full rangeof facial attractiveness. 3. Measuringfaces.Jones measuresrelativeeye width, relativenose height,and relativelip heightfrom photographsand uses these measurementsto produce equations that predictage on the basis of the size of these various features.These measurementsare problematic forseveral reasons. First,thousands of facial measurements are possible (Farkas I98I), and Jones'sselection ofparticular to features measureseems to be guidedonly loosely by a prioritheoreticalconsiderations relatedto neoteny.His theoreticaldiscussion of changes in facial structure a functionof age does not mention cheek as width,yet he measures it and then later omits it when he findsthatit is not relatedto age. Furthermore, choice of cheek width as a neotenous featureis not consistent with the features chosen by Cunningham(Cunningham

i selmanI994) andothers (Berry99I) havedemonstrated

and Cunningham, Barbee, Pike I990).

According

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in width,foreheadheight,eye placement,and eye separa- may or may not serve as foodforthought this area of tion. Cheek width is a "mature" featureaccordingto inquiry. i. Does attractiveness necessarilyequate with sexual Cunningham's criteria.If Jonesconsiders "neotenous" For and "mature" featuresto be opposite ends of a contin- attractiveness? sexual selection (or any kind ofDaruum of age appearance,then he should also have mea- winian selection) to operate,the ultimate outcome has fitness.In the case of femaleneoteny, chin width,eyebrowthick- to be differential suredchin area, chin length, ness, and cheekbone prominence,the other "mature" Jones argues that this process is actualized through to females featuresmeasured by Cunningham et al. We wonder males' beingmore attracted youthful-looking what resultswould be obtainedif these otherpotential because, on average,these females make forreproducmeasures were used and whethera neotenous-feature tively more valuable mates. Hence, attractivenessis approach can providemore than post-hocexplanations clearlyequated here with sexual attractiveness. are We and predictions.For neotenyto be useful as a theoryof told, however,that neotenyalso makes us perceiveinresearchers attractiveness, who employ it should (i) be fants, animals, and even cars as "cute"-a quality but consistentin theiruse of the same theoretically driven closely analogous to attractiveness withouta sexual measuresofneotenousfeatures and (2) not conveniently component. The article does not specifyhow attracomit featuresthat theyfindto be unrelatedto age. tiveness was definedfor the raters.If Jones'ssubjects Second, measurements obtained from photographs had been given a series of picturesof children(or dogs) may not be accurate or reliable (Farkaset al. I980). Evi- to rate accordingsimply to their "attractiveness," my from dence indicatesthatwhen measurements takenfrom guess is thatwe would findan age gradient are running faces as opposed to photographs,attractivefaces are youngerto older, with babies (or puppies) being rated more likely than less attractive faces to have facial fea- more attractive. The implication is that the female-attractivenesstureswithin + i standarddeviationofthe mean (Farkas, heremayreflect, least in at Munro, and Kolar I987). Farkas et al. suggestthat "the neotenyassociation reported face with most measurementsin the range of + i SD part,somethingother than an attractiveness-fecundity may be close to the 'ideal face"' (p. I28). Additionally, relationship.I am willing to take Jones'sword that the Jones'sestimationthathis stimuliwould have thefacial female models' predictedage of about 7 years does not of and thatthe mean that theirfaces are identicalto those of 7-year-old proportions first- second-graders suggests lower equations he has developed on the basis of measure- girls,but still, the models' "age" is considerably than that of an average sample of 20-year-old ments do not accuratelyestimateneoteny. women men should have been se4. Coda. Jones claims that across five populations whose very high fecundity more neotenous faces are perceivedas more attractive. lected to find extremelyattractive.Again, mightneomeasured(notincluding tenyelicit somethingotherthan just sexual attraction? However,ofthe 42 correlations the data frompooled samples), only i i (approximately The answer may lie in Jones'sn.3, where he mentions at How thatneotenous featuresprobablyact as a release forpa26%) were significant the .os level or greater. are we to account forthe 74% of the correlations that rentalbehavior.Among otherthings,such behaviorenfor indicated no significantrelationshipbetween attrac- tails providing and givingprotectionto individuals dependentupon oneself.It could thus tiveness and neoteny?lAlthoughJoneshas providedus who are relatively with interesting cross-cultural data, thus farthese data be argued that in the past neotenous adult females benefited disproportionatelyfrom male provisionraise more questions than theyanswer. ing-in which case neotenywould not be the resultof sexual selection. 2. Is female physical attractiveness DANIEL PERUSSE really a human Departmentof Anthropology, University Montreal, anomaly? Jones develops his argumentfor a human of Montreal, Quebec, Canada H3 C 317 anomaly by pointingout that (a) in most sexually seumontreal. lected species males show greater of development sexual (perussed@ere. ca). 5 v 95 than femalesand (b) this pattern reversed is advertising Jonesis to be commendedfora particularly clear,well- in the case of humans, where men are more concerned This seeming designed,and interesting study.As he pointsout, sexual thanwomen with physicalattractiveness. selectiontheory been neglectedin relationto human exception is explained by (c) the human female agehas This paperpresents some ofthefirst morphology. empir- related variance in fecundity.Hence, female physical linked to highfecundity youthwould have and ical testsofexplicitevolutionary hypotheses concerning features and femalefacial attractiveness providesreasonableevi- evolved to be attractiveto males-that is, would have dence in supportof the author'sclaim thatneotenyacts become criteriaformate choice by males. If we really in as a kind of "superstimulus"signaling femalefecundity. want to findsome originality our species, I thinkwe Finding Jones's study generallysound and his results ought to look at a ratherthan b: as a sexually selected plausible, I will limit myselfto raisingtwo issues that species, human males are exceptional in the degreeto which male-male competitionis played out not in sexof to our I. If we restrict comparison femalemeasures, the 28 ual advertising but in control over resources (e.g., b the at 40%, aresignificant the PerusseI993, I994). Concerning and c, however, measured onlyi i, roughly correlations human situationfitsstrikingly well what seems a quasi.os level or better.

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male or female,are more interested universal pattern:females generallydo advertisetheir face. Even children, currentfecunditythrougha host of visual, olfactory, in children'sfaces than in adult ones. The typicalreacpostural,behavioral,and othercues. In nonhumanpri- tion to the Kindchenschema is a wish to protectand mates, forexample, female physical signalingof fecun- pamperthe object,whetherit is Donald Duck, a kitten, dityhas been observedin at least 55 species out of 79 or a baby. Since "nurturing behaviour" is a component Hrdyand WhittenI987). Such sex- of courting animals and humans (e.g.,birdsfeedeach in (reviewedin Blaffer ual advertisingincludes vulval or labial swelling and otherduringcourtship), this could have entailed a prefper- erenceforchildlikecraniofacial clitoralreddening, reddening/pinkening/whitening, features. KonradLorenz chest blis- once said thatin his opinion the Neanderthalswere not face reddening, ineal swellingand reddening, differ- exterminatedbut simply found sapiens sapiens cuter and others.I fail to see the fundamental tering, them as mates. The preference neoteence between the human case and thatof otherspecies. and preferred for The anomaly is only apparent,stemmingfromthe fact nous-facedmates might therefore found in males be that human females go into estrus with much greater and females,althoughit mightnot be so markedin the than their animal counter- tradition-bound frequencyand asynchrony females. Another approach to the exfor parts-to the extent that men are likely to have been planation of the preference the Kindchenschemain selected to find physical featuresof repeatedlyfecund the male mate-selection processis the trainedeye ofthe in hunter.Our perception beautyand symmetry based however, Ultimately, attractive. womenpermanently of is all these species males are attractedby those female on the necessityforthe hunterto assess his prey.Why do we prefer bite into a crunchy, covarywith fecundity. to physicaltraitsthat strongly juicy apple thaninto A phenomenon that might come closer to a human a shriveledone that may have lost none of its flavor? Fromthe day we are bornwe are told thatnew things of attraction men to anomaly would be the differential women who possessed attractivephysical featuresin are betterthan old ones and that we should exchange degrees notwithstandingthe fact that they old fornew immediately.Fromm(I992) pointsout that different were of equal fecundity as (i.e., same-agebut differentially we regardour partners consumergoods and tryto get Blaffer the best we can afford the partner on differentially attractive). neotenousand therefore marketin relation Hrdy and Whitten (I987), however, point out that at to our own market values. Since Sigall and Landy's least fourprimatespecies show variance in femalesex- (I973) workwe have known thatmen are judgedby the In ual advertising. threeofthose species,the varianceis appearanceof theirspouses. This is why women in parIt age-related. may be of interestthat the firsttwo are ticular become status symbols; it goes without saying crab-eatingand Japanese macaques and the third is thata man can expresshis highstatusonlywitha young and good-lookingpartner.In nonhuman primates,as Homo sapiens. Goodall (i99I) points out, males prefer experiencedfemales to young ones. The preference childlike feafor tures seems to be a human characteristic. SCHWEDER Perhapsthis BARBARA has matingstrategy reproductive value onlyin monogaTulbingerkogel A-3001 Mauerbach, Austria. 56, mous species. It would be interesting know whether to 7 III 95 monogamous animals have developed a strategyfor has of The pattern the ideal partner variouscomponents, judgingthe ages of potentialpartners. and some innate,some imprinted, some activelylearned It fromthe environment. is not easy to keep them apart SYMONS and to focus on a single one. In everypopulationthere DONALD University California, of are tendencies toward maintainingtraditionsand ten- DepartmentofAnthropology, dencies toward trying new strategies. Females are Santa Barbara, Calif. 93I06, U.S.A. io iv 95 than males in the known to be more tradition-bound mate-selectionprocess. Some female birds are said to Jonesmakes a lucid, succinct, and persuasive case for model whereasthe males oftheir an adaptationistperspectiveon human physical attrachave an innatepartner after birth(Daly and Wilson I983). tiveness. Althoughmy comments are informed the by species are imprinted I interAnd in humans it seems that femalesare more content same perspective, will proposesome alternative (see SymonsI995 fora morethorough presenwith the averagemale face thanmales are withthe aver- pretations age female face. Thelen (I983) speaks of minority-type tation and forreferences).' contribution the to Jones'sresearch is a significant on human mate preference the partofmen wherephysical appearanceis concerned.Jones'sdata leave no doubt "adaptationistprogram,"the goal of which is to partithathuman males are more attracted neotenouspro- tion organisms into functional components-that is, by devices. Nevertheproblem-solving portionsin femalefaces thanby the averagefemaleface, into special-purpose to but I do not thinkit is sufficient explainthisin terms less, his main hypothesis-that the human male's preference for "neotenous" female facial featuresis a byof its indicationof fecundity. youth-is in a sense The response to a childlike appearance (Kindchen- productof selection forpreferring than"constraintist." does not imply It schema) is innate. Eibl-Eibesfeldt (I984) points out that less adaptationist between its signal characteris based on the difference and its characteristic featuresand those of the averageadult i. I thankNancyEtcoff YonieHarris their for metacomments.

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Volume 36, Number S, December 1995 exists in the population'sgene pool thatcauses its bearX ers to prefer on average. Now, the myriadvagaries of ontogeneticdevelopmentessentiallyguaranteethat individuals bearing this allele will, in fact, exhibit a rangeofpreferences distributed aroundX. Selectionwill favorthe allele if deviationsfromX in eitherdirection depressfitnessto the same extent.But thisis not always the case. An "asymmetricalfitnessdistribution" exists whenever a unit of deviationfromthe optimumin one direcmore thandoes a unit ofdeviation tion depressesfitness in the other.Extremeexamples have been referred as to effect":The most nutritiousgrass (the the "cliff-edge theoreticaloptimum) may grow rightat the edge of a but the cow that always triesto graze there steep cliff, is unlikelyto have the highestfitness the herd.When in fitnessdistribution an asymmetric exists,selection can be expectedto favora preference that diverges fromthe theoreticaloptimum (away fromthe cliff edge). For example, perhapsin ancestralhuman populationsfemales with relatively gracilejaws (and,hence, relatively short lower faces and large eyes) typicallyhad slightlyless mate value than females with average jaws, but selection nonetheless favoredmales who preferred gracile ratherthan averagejaws because the fitnesspenaltyfor choosing females with robustjaws was comparatively large. As Jones'stheoreticalpresentationmakes clear, the general hypothesisthat human males evolved speciesmechanisms designed to assess fetypical preference male mate value does not implythat males universally will develop the same absolute standardsof female attractiveness. The ideal stimulus values of some female can features be specified an absolutesense in phenotypic (e.g., unwrinkledand unblemishedskin), and these are The ideal likelyto be perceivedas attractive universally. stimulus values of otherphenotypic features, however, cannot be specifiedin an absolute sense because they varyamong human populations (e.g.,skin color),hence the attractivenessof these featuresis likely to be assessed relative to local phenotypesratherthan absolutely.For example,a psychologicalmechanismthatinstantiatedan obligatepreference a specificskin color for could not possibly have been universally adaptive amonghumans. What could have been universally adaptive, however, is a mechanism that calibratedpreferences facultatively, about using as input information local skin colors (which in ancestral populations representedadaptationsto local conditions). A species-typical male psychologicalmechanismthat instantiatesthe rule "Preferfemale skin that is a bit lighter than the adult female average" (in ancestral nubilpopulations relative lightnessprobablysignified and high estrogenlevels) would result ity,nulliparity, in very different absolute skin color ideals in Nigeria and Norway. Nigerian men would perceiveNorwegian women as much too light,and Norwegianmen would perceive Nigerian women as much too dark. This line ofreasoningalso implies,however,thatmen m4ysometimes perceive women of anotherpopulation as more

that in ancestral human populations neotenous facia features per se indexed relatively high female mat value. If the human male's preference neotenous facia for featuresis merely a by-product, presumablywoul it have entailed at least some costs in ancestralpopula tions. For example, assuming that Jones'shypothesisi correct,an ancestral male given the opportunity t choose between two potential mates of the same agc one of whom (A) had a more neotenous face than th other(B),would have been willingto pay a higherbride priceforA because ofhermore attractive face,althoug] would have represented B, at a lower bride-price, bettc value; or he might have failed to acquire B's superic weaving skills, which would have contributedsomc and insteadacquiredA's moregrac thingto his fitness, ile jaw, largereyes, smallernose, and fullerlips, whicl to would have cor according the by-product hypothesis, tributed nothing;or he mighthave chosen an older fe male with neotenous features over a younger female(c highermate value) with averagefeatures. While many factors can constrain selection fror it achievingdesignperfection, is nonethelessworthask ing,when considering any given constraintist hypothc sis, why selection didn't do better.If neotenous facio in features themselvesdid not index relatively high fe male mate value, why didn't selection favor male whose preferences were a function veridicalage cues of And why are only certain facial features,ratherthai when neotenous?In short,althougl all, most attractive mor Jones'shypothesismay well prove to be correct, relentlessly adaptationist possibilities are worth ex ploring. Perhapsthe facial proportions thatJones interprets ii termsof age cues also indexed some otheraspect(s) c female mate value. One possibilityis hormonalstatus which Jonesconsidersunlikely.Yet high androgenlev els in women are positivelycorrelated withreproductiv, and observable indices of higi system dysfunctions, androgenlevels-such as acne, hirsutism,and a higi ratio-seem to be systematically waist-to-hip perceivec as unattractive. my eye, the faces in Jones'sfigure To more in "masculinity"than in age. appear to differ Also, there is a clear adaptationistrationale for ex pectingfemale mate value in ancestralpopulations t have been a negativefunctionof parity, and certainfa cial proportions Maternalbone forma may indexparity. tionratesare elevatedduring whichmayper pregnancy, manently lengthen the mother's face, and a growtl hormone (hGH-V) is expressedin the placenta and se cretedin large amounts into the maternalcirculation which may permanently "coarsen" her facial features. Jonesproposes that neotenous female facial feature are supernormal But ther, stimuli,and he may be right. is a more general (and more adaptationist)reason tha a "preference mechanism"-in any species and in an, a domain-might be shaped by selection to identify ideal a stimulus other than the theoreticaloptimum Suppose that,in a particularcase, the theoreticalopti mum stimulusvalue is X. Further suppose thatan allele

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Sexual Selection and Facial Neoteny | 743

in attractive, certainrespects,thanwomen of theirowr population.A classic example is reported Wagatsumc by mer contact,Japanese (in the paperJonescites). On first perceivedwhite Westernwomen as less physicallyat includ tractivethan Japanesewomen in most features, facial hair,and eye color. But the mer ing skin texture, perceivedWesternwomen's typicalskin color as morn because it was a bit lighterthan the adul attractive, femaleaverageand, hence, close to theirideal Japanese If thereis significant interpopulation variationin fa cial proportions, perception neotenymay be anal the of ogous to the perceptionof skin color. That is, humar males may have been selected to prefer female facia: featuresthat are relatively neotenous, by local stan dards,ratherthan to prefer certainabsolute facial pro portions.If so, males will not necessarilyprefer femalc featuresthat are neotenous by the standardsof ever) human population. Surelyit is possible fora woman'" eyes to be too large,her lower face too short,her nose too small, and her lips too full (imagineBettyBoop as real woman). In fact,Jones'sdata implya ceiling effec forthe attractiveness facial neotenyeven withinpop of ulations. In sum, I propose that accurate predictionsabout a given male's perceptionsof female facial attractiveness can be derivedonly from(a) knowledgeof the designs of species-typical preference mechanismsin the human male brain and (b) knowledge of the female faces thai the male has been exposed to (because information con tained in these faces will have calibratedsome of his preference mechanisms).Intrapopulation agreement and interpopulation disagreementin attractiveness ratings are not evidence thatpeople "learn" standards attracof tivenessfromone another-any more than intrapopulation homogeneity and interpopulation in heterogeneity skin color is evidence thatpeople "learn" theirskin col ors fromone another.

moderate correlatomical landmarksand demonstrate tions between these indices and ratingsof attractiveis and ness. Althoughthe averagenesseffect important deserves theoreticaltreatment its own right(Koesin abundantevidence,at least forfemales,thatfaces close to the population average are not the most attractive possible faces (Alley and Cunningham i990 and references in appendix). is A different strategy necessaryto test the neoteny contra Mushypothesis.In this case it is appropriate, selman et al., to "omit features... not correlated with fromdifferent features age." It also helps if information can be combinedto providean overallindex ofneoteny. In studies i and 2 I focuson just threemeasuresoffacial and proportions (relativeeye width,relativenose height, relativelip height)because thereare stronggroundsin on the standardliterature facial agingforexpectingthe relative dimensions of the three major facial features (eyes,nose, and lips) to change with increasingage and consistentevidence in my samples that the measures do changewith age in the expectedfashion.Alternative measures of the relativedimensionsof the threemajor facialfeatures less satisfactory. heightand width are Eye with increasing of mouth do not decrease significantly age. Eye heightgives some signs ofbeingpositivelycorof across samrelatedwith ratings femaleattractiveness as ples but may be as much an expressivefeature a neowithage; including tenous one. Nose widthis correlated it in age regressionsfor females produces an age predictorequation similarto Equation 2.1 The corresponding index of neoteny predicts attractivenessabout as

lag

I990,

I990), thereis also Langloisand Roggman

well as Equation5 (Jones I994).

Reply
Ithaca,N.Y. 14853, U.S.A. I5 vi 95
DOUG JONES

Is therea link betweenneotenyand attractiveness? Virtually all commentatorsagree that the evidence prethe sentedin thispapersupports proposition thatfemale facial neoteny is linked to male perceptionsof attractiveness. The main dissent comes from Musselman, Langlois, and Roggman,who point to evidence for aD averageness or "prototype"effect:faces with proportions close to those of the averageface are perceivedas more attractive.In testing this hypothesis it makes fromas many facial sense to incorporateinformation 1 - ih p-lr. rofop-.t11.Q measurementsas possible. Thus Jonesand Hill (I9931 n r e.hQP nr%-n.;i:Ar a lt n -iPr. and Jones(n.d.) constructindices of the "averageness" of facial proportionswhich combine several hundred i. The equationis Age = - I68 * log[EW/FH] 87 * log[NH/FH] + ana- + 9 5 *log[NW/FH]. measurementsof distances between photographic
TATt-VlIIA

as Pooling of correlationcoefficients, in table 2, is a standardtechnique fortestinghypothesesacross multiple samples. What mattersis not the numberofsamples results but the consisforwhich there are significant tency with which males in five populations of raters neotenous featuresin the pooled samples. prefer and Why neoteny? Age,fecundity, sensorybias. I have proposed not only that there is a connectionbetween but that this facial neoteny and female attractiveness of connectionis a by-product the universalmale percepwomen are morephysically tion thatyounger attractive than older women and that estheticresponsesto signs of aging are largelygenetic adaptations to age-related Several commentators point out changes in fecundity. potential extensions or limitations of this line of argument. I argue that human beings are anomalous in that males are more concerned with female attractiveness than vice versa, but Perusse points out that males in many primatespecies are attentiveto cyclical and seaHis point is well sonal indicatorsof female fecundity. of taken. A more complete treatment the evolutionary basis of attractionto markersof youth and fecundity

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Volume 36, Number S, December 1995 they are also commonly attractedto secondarysexual charactersin the female figurethat appear at the time of puberty. Finally,Symonsproposesthatthe traitsI have labeled information about female neotenous may actually carry they carry fecundityover and above the information about age. In my paper I expressmild skepticismabout this possibility,partlybecause thereis no evidence in my samples that indices offacial neotenyare correlated indicatorsof fecundity with potential non-age-related ratio.But Sysuch as age at menarcheand waist-to-hip mons's (I995) recentworkon this subjecthas persuaded thatestrome thatwe need directtestsofthepossibility on ratiosand parityhave effects facial atgen/androgen of tractiveness over and above the effects aging.2 Attractivenessand morphological evolution. Brace to suggestsseveral reasons fordoubtingthat attraction on neotenousfeaturescould have had any effect human morphology sexual selection. He claims that (i) in via to the evolutionary past all women had the opportunity so reproduce, therewas no scope forsexual selectionon are females; (2) some changes in skull morphology not of readilyvisible and could not have been targets sexual that neotenyis "a selection; (3)Shea has demonstrated bankruptconcept" in the contextof human evolution; and (4) the "probablemutationeffect" providesa sufficient explanation for declining robustness since the Middle Pleistocene. Brace's claim that all femaleshave the same opportuby nityto reproduceis refuted an immense body of reecology (Bonsearch in demographyand reproductive gaarts I983, Ellison I990). Female fecundityshows considerablevariationboth within and between socieof ties; levels of ovarianfunctionand probabilities convariables, ceptiondependon a numberofenvironmental work efnutrition, includingfrequencyof intercourse, and pathogenlevels. Additionalvariancein female fort, and offspring fitnessresultsfromvariance in mortality In nutrisurvivorship. turn,frequenciesof intercourse, risks commonlydetional levels, and infantmortality and pend in parton women's success in finding keeping desirablemates. Researchin modernindustrial societies of on demonstratesan effect attractiveness a woman's of probability marriageand on the social status and attractiveness her husband (reviewedin JacksonI992). of Whetherquantitative research in traditionalsocieties will show similar correlationsbetween attractiveness and mating success is an open question; considerable qualitative evidence points in this direction. Studies of assortativematingcommonlyshow assortment even forthe traitsnot easily assessable by potential mates, such as lung volume and blood pressure (Spuhler I968). Presumablyassortmentresults not befemalephotographic 2. None of the Brazilianor U.S. American when or beingpregnant subjectshad had any children reported to Thus parity se could not have contributed per photographed. senin although in differences facialattractiveness thesesamples, of cculdhave. suggestive parity facialporportions bias against sory

caveats about applying some important Laland offers Understandtheoryto human psychology. evolutionary ing how the human bodyand brainhave been shaped by about selecnatural selection oftenrequiresinferences fromthose which most of tive pressuresverydifferent may select post hoc us face today.At worst,researchers models ofPleistocenehuman or from among alternative otherape, primate,or social carnivorebehaviorwhichHowadaptivestory. evermodel best fitssome preferred safegroundin makinginferever,we are on particularly ences about the selectivepressureson mate choice with because dechanges in fecundity, respectto age-related mographicevidence suggests that humans have faced much the same relationshipbetween age of mate and of relativeprobability conceptionnot just throughthe transition Pleistocene but rightup to the demographic of the past few centuries. Symons notes that according to my hypothesisatis tractionto neotenous facial features not adaptiveper of se but a by-product attractionto youngerwomen. It is a result,in otherwords,of "sensorybias." This is not an argumentagainst the hypothesisif we accept that build ioo% accuratepercepnaturalselectionwill rarely sensorybias will be ubiquitual systemsand therefore tous. Sensory systems commonly respond to stimuli other than those to which they are adapted and often respond especially stronglyto "supernormal" stimuli stimof thatpresentthe distinguishing features a target form(Ryan I990, Enquist and Arak ulus in exaggerated to tions is not an alternative the sensory-bias argument but a variationon it. It assumes evolutionary constraints such that natural selection cannot build sensory systems that always respond to stimulus X but never to nonadaptiveor less adaptive stimuli similarto X. Alternativeexplanations.SeveralcomWhyneoteny? mentatorsconsider alternativeexplanationsforthe attractionto neoteny.One popular explanationformen's attraction youthful to featuresis that such featuresare signs of powerlessness and submissiveness. I review and some reasons forbeing skeptical of this possibility, Jankowiakoffers groundsforsupcogent ethnographic betweenbeautyand power thatthe relationship porting My is the opposite of that proposedby the hypothesis. his own observationsin Brazil corroborate account of sexuality in China. Far fromsupposing that old men involved in relationshipswith youngwomen will typiby cally dominate theirpartners virtueof theirage and experience,Brazilians often suggest that men in such are and relationships especiallyvulnerableto cuckoldry economic exploitation. to Perusse and Schwederpropose that attraction neoof tenous features may be a by-product the parentalfeelings aroused by infantilefeatures,but this hypothesis does not seem to explainwhymen are moreconsistently than women. Nor to attracted neotenous facial features does it seem to explainwhy,althoughmen are attracted form feathe to femalefaces thatpresentin exaggerated tures that distinguishyoung women fromold women,
I993).

from funcfitness The argument asymmetrical

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Sexual Selection and Facial Neoteny 1745

cause people choose mates on the basis of lung volume per se but because theychoose on the basis ofcorrelated traits. Choice of partnerson the basis of perceptible imtraitsis likely to have consequences forcorrelated perceptibletraits. Shea (i988) criticizesthe hypothesisthat human beincrease in neotenyas a ings show an across-the-board result of a general slowdown in rates of morphological He change associated with delayedmaturation. also argues thatallegedlyneotenoustraitsin adult humans are not generally homologouswith similartraitsin juvenile specific nonhumanapes. This has no bearingon whether anatomical regions have a neotenous appearance as a resultof sexual selection (or othersocial selection). Trends toward craniofacial neoteny in the past ioo,ooo years may result fromecological selection for and increasedmechanical efficiency reducedmetabolic sexual or othersocial selection,or from direccost,from Of tional mutation-Brace's probable mutation effect. these three possibilities,the last is probablythe least plausible: a strongpredictionof the last hypothesisis increasingtrait variance over time, but Frayer(I977) findsthe opposite trend,towardreducedvariance. Whethersexual selection has contributed the evoto lution of human craniofacialneotenyis an open question. Brace providesno good reason to doubtit. Instead, his replydemonstrates failureto take sexual selection a this failureputs Brace in good seriously.Admittedly, the company.When Darwin attributed showyplumage of some male birdsto female choice, Wallace proposed of thatsuch plumage resultedinsteadfrom overflow the JulianHuxley arguedthat the elaborate surplusenergy. courtshipdances of some birds were not a productof of sexual selection but a by-product emotional excitement.Mayrsuggested thatthe extraordinary complexity and diversityof male genitalia in a number of groups resultednot fromfemale choice but frompleiotropyin thatgenes selected fortheireffects otherpartsof the body producedincreasedgenitalic complexityas a side effect. More recentresearchhas made it clear thatWallace, Huxley, Mayr, and many other major figuresin had a blind spot about sexual selectheory evolutionary lar blind spot seems to have kept many biological anthropologistsfrom seriously investigatingthe sexual selection of human morphology. Attractiveness, learning,and culture.Symonsand Laland considerthe topic of variationin standardsof attractiveness. Symonsnotes thatifpeople calibratetheir to standardsof attractiveness the local populationaverface is a neoteage-for example,if the most attractive nous version of the averageface-then standardsof attractivenesswill vary with population differences in facial proportions. Such variationwill involve learning Ache but not culture. Consistentwith this possibility, with each otherthan and Hiwi show stronger agreement with Brazilians, U.S. Americans,or Russians in stanThis agreementis more dards of facial attractiveness. the mechlikelyto reflect operationofa "face-averaging

anism" than shared culture,because the two populations have similar facial proportions but little in common culturally(Jonesand Hill I993). While habituationto local physical featuresmay account for a great deal of variation across populations, Laland rightly notes that many local preferences exfor aggerated characters cannotbe explainedin thisfashion. He has pioneeredmodels ofthe coevolutionofgenesand culturally transmitted standards attractiveness of which are likely to illuminate cross-culturaldivergencein standardsof beauty.I would add only that the theoretical bare bones of these models will need to be fleshed out with more empiricalwork on psychologicalmechanisms and culturalcontexts. Future research will further clarifythe relationship between neotenyand facial attractiveness. More generally, I suggest that the study of sexual esthetics may develop in the same fashion as the study of language. Both languageacquisition and the developmentof standards of sexual attractiveness probablyregulatedby are specialized "mental organs"shapedbynaturalselection. But both are presumably influenced well by nonadapas tive sensoryand cognitivebiases and by social factors, includingperceptionsof what is popular and what is da cwri atPAcl wiAth
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Cited References
T. R., AND M. CUNNINGHAM. I99I. Averaged faces are attractive, veryattractive but facesare notaverage. Psychological Sciencee2:I 23-25. M. I994. Sexual selection. Princeton: ANDERSSON, Princeton Press. University D. P. i982. and behavior. BARASH, Sociobiology New York: NorthHolland.[LEM, JHL, LAR] Elsevier M. M. I995. Courtship and continued BARTLEY, progress: Julian Huxley'sstudieson birdbehavior. Journal theHistory Biof of ALLEY,

tion(Eberhard I995). A simiI985, Cronin i99i, Bartley

in R. G. I985. Growth the agingcraniofacial skeleCenterforHuman Growth Development, ton.AnnArbor: and of University Michigan. BERNDT, R. M. I95 I. Sexual behavior western in Arnhem Land. in FundPublications Anthropology Viking I6. BERRY, D. S. I99I. Attractive facesare not all created equal: Joint effects facialbabyishness attractiveness social of and on and perception. Personality Social Psychology Bulletin
BEHRENTS, ARTHUR. I986. Perceiving characterin faces:The impactofage-related craniofacial changes on social perception. Psychological BulletinI00:3-I8. BERSCHEID, E., AND E. WALSTER. I974. Physical attractiveness.Advancesin Experimental Social Psychology 7:I57-2I5. AND M. BORGERHOFF MULDER. EdiBETZIG, L., P. TURKE, tors.I988. Human reproductive A behavior: DarwinianperNew York:Cambridge spective. University Press. BLAFFER HRDY, S., AND P. L. WHITTEN. of I987. "Patterning in sexual activity," Primate societies.EditedbyB. B. Smutset al. Chicago:University ChicagoPress.[DP] of and behavior: analyAn BONGAARTS, biology, j. I983. Fertility, sis of theproximate determinants. New York:AcademicPress. S. A. I986. Radiancefrom thewaters:Ideals offemiBOONE, nine beautyin Mende artand culture. New Haven: Yale UniPress. versity BERRY, I7:523-3I. D. [LEM, JHL, LAR] S., AND L. Z. MC

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Volume 36, Number 5, December 1995
ENLOW, D. H. I990.

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withbehavioral aryhypotheses data.Human Nature5:255-78. attractiveness: Resultsfrom fivepopulations. Ph.D. diss.,Uniof AnnArbor, versity Michigan, Mich. [DP] T. I988. Bodystyles. POLHEMUS, Bedfordshire: Lennard. . n.d. Physicalattractiveness thetheory sexual seand of versusnonindependent lection:Resultsfrom populations. AnnArbor: five Museumof P RUE T T - JONES, S. I99 2. Independent matechoice:Do females copyeach other? American Naturalof In Anthropology, University Michigan. press. ist I40:I000-I009. Criteria physical of JONES, D., AND K. HILL. I993. attractivePURCELL, D. G., AND A. L. STEWART. I988. The faceness in fivepopulations. Human Nature 4:27i-96. detection effect: Configuration enhancesdetection. D. I994. Evolutionary Perception KENRICK. social psychology: Fromsexual and Psychophysics 43:355-66. [LEM, JHL, LAR] selectionto social cognition. Advancesin Experimental Social Psychology 3I:75-I2I. RHODES, G. I986. Memory forlateralasymmetries wellin J. I990. Koinophilia knownfaces:Evidenceforconfigural KOESLAG, groups sexual creatures intospeinformation memory in of cies,promotes representations faces.Memory and Cognition I4:209-I9. stasis,and stabilizessocial behavior. Journal of Theoretical BiologyI44:I5-35. [LEM, JHL, LAR] K. N. I994. Sexual selection LALAND, witha culturally Merkmale transmit- RIEDL, B. I. M. I990. Morphologisch-metrische des tedmating preference. mannlichen weiblichen und Theoretical Population Biology Partnerleitbildes ihrer in Bedeutungfurdie Wahldes Ehegatten. Homo 4I:72-85. 45:I-I5. P. C. I974. Foundations interpersonal LANGLOIS. I990. Attractive faces ROSENBLATT, of atJ. H., AND L. A. ROGGMAN. are onlyaverage. traction. New York: Academic Press. [KNL] Psychological Science i:ii5-2i. [LEM, JHL, LAR] "Sexual selection, RYAN, M. J. I990. and sensory systems, senin R. J. CASEY, soryexploitation," Oxford in LANGLOIS, J. H., L. A. ROGGMAN, AND J. M. surveys evolutionary biology, pp. I57-95. Oxford: RITTER. I987. 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ANTHROPOLOGY

Volume 36, Number s, December 1995
E. i92i. The history humanmarriage. of Vol. London:Macmillan. G. C. I975. Sex and evolution. WILLIAMS, Princeton: Princeton Press. University WESTERMARCK,

NewperA. B. I976. Womenof value,men ofrenown: of Austin:University Texas exchange. in spectives Trobriand Press. and social beM. J. I99I. "Sexual selection WEST-EBERHARD, EditedbyL. Tigerand in havior," Man and beast revisited. D.C.: Smithsonian Instipp. M. Robinson, I59-72. Washington, tution.
WEINER,

I.

YOUNG,

in figural information faceperception. Perception I6:747-59.
[LEM, JHL, LAR]

A. W.,

D.

HELLAWELL,

AND

D.

C. HAY.

I987. Con-

Calendar
I996

Tamil Nadu, InConference, ation, 4th International and dia. Theme: Globalization, Self-Determination, in Justice Development. Write:PeterPenz, Faculty 4700 Studies,York University, of Environmental Keele St., NorthYork, Ont., Canada M3J IP3. April I 8-20. Delta Blues Symposium:The Blues II, inArk., State University, conference, terdisciplinary U.S.A. Deadline forabstractsJanuary I996. i5, Write:Delta SymposiumCommittee,Departmentof Englishand Philosophy,P.O. Box I890, Arkansas Ark. 72467-I890, State University, State University, or U.S.A. (e-mail: wclement@toltec.astate.edu rburns@quapaw.astate.edu). May 29-June 2. 3d EuropeanEthnopharmacological of Conference the Colloquium and iSt International and Anthropology Historyof Health and Disease, Genoa, Italy. Write: Comitato OrganizzazioneEtnoIstitutodi Antropologia Antropologia, farmacologia

DevelopmentEthics AssociJanuary i-8. International

Fisica, Universitadi Genova, Via Balbi, 4, i6i26 Genova, Italy (fax: [39] I0 2095987). me 13-I6. gth International Oral HistoryConference, Goteborg, Sweden. Theme: Communicating Experience.Deadline forabstractsDecember I. Write: Sven B. Ek or Birgitta SkarinFrykman, Department of Ethnology, of University Goteborg, VastraHamngatan 3, S-4II I 7 Goteborg,Sweden. eptember8-14. Lithic Technology:FromRaw Material Procurement Tool Production, to workshopin connectionwith the I 3th International Congressof the Union of Prehistoric and Protohistoric Sciences, Forli,Italy. Write:Sarah Milliken, c/o Segreteria XIII CongressoU.I.S.P.P., Via Marchesi, I, 47ioo Forli,Italy (fax: [39] 543 35805). )ctober19-20. Interface '95: 2oth Annual Humanities and TechnologyConference, Atlanta,Ga., U.S.A. Write:JulieNewell, Social and International Studies, SouthernCollege of Technology,100 S. Marietta Parkway, Marietta, Ga. 30060-2986, U.S.A.

Errata
In Manzi and Passarello's reporton the Neandertals size and shape values forthe leftthirdmolar ofBreuil 3 fromGrotta Breuil in the April issue (CA 36:355-66), shouldhavebeen I I 9.9 and I I 3.9 respectively. to two corrections table i (p. 36i) were overlooked;the