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Review of General Psychology © 2010 American Psychological Association

2010, Vol. 14, No. 3, 240 –250 1089-2680/10/$12.00 DOI: 10.1037/a0020451

The Science of Sex Appeal: An Evolutionary Perspective
Gordon G. Gallup, Jr. David A. Frederick
University at Albany, State University of New York University of California at Los Angeles

Growing evidence shows that features we find attractive in members of the opposite sex signal important
underlying dimensions of health and reproductive viability. It has been discovered that men with
attractive faces have higher quality sperm, women with attractive bodies are more fertile, men and
women with attractive voices lose their virginity sooner, men who spend more money than they earn have
more sex partners, and lap dancers make more tips when they are in the fertile phase of their menstrual
cycle. This paper highlights recent evidence showing that the way we perceive other people has been
shaped by our evolutionary history. An evolutionary approach provides a powerful tool for understanding
the consistency and diversity of mating preferences and behaviors across individuals and cultures.

Keywords: evolutionary psychology, facial attractiveness, body configuration, voice, menstrual cycle,
muscularity, body fat, dishonest signals

A cursory glance at the women featured in popular men’s mone and steroids enable people to grow taller and stronger.
magazines, such as Maxim or Playboy, suggests that men are Cosmetics, such as make-up, enable people to cover blotches,
attracted to young women with smooth skin, long soft hair, large wrinkles, and imperfections in the skin.
eyes, slender bodies, long legs, curved hips, large pronounced Why do so many people spend so much time, effort, and money
breasts, rounded buttocks, and flat stomachs (Spitzer, Henderson, on their appearance? Why do we find some people more attractive
& Zivian, 1999; Voracek & Fisher, 2002). In contrast, a reader than others? If beauty is only skin deep, why should it matter how
glancing through Cosmopolitan would conclude that women are people look? What follows is a review of the science of sex appeal,
attracted to tall athletic men with moderately muscular arms and targeting recent findings that illustrate the conceptual and heuristic
legs, broad shoulders, little body fat, square and powerful-looking value of an evolutionary perspective.1 We first briefly outline how
jaws, and toned abdominal muscles (e.g., Frederick, Fessler, & evolution shapes the way we process information about other
Haselton, 2005). Notably, however, some body types are rarely people, and then focus on why we find certain faces, body types,
represented. Women’s magazines almost never feature men who and voices appealing. We show how an evolutionary perspective
are very short, elderly, and fat, with wrinkly skin, open sores and enables us to understand and predict ways that women’s prefer-
wounds, and mouths with stained and missing teeth. ences for some of these traits shift across the menstrual cycle. We
The pressure to be attractive can leave many people feeling conclude with a discussion of how individuals have developed
dissatisfied with their appearance (Peplau et al., 2009; Frederick, cultural and technological innovations to enhance certain aspects
Forbes, Grigorian, & Jarcho, 2007), and across cultures and his- of their appearance.
tory, men and women have devised a wide variety of techniques to
enhance their attractiveness (Etcoff, 1999). Corsets give women The Impact of Evolution
the appearance of narrower waists relative to their hips and modern
gyms allow men to enhance their muscularity and shoulder width. It is important to understand that we do not experience the world
Cosmetic surgery enables women to create the appearance of or other people directly. Rather, our experience is a byproduct of
youthful faces, large shapely symmetrical breasts, narrow waists sensory input acting on the nervous system. Our sensory receptors
larger buttocks, and more symmetrical faces. Human growth hor- are evolved neurological mechanisms that convert mechanical,
chemical, thermal, or electromagnetic energy into nerve impulses.
These nerve impulses in turn activate evolved parts of the brain
that translate these impulses into experience.
Gordon G. Gallup, Jr., University at Albany, State University of New
The age-old question, “if a tree fell over in the woods and no one
York; David A. Frederick, Department of Psychology, University of Cal-
ifornia at Los Angeles; FPR-UCLA Center for Culture, Brain, and Devel-
was there to hear it, would it make a noise?” has a clear and
opment; and UCLA Center for Behavior, Evolution, and Culture. definitive answer from a neurobiological perspective. No doubt a
We thank Kelly Carrone, Melissa Fales, Andrew Galperin, Kelly Gild- felled tree would produce intense air borne vibrations, but in order
ersleeve, Martie Haselton, Susan Hughes, Christina Larson, Matthew Pas- to be “heard” or to make a “noise” these vibrations would have to
tizzo, Loni Petricone, and Nathan Pipitone for helpful comments on pre- impinge on an ear and trigger nerve impulses that activate relevant
vious drafts of this paper.
Correspondence concerning this article should be addressed to Gordon
1
G. Gallup, Jr., Department of Psychology, University at Albany, State In biological terms, traits that are more typical of women are consid-
University of New York, Albany, NY 12222. E-mail: gallup@albany.edu; ered “feminine” whereas traits that are more typical of males are consid-
or David A. Frederick, 3rd Floor Mailroom, 1285 Franz Hall, 405 Hilgard ered “masculine.” For example, testosterone “masculinizes” men’s faces
Ave, University of California, Los Angeles, CA 90095. E-mail: by making their jaws squarer, and estrogen “feminizes” women’s faces by
enderflies1@aol.com making their jaws more rounded.

240

2005. unprotected scrotal sack. orientation. The semen evolution to develop receptors and neural mechanisms that medi. Henderson and Anglin (2003) discov- is not an inherent property of rotting flesh. 2005). 2002). and 2) refrain from the photos for attractiveness. Although we take sound for granted. Miller. generating language for The way men rate facial attractiveness in women predicts es- the purposes of communication. Men with faces that women found attractive tended integrity of the testicles and this effect is mediated in part by to have significantly more fertile. It turns out that people with attractive faces also tend to have whether ecological conditions are harsh or stable. Thornhill & Gangestad. ethnic group status. 2002). attractive. of genes that help code for the development of the immune system. having copies of different to target recent findings that illustrate the value of an evolutionary alleles (genes) rather than having multiple copies of the same perspective. and age (see Thornhill & Gangestad. Arden. 2005). and fertility. Hoss. & Wu. Sugar is a good source of without the benefit of prior experience or socialization spend more calories. There are heightened sensitivity to testicular pain (Gallup. A turn prompts each successive generation of males to 1) learn to number of women who did not know these men were asked to rate avoid situations that put the testicles at risk. and sperm mo- tions. Gottfredson. and recognizing who is a kin member and discour. 1994). culture. Hall. (2009) reasoned that individ- independent of their gender. 2006). organisms that evolved robust and healthy overall may have more attractive faces and solutions to these problems had genes that became better repre. sugar as sweet/pleasant had an adaptive advantage in being able to Recent discoveries show surprising links between faces and identify and preferentially exploit food sources that were rich in underlying differences in health. enhanced ability to produce sperm. which may boost their testos- All of these examples are linked by a common theme. disease. and how consistent the link is (Weeden & Sabini. Murphy. which is a ubiquitous measure of health (see DeLamater & Hyde. the body in a thin. respond flexibly in response to changing environmental inputs. Researchers experience for men? Unlike women whose gonads are embedded are currently attempting to isolate the degree to which facial deep inside the body. & Pierce (2009) found small. descended testicles are inherently more vulnerable than ovaries. which aspects of health it relates cavity during embryonic development and usually wind up outside to. Even neonates the nervous system (Barash. & Sammis. sexual uals would be able to accurately gauge a person’s strength based . Sell et al. Finn. Organisms that developed receptors and neurological time looking at photos of faces that adults find attractive (Ramsey.. What follows is a representative review of sex appeal designed Being heterozygous at this loci—that is. By sampling calories. such as entering puberty earlier or later depending on tility. several intriguing explanations for this pattern. 1979). Barbee. Faces also contain cues to fertility. (2003) took facial Males with descended testicles were selected during the course of photos and semen samples from male college students.. In other the women rated the men’s faces were correlated with the semen words. 1999). sperm motility. Sounds only exist in high degree of cross-cultural consensus about which faces are your head. a byproduct of how people are educated and raised? A person’s face even conveys information about their body and Our perception of faces often prevails in spite of our experience. mechanisms that enabled them to experience foods containing Langlois. alleles—provides resistance to a wider variety of potential para- sites and threats to the immune system. ered that both men and women with faces that contemporary Why is being kicked in the groin an excruciatingly painful college students rated as being attractive lived longer. When terone and sperm production. According to this view tractive faces also show fewer subtle morphological deviations attractive faces and sex appeal are learned social constructions from bilateral symmetry. vitality. Many of these systems serve clear adaptive func. In an size that many such evolved systems exist and that these systems archival sample of 425 Army veterans. samples were assayed for sperm count. Both men and women with at- experience and cultural influences. 1998). testicles migrate down through the body attractiveness is linked to health. Faces can contain cues to underlying genetic resistance to aging incest with close genetic relatives (for an overarching sum. rapidly acquiring higher IQ scores (Zebrowitz. Just like sound. fears of snakes and dangerous predators.g. The Major Histocompatibility Complex is an important set mary see Buss. 2004). Rhodes. their behavior.. garner more interest from women. and internal states (e. and other carrion photographs taken from old high school yearbooks and matching eaters could talk. men who are heterozygotic at the Major His- tocompatibility Complex are judged by women as being more Many people think facial attractiveness is a byproduct of attractive (Roberts et al. Druen. sperm count. but the same is true for the way Asians rate placed on your tongue. hyenas. Given the potential dangers of failing to detect a Growing evidence shows that how people rate faces is largely physically powerful rival. which in morphology to generate a composite index of sperm quality. if vultures. Rubenstein. sperm quality: sperm concentration. Are attractive features simply and fitness (Grammer & Thornhill. between general intelligence and several important parameters of Pinker. Caucasian faces (Cunningham Roberts. and sperm ate the experience of pain in response to testicular insults. stores run low. Attractive men may 2009). 2006. Barrett & Kurzban. Not only do Caucasians rate Asian faces much the same Why is sugar sweet? Contrary to the impression it creates when way Asians do. but significant positive correlations ecologies. they would be the first to tell you that foul odor them with obituary records. higher quality sperm. 2006. or men who are generally more there were recurrent adaptive problems. for a meta-analysis see Langlois et al. 2000). For similar reasons. The researchers discovered that how engaging in behaviors that might endanger the testicles. sented in future generations. 2006). EVOLUTION OF SEX APPEAL 241 parts of the auditory cortex. There is a sound is not an inherent feature of the world. Soler et al. the experience of sweet is an evolved property of 1995. Consistent with an evolu- Attractive Faces tionary perspective. seeking out food when calorie trogen levels and female reproductive health (Law-Smith et al. Even intelligence and semen quality may be related. Evolutionary psychologists hypothe. & Griffin. men have evolved to behave in ways that maintain the assay results. & Rhodes.. As a consequence. sweet is not an inherent property of sugar.

68. In addition. Since impreg- nation produces hormonal changes that preclude reimpregnation. & Arons. along with their and can be easily measured and converted into waist-to-hip ratios babies. Waldenstrom. Sell et input is not necessary to develop or maintain this preference al. the because faces contain embedded features that are honest biological authors found compelling evidence for unique activation of neural signals of reproductive fitness. of the 25 highest and 25 lowest ranking companies listed in the Rossner. Platek and Singh (2010) have uncovered neurobiological evi- tractive faces tend to have more accentuated hour glass figures dence for the effects of female body configuration on interpersonal (narrow waists and broad hips). Paralleling these findings. college stu. with This research extends the findings of Shoup and Gallup (2008) optimal WHRs being about . In another study using these images. Ziomkiewicz. Neave. body parts to accentuate their WHRs.7 (Singh. each accounting for about 10% of Singh & Singh. men who that women rated as attractive had more pronounced wedge shaped were born blind or developed blindness later in life were presented torsos (broad shoulders and narrow hips). Playboy models (Tovee et al. those with faces showing average WHRs of about . Brewer. As a consequence. For multibillion dollar corpo- An hour glass figure serves as an important reproductive signal rations this translates into earning differences of hundreds of in other ways (Hughes & Gallup. Women with low WHRs are also Fortune 500. a feminine feature men prefer.. 2000). shows that attractive. more reproductively viable descendants. indicating that visual grip strength scores (see also Fink. Using facial photos of male cek & Fisher. attraction. 1997) and female film stars (Vora- grip strength. from the gene pool. 2010). Without revealing the more regular menstrual cycles (Van Hoof et al. related to reproductive success.242 GALLUP AND FREDERICK on their facial features. dominance. .S. and sexual history. 1993). it was possible to make ness is. Jes- prompt us to experience the faces of reproductively viable people sop. As a con- sequence. with each college students that were cropped at the neck. Attractiveness is not an inherent feature of a person’s Why Do People Find Women With Hour Glass face. Even those who were born many women prefer. with individuals with attractive faces had a better chance of leav. maturity) and global ratings of leadership were both significantly gall bladder disease. They have a lower risk of heart disease. just by seeing their faces. Some people have mature WHRs are more reproductively viable. and are source or identity of the pictures. attractive faces are an evolved Figures Attractive? property of the human nervous system. Women tend to have in subsequent generations. Their research was based on a unique sample of before A recent study by Jokela (2009) based on an archival sample of and after photos of women who underwent cosmetic reconfigura- high school yearbook photos and follow-up surveys from tion surgery. Dixson. ing healthier. the human reward centers in men’s brains when they saw the postsurgical nervous system has been shaped by natural selection in ways that images. Lipson.. and Samoa. dergo relatively unencumbered childbirth. 2007. (Karremans. Attractiveness is also influenced by what is going on below the genes being carried by men that mated preferentially or exclu- neck. were unable to deliver their babies perished. effectively eliminating their genes and those of their mate’s (WHRs) and shoulder-to-hip ratios (SHRs). & Dixson (2010) discovered that men found these as more attractive. Archer. diabetes.. The abdominal enlargement that occurs as pregnancy Body Configuration progresses leads to the loss of an hour glass figure. Why? Evolution is not about survival. their genes became better represented in subsequent generations. & Seydel. stroke. Indonesia. Singh. There are reliable sex differences in body configuration sively with pregnant women would not have been well represented (technically. the Using functional MRI to compare and contrast the way men reason there is good consensus as to which faces are attractive is responded to pre. woman’s waist relative to her hips is an indication of whether she is pregnant. Large samples of who found that men’s faces were cues to their body morphology. whereas men tend to have broader shoul.and postsurgical images of these women. Consistent with this hypothesis. Broad hips are also indicative of un- narrower waists and broader hips that often create the impression derlying pelvic skeletal morphology that enables women to un- of an hour-glass figure. 2006) both converge on this value. Women who differences in body configuration begin to emerge during puberty. They ovulate more often faces others have likable faces. these same males had stronger blind preferred the shape of the low WHR. and facial healthier. a masculine1 feature with mannequins varying in WHR. kidney disease. Andean pastoralists. and various cancers (see related to corporate profitability.000 people who graduated in 1957. In a recent study.. and more sex partners. 1993) and in vitro fertilization (Wass. the differences in company profits. 2009). People who mated preferentially community samples in such diverse areas as Cameroon. or faces that (Jasienska. 2003). in fact. they found Waist-to-Hip Ratios that participants could judge the measured physical strength of Bolivian horticulturists. Ratings of power (competence. trustworthy faces. The configuration of a millions of dollars on an annual basis. it is surgically altered bodies with lower WHRs more attractive using about the perpetuation of genes. These sex not an option during human evolutionary history. Women with low WHRs (narrow waists and broad hips) are dents. Just like foods that taste good. Frankenhuis. Thus. Caesarian sections were ders and narrower hips that lead to wedge shaped torsos. 1997). consistently rated by men in many cultures as more attractive. have convey a sense of power and leadership. people with more attractive faces have more children. More often than not before and after comparisons while holding body mass constant. and U. & Hellberg. Rule and Ambady (2008) asked more likely to conceive as a result of artificial insemination (Zaad- college students to rate photographs of the chief executive officers stra et al. 2006). By using patients who had fat transplanted from other over 10. Ellison. sexually dimorphic differences). 2004). By now it should come as no surprise that women with low Faces also vary in other domains. & Thune. Morgan. & Manning (2007) report that women with at.

see Sugiyama. extreme weight gain is culturally acceptable for infants tend to have higher IQ scores (Mortensen. Adequate signs of appearing pregnant. 2005). slightly better fertility. Thus. a recent analysis of Reuben’s paintings shows comes to body fat (Tovee. In suggested by Cashdan (2009) is that in harsh ecological conditions.. an ostensible benefit of the presence Africa and the South Pacific adolescents from elite families reside of these fatty acids in mother’s milk. This obesity can be thought of as a pregnancy mimic. Breast enlargement beginning at puberty. (e. Oliver. actually have the lowest mortality rates (Flegal. the capacity sial (Rigby. one thing is clear: the extreme thinness prized in many advantage of periods where food was plentiful in order to bridges Western cultures is a recent phenomenon and not inherently times when it was scarce (i. Swami. In fact. production. 2006). Following Ford and Beach’s (1951) ethnography documenting chain polyunsaturated fatty acids that are important for neurolog. resources may be valued more highly than body types that indicate financial. inevitable abdominal enlargement that accompanies pregnancy struating sooner. The attraction that many men have to enlarged breasts may be a Perhaps one of the biggest mysteries in evolutionary psychology is consequence of the connection between female body fat and re- the preference for thinness in industrialized societies.. presence of more gluteofemoral fat consisting of unique long. Saguy.K. Research at the Centers for Disease Control has shown that individuals who are overweight Body Fat (BMI ⫽ 25–29). such as rural Malaysia (Swami et al. & Gaesser. & Reinisch. lower WHRs) had significantly higher resources are not as plentiful. weight counterparts. & Furnham. whereas Whites in the U. sexuality. Oliver. 2010). women with relatively high fat stores (Lassek & Gaulin. Saguy. Despite healthier than being in the overweight category (for reviews see the preoccupation many women in the Western world have with Campos. Ernsberger. 2005). each of these conditions involve the gradual loss of an quarter of a woman’s body weight that they actually become hour glass figure. In less socioeconomically developed societies where more gluteofemoral fat (i. 2002). EVOLUTION OF SEX APPEAL 243 Evidence also shows that the configuration of a woman’s waist exposure to Western media only explained a small percentage of the relative to her hips may be an index of the availability of special variation in preferences. 2002). they women with higher WHRs might be preferred because the testos.. periods of feast or famine). shown that the onset of puberty in girls is largely independent of The other point to consider in the context of the contemporary age and requires reaching a point where approximately one sixth aversion to body fat among women in the Western world is that (17%) of their body weight is represented by body fat. Moreover. reproductive impairment for women than being overweight— but ductive function. & Gail.e. & Mangalparsad. Swami.” where they are fed high-fat diets in order to Finally. Life . obese) women go through menopause later than their normal 2008). and attractiveness. particularly in the period preceding marriage. But it is not until body fat stores reach about one and obesity. widespread preferences for plumpness in 81% of nonindustrialized ical development. in “milking huts. than in South compete for and obtain resources and status. not .K. A recent cross.. This may be one of the reasons why overweight (but not changes that arrest fertility.78. One possible solution to this conundrum attractive. there is growing evidence that gen. tem- body fat as a means of estimating the availability of these “brain poral.e.70 (e. Although the Make no mistake about it. women. Lower body fat on women’s levels of body fat were rated most attractive in rural nonindustrialized hips and buttocks can be distinguished from upper body fat by the sites. Williamson. that women on average had a WHR of . 2006. fat is highly adaptive. Research reviewed by Rose Frisch (2008) has not obese (Frisch. South African Zulus find heavier bodies more Gray. 2007). stores of body fat may be particularly valuable to women during pregnancy and lactation because these conditions can increase Breast Enlargement caloric demands by 10% to 15%.g. In parts of Sanders. under conditions nowadays in which food is plentiful. as in the case of anorexia nervosa.g. maintaining a slender figure. In harsh ecological Africa. is largely due to the was preferred in all industrialized countries studied. and individual differences in attitudes toward obesity and building” resources.. that is inde- cultural study of 41 sites across 10 world regions found that thinness pendent of a woman’s lactational status. 2010). Campos. plumpness is linked with psycho- cognitive test scores. prefer thinner bodies. found that these Zulus adopted preferences that were much more terone levels needed to create higher WHR enable women to better similar to those of individuals living in the U. Lassek and Gaulin found that women with body weight. conditions. Jones.. 2004. Although hardly logical traits of fertility.70 across history and all cultures al. Michaelsen. Einon. For example. and so did their children. Furnham. and reported localized accumulation of adipose tissue or fat in the breasts. 2006. decades of research has documented cross-cultural. Flegal et al. body types that indicate an ability to obtain scarce particularly body fat. A recent paper illustrates the role of culture and context when it 2009). but not obese.. Swami. Some body fat may be beneficial. Graubard. it is important to acknowledge that preferences for become properly plump in preparation for marriage (see Swami et WHR do not always center on . 2008). 2008). to store energy in the form of body fat enabled organisms to take 2006). Because of the appears to be the reason overweight little girls often begin men. In contrast. Using WHRs to index the ratio of upper to lower societies..K. Ernsberger. & Furnham. the same is true for obesity (Frisch. and ecological contexts. Because food exact nature of the link between body fat and health is controver- was not always plentiful in the ancestral environment. just as pregnancy leads to hormonal fertile. are malleable based on changing social. their study of South African Zulus who moved to the U. respectively (Dufour & Sauther. being thin and emaciated carries a far greater risk of eralized stores of body fact are critically related to female repro. & Gaesser. Traditionally. These findings suggest that preferences for some traits. In many of conclusive it is interesting that people who were breast fed as these societies. Extreme weight loss due to exercise and/or women’s preoccupation with dieting and weight loss may be starvation and the resulting depletion of body fat often leads to driven in no small part by an unconscious attempt to minimize any infertility in women.

In almost all of the testing sites. which cause breasts to droop and the International Body Project (2010) examined preferences for become less firm.. Pat- rocal (Frederick. bodies that varied systematically in muscularity or body fat. humans show an exaggerated sex difference in overall reinforced the development and maintenance of long-term pair muscle mass and physical strength. Additionally. 1997). and clear skin In support of the hypothesis that muscularity is attractive to women. there is suggestive evidence that the incidence of muscularity and other masculine features may function to signal infestation by human public lice has decreased along with the superior health and immune system competence. re. Frederick. and in male-male competition (Lassek & signal or advertisement for health and reproductive viability. There is reason to believe that human pubic hair evolved as a affairs. Participants rated a series of images of male effects on men’s satisfaction with their partner’s breasts are recip. women rate males with wedge shaped torsos as more tractiveness: very low and very high levels of muscle mass were attractive (Dijkstra & Buunk. Shoulder-to-Hip Ratios women preferred men who were more muscular than average and men wanted to be more muscular than average (in the few remain- Broad shoulders and narrow hips in men are the sexually di. Since was more muscular than their committed sex partner. et al. aggressive behavior which may pose a risk energy stores and ovulatory potential (Gallup. hour glass figures. Men with high research suggests that male muscularity may be an important cue SHRs tend to be stronger and more aggressive in high school of attractiveness. and the across 26 countries. is that much of the existing psycholog- elevated levels of salivary estradiol and higher fertility (Jasienska ical research on male attractiveness is limited to Western contexts. so the evolved propensity to attend to overall levels of male physical surface area of the body could serve as a radiator to keep the brain prowess. there men rate females with narrow waists and broad hips as more was a clear inverted-U association between muscularity and at- appealing. edged pleasant and even erotic properties of skin-to-skin contact spring (see Frederick & Haselton. however. function and increased muscle mass entails high energy demands. SHRs signal various embedded dimen. Swami. and the 56 members of women age and nurse children. and as a consequence enlarged breasts may have evolved high levels of testosterone can impair health. to the woman herself). Indeed. The loss of body hair is a distinguishing feature that sets humans apart from other primates. 2009. & Gallup. as well as produce to compensate for the loss of ovulatory cues by signaling ample unpredictable. there were significant cross- developed in this paper.2 The widely acknowl- advantages associated with muscularity can be passed onto off. Indeed. and insects are women and more successful in competitions with males (Frederick visible signs of current disease and parasitic infection. cent evidence shows that women with large breasts actually have One problem. 1982). exposing smooth unblemished skin may have become a acquiring resources. however. and that local ecological factors can shape the (Gallup. men that mated preferentially with females who did which states that women will experience less attraction to individ. In every culture without exception. and Ghana want to be more muscular. The ability to recent increase in pubic hair removal by women in some parts of build greater muscle mass is partially heritable. students desire greater muscularity in order to be more attractive to The presence of blotches. Gaulin. Exposure to Western media. Because testosterone compromises immune eliminate body hair as a refuge for parasites (Pagel & Bodmer. effective body hair removal was not an option during most of human they introduced the “inverted-U hypothesis of masculinized traits. Just as for women who have degree to which is it important. terns of preference for muscularity were significantly more prev- alent than preference for body fat. & Lever.” evolutionary history. and consistent with the thesis not preferred. Finally. et al. ing sites. the effects were in this same direction but failed to reach morphic counter point to an hour glass figure in women. Men with broad shoulders and narrow hips (high attractive. Many & Buchanan. Looking across primate made possible by the loss of body hair could likewise have species. There are any number of reasons why Muscularity this may have happened. showed only SHRs) have higher testosterone levels (Kasperk et al. 2005). 2004). Puts. Frederick and Haselton (2007) found that muscular male 2 Pubic hair removal is a peculiar anomaly from an evolutionary per- college students reported more sexual partners and short-term spective. There may also have been selective pressure to to build and maintain. 2010). and male college adaptations to resist infection (Symons. not have pubic hair would have been pedophiles. Skin quality can be an important cue for health and genetic Ukraine. cultural differences in the degree of muscularity that women found sions of fitness. Body weak relationships to these preferences in most settings. 2008). Research in the United States shows that women muscularity in 41 community and college samples in sites tend to become less satisfied with their breasts as they age.. Hughes. 2003). Just as statistical significance). open wounds and sores. 1995. 2007). White. Notably. & Gallup (2004) found that men with more accentuated wedge shaped torsos began having sex Skin at earlier ages and have more sex partners. diseases or parasites can produce yellowing of skin. Sugiyama. Many men in the United States.. 2007). 2007). This configuration is also related to physical prowess. .244 GALLUP AND FREDERICK span developmental changes in the shape and configuration of the uals with low levels of masculinity (because it is a cue of lower breasts are correlated with changes in a woman’s capacity to testosterone and dominance) and also very high levels (because ovulate. 1990). volatile. 2006). Dispenza. however. The value of breasts as a cue to fertility changes as To remedy this problem. and that women’s most recent short-term sexual partner salient puberty signal and a means of preserving sex pheromones. One possibility was the need to more Frederick and Haselton (2007) proposed that women have an effectively cool an ever expanding brain (Falk. Muscularity requires extensive calories and testosterone from overheating. Peplau. provided an advantage to ancestral males in terms of hunting. suggesting that muscle mass bonds between mates and between mothers and infants as well. 2001). suggesting that the world (Armstrong & Wilson.

Voice. and that this preference would be stron. 2008). and Sexual Behavior Voice Attractiveness In another study voice samples were taken from college stu- There are two approaches to the study of attractiveness. as soon as the caller starts to Women’s voices not only change with the onset of puberty. as well as one’s own attractiveness and 1999). & Pipi- prevalence across cultures influences different mating preferences. Sugiyama. tone. one would expect preferences for consequence of being better able to withstand prenatal and devel- smooth. & Perrett. Men with attractive focuses on proximate rather than ultimate mechanisms. even if you do not know who it is.. tempts to identify particular features such as pitch that distinguish and women with attractive voices likewise had more accentuated attractive from less attractive voices (e. Rhodes. it was discovered that ratings of voice attractive- fertility (Symons. EVOLUTION OF SEX APPEAL 245 denotes an absence of communicable diseases. . Prior to the invention of artificial illumination. 2006). associative learning. Both men and women with Burt. health and fertility are Gangestad. The other approach havior (Hughes. finger length. even after controlling for a variety of other variables. Frederick. Jones. the sound of a person’s voice functions as a medium for the transmission of Voice and Fertility biologically relevant information.g. or bilateral symmetry that contributes to attractive voices also reported having sex at earlier ages. and Buss (2005) examined how parasite compromised (for a recent review see Gallup. This holds what they say for samples collected during the ovulatory phase. The one dents. Ratings of voice attractiveness peaked from 1 to 10 as their voice is recorded. 2005). and were more likely to have engaged in sexual infidelity. The evidence shows that just like a person’s face. each side of the body and compared in a sample of male and sistent with this prediction. the same is true of faces. On the other hand. In a recent longitudinal study. teria (Fessler & Haley. their SHRs and WHRs were measured. and at- voices had more accentuated wedge shaped torsos (higher SHRs). which is the constant and it keeps the content neutral. ability to attract mates (Pillsworth. 2005). Ratings of voice principle means of communication at night.e. and they were featured in this paper involves looking at different traits as evolved invited to complete an anonymous survey about their sexual be- signals for important biological information. however. Harrison. people with attractive voices tended to diminishes much more rapidly for women. but rather to how they say it. Who would have known that Cameron. The capacity to produce ness were inversely proportional to the magnitude of fluctuating offspring decreases as men and women age. Con.. The emotion of disgust may have evolved initially because it logical development (Livshits & Kobyliansky. One can imagine The Sound of Symmetry the benefits of coming equipped with a system that inhibits sexual arousal when encountering people covered with infected boils that The extent to which features on one side of the body match are oozing and bursting with puss. unwrinkled skin. 2008).g. sound was our Some of these effects were quite pronounced. when listeners point of maximum fertility. status the person has.. If physical attractiveness is a cue to health and resistance to In the initial voice study on this topic seven representative parasites.. Evolutionary theory would Just as attractive voices signal subtle. Talking on the telephone approximates what it the mere sound of a person’s voice would be related to their sexual used to be like to have a conversation at night because on the behavior? telephone you do not see the person you are speaking with. that as random deviations from bilateral symmetry increase.g. When other students who did not know the speakers were asked to Skin quality can also provide an index to age and therefore rate their voices. rarely noticed differences suggest. multiple voice People who participate in much of the research described below samples were taken from females at different points in their cycle are typically asked to speak into a microphone and simply count (Pipitone & Gallup. as well as parasitic infections. opmental insults/perturbations. When you answer the phone. Feinberg. Given the association show fewer deviations from bilateral symmetry as an ostensible between age and fertility. physical attractiveness was ranked as female undergraduates (Hughes. 2006). tives as they do not show the same hormonal fluctuations. that this preference would be balanced with in fluctuating asymmetry that bear in important ways on health and other cues besides youth when choosing a partner. 2002). or socializa. 2008). rather than learning this pri. more sex attractive faces (e. known as fluctuating asymmetry (FA). and this capacity asymmetry. Haselton. 1991). menstrual cycle. People with faces that are more to obtain resources. partners. such as ability fertility. and the social influence and symmetrical are also rated as being more attractive (Perrett et al. 2004). 1992). hour glass figures (lower WHRs). parenting ability. but there is growing evidence others that are teeming with potentially deadly parasites and bac. Body Configuration. and this was undoubt- attractiveness accounted for almost 25% of the variance (i. ger for female bodies than for males. That way. they speak you usually know 1) whether it is a male or a female. wrist diameter) were measured on preference for physical attractiveness when choosing a mate. prevalent. resist minor insults and perturbations that occur during embryo- tion. and 2) also vary as a function of where they happen to be in their whether it is an adult or a child. Dispenza.. Each more important in a mate in societies where parasites were more student’s voice was also recorded as they counted from 1 to 10. & Gallup. then high parasite prevalence should evoke a stronger features (e. those on the other has been taken as evidence for the ability to marily through trial and error. Little. & Gallup. In other words. No one served as a motivational system that prompts people to avoid shows perfect bilateral symmetry. there were no cycle are asked to rate these recordings they will not be responding to dependent voice changes for women using hormonal contracep- what subjects say. edly one of the reasons we developed spoken language based on individual differences) in number of sex partners among the fe- sounds as opposed to sign language based on gestures (Gallup & male college students in this study. 1995.

Geoffrey Miller and his associates (Miller. 2007). and also make a telephone conversation an important element in the olfactory cues that men found more appealing during high fertility. suggesting that they were responding to voices rated as sounding more dominant could be distinguished cues related to the women’s fertility status (Pillsworth & Haselton.246 GALLUP AND FREDERICK Apicella. Finger Digit Ratios. and/or they had changing vocal. more masculine finger digit ratios indicative of greater show more skin and to wear miniskirts when they are ovulating prenatal exposure to testosterone. But what is particularly interesting about this for individuals with traits that are cues to underlying genetic effect is that the relationship between attractive voices and attrac. Unlike chimpanzees. but this tion. The effect was stronger the closer 2D/4D (Manning. The earnings were related to their menstrual cycle. Pillsworth. and Strength partner does not possess these traits. maturity. tive faces is dependent upon the maturational/hormonal status of human females do not exhibit obvious changes where their genitals the listener. The evolutionary logic behind this Given the foregoing. cycling earned approximately $335 per 5-hour shift when ovulat- ration. Whereas a deaf date would be a date sound unheard. $260 in the luteal phase. & Frederick. they had person first. . & Jordan. status. Fienberg. discovered that female women were to the day of ovulation. however. have tested whether there are other more subtle cues of voices by prepubescent girls failed to predict independent ratings ovulatory status. influences. 2002). the sound of your voice conveys information tips on average during the fertile phase of their menstrual cycle about what happened to you during prenatal development. without independent evidence of paternity it remains possible that men with more masculine voices merely think they In many species when females ovulate they enter estrus. Why did their emerging evidence would suggest that before you agree to a blind income increase so dramatically when they were ovulating? The date it might be prudent to have a telephone conversation with that possibilities include that the women were more sexual. (Haselton. a period have more children and/or they may be more prone to exaggerate of heightened interest in or receptivity to sex. particularly if their current long-term Voice. 2008). 2007) found that lap dancers made 80% more To summarize. sexiness. Sell et al. while the lap dancers were aware of differ- ductively relevant information can be used to make a distinction ences in the amount of money different dancers made. Although suggestive of a link between fertility The Menstrual Cycle and voice. In a series of studies. physical.. The length of the 2006. ing. Evolutionary psychologists. producing high they were in their fertile phase. occurred primarily for women whose partners were relatively less voices can be rated along any number of dimensions (e. second digit or index finger relative to the fourth digit or ring Changes in women’s sexual motivations can have interesting finger (expressed in terms of finger digit ratios: 2D/4D) appears to effects on women’s self-ornamentation. they were in their cycle compared to those taking hormonal con- The fact that the human voice is such a rich source of repro. attractive. your hormonal statues. When females are most likely to attractive voices also have more attractive faces (Saxton. Nat- whether you are on birth control pills and if not. both the perception as well as the sociated with testosterone. blind to the women’s fertility status. it should come as no surprise that men with preference is straightforward. your strength. Pastizzo. & conceive. none realized that their unseen. Mortezaie. creased sexual desire for other men when ovulating. Thornhill & Gangestad. and general robustness are stron- production of voice would appear to be subject to hormonal gest when they are in the ovulatory phase of their cycle. Indeed. People exposed to high levels and low fertility phases of their menstrual cycles. as well as between “blind dates” and “deaf dates. and Marlowe (2007) found that voice was decision making process about dating a person you have never associated with self-reports of reproductive success among male met. One factor driving these college students with dominant sounding voices had significantly results is that women in committed relationships are more likely to lower. Puts (2005) has shown. Independent of testosterone during prenatal life tend to have ring fingers that are judges. hunter-gatherers. and Gallup (2008) found that women with partners were ovulating. your age. whereas exposure to indicated that women were trying to look more attractive when high levels of estrogen has the opposite effect. from others based on their finger morphology. (2010) recently found that men’s voices contain cues of In a landmark study. intelligence.g. the growing popularity of Internet dating may greater energy. dominance). that men females often mate preferentially with males displaying costly with low pitch voices tend to be preferred as short-term mates traits linked to testosterone or dominance status (for a review see when women are in the ovulatory phase of their menstrual cycle. Bleske. ap. benefits that can be passed on to offspring. the effect only holds when listeners are post-pubertal enlarge dramatically during estrus. rated their photos and longer than their index fingers (low 2D/4D). re- searchers found that women with dating partners experience in- Voices not only predict facial appearance and body configura. 2006). physical strength. In addition to attractiveness. Not unrelated to these findings. Ratings of the same men’s however. and $185 when menstruating. women were be influenced by exposure to different levels of sex hormones photographed when they came into the laboratory during the high during embryological development. less attractive men were more likely to proachability. engage in loving and jealous mate guarding behaviors when their Hughes. One function of these shifts may be to motivate women to seek out a short-term sexual partner who can pass on advantageous traits to their offspring. traceptives. Thus. they reveal other differences. your sexual behavior.” A blind date is a date sight day to day fluctuations in their own tips. In such species. Haselton & Gangestad. your body configu. Moreover. 2006). women (adolescent and adult females). where you are in urally cycling lap dancers also made more tips regardless of where your menstrual cycle. claims of paternity. where men with lower pitch voices claimed to have more children. Hughes et al. In one study. they have been selected to show increases in preferences Roberts. Caryl. Those that were normally says something about your gender. and whether women’s preferences for traits as- of the speaker’s faces. In parallel. It also than when they were menstruating. Tybur.

Hughes et al. In theory. Not only do females place more emphasis on kissing. 2005). 1997) Honest Versus Dishonest Signals during ovulation. and credit. the ovulatory phase. 2009). As a case in relationship barometer. and especially those who spend more view kissing as more important at all stages of the relationship than money than they make have significantly more sex partners than do men.g. appearance but leave your reproductive potential unaffected. Cycle dependent changes in attractiveness such as faces are honest signals since they are correlated (positively in the these likely functioned in the ancestral environment to synchronize former case and negatively in the latter) with a person’s capacity insemination with ovulation as a means of increasing the chances to reproduce. By comparison. interesting but not altogether unexpected sex differences in kiss. and imperfections in the skin. this analysis. trying to gain A study by Kruger (2008) provides a compelling example of sexual favors). Buss. Men. & Gallup quent generations is to pair them with someone who is reproduc- (2007) discovered that the majority of both male and female tively viable. and sexy bodies are At the moment of a kiss there is a very rich and complicated all evolved features of the human nervous system. . Cosmetic surgery may improve your appearance. 2005). Many females would not consider having sex with of reproductive competition. on the other logical cues. Thus. Preti. dishonest signals in a different domain. may be a cue for an inability to resist pathogens. often use kissing as means to an end (e. dishonest courtship. women who spend rather than save. . Artificial Kissing sweeteners improve the flavor of food. dishonest signals often improve your of impregnation. When it comes to the use of dishonest signals in the arena as after sex. as already noted. EVOLUTION OF SEX APPEAL 247 As Miller’s study suggests. Dental disease.000 techniques ought to be dependent upon the extent to which they college students. Cosmetics. Kissing may also be a detect and avoid instances of reproductive deception. & Gallup. “I love you. such as make-up. 1995. 2003).” survey indicated they would be happy to have sex without kissing. Clearly one of exchange of visual. men can use conspicuous spending to exaggerate their more prone to emphasize the importance of kissing during as well earnings. Gallup. Whereas foods that taste sweet are usually a significant source of calories. 1970) and appears to be part of an but your reproductive capacity remains unchanged.” suggesting that the people at Maybelline to use kissing in long term relationships to update and monitor the are implicitly aware that their product can be used to mask bio- status of the relationship with their partner. Scutt. most males in this to “Maybelline. In contrast. Using a large questionnaire administered to over 1. enable people to cover blotches. Hughes. According to evolved human courtship strategy (Hughes. sultry voices. but they continue maybe it’s Maybelline. hand. and poor teeth would have been a significant handicap. lesions of the mouth. or feigning good intentions for purposes relationships that the amount of reported kissing is related to of gaining sexual favors is a common male sexual strategy. the existence of dishonest signals would be expected and men were far more likely than women to agree to have sex to create selective pressure for adaptations that enable people to with someone who was not a good kisser. 2008). Gulledge. function to assess the health. placed more emphasis on kissing in wrinkles. analysis. and reproduc- ered that their faces were rated more attractive when they were in tive viability of a prospective mate. 2002). Harrison. Likewise. The reason “first impressions count” is because many college students reported having found themselves attracted to of the features of interpersonal attraction are related to important someone on one or more occasions. and breast asymme- tries diminish (Manning. inability to chew (Geher. Sug- iyama. for relationship satisfaction (Gulledge. Tasty foods. and chemical information the most effective means of piggybacking one’s genes into subse- based on olfactory and gustatory cues. saccharine is the equivalent of a face lift. . but leave its nutritional Kissing between romantic partners occurs in over 90% of hu. reliant on kissing as a mate assessment device. & Leinster. 2001) as well as vaginal odor samples (Doty. women’s WHRs become more accentuated (Kirchengast & Gartner. Women are not only more metics company Maybelline reads “Maybe she’s born with it . attractive faces. Oral (Haselton. Whitehouse. Roberts et al. albeit subtle effects there is reason to suspect the existence of unconscious. cycle variation in fertility can have important. there is evidence that menstrual kissed them for the first time they were no longer interested. hard-wired on how women are perceived. (2007) found evidence for some make you look more reproductively viable. particularly without further evidence of sincerity food. Female body odor (Singh & Bronstad. Because men with attractive faces have higher quality recordings taken during midcycle are more attractive (Pipitone & sperm and women with facial wrinkles are less likely to conceive. 1975) Honest signals are those that reflect a person’s reproductive are also rated as being more pleasant and. value unaffected. Ford. Oubaid. health may indicate heritable differences in susceptibility to peri- odontal disease and environmental stresses (Symons. postural. There is evidence for couples in committed point. & Stahmann. & Huggins. Kruger found that men While both sexes rate kissing as a highly romantic act. Even common haliotisis. man societies (Eibl-Eibesfeldt. so let’s go to bed. they those who are more frugal. commitment skepticism is common among women ner’s health. (2005) photographed mechanisms that are triggered at the moment of a kiss which may women at different points during the menstrual cycle and discov.” Consistent with this Another function of kissing may be to assess a potential part. the effectiveness of all appearance enhancement 2007). for example. genetic compatibility. & Angleitner. example. tactile. ing. and as a way of attempting reconciliation.. The tagline for the cos- romantic relationships than did men. which stems from bacterial Concluding Remarks growth in the mouth. only to discover that after they underlying dimensions of fitness. “Mastercard” has become an analog someone they never kissed. Women. Harrison. voice potential. Nowadays with the advent of easy are more likely to insist on kissing before a sexual encounter.

. 113. & Lipinski. Patterns of sexual behavior. B. women dress to impress. G. & Gail. W. Current Anthropology. J. Fessler. 33. Gottfredson. F. outside-inside dichotomy in disgust and bodily experience. social construc. (1975). A. 49... J. 40 – 45. April).. Frederick.. Frederick. N.. 1099 –1107. G. 81– 86.. Haselton. D. V. G. Psychological Inquiry. Barbee. G. ior. Review of General Psychology. (2006). Forbes. R. A. Buchanan. A. H. C. G. J.. 277–282. A. Science. M. M. D. 509 –518. A. G. Comparative and evolutionary Geher. G. Cognition Hormones and Behavior. Grigorian. Journal of Family Therapy. . Intelligence. Evolution Harper & Row. 19.206 undergraduates. 233–242. (1998)... (2007). Ratings of voice Ford. A. Oubaid.. & Seydel. G. G. & Buunk. M. D. (2006). V. Association. R. D. A. 233–242. and Human Behavior. Physical attractiveness: The Project (2010. & Stahmann. (1999).248 GALLUP AND FREDERICK Although beauty is still in the eye of the beholder. (1970).. M.. & Hyde. the muscular ideal: Men’s body satisfaction in the United States. M. physical affection types and relationship satisfaction. K. B. Evolutionary intensity and pleasantness of human vaginal odors during the menstrual foundations of cultural variation: Evoked culture and mate preferences. Peplau. & Huggins. Berezovskaya. Survival of the prettiest: The science of beauty. and sexual behavior in males. G. J. L. P. S. (2007).. Permanent breast enlargement in human females: C. P. G.. New York: Harper & Row. Dijkstra. (2007). C. P. and Ghana. K.. Sex. P. P. T. 1316 –1318. (2008). The Barbie mystique: human nature. G... ioral and Brain Sciences. Gallup. P. Hughes. ior.. Do References representations of male muscularity differ in men’s and women’s mag- azines? Body Image. First Gulledge. 597-60l.. 3–19. 69. Barash. Evolution and Human Behav. Finn. (2007). A.. Haselton. E. 265–266. Consistency and variability in the cross-cultural perception of female Gallup. 295–304... I. M. Accuracy and oversexualization in cross-sex mind-reading: dimensions of the energetic of human pregnancy and lactation. & Gangestad. Doty.and estrogen-dependent traits. on the whole. A. G.. 7.. Satisfaction with breast size and shape across the lifespan. & Jarcho. A. J. F. L. & Marlowe. M. Desiring history of the species. J. E. Jr. B. M. 20. A.. Gallup. The epidemiology of overweight and obesity: Public health crisis or Frisch. 20.. Journal of Comparative Psychology. & Haselton. J. (2007).. Handgrip strength androgen.. M. M.. longevity. & Thornhill. Journal of Human Biology. B.. N. Pillsworth. & Anglin. (2009). and Emotion. Sex Arden. tiveness of human male voices. 57.... G. Intelligence Roles. 317–327. D. & Perrett. M. D.. H. G. The predicts reproductive success in male hunter-gatherers. (1992). & Sammis. 93–98. L. (2005).. D. J.. Journal of Sex Research. Journal of Personality and Social Psychol. Ukraine. Is our mental machinery “colored” by a visual bias? Metaphor and ogy. A.. C. 35. objectification and body satisfaction among 2. & Gallup. Symbolic Activity. G. (2001). and obesity. K. 5.. E. M. 31. A. 584 – 602. M. Haselton. Jr. 24. Voice pitch Frederick.. (2007). (l982). M. D. Miller. L. G... F. Brain evolution in Homo: The “radiator” theory. cycle. 190. D. Hormones and Behav- 82– 87. views. E. G. Guarding the perimeter: The women’s desires and men’s mate guarding across the ovulatory cycle. D... (2008). Dufour. sexes. S. 517–526. Manipulations of fundamental and formant frequencies influence the attrac. Morphology evoking nature of a rival’s body build. A. S. BiologyLet. C..-H.. G. M. & Gallup. White. & Haselton. D. Evolutionary Psychology. Burt. Modularity in cognition: Framing the Journal of Sexual Health. G. M. R. Ovulatory shifts in human female ornamenta- physical strength to women. G. R. J. (2003). 12. & Wu. that the eye of the beholder has been shaped by the evolutionary Haselton.. Personality and Social Psy- pubic louse? Sexually Transmitted Infections. I. A. Frederick. G.. L. predicts body morphology. A.. Jones. M. & Lever. (2007). Neave. 59.. N. (2007). 11. 17. Why is muscularity sexy? Armstrong. & Pipitone. Etcoff. S. Cancun. International Barrett. puberty and fertility. C.... 331–347. 35. debate. Druen. I. P. (1990). M. Waist-to-hip ratio across cultures: Trade-offs between Gallup. D.. Grammer. Male facial appearance signals Frederick. & Buss. Psychological Review. J.. & Cameron. Jr. 2. R. (2009). & Kurzban. Did the “Brazilian” kill the Tests of the fitness indicator hypothesis. UCLA Body Project I: Gender and ethnic differences in self- ters. aggression. lies. 628 – 647. R. D. Jr. A. 108.. 51. Biological Re- moral panic? International Journal of Epidemiology. (2006). Little. (2005). 37. A. (2006). D. F.. Haselton. The whisperings within: Evolution and the origin of Frederick. Behav. 22. Gallup. Saguy. (2008). (2006). G.. Cashdan. Psychology. (2002). 1861–1867. Jr. 28... Modality specific metaphors: physical attractiveness. 333–381.. (2005). H.. G. 561–568. Body fat. Cunningham. Journal of Evolutionary across 10 world regions: Results from the International Body Project I. and behavior: Phrenology revisited. (2008). ness. 335–341. B. The American Feinberg. Mortezaie. Henderson.. & Wilson. attractiveness and sexual selection: The role of symmetry and average- ior patterns. M. Oliver. (1951). American Journal of Human Biology.... G. Gulledge.. Preferences for muscularity in 26 countries objective ornament and subjective self-ratings. P. tion: Near ovulation. D. L. Feinberg. D. chology Bulletin. 14. A. R. Sadeghi-Azar. & Angleitner. Buss. 161–188. 75–96.. 25. and strategic interference: The psychology of deception between the Falk.. 351–356.. D. Archer. A.. Flegal. H. G. C. Graubard. Sex differences in the jealousy. Paper presented at the annual conference of the Western Psychological Campos. Ernsberger. Evolutionary Psy- 10 –18. D. (2009).. G. chology. (2005). Ford.. R. Their ideas of beauty are. Psychology of Men & Masculinity. D. G. Conditional expression of Fessler. (1979). T. Roberts. G. Human (Homo sapiens) facial Eibl-Eibesfeltd. On the origin of tionism in the study of human sexuality. Journal of Human Evolution. Essentialism vs. 8. (2004). 31. R. 293. N. M.. M. F. Apicella. 7. Evolution and Human Behavior. Peplau. Love and hate: On the natural history of behav. Mexico. 49. D. Frederick.. 423– 429. A. & Haley. R. 55– 60. Swami. M. R. M. (2003). & Manning. overweight. Bleske-Rechek.. descended scrotal testicles: The activation hypothesis. J. Evolution and Human Behavior. S. 3. W.. L. 103–117. and semen quality are positively correlated. M. DeLamater. the same as ours: A sociobiological analysis. M. Animal Behaviour.. 200 –211. 297–304. & Pierce. S. (1995). B. K. 1167–1183. Changes in the Gangestad. & Fink. M. (1994). 261–279.. Jr. R. Journal of the American Medical Association. 682– 684. 29 –38. D. R. Williamson. B. Facial attractiveness predicts Excess deaths associated with underweight. D. A. K. & the 56 Members of the International Body Brewer. New York: Methuen. & Beach. Romantic Anchor: New York. 82. (2006). 13. 3–23. 68. & Gaesser. New York: attractiveness predict sexual behavior and body configuration. we now know Frederick. American An adaptationist approach. E. Personality and Social Psychology Bulletin. D. 7. Dispenza. & Sauther. Preti. R.

Jones. G. (1997). 388 –397. D. S. S. Evolution and Human Behavior. 24. I. Role of body fat and body shape on Pagel. D. A. 328. C. Perrett. Perret. Ziomkiewicz. (2006). women’s preferences for male voice pitch.. C. N. V.. .. J. Sznycer. (2008). Evolution and Human Behavior. 441– 466. Harrison. (2009). Evolution and Human Behavior. J. Changes in fat distribution (WHR) 6-month-old infants. Petrie. Evolution and Human Behavior.. S. G. E. Developmental Science. O. theoretical review. A. Skeletal site-dependent expression of the in humans. M. Kasperk. havior. K. A. 109 –111. and Shoup. Annual qium. Large breasts and narrow waists indicate high reproductive New York: Penguin.. J... 31. . 93–108. Jr. (2002). M. & Hillier. EVOLUTION OF SEX APPEAL 249 Hughes. Mate preferences among the Shuar of Ecuador: symmetry: Voice as a marker of developmental instability. (2008). (2010). (1997). P. E. G. M. & Galllup. Waist-hip ratio and cognitive chief excecutive officer’s appearance predict company profits. 26. Hughes. S. G. N. 176 –181. M. Pipitone. (2002).. S. J. Evolution and Human Behavior. G.. The Journal of the American Medical Association. M.. lesbian adults. Pastizzo. J. Rubenstein. G. D. B. R. A. D. Puts. Jr. Langlois. Helmboldt. C. about their bodies and their behavior: What you see is what you get. (2009).. S. J. Cosmides. L... C... Digit ratio: A pointer to fertility.. J. Manning. Sell. Archives of Sexual Behavior. & Gartner. Evolutionary Psychology. C. Bio. attractiveness judgements of children. Human adaptations for the visual assessment of strength and sible measure of developmental homeostasis in humans: A review. S. Journal of Personality and Social Psychology. C. H. 6. Frederick. The association between duration of breastfeeding and adult Cross-cultural consensus for waist-hip ratio and women’s attractiveness. (2010). Evolution Platek. Male sexual attractiveness symmetry revisited: Subjective and objective analyses of voice. D. & Gallup. G. doi:10.. Krauss. The sound of Pillsworth. M. (2010). judgement of female health and attractiveness: An evolutionary perspec- Proceedings of the Royal Society of London: B. B. O. S.. T. J. 47–57. 26. N. Optimal waist-to-hip ratios in women and Human Behavior.: Rutgers University Press. Singh. M. M. . Scutt. (2003). L. Ovulatory cycle effects 293–307. Lipson.. W. G. M. D. Larson. humans: Evidence from the last 20th century United States. (1999). (2007).. G. Borcsok. Gosling. (2005). Burt. e9042. 713. (2003). Michaelsen.2010. 7. Jr. Physical attractiveness and reproductive success in 268 –274. S. Sex differ. 390 – 423. New Brunswick. I. Adaptations in humans for F. Costs and benefits of fat-free muscle mass in men: Relationship to mating success. 268. L.. J. Series B. & Arons.. to ovulation. Evolution and Singh. Blind men Puts. (2010). Sell. Singh. von Reuden. T. 575–584. 342–350. & Reinisch. G.. R. Manning. D. Cornwell. D. Women’s voice attractiveness logical Sciences... Hallam. 797– 801.. (2000). 26. Niethard. D. 247–258. Penton-Voak. G. C. Origins of a stereotype: Categorization of facial attractiveness by Kirchengast. Jr. A.. 201–211. Beauty and the beast: Mechanisms of sexual selection F. H. Jessop. J. 38.. G. E.. D. Adaptive significance of female attractiveness: Role of Human Behavior. Psychological Bulletin. (2002). S.. S. 276. male mate retention. W. K. (2002). 30. Jones. & Singh. & Roberts. 1213–1217. A... 65. 35.. behavior. J. activate neural reward centers in men. A. Caryl. Feinberg... Fluctuating asymmetry as a pos.. Proceedings of the Royal Society of London. 29. L. Evolutionary Psychology.. 5. L. Sex differences in morpho.. T.. 26 –34. 57. varies across the menstrual cycle.. (2010).. Lever.. A. T. J. 273.... D. G.. Mating context and menstrual cycle phase affect prefer a low waist-to-hip ratio.. Perrett. L. L. G. C. 157–175. D. M. (2002). Jokela. S.. A. Langlois.. Interna- mating intentions and mating success. 63... (1993). Journal predicts differential ovulatory shifts in female extra-pair attraction and of Nonverbal Behavior. (2008). 612– 631. Maisel. K. Tybur. (2009). Yee. Kruger. R. Jr.. C. resources.. & Gallup. M.. 29. M. Evolution Trait rankings and peer evaluations. J. (2008). Calcified Ramsey. 256 –267. dietary require. Psychological Topics. F. E. M.. Male financial consumption is associated with higher Rigby. (2004). & Haselton. M. Tissue.. Proceedings of the Royal Society Human Biology. The evolutionary psychology of facial beauty. D. 2365–2371. Human Behavior and Evolution. The face of success: Inferences from Lassek. 173–178. & Kobyliansky.. A. D. & Gaulin. ences in romantic kissing among college students: An evolutionary & Edwards.. Heuthe. S.. Rowland. G. 19. waist-to-hip ratio.. & Gaulin. F. Ethology.. S. R. and native immunity. D.. 31. N. Sanders. Body image satisfaction in heterosexual. J. Evolution and Human Behavior. G.. G.. I. G. MHC-herterozygosity and human facial & Smoot. & Singh. 287. (2006). 61. Hughes. 5. Breast asymmetry and phenotypic quality in women.. M. Evolution and Human Behavior. S. Men’s faces convey information health. G. & Griffin.. I. S. Law-Smith. gay. & Dixson. A. R. D. P. J. Society B. Proceedings of the Royal to oestrogen levels in women. 603– tional Journal of Epidemiology. 295–307. & Gallup.. Commentary: Counterpoint to Campos et al. Jr. 199 –226. . 271. J. 2.. & Ambady. Saxton. C.. 117–119. Lee.. 27. 469 – 479. Facial appearance is a cue assessing physical strength from the voice.. J. Harrison. M. Cloos. 464 – 473. Carter. M. 270. 28. Colleqium Antropolo- Rhodes. (2006). (2005). perspective.. & Bodmer. 322– togeny of mate choice. intelligence. 223–236. G. M. B-Biological Sciences. J. D. H. J. androgen receptor in human osteoblastic cell populations. Tooby.. Hoss. 213–216. & Leinster. W. A.. F. & Gallup. M. Dixson.. D. Whitehouse. Proceedings of the Royal Society of London. A.. N. L. Jasienska. 30. Ellison. tive. Miller. & Thune. von Rueden. A. 23. 18. M. J... Rubenstein. PLoS ONE. S. A... Pinker.. Evolution and Human Behavior. (2008). Proceedings of the Royal Society B.. and body weight across the menstrual cycle. potential. 112.. 32. A. Maxims or myths of beauty? A meta-analytic and attractiveness. I. D. Symmetry and human facial attractiveness. (2008). K. 31. M.. on tip earnings by lap dancers: Economic evidence for human estrus. M... 29. Evolution and Human Behavior. 612. & Gallup.. B. 126. Review of Psychology. Female body odour is a potential cue Evolution and Human Behavior. Mortensen. G. 375–381. (2004).1098/rspb. (2008). (2006). 79 – 80. and Human Behavior. P. Moore. Singh. & logical predictors of sexual behavior: Shoulder to hip and waist to hip Ghavami. Cosmides. C. & Bronstad. Frankenhuis. Hughes. The blank slate: The modern denial of human nature. and ratios. I. A. 20. Evolution and Human Be- 182–186. Evolutionary Psychology.. Sznycer. 331–350. S. & Gurven. Kalakanis. & Gurven. M. Morgan. C. 1179 –1185. (2009). & Ziegler. D. Little. Karremans. A naked ape would have fewer parasites.. D. E. fighting ability from the body and face. 173–180.. R. D. G. J. Vocal and facial Lassek W. Roberts. A. N. The sound of Pillsworth. adolescents and adults: The on- ments. Psycho- ability: Is gluteofemoral fat a privileged store of neurodevelopmental logical Science. (1991). & Jordan.. N. (2001). M. R. G. A. (2006). T. Rule. Livshits. 6. Bryant. Peplau. B.0769 135–140. E. B. (2006). (2007). J. A. A. S. Tooby.. Evolution and Human Behavior...

.. Received December 30. (1999). J. 27. (2006). 131–144. R.. 313–325.. (2007). (2006).. Facial sexual dimorphism... origins. Nunez. The evolutionary biology of human female sexuality. R. Murphy. Jones. The female nude in Rubens: Androgenousness. Evolution and Human Behavior. A. breast size. 545–565... M. 24. Tovee. D. A. Imagination. 2010. 2009 Tovee. body fat distribution in females impairs the pregnancy rate of in-vitro Swami. J.. J. L. I. different culture. R. 27. V. ences in population versus media body sizes: A comparison over four Koppenaal. W. M. J. T. D. M. W. & Furnham. Beauty is in the adaptations of the beholder. Buss. DOI: 10. M. A. V. (2006). 2057–2060.. Gray. Body weight ideals and body dissatis- change in body measures: Trend analysis. S.. C. M. 3. 105-112). Zaadstra. Swami.). Hall.. M.. & Furnham. body mass index. A. British Medical Journal. 40... Thornhill. 85. pp. & Cohen.... curvaceousness. NJ: Wiley. D. Archives of Sexual Behavior. or oligomenorrhea. Waas. Human Reproduction. S... Waldenstrom.. M. F.. & Rhodes.. for women’s profile waist-to-hip ratio.. British Journal of Psychology. Pinkerson (Eds. Zebrowitz. Allik. Swami.. Physical attractiveness in adaptationist perspec. Chicago: University of Chicago Press. Vol. otropin concentration. M. (1997). “The “I’s” Have It: A Framework for Serious Educational Game Design” by Leonard A. and susceptibility to disease in men and women.. R. and Personality. and Metabolism. J. faction in 26 countries across 10 world regions: Results of the Interna- 1447–1448. A. and ethnic group in Weeden. Symons. Furnham. Alcalay. (2009). Evolution and Human Behavior.. Facial attractiveness. . Insulin. B. Mason. androgen. Frederick. Fat and female 80 –118). V. B. A. D. Emery. Abramson & S. . 297–304. M. Swami. Te Velde. W. Hirasing.. (2002). Kaptein. S. Rossner.. M. H. L.. (2003). & Schoemaker. T. 28. In P. In D. 238 –249. & Gangestad. Henderson. Van Hoof. No. A. A. (pp... and waist-to-hip ratio in the first Sugiyama. L. M. Hoboken. Accepted June 5. the final acceptance date was incorrect. Seidell. New York: Oxford University. 131. V. (1997).. (2008). R. C. J. J. 1394 –1400. P. McCluskey.. . Shapely centrefolds? Temporal .. (1993)... (1995). Einon. J. Personality and Social Psychology Bulletin. Voracek. Men’s preference fertilization-embryo transfer. & Fisher. & Karbaat. 452– 460. L. A. P. sexual culture (pp. L. S.. Evolution and Human Behavior.. B. Voorhorst. years after menarche in girls with regular menstrual cycle. western societies: A review. 35. Van Noord. & Zivian. F.. & Gangestad. L.. B. 306. Facial attractiveness in men provides clues to semen Changing perceptions of attractiveness as observers are exposed to a quality. Gender differ. M. (2000). 325. Success is all in the measures: 309 –325. M. 199 –207. tional Body Project I. Revision received June 5. M. Psychological Bulletin. 2010. A.. L. Physical attractiveness and health in Britain and South Africa. R. and starring frequencies in adult Disconfirmatory evidence of the waist-to-hip ratio hypothesis of female media actresses. Supermodels: Stick insects or hourglasses? Lancet. D. 2010 䡲 Correction to Annetta (2010) In the article. 1474 –1475. G. Trends in British Medical Journal. R. Sex Roles. Medina. 484 – 487. & Mangalparsad. Anderson. C.. H. & Gangestad. Nunez.250 GALLUP AND FREDERICK Soler. R. J. M. (1999). The final acceptance date should be January 15. 14. 2010 350. The handbook of evolutionary psychology. J. J. E. (2005). K. 100. Sexual nature. & Fisher. J. and gonad- decades.. P.. irregular tive. physical attractiveness. fecundity: Prospective study of body fat distribution in conception rates. 443– 456. Aavik. J. menstrual cycles. Sancho. 36. Annetta (Review of General Psychology. E. N... Voracek. Vrieswijk. Thornhill. Tovee. (2005). Personality and Social Psychology Bulletin. V. Nunez. M. . S. D.. Cognition. 12. 26. Gutierrez. 635– 653. Looking smart and looking good: Facial cues to intelligence and their developmental stability. (2002).1037/a0020505 . Habbema. Spitzer. & Sabini. . An adroid 139 –147. S. M. Zivcic-Becirevic. Thornhill. & Hellberg. E. 2. A. Cognitive Sciences. Journal of Clinical Endocrinology 292–343). J. (2010). D.