The Biology of Beauty

Nancy Etcoff, Survival of the Prettiest: The Science of Beauty (New York: Anchor Books 2000) 

Beauty is in the eye of the beholder.  

This much is true by very definition. After all, the Oxford English Dictionary defines beauty as: 

A combination of qualities, such as shape, colour, or form, that pleases the aesthetic senses, especially the sight’. 

If beauty is in the eye of the beholder, then the ‘eye of the beholder’ has been shaped by a process of natural, and sexual, selection to find certain things beautful — and, if beauty is in the ‘eye of the beholder’, then sexiness is located in a different part of the male anatomy but similarly subjective

Thus, beauty is defined as that which is pleasing to an external observer. It therefore presupposes the existence of an external observer, separate from the person or thing that is credited with beauty, from whose perspective the thing or individual is credited with beauty.[1]

Moreover, perceptions of beauty do indeed differ.  

To some extent, preferences differ between individuals, and between different races and cultures. More obviously, and to a far greater extent, they also differ as between species.  

Thus, a male chimpanzee would presumably consider a female chimpanzee as more beautiful than a woman. The average human male, however, would likely disagree – though it might depend on the woman. 

As William James wrote in 1890: 

To the lion it is the lioness which is made to be loved; to the bear, the she-bear. To the broody hen the notion would probably seem monstrous that there should be a creature in the world to whom a nestful of eggs was not the utterly fascinating and precious and never-to-be-too-much-sat-upon object which it is to her” (Principles of Psychology (vol 2): p387). 

Beauty is therefore not an intrinsic property of the person or object that is described as beautiful, but rather a quality attributed to that person or object by a third-party in accordance with their own subjective tastes. 

However, if beauty is then indeed a subjective assessment, that does not mean it is an entirely arbitrary one. 

On the contrary, if beauty is indeed in the ‘eye of the beholder’ then it must be remembered that the ‘eye of the beholder’—and, more importantly, the brain to which that eye is attached—has been shaped by a process of both natural and sexual selection

In other words, we have evolved to find some things beautiful, and others ugly, because doing so enhanced the reproductive success of our ancestors. 

Thus, just as we have evolved to find the sight of excrement, blood and disease disgusting, because each were potential sources of infection, and the sight of snakes, lions and spiders fear-inducing, because each likewise represented a potential threat to our survival when encountered in the ancestral environment in which we evolved, so we have evolved to find the sight of certain things pleasing on the eye. 

Of course, not only people can be beautiful. Landscapes, skylines, works of art, flowers and birds can all be described as ‘beautiful’. 

Just as we have evolved to find individuals of the opposite sex attractive for reasons of reproduction, so these other aspects of aesthetic preference may also have been shaped by natural selection. 

Thus, some research has suggested that our perception of certain landscapes as beautiful may reflect psychological adaptations that evolved in the context of habitat selection (Orians & Heerwagen 1992).  

However, Nancy Etcoff does not discuss such research. Instead, in ‘Survival of the Prettiest’, her focus is almost exclusively on what we might term ‘sexual beauty’. 

Yet, if beauty is indeed in the ‘in the eye of the beholder’, then sexiness is surely located in a different part of the male anatomy, but equally subjective in nature. 

Indeed, as I shall discuss below, even in the context of mate preferences, ‘sexiness’ and ‘beauty’ are hardly synonyms. As an illustration, Etcoff herself quotes that infamous but occasionally insightful pseudo-scientist and all-round charlatan, Sigmund Freud, whom she quotes as observing:  

The genitals themselves, the sight of which is always exciting, are nevertheless hardly ever judged to be beautiful; the quality of beauty seems, instead, to attach to certain secondary sexual characters” (p19: quoted from Civilization and its Discontents). 

Empirical Research 

Of the many books that have been written about the evolutionary psychology of sexual attraction (and I say this as someone who has read, at one time or another, a good number of them), a common complaint is that they are full of untested, or even untestable, speculation – i.e. what that other infamous scientific charlatan Stephen Jay Gould famously referred to as just so stories

This is not a criticism that could ever be levelled at Nancy Etcoff’s ‘Survival of the Prettiest’. On the contrary, as befits Etcoff’s background as a working scientist (not a mere journalist or popularizer), it is, from start to finish, it is full of data from published studies, demonstrating, among other things, the correlates of physical attractiveness, as well as the real-world payoffs associated with physical attractiveness (what is sometimes popularly referred to as ‘lookism’). 

Indeed, in contrast to other scientific works dealing with a similar subject-matter, one of my main criticisms of this otherwise excellent work would be that, while rich in data, it is actually somewhat deficient in theory. 

Youthfulness, Fertility, Reproductive Value and Attractiveness 

A good example of this deficiency in theory is provided by Etcoff’s discussion of the relationship between age and attractiveness. Thus, one of the main and recurrent themes of ‘Survival of the Prettiest’ is that, among women, sexual attractiveness is consistently associated with indicators of youth. Thus, she writes: 

Physical beauty is like athletic skill: it peaks young. Extreme beauty is rare and almost always found, if at all, in people before they reach the age of thirty-five” (p63). 

Yet Etcoff addresses only briefly the question of why it is that youthful women or girls are perceived as more attractive – or, to put the matter more accurately, why it is that males are sexually and romantically attracted to females of youthful appearance. 

Etcoff’s answer is: fertility

Female fertility rapidly declines with age, before ceasing altogether with menopause

There is, therefore, in Darwinian terms, no benefit in a male being sexually attracted to an older, post-menopausal female, since any mating effort expended would be wasted, as any resulting sexual union could not produce offspring. 

As for the menopause itself, this, Etcoff speculates, citing scientific polymath, popularizer and part-time sociobiologist Jared Diamond, evolved because human offspring enjoy a long period of helpless dependence on their mother, without whom they cannot survive. 

Therefore, after a certain age, it pays women to focus on caring for existing offspring, or even grandchildren, rather than producing new offspring whom, given their own mortality, they will likely not be around long enough to raise to maturity (p73).[2]

This theory has sometimes been termed the grandmother hypothesis.

However, the decline in female fertility with age is perhaps not sufficient to explain the male preference for youth. 

After all, women’s fertility is said to peak in their early- to mid-twenties.[3]

However, men’s (and boy’s) sexual interest, if anything, seems to peak in respect of females, if anything, somewhat younger, namely in their late-teens (Kenrick & Keefe 1992). 

To explain this, Douglas Kenrick and Richard Keefe propose, following a suggestion of Donald Symons, that this is because girls at this age, while less fertile, have higher reproductive value, a concept drawn from ecology, population genetics and demography, which refers to an individual’s expected future reproductive output given their current age (Kenrick & Keefe 1992). 

Reproductive value in human females (and in males too) peaks just after puberty, when a girl first becomes capable of bearing offspring. 

Before then, there is always the risk she will die before reaching sexual maturity; after, her reproductive value declines with each passing year as she approaches menopause. 

Thus, Kenrick and Keefe, like Symons before them, argue that, since most human reproduction occurs within long-term pair-bonds, it is to the evolutionary advantage of males to form long-term pair-bonds with females of maximal reproductive value (i.e. mid to late teens), so that, by so doing, they can monopolize the entirety of that woman’s reproductive output over the coming years. 

Yet the closest Etcoff gets to discussing this is a single sentence where she writes: 

Men often prefer the physical signs of a woman below peak fertility (under age twenty). Its like signing a contract a year before you want to start the job” (p72). 

Yet the theme of indicators of youth being a correlate of female attractiveness is a major theme of her book. 

Thus, Etcoff reports that, in a survey of traditional cultures: 

The highest frequency of brides was in the twelve to fifteen years of age category… Girls at this age are preternaturally beautiful” (p57). 

It is perhaps true that “girls at this age are preternaturally beautiful” – and Etcoff, being female, can perhaps even get away with saying this without being accused of being a pervert or ‘paedophile’ for even suggesting such a thing. 

Nevertheless, this age “twelve to fifteen” seems rather younger than most men’s, and even most teenage boys, ideal sexual partners, at least in western societies. 

Thus, for example, Kenrick and Keefe inferred from their data that around eighteen was the preferred age of sexual partner for most males, even those somewhat younger than this themselves.[4]

Of course, in primitive, non-western cultures, women may lose their looks more quickly, due to inferior health and nutrition, the relative unavailability of beauty treatments and because they usually undergo repeated childbirth from puberty onward, which takes a toll on their health and bodies. 

On the other hand, however, obesity is more prevalent in the West, decreases sexual attractiveness and increases with age. 

Moreover, girls in the west now reach puberty somewhat earlier than in previous centuries, and perhaps earlier than in the developing world, probably due to improved nutrition and health. This suggests that females develop secondary sexual characteristics (e.g. large hips and breasts) that are perceived as attractive because they are indicators of fertility, and hence come to be attractive to males, rather earlier than in premodern or primitive cultures. 

Perhaps Etcoff is right that girls “in the twelve to fifteen years of age category… are preternaturally beautiful” – though this is surely an overgeneralization and does not apply to every girl of this age. 

However, if ‘beauty’ peaks very early, I suspect ‘sexiness’ peaks rather later, perhaps late-teens into early or even mid-twenties. 

Thus, the latter is dependent on secondary sexual characteristics that develop only in late-puberty, namely larger breasts, buttocks and hips

Thus, Etcoff reports, rather disturbingly, that: 

When [the] facial proportions [of magazine cover girls] are fed into a computer, it guesstimates their age to be between six and seven years of age” (p151; citing Jones 1995). 

But, of course, as Etcoff is at pains to emphasize in the next sentence, the women pictured do not actually look like they are of this age, either in their faces let alone their bodies. 

Instead, she cites Douglas Jones, the author of the study upon which this claim is based, as arguing that the neural network’s estimate of their age can be explained by their display of “supernormal stimuli”, which she defines as “attractive features… exaggerated beyond proportions normally found in nature (at least in adults)” (p151). 

Yet much the same could be said of the unrealistically large, surgically-enhanced breasts favored among, for example, glamour models. These abnormally large breasts are likewise an example of “supernormal stimuli” that may never be found naturally, as suggested by Doyle & Pazhoohi (2012)

But large breasts are indicators of sexual maturity that are rarely present in girls before their late-teens. 

In other words, if the beauty of girls’ faces peaks at a very young age, the sexiness of their bodies peaks rather later. 

Perhaps this distinction between what we can term ‘beauty’ and ‘sexiness’ can be made sense of in terms of a distinction between what David Buss calls short-term and long-term mating strategies

Thus, if fertility peaks in the mid-twenties, then, in respect of short-term mating (i.e. one-night stands, casual sex, hook-ups and other one-off sexual encounters), men should presumably prefer partners of a somewhat greater age than their preferences in respect of long-term partners – i.e. of maximal fertility rather than maximum reproductive value – since in the case of short-term mating strategies there is no question of monopolizing the woman or girl’s long-term future reproductive output. 

In contrast, cues of beauty, as evinced by relatively younger females, might trigger a greater willingness for males to invest in a long-term relationship. 

This ironically suggests, contrary to contemporary popular perception, males’ sexual or romantic interest in respect of relatively younger women and girls (i.e. those still in their teens) would tend to reflect more ‘honourable intentions’ (i.e. more focussed on marriage or a long-term relationship rather than mere casual sex) than does their interest in older women. 

However, as far as I am aware, no study has ever demonstrated differences in men’s preferences regarding the preferred age-range of their casual sex partners as compared to their preferences in respect of longer-term partners. This is perhaps because, since commitment-free casual sex is almost invariably a win-win situation for men, and most men’s opportunities in this arena likely to be few and far between, there has been little selection acting on men to discriminate at all in respect of short-term partners. 

Are There Sex Differences in Sexiness? 

Another major theme of ‘Survival of the Prettiest’ is that the payoffs for good-looks are greater for women than for men. 

Beauty is most obviously advantageous in a mating context. But women convert this advantage into an economic one through marriage. Thus, Etcoff reports: 

The best-looking girls in high school are more than ten times as likely to get married as the least good-looking. Better looking girls tend to ‘marry up’, that is, marry men with more education and income then they have” (p65; see also Udry & Eckland 1984; Hamermesh & Biddle 1994). 

However, there is no such advantage accruing to better-looking male students. 

On the hand, according to Catherine Hakim, in her book Erotic Capital: The Power of Attraction in the Boardroom and the Bedroom (which I have reviewed here, here and here) in the workplace, the wage premium associated with being better looking is actually, perhaps surprisingly, greater for men than for women. 

For Hakim herself: 

This is clear evidence of sex discrimination… as all studies show women score higher than men on attractiveness” (Money, Honey: p246). 

However, as I explain in my review of her book, the better view is that, since beauty opens up so many other avenues to social advancement for women, notably through marriage, relatively more beautiful women corresponding reduce their work-effort in the workplace since they have need of pursuing social advancement through their careers when they can far more easily achieve it through marriage. 

After all, by bother to earn money when you can simply marry it instead. 

According to Etcoff, there is only one sphere where being more beautiful is actually disadvantageous for women, namely in respect of same-sex friendships: 

Good looking women in particular encounter trouble with other women. They are less liked by other women, even other good-looking women” (p50; citing Krebs & Adinolfy 1975). 

She does not speculate as to why this is so. An obvious explanation is envy and dislike of the sexual competition that beautiful women represent. 

However, an alternative explanation is perhaps that beautiful women do indeed come to have less likeable personalities. Perhaps, having grown used to receiving preferential treatment from and being fawned over by men, beautiful women become entitled and spoilt. 

Men might overlook these flaws on account of their looks, but, other women, immune to their charms, may be a different story altogether.[5]

All this, of course, raises the question as to why the payoffs for good looks are so much greater for women than for men? 

Etcoff does not address this, but, from a Darwinian perspective, it is actually something of a paradox which I have discussed previously

After all, among other species, it is males for whom beauty affords a greater payoff in terms of the ultimate currency of natural selection – i.e. reproductive success. 

It is therefore male birds who usually evolve more beautiful plumages, while females of the same species are often quite drab, the classic example being the peacock and peahen

The ultimate evolutionary explanation for this pattern is called Bateman’s principle, later formalized by Robert Trivers as differential parental investment theory (Bateman 1948; Trivers 1972). 

The basis of this theory is this: Females must make a greater minimal investment in offspring in order to successfully reproduce. For example, among humans, females must commit themselves to nine months pregnancy, plus breastfeeding, whereas a male must contribute, at minimum, only a single ejaculate. Females therefore represent the limiting factor in mammalian reproduction for access to whom males compete. 

One way in which they compete is by display (e.g. lekking). Hence the evolution of the elaborate tail of the peacock

Yet, among humans, it is females who seem more concerned with using their beauty to attract mates. 

Of course, women use makeup and clothing to attract men rather than growing or evolving long tails. 

However, behavior is no less subject to selection than morphology, so the paradox remains.[6]

Indeed, the most promising example of a morphological trait in humans that may have evolved primarily for attracting members of the opposite sex (i.e. a ‘peacock’s tail’) is, again, a female trait – namely, breasts

This is, of course, the argument that was, to my knowledge, first developed by ethologist Desmond Morris in his book The Naked Ape, which I have reviewed here, and which I discuss in greater depth here

As Etcoff herself writes: 

Female breasts are like no others in the mammalian world. Humans are the only mammals who develop rounded breasts at puberty and keep them whether or not they are producing milk… In humans, breast size is not related to the amount or quality of milk that the breast produces” (p187).[7]

Instead, human breasts are, save during pregnancy and lactation, composed predominantly of, not milk, but fat. 

This is in stark contrast to the situation among other mammals, who develop breasts only during pregnancy. 

Breasts are not sex symbols to other mammals, anything but, since they indicate a pregnant or lactating and infertile female. To chimps, gorillas and orangutans, breasts are sexual turn-offs” (p187). 

Why then does sexual selection seem, at least on this evidence, to have acted more strongly on women than men? 

Richard Dawkins, in The Selfish Gene (which I have reviewed here), was among the first to allude to this anomaly, lamenting: 

What has happened in modern western man? Has the male really become the sought-after sex, the one that is in demand, the sex that can afford to be choosy? If so, why?” (The Selfish Gene: p165). 

Yet this is surely not the case with regard to casual sex (i.e. hook-ups and one-night stands). Here, it is very much men who ardently pursue and women who are sought after. 

For example, in one study at a University campus, 72% of male students agreed to go to bed with a female stranger who propositioned them to this effect, yet not a single one of the 96 females approached agreed to the same request from a male stranger (Clark and Hatfield 1989). 

(What percentage of the students sued the university for sexual harassment was not revealed.) 

Indeed, patterns of everything from prostitution to pornography consumption confirm this – see The Evolution of Human Sexuality (which I have reviewed here). 

Yet humans are unusual among mammals in also forming long-term pair-bonds where male parental investment is the norm. Here, men have every incentive to be as selective as females in their choice of partner. 

In particular, in Western societies practising what Richard Alexander called socially-imposed monogamy (i.e. where there exist large differentials in male resource holdings, but polygynous marriage is unlawful) competition among women for exclusive rights to resource-abundant alpha males may be intense (Gaulin and Boser 1990). 

In short, the advantage to a woman in becoming the sole wife of a multi-millionaire is substantial. 

This, then, may explain the unusual intensity of sexual selection among human females. 

Why, though, is there not evidence of similar sexual selection operating among males? 

Perhaps the answer is that, since, in most cultures, arranged marriages are the norm, female choice actually played little role in human evolution. 

As Darwin himself observed in The Descent of Man as an explanation as to why intersexual selection seems, unlike among most other species, to operated more strongly on human females than on men:

Man is more powerful in body and mind than woman, and in the savage state he keeps her in a far more abject state of bondage than does the male of any other animal; therefore it is not surprising that he should have gained the power of selection” (The Descent of Man).

Instead, male mating success may have depended less upon what Darwin called intersexual selection and more upon intrasexual selection – i.e. less upon female choice and more upon male-male fighting ability (see Puts 2010). 

Male Attractiveness and Fighting Ability 

Paradoxically, this is reflected even in the very traits that women find attractive in men. 

Thus, although Etcoff’s book is titled ‘The Evolution of Prettiness’, and ‘prettiness’ is usually an adjective applied to women, and, when applied to men, is—perhaps tellingly—rarely a complement, Etcoff does discuss male attractiveness too.  

However, Etcoff acknowledges that male attractiveness is a more complex matter than female attractiveness: 

We have a clearer idea of what is going on with female beauty. A handsome male turns out to be a bit harder to describe, although people reach consensus almost as easily when they see him” (p155).[8]

Yet what is notable about the factors that Etcoff describes as attractive among men is that they all seem to be related to fighting ability. 

This is most obviously true of height (p172-176) and muscularity (p176-80). 

Indeed, in a section titled “No Pecs, No Sex”, though she focuses on the role of pectoral muscles in determining attractiveness, Etcoff nevertheless acknowledges: 

Pectoral muscles are the human male’s antlers. Their weapons of war” (p177). 

Thus, height and muscularity have obvious functional utility. 

This in stark contrast to traits such as the peacock’s tail, which are often a positive handicap to their owner. Indeed, one influential theory of sexual selection contends that it is precisely because they represent a handicap that they have evolved as a sexually-selected fitness indicator, because only a genetically superior male is capable of bearing the handicap of such an unwieldy ornament, and hence possession of such a handicap is paradoxically an honest signal of health. 

Yet, if men’s bodies have evolved more for fighting than attracting mates, the same is perhaps less obviously true of their faces. 

Thus, anthropologist David Puts proposes: 

Even [male] facial structure may be designed for fighting: heavy brow ridges protect eyes from blows, and robust mandibles lessen the risk of catastrophic jaw fractures” (Puts 2010: p168). 

Indeed, looking at the facial features of a highly dominant, masculine male face, like that of Mike Tyson, for example, one gets the distinct impression that, if you were foolish enough to try punching it, it would likely do more damage to your hand than to his face. 

Thus, if some faces are, as cliché contends, highly ‘punchable’, then others are presumably at the opposite end of this spectrum. 

This also explains some male secondary sexual characteristics that otherwise seem anomalous, for example, beards. These have actually been found in some studies “to decrease attractiveness to women, yet have strong positive effects on men’s appearance of dominance” (Puts 2010: p166). 

David Puts concludes: 

Men’s traits look designed to make men appear threatening, or enable them to inflict real harm. Men’s beards and deep voices seem designed specifically to increase apparent size and dominance” (Puts 2010: p168). 

Interestingly, Etcoff herself anticipates this theory, writing: 

Beautiful ornaments [in males] develop not just to charm the opposite sex with bright colors and lovely songs, but to intimidate rivals and win the intrasex competition—think of huge antlers. When evolutionists talk about the beauty of human males, they often refer more to their weapons of war than their charms, to their antlers rather than their bright colors. In other words, male beauty is thought to have evolved at least partly in response to male appraisal” (p74) 

Of course, these same traits are also often attractive to females. 

After all, if a tall muscular man has higher reproductive success because he is better at fighting, then it pays women to preferentially mate with tall, muscular men so that their male offspring will inherit these traits and hence themselves have high reproductive success, helping the spread the women’s own genes by piggybacking on the superior male’s genes.  

This is a version of sexy son theory

In addition, males with fighting prowess are better able to protect and provision their mates. 

However, this attractiveness to females is obviously secondary to the primary role in male-male fighting. 

Moreover, Etcoff admits, highly masculine faces are not always attractive. 

Thus, unlike the “supernormal” or “hyperfeminine” female faces that men find most attractive in women, women rated “hypermasculine” faces as less attractive (p158). This, she speculates, is because they are perceived as overaggressive and unlikely to invest in offspring

As to whether such men are indeed less willing to invest in offspring, this Etcoff does not discuss and there appears to be little evidence on the topic. But the association of testosterone with both physiological and psychological masculinization suggests that the hypothesis is at least plausible

Etcoff concludes: 

For men, the trick is to look masculine but not exaggeratedly masculine, which results in a ‘Neanderthal’ look suggesting coldness or cruelty” (p159). 

Examples of males with perhaps overly masculine faces are perhaps certain boxers, who tend to have highly masculine facial morphology (e.g. heavy brow ridges, deep set eyes, wide muscular jaws), but are rarely described as handsome. 

For example, I doubt anyone would ever call Mike Tyson handsome. But, then, no one would ever call him exactly ugly either – at least not to his face. 

An extreme example might be the Russian boxer Nikolai Valuev, whose extreme neanderthal-like physiognomy was much remarked on. 

Another example that sprung to mind was the footballer Wayne Rooney (also, perhaps not uncoincidentally, said to have been a talented boxer) who, when he first became famous, was immediately tagged by the newspapers, media and comedians as ugly despite – or indeed because of – his highly masculine, indeed thuggish, facial physiognomy

Likewise, Etcoff reports that large eyes are perceived as attractive in men, but these are a neotenous trait, associated with both immature infants and indeed with female beauty (p158). 

This odd finding Etcoff attributes to the fact that large eyes, as an infantile trait, evoke women’s nurturance, a trait that evolved in the context of parental investment rather than mate choice

Yet this is contrary to the general principle in evolutionary psychology of modularity of mind and the domain specificity of psychological adaptations, whereby it is assumed that that psychological adaptations for mate choice and for parental investment represent domain-specific modules with little or no overlap. 

Clearly, for psychological adaptations in one of these domains to be applied in the other would result in highly maladaptive behaviours, such as sexual attraction to infants and to your own close biological relatives.[9]

In addition to being more complex and less easy to make sense of than female beauty, male physical attractiveness is also of less importance in determining female mate choice than is female beauty in male mate choice

In particular, she acknowledges that male status often trumps handsomeness. Thus, she quotes a delightfully cynical, not especially poetic, line from the ancient Roman poet Ovid, who wrote: 

Girls praise a poem, but go for expensive presents. Any illiterate oaf can catch their eye, provided he’s rich” (quoted: p75). 

A perhaps more memorable formulation of the same idea is quoted on the same page from a less illustrious source, namely boxing promoter, numbers racketeer and convicted killer Don King, on a subject I have already discussed, namely the handsomeness (or not) of Mike Tyson, King remarking: 

Any man with forty two million looks exactly like Clark Gable” (quoted: p75). 

Endnotes

[1] I perhaps belabor this rather obvious point only because one prominent evolutionary psychologist, Satoshi Kanazawa, argues that, since many aspects of beauty standards are cross-culturally universal, beauty standards are not ‘in the eye of the beholder’. I agree with Kanazawa on the substantive issue that beauty standards are indeed mostly cross-culturally universal among humans (albeit not entirely so). However, I nevertheless argue, perhaps somewhat pedantically, that beauty remains strictly in the ‘eye of the beholder’, but it is simply that the ‘eye of the beholder’ (and the brain to which is attached) has been shaped by a process of natural selection so as to make different humans share the same beauty standards. 

[2] While Jared Diamond has indeed made many original contributions to many fields, this idea does not in fact originate with him, even though Etcoff oddly cites him as a source. Indeed, as far as I am aware, it is even especially associated with Diamond. Instead, it may actually originatea by another, lesser known, but arguably even more brilliant evolutionary biologist, namely George C Williams (Williams 1957). 

[3] Actually, pregnancy rates peak surprisingly young, perhaps even disturbingly young, with girls in their mid- to late-teens being most likely to become pregnant from any single act of sexual intercourse, all else being equal. However, the high pregnancy rates of teenage girls are said to be partially offset by their greater risk of birth complications. Therefore, female fertility is said to peak among women in their early- to mid-twenties.

[4] This Kenrick and Keefe inferred from, among other evidence, an analysis of lonely hearts advertisements, wherein, although the age of the female sexual/romantic partner sought was related to the advertised age of the man placing the ad (which Kenrick and Keefe inferred was a reflection of the fact that their own age delimited the age-range of the sexual partners whom they would be able to attract, and whom it would be socially acceptable for them to seek out) nevertheless the older the man, the greater the age-difference he sought in a partner. In addition, they reported evidence of surveys suggesting that, in contrast to older men, younger teenage boys, in an ideal world, actually preferred somewhat older sexual partners, suggesting that the ideal age of sexual partner for males of any age was around eighteen years of age (Kenrick & Keefe 1992).

[5] Etcoff also does not discuss whether the same is true of exceptionally handsome men – i.e. do exceptionally handsome men, like beautiful women, also have problems maintaining same-sex friendships. I suspect that this is not so, since male status and self-esteem is not usually based on handsomeness as such – though it may be based on things related to handsomeness, such as height, athleticism, earnings, and perceived ‘success with women’. Interestingly, however, French novelist Michel Houellebecq argues otherwise in his novel, Whatever, in which, after describing the jealousy of one of the main characters, the short ugly Raphael Tisserand, towards an particularly handsome male colleague, writes: 

Exceptionally beautiful people are often modest, gentle, affable, considerate. They have great difficulty in making friends, at least among men. They’re forced to make a constant effort to try and make you forget their superiority, be it ever so little” (Whatever: p63) 

[6] Thus, in other non-human species, behaviour is often subject to sexual selection, in, for example, mating displays, or the remarkable, elaborate and often beautiful, but non-functional, nests built by male bowerbirds, which Geoffrey Miller sees as analogous to human art. 

[7] An alternative theory for the evolution of human breasts is that they evolved, not as a sexually selected ornament, but rather as a storehouse of nutrients, analogous to the camel’s humps, upon which women can draw during pregnancy. On this view, the sexual dimorphism of their presentation (i.e. the fact that, although men do have breasts, they are usually much less developed than those of women) reflects, not sexual selection, but rather the calaric demands of pregnancy. 
However, these two alternative hypotheses are not mutually incompatible. On the contrary, they may be mutually reinforcing. Thus, Etcoff herself mentions the possibility that breasts are attractive precisely because: 

Breasts honestly advertise the presence of fat reserves needed to sustain a pregnancy” (p178.) 

On this view, men see fatty breasts as attractive in a sex partner precisely because only women with sufficient reserves of fat to grow large breasts are likely to be capable of successfully gestating an infant for nine months. 

[8] Personally, as a heterosexual male, I have always had difficulty recognizing ‘handsomeness’ in men, and I found this part of Etcoff’s book especially interesting for this reason. In my defence, this is, I suspect, partly because many rich and famous male celebrities are celebrated as ‘sex symbols’ and described as ‘handsome’ even though their status as ‘sex symbols’ owes more to the fact they are rich and famous than their actual looks. Thus, male celebrities sometimes become sex symbols despite their looks, rather than because of them. Many famous rock stars, for example, are not especially handsome but nevertheless succeed in becoming highly promiscuous and much sought after by women and girls as sexual and romantic partners. In contrast, men did not suddenly start idealizing fat or physically unattractive female celebrities as sexy and beautiful simply because they are rich famous celebrities.
Add to this the fact that much of what passes for good looks in both sexes is, ironically, normalness – i.e. a lack of abnormalities and averageness – and identifying which men women consider ‘handsome’ had, before reading Etcoff’s book, always escaped me.
However, Etcoff, for her part, might well call me deluded. Men, she reports, only claim they cannot tell which men are handsome and which are not, perhaps to avoid being accused of homosexuality

Although men think they cannot judge another man’s beauty, the agree among themselves and with women about which men are the handsomest” (p138). 

Nevertheless, there is indeed some evidence that judging male handsomeness is not as clear cut as Etcoff seems to suggests. Thus, it has been found that, not only do men claim to have difficulty telling handsome men from ugly men, but also women themselves are more likely to disagree among themselves about the physical attractiveness of members of the opposite sex as compared to men (Wood & Brumbaugh 2009Wake Forest University 2009). 
Indeed, not only do women not always agree with one another regarding the attractiveness of men, sometimes they can’t even agree with themselves. Thus, Etcoff reports: 

A woman makes her evaluations of men more slowly, and if another woman offers a different opinion, she may change her mind” (p76). 

This indecisiveness, for Etcoff, actually makes good evolutionary sense:

If women take a second look, compare notes with other women, or change their minds after more thought, it is not out of indecisiveness but out of wisdom. Mate choice is not just about fertility—most men are fertile most or all of their lives—but about finding a helpmate to bring up the baby” (p77). 

Another possible reason why women may consult other women as to whether a given man is attractive or not is sexy son theory
On this view, it pays for women to mate with men who are perceived as attractive by other women because then any offspring whom they bear by these men will likely inherit the very traits that made the father attractive to women, and hence themselves be attractive to women and hence be successful in spreading the woman’s own genes to subsequent generations. 
In other words, being attractive to other women is itself an attractive trait in a male. However, sexy son theory is not discussed by Etcoff.

[9] Another study discussed by Etcoff also reported anomalous results, finding that women actually preferred somewhat feminized male faces over both masculinized and average male faces (Perrett et al 1998). However, Etcoff cautions that: 

The Perrett study is the only empirical evidence to date that some degree of feminization may be attractive in a man’s face” (p159). 

Other studies concur that male faces that are somewhat, but not excessively, masculinized as compared to the average male face are preferred by women. 
However, one study published just after the first edition of ‘Survival of the Prettiest’ was written, holds the possibility of reconciling these conflicting findings. This study reported cyclical changes in female preferences, with women preferring more masculinized faces only when they are in the most fertile phase of their cycle, and at other times preferring more feminine features (Penton-Voak & Perrett 2000). 
This, together with other evidence, has been controversially interpreted as suggesting that human females practice a so-called dual mating strategy, preferring males with more feminine faces, supposedly a marker for a greater willingness to invest in offspring, as social partners, while surreptitiously attempting to cuckold these ‘beta providers’ with DNA from high-T alpha males, by preferentially mating with the latter when they are most likely to be ovulating (see also Penton-Voak et al 1999Bellis & Baker 1990). 
However, recent meta-analyses have called into question the evidence for cyclical fluctuations in female mate preferences (Wood et al 2014; cf. Gildersleeve et al 2014), and it has been suggested that such findings may represent casualties of the so-called replication crisis in psychology
While the intensity of women’s sex drive does indeed seem to fluctuate cyclically, the evidence for more fine-grained changes in female mate preferences should be treated with caution. 

References 

Bateman (1948), Intra-sexual selection in DrosophilaHeredity, 2(3): 349–368. 
Bellis & Baker (1990). Do females promote sperm competition?: Data for humansAnimal Behavior, 40: 997-999. 
Clark & Hatfield (1989) Gender differences in receptivity to sexual offers. Journal of Psychology & Human Sexuality, 2(1), 39–55 
Doyle & Pazhoohi (2012) Natural and Augmented Breasts: Is What is Not Natural Most Attractive? Human Ethology Bulletin 27(4):4-14. 
Gaulin & Boser (1990) Dowry as Female Competition, American Anthropologist 92(4):994-1005. 
Gildersleeve et al (2014) Do women’s mate preferences change across the ovulatory cycle? A meta-analytic reviewPsychological Bulletin 140(5):1205-59. 
Hamermesh & Biddle (1994) Beauty and the Labor Market, American Economic Review 84(5):1174-1194.
Jones 1995 Sexual selection, physical attractiveness, and facial neoteny: Cross-cultural evidence and implications, Current Anthropology, 36(5):723–748. 
Kenrick & Keefe (1992) Age preferences in mates reflect sex differences in mating strategies. Behavioral and Brain Sciences 15(1):75-133. 
Orians & Heerwagen (1992) Evolved responses to landscapes. In Barkow, Cosmides & Tooby (Eds.), The Adapted Mind: Evolutionary Psychology and the Generation of Culture (pp. 555–579). Oxford University Press. 
Penton-Voak et al (1999) Menstrual cycle alters face preferencesNature 399 741-2. 
Penton-Voak & Perrett DI (2000) Female preference for male faces changes cyclically: Further evidence. Evolution and Human Behavoir 21(1):39–48. 
Perrett et al (1998) Effects of sexual dimorphism on facial attractiveness. Nature 394(6696):884-7. 
Puts (2013) Beauty and the Beast: Mechanisms of Sexual Selection in Humans. Evolution and Human Behavior 31(3):157-175. 
Wake Forest University (2009) Rating Attractiveness: Consensus Among Men, Not Women, Study Finds. ScienceDaily. ScienceDaily, 27 June 2009. 
Trivers (1972) Parental investment and sexual selectionSexual Selection & the Descent of Man, Aldine de Gruyter, New York, 136-179. Chicago. 
Williams (1957) Pleiotropy, natural selection, and the evolution of senescence. Evolution. 11(4): 398–411. 
Wood & Brumbaugh (2009) Using Revealed Mate Preferences to Evaluate Market Force and Differential Preference Explanations for Mate Selection, Journal of Personality and Social Psychology 96(6):1226-44.
Udry & Eckland (1984) Benefits of Being Attractive: Differential Payoffs for Men and Women, Psychological Reports 54(1):47–56.
Wood et al (2014). Meta-analysis of menstrual cycle effects on women’s mate preferencesEmotion Review, 6(3), 229–249.  

Donald Symons’ ‘The Evolution of Human Sexuality’: A Founding Work of Modern Evolutionary Psychology

The Evolution of Human Sexuality by Donald Symons (Oxford University Press 1980). 

Research over the last four decades in the field that has come to be known as evolutionary psychology has focused disproportionately on mating behaviour. Geoffrey Miller (1998) has even argued that it is the theory of sexual selection rather than that of natural selection which, in practice, guides most research in this field. 

This does not reflect merely the prurience of researchers. Rather, given that reproductive success is the ultimate currency of natural selection, mating behaviour is, perhaps along with parental investment, the form of behaviour most directly subject to selective pressures.

Almost all of this research traces its ancestry ultimately to Donald Symons’ ‘The Evolution of Human Sexuality’ by Donald Symons. Indeed, much of it was explicitly designed to test claims and predictions formulated by Symons himself in this very book.

Age Preferences

For example, in his discussion of the age at which women are perceived as most attractive by males, Symons formulated two alternative hypotheses. 

First, if human evolutionary history were characterized by fleeting one-off sexual encounters (i.e. one-night standscasual sex and hook-ups), then, he reasoned, men would have evolved to find women most attractive when the latter are at the age of their maximum fertility

For women, fertility is said to peak around when a woman reaches her mid-twenties since, although women still in their teens have high pregnancy rates, they also experience greater risk of birth complications

However, if human evolutionary history were characterized instead by long-term pair bonds, then men would have evolved to be maximally attracted to somewhat younger women (i.e. those at the beginning of their reproductive careers), so that, by entering a long-term relationship with the woman at this time, a male is potentially able to monopolize her entire lifetime reproductive output (p189). 

More specifically, males would have evolved to prefer females, not of maximal fertility, but rather of maximal reproductive value, a term borrowed from demography and population genetics which refers to a person’s expected future reproductive output given their current age. Unlike fertility, a woman’s reproductive value peaks around her mid- to late-teens.  

On the basis of largely anecdotal evidence, Symons concludes that human males have evolved to be most attracted to females of maximal reproductive value rather than maximal fertility.  

Subsequent research designed to test between Symons’s rival hypotheses has largely confirmed his speculative hunch that it is younger females in their mid- to late-teens who are perceived by males as most attractive (e.g. Kenrick and Keefe 1992). 

Why Average is Attractive

Symons is also credited as the first person to recognize that a major criterion of attractiveness is, paradoxically, averageness, or at least the first to recognize the significance of, and possible evolutionary explanation for, this discovery.[1] Thus, Symons argues that: 

“[Although] health and status are unusual in that there is no such thing as being too healthy or too high ranking… with respect to most anatomical traits, natural selection produces the population mean” (p194). 

On this view, deviations from the population mean are interpreted as the result of deleterious mutations or developmental instability, and hence bad genes.[2]

Concealed Ovulation

Support has even emerged for some of Symons’ more speculative hunches.

For example, one of Symons’ two proposed scenarios for the evolution of concealed ovulation, in which he professed “little confidence” (p141), was that this had evolved so as to impede male mate-guarding and enable females select a biological father for their offspring different from their husbands (p139-141).

Consistent with this theory, studies have found that women’s mate preferences vary throughout their menstrual cycle in a manner compatible with a so-called ‘dual mating strategy’, preferring males evidencing a willingness to invest in offspring at most times, but, when at their most fertile, preferring characteristics indicative of genetic quality (e.g. Penton-Voak et al 1999). 

Meanwhile, a questionnaire distributed via a women’s magazine found that women engaged in extra-marital affairs do indeed report engaging in ‘extra-pair copulations’ (EPCs) at times likely to coincide with ovulation (Bellis and Baker 1990).[3]

The Myth of Female Choice

Interestingly, Symons even anticipated some of the mistakes evolutionary psychologists would be led into.

Thus, he warns that researchers in modern western societies may be prone to overestimate the importance of female choice as a factor in human evolution, because, in their own societies, this is a major factor, if not the major factor, in determining marriage and sexual and romantic relationships (p203).[4]

However, in ancestral environments (i.e. what evolutionary psychologists now call the Environment of Evolutionary Adaptedness or EEA) arranged marriages were likely the norm, as they are in most premodern cultures around the world today (p168).[5]

Thus, Symons concludes: 

There is no evidence that any features of human anatomy were produced by intersexual selection [i.e. female choice]. Human physical sex differences are explained most parsimoniously as the outcome of intrasexual selection (the result of male-male competition)” (p203). 

Thus, human males have no obvious analogue of the peacock’s tail, but they do have substantially greater levels of upper-body strength and violent aggression as compared to females.[6]

This was a warning almost entirely ignored by subsequent generations of researchers before being forcefully reiterated by Puts (2010)

Homosexuality as a ‘Test-Case

An idea of the importance of Symons’s work can be ascertained by comparing it with contemporaneous works addressing the same subject-matter.

Edward O Wilson’s On Human Nature was first published in 1978, only a year before Symons’s ‘The Evolution of Human Sexuality’. 

However, whereas Symons’s book set out much of the theoretical basis for what would become the modern science of evolutionary psychology, Wilson’s chapter on “Sex” has dated rather less well, and a large portion of chapter is devoted to introducing a now faintly embarrassing theory of the evolution of homosexuality which has subsequently received no empirical support (see Bobrow & Bailey 2001).[7]

In contrast, Symons’s own treatment of homosexuality is innovative. It is also characteristic of his whole approach and illustrates why ‘The Evolution of Human Sexuality‘ has been described by David Buss as “the first major treatise on evolutionary psychology proper” (Handbook of Evolutionary Psychology: p251).

Rather than viewing all behaviours as necessarily adaptive (as critics of evolutionary psychology, such as Stephen Jay Gould, have often accused sociobiologists of doing),[8] Symons instead focuses on admittedly non-adaptive (or, indeed, even maladaptive) behaviours, not because he believes them to be adaptive, but rather because they provide a unique window on the nature of human sexuality.

Accordingly, Symons does not concern himself with how homosexuality evolved, implicitly viewing it as a rare and maladaptive malfunctioning of normal sexuality. Yet the behaviour of homosexuals is of interest to Symons because it provides a window on the nature of male and female sexuality as it manifests itself when freed from the constraints imposed by the conflicting desires of the opposite sex.

On this view, the rampant promiscuity manifested by many homosexual men (e.g. cruising and cottaging in bathhouses and public lavatories, or Grindr hookups) reflects the universal male desire for sexual variety when freed from the constraints imposed by the conflicting desires of women. 

This desire for sexual variety is, of course, obviously reproductively unproductive among homosexual men themselves. However, it evolved because it enhanced the reproductive success of heterosexual men by motivating them to attempt to mate with multiple females and thereby father multiple offspring.

Thus, a powerful ruler like with a large harem like Ismail the Bloodthirsty’ of Morocco could reputedly father as many as 888 offspring.

In contrast, burdened with pregnancy and lactation, women’s potential reproductive rate is more tightly constrained than that of men. They therefore have little to gain reproductively by mating with multiple males, since they can usually gestate, and nurse, only one offspring at a time.

It is therefore notable that, among lesbians, there is little evidence of the sort of rampant promiscuity common among gay men. Instead, lesbian relationships seem to be characterized by much the same features as heterosexual coupling (i.e. long-term pair-bonds).

The similarity of heterosexual coupling to that of lesbians, and the striking contrast with that of male homosexuals, suggests that it is women, not men, who exert decisive influence in dictating the terms of heterosexual coupling.[9]

Thus, Symons reports:

There is enormous cross-cultural variation in sexual customs and laws and the extent of male control, yet nowhere in the world do heterosexual relations begin to approximate those typical of homosexual men This suggests that, in addition to custom and law, heterosexual relations are structured to a substantial degree by the nature and interests of the human female” (p300). 

This conclusion is, of course, diametrically opposite to the feminist contention that it is men who dictate the terms of heterosexual coupling and for whose exclusive benefit such relationships are structured.

It also suggests, again contrary to feminist assumptions of male dominance, that most men are ultimately frustrated in achieving their sexual ambitions to a far greater extent than are most women. 

Thus, Symons concludes: 

The desire for sexual variety dooms most human males to a lifetime of unfulfilled longing” (p228). 

Here, Symons anticipates Camille Paglia who was later to famously observe: 

Men know they are sexual exiles. They wander the earth seeking satisfaction, craving and despising, never content. There is nothing in that anguished motion for women to envy” (Sexual Personae: p19). 

Criticisms of Symons’s Use of Homosexuality as a Test-Case

There is, however, a potential problem with Symons’s use of homosexual behaviour as a window onto the nature of male and female sexuality as they manifest themselves when freed from the conflicting desires of the opposite sex. The whole analysis rests on a questionable premise – namely that homosexuals are, their preference for same-sex partners aside, otherwise similar, if not identical, to heterosexuals of their own sex in their psychology and sexuality.

Symons defends this assumption, arguing: 

There is no reason to suppose that homosexuals differ systematically from heterosexuals in any way other than their sexual object choice” (p292). 

Indeed, in some respects, Symons seems to see even “sexual object choice” as analogous among homosexuals and heterosexuals of the same sex.

For example, he observes that, unlike women, both homosexual and heterosexual men tend to evaluate prospective mates primarily on the basis their physical appearance and youthfulness (p295). 

Thus, in contrast to the failure of periodicals featuring male nudes to attract a substantial female audience (see below), Symons notes the existence of a market for gay pornography parallel in most respects to heterosexual porn – i.e. featuring young, physically attractive models in various states of undress (p301).

This, of course, contradicts the feminist notion that men are led to ‘objectify’ women only due to the sexualized portrayal of the latter in the media.

Instead, Symons concludes: 

That homosexual men are at least as likely as heterosexual men to be interested in pornography, cosmetic qualities and youth seems to me to imply that these interests are no more the result of advertising than adultery and alcohol consumption are the result of country and western music” (p304).[10] 

However, this assumption of the fundamental similarity of heterosexual and homosexual male psychology has been challenged by David Buller in his book, Adapting Minds: Evolutionary Psychology and the Persistent Quest for Human Nature.

Buller cites evidence that male homosexuals are ‘feminized’ in many aspects of their behaviour and morphology.

For example, one study reported that, despite stereotypically more likely to ‘hit the gym’, gay man nevertheless had relatively less muscular development than heterosexual men, and lower shoulder-to-hip ratios on average (Evans 1972). Another particularly interesting recent study found that male homosexuals have more female-typical occupation interests than do heterosexual males (Ellis & Ratnasingam 2012).

Likewise, one of the few consistent early correlates of homosexuality is gender non-conformity in childhood and some evidence (e.g. digit ratios, the fraternal birth order effect) has been interpreted to suggest that the level of prenatal exposure to masculinizing androgens (e.g. testosterone) in utero affects sexual orientation (see Born Gay: The Pyschobiology of Sexual Orientation).

Indeed, Symons himself mentions the evidence of an association between homosexuality and levels of masculinizing androgens in utero (albeit in respect of lesbians rather than of male homosexuality) just a few pages before his discussion of the promiscuous behaviours of male homosexuals (p289).

As Buller also notes, although gay men seem, like heterosexual men, to prefer youthful sexual partners, they also appear to prefer sexual partners who are, in other respects highly masculine.[11]

Thus, Buller observes: 

“The males featured in gay men’s magazines embody very masculine, muscular physiques, not pseudo-feminine physiques” (Adapting Minds: p227).

Indeed, the models in such magazines seem in most respects similar in physical appearance to the male models, pop stars, actors and other ‘sex symbols’ and celebrities fantasized about by heterosexual women and girls.

How then are we to resolve this apparent paradox?

One possible explanation that some aspects of the psychology of male homosexuals are feminized but not others – perhaps because different parts of the brain are formed at different stages of prenatal development, at which stages the levels of masculinizing androgens in the womb may vary. 

Thus, Glenn Wilson, writing in 1989 and citing the work of Ellis & Ames (1987), reports that:

The masculinization/feminization effects occur in different parts of the brain and, more importantly, at different times during pre-natal development. Indications are that sex orientation in humans depends critically upon the hormone balance prevailing during the third and fourth months of pregnancy, while secondary sex characteristics and sex-typical behaviour patterns are influenced more by hormones circulating during the fifth and sixth months of pregnancy” (The Great Sex Divide: p79).

Indeed, there is even some evidence that homosexual males may be hyper-masculinized in some aspects of their physiology.

For example, it has been found that homosexual males report larger penis-sizes than heterosexual men (Bogaert & Hershberger 1999). 
 
Researchers Glenn Wilson and Qazi Rahman propose, may be because: 

If it is supposed that the barriers against androgens with respect to certain brain structures (notably those concerned with homosexuality) lead to increased secretion in an effort to break through, or some sort of accumulation elsewhere… then there may be excess testosterone left in other departments” (Born Gay: The Psychobiology of Sex Orientation: p80). 

Another possibility is that male homosexuals actually lie midway between heterosexual men and women in their degree of masculinization.  

On this view, homosexual men come across as relatively feminine only because we naturally tend to compare them to other men (i.e. heterosexual men). However, as compared to women, they may be relatively masculine, as reflected in the male-typical aspects of their sexuality focused upon by Symons.

Interestingly, this latter interpretation suggests the slightly disturbing possibility that, freed from the restraints imposed by women, heterosexual men would be even more indiscriminately promiscuous than their homosexual counterparts.

Evidence consistent with this interpretation is provided by one study from the 1980s which found that, when approached by a female stranger (also a student), on a University campus, with a request to go to bed with them, fully 72% of male students agreed (Clark and Hatfield 1989). 

In contrast, in the same study, not a single one of the 96 females approached by male strangers with the same request on the same university campus agreed to go to bed with the male stranger.

Yet what percentage of the female students subsequently sued the university for sexual harassment was not reported.

Pornography as a “Natural Experiment

For Symons, fantasy represents another window onto sexual and romantic desires. Like homosexuality, fantasy is, by its very nature, unconstrained by the conflicting desires of the opposite sex (or indeed by anything other than the imagination of the fantasist). 

Symons later collaborated in an investigation into sexual fantasy by means of a questionnaire (Ellis and Symons 1990). 

However, in the present work, he investigates fantasy indirectly by focusing on what he calls “the natural experiment of commercial periodical publishing” – i.e. pornographic magazines (p182).

In many respects, this approach is preferable to a survey because, even in an anonymous questionnaire, individuals may be less than honest when dealing with a sensitive topic such as their sexual fantasies. On the other hand, they are unlikely to regularly spend money on a magazine unless they are genuinely attracted by its contents.

Before the internet age, softcore pornographic magazines, largely featuring female nudes, commanded sizeable circulations, despite the not insubstantial stigma attached to their purchase. However, their readership (if indeed ‘readership’ is the right words, since there was typically little reading involved, save of the ‘one-handed’ variety) was almost exclusively male.

In contrast, there was little or no female audience for magazines containing pictures of naked males. Instead, magazines marketed towards women (e.g. fashion magazines) contain, mostly, pictures of other women.

Indeed, when, in the 1970s, attempts were made, in the misguided name of feminism and ‘women’s liberation, to market magazines featuring male nudes to a female readership, one such title, Viva, abandoned publishing male nudes after just a few years due to lack of interest or demand, then subsequently went bust just a few years after that, while the other, Playgirl, although it remained in publication for many years and did not entirely abandon male nudes, was notorious, as a consequence, for attracting a readership composed in large part of homosexual men.

Symons thus concludes forcefully and persuasively: 

The notion must be abandoned that women are simply repressed men waiting to be liberated” (p183). 

Indeed, though it has been loudly and enthusiastically co-opted by feminists, this view of women, and of female sexuality – namely women as “repressed men waiting to be liberated” – represents an obviously quintessentially male persepective. 

Indeed, taken to extremes, it has even been used as a justification for rape.

Thus, the curious, though recurrent, sub-Freudian notion that female rape victims actually secretly enjoy being raped seems to rest ultimately on the assumption that female sexuality is fundamentally the same as that of men (i.e. indiscriminately enjoying of promiscuous sex) and that it is only women’s alleged sexual ‘repression’ that prevents them admitting as much.[12]

Romance Literature 

Unfortunately, however, there is notable omission in Symons’s discussion of pornography as a window into male sexuality – namely, he omits to consider whether there exists any parallel artistic genre that offers equivalent insight into the female psyche.

Later writers on the topic have argued that romance novels (e.g. Mills and Boon, Jane Austin), whose audience is as overwhelmingly female as pornography’s is male, represent the female equivalent of pornography, and that analysis of the the content of such works provides insights into female mate preferences parallel to those provided into male psychology by pornography (e.g. Kruger et al 2003; Salmon 2004; see also Warrior Lovers: Erotic Fiction, Evolution and Female Sexuality, co-authored by Symons himself).

Thus, popular science writer Matt Ridley reports:

Two industries relentlessly exploit the sexual fantasizing of men and women: pornography and the publishing of romance novels: Pornography is aimed almost entirely at men. It varies little from a standard formula all over the world… The romance novel, by contrast, is aimed entirely at a female market. It, too, depicts a fictional world that has changed remarkably little except in adapting to female career ambitions and to a less inhibited attitude toward the description of sex” (The Red Queen: p270-271)

Symons touches upon this analogy only in passing, when he observes that:

Heterosexual men are, of course, aware that the female sexuality portrayed in men’s magazines reflects male fantasy more than female reality, just as heterosexual women are aware that the happy endings of stories in romance magazines exist largely in the realm of fantasy” (p293).

Yet, while feminists perpetually complain about how pornography supposedly creates unrealistic expectations of women and girls and puts undue pressure on women and girls to live up to this male fantasy, few men complain about how the equally unrealistic portrayal of men in romance literature creates unrealistic expectations of boys and men and puts undue pressure on boys and men to live up to a female fantasy.

Female Orgasm as Non-Adaptive

An entire chapter of ‘The Evolution of Human Sexuality’, namely Chapter Three (entitled, “The Female Orgasm: Adaptation or Artefact”), is devoted to rejecting the claim that the female orgasm represents a biological adaptation.

This is perhaps excessive. However, it does at least conveniently contradicts the claim of some critics of evolutionary psychology, and of sociobiology, such as Stephen Jay Gould that the field is ‘ultra-Darwinian’ or ‘hyper-adaptionist’ and committed to the misguided notion that all traits are necessarily adaptive.[13]

In contrast, Symons champions the thesis that the female capacity for orgasm is a simply non-adaptive by-product of the male capacity to orgasm, the latter of which is of course adaptive.

On this view, the female orgasm (and clitoris) is, in effect, the female equivalent of male nipples (only more fun).

Certainly, Symons convincingly critiques the romantic notion, popularized by Desmond Morris among others, that the female orgasm functions as a mechanism designed to enhance ‘pair-bonding between couples.

However, subsequent generations of evolutionary psychologists have developed less naïve models of the adaptive function of female orgasm.

For example, Geoffrey Miller argues that the female orgasm, and clitoris, functions as an adaptation for mate choice (The Mating Mind: p239-241).

Of course, at first glance, experiencing orgasm during coitus may appear to be a bit late for mate choice, since, by the time coitus has occurred, the choice in question has already been made. However, given that, among humans, most sexual intercourse is non-reproductive (i.e. does not result in conception), the theory is not altogether implausible.

On this view, the very factors which Symons views as suggesting female orgasm is non-adaptive – such as the relative difficultly of stimulating female orgasm during ordinary vaginal sex – are positive evidence for its adaptive function in carefully discriminating between suitors/lovers to determine their desirability as father for a woman ’s offspring.

Nevertheless, at least according to the stringent criteria set out by George C Williams in his classic Adaptation and Natural Selection, as well as the more general principle of parsimony (also known as Occam’s Razor), the case for female orgasm as an adaptation remains unproven (see also Sherman 1989; Case Of The Female Orgasm: Bias in the Science of Evolution).

Out-of-Date?

Much of Symons’ work is dedicated to challenging the naïve group-selectionism of Sixties ethologists, especially Desmond Morris. Although scientifically now largely obsolete, Morris’s work still retains a certain popular resonance and therefore this aspect of Symons’s work is not entirely devoid of contemporary relevance.

In place of Morris‘s rather idyllic notion that humans are a naturally monogamous ‘pair-bonding’ species, Symons advocates instead an approach rooted in the individual-level (or even gene-level) selection championed Richard Dawkins in The Selfish Gene (reviewed here).

This leads to some decidedly cynical conclusions regarding the true nature of sexual and romantic relations among humans.

For example, Symons argues that it is adaptive for men to be less sexually attracted to their wives than they are to other women – because they are themselves liable to bear the cost of raising offspring born to their wives but not those born to other women with whom they mate (e.g. those attached to other males).

Another cynical conclusion is that the primary emotion underlying the institution of marriage, both cross-culturally and in our own society, is neither love nor even lust, but rather male sexual jealousy and proprietariness (p123). 

Marriage, then, is an institution borne not of love, but of male sexual jealousy and the behaviour known to biologists as mate-guarding.

Meanwhile, in his excellent chapter on ‘Copulation as a Female Service’ (Chapter Eight), Symons suggests that many aspects of heterosexual romantic relationships may be analogous to prostitution.

As well as its excessive focus on debunking sixties ethologists like Morris, ‘The Evolution of Human Sexuality’ is also out-of-date in a more serious respect Namely, it fails to incorporate the vast amount of empirical research on human sexuality from a sociobiological perspective which has been conducted since the first publication of his work.

For a book first published thirty years ago, this is inevitable – not least because much of this empirical research was inspired by Symons’ own ideas and specifically designed to test theories formulated in this very work.

In addition, potentially important new factors in human reproductive behaviour that even Symons did not foresee have been identified, for example role of levels of fluctuating asymmetry functioning as a criterion for, or at least correlate of, physical attractiveness.

For an updated discussion of the evolutionary psychology of human sexual behaviour, complete with the latest empirical data and research, readers should consult the latest edition of David Buss’s The Evolution Of Desire: Strategies of Human Mating.

In contrast, in support of his theories Symons relies largely on classical literary insight, anecdote and, most importantly, a review of the ethnographic record.

However, this latter focus ensures that, in some respects, the work remains of more than merely of historical interest.

After all, one of the more legitimate criticisms levelled against recent research in evolutionary psychology is that it is insufficiently cross-cultural and, with several notable exceptions (e.g. Buss 1989), relies excessively on research conducted among convenience samples of students at western universities.

Given costs and practicalities, this is inevitable. However, for a field that aspires to understand a human nature presumed to be universal, such a method of sampling is highly problematic, especially given what has recently been revealed about the ‘WEIRD-ness’ of western undergraduate samples.

The Evolution of Human Sexuality’ therefore retains its importance for two reasons. 

First, is it the founding work of modern evolutionary psychological research into human sexual behaviour, and hence of importance as a landmark and classic text in the field, as well as in the history of science more generally. 

Second, it also remains of value to this day for the cross-cultural and ethnographic evidence it marshals in support of its conclusions. 

Endnotes

[1] Actually, the first person to discover this, albeit inadvertently, was the great Victorian polymath, pioneering statistician and infamous eugenicist Francis Galton, who, attempting to discover abnormal facial features possessed by the criminal class, succeeded in morphing the faces of multiple convicted criminals. The result was, presumably to his surprise, an extremely attractive facial composite, since all the various minor deformities of the many convicted criminals whose faces he morphed actually balanced one another out to produce a face with few if any abnormalities or disproportionate features.

[2] More recent research in this area has focused on the related concept of fluctuating asymmetry.

[3] However, recent meta-analyses have called into question the evidence for cyclical fluctuations in female mate preferences (Wood et al 2014; cf. Gildersleeve et al 2014), and it has been suggested that such findings may represent casualties of the so-called replication crisis in psychology. It has also been questioned whether ovulation in humans is indeed concealed, or is actually detectable by subtle cues (e.g. Miller et al 2007), for example, changes in face shape (Oberzaucher et al 2012), breast symmetry (Scutt & Manning 1996) and body scent (Havlicek et al 2006).

[4] Another factor leading recent researchers to overestimate the importance of female choice in human evolution is their feminist orientation, since female choice gives women an important role in human evolution, even, paradoxically, in the evolution of male traits.

[5] Actually, in most cultures, only a girl’s first marriage is arranged on her behalf by her parents. Second- and third-marriages are usually negotiated by the woman herself. However, since female fertility peaks early, it is a girl’s first marriage that is usually of the most reproductive, and hence Darwinian, significance.

[6] Indeed, the human anatomical trait in humans that perhaps shows the most evidence of being a product of intersexual selection is a female one, namely the female breasts, since the latter are, unlike the mammary glands of most other mammals, permanently present from puberty on, not only during lactation, and composed primarily of fatty tissues, not milk (Møller 1995; Manning et al 1997; Havlíček et al 2016). 

[7] Wilson terms his theory “the kin selection theory hypothesis of the origin of homosexuality” (p145). However, a better description might be the ‘helper at the nest theory of homosexuality’, the basic idea being that, like sterile castes in some insects, and like older siblings in some bird species where new nest sites are unavailable, homosexuals, rather than reproducing themselves, direct their energies towards assisting their collateral kin in successfully raising, and provisioning, their own offspring (On Human Nature: p143-7). The main problem with this theory is that there is no evidence that homosexuals do indeed devote any greater energies towards assisting their kin in raising offspring. On the contrary, homosexuals instead seem to devote much of their time and resources towards their own sex life, much as do heterosexuals (Bobrow & Bailey 2001).

[8] As we will see, contrary to the stereotype of evolutionary psychologists as viewing all traits as necessarily adaptive, as they are accused of doing by the likes of Gould, Symons also argued that the female orgasm and menopause are non-adaptive, but rather by-products of other adaptations.

[9] This is not necessarily to say that rampant, indiscriminate promiscuity is a male utopia, or the ideal of any man, be he homosexual or heterosexual. On the contrary, the ideal mating system for any individual male is harem polygyny in which the chastity of his own partners is rigorously policed (see Laura Betzig’s Despotism and Differential Reproduction: which I have reviewed here). However, given an equal sex ratio, this would condemn other males to celibacy and perpetual ‘inceldom. Similarly, Symons reports that “Homosexual men, like most people, usually want to have intimate relationships”. However, he observes:

Such relationships are difficult to maintain, largely owing to the male desire for sexual variety; the unprecedented opportunity to satisfy this desire in a world of men, and the male tendency towards sexual jealousy” (p297).  

It does indeed seem to be true that homosexual relationships, especially those of gay males, are, on average, of shorter duration than are heterosexual relationships. However, Symons’ claim regarding “the male tendency towards sexual jealousy” is questionable.
Actually, subsequent research in evolutionary psychology has suggested that men are no more prone to jealousy than women, but rather that it is sorts of behaviours which most intensely provoke such jealousy that differentiate the sexes (Buss 1992). Moreover, many gay men practice open relationships, which seems to suggest a lack of jealousy – or perhaps this simply reflects a recognition of the difficulty of maintaining relationships given, as Symons puts it, “the male desire for sexual variety [and] the unprecedented opportunity to satisfy this desire in a world of men”. 

[10] Indeed, far from men being led to objectify women due to the portrayal of women in a sexualized manner in the media, Symons suggests:

There may be no positive feedback at all; on the contrary, constant exposure to pictures of nude and nearly nude female bodies may to some extent habituate [i.e. desensitize] men to these stimuli” (p304).

[11] Admittedly, some aspects of body-type typically preferred by gay males (especially the so-called twink ideal) do reflect apparently female traits, especially a relative lack of body-hair. However, lack of body-hair is also obviously indicative of youth. Moreover, a relative lack of body-hair also seems to be a trait favoured in men by heterosexual women. For a discussion of the relative preference on the part of (heterosexual) females for masculine versus feminine physical appearance in male sex partners, see here.

[12] Thus, some men might indeed welcome being ‘raped’, albeit only under highly unusual circumstances – namely by an attractive opposite-sex partner (or, in the case of homosexual men, an attractive same-sex partner) to whom they are sexually attracted. Thus, Kingsley Browne, in his excellent Biology at Work (which I have reviewed here) quotes the perhaps remarkable finding that:

A substantial number of men ‘viewed an advance by a good-looking woman who threatened harm or held a knife as a positive sexual opportunity’” (Biology at Work: p196; quoting Struckman-Johnson & Struckman-Johnson 1994).

Of course, large numbers of women also report rape fantasies (Bivona & Critelli 2009). Yet this does not, of course, mean they would actually welcome real sexual assault, which would almost certainly take a very different form from the fantasy. In practice, therefore, members of neither sex are ever likely to welcome sexual assault in the form which it is actually likely to actually come.

[13] Incidentally, Symons also rejects the theory that the female menopause is adaptive, a theory which has subsequently become known as the grandmother hypothesis (p13). Also, although it does not directly address the issue, Symons’ discussion of human rape (p276-85), has also been interpreted as implicitly favouring the theory that rape is a by-product of the greater male desire for commitment free promiscuous sex, rather than the product of a specific rape adaptation in males (see Palmer 1991; and A Natural History of Rape: reviewed here). 

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