favoring survival. The costly structure is thus a doubly honest ad: only a superior animal can afford its cost, and it makes the animal even more superior.

For instance, the antlers of male deer represent a big investment of calcium, phosphate, and calories, yet they are grown and discarded each year. Only the most well-nourished males-ones that are mature, socially dominant, and free of parasites-can afford that investment. Hence a female deer can regard big antlers as an honest ad for male quality, just as a woman whose boyfriend buys and discards a Porsche sports car each year can believe his claim of being wealthy. But antlers carry a second message not shared with Porsches. Whereas a Porsche does not generate more wealth, big antlers do bring their owner access to the best pastures by enabling him to defeat rival males and fight off predators.

Let us now examine whether any of these three theories, devised to explain the evolution of animal signals, can also explain features of human bodies. But we first need to ask whether our bodies possess any such features requiring explanation. Our first inclination might be to assume that only stupid animals require genetically coded badges, like a red dot here and a black stripe there, in order to figure out each other's age, status, sex, genetic quality, and value as a potential mate. We, in contrast, have much bigger brains and far more reasoning ability than any other animal. Moreover, we are uniquely capable of speech and can thereby store and transmit far more detailed information than any other animal can. What need have we of red dots and black stripes when we routinely and accurately determine the age and status of other humans just by talking to them? What animal can tell another animal that it is twenty-seven years old, receives an annual salary of $125,000, and is second assistant vice president at the country's third largest bank? In selecting our mates and sex partners, don't we go through a dating phase that is in effect a long series of tests by which we accurately assess a prospective partner's parenting skills, relationship skills, and genes?

The answer is simple: nonsense! We too rely on signals as arbitrary as a widowbird's tail and a bowerbird's crest. Our signals include faces, smells, hair color, men's beards, and women's breasts. What makes those structures less ludicrous than a long tail as grounds for selecting a spouse— the most important person in our adult life, our economic and social partner, and the coparent of our children? If we think that we have a signaling system immune to cheating, why do so many people resort to makeup, hair dyes, and breast augmentation? As for our supposedly wise and care-ful selection process, all of us know that when we walk into a room full of unfamiliar people, we quickly sense who attracts us physically and who doesn't. That quick sense is based on “sex appeal,” which just means the sum of the body signals to which we respond, largely unconsciously. Our divorce rate, now around 50 percent in the United States, shows that we ourselves acknowledge the failure of half of our efforts to select mates. Albatrosses and many other pair-bonded animal species have much lower “divorce” rates. So much for our wisdom and their stupidity!

In fact, like other animal species, we have evolved many body traits that signal age, sex, reproductive status, and individual quality, as well as programmed responses to those and other traits. Attainment of reproductive maturity is signaled in both human sexes by the growth of pubic and axillary hair. In human males it is further signaled by the growth of a beard and body hair and by a drop in the pitch of the voice. The episode with which I began this chapter illustrates that our responses to those signals can be as specific and dramatic as a gull chick's response to the red spot on its parent's bill. Human females additionally signal reproductive maturity by expansion of the breasts. Later in life, we signal our waning fertility and (in traditional societies) attainment of wise elder status by the whitening of our hair. We tend to respond to the sight of body muscles (in appropriate amounts and places) as a signal of male physical condition, and to the sight of body fat (also in appropriate amounts and places) as a signal of female physical condition. As for the body signals by which we select our mates and sex partners, they include all those same signals of reproductive maturity and physical condition, with variation among human populations in the sig-nals that one sex possesses and that the other sex prefers.

For instance, men vary around the world in the luxuriance of their beard and body hair, while women vary geographically in the size and shape of their breasts and nipples and in their nipple color. All of these structures serve us humans as signals analogous to the red dots and black stripes of birds. In addition, just as women's breasts simultaneously perform a physiological function and serve as a signal, I shall consider later in this chapter whether the same might be true for men's penises.

Scientists seeking to understand the corresponding signals of animals can carry out experiments involving mechanical modifications of an animal's body, such as shortening a widowbird's tail or painting over a gull's red spot. Legal obstacles, moral compunctions, and ethical considerations prevent us from performing such controlled experiments on humans. Also preventing us from understanding human signals are our own strong feelings that cloud our objectivity about them, and the great degree of cultural variation and individually learned variation in both our preferences and our bodies' self-modifications. However, such variation and self-modification can also help us gain understanding by serving as natural experiments, albeit ones lacking experimental controls. At least three sets of human signals seem to me to conform to Kodric-Brown's and Brown's truth-in-advertising model: men's body muscle, facial “beauty” in both sexes, and women's body fat.

Men's body muscle tends to impress women as well as other men. While the extreme muscle development of professional bodybuilders strikes many people as grotesque, many (most?) women find a well-proportioned muscular man more attractive than a scrawny man. Men also use the muscular development of other men as a signal-for example, as a way of quickly assessing whether to get into a fight or to retreat. A typical example involves a magnificently muscular instructor named Andy at the gymnasium where my wife and I exercise. Whenever Andy lifts weights, the eyes of all the women and men in the gym are on him. When Andy explains to a customer how to use one of the gym's exercise machines, he begins by demonstrating the machine's operation himself while asking the customer to place a hand on the relevant muscle on Andy's body so that the customer can understand the correct motion. Undoubtedly, this means of explanation is pedagogically useful, but I am sure that Andy also enjoys the overwhelming impression that he leaves.

At least in traditional societies based on human muscle power rather than on machine power, muscles are a truthful signal of male quality, like a deer's antlers. On the one hand, muscles enable men to gather resources such as food, to construct resources such as houses, and to defeat rival men. In fact, muscles play a much larger role in a traditional man's life than do antlers in the life of a deer, which uses antlers only in fighting. On the other hand, men with other good qualities are better able to acquire all the protein required to grow and maintain big muscles. One can fake one's age by dyeing one's hair, but one cannot fake big muscles. Naturally, men did not evolve muscles solely to impress other men and women, in the way that male bowerbirds evolved a golden crest solely as a signal to impress other bowerbirds. Instead, muscles evolved to perform functions, and men and women then evolved or learned to respond to muscles as a truthful signal.

A beautiful face may be another truthful signal, although the underlying reason is not as transparent as in the case of muscles. If you stop to think about it, it may seem absurd that our sexual and social attractiveness depends on facial beauty to such an inordinate degree. One might reason that beauty says nothing about good genes, parent-ing qualities, or food-gathering skills. However, the face is the part of the body most sensitive to the ravages of age, disease, and injury. Especially in traditional societies, individuals with scarred or misshapen faces may thereby be advertising their proneness to disfiguring infections, inability to take care of themselves, or burden of parasitic worms. A beautiful face was thus a truthful signal of good health that could not be faked until twentieth-century plastic surgeons perfected facelifts.

Our remaining candidate for a truthful signal is women's body fat. Lactation and child care are a big energy drain on a mother, and lactation tends to fail in an undernourished mother. In traditional societies before the advent of infant formulas and before the domestication of milk-producing hoofed animals, a mother's lactational failure would have been fatal to her infant. Hence a woman's body fat would be a truthful signal to a man that she was capable of rearing his child. Naturally, men should prefer the correct amount of fat: too little could be a harbinger of lactational failure, but too much could signal difficulties in walking, poor food-gathering ability, or early death from diabetes.

Perhaps because fat would be difficult to discern if it were spread uniformly over the body, women's bodies have evolved with fat concentrated in certain parts that are readily visible and assessed, although the anatomical location of those fat deposits varies somewhat among human populations. Women of all populations tend to accumulate fat in the breasts and hips, to a degree that varies geographically. Women of the San population native to southern Africa (the so-called Bushmen and Hottentots) and women of the Andaman Islands in the Bay of Bengal accumulate fat in the buttocks, producing the condition known as steatopy-gia. Men throughout the world tend to be interested in women's breasts, hips, and buttocks, giving rise in modern societies to yet another surgical method of fake signals, breast enhancement. Of course, one can object that some individual men are less interested than other men in these signs of female nutritional status, and that the relative popularity of skinny and plump fashion models fluctuates from year to year as fads. Nevertheless, the overall trend in male interest is clear.

Suppose one were again playing God or Darwin and deciding where on a woman's body to concentrate body fat as a visible signal. The arms and legs would be excluded because of the resulting extra load on them during walking or use of the arms. That still leaves many parts of the torso where fat could be safely concentrated without impeding movement, and in fact I just mentioned that women of various populations have evolved three different signaling areas on the torso. Nevertheless, one has to ask whether the evolutionary choice of signaling area is completely arbitrary, and why there are no populations of women with other signaling locations, such as the belly or the middle of the back. Paired fat deposits on the belly would seem to create no more difficulties for locomotion than do our actual paired deposits in the breasts and buttocks. It is curious, however, that women of all populations have evolved fat deposition in the breasts, the organs whose lactational performance men may be attempting to assess by fat deposit signals. Hence some scientists have suggested that large fatty breasts are not only an honest signal of good overall nutrition but also a deceptive specific signal of high milk-producing ability (deceptive because milk is actually secreted by breast glandular tissue rather than by breast fat). Similarly, it has been suggested that fat deposition in the hips of women worldwide is also both an honest signal of good health and a deceptive specific signal suggesting a wide birth canal (deceptive because a truly wide birth canal would minimize the risk of birth traumas but mere fat hips would not).

At this point, I have to anticipate several objections to my assumption that the sexual ornamentation of women's bodies could have any evolutionary significance. Whatever the interpretation, it is of course a fact that women's bodies do possess structures functioning as sexual signals, and that men tend to be especially interested in those particular parts of women's bodies. In those respects women resemble females of other primate species living in troops that contain many adult males and adult females. Like humans, chimpanzees and baboons and macaques live in troops and have sexually ornamented females (as well as males). By contrast, female gibbons and the females of other primate species that live as solitary male-female pairs bear little or no sexual ornamentation. This correlation suggests that if and only if females compete intensively with other females for males' attention-for example, because multiple males and females encounter each other daily in the same troop-then females tend to evolve sexual ornamentation in an ongoing evolutionary contest to be more attractive. Females who do not have to compete on such a regular basis have less need of expensive body ornamentation.

In most animal species (including humans) the evolutionary significance of male sexual ornamentation is undisputed, because males surely compete for females. However, scientists have raised three objections to the interpretation that women compete for men and have evolved bodily ornaments for that purpose. First, in traditional societies at least 95 percent of women marry. This statistic seems to suggest that virtually any woman can get a husband, and that women have no need to compete. As one woman biologist expressed it to me, “Every garbage can has a lid, and there is usually a bad-looking man for every bad-looking woman.”

But that interpretation is belied by all the effort that women consciously put into decoration and surgical modification of their bodies so as to be attractive. In fact, men vary greatly in their genes, in the resources that they control, in their parenting qualities, and in their devotion to their wives. Although virtually any woman can get some man to marry her, only a few women can succeed in getting one of the few high-quality men, for whom women must compete intensely. Every woman knows that, even though some male scientists evidently don't.

A second objection notes that men in traditional societies had no opportunity to choose their spouse, whether on the basis of sexual ornamentation or any other quality. Instead, marriages were arranged by clan relatives, who did the choosing, often with the motive of cementing political alliances. In reality, though, bride prices in traditional societies, such as the New Guinea societies where I work, vary according to a woman's desirability, the woman's health and probable mothering qualities being important considerations. That is, although a bridegroom's views about his bride's sex appeal may be ignored, his relatives who actually select the bride do not ignore

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