expecting me to conclude that American men aren't good for much. Of course that's not what I conclude. I acknowledge that many (most? by far the most?) American men are devoted husbands, work hard to increase their income, devote that income to their wives and kids, do much child care, and don't philander.

But, alas, the Ache findings are relevant to at least some men in our society. Some American men do desert their wives and children. The proportion of divorced men who renege on their legally stipulated child support is scandalously high, so high that even our government is starting to do something about it. Single parents outnumber copar-ents in the United States, and most single parents are women.

Among those men who remain married, all of us know some who take better care of themselves than of their wives and children, and who devote inordinate time, money, and energy to philandering and to male status symbols and activities. Typical of such male preoccupations are cars, sports, and alcohol consumption. Much bacon isn't brought home. I don't claim to have measured what percentage of American men rate as show-offs rather than providers, but the percentage of show-offs appears not to be negligible.

Even among devoted working couples, time budget studies show that American working women spend on the average twice as many hours on their responsibilities (defined as job plus children plus household) as do their husbands, yet women receive on the average less pay for the same job. When American husbands are asked to estimate the number of hours that they and their wives each devote to children and household, the same time budget studies show that men tend to overestimate their own hours and to underestimate their wife's hours. It's my impression that men's household and child-care contributions are on the average even lower in some other industrialized countries, such as Australia, Japan, Korea, Germany, France, and Poland, to mention just a few with which I happen to be familiar. That's why the question what men are good for continues to be debated within our societies, as well as between anthropologists.

CHAPTER 6. MAKING MORE BY MAKING LESS: The Evolution of Female Menopause

Most wild animals remain fertile until they die, or until close to that time. So do human males: although some men become infertile or less fertile at various ages for various reasons, men experience no universal shutdown of fertility at any particular age. There are innumerable well-attested cases of old men, including a ninety-four- year-old, fathering children.

But human females undergo a steep decline in fertility from around age forty, leading to universal complete sterility within a decade or so. While some women continue to have regular menstrual cycles up to the age of fifty-four or fifty-five, conception after the age of fifty was rare until the recent development of medical technologies using hormone therapy and artificial fertilization. For example, among the American Hutterites, a strict religious community that is well nourished and opposed to contraception, women produce babies as fast as is biologically possible for humans, with a mean interval of only two years between births, and a mean final number of eleven children. Even Hutterite women stop producing babies by age forty-nine.

To laypeople, menopause is an inevitable fact of life, albeit often a painful one anticipated with foreboding. But to evolutionary biologists, human female menopause is an aberration in the animal world and an intellectual paradox. The essence of natural selection is that it promotes genes for traits that increase the number of one's descendants bearing those genes. How could natural selection possibly result in every female member of a species carrying genes that throttle her ability to leave more descendants? All biological traits are subject to genetic variation, including the age of human female menopause. Once female menopause somehow became fixed in humans for whatever reason, why did not its age of onset gradually become pushed back until it disappeared again, because those women who experienced menopause later in life left behind more descendants?

To evolutionary biologists, female menopause is thus among the most bizarre features of human sexuality. As I shall argue, it is also among the most important. Along with our big brains and upright posture (emphasized in every text of human evolution), and our concealed ovula-tions and penchant for recreational sex (to which texts devote less attention), I believe that female menopause was among the biological traits essential for making us distinctively human-a creature more than, and qualitatively different from, an ape.

Many biologists would balk at what I have just said. They would argue that human female menopause does not pose an unsolved problem, and that there is no need to discuss it further. Their objections are of three types.

First, some biologists dismiss human female menopause as an artifact of a recent increase in human expected life span. That increase stems not just from public health measures within the last century but possibly also from the rise of agriculture ten thousand years ago, and even more likely from evolutionary changes leading to increased human survival skills within the last forty thousand years. According to this view, menopause could not have been a frequent occurrence for most of the several million years of human evo-lution, because (supposedly) almost no women or men survived past the age of forty. Of course, the female reproductive tract was programmed to shut down by age forty, because it would not have had the opportunity to operate thereafter anyway. The increase in human life span has developed much too recently in our evolutionary history for the female reproductive tract to have had time to adjust-so goes this objection.

However, this view ignores the fact that the human male reproductive tract, and every other biological function of both women and men, continue to function in most people for many decades after age forty. One would therefore have to assume that every other biological function was able to adjust quickly to our new long life span, leaving unexplained why female reproduction was uniquely incapable of doing so. The claim that formerly few women survived until the age of menopause is based on paleode-mography, that is, on attempts to estimate age at time of death in ancient skeletons. Those estimates rest on un-proven, implausible assumptions, such as that the recovered skeletons represent an unbiased sample of an entire ancient population, or that ancient adult skeletons really can be aged accurately. While paleodemographers' ability to distinguish the ancient skeleton of a ten-year-old from that of a twenty-five-year-old is not in question, the ability they claim to distinguish an ancient forty-year-old from a fifty-five-year-old has never been demonstrated. One can hardly reason by comparison with skeletons of modern people, whose different lifestyles, diets, and diseases surely make their bones age at different rates from the bones of ancients.

A second objection acknowledges human female menopause as a possibly ancient phenomenon but denies that it is unique to humans. Many or most wild animals exhibit a decrease in fertility with age. Some elderly individuals of a wide variety of wild mammal and bird species are found to be infertile. Many elderly female individuals of rhesus macaques and certain strains of laboratory mice, living in laboratory cages or zoos where their lives are considerably extended over expected spans in the wild by gourmet diets, superb medical care, and complete protection from enemies, do become infertile. Hence some biologists object that human female menopause is merely part of a widespread phenomenon of animal menopause. Whatever that phenomenon's explanation, its existence in many species would mean that there is not necessarily anything peculiar about menopause in the human species requiring explanation.

However, one swallow does not make a summer, nor does one sterile female constitute menopause. That is, detection of an occasional sterile elderly individual in the wild, or of regular sterility in caged animals with artificially extended life spans, does nothing to establish the existence of menopause as a biologically significant phenomenon in the wild. That would require demonstrating that a substantial fraction of adult females in a wild animal population become sterile and spend a significant portion of their life spans after the end of their fertility.

The human species does fulfill that definition, but only one or possibly two wild animal species are definitely known to do so. One is an Australian marsupial mouse in which males (not females) exhibit something like menopause: all males in the population become sterile within a short time in August and die over the next couple of weeks, leaving a population that consists solely of pregnant females. In that case, however, the postmenopausal phase is a negligible fraction of the total male life span. Marsupial mice do not exemplify true menopause but are more appropriately considered an example of big-bang reproduction, alias semelparity-a single lifetime reproductive effort rapidly followed by sterility and death, as in salmon and century plants. The better example of animal menopause is provided by pilot whales, among which one-quarter of all adult females killed by whalers proved to be postmenopausal, as judged by the condition of their ovaries. Female pilot whales enter menopause at the ago of thirty or forty years, have a mean survival of at least fourteen years after menopause, and may live for over sixty years.

Menopause as a biologically significant phenomenon is thus not unique to humans, being shared at least with one species of whale. It would be worth looking for evidence of menopause in killer whales and a few other species as possible candidates. But still-fertile elderly females are often encountered among well-studied wild populations of other long-lived mammals, including chimpanzees, gorillas, baboons, and elephants. Hence those species and most others are unlikely to be characterized by regular menopause. For example, a fifty-five-year-old elephant is considered elderly, since 95 percent of elephants die before that age. But the fertility of fifty-five- year-old female elephants is still half that of younger females in their prime.

Thus, female menopause is sufficiently unusual in the animal world that its evolution in humans requires explanation. We certainly did not inherit it from pilot whales, from whose ancestors our own ancestors parted company over fifty million years ago. In fact, we must have evolved it since our ancestors separated from those of chimps and gorillas seven million years ago, because we undergo menopause and chimps and gorillas appear not to (or at least not regularly).

The third and last objection acknowledges human menopause as an ancient phenomenon that is unusual among animals. Instead, these critics say that we need not seek an explanation for menopause, because the puzzle has already been solved. The solution (they say) lies in the physiological mechanism of menopause: a woman's egg supply is fixed at her birth and not added to later in her life. One or more eggs are lost by ovulation at each menstrual cycle, and far more eggs simply die (termed atresia). By the time a woman is fifty years old, most of her original egg supply has been depleted. Those eggs that remain are half a century old, increasingly unresponsive to pituitary hormones, and too few in number to produce enough estra-diol to trigger the release of pituitary hormones.

But there is a fatal counterobjection to this objection. While the objection is not wrong, it is incomplete. Yes, depletion and aging of the egg supply are the immediate causes of human menopause, but why did natural selection program women such that their eggs become depleted or unresponsive in their forties? There is no compelling reason why we could not have evolved twice as large a starting quota of eggs, or eggs that remain responsive after half a century. The eggs of elephants, baleen whales, and possibly albatrosses remain viable for at least sixty years, and the eggs of tortoises are viable for much longer, so human eggs could presumably have evolved the same capability.

The basic reason why the third objection is incomplete is because it confuses proximate mechanisms with ultimate causal explanations. (A proximate mechanism is an immediate direct cause, while an ultimate explanation is the last in the long chain of factors leading up to that immediate cause. For example, the proximate cause of a marriage breakup may be a husband's discovery of his wife's extramarital affairs, but the ultimate explanation may be the husband's chronic insensitivity and the couple's basic incompatibility that drove the wife to affairs.) Physiologists and molecular biologists regularly fall into the trap of overlooking this distinction, which is fundamental to biology, history, and human behavior. Physiology and molecular biology can do no more than identify proximate mechanisms; only evolutionary biology can provide ultimate causal explanations. As one simple example, the proximate reason why so-called poison-dart frogs are poi-sonous is that they secrete a lethal chemical named batra-chotoxin. But that molecular biological mechanism for the frogs' poisonousness could be considered an unimportant detail because many other poisonous chemicals would have worked equally well. The ultimate causal explanation is that poison-dart frogs evolved poisonous chemicals because they are small, otherwise defenseless animals that would be easy prey for predators if they were not protected by poison.

We have already seen repeatedly in this book that the big questions about human sexuality are the evolutionary questions about ultimate causal explanation, not the

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