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People from different parts of the world can be distinguished at a glance by so-called racial characteristics. But those same traits - ones such as the colour of our skin and hair and eyes, or the shapes of our breasts and genitalsplay a big role in how we select our mates and sex partners. Thus, our outward appearances and our beauty standards have evolved in tandem to different local end points.

'White man! Lookim this-feller line three-feller man. This-feller number-one he belong Buka Island, na 'nother-feller number-two he belong Makira Island, na this-feller number-three he belong Sikaiana Island. Yu no savvy? Yu no enough lookim straight? I think, eye-belong-yu he bugger-up finish? . No, damn it, my eyes-belong-me were not ruined beyond repair. It was my first visit to the Solomon Islands in the Southwest Pacific, and I told my scornful guide through the medium of pidgin English that I saw perfectly well the differences between those three men in a row over there. The first one had jet-black skin and frizzy hair, the second had niuch lighter skin and frizzy hair, and the third had straighter hair and more slanty eyes. The only thing the matter with me was that I had no experience of what people from each particular Solomon island looked uke. By the end of my first trip through the Solomons, I too could match People to their islands by their skin and hair and eyes. In those variable features, the Solomons are a microcosm of humanity.

Simply by looking at a person, even laymen can often tell what part of the world that person comes from, and trained anthropologists may be able to 'place' him or her in the right part of the right country. For example, given one person each from Sweden, Nigeria, and Japan, none of us would have any trouble deciding at a glance which person was from which country. The most visibly variable features in clothed people are of course skin colour, the colour and form of the eyes and hair, body shape, and (in men) the amount of facial hair. If the people to be identified were undressed, we might also notice differences in amount of body hair, the size and shape and colour of a woman's breasts and nipples, the form of her labia and buttocks, and the size and angle of a man's penis. All those variable features contribute to what we know as human racial variation. Those geographic differences among humans have long fascinated travellers, anthropologists, bigots, and politicians, as well as the rest of us. Since scientists have solved so many arcane questions about obscure unimportant species, surely you might expect them to have answered one of the most obvious questions about ourselves: 'Why do people from different areas look different? Our understanding of how humans came to differ from other animals would remain incomplete if we did not also consider how, in the process, human populations acquired their most visible differences from each other. Nevertheless, the subject of human races is so explosive that Darwin excised all discussion of it from his famous 1859 book On the Origin of Species. Even today, few scientists dare to study racial origins, lest they be branded racists simply for being interested in the problem. There is another reason why we do not understand the significance of human racial variation: it proves to be an unexpectedly difficult problem. Twelve years after Darwin wrote his book attributing the origin of species to natural selection, he wrote another book 898 pages long, attributing the origin of human races to our sexual preferences which I described in the last chapter, and entirely rejecting a role of natural selection. Despite that verbal overkill, many readers were unconvinced. To this day, Darwin's theory of sexual selection (as he called it) remains controversial. Instead, modern biologists generally invoke natural selection to explain the visible differences among human racesespecially the differences in skin colour, whose relation to sun exposure seems obvious. However, biologists cannot even agree on why natural selection led to dark skin in the tropics. I shall explain why I believe natural selection to have played only a secondary role in our racial origins, and why Darwin's preference for sexual selection seems to me correct. I therefore consider visible human racial variation to be largely a byproduct of the remodelled human life-cycle that forms the subject of Part Two of this book.

Firstly, to place matters in perspective, let's realize that racial variation is not at all confined to humans. Most animal and plant species with sufficiently wide distributions, including all higher ape species except the geographically localized pgymy chimp, also vary geographically. So marked is variation in some bird species, such as North America's white-crowned sparrow and Eurasia's yellow wagtail, that experienced birdwatchers can identify an individual bird's approximate birthplace by its plumage pattern. Variation in apes encompasses many of the same characteristics that vary geographically in humans. For example, among the three recognized races of gorillas, western lowland gorillas have the smallest bodies and rather grey or brown hair, while mountain gorillas have the longest hair, and eastern lowland gorillas share black hair with mountain gorillas. Races of white-handed gibbons similarly vary in hair colour (variously black, brown, reddish, or grey), hair length, tooth size, protrusion of the jaws, and protrusion of the bony ridges over the eyes. All these traits that I have just mentioned as varying among gorilla or gibbon populations also differ among human populations. How does one decide whether recognizably distinct animal populations from different localities constitute different species, or belong instead to the same species and just constitute different races (also known as subspecies)? As explained in Chapter Two, the distinction is based on interbreeding under normal circumstances: members of the same species may interbreed normally if given the opportunity, while members of different species do not. (But closely related species that would not normally interbreed in the wild, like lions and tigers, may do so if a male of one is caged with a female of the other and given no other choice.) By this criterion, all living human populations belong to the same species, since some interbreeding has occurred whenever humans from different regions have come into contacteven people as dissimilar in appearance as African Bantus and Pygmies. With humans as with other species, populations may intergrade into each other, and it becomes arbitrary to decide which populations to group as races. By the same criterion of interbreeding, the large gibbons known as siamangs are a distinct species from the smaller gibbons, since both occur together in the wild without hybridizing. This is also the criterion for considering Neanderthals possibly as a species distinct from Homo sapiens, since hybrid skeletons have not been identified despite apparent Cro-Magnon/Neanderthal contact (see Chapter Two).

Racial variation has characterized humans for at least the past several thousand years, and possibly much longer. Already around 450 BC, the Greek historian Herodotus described the Pygmies of West Africa, the black-skinned Ethiopians, and a blue-eyed red-haired tribe in Russia. Ancient paintings, mummies from Egypt and Peru, and bodies of people preserved in European peat bogs confirm that people several thousand years ago differed in their hair and facial features much as they do today. Origins of modern races can be pushed back still further, to at least ten thousand years ago, since fossil skulls of that age from various parts of the world differ in many of the same respects that modern skulls from the same regions differ. More controversial are the studies of some anthropologists, contested by others, reporting continuity of racial skull characteristics for hundreds of thousands of years. If those studies are correct, then some of the human racial variation that we see today may predate the Great Leap Forward, and may have gone back to the times of Homo erectus. Now let's turn to the question of whether natural selection or sexual selection has made the larger contribution to those visible geographic differences of ours. Take first the arguments about natural selection, the selection of traits that enhance survival. No scientist denies today that natural selection does account for many of the differences between species, such as why lions have paws with claws while we have grasping fingers. No one denies either that natural selection explains some geographic variation ('racial variation') within some animal species. For instance, Arctic weasels that live in areas covered by winter snow change colour from brown in summer to white in winter, while more southerly weasels stay brown all year. That racial difference enhances survival, because white weasels against a brown background would be glaringly conspicuous to their prey if they were not camouflaged against snow.

By the sajne token, natural selection surely explains some geographic variation in humans. Many black Africans but no Swedes have the sickle-cell haemoglobin gene, because the gene protects against malaria, a tropical disease that would otherwise kill many Africans. Other localized human traits that surely evolved through natural selection include the big chests of Andean Indians (good for extracting oxygen from thin air at high altitudes), the compact shapes of Eskimos (good for conserving heat), the slender shapes of southern Sudanese (good for losing heat), and the slit-like eyes of northern Asians (good for protecting eyes against cold and against sun glare off the snow). All these examples are easy to understand. Can natural selection similarly explain the racial differences that we think of first, those in skin colour and eye colour and hair? If so, one might expect that the same trait (for instance, blue eyes) would reappear in different parts of the world with similar climates, and that scientists would agree on what the trait is good for.

Seemingly the simplest trait to understand is skin colour. Our skins run the spectrum from various shades of black, brown, copper, and yellowish to pink with or without freckles. The usual story to explain this variation by natural selection goes as follows. People from sunny Africa have blackish skins. So too (supposedly) do people from other sunny places, like southern India and New Guinea. Skins are said to get paler as one moves north or south from the equator, until one reaches northern Europe, with the palest skins of all. Obviously, dark skins evolved in those people who were exposed to much sunlight. That is just like the skins of whites tanning under the summer sun (or in tanning salons!), except that tanning is a reversible response to sun rather than a permanent genetic one. It is equally obvious what good a dark skin does in sunny areas: it protects against sunburn and skin cancer. Whites who spend lots of time outdoors in the sun tend to get skin cancer, and they get it on exposed parts of their body like their head and hands. Does that not all make sense?

Yes, but it is really not so simple at all. To begin with, skin cancer and sunburn cause little debilitation and few deaths. As agents of natural selection, they have an utterly trivial impact compared to infectious diseases of childhood. Hence many other theories have been proposed to explain the supposed pole-to-equator gradient in skin colour.

One favourite competing theory notes that the sun's ultraviolet rays promote vitamin D formation in a layer of our skin beneath the main pigmented layer. Thus, people in sunny tropical areas might have evolved dark skin to protect them against the risk of kidney disease caused by too much vitamin D, while people in Scandinavia with its long dark winters evolved pale skins to protect them against the risk of rickets caused by too little vitamin D. Two other popular theories are that dark skins are to protect our internal organs against overheating by the tropical sun's infrared rays, orjust the oppositedark skins help keep tropical people warm when the temperature drops. And if those four theories are not enough for you, consider four more: that dark skins provide camouflage in the jungle, or that pale skins are less sensitive to frostbite, r that dark skins protect against beryllium poisoning in the tropics, or that pale skins cause deficiency of another vitamin (folic acid) in the tropics.

With at least eight theories in the running, we can hardly claim to understand why people from sunny climates have dark skins. That in rtself does not refute the idea that, somehow, natural selection caused the evolution of dark skins in sunny climates. After all, dark skins could have multiple advantages, which scientists may sort out some day. Instead, the heaviest objection to any theory based on natural selection is that the association between dark skins and sunny climates is a very imperfect one. Native peoples had very dark skins in some areas receiving relatively little sunlight, like Tasmania, while skin colour is only medium in sunny areas of tropical Southeast Asia. No American Indians have black skins, not even in the sunniest parts of the New World. When one takes cloud cover into account, the world's most dimly lit areas, receiving a daily average of under three-and-a-half hours of sunlight, include parts of equatorial West Africa, southern China, and Scandinavia, inhabited respectively by some of the world's blackest, yellowest, and palest peoples! Among the Solomon Islands, all of which share a similar climate, jet-black people and lighter people replace each other over short distances. Evidently, sunlight has not been the sole selective factor that moulded skin colour.

The first response of anthropologists to these objections is to raise a counter-objection, the time factor. This argument tries to explain away the cases of pale-skinned people in the tropics by claiming that those particular peoples migrated to the tropics too recently to have evolved black skins. For example, the ancestors of American Indians may have reached the New World only 11,000 years ago (Chapter Eighteen): perhaps that has not been long enough to evolve black skins in the tropical Americas. But if you are going to evoke the time factor to explain away objections to the climate theory of skin colour, then you also have to consider the time factor for peoples who supposedly support that theory. One of the prime supports of the climate theory is the pale skin of Scandinavians, living in the cold, dark, foggy North. Unfortunately, Scandinavians have been in Scandinavia for an even shorter time than American Indians have been in the Amazon. Until about 9,000 years ago, Scandinavia was covered by an ice-sheet and could hardly have supported any people, pale-skinned or dark-skinned. Modern Scandinavians reached Scandinavia only around 4,000 or 5,000 years ago, as a result of the expansion of farmers from the Near East (Chapter Ten) and of Indo-European speakers from southern Russia (Chapter Fifteen). Either Scandinavians acquired their pale skins long ago in some other area with a different climate, or else they acquired them in Scandinavia within half the time that Indians have spent in the Amazon without becoming dark-skinned.

The sole people in the world about whom we can be certain that they spent the last 10,000 years in the same location were the natives of Tasmania. Lying south of Australia, at the temperate latitude of Chicago or Vladivostok, Tasmania used to be connected to Australia until it was cut off by rising sea levels 10,000 years ago and became an island. Since modern Tasmanian natives did not have boats capable of going more than a few miles, we know that they were derived from colonists who walked out to Tasmania at the time of its connection to Australia, and who remained there continuously until they were exterminated by British colonists in the Nineteenth Century (Chapter Sixteen). If any people had enough time for natural selection to match their skin colour to their local temperate-zone climate, it was the Tasmanians. Yet they had blackish skins, supposedly adapted to the Equator.

If the case for natural selection of skin colour seems weak, that for hair colour and eye colour is virtually non-existent. There are no consistent correlations with climate, and not even any half-plausible theories for the supposed advantage lent by each colour type. Blonde hair is common in cold, wet, dimly lit Scandinavia and also among Aborigines of the hot, dry, sunny desert of central Australia. What do those two areas have in common, and how does being blonde help both Swedes and Aborigines to survive? Do freckles and red hair help Irishmen catch leprechauns? Blue eyes are common in Scandinavia and supposedly help their owners see farther in dim, misty light, but that speculation is unproven, and all my friends in the even dimmer, mistier mountains of New Guinea see just fine with their dark eyes.

The racial traits for which it seems most absurd to seek an explanation based on natural selection are our variable genitalia and secondary sex characteristics. Are hemispherical breasts an adaptation to summer rainfall and conical breasts an adaptation to winter fog, or vice versa? Do the protruding labia minora of Bushmen women protect them against pursuing lions, or reduce their water losses in the Kalahari Desert? You surely don't think that men with hairy chests can thereby keep warm while going shirtless in the Arctic, do you? If you do think so, then please explain why women do not share hairy chests with men, since women also have to keep warm. Facts such as these were what made Darwin despair of imputing human racial variation to his own concept of natural selection. He finally Qismissed the attempt with a succinct statement: 'Not one of the external differences between the races of man are of any direct or special service to him. When Darwin came up with a theory that he preferred, he termed it sexual selection' to contrast with natural selection, and he devoted an entire book to explaining it. The basic notion behind this theory is easily grasped. Darwin noted many animal features that had no obvious survival value but that did play an obvious role in securing mates, either by attracting an individual of the opposite sex or by intimidating a rival of the same sex. Familiar examples are the tails of male peacocks, the manes of male lions, and the bright red buttocks of female baboons in oestrus. If an individual male is especially successful at attracting females or intimidating rival males, that male will leave more descendants and will tend to pass on his genes and traitsas a result of sexual selection, not natural selection. The same argument applies to female traits as well.

For sexual selection to work, evolution must produce two changes simultaneously: one sex must evolve some trait, and the other sex must evolve in tandem a liking for that trait. Female baboons could hardly afford to flash red buttocks if the sight revolted male baboons to the point of their becoming impotent. As long as the female has it and the male likes it, sexual selection could lead to any arbitrary trait, just as long as it does not impair survival too much. In fact, many traits produced by sexual selection do seem quite arbitrary. A visitor from outer space who had yet to see humans could have no way of predicting that men rather than women would have beards, that the beards would be on the face rather than above the navel, and that women would not have red and blue buttocks.

That sexual selection really can work, at least in birds, was proved by an elegant experiment carried out by the Swedish biologist Make Andersson on the long-tailed widowbird of Africa. In this species the male's tail in the breeding season grows to 20 inches long, while the female's tail is only 3 inches. Some males are polygamous and acquire up to six mates, at the expense of other males who get none. Biologists had guessed that a long tail served as an arbitrary signal by which males attracted females to join their harem. Andersson's test was to cut off part of the tail from nine males until their tails were only 6 inches long. He then glued those cut segments to the tails of nine other males to give them 30-inch tails, and he waited to see where the females built their nests. It turned out that the males with the artificially lengthened tails attracted on the average over four times as many mates as the males with artificially shortened tails. Perhaps our first reaction to Andersson's experiment is: those dumb birds! Imagine a female selecting a particular male to father her offspring merely because his tail is longer than other males' tails! But before we get too smug, let's consider again what we learned in the last chapter about how we humans select our own mates. Are our criteria such good indicators of genetic worth? Do not some men and women set disproportionate value on the size or form of certain body parts, which are really nothing more than arbitrary signals for sexual selection? Why did we evolve to pay any attention at all to a beautiful face, which is useless to its owner in the struggle for survival?

In animals some of the traits that vary racially are ones produced by sexual selection. For instance, lions' manes vary in length and in colour. Males of the astrapia birds of paradise in New Guinea have fancy tails to display to females, but different populations evolved tails of different shapes and colours. From west to east, the tails are broad and purple, short and white-based, very long and white, long and purple, and broad and purple again. Similarly, snow geese occur in two colour phases, a blue phase commoner in the western Arctic and a white phase commoner in the eastern Arctic. Birds of each phase prefer a mate of the same phase. Could human breast shape and skin colour similarly be the outcome of sexual preferences that vary arbitrarily from area to area? After 898 pages Darwin convinced himself that the answer to this question was a resounding 'yes'. He noted that we pay inordinate attention to breasts, hair, eyes, and skin colour in selecting our mates and sex partners. He noted also that people in different parts of the world define beautiful breasts, hair, eyes, and skin by what is familiar to them. Thus, Fijians, Hottentots, and Swedes each grow up with their own learned, arbitrary beauty standards, which tend to maintain each population in conformity with those standards, since individuals deviating too far from the standards would find it harder to obtain a mate. Darwin died before his theory could be tested against rigorous studies of how people actually do select their mates. Such studies have proliferated in recent decades, and I summarized the results in Chapter Five. There I showed that people tend to marry individuals who resemble themselves in every conceivable character, including hair and eye and skin colour. To explain that seeming narcissism of ours, I reasoned that we develop our beauty standards by imprinting on the people we see around us in childhoodespecially on our parents and siblings, the people of which we see the most. But our parents and siblings are also the people to whom we bear the strongest physical resemblance, since we share their genes. Thus, if you are a fair-skinned, blue-eyed blonde who grew up in a family of fair-skinned, blue-eyed blondes, that is the sort of person whom you will consider most beautiful and will seek as a mate. In the meantime, my dark-skinned, dark-haired New Guinea friends were growing up with other New Guineans and learning to regard fair-skinned, blue-eyed blondes as grotesquely revolting.

To test that imprinting theory of human mate choice rigorously, one would have to do experiments like shipping some Swedish babies to adoptive parents in New Guinea, or painting some Swedish parents permanently black. Then, after waiting twenty years for the babies to grow up, one could study whether they preferred Swedes or New Guineans as sex partners. Alas, once again, the Search for Truth about humans founders on practical problems, but such tests can be performed with full experimental rigour on animals. Take snow geese, for example, with their blue or white colour phases. Do white geese learn or inherit their preference in the wild for white geese over blue ones? Canadian biologists hatched gosling eggs in an incubator, then put the goslings into a nest of goose 'foster-parents'. When those goslings grew up, they chose a mate with the colour of the foster-parents. When goslings were reared in a large mixed flock of both blue and white birds, they showed no preference between blue and white prospective mates on reaching adulthood. Finally, when the biologists dyed some white parents pink, their offspring came to prefer pink-dyed geese. Thus, geese do not inherit but learn a colour preference, by imprinting on their parents (and on their siblings and playmates). How, then, do I think that people in different parts of the world evolved their differences? Our insides remained invisible to us and were moulded only by natural selection, with results such as that of tropical Africans but not Swedes evolving the anti-malarial defence of a sickle-cell haemoglobin gene. Many visible features of our outsides also got moulded by natural selection. But, just as in animals, sexual selection had a big effect in moulding the external traits by which we pick our mates.

For us humans those traits are especially the skin, eyes, hair, breasts, and genitals. In each part of the world those traits evolved in tandem with our imprinted aesthetic preferences to reach different, somewhat arbitrary results. Which particular human population ended up with any given eye or hair colour may have been partly an accident of what biologists term the 'founder effect'. That is to say, if a few individuals colonize an-empty land and their descendants then multiply to fill the land, the genes of those few founding individuals may still dominate the resulting population many generations later. Just as some birds of paradise ended up with yellow plumes and others with black plumes, so some human populations ended up with yellow hair and others with black hair, some with blue eyes and others with green eyes, some with orange nipples and others with brown nipples.

I do not mean thereby to claim that climate has nothing whatsoever to do with skin colour. I acknowledge that tropical peoples tend on the average to have darker skins than temperate-zone peoples, though there are many exceptions, and that this is probably due to natural selection, though we are unsure of the exact mechanism. Instead, I am saying that sexual selection has been strong enough to render the correlation between skin colour and sun exposure quite imperfect.

If you are still sceptical about how traits and aesthetic preferences can evolve together to different and arbitrary end points, just think about our changing fashion preferences. When I was a schoolboy in the early 1950s, women rated men with crew-cuts and clean-shaven faces a's handsome. Since then, we have seen a parade of men's fashions, including beards, long hair, earrings, purple-dyed hair, and the Mohawk hair style. A man daring to flaunt any of those fashions in the 1950s would have revolted the girls and enjoyed zero mating success. That is not because crew-cuts were better adapted to atmospheric conditions of Stalin's last years, while a purple Mohawk has higher survival value in our post-Chernobyl era. Instead, men's appearances and women's tastes changed in tandem, and the changes occurred far more rapidly than evolutionary changes in skin colour, since no gene mutations were required. Either women came to like crew-cuts because good men had them, or men adopted crew-cuts because good women liked them, or something of both happened. The same goes for women's appearances and men's tastes.

To a zoologist, the visible geographic variability that sexual selection produced in humans is impressive. I have argued in this chapter that much of our variability is a by-product of a distinctive feature of the human life-cycle, our choosiness with respect to our spouses and sex partners. I do not know of any other wild animal species in which eye colour of different populations can be green, blue, grey, brown, or black, while skin colour varies geographically from pale to black and hair is either red, yellow, brown, black, grey, or white. There may be no limits, except those imposed by evolutionary time, on the colours with which sexual selection can adorn us. If humanity survives another 20,000 years, I predict that there will be women with naturally green hair and red eyesplus men who think such women are the sexiest.

FIVE HOW WE PICK OUR MATES AND SEX PARTNERS | The rise and fall of the third chimpanzee | SEVEN WHY DO WE GROW OLD AND DIE?