It’s possible, we submit, to carry the fear of anthropomorphism too far. There are excesses worse than a surfeit of sentiment. There must be some interior state, some thoughts and feelings among the monkeys and apes, and if they are genetically our close relatives, if their behavior is so similar to ours as to be familiar, it’s not unreasonable to attribute to them feelings similar to ours as well. Of course, until better communication with them is established, or until we understand much more about how their brains and hormones work, we can’t be sure. But it’s plausible, it’s an effective teaching tool, and in this book on a few occasions we attempt to portray what it might be like inside the head of another animal.

——


By now, the reader will have at least suspected that the interior monologues of the preceding chapter—the first and third by a middle-ranking female, the second by a high-ranking male—are not intended to refer, exactly, to humans. Instead, we’ve tried to depict what it’s like to be a chimp in chimp society. Systematic, long-term observation of chimp groups in the wild is a new field of science. We’ve relied chiefly on the courageous, insightful, and pioneering work of Jane Goodall at the Gombe Reservation in Tanzania, as well as studies by Toshisada Nishida and his colleagues in the Mahale Mountains, also in Tanzania, and by Frans de Waal, who investigated a troop of chimpanzees in a two-acre enclosure in the Arnhem Zoo in the Netherlands.25 Every event dramatized in the last chapter is based on the accounts of these scientists. Their observations speak to us of a way of life that is unmistakably familiar, rich with the Sturm und Drang of human relations. Of course, no human has ever been inside a chimp’s mind, and we cannot be sure how they think. We have taken liberties. We make no apologies for doing so, but stress that it is intended only as a way to think about the chimps.

We must be careful of circular reasoning here—foisting human mental and emotional processes on the chimps, and then triumphantly concluding at the end of our narrative how much like us they are. If we’re to understand ourselves better by looking closely at chimpanzees, we’ll have to give great weight to what they do and comparatively little to what we imagine is going on inside their heads. We must be careful not to deceive ourselves. The behaviorists were not wholly misguided.

We didn’t mention that chimps sleep in trees and that they spend a great deal of time grooming one another. Although chimps do not seem as much transfixed by oral sex as some other primates (cunnilingus is an almost invariable part of foreplay among the orangs26), we used the now-popular phrase to “suck up” to someone because it seems to us, in its present English-language associations at least, to approximate some of the nuance of chimpanzee submission. (The gestural vocabulary of chimpanzee submission does include kissing the alpha’s thigh.)

Many behavioral differences exist between chimps and humans, just as between chimps and gorillas or between gibbons and orangutans. But we are struck by how much the core of chimpanzee social life in the wild resembles some forms of human social organization, especially under great stress—in prisons, say, or urban and motorcycle gangs, or crime syndicates, or tyrannies and absolute monarchies. Niccolò Machiavelli, chronicling the maneuvering necessary to get ahead in the seamy politics of Renaissance Italy—and shocking his contemporaries, especially when he was honest—might have felt more or less at home in chimpanzee society. So might many dictators, whether they style themselves of the right or left persuasion. So might many followers. Beneath a thin varnish of civilization, it sometimes seems, there’s a chimp struggling to bust out—to take off the absurd clothes and the restraining social conventions and let loose. But this is not the whole story.

They’re a little shorter, somewhat hairier, much stronger, and a lot more sexually active than most humans are. They have brown hair and brown eyes. In their natural habitats, they may live to be forty or fifty years old—which is longer than the average in any human society before the Industrial and Medical revolutions. But their average life expectancy is much less. Unlike modern humans, females past infancy are not likely to live as long as males. They alternate between walking on two feet and on all fours, using their knuckles. Chimpanzee males tend to have short fuses. They give off a faint but characteristic odor when they’re nervous or excited, revealing emotions they sometimes try to hide. Chimps are not ashamed of displaying their sexual parts. By our lights they’re a lot dumber than we are, but they do use and even make tools. They apparently hold grudges, nurse resentments, and harbor thoughts of revenge. They plan future courses of action.

Family ties may be strong and lasting. Aged mothers will rush to the defense of their children, even full-grown sons. Orphaned infants are tenderly raised by older siblings. They experience prolonged grief at the loss of a loved one. They suffer from bronchitis and pneumonia, and can be infected with almost any human disease, including the AIDS virus. The elderly turn gray, get wrinkles, lose teeth and hair. Chimps get drunk. They’re able to learn more words of a human language than we have of any chimp language. When they look in the mirror, they recognize themselves. They are, at least to some degree, self-aware. Infants get cranky and irritable when they’re weaned. Chimps form friendships, often with comrades-in-arms who hunt together and guard their turf against intruders. They share food with relatives and friends.

When raised among humans, they have been known to masturbate to pictures of naked people. (This is probably true only of those who, through prolonged contact, have come to consider themselves human. Wild chimps would no more masturbate to erotic images of humans than vice versa.) They keep secrets. They lie. They both oppress and protect the weak. Some, despite many setbacks, persistently strive for social advancement and career opportunities. Others, less ambitious, are more or less content with their lot.

Among much other innate knowledge, they are born with an understanding about how to make a bed of leaves each night up there in the trees. They are much better climbers than we, partly because they haven’t lost, as we have, the ability to grasp branches with their feet. The youngsters love to climb trees and rival one another in spectacular feats of gymnastic derring-do. But when an infant has climbed too high, its mother—socializing with her friends at the base of the tree—decisively taps the trunk and the baby obediently scampers down.

The forest is crisscrossed with a network of trails made by generations of chimps going about their daily business. Each knows the local geography at least as well as the average human city-dweller knows the neighborhood streets and shops. They almost never get lost. Here and there along the trails are trees with acoustically resonant trunks. When a party of foragers spies such a tree, many run forward and drum away—both sexes, children as well as adults. There are no strings, woodwinds, or brass yet, but the percussion section is in place.

Chimps recognize one another’s individual voices, and a distinctive pant-hoot may summon an ally or relative from a considerable distance. In answering a pant-hoot from, say, an adjacent valley, they lift their heads and purse their lips as if they were on stage at La Scala. Up close, they have an uncanny ability—“uncanny” means only that we haven’t been smart enough to figure it out yet—to communicate with one another, not just about such straightforward matters as sex or dominance, but about much more subtle matters, such as hidden dangers, or buried food supplies. A classic set of experiments was done by the psychologist E. W. Menzel:[Menzel] maintained four to six young chimpanzees in a large outdoor enclosure that was also connected to a smaller holding cage. He restrained all but one animal in the holding cage, while showing this chosen “leader” the hidden location of either an amount of food or an aversive stimulus such as a stuffed snake. The leader was then returned to the holding cage, and the whole group was released. According to Menzel’s reports, the variable behavior of the animals indicated that they “seemed to know approximately where the object was, and what sort of object it was, long before the leader reached the spot where it had been hidden” … If the goal was food, they ran ahead looking in possible hiding places; if it was a stuffed alligator or snake, they emerged from their cage showing piloerection [their hair standing on end] and staying close to their companions. If the hidden item was an alligator or snake, they became very cautious in their approach and often mobbed the area, hooting in the direction of the hidden item and hitting at it with sticks. If the hidden item was food, the animals searched the area intensively and showed little fear or distress. The behaviors occurred even if the aversive stimulus had been removed before the animals were released from the holding cage, so it was not the item itself that produced these reactions.In the food tests, one male (Rocky) began to monopolize the food supply when it was located. When Belle, a female, served as leader, she attempted to avoid indicating the location of the food cache, but Rocky could often extrapolate from her line of orientation and find the food. If Belle were shown two caches, one large and one small, she would lead Rocky to the small one and, while he was busy eating, run to the larger one which she would share with other individuals. Menzel concluded that chimpanzees could communicate the direction, amount, quality, and nature of the goal, as well as attempt to conceal at least some of this information, but precisely how chimpanzees achieve such communication is still not known.27


The only possibilities seem to be gestures and speech.

Chimps have hundreds of different kinds of food and crave dietary variety. They eat fruit, leaves, seeds, insects, and larger animals, sometimes dead ones. Caterpillars are delicacies, and the discovery of an infestation of caterpillars becomes a memorable gastronomic event. They’re known to eat soil from cliff faces, presumably to provide mineral nutrients such as salt. Mothers will offer choice tidbits of food to their infants and will snatch unusual, possibly dangerous, foods from their mouths. In the wild, adults share food occasionally, often in response to begging. There are no set mealtimes; they snack throughout the day. As a foraging party moves on, one of its members may carry a branch still laden with berries or leaves to munch as she rambles.

When in the middle of the night, in their beds of leaves in the high branches, they are awakened by the cries of predators, they clutch each other in fright, their urine and feces raining down on the forest floor below.

They love to play, children (whose energy is prodigious) more than adults, but even adult play is common—especially when there’s enough to eat and large numbers of chimps gather together. Play often involves, but is not restricted to, mock fighting.

Chimp males are protective toward females and the young. They will readily risk their own lives to protect “women and children” from attack, or to rescue a youngster in trouble. Goodall writes, “Often it seems that a male cannot resist reaching out to draw an infant into a close embrace, to pat him, or to initiate gentle play.”28 When a male is discovered in flagrante delicto with a female, which is often, an infant may rush up to punch the male in the mouth or jump on the back of the female, most often his mother.* In such situations the male’s tolerance frequently exceeds human limits.

But in a display competition for dominance, all this good-natured equanimity vanishes, and a male who ordinarily is protective of infants may pick up a small, innocent bystander and slam it to the ground in his rage. When an unfamiliar female is discovered in their territory, chimps are known to seize her infant by the ankles and smash it against the rocks.29

Chimps tend to pick on the runt of the litter, and to displace their own anger away from higher-ranking chimps (who might do them harm) to those who are milder-tempered, younger, weaker, and female. At Gombe in 1966 there was a polio epidemic which resulted in the partial paralysis of full-fledged members of the group. Crippled by their disease, they were forced to move in odd ways, dragging limbs. Other chimps were at first afraid; then they threatened the afflicted, and then attacked them.

Because aggression is episodic and friendly relations so much more common, some early field observers were tempted by the notion that chimps in a state of nature (that is, unimprisoned) are non-violent and peace-loving. This is not the case. In hunting other animals, in working the dominance hierarchy, in hustling the females, in peevish moments, and in skirmishes with other groups of chimps (the Strangers, in our narrative), they show themselves capable of great violence.

Meat contains essential amino acids and other molecular building blocks more difficult to acquire from plants. Both sexes are ravenous for meat. On rare occasions, females will attack other females in their group and steal and eat their infants. Once the little one is in hand, there are no ill feelings directed to the mother of the tiny victim. In one case, a female approached those who were eating her baby; one of the diners responded by putting out her arms to embrace and comfort the grieving mother. Chimps are known to hunt mice, rats, small birds, a twenty-kilogram adolescent bush pig, monkeys such as baboons and colubuses, and other chimps.

A successful hunt is accompanied by enormous excitement. The spectators scream, hug, kiss, and pat one another reassuringly. Those actually involved in the kill immediately begin feeding, or attempt to carry off tasty body parts. The forest is filled with screeches, barks, pants, and hoots—which attract additional chimps, sometimes from a considerable distance. Generally males help themselves to bigger portions than females. Those of high rank are more likely to distribute the spoils, and one way or another most who are actually present at the kill gain a share. Newcomers plead for morsels. Pieces will be stolen, and the chimp whose prize has been taken will be furious, perhaps indulging in a temper tantrum. Portions of meat are taken to bed for midnight snacks.

A rat may be eaten head first. A monkey or young antelope is often killed by having its head smashed against a rock or tree trunk, or by giving it a vampirish bite in the back of the neck. Almost always the brains are eaten first. This is often the prize of the hunter who performs the actual kill. Other tasty body parts include the genitals of male victims and the fetuses of pregnant female victims. Goodall reports the final, attenuated, scream of a young bush pig as a chimp, like some ancient Aztec priest, tore out its living heart. Cooking has not yet been invented, nor flatware, nor table manners, nor squeamishness. This is a world of red blood and raw meat.

Janis Carter describes30 a juvenile chimp and a colubus monkey, about its own size, grooming one another; but when the colubus is seized by the tail and killed by a passing adult chimp, who bashes its head against a tree, the juvenile readily enough joins in devouring its erstwhile playmate. Most of the monkey (and small mammal) victims of chimp predation are infants and juveniles, often snatched from their mothers’ arms. Sometimes the mother tries to rescue the infant and is herself eaten.

In this world there is no mercy shown to food, even if it’s still ambulatory. Food is for eating. Those who are moved to mercy eat less and leave fewer offspring. Clearly the chimps do not recognize monkeys, or chimps of other groups, or even members of their own group as deserving of mercy or other moral considerations. They may be heroic in defending their own young, but they do not show the least compassion for the young of other groups of species. Perhaps they consider them “animals.”

Hunting is a cooperative endeavor. Cooperation is essential for making the larger kills—and also for avoiding their dangers, such as an enraged bush pig charging, tusks first, to save its young. The hunters exhibit real teamwork. One chimp may softly call to another when it has detected prey in the underbrush. They smile to one another. The victim is flushed out of its cover toward other chimps who are lying in wait. Escape routes are blocked off. Ambushes are refined. Plays are called. The chimps—so passionate after the kill—were coolly planning it all out beforehand.

——


In densely forested habitats, the territory controlled by a given chimpanzee group is only a few kilometers wide. In sparsely wooded regions, it can be as much as thirty kilometers across. These are the territories that a chimp group considers its turf, its home, its fatherland or motherland, to which something like patriotic sentiments are owed. It is not to be trespassed by strangers. It’s a jungle out there. The typical day range of a chimp combat patrol is a few kilometers. So if they live in heavy forest, they can fairly readily patrol a good portion of the border in a single day. But if the vegetation and food supply are more sparse and their territory accordingly larger, it may be a few days’ journey from one end to the other, and longer if they go around the perimeter.A patrol is typified by cautious, silent travel during which the members of the party tend to move in a compact group. There are many pauses as the chimpanzees gaze around and listen. Sometimes they climb tall trees and sit quietly for an hour or more, gazing out over the “unsafe” area of a neighboring community. They are very tense and at a sudden sound (a twig cracking in the undergrowth or the rustling of leaves) may grin and reach out to touch or embrace one another.During a patrol the males, and occasionally a female, may sniff the ground, treetrunks, or other vegetation. They may pick up and smell leaves, and pay particular attention to discarded food wadges, feces, or abandoned tools on termite heaps. If a fairly fresh sleeping nest is seen, one or more of the adult males may climb up to inspect it and then display around it so that the branches are pulled apart and it is partially or totally destroyed.Perhaps the most striking aspect of patrolling behavior is the silence of those taking part. They avoid treading on dry leaves and rustling the vegetation. On one occasion vocal silence was maintained for more than three hours … [When] patrolling chimpanzees return once more to familiar areas, there is often an outburst of loud calling, drumming displays, hurling of rocks, and even some chasing and mild aggression between individuals … Possibly this noisy and vigorous behavior serves as an outlet for the suppressed tension and social excitement engendered by journeying silently into unsafe areas.31


In this description by Jane Goodall of a patrol at Gombe, we are taken by the ability of the chimps to overcome their fear, to exercise self-control by inhibiting their usual noisy interchanges, but particularly by their deductive abilities. These chimps are tracking. They are weighing the evidence of branches, footprints, droppings, artifacts. We might expect that, when food is in short supply, group differences in tracking skills help determine who lives and who dies. Not just strength and aggressiveness are being selected here, but something akin to reasoning and quick-wittedness. And stealth. When one human who lived with a troop for a long time tried to accompany a patrol as it set out, they looked at him reprovingly. He was just too clumsy. He could not, as they do, slip silently through the forest.

So the long-range combat patrol wends its way toward the borders of their turf If it’s more than a day’s walk, they’ll set up camp at night and continue on their way tomorrow. What happens if they encounter members of another group, Strangers from the adjacent territory? If it’s just one or two intruders, they’ll attempt to attack and kill them. There’s much less disposition here toward threat displays and intimidation. But if two parties of roughly equal strength encounter one another, now there are a great many threat displays, rocks and sticks are thrown, trees are drummed. “Somebody hold me back, I’m gonna break his knees,” you can almost hear them saying. They practice threat assessment If the patrol senses an obviously larger number of Strangers, it is likely to beat a hasty retreat. At other times chimp patrols may penetrate enemy territory or even raid its populated core area—for many purposes, including copulating with unfamiliar females. The combination of tracking, stealth, danger, teamwork, fighting hated enemies, and the opportunity for sex with strange females is enormously attractive to the males.

The delight shown by the members of a patrol after having successfully returned from dangerous—perhaps enemy-held—territory is little different from what happens when chimps unexpectedly encounter a substantial cache of food. They screech, kiss, hug, hold hands, pat one another on the shoulders and the rump, and jump up and down. Their camaraderie is reminiscent of teammates in mutual embrace just after winning the national title At the start of a heavy rain, male chimps often perform a spectacular dance. On coming upon a stream or waterfall they display, seize vines, swing from one tree to another, and cavort high above the water in a breathtaking acrobatic performance that may last for ten minutes or more. Perhaps they are awed by the natural beauty or entranced by the white noise. Their evident joy sheds a revealing light on the eighteenth-century doctrine32 that humans are right to enslave other animals because we are unmatched in our capacity to be happy.

The prescription offered by Sewall Wright for a successful evolutionary response to a changing environment closely matches many aspects of chimp society. The species is divided into free-ranging groups, generally comprising between ten and one hundred individuals. They have different territorial ranges, so that if the environment alters the impact will be at least slightly different from group to group. A staple food at one end of a vast tropical forest may be a rare delicacy at the other. A blight or infestation that might result in serious malnutrition or famine for chimps in one part of the forest might have negligible consequences in another. Each territorial group is enough inbred that the gene frequencies differ systematically, group to group. And yet the pattern of inbreeding is relieved by exogamy (outbreeding). There are key sexual encounters with chimps from adjacent territories, initiated either when a patrol penetrates into alien territory or when a foreign female wanders over. These unions provide genetic communication, group to group, so that if in an adaptive crisis one group were more fit than the others, the adaptation would rapidly spread to the entire chimpanzee population through a sequence of sexual contacts—perhaps hundreds of copulations in a chain linking the remotest groups of a vast tropical forest. If there’s a modest environmental crisis, the chimpanzees are ready.

If this is indeed the explanation, at least in part, of the territoriality, ethnocentrism, xenophobia, and occasional exogamy that characterize chimpanzee society, we do not imagine that individual chimps understand the reasons for their behavior. They simply can’t stand the sight of strangers, find them hateful and deserving of attack—except, of course, for the chimps of the opposite sex, who are unaccountably exciting. The females occasionally run away with strange males, no matter what crimes they may earlier have committed against their land and kin. Perhaps they feel something of what Euripides makes Helen of Troy feel:What was there in my heart, that I forgotMy home and land and all I loved, to run awayWith a strange man?…


Ah, husband still, how shall thy hand be bentTo slay me? Nay, if Right be come at last,What shalt thou bring but comfort for pains past,And harbour for a woman storm-driven:A woman borne away by violent men …33


Mothers know who their sons are and so can preferentially resist their (very rare) sexual advances. But fathers are not sure who their daughters are, and vice versa. Thus, when a female comes of age in a small group, there’s a significant chance of an incestuous union, further inbreeding, more infant mortality, and fewer of their genetic sequences passed on to future generations. So around the time of first ovulation, a female often feels an inexplicable urge to visit the neighboring territory. This can be a dangerous undertaking, as she probably understands full well. The compulsion, then, must be strong, which in turn underscores the evolutionary importance of her mission. Combine this not uncommon itch to wander at first ovulation with the rarity of brother-sister and, especially, mother-son unions and it’s clear that a high-priority, well-functioning incest taboo is operating among the chimps.

There’s one aspect of chimpanzee territoriality not shared by other apes—all of whom are divided into territorial, xenophobic groups, with a little exogamy thrown in: Unlike encounters within the group, where bluff and intimidation play major roles and only rarely does anyone get seriously hurt, when two chimp groups interact there can be real violence. No main force combat has ever been observed. They prefer guerrilla tactics. One group will pick off the members of the other in ones and twos until there’s no longer a viable force left to defend the adjacent territory. Chimpanzee groups are constantly skirmishing to see if it’s possible to annex more turf. If the penalty for failure in combat is death for the males and alien sexual bondage for the females, the males soon find themselves caught up in a powerful selection for military skills. Genes for those skills must have been racing through the tropical forests, by exogamous mating, until nearly all chimps had them. If they didn’t, they died.

Moreover, the skills that make you good on patrol and good in skirmishes also make you good in the hunt. If your combat skills are honed, you can also supply your friends, loved ones, and concubines—to say nothing of yourself—with more of that delicious red meat. Except for the part about the good eating, being a male chimp is a little like being in the army.


* “[An ape’s] face resembles that of a man in many respects … [I]t has similar nostrils and ears, and teeth like those of man, both front teeth and molars … [I]t has hands and fingers and nails like man, only that all these parts are somewhat more beastlike in appearance. Its feet are exceptional. . like large hands … [T]he internal organs are found on dissection to correspond to those of man”4* Savage also wrote the first systematic account of gorillas in the wild, and was responsible for the modern use of the ancient North African word “gorilla”. He took pains to repudiate popular notions of gorillas carrying off attractive women for unspeakable purposes—the theme echoed a century later to enormous public acclaim in the motion picture King Kong.* The soldiers of Alexander the Great—not otherwise known for their prudishness—are said, in their India campaign, to have put monkeys to death for their “lasciviousness.”13* Jean-Jacques Rousseau, in 1753, had gone further and classified chimps and men as members of the same species, the power of speech being at the beginning, in his view, not “natural to man.”15 Congreve had toyed with something similar.* A young mother will not usually come again into estrus until she weans her infant. The little one, understandably enough, may interpret weaning as rejection. The mother’s renewed sexual interest in adult (and sub-adult) males probably compounds the infant’s agony and resentment. Perhaps we also share the Oedipus complex with the apes.




Chapter 16




LIVES OF THE APES


I hear the apes howl sadly


In dark mountains.


The blue river


Flows swiftly through the night.


MENG HAU-RAN


(Tang Dynasty, early 730s),


“Written for Old Friends in


Yang-jou City While Spending the


Night on the Tung-lu River”1


The alpha male is sitting bolt upright, jaw set, staring confidently into middle distance. The hair on his head, shoulders, and back is standing on end, which gives him an even more imposing aspect. Before him crouches a subordinate, in a bow so deep that his gaze must be fixed on the few tufts of grass directly before him. If these were humans, this posture would be recognized as much more than deference. This is abject submission. This is abasement. This is groveling. The alpha’s feet may, in fact, be kissed. The supplicant could be a vanquished provincial chieftain at the foot of the Chinese or Ottoman Emperor, or a tenth-century Catholic priest before the Bishop of Rome, or an awed ambassador of a tributary people in the presence of Pharaoh.2

Calm and assured, the alpha male does not scowl at his nearly prostrate subordinate. Instead, he reaches out and touches him on the shoulder or head. The lower-ranking male slowly rises, reassured. Alpha ambles off, touching, patting, hugging, occasionally kissing those he encounters. Many reach out their arms and beg for contact, however brief. Almost all—from highest to lowest rank—are visibly buoyed by this king’s touch. Anxiety is relieved, perhaps even minor illnesses cured, by the laying-on of hands.

Regal touching, one after the other, in a sea of outstretched hands seems familiar enough to us—reminiscent, say, of the President striding down the central aisle of the House of Representatives just before the State of the Union address, especially when he’s riding high in the polls. The future King Edward VIII on his world tour, Senator Robert Kennpdy in his presidential campaign, and countless other political leaders have returned home black and blue from the grasp of their enthusiastic followers.

The alpha male will intervene to prevent conflict, especially between hotheaded young males pumped up on testosterone, or when aggression is directed at infants or juveniles. Sometimes a withering glance will suffice. Sometimes the alpha will charge the pair and force them apart. Generally, he approaches with a swagger, arms akimbo. It’s hard not to see here the rudiments of government administration of justice. As in all primate leadership positions, an alpha male must accept certain obligations. In return for deference and respect, for sexual and dining privileges, he must render services to the community, both practical and symbolic. He adopts an impressive demeanor, even something approaching pomp, in part because his subordinates demand it of him. They crave reassurance. They are natural followers. They have an irresistible need to be led.

Beyond the reaching out of hands there are many styles of submission, of which the most common is, in the scientific literature, demurely called “presenting.” What is it that’s being presented? The subordinate animal, male or female—but here we’re discussing males in the dominance hierarchy—wishing to pay its respects to the alpha male crouches down and elevates its anogenital region toward the leader, moving its tail out of the way. It sometimes gives a little bump and grind. It may whimper and, grinning over its shoulder, approach the alpha, raised rump first. The subordinate’s need to pay respect in this manner is so great that it may even present to an alpha who’s fast asleep.

The alpha (if awake) moves forward, grasps the submissive animal from behind, closely embraces it, and not infrequently makes a few pelvic thrusts. Since this is the invariable posture and motion of chimp copulation, there can be no mistaking the symbolic significance of the exchange: The subordinate animal asks please to be fucked, and the dominant animal, perhaps a little reluctantly, complies.

In most cases these actions are only symbolic. There is no intromission and no orgasm. They fake it. You wish to pay respect to a high-ranking male, but Nature has not equipped you with appropriate spoken language. Still, there are many postures and gestures in your everyday life that have a meaning readily apprehended by everyone. If females must comply with nearly every proferred sexual invitation, the sex act itself is a vivid, powerful, and unambiguous symbol of submission. Indeed, presenting is a mark of deference and respect among all the apes and monkeys, and among many other mammals as well.

The anger of a high-ranking male is fearsome. His arousal is obvious to any bystander, because all the hair on his body is standing on end. He may charge, intimidate, and tear branches from trees. If you’re not prepared to meet him in single combat, you might want to appease him, to keep him happy. You closely monitor the slightest raising of a single one of his hairs. Not only are you perpetually compliant (“I’m yours whenever you want me”), but just for your own comfort you need frequent reassurance that he’s not angry with you. When he is angry, he exaggerates his size and ferocity and displays the weapons that he will bring to bear if the adversary does not submit. He uses his displays to keep more junior males in line, and they use theirs to advance within the hierarchy. Displays may serve as a response to a challenge, or just as a general reminder to the community at large that here’s someone not to be trifled with. Of course, it’s not all bluff; if it were, it wouldn’t work. There must be a credible threat of violence. A kind of menace maintenance is required. If push comes to shove there may be serious fighting. But much more often the display has a ritual and ceremonial character. (Almost always the alpha wins, and if, on occasion, he loses, that doesn’t usually mean that the hierarchy has been inverted; for that to happen, a consistent pattern of defeat is needed.)

The lesson being communicated is deterrence, pure and simple: “Cross me and you’ll have to deal with this stature, these muscles, these teeth (note my canines), this rage.” Chimpanzee strategy is encapsulated in the earliest comprehensive account we have of human military affairs, the sixth-century B.C. work The Art of War, by Sun Tzu: “The supreme act of war is to subdue the enemy without fighting.”3 Deterrence is old. And so is its prerequisite, imagination.

So law and order are maintained, and the status of the leadership preserved through the threat (and, if necessary, the reality) of violence; but also through patronage delivered to constituents, and through the widespread craving to have a hero to admire, who can tell you what to do—especially when there’s a threat from outside the group. Violence and intimidation alone would not suffice—although there may be those who enjoy being chastised and bullied, who perhaps look on it as a form of affection.

Male chimps are obsessively motivated to work their way up the dominance ladder. This involves courage, fighting ability, often size, and always real skill in ward-heeler politics. The higher his rank, the fewer the attacks on him by other males and the more gratifying instances of deference and submission. But the higher his rank the more he will be obliged to take pains to reassure subordinates. The dominance hierarchy makes for a stable community not only because the high-ranking males break up fights among their subordinates, but also because the very existence of hierarchy, along with the genetic tradition of compliance, inhibits conflict. One powerful motivation to be high-ranking is that the top echelons often have preferential sexual access to ovulating females. As in all mammals, this behavior is mediated by testosterone and related steroid hormones. Leaving more offspring is what natural selection is about. For this reason alone, hierarchy makes evolutionary sense.

The alpha male, merely by virtue of his exalted status, stimulates the formation of cabals to depose him. A lower-ranking male may challenge the alpha by bluff, intimidation, or real combat, as a step towards reversing their relative status. Especially under crowded conditions, females can play a central role in encouraging and helping to implement coups d’état. But the alpha male is often prepared single-handedly to take on coalitions of two, three, or four opponents.

Alphas enforce authority; betas and others sometimes challenge it—not on abstract philosophical grounds, but as a means to selfish ends. We might guess that both warring inclinations are built into us too, a different balance in different people, with much depending on the social environment. The roots of tyranny and freedom trace back to long before recorded history, and are etched in our genes.

Over a period of years in a typical small chimpanzee group, half a dozen different males may become alpha in succession—because of death or illness of the dominant male, or because of challenges from below. On the other hand, an alpha male maintaining his status for a decade is not unknown either. Perhaps coincidentally, these terms of office are roughly those typical of human governments—ranging, respectively, from Italy, say, to France. Political assassination—that is, dominance combat in which the loser dies—is rare.

In combat, males are more likely to hit, kick, stomp, drag, and wrestle. Or throw stones and beat with clubs, if any are handy. Females are more likely to pull hair and scratch, and to grapple and roll. For all their baring of teeth, males rarely bite anyone in the group, because their canines can do terrible damage. They may flash the razors and switch-blade knives, but they hardly ever draw blood. Females with much less prominent canines have fewer inhibitions. Any given fight is likely to stimulate other fights among unrelated or even nonaligned parties. One combatant may poignantly appeal for aid from passersby, who may, in any case, be attacked for no apparent reason. Any conflict seems to raise the testosterone level in all the male bystanders. Everyone’s hair stands on end. Perhaps long-standing resentments flare. General mayhem often results.

Chimps will place their fingers between the teeth of a high-ranking male and derive reassurance when the fingers are returned intact. At times of rising group tension, male chimps may touch or heft each other’s testicles, as the ancient Hebrews and Romans are said to have done upon concluding a treaty, or testifying before a tribunal. Indeed the root of “testify” and “testimony” is the Latin word, testis. The significance of the gesture, less common now that men wear pants, is not only transcultural, but trans-species.

——


From infancy, chimps are groomed, chiefly by their mothers. They in turn clutch their mothers’ fur from the moment of their birth. The infant revels in the physical contact, deriving deep and long-term psychological benefits from it. Even if their physical needs are attended to, monkeys and apes that, as infants, don’t receive something like hugging and grooming, grow up to be socially, emotionally, and sexually incompetent. As the infant matures, grooming behavior is slowly transferred to others. Most adults have many grooming partners. In a grooming pair, one partner mainly does, the other is mainly done to. But even the alpha will play either role. One individual will sit serenely while the other combs through its hair, rubs all its parts, and occasionally finds a parasite (a louse or a tick—maybe getting ripped on butyric acid), which it promptly eats. Sometimes the chimps hold hands the whole time. Jittery full-grown males will return to their mothers to be groomed and reassured. Males who become irritable with one another often hastily repair to mutual grooming to calm each other down. It may have been selected for long ago, as an improvement in chimp hygiene and public health, but grooming has now become a centrally important social activity, probably lowering testosterone and adrenaline titers.

The closest human counterpart may be the back rub or the body massage, which have been raised to art forms in cultures as diverse as modern Japan and Sweden, Ottoman Turkey and Republican Rome—where, in characteristic human fashion, a specialized tool, the strigil, was employed to rub the back. Gentlemen in Restoration England idled away the hours by collectively combing their wigs. Where body lice are a problem, human parents carefully and routinely go through their children’s hair. The emotional power of being groomed by the alpha male is perhaps akin to the laying-on of hands by shamans, healing ministers, chiropractors, charismatic surgeons, and kings.

Despite the importance of the male dominance hierarchy, it is by no means the only important chimp social structure, as the grooming pairs indicate. A mother and her children, or two grown siblings, have special, lifelong, mutually supportive bonds. A high-ranking son may be to a mother’s social advantage. There are also long-term relationships between unrelated individuals of the same sex that might certainly be called friendships. Largely outside the male hierarchy, there’s an intricate set of female bonds that often depend on the number and status of relatives and friends. These extrahierarchical alliances provide important means of mitigating or reordering a dominance hierarchy: If the alpha male is undefeated in one-on-one confrontation, an alliance of two or three lower-ranking males with supporting females may conceivably put him to flight. High-ranking males are known to establish alliances with promising younger males, perhaps co-opting them to prevent future putsches. Occasionally females will step in to defuse a tense encounter.

Alliances are made and broken. Loyalties shift. There is courage and devotion, perfidy and betrayal. No dedication to liberty and equality is evident in chimpanzee politics, but machinery is purring to soften the more hard-hearted tyrannies: The focus is on the balance of power. Frans de Waal writes:The law of the jungle does not apply to chimpanzees. Their network of coalitions limits the rights of the strongest; everybody pulls strings.4


In this complex, fluid social life great benefits accrue to those skilled in discerning the interests, hopes, fears, and feelings of others. The alliance strategy is opportunistic. Today’s allies may be tomorrow’s adversaries and vice versa. The only constant is ambition and fixity of purpose. Lord Palmerston, the nineteenth-century British Prime Minister—who described his nation’s foreign policy as no permanent national alliances, only permanent national interests—would have been right at home among the chimps.

Males have special reasons to avoid permanent rivalries. In the hunt and in patrols into enemy territory, they rely on one another. Mistrust would endanger their effectiveness. They need alliances to work their way up the promotion ladder or to maintain themselves in power. So, while males are much more aggressive than females, they are also much more highly motivated toward reconciliation.

When Calhoun crowded his rats together he found a wholesale change in their behavior, almost as if their collective strategy was now to kill off enough of themselves and to lower the birth rate enough that the population in the next generation would be reduced to manageable numbers. Given all the chimp propensities that we’ve chronicled (and the fact, described in the next chapter, that baboons can go into a murderous, annihilating group frenzy when packed together), you might expect that chimps behave badly when overcrowded, as in zoos. In close confines a male chimp cannot escape from an attack, cannot lead a female into the bushes away from the controlling gaze of the alpha male, cannot enjoy the excitement of the hunt or the patrol or contact with females from adjacent territories. You might expect frustration levels to rise, and hierarchical encounters now to involve less bluff and more real combat. If you’re not ready for a fight to the death, you’d better, you might think, find some way to mollify, appease, show deference, pay your respects, perform services, be useful—and genuflect at every step so the alpha harbors no possible misgivings about whether you know your place.

Surprisingly, just the opposite is true. In zoo after zoo, males—and especially high-ranking males—exhibit a degree of measured restraint under crowded conditions that would be unthinkable if they were free. Imprisoned chimps are much more likely to share their food. Captivity somehow brings forth a more democratic spirit. When jammed together, chimps make an extra effort to get the social machinery humming. In this remarkable transformation it is the females who are the peacemakers. When, after a fight, two males are studiously ignoring one another—as if they were too proud to apologize or make up—it is often a female who jollies them along and gets them interacting. She clears blocked channels of communication.

At the Arnhem colony in the Netherlands, every adult female was found to play a therapeutic role in communication and mediation among the petulant, rank-conscious, grudge-holding males. When real fights were about to break out and the males began to arm themselves with rocks, the females gently removed the weapons, prying their fingers open. If the males rearmed themselves, the females disarmed them again. In the resolution of disputes and the avoidance of conflict,* females led the way.5

So, it turns out that indeed chimps are not rats: Under crowded conditions they make extraordinary efforts to be more friendly, to be slower to anger, to mediate disputes, to be polite—and the female role in calming the testosterone-besotted males is crucial. This is an important and encouraging lesson about the dangers of extrapolating behavior from one species to another, especially when they are not very closely related. Since humans are much more like chimps than like rats, we can’t help wondering what would happen if women played a role in world politics proportionate to their numbers. (We’re not talking about those occasional women Prime Ministers who have risen to the top by besting the men at their own games, but about proportional representation of women at all levels of government.)

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Students of the chimpanzee call it “courtship.” It’s a set of ritualized gestures by which the male signals to the female his sexual intentions. But in ordinary usage courtship is a word describing a patient human attempt, over long periods of time, and often with great gentleness and subtlety, to build trust and to create the foundations for a long-term relationship. The male chimpanzee’s courtship communication is much briefer and more to the point, much closer to “Let’s fuck.” He may swagger, shake a branch, rustle some leaves, fix her with his stare, and reach out an arm toward her. His hair will be erect. And not just his hair. An erect penis—bright red, contrasting vividly with his black scrotum—is an invariable part of chimpanzee “courtship,” which you might think is a good thing because most of the other symbolic desiderata of courtship are barely distinguishable from those used in intimidating other males. In chimpish, “Let’s fuck” sounds almost exactly like “I’m gonna kill you.” The significance of this similarity has not been lost on the females. They comply. A typical female rejection rate to an unrelated male’s sexual overture is about 3%.

In chimpanzee etiquette, the correct response to the male courtship display is to crouch down on the ground and lift your behind invitingly. If the social niceties should elude you at first, the male will shortly set you straight. Recalcitrant females are attacked. All males in the group expect sexual access to all females, subject to necessary exclusions enforced by jealous, higher-ranking males. (Adolescent females are available for copulation even to infant males, who are sometimes ardent lovers.) Again, a significant exception is mothers and sons; although the son may give it a try, the mother tends to resist vigorously.

It’s natural for us to think of the instant submission and compliance of these female apes as exacted under threat of bodily harm, as rape pure and simple, even if the female is not bitten or bruised. But this cannot be the whole story, because female primates raised alone will, on going into first estrus, present themselves readily to many passing males, to humans, and, occasionally, even to furniture. Not just some degree of compliance is hardwired and built in, but so is real sexual enthusiasm. As in the hamsters-in-motorcycle-jackets experiment, the females, if given a chance, often show a marked preference for the higher-ranking males: The Big Guy, he’s all right. Perhaps also the males present themselves to those of higher rank not so much as a humiliating means of social advancement but because they genuinely enjoy submission.

As with most animals, the chimp male enters the female’s vagina from behind. Often the male is in a crouching or seated position, with his hands on her waist or buttocks as she positions herself on him. To a human observer their faces are strangely expressionless. Much has been made about the difference between chimp and human sexual practices—almost certainly in an attempt to deny the closeness of the kinship. But the favorite ancient Roman sexual practice was chimp-like, the male seated on a small stool and the female, often her back to his front, settling herself down on him. The style of our hunter-gatherer ancestors (if we may judge from contemporary examples) is also more like the chimps: They are often recumbent on their sides, the male embracing the female from behind. As a fashionable human sexual practice, perhaps the “missionary position” is not much older than missionaries—although, as we’ll see later, there’s one other animal that adopted it long before they did.

By human standards chimp sexual life is a perpetual open-air orgy—compulsive, unending, and always with the male grasping the female from behind. The average copulation rate is one or two an hour. Every hour. For each mature chimp. In estrus, of course, it’s more. When the females are ovulating and able to be impregnated, their vulvas and allied nether parts swell extravagantly and turn bright pink.* In estrus, they’re walking sexual advertisements, and are then far more alluring. Because estrous periods are to some degree synchronized, there are times when a chimpanzee group is a sea of bobbing, compliant, soliciting swollen red rumps. Olfactory cues also signal their sexual availability. In marginal cases a passing male, unable to determine just by looking if she’s ovulating, may simply insert his finger into her vulva and take a sniff.

Chimpanzee sex isn’t a long and drawn-out business. Maybe eight or nine thrusts, each taking less than a second, and they’re done. The males have, by human standards, impressive recovery rates, including documented sequences of many ejaculations at five-minute intervals. Females in estrus are especially attractive in the early morning, probably because of the long and stressful celibacy imposed on the males by the necessity of having to sleep at night. As a kind of community property for the males, she may be taken every ten minutes by one male after another through mid-morning, by which time they may tire a little.

Occasionally a heroic or foolish female will refuse the male despite his transfixing stare, threatening gestures, and other signs of arousal. When he makes his approach she may scream and run away from him. Generally she doesn’t get far. When some hesitation is discerned, young males will ostentatiously search for a rock, or actually find one and make as if to throw it at her. This serves almost always as a convincing argument. One of the earliest studies of chimp sexual behavior suggested that female compliance occurs “by reason of the dominance or impulsiveness of the male and the desire of the female to avoid risk of physical injury by obeying his command.”7

Despite their apparently unrestrained sexual behavior, chimps get jealous. A male who rejected the solicitation of a female in estrus, but instead copulated with her daughter, was slapped in the face by the outraged mother. Cruising migrant females from the next territory are threatened or attacked by the local females—especially if the visitors go so far as to groom with one of the resident males. The male may also blaze with sexual jealousy over a particular female’s behavior—but, almost without exception, only when she is vividly pink and swollen and able to conceive. High-ranking males will then chase away aroused lower-ranking males. Although it’s unlikely he’s thought this out, his motive, it seems very clear, is to monopolize her around the time of ovulation so that no one but he can father her children.* As far as he’s concerned, the rest of the time she can do as she pleases.

Possessiveness is hard to maintain, though, at the core of the territory where the chimp population density is high. Even the most vigilant and high-ranking males will be distracted—by hunting, say, or challenges from lower ranks, or insufficient deference, or by grooming, or by the necessity of adjudicating disputes. And during such an intervention—it may last only a few minutes—other males, patiently awaiting their chance, pounce on the off-limits female, especially if she’s in estrus. Kleptogamy is on their minds. In zoos a female will, as soon as the alpha male is removed from her cage, present herself to lower-ranking males, even if this requires adroit positioning so the act can be performed through the bars of two adjacent cages. Both in the wild and in captivity, when the cuckolded male discovers what has happened, he attacks the female. Perhaps he knows that she was all too willing. Besides, it’s much safer than attacking a rival male.

Even when the alpha is present, a subordinate male may catch the eye of a female who strikes his fancy and then gaze pointedly toward some nearby bushes. Nonchalantly, he then ambles off, often followed after a discreet interval by the female. Sometimes their infidelity is observed. Motivated by jealousy or by the wish to ingratiate himself to the leader, the informer rushes up to the alpha in great excitement, takes his arm, points, and leads him to the treacherous couple. At other times the female may inadvertently reveal what is going on by uttering a high-pitched scream at the moment of her orgasm. After being discovered in this way more than once, females do not usually abandon the risky practice of clandestine rendezvous; instead, they learn to suppress the scream, converting it into a kind of husky pant.

Frans de Waal reports that, following a long grooming session between a high-ranking and a low-ranking male,a subordinate male may invite the female and enjoy a copulation without interference by the others. These interactions give the impression that males obtain “permission” for an undisturbed mating by paying a price in grooming currency … Perhaps sexual bargaining represents one of the oldest forms of tit for tat, one in which a tolerant atmosphere is created through appeasing behavior.9


To achieve reliable sexual monopoly during her estrus, the ardent male must usher the female away from the multitude. Scientists who study chimps call this “consortship,” and distinguish it from “courtship.” The proposition is put to the female as follows: He takes a few steps away and looks at her over his shoulder. If she does not instantly follow, he shakes a nearby branch. If this provides insufficient inducement, he will chase her and, if need be, attack her. More often she goes quietly, especially if he’s high-ranking. Then, off somewhere alone in the forest, he has her to himself. It is a distant intimation of monogamy.

Consortship typically lasts for weeks, and is not without its perils. The happy couple may be attacked by predators or patrols from the neighboring territory; and the male’s status in the dominance hierarchy may be undergoing active review during his absence. Jane Coodall reports a few cases in which the young female’s mother invites herself along on the consortship; “as far as the male is concerned,” she is a “most unwelcome chaperone.” Here, where conception is most likely, the incest taboo is particularly vivid—no case is known of a male chimp ever inviting his own mother or sister to be his consort.

Why do the females put up with all this? Certainly males are larger and stronger than females and can and will hurt them, if that’s what’s needed to get their way. But this is only in one-on-one interactions. Why don’t females band together to defend themselves against a sexually predatory male? If two or three aren’t enough, six or eight would be. This is known, but rare, in the wild. (It is the custom among the chimps in the Tai National Forest in the Ivory Coast.) But it’s more common when they’re in closer quarters, as in the Arnhem colony in the Netherlands. Here the social conventions are different. If a male solicits a female and she’s uninterested, she so indicates, and that, usually, is that. If he makes himself obnoxious, he may be attacked by one or more other females. It is astonishing that so striking a characteristic of chimpanzee life in the wild as male sexual oppression of females can to such an extent be reversed merely because they’re all crowded together in a minimum security prison. We’ve already seen how, under these conditions, restraint, coalition building, and peacemaking by females come to the fore. Societies in which females have something approaching equality are also societies that benefit from their political skills.

In a state of freedom—where it’s possible to avoid your rivals by taking your sweetheart on a little trip into the country, and where you can escape a bully by running away—the circumspection required in crowded conditions is relaxed. Here testosterone is at full throttle and gentlemanly behavior is uncommon. The primate expert Sarah Blaffer Hrdy10 speculates that, among wild chimpanzees, female compliance to male sexual demands is the single mother’s desperate strategy for safeguarding her children. The males, Hrdy proposes, nursing their resentment at any rejection, might attack the children of an unresponsive mother (perhaps at a later time), or at least not protect them against attack by others.* In the brutal world of the chimpanzee, she suggests, the female does what the males ask in order to bribe them, so they will not kill (and, who knows, if they’re in a good mood might even help save) her children.* If Hrdy is right, perhaps the males are not oblivious of the bargain struck. Do they threaten the children in order to make the mothers come around? Do they attack children at random as a cautionary lesson for any mothers toying with noncompliance? Have chimp males organized a protection racket, with the females and the young as their victims?

Let’s leave aside the possibility of conscious extortion, and think for just another moment about Hrdy’s speculation. The females don’t provide food for the males. They don’t seem to be any better at grooming than the males. Perhaps the only commodity—certainly the most valuable commodity—they can offer to protect their children is their bodies. So they make the best of a desperate situation. Now a male is less likely to attack and more likely to protect her baby. But when circumstances change, when aggression is inhibited because of crowding, the females can finally say “No”—without having their heads handed to them for it.

Again, we must not imagine that chimps think all this through. They must have some other, more immediate reinforcement of their behavior. Hrdy raises the question of the selective advantage of orgasms, especially multiple orgasms, among female apes and humans. In a monogamous couple, what evolutionary benefit does it confer? she asks, and argues that none is apparent. But if instead we imagine the female copulating with many males in order that none of them harm her offspring, then, Hrdy conjectures, the orgasm—reinforcing successive matings with many partners—plays a vital role.

To what extent female sexual compliance is coerced by the males and to what extent it is entered into voluntarily and exuberantly is still not clear.

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Nucleic acids compete, individual organisms compete, social groups compete, perhaps species compete. But there is also competition on a very different level: Sperm cells compete. In a single human ejaculation there are some 200 million sperm cells, the fittest among them with tails lashing, racing against each other, speeding along at an average clip of five inches per hour, each striving—or so it seems—to be first to reach the egg. A surprising number, though, from normal, fertile males have deformed heads, multiple heads or tails, kinked tails, or are just motionless, dead in the water. Some swim straight, others in convoluted paths that may turn back on themselves. The egg may actually choose among sperm cells. Chemically, it cries out to them, egging them on. Sperm cells are equipped with a sophisticated array of odor receptors, some oddly similar to those in the human nose. When the sperms obediently arrive in the vicinity of the calling egg, they don’t seem to have sense enough to stop swimming and thrashing, and molecules on the eggs surface may cast out a kind of fishing line, hook the sperm, and reel it in. The fertilized egg then promptly establishes a barrier that turns away all future sperm cells who may come blundering in. These modern findings are rather different from the conventional view of the passive egg waiting to be claimed by the champion sperm.13

But there is, in an ordinary impregnation, something like one success and 200 million failures. So conception, while controlled to a significant degree by the egg, is still in part the result of a competition among sperm cells for speed, range, trajectory, and target recognition, at least.*

Odds anywhere approaching 200-million-to-1 in every conception, continued once a generation through geological ages, imply an extremely strong selection of sperm. Leaner, more streamlined sperm cells with more swiftly lashing flagellas that can swim straight and that have superior chemical sensors will probably arrive first; but that has very little to do with the characteristics, once grown up, of the individual so conceived. Getting to the egg first with genes for boorishness, say, or stupidity, seems a dubious evolutionary benefit. A great deal of effort would appear to be squandered in natural selection among the sperm cells.14 But then it seems odd that so many sperm cells are dysfunctional. We do not understand why this should be.

Many other factors affect which sperm succeeds: Who’s conceived must depend on the progress of the egg into the fallopian tubes, the precise moment of ejaculation, the position of the parents, their rhythm of motion, subtle distractions or encouragements, cyclical hormonal and metabolic variables, and so on. At the heart of reproduction and evolution, again we find a surprisingly strong random component.

The monkeys and apes are preeminent among animals where many males mate, one after the other, with the same female. They can hardly contain themselves, jumping up and down with excitement, awaiting their turn. In chimpanzees, as we’ve noted, there may be dozens of copulations in quick succession with an ovulating female. So the act itself cannot be prolonged or rich in nuance. Several pelvic thrusts, roughly one a second, and it’s over. For an average male there’s a copulation maybe once an hour, all the livelong day. For females in estrus it’s much more than that.

In ten or twenty minutes many males may have copulated with the same female. So consider the sperm cells of these various male chimps, racing against one another. Essentially, they set out from the same starting line. The probability of insemination by a given male is proportional to the number of sperm cells delivered, other things being equal; and thus the chimps with the largest number of sperm cells per ejaculation, the chimps able to copulate the most times in succession before exhaustion sets in, have an advantage. Having more sperm cells requires larger testicles. The very large testicles of male chimps amount to about a third of a percent of their entire body weight—twenty or more times the endowment, relatively speaking, of primates who are monogamous or who live in breeding units of one male and several females. In general it is found that males have considerably larger testicles for their body size in species where many males mate with each female. Not only is there selection for testicular volume, but also for an interest in copulation. This may be one of the routes—there are many mutually reinforcing trajectories, as we’ve described—to the highly sexual social proclivities of our primate order. Because men, compared to male chimps, have such relatively small testicles, we might guess that promiscuous societies were uncommon in the immediate human past. But a few million years ago, say, our ancestors may have been substantially more indiscriminate sexually and substantially better endowed.

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A mother and her adult daughter who have been foraging separately for a few hours may merely look at each other and give a few grunts when they meet; but if they have been separated for a week or more, they are likely to fling their arms around each other with grunts or little screams of excitement, then settle down for a session of social grooming.15


Chimpanzee females and their young have deep bonds of affection, while the adolescent and adult males seem more often mesmerized by rank and sex. The young revel in rough-and-tumble play together. Infants whimper and scream if they find themselves out of sight of their mothers. Youngsters will come to the aid of their mother if she is being attacked, and vice versa. Siblings may show each other special, affectionate consideration throughout their lives, and take care of the young during childhood if—as is common—the mother has died before the children are grown. Occasionally chimps of either sex will endanger themselves to help others, even those who are not close relatives. Male bonding on a hunt or patrol is palpable. Clearly there are opportunities—especially when the testosterone titers are low—for civil, affectionate, even altruistic behavior in chimpanzee society.

Adult males, despite the dominance hierarchy, spend considerable time alone. After the birth of their first baby or two, most females spend their entire lives with others. So females are both required to develop more refined social skills and have more opportunity to do so. As is usual among monkeys and apes—with rare exceptions—only one child is born at a time. Except when they’re in estrus, their time is spent mainly with the children. This is key for the next generation: As we’ve mentioned, apes and monkeys that are not regularly cared for, nursed, held, fondled, and groomed by an adult tend to become socially awkward, sexually inept, and disastrous as parents when they grow up.

Females are not born knowing how to be competent mothers; they must be taught by example. The investment of time required of the mother is substantial: The young are not weaned until they’re five or six years old, and enter puberty around age ten. For much of the time until weaning they’re unable to care for themselves. They’re very good, though, in clutching their mother’s hair as they ride upside-down on her belly and chest. So long as they allow the infant to nurse whenever it wants, perhaps several times an hour, chimp mothers are usually infertile, and unattractive to males. This is called “lactational anestrus.” Without the males constantly hassling them for sex, they’re able to spend much more time with the kids.

Chimp mothers use corporal punishment only very rarely. Infants learn the conventional modes of threat and coercion by closely observing older male role models. Infant males soon attempt to intimidate females. This may take some effort; females, especially high-status females, may not take kindly to being bullied by some young whippersnapper. The upstart’s mother may help him in his efforts at intimidation. But before reaching adulthood nearly every male has obtained submission from nearly every female. Nursing male infants—including those still years away from weaning—routinely and successfully copulate with adult females. Adolescent males emulate adult males carefully (aping every nuance of their intimidation displays, for example), wish to be their apprentices and acolytes, are simultaneously nervous and submissive and hopeful in their presence. They’re looking for heroes to worship. It even happens that an adolescent who’s been cruelly attacked by an adult male will leave his mother and follow the aggressor everywhere, submission signals flashing, longing for acceptance at some future and glorious time.

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From a human perspective chimpanzee social life has many nightmarish flourishes. And yet, despite its excesses, it’s hauntingly familiar. Many spontaneous groupings of men are oriented around hierarchy, combat, blood sports, and loveless sex. The combination of dominant males, submissive females, differential but scheming subordinates, a driving hunger for “respect” up and down the hierarchy, the exchange of current favors for future loyalty, barely submerged violence, protection rackets, and the systematic sexual exploitation of all available adult females, has some marked points of similarity with the lifestyles and ambiance of absolute monarchs, dictators, big-city bosses, bureaucrats of all nations, gangs, organized crime, and the actual lives of many of the figures in history adjudged “great.”

The horrors of everyday life among the chimpanzees recall similar events in our history. We find humans behaving like chimps at their worst in endless succession in the daily press, in modern popular fiction, in the chronicles of the most ancient civilizations, in the sacred books of many religions, and in the tragedies of Euripides and Shakespeare. A summary of human nature based on the plays of Shakespeare would define “man,” wrote Hippolyte Taine, asa nervous machine, governed by a mood, disposed to hallucinations, transported by unbridled passions, essentially unreasoning … and led at random, by the most determinate and complex circumstances, to pain, crime, madness, and death.16


We’re not descended from chimps (or vice versa); so there’s no necessary reason why any particular chimp trait need be shared by humans. But they’re so closely related to us that we might reasonably guess that we share many of their hereditary predispositions—perhaps more effectively inhibited or redirected, but smoldering in us nevertheless. We’re constrained by the rules that, through society, we impose on ourselves. But relax the rules, even hypothetically, and we can see what’s been churning and fermenting inside us all along. Beneath the elegant varnish of law and civilization, of language and sensibility—remarkable accomplishments, to be sure—just how different from chimpanzees are we?

For example, consider the crime of rape. Many men find depictions of rape arousing—especially if the woman is portrayed as enjoying it despite her initial resistance. Most American high school and college students (of both sexes) believe that a man is justified in forcing a woman to have sex—at least when the woman behaves provocatively.17 More than a third of American college men acknowledge some propensity to commit rape if they were guaranteed they could get away with it.18 The percentage goes up if some euphemism such as “force” appears in the question instead of “rape.” The actual risk of an American woman being raped in her lifetime is at least one chance in seven; almost two-thirds of the victims have been raped when they were minors.19 Perhaps men in other nations are less fascinated with rape than Americans are; perhaps mature men, with lower testosterone titers, are less comfortable with rape than adolescents are.20 But it would be hard to argue that there’s no biological predisposition for men to rape.

While a range of causal factors have been proposed, most rapists turn out to be not slavering psychopaths, but ordinary men given the opportunity and acting on impulse,21 sometimes repeatedly and compulsively. Some students of the subject see rape as a biological strategy (entered into without his conscious understanding) to propagate the rapist’s genes;22 others see it as a means for men (again largely unconsciously) to maintain through intimidation and violence their domination over women.23 The two explanations do not seem mutually exclusive; and both seem to be operative in chimp society. Also, a significant minority of women are aroused by fantasies of rape, and, in one study, women who have been raped by an acquaintance seem disturbingly more likely to continue dating their assailants than those who were subjected only to attempted rape by an acquaintance.24 This is at least reminiscent of the compliance pattern of female chimps.

Over a set of hereditary predispositions human society lays down a kind of stencil that permits some to be fully expressed, some partially, and some hardly at all. In cultures where women have roughly comparable political power with men rape is rare or absent.25 However strong any genetic propensity toward rape might be, social parity appears to be a highly effective antidote. Depending on the structure of the society, many different brews of human proclivities can be elicited.

——


Chimpanzee society has an identifiable set of rules that most of its members live by: They submit to those of higher rank. Females defer to males. They cherish their parents. They care for their young. They have a kind of patriotism, and defend the group against outsiders. They share food. They abhor incest. But they have, so far as is known, no lawgivers. There are no stone tablets, no sacred books in which a code of conduct is laid out. Nevertheless, there is something like a code of ethics and morals operating among them—one that many human societies would find recognizable and, as far as it goes, congenial.


* Among the males. Among their own gender, females may carry grudges for years, and refuse to be reconciled.* That this might have something to do with sex was first proposed, in the face of considerable Victorian skepticism and unease, by the ever-insightful Charles Darwin.6* Similar behavior is known among other social animals—in gorillas, for example, where the alpha permits a female to mate with lower-ranking males, but only if she’s pregnant. Among wolves, only the alpha male and the alpha female breed, but the female mates with other members of the pack when she’s not in heat.8* This is not just an unpleasant circumstance of chimpanzee life; it occurs among gorillas, baboons, and many other apes and monkeys. Over a fifteen-year-long study of gorillas near the Virunga volcano in Rwanda, more than a third of all infant mortality was directly due to killing by gorilla males. Infanticide for them is a way of life.11* Something similar is observed in other, quite different non-monogamous species—for example, hedge sparrows. The alpha male works hard to prevent copulation by betas, but only in the females’ fertile period. However the female, even in the fertile period, may dart away on occasion for surreptitious matings with the betas. Only in this case will a beta help feed her chicks. Again the females are using male preoccupation with sex to induce them to help her little ones.12* A sperm that carries the smaller Y chromosome—the one that makes a male—weighs slightly less than one that carries the bigger X chromosome that makes a female; if lighter sperms travel faster, this may be why slightly more males are conceived than females.




Chapter 17




ADMONISHING THE CONQUEROR


Perhaps no order of mammals presents us with


so extraordinary a series of gradations as this


[step by step, from humans to apes to monkeys


to lemurs]—leading us insensibly from the


crown and summit of the animal creation down


to creatures, from which there is but a step, as


it seems, to the lowest, smallest, and least


intelligent of the placental Mammalia. It is as if


nature herself had foreseen the arrogance


of man, and with Roman severity had provided


that his intellect, by its very triumphs, should


call into prominence the slaves, admonishing


the conqueror that he is but dust.


T. H. HUXLEY


Evidence as to Mans Place in Nature1


The Archbishop of York is Primate of England. The Archbishop of Armagh is Primate of Ireland. The Archbishop of Warsaw is Primate of Poland. The Pope is Primate of Italy. The Archbishop of Canterbury is Primate of the planet, at least as far as his Anglican communicants are concerned. These ancient titles come from the medieval Latin word primus, which in turn derives from older Latin words meaning “principal” and “first.” The ecclesiastic use was straightforward: A primate of a region was the chief (“first”) of all its bishops. In recent centuries the title has devolved often to little more than an honorific. Other titles have taken precedence. But “Prime Minister” and “President” and “Premier” come from similar linguistic roots, all meaning “first.”

When Linnaeus was drawing up the family tree of life on Earth he was, as we’ve noted, afraid to include humans among the apes. But despite widespread opposition, it was impossible to deny some deep connections of monkeys, apes, and humans.* So all were classified into the order (for him, one taxon higher than genus) that he called primates. Scientists who study non-human primates—of course, they’re all primates themselves—are called primatologists.

This other meaning of “primate” also derives from the Latin for “first.” It’s hard to see by what standard a squirrel monkey, say, could be considered “first” among the lifeforms of Earth. But if a case is made that humans are “first,” then the tarsiers, bushbabies, mandrills, marmosets, sifakas, aye-ayes, mouse lemurs, pottos, lorises, spider monkeys, titis, and all the rest are dragged in along with us. We’re “first.” They’re our close relatives. So they, in some sense, must be “first” also—an undemonstrated and suspect conclusion in a biological world that runs from virus to great whale. Perhaps, instead, the argument goes the other way, and the humble status of most members of the primate tribe casts doubts on the lofty title we have appropriated to ourselves. It would make things so much easier for our self-esteem if those other primates weren’t—anatomically, physiologically, genetically, and in their individual and social behavior—so much like us.

Surely there is at least a hint in the word “primate,” not just of self-congratulation, but of the idea, fully realized in the practices of our own time, that we humans arrogate command and control of all life on Earth into our own hands. Not primus inter pares, first among equals, but just plain primus. We’ve found it convenient, even reassuring, to believe that life on Earth is a vast dominance hierarchy—sometimes called “The Great Chain of Being”—with us as the alphas. Sometimes we claim that it wasn’t our idea, that we were commanded by a Higher Power, the most Alpha of Alphas, to take over. Naturally, we had no choice but to obey.

About two hundred species of primates are known. Conceivably, in the quickly dwindling tropical rainforest another species or two—nocturnal or elegantly camouflaged—may have so far escaped our notice. There are about as many species of primates as there are nations on Earth. And like the nations, they have their different customs and traditions, which we sample in this chapter.

——


Take the baboons—“the people who sit on their heels,” as the !Kung San people of the Kalahari Desert respectfully call them. Hamadryas baboons are different from savanna baboons (from whom they diverged perhaps 300,000 years ago), and free baboons behave very differently from baboons crowded together in zoos (the latter “insolently lascivious,” as an eighteenth-century naturalist described them). One telltale trait they all have in common: Sharing meat is virtually unknown among baboon males of either species, although it’s fairly widespread among the chimps.

At sunrise the baboons rouse themselves from their sleeping cliffs and break up into a number of smaller groups. Each group wends its separate way over the savanna, foraging, scampering, playing, intimidating, mating—all in a day’s work. But at the end of the day, all the groups converge on the same distant waterhole, and it may be a different waterhole on different days. How do the groups, out of sight of one another for most of the day, know to wind up at the same waterhole? Have the leaders negotiated the matter as the sun was rising over the sleeping cliffs?

Adult male hamadryas baboons are almost twice as large as the females. They display a leonine mane, enormous, almost fanglike canine teeth, and a ruthless character. These males were deified by the ancient Egyptians. They utter deep and prolonged grunts as they copulate. Their faces are “the color of raw beef steak—as different from the mousey grey-brown females as if they belonged to two different species.”2 As females approach sexual maturity, they are chosen by particular males and herded into harems. Squabbling among competing males over ownership of the females may have to be worked out. A high priority of the males is maintaining and improving their status in the dominance hierarchy.

Hamadryas harems characteristically comprise from one to ten females; the males are concerned to keep peace among the females and to make sure that they do not so much as glance at another male. This is a bondage with little hope of escape. A female must follow her male about for the rest of her days. She must be sexually submissive: the least reluctance and she is bitten in the neck. It is not unknown for a hamadryas female to have her skull punctured and crushed in the massive jaws of the male for a minor infraction of the behavioral code he ruthlessly enforces.3 Conflict and tension around her are high when she’s ovulating, and somewhat muted when she’s pregnant or nursing the young. Unlike the chimps, you can see sexual coercion in the very posture of the baboon copulatory style: The male typically grasps the female’s ankles with his prehensile feet while mating, guaranteeing that she cannot run away. Compared to hamadryas behavioral norms, chimps live in an almost feminist society.

In a quarrel among females, one will sometimes threaten her rival with her teeth and forearms and, at the same time, alluringly present her rump to the male; with this postural offer of a deal, she sometimes induces him to attack her adversary. Subordinate male savanna baboons, as well as barbary apes, may use an infant—an unrelated infant, a bystander infant, or maybe an infant he is baby-sitting—as a hostage or shield or placatory object when approaching a high-ranking male. This tends to calm the alpha down if he’s in a grumpy mood.

The hamadryas male’s larger size and ferocious temperament doubtless are useful when the troop is imperilled by predators, or in conflict with other groups. But, as in the rest of the animal kingdom, when there are conspicuous differences in stature between the sexes (usually, it’s the males who are bigger), there’s exploitation and abuse of the smaller and weaker (usually the females).* Another distinction of the hamadryas baboons is that, alone among nonhuman primates so far as we know, two groups have been observed to ally themselves in combat against a third.4

Among savanna baboons, where the size difference between the sexes is not so striking, there are no harems. They are great walkers; it’s not unknown for a troop to cover twenty miles a day. Unlike chimps and hamadryas baboons, here it’s the male who leaves the natal troop around puberty—again, probably as an evolutionary device to avoid incest, and genetically to connect semi-isolated populations. When he attempts to enter a new troop, objections are likely to be raised by the resident males. Acceptance by the group often requires the time-honored method of submission, bluff, coercion, and alliance-making in the male hierarchy. But in many cases another strategy works well: make friends with a particular female in the troop and her children. He grooms her. He baby-sits and cares for her young. No killing off the young here in order to bring her into ovulation, as with rats and lions. If all goes well, she sponsors his entry into the troop. We can imagine a certain exhilaration as, gingerly, he attempts to enter the new community, his gaffes and old enemies left behind, a clean slate before him, and success dependent almost entirely on his social skills.

The males are more flighty and tempestuous than the females. Social stability is mainly provided on the female side. Indeed, since savanna baboon males are transients, the only hope for coherent group structure lies with the females. In all things, female baboons are comparatively conservative; it is the testosterone-pumped males who take the risks.

The female dominance hierarchy is largely hereditary. Daughters of alpha females are given unusual deference, even as juveniles, and have a good chance of achieving alpha status when they grow up. Every close relative of the dominant female may outrank every other member of the troop—a royal family. Submission and dominance in the female hierarchy of savanna baboons and many other monkey species is conveyed in the time-honored idiom of presenting and mounting, the heterosexual metaphor again adapted to another purpose.

——


For reasons not fully understood but worth speculating on, much more attention—at least in public discussion and until recently—has been given to hamadryas baboons than to their savanna cousins. Sometimes the impression has been left that hamadryas behavior is representative of all nonhuman primates, or even all primates. For example, the hamadryas males, in a species in which nothing else is owned, have a clear sense of females as private property. But this is by no means true of all primates. The hamadryas baboons, it turns out, provide perhaps the most extreme example of hierarchy and brutality in the entire order of the primates. This behavior was especially marked under a set of cruel circumstances devised by humans who meant them no harm:

Living with apes or monkeys in the wild did not much appeal to primatologists until recently. More typical was an expedition back to his native South Africa by Solly Zuckerman, anatomist to the Zoological Society of London:On the 4th of May, 1930, I succeeded in collecting on a farm near Grahamstown in the Eastern Province twelve adult females from one troop of baboons. Four of these were non-pregnant. Five were pregnant; one had an embryo 2.5 mm. in length; another one of 16.5 mm.; the third one of 19 mm.; the fourth one of 65 mm.; and the fifth an apparently full term male foetus with a crown-rump length of 230 mm. Three were lactating, and their babies were caught alive. One infant was estimated to be four months old, and the other two were each about two months.5


He dutifully noted how much fresh semen there was at various depths within the reproductive tracts of his female victims; “collected,” it turns out, is a euphemism for “killed.” Baboons had been officially declared “vermin” in South Africa, because they’re smart enough to defeat the efforts of farmers to safeguard their crops. A bounty was paid for each dead baboon. So a few baboons “collected” for science hardly mattered, compared with the wholesale slaughter being organized by the farmers. Through such studies Zuckerman “had the luck to discover from post-mortem study that ovulation in mature females occurs in the middle of the monthly sexual cycle.”6 The corresponding discovery about the human menstrual cycle was made around the same time.

He had long been interested in the standing of humans among the primates, and was dissecting baboons in South Africa while still a teenager.7 But he was not wholly unmoved by the plight of the hunted baboons, and later quoted this early-twentieth-century account:Hugging her baby tight to her breast, she regarded us with a world of sadness in her eyes, and with a gasp and shudder she died. We forgot for the moment that she was but a monkey, for her actions and expression were so human, that we felt we had committed a crime. Muttering an oath, my friend turned and walked rapidly off, vowing that this was the last time he would shoot a monkey. “It isn’t sport, it’s downright murder,” he declared, and I fervently agreed with him.8


If you wanted to meet a baboon—and you lived in a country where they didn’t roam about in the wild—you could always go to the local zoo and see the bedraggled and deracinated inmates, lifers pent up in tiny cubicles. After World War I, some European zoos thought it would be better, as well as more “humane,” if a large number of baboons could be gathered together in a partly open enclosure admitting observation by city-bound primatologists. The London Zoo was among them, and Dr. Zuckerman was playing a central role in the organization of one of these multiyear experiments:

In the spring of 1925, about one hundred hamadryas baboons were introduced into moat-bordered Monkey Hill, about 33 by 20 meters in area. So each baboon had, on average, less than 7 square meters, or some 60 square feet, indeed about the size of a small prison cell. It had been intended that this be an all-male group, but through an “accidental inclusion” six of the hundred baboons proved to be female. After a time, the oversight was rectified and the group was augmented by a further thirty females and five males. By late 1931, 64% of the males were dead, and 92% of the females:Of the thirty-three females that died, thirty lost their lives in fights, in which they were the prizes fought for by the males. The injuries inflicted were of all degrees of severity Limb-bones, ribs, and even the skull, have been fractured. Wounds have sometimes penetrated the chest or abdomen, and many animals showed extensive lacerations in the ano-genital region … The fight in which the last of these females lost her life was so protracted and repellent—from the anthropocentric point of view—that the decision was made to remove the five surviving females from the Hill … The very high percentage of females killed in the London Colony suggests. . that the social group of which they formed a part was in some way unnatural.9


Despite this last qualification, the hamadryas colony at the London Zoo reinforced a widespread belief in an unconstrained Darwinian struggle for existence. Even though baboons would quickly have exterminated themselves from the world if the events at Monkey Hill were characteristic of life in the wild, many people felt that they had now glimpsed Nature in the raw, a brutal Nature, red in claw and fang, a Nature from which we humans are insulated and protected by our civilized institutions and sensibilities. And Zuckerman’s vivid descriptions of the unrestrained sex lives of the baboons—he was one of the first to stress that baboon social organization may be determined largely by sexual considerations—increased the contempt that many humans felt toward the other primates.

What had gone wrong on Monkey Hill? First, almost all of the baboons introduced into the “colony” were unknown to one another. There was no long-term mutual habituation, no prior establishment of dominance hierarchies, no common understanding in these harem-obsessed males of who was to have many females and who none at all. No kinship-based female dominance hierarchy had been established. Unlike the situation in the wild, there were many more males than females. Finally, these baboons were crowded together to a degree rarely experienced in their natural state.

Because of their powerful jaws and spectacular canines, baboon males within a troop hardly ever fight among themselves in earnest, although corporal punishment is visited on the females for the slightest infractions. But in the London Zoo, dominance hierarchies had to be established, dedicated attempts were made to steal females, escape from a formidable attacker was cut off by the moat, and the calming influence of many sexually compliant females was almost entirely wanting. The result was carnage. In all six and a half years, only one infant survived. When the males would fight over them, the adult females would listlessly wait, as if “paralysed.” The battered, lacerated, punctured females would be sexually used by a quick succession of males.

But the females were not mere passive instruments:[W]hen her overlord’s back was turned she quickly presented to the bachelor attached to her party, who mounted for a moment. The overlord then slightly turned his head, whereupon the female rushed to him, her body low to the ground, presenting and squealing, and threatening her seducer with grimaces and with quick thrusts of her hands on the rocks. This behaviour immediately stimulated an attack by the overlord … Closely pursued, the bachelor fled. On another occasion the same female was left alone for forty seconds while her overlord chased a bachelor around Monkey Hill. In that space of time she was mounted and penetrated by two males to whom she had presented. Both of these immediately made off after their contact with the female, who again responded to the return of her male in the manner described above.10


When females were killed, the males would continue to drag them around, one male after another, to fight over them, and to copulate with their corpses. When the keepers, grimly watching this necrophilial tableau unfold, felt it necessary—for “anthropocentric” reasons—to enter the compound and remove the dead body, the males, in concert, would violently object and resist. Zuckerman, writing back then in the 1920s, used and may have coined the phrase “a sexual object”11 in describing the lot of the female baboon.

We’ve seen in Calhoun’s experiments with rats that—even when there’s plenty of food, even when there are as many males as females—severe crowding induces violent and other modes of behavior that many would describe as aberrant and maladaptive. We’ve also seen in the Arnhem chimp colony how, under similar circumstances, new modes of behavior come to the fore to inhibit violence. From the baboons in the London Zoo we learn that if you take a species given to sexual violence in the best of conditions, provide a small number of sexual prizes to be fought over, arrange to have no pre-existing social order in which the animals know where they fit, and now crowd everybody together with no hope of escape, mayhem is the likely outcome. Monkey Hill reveals a deadly intersection of sex, hierarchy, violence, and crowding that may or may not apply to other primates.*

In Nature, as Zuckerman recognized, hamadryas baboons live much more peaceably. Dominant males are surrounded by a small corona of females, their offspring, and a few affiliated “bachelor” males. These harems wander over the landscape in bands, collecting food. Hundreds of baboons, a kind of gathering of the tribes, camp out each night near one another on sleeping cliffs. Fights to the death for possession of the females (or for any other reason) hardly ever happen. Everyone knows his, and especially her, place. The females are of course routinely abused, bitten on average once a day, but not so deeply as to draw blood. They are certainly not all killed off because they might be interested in other males, as happened in the London Zoo.

Hamadryas baboons in very small groups behave very differently: A bachelor baboon male watches a couple—on their first date—in an adjacent cage. Days go by, and he is forced to observe their deepening sexual relationship while he sits alone. When he’s then introduced into their cage, he makes no effort to attack the male or to lure the female away. He respects their relationship. He looks away when they have sex. He is a model of rectitude and circumspection, even if he’s of larger stature than either.12

Unsurprisingly, there are ways of arranging a primate society so its structure collapses and almost everybody dies. Shall we think of primates who find themselves in such circumstances as criminals? Are they accountable for their actions? Do they have free will? Or shall we attribute the bulk of the responsibility for what happens to those through whose miscalculation the social environment was established? For a society to be successful, it must be consonant with the nature and character of the individuals who must live in it. If those contriving social structures overlook who these individuals are, or sentimentalize their nature, or are incompetent social engineers, disaster can result.

Zuckerman has consistently argued that almost nothing about human nature or evolution can be learned by studying monkeys and apes—quite the opposite of many students of animal behavior who believe that understanding primates might provide a direct route to understanding humans: “[M]y unbending critical attitude to attempts to explain human behavior by analogies from the animal world must have been acquired at a very early age.”13 He describes Konrad Lorenz, Desmond Morris, and Robert Ardrey—who popularized, with at least some excesses, the idea that we have something to learn about ourselves from studying other animals—as “three writers who are equally adept at devising superficial analogies.”14

As “Prosector” of the London Zoo—the officer in charge of animal autopsies—Zuckerman later submitted the manuscript of a book, entitled The Social Life of Monkeys and Apes, for approval to his superior in the zoo’s dominance hierarchy. It was promptly rejected on grounds of displaying an undecorous explicitness on matters sexual (for example, “The overlord’s attention is caught by the perineal region of one of his females, usually when her sexual skin is swollen. He bends his head forward, his hand reaches out, his lips and tongue move and, having thus stimulated the sexual response in the female, he mounts and copulates”15). Zuckerman offered the book for publication anyway. In his autobiography, From Apes to Warlords, published forty-six years later—amidst much vivid detail of those years—he makes only the most tangential reference to the events at Monkey Hill.

At the start of World War II Zuckerman studied the consequences of aerial bombardment on civilian populations—his anatomical knowledge could there be put to good use. He soon moved on to analyzing the effectiveness of aerial bombing in the accomplishment of strategic goals, where his skeptical proclivities came in handy: The RAF’s Bomber Command (and the U.S. Army Air Corps), he found, had consistently exaggerated the potential of massive aerial bombardment to lessen the enemy’s will to fight and to shorten the war.

After the war Zuckerman headed the London Zoo, and through a few turns in his career wound up to be the principal scientific adviser to the British Ministry of Defence, where his expertise in understanding dominance hierarchies may have been germane. Created a Life Peer, Lord Zuckerman worked for many years to slow the nuclear arms race.

——


Baboons as a whole represent only one small corner in the vast arena of primate behavior. We could just as easily have focusedon any of a number of lemur species, species in which females rather routinely dominate males. We could have decided to make an example of the shy and nocturnal owl monkey … where males and females cooperate in child care with the male playing the major role in carrying and protecting the infant, or we could have focused on the gentle South American monkeys known as “muriqui” … who specialize in avoiding aggressive interactions, or any of a host of other primate species in which we now know that females play an active role in social organization.16


Consider the gibbon. Its preternaturally long arms permit it to make great balletic leaps through the canopy of the forest—sometimes ten meters or more from branch to branch—that put champion human gymnasts to shame. Gibbons are, apparently without exception, monogamous. They marry for life. They produce haunting songs heard a kilometer or more away. Adult males often sing long solos in the darkness just before sunrise. Bachelors sing longer than old married males, and at a different time of day. Wives prefer duets with their husbands. Widows bear their grief in silence and sing no more.

Gibbons are also territorial and their matins serve to keep intruders away. A nuclear family, typically parents and two children, tends to control a small turf. Defense of the home territory is accomplished not so much by throwing stones or raining blows as by singing anthems. Perhaps there are cadences, timbres, frequencies, and amplitudes that other gibbons, contemplating a little poaching, find especially impressive and daunting. At least sometimes, an aging father will confer responsibility for territorial defense on his adolescent son, passing the patriotic torch on to the younger generation. In other equally poignant instances, adolescents are banished from the home territory by the parents, perhaps to avoid the temptations of incest. Adult males and females behave pretty much alike, and have nearly equal social status. Primatologists describe the females as “codominant,” and the partners in a marriage as “relaxed” and “tolerant.”17

Gibbon life seems downright operatic. It’s easy to conjure up feverish love solos, duets sung in praise of marital felicity, and ritual intimidation chants cast into the forest night: “We’re here, we’re tough, we sing good songs. Better leave our turf alone.” Perhaps there are gibbon Verdis singing power-transfer arias, rich with pathos, soulful lamentations on the passing of glory and of time.

Or consider the bonobo. This is a reclusive species or subspecies of chimpanzee that lives in a single group in Central Africa, south of the Zaire River.18 Bonobos have certain traits that render them conventionally ineligible for the local zoo, which may be one reason that they’re not nearly so well known as the common chimp we’ve described in the preceding chapters. Bonobos, given the Linnaean name Pan paniscus, are also called pygmy chimpanzees; they’re smaller and more slender and their faces protrude less than the usual variety, Pan troglodytes, which we’ll here and there continue to describe simply as chimpanzees.* Bonobos often stand up and walk on two legs. (They have a kind of webbing of skin between their second and third toes.) They stride with their shoulders squared and do not slouch as much as chimps do. “When bonobos stand upright,” writes de Waal, “they look as if they had walked straight out of an artist’s impression of prehistoric man.”19

Unlike chimp females, among whom estrus is advertised and is a time of pronounced sexual receptivity, bonobo females display genital swellings about half the time; and they’re nearly always attractive to the adult males. We recall that common chimps, Pan troglodytes, like almost all animals, have sex with the male entering the female’s vagina from behind, his front against her back. But in bonobos, about a quarter of the time, the matings are face-to-face. This is the position the females seem to prefer, probably because their clitorises are large and positioned far forward compared to chimps. Bonobos indicate their mutual attraction by prolonged gazing into one another’s eyes, a practice which precedes almost all their matings, and which is unknown among common chimps. The initiation of sexual activity among the bonobos is mutual, unlike the chimps, where it is peremptory and nearly always by the males. While in general, especially in larger social contexts, male bonobos dominate females, this is not always the case, especially when they’re alone together. At night, in the forest canopy, a male and a female will sometimes snuggle up together in the same nest of leaves. Adult chimps never do.

The sexual activity of common chimps, which by human standards seems obsessive to the point of mania, is almost puritanical by bonobo standards. The average number of penile thrusts in an average copulation—a measure of sexual intensity that primatologists are drawn to, in part because it can be quantified—is around forty-five for bonobos, compared to less than ten for chimps. The number of copulations per hour is 2½ times greater for bonobos than for chimps—although these observations are for bonobos in captivity, where they may have more time on their hands or more need for mutual comfort than when they are free. Less than a year after giving birth, bonobo females are ready to resume their lives of sexual abandon; it takes three to six years for chimp females.20

Bonobos use sexual stimulation in everyday life for many purposes besides mere satisfaction of the erotic impulse—for quieting infants (a practice said once to have flourished also among Chinese grandmothers), as a means of resolving conflict among adults of the same sex, as barter for food, and as a generic, all-purpose approach to social bonding and community organization. Less than a third of the sexual contacts among bonobos involve adults of opposite sexes. Males will rub rumps together or engage in oral sex in ways unheard of among the more prudish chimps; females will rub their genitalia together, and sometimes prefer it to heterosexual contacts. Females characteristically engage in genital rubbing just before they’re about to compete for food or for attractive males; it seems to be a way of reducing tension. In times of stress, a bonobo male will spread his legs and present his penis to his adversary in a friendly gesture.

Despite these differences in nuance, bonobos are still chimpanzees. There’s a male dominance hierarchy, although not nearly as pronounced as among common chimps; dominant males have preferential access to females, although males do not always dominate females; there are submissive gestures and greetings; the size of groups is about the same as with chimps, a few dozen; adolescent females wander over to adjacent groups; the males preferentially hunt animal prey, although apparently not in hunting parties; males are proportionately larger than the females by about the same ratio as among chimps; and encounters between groups sometimes become violent—although groups may also on encountering one another, behave very peaceably and laid-back. Infanticide and all other killing of bonobo by bonobo are, so far, unknown. Their standard initial response on meeting unfamiliar humans, as we ourselves experienced, is a very chimp-like, and adequately intimidating, charging display.

Grooming is most frequent between males and females and least common between males and males, the reverse of chimp practice. The grin serves not mainly as a gesture of submission, but performs a range of functions similar to those of the human smile. Male bonding is much weaker than in chimp society, and the social position of females much stronger. Certain mothers and sons associate closely until the son becomes an adult; among chimps the relationship tends more often to be broken off when the young male reaches adolescence. Social skills for resolving conflicts are much more highly developed among the bonobos than among the chimps, and dominant individuals are much more generous in making peace with their adversaries.

If we feel a certain revulsion at having hamadryas baboons as relatives, we may take some comfort from our connection with the gibbons and the bonobos. Indeed, we’re far more closely related to the apes than to the monkeys. Chimps and bonobos are certainly members of the same genus and, according to some taxonomic classifications, even the same species. Given that, it’s startling how different they are from one another. Perhaps many of the distinctions between the two—ranging from the frequency, increased variety, and social utility of sex to the relatively higher status of females—are due to the evolution in the bonobos of a new step: abandoning the monthly badge of ovulation, graduating from estrus. Perhaps when ovulation is not evident at a glance or a sniff, females can be viewed as more than sexual property.

The primates are so rich in potential that even a small change in anatomy or physiology may provide an aperture to a universe never dreamt of in the rude sleeping pallets made each night in the low branches of the once-vast tropical forests.


SOME SKETCHES FROM LIFE


Monkeys:

Monkeys are liable to many of the same non-contagious diseases as we are … Medicines produced the same effect on them as on us. Many kinds of monkeys have a strong taste for tea, coffee, and spirituous liquors: they will also, as I have myself seen, smoke tobacco with pleasure. Brehm asserts that the natives of north-eastern Africa catch the wild baboons by exposing vessels with strong beer, by which they are made drunk. He has seen some of these animals, which he kept in confinement, in this state; and he gives a laughable account of their behaviour and strange grimaces. On the following morning they were very cross and dismal; they held their aching heads with both hands, and wore a most pitiable expression: when beer or wine was offered them, they turned away with disgust, but relished the juice of lemons. An American monkey, an Ateles, after getting drunk on brandy, would never touch it again, and thus was wiser than many men. These trifling facts prove how similar the nerves of taste must be in monkeys and man, and how similarly their whole nervous system is affected.21


Eastern Mountain Gorillas:

When two animals meet on a narrow trail the subordinate gives the right-of-way; subordinates also yield their sitting place if approached by superiors. Sometimes the dominant animal intimidates the subordinate by starting at it. At most it snaps its mouth or taps the body of the other animal with the back of its hand.22


Monkeys:

[P]hallic threatening, derived from a sexual domination gesture (mounting), … has been described among many monkey species in both the Old World and the New. Among guenons and baboons, a few males always sit with their back to the group keeping guard, displaying their strikingly colored penis and their sometimes similarly strikingly colored testicles. If a stranger to the group approaches too closely, the guards actually have an erection; so-called “rage copulations” also take place.23


Squirrel monkeys:

The displaying monkey vocalizes, spreads one thigh, and directs the fully erect penis toward the head or chest of the other animal. The display is seen in its most dramatic form when a new male is introduced into an established colony of squirrel monkeys … Within seconds all males begin to display to the strange monkey, and if the new male does not remain quiet with its head bowed, it will be viciously attacked.24


Brown capuchin monkeys:

An estrous female will shadow the dominant male for days. At frequent intervals she approaches him closely, grimaces at him while giving a distinctive vocalization, pushes him on the rump, and shakes branches at him. When she is ready to copulate, she charges him, he runs away, she follows, and when he stops running, they mate.25


Orangutans:

At midcycle, a female orangutan will seek out the dominant male in her vicinity. At other times during her cycle, young males and subordinate males will sometimes cluster around her and it appears that she is being forced to mate with them. She resists, she screams, she fights, but they mate with her anyway. It is either a good act, or it’s the equivalent to rape. Primatologists try not to use that term. People tend to get upset.26


Lemurs:

In Lemur catta, the incidence of aggression within groups is high, particularly between males. Aggression takes the form of-chasing, cuffing, scent marking, and, in males, stink fighting … Acts of submission include retreat or cowering as a dominant animal approaches, and low-ranking males habitually walk with lowered head and tail carriage, lagging behind the group and generally avoiding other animals. Females are much less frequently aggressive than males, and the female dominance hierarchy is less easy to detect, although the few agonistic encounters observed suggest that it is stable. Yet, “at any time … a female may casually supplant any male or irritably cuff him over the nose and take a tamarind pod from his hand.”27


Monkeys:

In most monkeys with multimale groups, tolerant or cooperative relationships among males are rare or unknown. Male-male grooming, for example, is virtually nonexistent in rhesus monkeys … [I]f grooming ever occurs, it is given entirely by subordinates to dominant males …, unlike the more reciprocal system in chimpanzees. As another example, Watanabe … studied alliance formation among Japanese macaques. Out of 905 cases only 4 alliances were between adult males. Relationships between males in these groups are thus primarily competitive.28


Stumptail monkeys:

The two newcomer adult females … were thus repeatedly mounted as well as bullied by the three subadult males and the higher ranking juvenile male throughout their stay. This forced mounting might be considered as rape, in the sense that the female was obviously unreceptive and unwilling. She kept crouching while the male forcibly lifted her hindquarters, shook and even bit her, and ignored her screams and dismount signals.29


Stumptail monkeys:

At the very moment that the round-mouthed expression appeared on the female’s face and the hoarse vocalizations were uttered, the equipment registered a sudden acceleration of her heart rate, from 186 to 210 beats per minute, and intense uterine contractions.

Actually, this experiment concerned reassurance behavior. The female’s partners were other females … [It] can be demonstrated that the sexual posture that stumptails often adopt during reconciliation is accompanied by physiological signs of orgasm. This is not to say that sexual climax is achieved during every reconciliation.… [Nature] has provided stumptails with a built-in incentive for making up with their enemies.30


Colobine monkeys:

[I]nfants are often passed around to other females from soon after birth. This pattern may continue for the first few months of life. In particular contrast to some macaques and baboons, every colobine infant has free access to every other infant, and females of all ranks have free access to all infants. Swapping of infants may be one of the roots of the [comparatively] nonaggressive colobine society …

A very interesting feature of colobine intertroop encounters is the fact that they have readily available means of avoiding such contact. As arboreal animals occupying upper story vegetation which provides a relatively unobstructed view of surroundings, and as possessors of loud, sonorous vocalizations, colobine groups could rather easily avoid contact. Nevertheless, contact is frequent. Colobines maintain troop separation by one or a combination of the following: variable movement patterns, the male whoop vocalization, and male vigilance behavior.

… Excitement is high during this stage, which includes tremendous leaping and running through the tree tops, as is evidenced by frequent defecation and urination. Another indication of high excitement and/or tension is the fact that males may have penile erections …

The most common dominant signals include grinning, staring, biting air, slapping the ground, lunging, chasing, bobbing the head, and mounting another animal. Submissive gestures include presenting the hindquarters, looking away, running away, turning one’s back to another animal, and being mounted … The higher the animal’s position in the dominance hierarchy, the wider the personal space it controls which a less dominant animal may not enter without first clarifying its intent.31


Monkeys:

[A]s long as the infant monkey should be riding on its mother, whether it is injured or even dead, its mother will continue to carry it. If she stops carrying it, an adult male is likely to go to her and to bark at her and in this sense make it clear to her that she should continue carrying the infant. We had one case in our small colony at Berkeley where a mother carried her dead infant for two days and dropped it, and then the dominant adult male of the troop picked the infant up and carried it for two more days before discarding it.32


Vervet monkeys:

In 1967, T. T. Struhsaker reported that East African vervet monkeys gave different-sounding alarm calls to at least three different predators: leopards, eagles, and snakes. Each alarm elicited a different, apparently adaptive response from other vervets nearby. Struhsaker’s observations were important because they suggested that nonhuman primates might in some cases use different sounds to designate different objects or types of danger in the external world …

Seyfarth, Cheney, and Marler … began by tape-recording alarm calls given by vervets in actual encounters with leopards, eagles, and snakes. Then they played tape-recordings of alarm calls in the absence of predators and filmed the monkeys’ responses.

[W]hile adult vervet monkeys restrict their eagle alarm calls to a small number of genuine avian predators, infants give alarm calls to many different species, some of which present no danger. Eagle alarms given by infants, however, are not entirely random and are restricted to objects flying in the air … From a very early age, therefore, infants seem predisposed to divide external stimuli into different classes of danger. This general predisposition is then sharpened with experience, as infants learn which of the many birds they encounter daily pose a threat to them …

[But] … experiments offer no proof that primates in the wild recognize the relationship between a vocalization and its referent.33


Squirrel monkeys:

The Gothic variety of the male squirrel monkey provides a most graphic example. He signals 1) his aim to dominate another male, 2) his intention to assault him, and 3) his amorous ideas about a female—all three—by shoving his erect phallus into the face of the other monkey while grinding his teeth. The courtship display is identical to the aggressive display. Ethologists have found this crossed-wire phenomenon in numerous reptilian and lower forms.34


Hamadryas baboons:

[Y]oung males … present in situations which provoke fear. They employ sexual approach in obtaining access to each other and to entice a fellow for play. They masturbate and mount each other. They mount and are mounted by adult males and by adult females, their heterosexual activities not provoking aggressive responses from the overlords. They engage in manual, oral and olfactory ano-genital examination with animals of their own age and with adults of both sexes. They frequently end a sexual act by biting the animal with whom they have been in contact. This end to sexual activity, which is not usually seen in the behaviour of adults, often appears to be playful.35


Baboons:

Sir Andrew Smith, a zoologist whose scrupulous accuracy was known to many persons, told me the following story of which he was himself an eye-witness; at the Cape of Good Hope an officer had often plagued a certain baboon, and the animal, seeing him approaching one Sunday for parade, poured water into a hole and hastily made some thick mud, which he skilfully dashed over the officer as he passed by, to the amusement of many bystanders. For long afterwards the baboon rejoiced and triumphed whenever he saw his victim.36


Baboons:

In Abyssinia, Brehm encountered a great troop of baboons who were crossing a valley: some had already ascended the opposite mountain, and some were still in the valley: the latter were attacked by the dogs, but the old males immediately hurried down from the rocks, and with mouths widely opened, roared so fearfully, that the dogs quickly drew back. They were again encouraged to attack; but by this time all the baboons had reascended the heights, excepting a young one, about six months old, who, loudly calling for aid, climbed on a block of rock, and was surrounded. Now one of the largest males, a true hero, came down again from the mountain, slowly went to the young one, coaxed him, and triumphantly led him away—the dogs being too much astonished to make an attack.37


Titis and other small monkeys:

Hidden among the tangled branches and vines of the Neotropical forests are the most paternal of primate fathers. The monogamously mated males of the small titi (Callicebus sp.) and night monkeys and the tiny Callimiconidae and Callitrichidae are unique in the intensity and duration of their relations with infants … Males of these species share all parental duties except nursing, and although the extent of participation is quite variable within species, they are generally the major caretakers of infants …

Males in these species are often strongly attracted to infants. Immediately after birth, they have been observed trying to sniff, touch, or hold the still-bloody newborn, and they sometimes even lick off the covering birth fluids … Within hours of birth, males carry infants on their backs, groom them, and protect them … Large portions of a male’s day are devoted to infant care, and the most devoted fathers return their infants to the mother only to suckle …

Males also permit infants to take food from their hands and mouths … The food items shared are those that infants have difficulty obtaining or processing themselves, such as large mobile insects or hard-shelled fruit …

Fiercely protective, males will defend infants against any real or imagined threat. In captivity, tiny lion tamarin males have flung themselves against intruders as intimidating as woolly monkeys, macaques, and humans.38



* Apes are bigger and smarter than monkeys, and lack tails. The apes are the chimpanzees, gorillas, gibbons, siamangs, and orangutans. The siamangs are about as closely related to gibbons as chimps are to humans.* The fact that in every human ethnic group and culture males have been on average larger than females has not escaped the notice of primatologists. It may have something to do with the penchant of men for sexism, coercion of women, rape, and harems when they can get away with it. The key question is to what extent anatomy is destiny, a point to which we will return.* Something similar happened when a number of fugitive Englishmen, without a well-established dominance hierarchy (the alpha male and his close followers had been put overboard in a small boat), along with a few Polynesian women settled tiny Pitcairn Island in 1790, after the mutiny on H M S Bounty.* Those who study chimps and bonobos, so the joke goes, are called panthropologists.




Chapter 18




THE ARCHIMEDES OF THE MACAQUES


Some ascribe this to his natural genius; while


others think that incredible effort and toil


produced these, to all appearances, easy and


unlabored results. No amount of investigation


of yours would succeed in attaining the proof,


and yet, once seen, you immediately believe


you would have discovered it—by so smooth


and so rapid a path he leads you to the


conclusion … Such was Archimedes.


PLUTARCH


“Marcellus,” in The Lives of the Noble


Grecians and Romans1


We humans have not evolved from any of the two hundred other primate species alive today; rather, we and they have evolved together from a succession of common ancestors. As we reconstruct the primate family tree, we discover who our closest relatives are. The behavior of the primates varies so widely, even between species in the same genus, that it really does make a difference for our view of ourselves which ones are our nearest relatives.

The answer, as we’ve already described, seems to be that the chimps are our closest kin, sharing some 99.6% of our active genes. We know from DNA sequencing, as you would of course suspect, that bonobos and ordinary chimps are a lot more like each other than either of them are like us.2 But 99.6% is very close. We must share many characteristics of both. (Indeed, there must be behavioral traits that we share with our most distant primate cousins.)

By using molecular and anatomical evidence, together with the record in the rocks, the entire family tree of primates can be drawn, at least approximately, and a timeline placed upon it. The evidence from the bones and from the molecules are not in perfect accord, although they are beginning to converge; in this book, we have given weight to gene sequencing and DNA hybridization data. According to the molecular evidence, gorillas branched off from the evolutionary line leading to us about 8 million years ago; the still unidentified, now-extinct common ancestor of humans and chimpanzees separated from the gorillas maybe a million years later. Very quickly thereafter, the lines to chimps and humans began evolving toward their separate destinies.3 On a planet that’s been inhabited a thousand times longer, that’s pretty recently, as recent as the last two weeks in the life of a fifty-year-old human. This doesn’t mean that humans and chimps themselves began 6 million years ago; only that our common twig in the evolutionary tree branched out then.

To understand a little more about our primate nature and its development, let’s cast our minds back toward the end of the Mesozoic Age, around 100 million years ago. That would be about a year ago in the life of a middle-aged person. There were mammals even then; but they were not easy to find. The daytime was ruled by the dinosaurs; among them, some of the most fearsome killing machines ever to evolve on land. Our mammalian ancestors, it is thought, were timid, weak, and small; they were in fact typically the size of mice. Like all reptiles and amphibians today, some of the dinosaurs may (this is still a controversial point) have been cold-blooded; if so, in the chill of the evening, especially in winter, they closed up shop—especially the smaller ones that both preyed on mouse-sized mammals and were more vulnerable to the cold. But the mammals themselves were warm-blooded, and so could stay out all night.

Imagine a moonlit darkness in which their adversaries lay senseless, strewn across the landscape in stupors of sleep. This was the chance for our ancestors to scamper about their humble business—catching grubs, nibbling leaves, mating, caring for the young. But to function well in the dark, they had to be very good at using senses other than sight; and in that epoch the mammalian brain evolved along with elaborate machinery for enhanced hearing and smell, their hedge against whatever dinosaurs hunted at night. .

Asleep in burrows during the day, our ancestors perhaps tossed fitfully, dreaming daymares filled with row after row of needle-like teeth and nimble, hair-raising scampers to safety. They may have been frightened all their lives, their hearts in their throats at every daylit step, longing for nightfall.

Sixty-five million years ago, a bolt from the blue—the impact of a small world—seems to have cataclysmically altered the planetary environment, wiping out the dinosaurs and permitting the mammals, wholly insignificant until then, to flourish and diversify. We do not know if there were primates so early, or if some other mammal quickly evolved into the first primate. We do know from fossil evidence that tiny monkey-like beings, weighing perhaps a few ounces, with teeth about a millimeter long, lived in what is today Algeria just after the extinction of the dinosaurs.4 By 50 million years ago (six months ago in the life of our fifty-year-old) there were arboreal primates living in subtropical Wyoming.5 The canine teeth of the males were twice as long as the females’. If we can judge by what this difference means in contemporary monkeys, the males bullied the females, established dominance hierarchies, competed with each other, and probably maintained harems. All that’s been with us since the beginning of the primate order.

The first primates are judged to have been much more like early mammals (with longer snout, eyes to the sides of the head, and claws) than are modern monkeys, apes, and humans. The so-called “lower” primates, or prosimians—lemurs or lorises, say—may be something like the earliest primates. You can see that they’re nocturnal at a glance: Their eyes are appealingly large for their faces, the larger aperture being an adaptation for night vision in a world illuminated only by the moon and the stars.

They probably communicated in part by spraying scents from specialized glands.* They had brains—large for their body size—to think with, stereoscopic vision to see with, and hands to manipulate the environment. Typical primate dominance hierarchy rituals had probably already appeared, including both sexes presenting their rears as a gesture of submission to the dominant male.

The early evolution of primates was marked by a profound transformation of creatures of the night into habitués of daylight; a corresponding suppression of the sense of smell6 and elaboration of vision; developing facial muscles so moods could be communicated by expressions; a still more powerful bond between mother and child; a longer period of infantile dependence; and an improving ability of the newer, higher brain centers of the cerebral cortex to moderate aggression and other behavior patterns emanating from the older, lower layers. All this in turn led to major changes in primate society: The less aggression, the more a true communal life is possible; the longer the childhood, the more parents can teach their young. Alliances and support groups, reconciliation, reassurance, forgiveness, remembering the past behavior of specific individuals, and planning future actions swiftly evolved. Our ancestors were by now well along a path toward greater alertness, intelligence, communications skills, love.

After the extinction of the dinosaurs, mammals moved out into the daylight. For a while, they must have felt safe and free. But the growing, multiplying, and diversifying mammals eventually became too good a meal to pass up. They began to eat each other. And new predators evolved, including birds of prey. The day shift became increasingly dangerous. For example, in a study of modern South American harpy eagles, 39% of the “prey items” returned to the nest turn out to be body parts of monkeys.7 In daylight you have to be on your toes. Mutual defense—scanning the skies, say, and air raid sirens when an eagle is spied—becomes vital.

Foraging baboons, faced with predators, typically respond by closing ranks and moving faster.8 Certain collective behavior that we readily describe as military constitutes an adaptive response of very ancient standing to the threat of predation. Competent predators can force the potential prey to evolve rapidly—toward binocular vision, arboreal acrobatics, mutual support, quickly disinhibited combat skills, intelligence, and general military virtues.

Monkeys are born with an ability to recognize the significance of various facial expressions—although just how to respond to such expressions depends on experience and training. There are single brain neurons that are preferentially triggered when the monkey sees the eyes or mouth or fur of another monkey. There is even a kind of brain cell specifically responsive to a crouching or bowing posture. Facial expressions and body posture have a meaning in the primates that’s hardwired, and not merely a matter of social convention. The male rhesus monkey’s come-hither look is to thrust out his chin and pucker his lips; if you’re a rhesus monkey (of either sex), it’s important, even early in your career, to know what this means.

One of the uses to which the evolving primate brain has been put is the storing up of grudges. Monkeys generally make up—often by ceremonially mounting each other—within minutes after a fight. Chimp males, with females frequently in a peacemaking role, may take hours or days. But among themselves the females are less forgiving; they may hold grudges for the rest of their lives. Humans of both sexes can take anywhere from moments to millennia. Even among monkeys, a smoldering resentment against an individual is often broadened to encompass his or her relatives. Among the many new social forms invented by the primates are feuds and vendettas, sometimes extending over many generations—intimations of the beginnings of history.

As in most mammals, primate aggression, dominance, territoriality, and the sex drive are mediated by testosterone circulating in the blood, and generated mainly by the testicles. Almost certainly this was true of the earliest primates, and long before. The more testosterone and other androgens the developing fetal brain receives, the more of these masculine characteristics the animal will exhibit when he grows up. The lower the testosterone levels in a male, the more subdued will be these proclivities and the more likely that he will present himself for mounting by other males. But the testosterone levels also respond to the mantle of leadership. When presented with females in estrus and no high-ranking males around, the testosterone level of lower-ranking males soars. Within certain limits, primates rise to the occasion. The office makes the monkey.

Males of many primate species (although, on average, not humans) show a marked preference for female sexual partners who have already produced offspring; younger females may have to make special efforts to be alluring.9 We have described the vigilance with which chimp alpha males guard their females, but only during ovulation. Nevertheless, sex has evolved in primates into something much more than simply the means for the replication and recombination of DNA sequences. Year-round, virtually compulsive sex with many partners—described by human observers as “promiscuous,” “depraved,” “perverse,” and “indiscriminate”—is there for a reason. It serves as a mechanism of socialization. This is clearest among the bonobos. Despite sexual jealousy, it holds the group together. It provides bonds of affection, common goals, means of identification with others, and a gentling of dangerous aggression. The essence of primate living arrangements is a gregarious communal life, which partakes of many recognizable aspects of human culture and society. One of the chief motivations for this communal life is sex.

Adult role models are essential among animals in which childhood learning plays so central a role. Dominance hierarchies soften violence (but not aggression) within the group. Cooperation is important in any hunt, critical in hunting large animals, and sometimes essential in evading predators. In a survey of thirty primate species in the wild, the probability that any given individual will be eaten by year’s end is found to be one chance in sixteen.10 Evading predators must be very high on the primate agenda—and communal life provides early warning and collective defense.

Vervet monkeys have ventured a little out of the comparative safety of the forest and into the open savanna, where there is less cover for them, and more danger. By playing recordings of their calls back to them, they reveal that they have specific, readily understood alarm cries that elicit specific actions—for a python or black mamba (whereupon all stand on tiptoes and peer anxiously about them in the grass), for a Martial eagle, (whereupon all look up into the sky and dive into deep foliage), and for a leopard (whereupon all quickly scramble up into the trees). Different predators elicit different cries and different evasive behavior. The responses are in part learned. Infants frantically sound the eagle alarm even when a non-raptor is spied flying overhead, and sometimes in response to a falling leaf. Gradually, they get better at making these distinctions. They learn from experience and from others. They have a range of other grunts, some of which scientists think they understand; vervets leave at least a superficial impression of conversing with each other. Gregariousness, by several different routes, spurs social intelligence, which seems to be, of all the species of life on Earth, most highly evolved in the primates.

The vervet fear of snakes is shared by baboons, chimps, and many other primates. You expose wild rhesus monkeys to snakes and objects that look like snakes and they jump out of their skins. Do the same experiment with laboratory-raised rhesus monkeys who have never seen a snake and, although some of them are afraid, you find that they’re much less distraught. In one experiment the wild chimps’ snake phobia became almost manageable when every time the chimp saw a snake it also was offered a banana.11 So is the fear of snakes not hereditary, but somehow taught by mothers to their babies? Or is there an inborn fear that’s softened in laboratory monkeys because they become habituated to harmless, snake-like objects—hoses, for example? Which is it: heredity or environment? Is knowledge of what a snake looks like, and that snakes mean primates no good, encoded in the DNA? Or are baby primates just watching adults closely and copying what they do?

Almost certainly the answer is a mix between the two. There seems to be an inborn snake-aversion program in the brains of primates. But this is not a closed program, inaccessible to new information from the outside world. Instead it’s an open program that can be modified by experience—for example, “I’ve seen a lot of snakes in my time that don’t do me much harm, so I’ll be a little more relaxed around them,” or, “Every time I see a snake, a banana miraculously appears; snakes have their good points too.” Most primate programs are open, adaptive, malleable, adjustable to new circumstances—and therefore necessarily partaking of ambivalence, complexity, inconsistency.

In a typical modern chronology,12 the line that would lead to us split off from Old World Monkeys about 25 million years (m.y.) ago; from the gibbons, 18 m.y. ago; from orangutans around 14 m.y. ago; from gorillas some 8 m.y. ago; and from the chimps approximately 6 m.y. ago. Bonobos and common chimps went their separate ways only about 3 m.y. ago. Our genus, Homo, is 2 million years old. Our species, Homo sapiens, is maybe 100,000 to 200,000 years old—the equivalent of the last day in the life of that fifty-year-old.

Committed to a communal social life, under intense selection pressure from predators, with brains evolving rapidly and education of the young effectively institutionalized, the primates have been developing new forms of intelligence. Their curiosity, experimental bent, and intellectual quickness are partly responsible for their success.

——


Here is an account, by a Japanese primatologist, of a remarkable set of events that transpired in a colony of macaques isolated on a small island called Koshima. Initially, in 1952, there were only twenty of the monkeys; the number almost trebled over the following decade. The natural food supply on Koshima was inadequate, so the monkeys had to be provisioned—with sweet potatoes and wheat dumped on the shore by the primatologists who were observing them.

As anyone knows who’s ever been to a picnic at the beach, sand sticks to food and makes it unpleasantly gritty. In September 1953 a one-and-a-half-year-old female named Imo figured out that she could rinse the sand off her sweet potatoes by dunking them in a nearby brook.After Imo, the next individual to learn potato washing was Imo’s playmate, who did so in October. Imo’s mother and another male peer began to wash in January 1954. In subsequent years (1955 and 1956), three of Imo’s lineage (younger brother, elder sister, and niece) and four animals from other lineages (two were a year younger and two were a year older than Imo) started to do so. Thus, with the exception of her mother, all the individuals that learned potato washing quickly were either peers or young close relatives of Imo …After 1959, features of information transfer changed. Sweet potato washing was no longer a new mode of behavior: when infants were born, they found most of their mothers and elders washing potatoes and learned this behavior from them as they learned the group’s usual food repertoire. Infants are taken to the edge of the water during the period when they are dependent on mothers’ milk. While their mothers wash potatoes, infants watch carefully and put into their mouths pieces of potatoes that mothers drop in the water. Most of the infants acquire potato washing around 1 to 2.5 years old …[I]n the second period (1959-present, the period of “precultural propagation”), acquisition of potato washing occurred independent of sex and age. During the second period, virtually all individuals … acquired this habit through their mothers or playmates when they were infants or juveniles.


But there was still the problem of sandy wheat—until Imo’s second epiphany:In 1956, when Imo was 4 years old, she took a handful of mixed wheat and sand to the brook. When it was dropped on the water, the sand sank and the floating wheat could be skimmed off the water’s surface, now clean again. This “placer-mining” technique* was also adopted by some of the other monkeys, and soon more and more animals learned it …Compared to potato washing, placer mining was quite slow to propagate …Placer mining appears to require more understanding of complex relations between objects and may be particularly difficult to learn because a monkey must “discard” his food first, while in potato washing he can keep the potato from the beginning to the end.13


Imo was a primate genius, an Archimedes or an Edison among the macaques. Her inventions spread slowly; macaque society, like traditional human societies, is very conservative. Perhaps the fact that she came from a high-ranking family in a species given to hereditary matriarchy aided acceptance. As is usually true, adult males were the slowest to catch on, obstinate to the last; a female invented the process, other females copied her, and then it was taken up by youngsters of both sexes. Eventually, infants learned it at their mother’s knee. The reluctance of the adult males must tell us something. They are fiercely competitive and hierarchy-ridden. They are not much given to friendships or even to alliances. Perhaps they felt impending humiliation—if they were to imitate Imo, they would be following her lead, becoming in some sense subservient to her, and thereby losing dominance status. They would rather eat sand.

No other group of macaques anywhere in the world is known to have made such inventions. By 1962, it is true, macaques on other islands and the mainland, recently provisioned with potatoes, began washing their food before eating it. But it is unclear whether this was due to independent invention or to cultural diffusion: In 1960, for example, Jugo—a macaque who had become adept at washing potatoes—swam from Koshima to a nearby island where he stayed for four years and may have trained the resident macaques.14 Perhaps there were other macaque Archimedes; perhaps not. Imo is the only one we know for certain.

It took a generation for these two obviously useful inventions to become widely accepted.15 The conservative, near immobility of popular prejudice, the reluctance to adopt a new practice even if its advantages are clear, is a tendency not restricted to Japanese macaques.16 Perhaps the stolidity of the adult males is partly a matter of learning abilities declining with age. Human teenagers seem so much more adept than their parents at, say, operating a personal computer or programming a videocassette recorder. But this doesn’t explain why adult female macaques learned so much more readily than their male counterparts.

We can see how such inventions made in different, nearly isolated, groups can lead to cultural differentiation even in monkeys. A much more innovative species of primate, in which various groups are in occasional contact, conflict, or competition, might, we would guess, devise spectacular new forms of culture and technology.

——


An early Algerian myth held that long ago apes could talk, but were rendered mute for their transgressions by the gods. There are many similar stories in Africa and elsewhere.17 In another widespread African story, apes can talk, but prudently refuse to do so—because talking apes, their intelligence in this way made manifest, will be put to work by humans. Their silence is proof of their intelligence. Occasionally the indigenous people would introduce a visiting explorer to a chimp with many remarkable skills and tell him that it could also speak. But, at least while the explorer was there, none ever did.

Lucy was a chimpanzee celebrity. She was one of the first of the apes to learn to use a human language. The mouth and throat of the chimp are not configured for speech as ours are. In the 1960s, the psychologists Beatrice and Robert Gardner wondered whether chimps might be intellectually capable of language but prevented from speaking by the limitations of their anatomy. Chimps have phenomenal dexterity. So the Gardners decided to teach a chimp named Washoe a gestural language, Ameslan, the American sign language used by hearing-impaired humans. Here each gesture can represent a word, rather than a syllable or a sound, and in this respect Ameslan is more like Chinese ideograms than the Greek, Latin, Arabic, or Hebrew alphabets.

Young female chimps proved to be adept pupils. Some of them eventually acquired vocabularies of hundreds of words. Julian Huxley—T. H. Huxley’s grandson, and a leading evolutionary biologist—had argued that “plenty of animals can express the fact that they are hungry, but none except man can ask for an egg or a banana.”18 Now there were chimps eagerly requesting bananas, oranges, chocolate candies, and much else, each represented by a different sign or symbol. Their communications were often clear, unambiguous, and apparently in context, as has been attested to by delighted audiences of hearing-impaired people watching films of signing chimps. They were able, it is said, to use their signs in a fairly consistent elementary grammar, and to invent from the words they knew phrases that they had never before encountered. Chimps were found to generalize a word such as “more” into new contexts—such as “more go” and “more fruit.”19 A swan evoked the spontaneous neologism, in independent and widespread use among humans, “water bird.”

Lucy was one of the first. It was she who signed “candy drink” after first tasting a watermelon, and “cry hurt food” after her first experience with a radish. She became, it is said, able to distinguish the meaning of “Lucy tickle Roger” from “Roger tickle Lucy.” Tickling is close to grooming. When idly turning the pages of a magazine, Lucy made the sign for “cat” when she turned to a picture of a tiger, and “drink” when she came upon a wine advertisement. Lucy had a human foster mother; she was, after all, only a few years old during the whole of her laboratory experience with language, and young chimps especially crave emotional support. One day, when her foster mother, Jane Temerlin, left the laboratory, Lucy gazed after her and signed, “Cry me. Me cry.”

Ameslan-literate apes have often been spied signing to themselves when they thought no one else was present. Perhaps this was just wordplay, trying to get the new skill down pat. Or perhaps it was an experiment to see if they could conjure “fruit,” say, out of the air with no humans present, just by producing the right words. It had worked well enough when humans were around.

To what extent Lucy and her fellows understood the gestural language they were using, and to what extent they were merely memorizing sequences of signs whose true meaning they failed to grasp, is a subject of scientific debate. To what extent young humans learning their first language do the one or the other is also subject to debate.

Perhaps only the hits were recorded and not the misses; that is, maybe Lucy and other chimps judged Ameslan-literate generated a wide range of signs more or less at random which, when they made contextual sense, were written up by the human observers and discussed at scientific meetings, but which, when irrelevant or unintelligible, were ignored. This is the anecdotal fallacy* that haunts this branch of science. But the anecdotes are plentiful and striking.

One of the most thoroughgoing examinations of the linguistic and grammatical abilities of apes was done by the psychologist Herbert Terrace and his colleagues, who recorded on videotape nearly twenty thousand signing attempts generated by a male chimp named Nim.20 He mastered over one hundred different gestural signs. Nim would regularly sign “Play me” or “Nim eat” in context and with apparent understanding. But there was no evidence, Terrace concluded, that Nim put more than two signs together in any consistent manner appropriate to the context. The average length of his sentences was less than two words long. His longest recorded sentence was “Give orange me give eat orange me eat orange give me eat orange give me you.” It does seem a little frantic, but oranges are tasty, chimps are not known for their patience, and anyone who has spent time with a small excited child will recognize the syntax. Note that four of the words are non-redundant (“give me orange you”), and that no words irrelevant to this urgent request are included among the sixteen. Emphasis through repetition is common in human languages. But the simplicity of chimp sentences has rendered their use of language unimpressive in the minds of many psychologists and linguists. Nim was also belittled for interrupting his trainers’ signing with his own, for being too imitative (repeating remarks of his trainer), and for not inventing grammatical rules such as the subject-predicate sequence.

This work has in turn been criticized. Chimps require close emotional ties for social tasks and, one would think, especially for something as difficult as language; instead, Nim had sixty different trainers over a four-year period. There is a tension between a loving, one-on-one environment that might be needed to teach language skills and the emotionally sterile protocols needed so that scientific results are, with high reliability, uncontaminated by the enthusiasm of the experimenters. It has frequently been found that apes sign most creatively in spontaneous circumstances in their everyday life, and not in experimental sessions. Too, there was great emphasis on drill in the Nim experiments, the very opposite of spontaneity. The complaint about Nim interrupting the signing of his trainer has itself been belittled, because Ameslan speakers may sign simultaneously without stepping on each other’s lines, an advantage of signing over speech. Delayed imitation is just what human children do when they first learn language. For all these reasons, just how much grammatical dexterity apes have is still an open question.21

But clearly chimps can use something like the rudiments of language with much greater facility than had been thought possible before the experiments of the Gardners. They can unambiguously associate certain signs with certain people, animals, or objects—unsurprising when there are monkeys with different alarm cries and evasion strategies for different species of predators. Chimps have mastered an elementary vocabulary of a few hundred words, comparable to what a normal human two-year-old can do. Chimps who have some knowledge of these signs and who are raised together have been known spontaneously to sign to one another. There is at least one case in which a young chimp, uninstructed by any human, is said to have learned dozens of signs from another chimp knowledgeable in Ameslan.22

“We may consider it proven,” said the psychologist William James, “that the most elementary single difference between the human mind and that of brutes lies in this deficiency on the brute’s part to associate ideas by similarity.” He held this to be a more fundamental cause of human uniqueness than reason, language, and laughter—all of which, he taught, emerge from recognizing similarities among ideas.23

Some chimps were taught a common symbol to describe any one of three foods, and another to describe any one of three tools. Then they were taught the individual names of other foods and other tools and asked to put them in the proper categories—not the new foodstuffs or tools themselves, but the arbitrary names of the new foodstuffs and tools. They did exceptionally well.24 How is this possible, unless chimps reason, form abstract ideas, and “associate ideas by similarity”? Another domesticated chimp, Viki Hayes, was given two piles of pictures, one of humans, the other of nonhumans, and then handed a stack of additional pictures and invited to categorize. Her performance was perfect, with one small exception: She placed the picture of herself among the humans.

The psychologist Sue Savage-Rumbaugh25 and her colleagues devised a keyboard with 256 lexigrams on its two sides. Each lexigram stands for something of interest to a chimp—“tickle,” “chase,” “juice,” “ball,” “bug,” “blueberry,” “banana,” “outdoors,” “videotape,” and so on. The lexigrams do not depict their referent; rather, they show geometrical or abstract figures that only by arbitrary convention are connected with what they stand for. The scientists tried to teach this lexigraphic language to an adult bonobo, but she was an indifferent student. Her six-month-old son, Kanzi, often accompanied his mother to these training sessions and was mainly ignored by the scientists. Two years later, having observed the laboratory routine in depth but never having been trained (for example, by being given a banana for typing the banana lexigram), Kanzi demonstrated that he was learning what they were trying to teach his mother. (His interest eventually became hard to miss: He would leap on her hand, her head, or the keyboard at the moment she was about to select a lexigram.) The focus of the study switched to him.

By age four he had mastered the board, and would routinely use lexigrams to request, confirm, imitate, choose an alternative, express an emotion, or just comment. He would indicate a future course of action and then do it. In combining two action lexigrams, he would predict (or better, reveal) the impending sequence of events; if he typed “chase, tickle” he would chase and then tickle the experimenter or another chimp, and only very rarely tickle before chasing. Kanzi typed “hide peanut,” and then did just that. It seems hard to deny that Kanzi has a mental image of his intended future actions, and in appropriate sequence. As time went on, he developed other grammatical rules, especially putting the action before the object, rather than vice versa (“bite tomato,” rather than “tomato bite”). Inventing grammar is much more impressive than merely being taught it.

Still, after some years about 90% of Kanzi’s utterances were only a single symbol;* rarely did they comprise more than two symbols. This is the same pattern found for Nim. Perhaps we are coming up against some fundamental limitations in chimp capacity for language.

Kanzi has shown, again via an accidental discovery, that he can understand hundreds of words of spoken English. Place earphones on his head, situate yourself in another room, make a request of him through a microphone, and the video camera reveals him doing what he is asked. Done this way, no gestural cues can be unconsciously communicated from human to ape. Typical of over 600 novel requests, perfectly complied with, were “Put the backpack in the car,” “Do you see the rock?… Can you put it in the hat?” “Take the mushrooms outdoors,” “Knife the orange,” “Eat the tomato,” and “I want Kanzi to grab Rose.” Even some of Kanzi’s errors are not so bad. Asked “Can you put the rubber band on your foot?” he promptly put it on his head.26 His performance was comparable to that of a 2½-year-old human who was tested in the same set of experiments. Other bonobos are also found to understand spoken English.

Kanzi loves to play ball. Hide a ball in one of seven designated sites in the laboratory’s fifty-five-acre forest, tell him by lexigram or spoken word where the ball is, and Kanzi with high accuracy makes for the site, searches, and finds the ball.27 In this case there is a reward for understanding spoken English. But in most cases Kanzi receives no reward except the approval of humans and perhaps some gratifying sense of the power of communication. The motives of a young child learning language may not be very different.

In a different laboratory, a chimp named Sarah was able to recognize that red characterized an apple more than green did (she had not been exposed to the Granny Smith variety), and a square with a stem was a better representation of an apple than a square without a stem. She was also able to associate the words for each of these properties of an apple with the word for apple—and these words were not in Ameslan, but in a symbolic language of plastic tokens she had been taught, the tokens not resembling the objects in question.28 (“Apple,” for example, was represented by a small blue triangle.) How is this possible, unless chimps are able to abstract and categorize?

Other experiments have shown chimps capable of reasoning by analogy and by transitive inference, described by the discoverers of this aspect of chimp thought as “ ‘A r B, B r C, therefore A r C,’ where r is some transitive relation, such as greater than.”29 (There may, for all we know, be critics who do not even understand the preceding sentence but who deny that chimps reason.) Still other experiments have been interpreted as showing that chimpanzees impute states of mind to others, or, as the psychologists David Premack and G. Woodruff put it, that chimps have “a theory of mind.”30

Where chimps are linguistically deficient, at least so far, is in grammar and syntax. They are bereft of subordinate clauses, articles and prepositions, tenses, conjugation of verbs, and the like—as are small humans first learning language. The absence of such grammatical machinery prevents the lucid expression of even fairly simple ideas; misunderstandings tend to accumulate. Compounded by small vocabularies, it’s a little like a middle-aged American, relying on barely remembered high-school French, attempting to be understood in rural Provence. A better analogy might be the “pidgin” languages that emerge at the interface between two or more fully realized but very different human languages; despite their linguistic facility, the speakers revert to something like chimpish. Oddly, no one has made a serious and systematic effort to teach apes grammar and syntax,31 so we can’t be sure it’s beyond their reach. “Until then,” writes a modern linguist, “one cannot entirely close off the possibility, unlikely as it may be, that apes could acquire language in its fullest sense.”32

Savage-Rumbaugh and her co-workers toy with the possibility that chimps and bonobos exhibit impressive facilities to learn something of human languages because they have their own languages, vocal or gestural, that we have not yet deciphered.33 In announcing the location of prey, or predators, or a hostile patrol, rudimentary language would be strongly favored by natural selection. Long before humans and chimps went their separate ways, considerable aptitudes for thought, invention, and language were probably percolating in our primate ancestors.

But partly because of Terrace’s work, and partly because of the perceived difficulty of doing clean, controlled, non-anecdotal experiments on so emotional a being as a chimpanzee, financial support for many of these studies has nearly disappeared. In one case, the colony where apes had been taught Ameslan had fallen on hard times. Years had passed. Support was drying up. No one seemed interested in conversing with the chimps anymore. The grounds had become weedy and overgrown. The inmates were about to be shipped to laboratories for medical experimentation. Before the end, they were visited by two people who had known them in the old days. “What do you want?” the visitors asked in Ameslan. “Key,” two chimps are said to have signed back from behind bars, one after the other. “Key.” They wanted out. They wanted to escape. Their request was not granted.34

——


When chimps approach sexual maturity, their behavior changes. Both sexes are then much stronger than humans and given to occasional, unpredictable bouts of rambunctiousness and violence. So as the chimps get older, almost inevitably the experimenters find themselves driven to use steel cages, collars, leashes, and electric cattle prods. The chimps must feel, bit by bit, betrayed by the humans and less inclined to cooperate in their strange language games. Accordingly, back in the days when the research was generously supported, it was thought prudent to terminate experiments on teaching chimps language—requiring, as it does, close face-to-face daily contact—when they begin to mature. As a result, we do not know what the linguistic abilities of an adult chimpanzee might be. Lucy, like some aging child actor, was forced into retirement just a little after puberty. The laboratory in which she had demonstrated her accomplishments in sign language was closed.

Jane Goodall, who had by then spent a decade and a half living with chimps in the wild, was astonished on meeting Lucy:Lucy, having grown up as a human child, was like a changeling, her essential chimpanzeeness overlaid by the various human behaviours she had acquired over the years. No longer purely chimp yet eons away from humanity, she was man-made, some other kind of being. I watched, amazed, as she opened the refrigerator and various cupboards, found bottles and a glass, then poured herself a gin and tonic. She took the drink to the TV, turned the set on, flipped from one channel to another then, as though in disgust, turned it off again. She selected a glossy magazine from the table and, still carrying her drink, settled in a comfortable chair. Occasionally, as she leafed through the magazine she identified [in Ameslan] something she saw …35


In the second half of her life, Lucy lived with other chimpanzees on a small island in Gambia. Her adjustment to Africa was slow and difficult, and she becamean emaciated, hairless wreck … She had been born and raised in the United States, and in pampered upper-middle-class circumstances … Lucy, the fastidious, toilet-trained chimpanzee princess … slept on a mattress, sipped soda, developed schoolgirl crushes, and would sit in the living room during the afternoon and leaf through magazines.36


But after a year or two in Gambia, thanks to the loving care of Janis Carter, she began to adjust. She had regular contact with humans and was often the first chimp to greet visitors to the island. She was used to humans. Her relations with other chimps were more strained. She had missed out on the rollicking childhood of a chimp in the wild.

In 1987 Lucy’s skeleton was discovered. The most likely reconstruction of events is that humans came to the island, killed Lucy, probably by shooting her, and skinned her body. Her hands and feet, the very organs that had made her famous, were missing.37 Those responsible have never been found.38


ON IMPERMANENCE


In the life of a man, his time is but a moment, his being an incessant flux, his senses a dim rushlight, his body a prey of worms, his soul an unquiet eddy, his fortune dark, and his fame doubtful. In short, all that is of the body is as coursing waters, all that is of the soul as dreams and vapours; life a warfare, a brief sojourning in an alien land; and after repute, oblivion. Where, then, can man find the power to guide and guard his steps? In one thing and one alone: the love of knowledge.


MARCUS AURELIUS, Meditations39



* Ring-tailed lemur males will smear a pheromone they generate onto their tails and then wave these prominent, black-and-white-banded appendages at each other, wafting the smell into the air. This is mainly a competition for females: Apparently the most aromatic lemur tends to win the most attractive female. In one lemur species, all adult males may have their tail waves answered in the same evening, because all adult females come into heat together, by the light of the silvery (and full) moon.* Used in panning for gold.* Also called the fallacy of the enumeration of favorable circumstances. No dishonest intent is implied; it is merely one of those failures of logic that humans are prey to. We tend not to be dispassionate observers.* One expert reviewer likens this sentence to saying that “90% of the materials dug from a gold mine are not gold ore”




Chapter 19




WHAT IS HUMAN?


Having proved mens & brutes bodies on one


type: almost superfluous to consider minds.


CHARLES DARWIN


Notebooks on Transmutation of Species1


We humans are the dominant species on the planet, a status affirmed by several standards—our ubiquity, our subjugation (politely called domestication) of many animals, our expropriation of much of the primary photosynthetic productivity of the planet, our alteration of the environment at the Earth’s surface. Why us? Of all the promising lifeforms—implacable killers, professional escape artists, prolific replicators, nearly invisible beings that no macroscopic predator can find—why did one primate species, naked, puny, and vulnerable, manage to subordinate all the rest and to make this world, and others, its domain?

Why are we so different? Or are we? Unambiguous definitions of humans—definitions that include almost all members of our species, but no one else—can be produced from anatomy or from DNA base sequencing. But they fail the purpose. They explain nothing that we recognize as fundamental about ourselves. Perhaps sometime in the future we will discover that unique sequences of As, Cs, Gs, and Ts encode for particular sequences of amino acids that constitute particular proteins that catalyze particular chemical reactions that motivate particular behavior that we might agree is characteristically human. But so far no such sequence has been found.

If, then, we can discern no clear-cut distinction in our chemistry (or anatomy) that explains our dominant role, the only ready alternative is to survey our behavior. It seems plausible that the sum of our everyday activities would be sufficiently defining, but a surprisingly large number of those activities can be performed by apes. For example, here’s a description of the accomplishments of Consul, the first chimp acquired, in 1893, by the zoo in Manchester, England:[He was] able to put on his own coat and hat, seat himself in his own carriage for a drive, sit at table with company, use his knife and fork with propriety, pass his plate for a fresh supply of food, use his serviette [napkin], wash his hands after meals, put coals on the fire, ring the bell for the maid, go into the kitchen for a romp with the girls, walk into his hotel, shake hands with his friends, kiss the barmaid, smoke his pipe, and mix his own drinks.2


True, Consul’s deportment may be dismissed as mere mimicry; but that may also be said of those of us who marvel at his abilities.

Is there anything we do that’s uniquely human—that all or almost all of us, of every culture, throughout history, do and that no other animal does? You might think something along these lines would be easy to find, but the subject is redolent with self-deception. We have too much of a stake in the answer to be unbiased.

Philosophers of marauding high-technology civilizations have often argued that humans deserve a category distinct from and above all the other animals.* It is not enough that humans have a different assortment of the qualities evident in the other animals—more of some traits, fewer of others. A radical difference in kind, not some fuzzy-edged difference in degree, is needed, longed for, sought.

Most of the philosophers adjudged great in the history of Western thought held that humans are fundamentally different from the other animals. Plato, Aristotle, Marcus Aurelius, Epictetus, Augustine, Aquinas, Descartes, Spinoza, Pascal, Locke, Leibniz, Rousseau, Kant, and Hegel were all proponents “of the view that man differs radically in kind from [all] other things”; except for Rousseau, they all held the essential human distinction to be our “reason, intellect, thought, or understanding.”3 Almost all of them believed that our distinction arises from something made neither of matter nor of energy that resides within the bodies of humans, but of no one else on Earth. No scientific evidence for such a “something” has ever been produced. Only a few of the great Western philosophers—David Hume, for instance—argued, as Darwin did, that the differences between our species and others were only of degree.

Many famous scientists, while fully accepting evolution, have parted company with Darwin on this question. For example, Theodosius Dobzhansky: “Homo sapiens is not only the sole tool-making and the sole political animal, he is also the sole ethical animal.”4 Or George Gaylord Simpson: “[M]an is an entirely new kind of animal … [T]he essence of his unique nature lies precisely in those characteristics that are not shared with any other animal,”5 especially self-awareness, culture, speech, and morality. The difference between humans and non-human animals according to a number of contemporary philosophers6 goes like this:Precisely because they are incapable of conceptual thought, animals … are not only (1) incapable of sentence-making that includes statements about the past and future, (2) unable to fabricate tools for remote future use, (3) devoid of a cumulative cultural inheritance that constitutes a long historical tradition, but they are also (4) incapable of any behavior that is not rooted in the perceptually apprehended present situation.


Apart from quibbles about how long is long in (3), every one of these confident assertions now appears false, on the basis of the sort of evidence we have presented or are about to present in this book. Even if we ourselves are not personally scandalized by the notion of other animals as close relatives, even if our age has accommodated to the idea, the passionate resistance of so many of us, in so many epochs and cultures, and by so many distinguished scholars, must say something important about us. What can we learn about ourselves from an apparent error so widespread, propagated by so many leading philosophers and scientists, both ancient and modern, and with such assurance and self-satisfaction?

One of several possible answers: A sharp distinction between humans and “animals” is essential if we are to bend them to our will, make them work for us, wear them, eat them—without any disquieting tinges of guilt or regret. With untroubled consciences, we can render whole species extinct—for our perceived short-term benefit, or even through simple carelessness. Their loss is of little import: Those beings, we tell ourselves, are not like us. An unbridgeable gap has thus a practical role to play beyond the mere stroking of human egos.7 Darwin’s formulation of this answer was: “Animals whom we have made our slaves, we do not like to consider our equals.”8

——


We now proceed, in Darwin’s footsteps,9 to examine some of the multitude of proffered definitions of ourselves, explanations of who we are. We will try to see whether they make sense, especially in the light of what we know about the other beings that share the Earth with us.

One of the earliest attempts at an unambiguous characterization of humanity was Plato’s: Man is a featherless biped. When news of this advance in the art of definition reached the philosopher Diogenes, so the story goes, he introduced a plucked chicken into the weighty deliberations of Plato’s celebrated Academy, asking the assembled scholars to salute “Plato’s man.” This is of course unfair, because chickens are ordinarily born with feathers, just as they are ordinarily born with two feet. How we mutilate them afterwards does not change their fundamental nature. But the academicians took Diogenes’ challenge seriously and added another qualification: Humans were redefined as featherless bipeds with broad flat nails.

Surely this does not get us very far to the essence of human nature. The Platonic definition might suggest, though, a necessary if not a sufficient condition, because standing on two legs is essential for freeing the hands, hands are the key to technology, and many people think our technology defines us. Still, raccoons and prairie dogs have hands and no technology, and bonobos walk upright a good part of their lives. We will address chimpanzee technology shortly.

——


In his classic justification of free enterprise capitalism, Adam Smith asserts that “the propensity to truck, barter, and exchange one thing for another … is common to all men, and to be found in no other race of animals.”10 Is this true? Private property was proposed as the central difference between humans and the other animals by Martin Luther in the sixteenth century, and by Pope Leo XIII in the nineteenth.11 Is this true?

Chimpanzees are fond of trade, and understand the idea very well: food for sex, a back rub for sex, betrayal of the leader for sex, spare my baby’s life for sex, virtually anything for sex. Bonobos take these exchanges to a new level. But their interest in barter is by no means restricted to sex:[Chimpanzees] are famous for their tradesmanship. Experimental studies indicate that the ability comes without any specific training. Every zookeeper who happens to leave his broom in the baboon cage knows there is no way he can get it back without entering the cage. With chimpanzees it is simpler. Show them an apple, point or nod at the broom, and they understand the deal, handing the object back through the bars.12


With regard to females at least, chimp males have a well-developed sense of private property (raised to institutional status among the hamadryas baboons), and a rudimentary sense of private property attaches to food and to some tools.

The Wealth of Nations was published in 1776, well before any serious study had been made of the lives of the apes, even in captivity. However, Smith’s argument about the uniqueness of trade among humans is embedded in a deeper misreading of the animal world:In almost every other race of animals, each individual, when it is grown up to maturity, is entirely independent and in its natural state has occasion for the assistance of no other living creature. But man has almost constant occasion for the help of his brethren, and it is in vain for him to expect it from their benevolence only. He will be more likely to prevail if he can interest their self-love in his favor and show them that it is for their own advantage to do for him what he requires of them.13


But the gregariousness of the primates is one of their hallmarks. Mutual aid in working both sides of the predator/prey relationship and in conflict with other groups of the same species is widespread, not just among the primates, but among most mammals and birds.

While selfishness, exploitation, and trade are commonplace in chimpanzee society, we cannot use this fact along with our kinship with chimps to justify laissez faire economics. Nor can we use it to discredit free market societies on the grounds of their being ape-like.* Cooperation, friendship, and altruism are also chimp traits, but this is not an argument for some competing socialist economic doctrine. Recall the macaques who would rather go hungry than administer an electric shock to other, not closely related macaques—going so far as to reject even substantial material incentives. Is this a rebuke to advocates of capitalism? At least as far back as Aesop, animal behavior has been used to buttress this or that economic theory. Even in our ideological debates, we make the other animals work for us.

——


“Man is a social animal,” wrote Aristotle, or, as it is sometimes translated, “Man is a political animal.” This was meant to be characteristic of humans, but not defining; again, a necessary but not a sufficient condition. The subtle and volatile factionalism of chimp and bonobo societies shows how far off the mark this is as a distinction of humanity. The social insects—ants, bees, termites—have much better organized and much more stable social structures than humans. Particular aspects of human social behavior fare no better, although a great many such definitions have been proposed: For example, humans tenderly cherish their young, but so do most other mammals and birds.

“Courage is the peculiar excellence of man,” Tacitus recorded the Roman aristocrat Claudius Civilis as saying.14 Even if the heroic exploits of mother birds shamming a broken wing, or of elephants and chimps saving their young from predators or rushing water, or of the beta hind staring the wolf in the eye so her companions can escape—even if such examples were unknown in the time of this Claudius, didn’t he know about dogs? He was put in chains and brought before Nero. History does not record how much of the “peculiar excellence” was available to him in his hour of need.

Another ancient definition of humans, tracing back to Aristotle, is a “rational animal.”15 This is the distinction pointed to by many of the key figures in Western philosophy. But the categorizing chimps, reasoning by analogy and transitive inference, the conversing bonobos, and the culturally innovative macaques remind us that other animals reason also; not as well as the great Western philosophers, to be sure—but the philosophers believed not in a difference of degree, but in a radical difference in kind.

“[M]an differs from irrational creatures in this, that he is master of his actions,” was a tenet of St. Thomas Aquinas in his Summa Theologica. But are we “masters” of our actions always and in all circumstances? Do other animals never exhibit “mastery”? In giving, as was his practice, selected pros and cons for the propositions discussed, Aquinas—debating “whether choice is to be found in irrational animals?”—mentions a case where a stag at a crossroads seemed to choose one path by excluding the alternatives. This is rejected as evidence of choice because “choice properly belongs to the will, and not to the sensitive appetite which is all that irrational animals have. Therefore, irrational animals are not able to choose.” He also held that “irrational animals” could not command, “since they are devoid of reason.” All this may have satisfied generations of philosophers, and established a tradition that influenced Descartes, but is it not clear that Aquinas—consider his starting point of “irrational animals”—was begging the question, assuming what he was trying to prove?16

“Actions directed towards a goal do not occur in any other animals at all,” in a like vein wrote Jakob von Uexküll, a once influential expert on animal behavior.17 But we need only think of the chimp holding a club behind his back and searching for his rival, or collecting stones to throw at an enemy, or the female prying his fingers open and removing the stones, to realize how much in error such statements are.

For the philosopher John Dewey, what distinguishes us is memory:With the animals, an experience perishes as it happens, and each new doing or suffering stands alone. But man lives in a world where each occurrence is charged with echoes and reminiscences of what has gone before, where each event is a reminder of other things.18


This claim is manifestly untrue for many animals, and chimps above all live in a world “charged with echoes and reminiscences.” The cat experiencing a hot stove avoids the stove thereafter; elephants and deer soon grow wary of hunters; dogs who have been beaten cower when the rolled-up newspaper is raised; even worms, even one-celled protozoa can be taught to run a simple maze. The dominance hierarchy is a frozen memory of past coercion. How oblivious of the real life of nonhuman animals is Dewey’s attempt to define us!

Many human sexual practices have been thought to be defining. Maybe it’s kissing: “Only mankind kisses. Only mankind has the reason, the logic, the happy faculty of being able to appreciate the charm, the beauty, the extreme pleasure, the joy, the passionate fulfilment of the kiss!” rhapsodizes a small book on the subject.19 But chimps routinely and exuberantly kiss.

Maybe what’s special about us is our reproductive posture: “It seems plausible to consider that face-to-face copulation is basic to our species.”20 But face-to-face copulation is common among the bonobos.

Concealed ovulation and female orgasm21 have been thought unique to humans, but bonobos do not garishly advertise their ovulations, and female chimps, bonobos, stumptail monkeys and, probably, many other primate females have orgasms—as determined in part by equipping them with physiological sensors before they mate, in the style of an experiment by Masters and Johnson.

Maybe it’s our mode of sexual coercion: “That rape … is an exclusively human character seems to be beyond serious doubt,” opined a scientist writing on primates in 1928.22 But rape is known among orangutans and stumptails, violent sexual coercion is a commonplace among baboons and chimps, and the doubt is serious indeed.

Maybe it’s the elaboration and duration of our sexual foreplay; in this at least some humans may lead the other primates.23 But this is learned behavior, as the prevalence of premature ejaculation, especially among adolescent boys, and the self-taught ability of many men to postpone ejaculation make clear. In the integration of sexual acts into everyday social life humans are probably down toward the bottom of the primate list. Most human cultures demand that even socially condoned sexual behavior be carried on in private;24 we can see something of the sort in chimp consortship, and in clandestine encounters out of sight of the dominant males.

Maybe our distinction is the traditional and striking gender-specific division of labor: The men hunt and fight; the women gather and nurture.25 But this cannot be a defining characteristic, because chimps have a similar division of labor: Patrols, group defense, and throwing missiles are all mainly male responsibilities; caring for the young and using tools to crack open nuts are mainly female responsibilities. Also, women’s and men’s jobs are in our time becoming increasingly indistinguishable.

Our long childhood, the years between birth and puberty, is essential for our education, but it is not as long as an elephant’s; and the progressively earlier arrival of sexual maturity in the human life cycle over the last few centuries is whittling down our childhood so that it is now only a little longer than the chimpanzees’ (who sexually mature around age ten). Play is so central to our growing up that it was once suggested26 to call our species Homo ludens (“the man who plays”). But play can be seen throughout the mammalian class, especially when maturity is long delayed.

The Roman philosopher Epictetus, a former slave, held the distinguishing characteristic of humans to be personal hygiene.27 He must have known about birds, cats, and wolves but argued that “when … we see any other animal cleaning itself, we are accustomed to speak of the act with surprise, and to add that the animal is acting like a man.” But he then complains that many men are “dirty,” “stinking,” and “foul” and do not share this “distinguishing” characteristic. Such a man is advised to “go into a desert … and smell yourself.”

Humans have been called the only animal that laughs. However, chimps smile and laugh a lot.28 The Athenian Stranger, in Plato’s Laws,29 says humans are “afflicted with the inclination to weep more than any other animal.” But this inclination varies widely from culture to culture, and whimpering and crying is a fact of daily life among the chimps, children and adults alike.30

Humans—who enslave, castrate, experiment on, and fillet other animals—have had an understandable penchant for pretending that animals do not feel pain. On whether we should grant some modicum of rights to other animals, the philosopher Jeremy Bentham stressed that the question was not how smart they are, but how much torment they can feel. Darwin was haunted by this issue:In the agony of death a dog has been known to caress his master, and every one has heard of the dog suffering under vivisection, who licked the hand of the operator; this man, unless the operation was fully justified by an increase of our knowledge, or unless he had a heart of stone, must have felt remorse to the last hour of his life.31


From all criteria available to us—the recognizable agony in the cries of wounded animals, for example, including those who usually utter hardly a sound*—this question seems moot. The limbic system in the human brain, known to be responsible for much of the richness of our emotional life, is prominent throughout the mammals. The same drugs that alleviate suffering in humans mitigate the cries and other signs of pain in many other animals. It is unseemly of us, who often behave so unfeelingly toward other animals, to contend that only humans can suffer.

Murder, cannibalism, infanticide, territoriality, and guerilla warfare are not unique to humans, as described in preceding chapters. Ants have slaves and domesticated animals and main force warfare.

“The use of punishment in the attempt to train their young in anything other than avoidance,” writes Toshisada Nishida, “seems exclusively limited to humans … No nonprimate mammals are known to teach by discouragement.”33 But his exception of the nonhuman primates says much. Also, many animals coerce and punish the young as part of the educational process, aiding smooth entrance into the dominance hierarchy. It’s a little like hazing and initiation rites in our species.

Humans have institutionalized marriage and advocated monogamy, at least as an ideal; but gibbons, wolves, and many species of birds practice monogamy and mate for life. The courtship dances of animals are surely a kind of marriage ceremony. The following characteristics are described as typical of human marriage:There is some degree of mutual obligation between wife and husband. There is a right of sexual access (often but not invariably exclusive). There is an expectation that the relationship will persist through pregnancy, lactation, and childrearing. And there is some sort of legitimization of the status of the couple’s children.34


But all of this is known in other animals, for example among the gibbons, plus primogeniture.

The nineteenth-century philosopher and theologian Ludwig Feuerbach—known for his influence on Karl Marx—proposed that the distinction of humans is recognition of ourselves as a species.35 But many animals readily distinguish members of their own species from members of all others—for example, through olfactory cues. And humans are notable for demonizing members of their own species, declaring them less than human, to disinhibit sanctions on murder—especially during wartime.

Humans are sometimes said to be better at making class distinctions than other primates are,36 but primate dominance hierarchies, some of them hereditary, seem to embrace a fineness of social discrimination that in some respects exceeds even our own.

We conclude that none of these sexual and social traits seem to work as defining characteristics of the human species. The behavior of other animals, especially the chimps and bonobos, renders such pretensions specious. They are just too much like us.

——


Knowledge and behavior patterns that are not hardwired into our genetic material, but rather are learned and passed on within a given group from generation to generation, are called culture. Could culture be the defining mark of humanity?

“Culture,” says a major article in The Encyclopaedia Britannica,is due to an ability possessed by man alone The question of whether the difference between the mind of man and that of the lower animals is one of kind or of degree has been debated for many years, and even today [1978] reputable scientists can be found on both sides of this issue. But no one who holds the view that the difference is one of degree has adduced any evidence to show that non-human animals are capable, to any degree whatever, of a kind of behaviour that all human beings exhibit.


The author then gives three examples of behavior that he thinks characterizes humans, and concludes, “There is no reason or evidence that will lead one to believe that any animal other than man can have or be brought to any appreciation or comprehension whatever of such meanings and acts.”37

And what are these three examples? One is “defining and prohibiting incest.” But this prohibition, at least for the father-daughter and mother-son varieties, is, as we’ve described, prevalent, indeed nearly invariable, among the primates—who have elaborate conventions to guarantee high levels of outbreeding. The taboo applies to many other animals as well. In studying Kenyan birds known as bee-eaters, the biologist Stephen Emlen carefully noted the identity and behavior of each bird; in eleven years of work he was unable to find even a single case of incest, either between siblings or between parent and offspring. (The other two examples given in the Britannica article are “classifying one’s relatives and distinguishing one class from another,” which chimps do well enough—at least for mother-child and sibling kinship—and “remembering the sabbath to keep it holy,” which is an institution unknown in many human cultures.)

Despite the common description of the incest prohibition as a taboo—that is, learned—it seems to be, to a considerable degree, innate. It serves as a hereditary ethical proscription, evolved for good genetic reasons, and reinforced by the conventions and rules of society (although, for all that, functioning imperfectly—very imperfectly in civilized society).

Clearly chimps have at least the rudiments of culture. In different forests, they must deal with different local geographies and ecologies. They remember over weeks—maybe over years—termite mounds, drumming trees, or, in one account, the site of a noteworthy combat. Such matters are common knowledge. Each group, with its own terrain and its own sequence of historical events, has its own miniature culture. Mutually isolated groups of chimps have different conventions in fishing for termites or driver ants, in using leaves as sponges for soaking up drinking water, in how they hold on to each other during grooming, in some aspects of the gestural language of courtship, and in hunting protocols.38 And thanks to Imo, the macaque genius who figured out how to separate the wheat from the sand, we even have some insight into the emergence and spread of new discoveries and new cultural institutions among the primates.

The celebrated philosopher Henri Bergson—an exponent of the “revolt against reason” and best known for the idea that some immaterial “vital impulse” permeates life and makes evolution go—wrote that “man … is alone in realizing that he is subject to illness.”39 But chimps have a vast pharmacopoeia all around them, and a kind of folk or herbal medicine. For example, for chimps both at Gombe and at Mahale, leaves of a plant called Aspilia are a kind of dietary staple, preferentially eaten in the early morning. Despite the wrinkled noses of those partaking (the taste is bitter), it’s consumed by both sexes, all ages, the healthy as well as the sick. But there’s something odd about it: The chimps eat these leaves regularly, but consume very few of them at any one time—so their nutritional value is in doubt. In the rainy season, though, when apes are plagued by intestinal worms and other illnesses, ingestion increases dramatically. Analysis of Aspilia leaves reveals the presence of a powerful antibiotic and an agent that kills nematodes. It’s a good guess they’re treating themselves. Among other examples, a chimp sick with an intestinal disorder ingested large amounts of the shoots of a plant, different from Aspilia and not ordinarily a part of its diet, which also proved to be rich in natural antibiotics.40

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