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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.