In most primate species, the distribution of food in the environment determines the distribution of females, and the distribution of females determines the distribution of males. When food is so dispersed that females do best by foraging on their own, males disperse to pair up with the lone females. This gives rise to monogamous couples. It is a fairly rare pattern among primates, limited to gibbons, some lemurs, and some African and South American monkeys.
When food comes in patches large enough for several females to share, they tend to band together in small groups to find the food, and to protect each other against predators, unwanted males, and competing female groups. As long as the female band is not too large, a single male can exclude other males from sexual access to the band, which thus becomes "his." This "harem system" of single-male polygyny is fairly common in primates, being found in hamadryas baboons, colobus monkeys, some langurs, and gorillas. The competition between males to guard the female groups creates very strong sexual selection pressures for male size, strength, aggressiveness, and large canine teeth.
When food comes in still larger patches, female groups can grow too large for any single male to defend them. The males must then form coalitions, resulting in a complex multi-male, multi-female group, as in some baboons, macaques, ring-tailed lemurs, howler monkeys, and chimpanzees. Our hominid ancestors probably lived in such groups, in which sexual selection gets more complicated. Sometimes, females in multi-male groups appear to use sperm-production ability as the main fitness indicator. A chimpanzee female might mate with every male in the group every time she becomes fertile. She lets their sperm fight it out in her reproductive tract, and the strongest swimmers with the best endurance will probably fertilize her egg.
In response to this sexual selection for good sperm, male chimpanzees have evolved large testicles, copious ejaculates, and high sperm counts. Female primates face a trade-off. They can select for the best-swimming sperm by mating very promiscuously, or they can select for the best courtship behavior by mating very selectively. Or they can do a little of both, selecting a small group of male lovers for their charm and then letting their sperm fight it out.
In species that do not get completely caught up in runaway sperm competition, females can favor various male behavioral traits. Multi-male groups obviously allow greater scope for females to choose between males. If they favor dominant males, males evolve through sexual selection to compete intensely for social status by individual force or by forming coalitions. If females favor kind males, males evolve through sexual selection to groom females, protect their offspring, and guard them from other males.
Given multi-male, multi-female primate groups, how does mate choice work? Female primates can exercise choice by joining groups that contain favored males, initiating sex with them during estrus, supporting them during conflicts, and developing long-term social relationships with them. Females can reject unfavored males by refusing to cooperate during copulation attempts, driving males away from the group, or leaving the group. But female mate choice criteria remain obscure for most primate species. In contrast to modern humans, female primates rarely favor males who can provide resources or paternal care of offspring. The sporadic male care that is observed, such as watching, carrying, and protecting infants, is better described as courtship effort than as paternal care. The male is unlikely to be the infant's father, but is simply trying to mate with the infant's mother by doing her a favor.
Primate researchers still know little about what traits are preferred by male and female primates. For example, we know less about female choice in other apes than we do about female choice in the Tungara frog, the guppy fish, or the African long-tailed widowbird. Nevertheless, three kinds of female preference have been reported in primates: preferences for high-ranking males capable of protecting females and offspring from other males; preferences for male "friends" that have groomed the female a lot and have been kind to her offspring; and preferences for new males from outside the group, perhaps to avoid genetic inbreeding. Each sort of preference could be explained in terms of female choice for good genes, or female choice for material and social benefits. Although male primates have evolved an astounding diversity of beards, tufts, and colorful hair styles, there has been very little research on female choice for male appearance.
Also, there has been virtually no research on primate sexual choice for personality or intelligence. Female primates are sometimes reported to show "irrational" or "capricious" preferences that cannot be explained on the basis of male dominance, age, or group membership. Sometimes two primates just seem to like each other based on unknown features of appearance, behavior, or personality. Female primates might well be choosing males for their personalities and notjust their status, but we do not know.
Most primates follow the general animal pattern of male sexual competition and female choosiness. But when the costs of male sexual competition and courtship are high, males also have incentives to be choosy When male mate choice becomes important, sexual selection affects females as well as males. In monogamous marmosets and tamarins, females compete to form pairs with quality males and drive off competing females. In single-male harem systems, the dominant male's sperm can become a limiting resource for female reproduction, and high-ranking females prevent low-ranking females from mating through aggression and harassment. In multi-male groups, females sometimes compete to form consortships and friendships with favored males. Such patterns of female competition suggest some degree of male mate choice. When the costs of sexual competition and courtship are high, males have an incentive to be choosy about how they spread their sexual effort among the available females. Males compete much more intensely for females who show signs of fertility such as sexual maturity, estrus swellings, and presence of offspring Like females, some male primates also develop special friendships with particular sexual partners. It may not be romantic love, but, at least among some baboon pairs, it looks pretty similar.
Our closest ape relatives, the chimpanzees and the bonobos, live in multi-male, multi-female groups in which sexual choice is dynamic, intense, and complicated. Under these relentlessly social conditions, reproductive success came to depend on social intelligence rather than brute strength. Both sexes compete, both sexes have dominance hierarchies, and both sexes form alliances. Sexual relationships develop over weeks and years rather than minutes. Many primatologists and anthropologists believe that our earliest hominid ancestors probably lived under similar social and sexual conditions. Constant sociosexual strategizing in mixed-sex groups was the legacy of our ape-like ancestors. It was the starting point, not the outcome, of sexual choice in human evolution.
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