Vervet monkeys give different alarm calls to different predators. Recordings of the alarms played back when predators were absent caused the monkeys to run into trees for leopard alarms, look up for eagle alarms, and look down for snake alarms. Adults call primarily to leopards, martial eagles, and pythons, but infants give leopard alarms to various mammals, eagle alarms to many birds, and snake alarms to various snakelike objects. Predator classification improves with age and experience.

Longevity is a major component of variation in fitness in long-lived iteroparous species [1-4]. Among female baboons, variation in breeding lifespan accounts for approximately 50% of the variation in lifetime fitness [5, 6]. However, we know little about the causes of variation in longevity in primates or other long-lived mammals. Savannah baboons form strong, equitable, and enduring relationships with specific female partners, particularly with close relatives and agemates [7-10]. The quality of females' social relationships influences their ability to cope with stressful events [11-13] and is associated with variation in female reproductive success [9, 14]. Here we show that dominance rank and the quality of close social bonds have independent effects on the longevity of female chacma baboons (Papio hamadryas ursinus). High-ranking females live longer than lower-ranking females. In addition, females who form stronger and more stable social bonds with other females live significantly longer than females who form weaker and less stable relationships. These data extend our understanding of the adaptive value of social bonds in baboons and complement a growing body of evidence that indicates that social bonds have adaptive value in a range of taxa, from mice to humans [9, 14-19].

Sociality has evolved in many animal taxa, but primates are unusual because they establish highly differentiated bonds with other group members. Such bonds are particularly pronounced among females in species like baboons, with female philopatry and male dispersal. These relationships seem to confer a number of short-term benefits on females, and sociality enhances infant survival in some populations. However, the long-term consequences of social bonds among adult females have not been well established. Here we provide the first direct evidence that social relationships among female baboons convey fitness benefits. In a group of free-ranging baboons, Papio cynocephalus ursinus, the offspring of females who formed strong social bonds with other females lived significantly longer than the offspring of females who formed weaker social bonds. These survival benefits were independent of maternal dominance rank and number of kin and extended into offspring adulthood. In particular, females who formed stronger bonds with their mothers and adult daughters experienced higher offspring survival rates than females who formed weaker bonds. For females lacking mothers or adult daughters, offspring survival was closely linked to bonds between maternal sisters. These results parallel those from human studies, which show that greater social integration is generally associated with reduced mortality and better physical and mental health, particularly for women.

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Convergent evidence from many species reveals the evolutionary origins of human friendship. In horses, elephants, hyenas, dolphins, monkeys, and chimpanzees, some individuals form friendships that last for years. Bonds occur among females, among males, or between males and females. Genetic relatedness affects friendships. In species where males disperse, friendships are more likely among females. If females disperse, friendships are more likely among males. Not all friendships, however, depend on kinship; many are formed between unrelated individuals. Friendships often involve cooperative interactions that are separated in time. They depend, at least in part, on the memory and emotions associated with past interactions. Applying the term "friendship" to animals is not anthropomorphic: Many studies have shown that the animals themselves recognize others' relationships. Friendships are adaptive. Male allies have superior competitive ability and improved reproductive success; females with the strongest, most enduring friendships experience less stress, higher infant survival, and live longer.

In animal communication natural selection favors callers who vocalize to affect the behavior of listeners and listeners who acquire information from vocalizations, using this information to represent their environment. The acquisition of information in the wild is similar to the learning that occurs in laboratory conditioning experiments. It also has some parallels with language. The dichotomous view that animal signals must be either referential or emotional is false, because they can easily be both: The mechanisms that cause a signaler to vocalize do not limit a listener's ability to extract information from the call. The inability of most animals to recognize the mental states of others distinguishes animal communication most clearly from human language. Whereas signalers may vocalize to change a listener's behavior, they do not call to inform others. Listeners acquire information from signalers who do not, in the human sense, intend to provide it.

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