2002 chimpanzees: Gilby and Wrangham 2008 Nishida 1979 Watts 2000a, b cf. 2009), muriquis, Brachyteles arachnoids: Strier et al. In male philopatric species, in contrast, adult males form the strongest social bonds, e.g., red colobus ( Colobus badius: Struhsaker and Leland 1976), spider monkeys ( Ateles geoffroyi: Slater et al. In female philopatric species adult females have very strong social bonds, e.g., rhesus macaques ( Kapsalis and Berman 1996), capuchins ( Cebus capucinus: Perry 1996), vervets ( Cercopithecus aethiops: Seyfarth 1980), savannah baboons ( Papio cynocephalus: Seyfarth 1976 Silk et al. There is also evidence that patterns of social relationships differ between adult males and females. 2014), whereas in patas monkeys ( Erythrocebus patas) the time spent playing decreased with age in both sexes ( Rowell and Chism 1986). Similarly, in chimpanzees ( Pan troglodytes) the percentage of time dedicated to social grooming of other group members increased with age for both male and female juveniles ( Lonsdorf et al. For example, both male and female Japanese macaques ( Macaca fuscata) showed a decrease in the variety of social behaviors exchanged with their mothers after the first year of life and mostly groomed each other as they grew older ( Nakamichi 1989). There is evidence that some aspects of sociality change through development in a similar way for both sexes. However, few studies so far have explored how social relationships develop through ontogeny, and especially whether sex differences exist in the development of these relationships. Given these premises, it is not surprising that group-living primates are characterized by a complex network of social relationships. Similarly, enduring social relationships enhanced reproductive success in male macaques ( Schülke et al. 2008) and increased offspring survival ( Silk et al. The quality of female baboons’ social relationships, for example, positively affected their ability to cope with stressful events ( Crockford et al. However, recent evidence has shown that the quality of social relationships plays a crucial role in enhancing primates’ fitness even when social relationships involve nonkin. The existence of strong social relationships among nonkin has long been considered an evolutionary puzzle ( Cheney 2011 Clutton-Brock 2009 Seyfarth and Cheney 2012). Although strong social relationships are typically observed between genetically related individuals ( Chapais 2001 Silk 2002a), long-term cooperative relationships are also common between unrelated individuals ( Seyfarth and Cheney 2012 Silk 2002b). Several studies have shown that primates form strong and enduring social bonds with their conspecifics ( Lehmann and Boesch 2009 Silk et al. We propose that play might serve as a trigger of sex differences in social behavior, with sex differences emerging early in development and increasing through time as males and females gradually grow into their adult social roles. The only notable exception to this pattern was play, which was more pronounced in males from the beginning of their lives. 2 yr old, suggesting that this might be a milestone in the development of sociality in rhesus macaques. Importantly, most developmental changes in sociality happened when individuals were ca. Moreover, both male and female individuals interacted mostly with maternal kin, although males also preferred paternal kin over nonkin. High-ranking individuals, especially older females, were generally preferred as social partners. Sex differences in social behavior varied through development, but also depended on rank, partner’s rank, and kin line, although not consistently. In contrast, males interacted more with males and age peers, especially around maturation. Females engaged in more social interactions than males, especially with other females, and were more involved in grooming around the time of maturation. In particular, sex differences in social behavior varied through development depending on the partner’s sex and age. Generalized linear mixed models revealed that social behaviors mostly followed different developmental trajectories in males and females and were highly dependent on the social context. Here we analyzed affiliative interactions (proximity, social grooming, play) combined with demographic and genetic data in semi-free-ranging rhesus macaques ( Macaca mulatta) on Cayo Santiago over their first 4 yr of life (from birth to sexual maturation) to understand how these interactions change through development in both sexes. However, little is known about how social relationships develop through ontogeny, and whether their development follows the same trajectory in males and females. Several studies have documented the importance of social bonding for the enhancement of individual fitness.
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