Bartlett TQ

Borgatti SP

Clutton-Brock TH

Grueter CC

Kawai M

Kirkpatrick RC

Lynch M

Mantel N

Pusey AE

Wang HP

Wang XW

A small number of primate species including snub-nosed monkeys (colobines), geladas (papionins) and humans live in multilevel societies (MLSs), in which multiple one-male polygamous units (OMUs) coexist to form a band, and non-breeding males associate in bachelor groups. Phylogenetic reconstructions indicate that the papionin MLS appears to have evolved through internal fissioning of large mixed-sex groups, whereas the colobine MLS evolved through the aggregation of small, isolated OMUs. However, how agonistic males maintain tolerance under intensive competition over limited breeding opportunities remains unclear. Using a combination of behavioural analysis, satellite telemetry and genetic data, we quantified the social network of males in a bachelor group of golden snub-nosed monkeys. The results show a strong effect of kinship on social bonds among bachelors. Their interactions ranged from cooperation to agonism, and were regulated by access to mating partners. We suggest that an ‘arms race’ between breeding males' collective defence against usurpation attempts by bachelor males and bachelor males' aggregative offence to obtain reproductive opportunities has selected for larger group size on both sides. The results provide insight into the role that kin selection plays in shaping inter-male cohesion which facilities the evolution of multilevel societies. These findings have implications for understanding human social evolution, as male–male bonds are a hallmark of small- and large-scale human societies.</jats:p

Xiao-Guang Qi

Kang Huang

Gu Fang

Cyril C. Grueter

Derek W. Dunn

Yu-Li Li

Weihong Ji

Xiao-Yan Wang

Rong-Tao Wang

Paul A. Garber

Bao-Guo Li

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Proceedings of the Royal Society B Biological Sciences

Male cooperation for breeding opportunities contributes to the evolution of multilevel societies

Male cooperation for breeding opportunities contributes to the evolution of multilevel societies

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