Multilevel societies (MLS), in which polygynous reproductive units are nested in a larger social matrix, represent a highly complex social system documented only in a small number of mammalian species. Using long-term behavioural data, satellite telemetry and social network analysis, we present a new framework for understanding the function and social dynamics of the golden snub-nosed monkey MLS. Here we show that several one-male units form a cohesive breeding band that associates with one or more all-male units to form a herd. Herds seasonally aggregate and exchange members, thus facilitating gene flow and inbreeding avoidance. This MLS evolved from the aggregation of independent one-male, multifemale units that characterize ancestral Asian colobines; the evolutionary pathway leading to this MLS contrasts with that proposed for African papionins, which appear to have undergone internal fissioning of multimale–multifemale groups. The results suggest that both environmental and phylogenetic factors are important in the evolution of a primate MLS.
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