Males collectively defend their one-male units against bachelor males in a multi-level primate society.

Group-level male-male co-operation, which has been documented in several primate and non-primate societies, may be mutualistically advantageous to the participants when confronted with threats such as takeovers and cuckoldry by external males. Co-operation among members of distinct social units-while universal among humans-is extremely rare in non-human primates. We present the first observations of collective action or co-operation among males of different one-male units (OMU) in a multi-level society of Rhinopithecus roxellana. A total of 59 instances of male co-operation were recorded. Male co-operation included coordinated chasing, joint vigilance, and patrolling behavior directed at lone adult males trying to enter an OMU. Male co-operation was significantly more frequent during the mating season when the risk of incursions and extra-group paternity was higher. Paternity of infants born in the subsequent birth season and kin relationships among resident males were identified using microsatellite genotype. All infants were sired by OMU males, which we interpret as possible evidence for their success at thwarting mating attempts by satellite males. OMU males were principally unrelated suggesting that male co-operation is best understood in terms of the mutual direct benefits individuals obtain through collective action. Our findings lend support to the bachelor threat hypothesis in which the cooperative behavior of several individuals is more effective than the lone action of a single individual in providing mate defense. Our research has implications for understanding male bonding, higher-level collective action, and the evolution of social co-operation in human societies.