1 Department of Anthropology, University of Texas at San Antonio, Texas, USA.
2 Department of Anthropology and Archaeology, University of Calgary, Calgary, Alberta, Canada.
3 Department of Biology, Concordia University, Montréal, Quebec, Canada.

Cooperative home range defense is common in primates, despite a collective action problem that arises when group members benefit from winning the intergroup encounter regardless of whether they participate. The costs associated with this collective action problem may be mitigated by residing in small groups, residing with kin, or by forming strong bonds with group members. The potential to decouple the effects of these variables provided an opportunity to investigate which of these three variables best explains coparticipation in intergroup encounters among adult and subadult female colobus at Boabeng-Fiema, Ghana. Because males are often the main participants, we also investigated the relationship between female-female coparticipation and adult and subadult male participation. We collected intergroup behaviors from 94 adult and subadult individuals in eight groups during 1 year. We quantified female grooming bond strength and approach rates using focal samples. We classified female dyads as close kin (i.e., halfsiblings or more closely related) or nonkin based on partial pedigrees and genotypes generated from 17 STR loci. Female-female coparticipation was higher in dyads with stronger grooming bonds but was not associated with dyadic kinship, approach rate, or age class. Female coparticipation decreased with increasing female group size as expected if there is a collective action problem. Females coparticipated less in groups with more males and male intergroup aggression, possibly because there is less need for female-female cooperation if males are participating in the intergroup encounter. Females in smaller groups may not only benefit from increased female-female cooperation during intergroup encounters, they are also likely to reside with a higher-quality alpha male, both of which may increase the likelihood of winning intergroup encounters. There may be strong selection for facultative female dispersal in populations like the Boabeng-Fiema colobus in which small groups are associated with multiple benefits and cooperation is not affected by kinship.