The demographic and reproductive context of male replacements in Cebus capucinus

Male primates may immigrate into groups by peacefully joining the residents and taking up low-ranking positions in the hierarchy, or they may enter by force, challenging the resident males and attempting to drive them from high rank or from the group. Here we address the questions of how, when, and why immigrating male white-faced capuchins (C. capucinus) at Santa Rosa replace the former resident males of our groups, rather than simply joining them. We present data on 15 male replacements in 6 study groups tracked from 1984 through March 2004. During 11 aggressive takeovers, resident males were nearly always outnumbered by coalitions of invading males; lone resident males were particularly vulnerable. Both residents and invaders were wounded and infants often perished during or soon after takeovers. Male replacements also occur when resident males abandon their groups and males from neighboring groups 'waltz in' to become resident. Three such 'waltz in' replacements occurred during the study period. If we combine takeovers with 'waltz in' cases, replacements occur about every 4 years in our study groups, almost invariably during the dry season months of January to April, about 3-6 months before the annual peak in conceptions. In the years that groups are subject to takeovers, group composition includes significantly lower proportions of adult males than in no-takeover years. We conclude that: (1) the mechanism of male replacement is usually aggressive takeover, but sometimes abandonment of the group by prior resident males occurs; and (2) aggressive takeovers are more likely to happen when the group is vulnerable because it has a lower proportion of adult males, particularly when all co-resident males have emigrated, leaving only the alpha male in residence. Our long-term study shows that adult males need coalition partners not only to gain entry to a group but also to maintain their membership within it.