Female participation in collective group defense in black howler monkeys (Alouatta pigra).

Many group-living animals actively defend a home range against neighboring groups. In many of these societies, males are the primary participants during group defense, while female participation ranges from seldom to frequent. Among howler monkeys (Alouatta spp.), loud calls (i.e., howling) are often used in the context of intergroup spacing as a form of cooperative group defense. Males initiate and lead these howling bouts, but females occasionally participate as well. During a 28-month study, I examined social and ecological factors influencing the participation of adult females in naturally occurring howling bouts of five multimale-multifemale groups of black howler monkeys (A. pigra) at Palenque National Park, Mexico. I calculated the percentage of time each female participated during howling bouts for which the participation of all resident females could be recorded ≥80% of the time (N = 287). At least one female was observed to participate in 53% of the vocal displays. Female participation was significantly greater during howling bouts that were part of visual intergroup encounters compared to spontaneous calls or calls in response to nearby calls when there was no visual contact with rival groups. Female calling behavior was not influenced by the presence of infants vulnerable to infanticide or by the proximity to food resources. Nonetheless, in four of the five study groups, one female called significantly more than the other resident female(s), suggesting that these females played a special role within the group's social dynamics, not previously recognized for this species.