Male chimpanzees, Pan troglodytes, engage in cooperative territorial defence and sometimes kill members of neighbouring communities. Observations of intergroup interactions suggest that escalation of aggression depends on numerical assessment, with lethal attacks occurring when numerical advantage reduces the costs of attacking. To gain a better understanding of the factors guiding participation in intergroup conflict, we conducted a series of playback experiments with the Kanyawara chimpanzee community of the Kibale National Park, Uganda. We tested whether the response to the playback of the 'pant-hoot' call of a single extragroup male depended on the number of adult males in the listening party, the location of the speaker relative to the territory edge, and each male's agonistic rank. These playbacks elicited cooperative responses, with the nature of the response depending on the number of adult males in the party. Parties with three or more males consistently joined in a chorus of loud vocalizations and approached the speaker together. Parties with fewer adult males usually stayed silent, approached the speaker less often, and travelled more slowly if they did approach. In contrast to many territorial species, the location of the simulated intruder did not affect the response. Although high-ranking males might be expected to benefit more from repelling outside males, both high- and low-ranking males showed a similar pattern of response. Each male responded as if he benefited from repelling intruders, but only if he had strength in numbers. This pattern of response is consistent with cooperation based on mutualism.
Bibliographical noteFunding Information:
Field research in Uganda was sponsored by Makerere University and the Uganda Wildlife Authority. We thank G. Isabirye-Basuta and J. Kasenene for logistic support and J. Mitani for providing call stimuli. We also thank M. Muller for providing male dominance rank data for 1998. J. Barwogeza, C. Katangole, D. Kateeba, F. Mugurusi, D. Muhangyi, C. Muruuli, P. Tuhairwe and the late J. Basigara provided invaluable assistance in the field. We thank S. C. Alberts, M. Gerald, R. Heinsohn, C. Hooven, M. Leighton, A. Marshall, M. Muller, C. Nunn, R. Sun and anonymous referees for discussion and comments on the manuscript. This work was supported by a Howard Hughes Medical Institute Predoctoral Fellowship (M.L.W.) and grants from the Leakey Foundation (M.D.H., R.W.W.), the Mellon Foundation (M.L.W.) and NSF awards BCS-9812781 (M.D.H., M.L.W.) and BCS-9807448 (R.W.W., M. Muller and P. Ellison). The research presented here was described in Animal Research Protocol No. 96-03, approved on 9 March 1999 by the Institutional Animal Care and Use Committee of Harvard University.