Behavioral Ecology and Sociobiology

, Volume 57, Issue 1, pp 50–61 | Cite as

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (Lemur catta)

  • Charles L. NunnEmail author
  • Robert O. Deaner
Original Article


Cooperation in animal social groups may be limited by the threat of “free riding,” the potential for individuals to reap the benefits of other individuals’ actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other group’s territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.


Lemur catta Territoriality Individual participation Cooperation 



We thank the DUPC staff and our summer research assistants for their support. We also acknowledge helpful discussions or comments from R. Barton, T. Clutton-Brock, C. Gilbert, P. Kappeler, R. Lewis, S. Patek, M. Pereira, C. van Schaik and anonymous reviewers. Support was provided by an NSF graduate student fellowship, Conservation International Primate Action Fund, Duke Arts and Sciences Research Council, and Sigma Xi. This paper is Duke Primate Center publication no. 752. The research presented here was described in Animal Research Protocol no. A124-98-3 and A144-99-3R1, approved on 26 March 1998 and 26 April 1999 by the Institutional Animal Care and Use Committee of Duke University.


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Copyright information

© Springer-Verlag 2004

Authors and Affiliations

  1. 1.Section of Evolution and EcologyUniversity of CaliforniaDavisUSA
  2. 2.Department of NeurobiologyDuke University Medical CenterDurhamUSA
  3. 3.Department of Integrative BiologyUniversity of CaliforniaBerkeleyUSA

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