1 Department of Anthropology, University of Toronto Scarborough, Toronto, Canada. arseneau.jean.m@gmail.com.
2 Department of Anthropology, University of Toronto Scarborough, Toronto, Canada.
3 Wildlife Research Center, Kyoto University, Kyoto, Japan.
4 Department of Anthropology, University of Calgary, Calgary, Canada.
5 Department of Anthropology, University of Texas San Antonio, San Antonio, USA.
6 Department of Biology, Concordia University, Montreal, QC, Canada.
7 Department of Anthropology, University of Texas San Antonio, San Antonio, USA. eva.wikberg@utsa.edu.

Intergroup aggression often results in the production of public goods, such as a safe and stable social environment and a home range containing the resources required to survive and reproduce. We investigate temporal variation in intergroup aggression in a growing population of colobus monkeys (Colobus vellerosus) to ask a novel question: "Who stepped-up to produce these public goods when doing so became more difficult?". Both whole-group encounters and male incursions occurred more frequently as the population grew. Males and females were both more likely to participate in whole-group encounters when monopolizable food resources were available, indicating both sexes engaged in food defence. However, only females increasingly did so as the population grew, suggesting that it was females who increasingly produced the public good of home range defence as intergroup competition intensified. Females were also more active in male incursions at high population densities, suggesting they increasingly produced the public good of a safe and stable social environment. This is not to say that males were chronic free-riders when it came to maintaining public goods. Males consistently participated in the majority of intergroup interactions throughout the study period, indicating they may have lacked the capacity to invest more time and effort.