Social instability raises the stakes during social grooming among wild male chimpanzees

Explaining cooperative behaviour is a fundamental issue for evolutionary biology. The challenge for any cooperative strategy is to minimize the risks of nonreciprocation (cheating) in interactions with im- mediate costs and delayed benefits. One of a variety of proposed strategies, the raise-the-stakes (RTS) strategy, posits that individuals establish cooperation by increasing investment across interactions from an initial interaction. This model has received little quantitative support, however, probably because individuals of many social species engage in repeated interactions from a young age. In some situations, however, such as following conflicts, after prolonged absences or during social instability, established relationships may become unreliable predictors of future behaviour, creating an environment for RTS. We investigated grooming interactions among wild male chimpanzees, Pan troglodytes, testing RTS in these specific contexts. We found evidence to support the view that male chimpanzees employed RTS during social instability, but not under the other conditions. However, we also found that the duration of episodes (discrete parcels) of grooming was negatively related to aggression risk and in consequence suggest that the patterning of grooming interactions indicative of RTS was less to do with preventing cheating, and more to do with avoiding the elevated risks of intramale aggression during the period of social instability. We interpret the apparent support for RTS in our data as a by-product of the way chimpanzees cope with fluctuating (here, elevated then diminishing) risks of aggression. We suggest that social instability raises the stakes for grooming by creating a more hazardous marketplace in which to trade.