Spatiotemporal environmental variation, risk aversion, and the evolution of cooperative breeding as a bet-hedging strategy.

In cooperatively breeding systems in which some individuals delay reproduction to help raise others' offspring, environmental variation in space and time influences individual reproductive strategies as well as interspecific patterns of sociality. Although most environmental explanations for cooperative breeding emphasize the mean fitness gains of living socially, the fittest individuals are not always those that produce on average the highest number of offspring. At times, variance in fecundity can influence fitness as much as mean fecundity, particularly in small populations like those of cooperative breeders. Cooperative breeding behavior could therefore be a risk-averse strategy to maximize fitness by reducing environmentally induced fecundity variance. Such a within-generation bet-hedging hypothesis for social evolution predicts that (i) variance in reproductive success should be related to environmental variation, (ii) variance in reproductive success should be related to the potential for cooperation in a group, and (iii) the potential for cooperation should be related to environmental variation. Using data from a 10-y study of cooperatively breeding superb starlings (Lamprotornis superbus) living in a temporally and spatially variable savanna ecosystem, I found that variance in reproductive success declined with increasing environmental quality (temporal variation), increasing territory quality (spatial variation), and increasing group size (potential for cooperation), which is itself related to environmental variation. To understand the adaptive value of cooperative breeding behavior in variable environments, researchers must consider both mean and environmentally induced variance in fecundity. Determining how spatiotemporal environmental variation drives risk-averse strategies may provide insights into the evolution of complex social behavior.