Critical issues in experimental studies of prosociality in non-human species

Prosociality and acts of altruism are defined as behaviours which benefit another with either no gain or some immediate cost to the self. To understand the evolutionary origins of these behaviours, in recent years, studies have extended to primate species; however, studies on non-primates are still scarce. In light of the fact that phylogenetic closeness to humans does not appear to correlate with prosocial tendencies, but rather differences in the propensity towards prosociality may be linked to allomaternal care or collaborative foraging, it appears that convergent selection pressures may be at work in the evolution of prosociality. It would hence seem particularly important to extend such studies to species outside the primate clade, to allow for comparative hypothesis testing of the factors affecting the evolution of prosocial behaviours. In the current review, we focus on the experimental paradigms which have been used so far (i.e. the prosocial choice task, helping paradigms and food-sharing tests) and highlight the strengths and weaknesses of each method. In line with the aim of encouraging a broader comparative approach to the topic of prosociality, particular emphasis is placed on the methodological issues that need to be taken into account. We conclude that although a number of the paradigms used so far may be successfully applied to non-primate species, there is a need to simplify the cognitive demands of the tasks and ensure task comprehension to allow for a 'fair' comparative approach of prosocial tendencies across species.