Foraging associations are related with helping interactions in a cooperatively breeding bird

Kin selection has been the main hypothesis explaining helping behaviour in cooperative breeders, with evidence being largely based on the observation that helpers tend to provide to related offspring. However, kin-biased help could conceal additional, mechanisms contributing to the maintenance of cooperation. Under pay-to-stay, group augmentation and partner choice hypotheses, a range of direct benefits can arise through helping. Here, we explored this potential mechanism by testing whether the social associations of breeding individuals were related with the help that they received from non-breeding individuals. We collected social associations from PIT-tagged sociable weavers, Philetairus socius, at RFID feeding stations, which allowed us to compare associations between breeders and either their helpers (mostly kin) or their other kin that did not help--before, during and after reproduction. Using correlative tests and data-driven simulations, we show that helpers have stronger foraging bonds with breeders than non-helping kin, and that these stronger bonds are present both prior and post breeding. Furthermore, helper-breeding female social affiliations were positively correlated with the amount of help provided. Our results suggest that direct benefits of social associations complement kin selection to determine helping decision, and that these in turn influence future social associations.