Best of both worlds? Helpers in a cooperative fairy-wren assist most to breeding pairs that comprise a potential mate and a relative

In cooperative breeders, individuals forego independent reproduction and help others raise offspring. Helping is proposed to be driven by indirect benefits from raising relatives, and/or direct benefits from raising additional recruits or helping itself. We propose that consideration of social context is also important, in particular the characteristics of the breeding pair: helping may also serve to lighten the workload of - or maintain social bonds with - breeders (e.g. kin, potential mates) who in turn can offer benefits to helpers (e.g. prolonged nepotism, future mating, future production of relatives). Here, we test this hypothesis, while controlling for potential direct and indirect benefits from raising offspring, in purple-crowned fairy-wrens (Malurus coronatus) exhibiting variation in social group composition, and thus, breeder value. We show that helper provisioning rates to the nest were explained by characteristics of breeders that helpers assisted, rather than benefits from raising offspring. The presence of at least one related breeder was a prerequisite to help, but helpers provisioned most if assisting a relative and potential mate. Neglecting to take group composition into account would have led to misinterpretation of our results. A comprehensive understanding of the evolution of cooperative breeding hence requires nuanced consideration of social context.