Indirect fitness benefits gained through kin-selected helping are widely invoked to explain the evolution of cooperative breeding behavior in birds. However, the impact of helpers on productivity of helped broods can be difficult to determine if the effects are confounded by territory quality or if the benefit of helpers is apparent only in the long term. In riflemen Acanthisitta chloris, helping and group membership are effectively decoupled as adult helpers are individuals that have dispersed from their natal territory and live independently from breeders in "kin neighborhoods." Nevertheless, helpers direct their care toward close relatives, suggesting that helping provides indirect fitness benefits. The aim of this study was to examine the benefits of helpers to recipient offspring in the rifleman, investigating both short- and long-term effects. The total amount of food delivered to nestlings in helped broods was greater than that received by broods without helpers. This did not result in any short-term increase in nestling mass or nestling body condition nor was there any reduction in length of the nestling period at helped nests. However, helpers were associated with a significant increase in juvenile recruitment, with twice the proportion of fledglings surviving to the next breeding season from helped broods relative to unhelped broods. Thus, helpers gain indirect fitness by improving the survival of kin, and in contrast to a previous study of riflemen, we conclude that kin selection has played a key role in the evolution of cooperative breeding in this species.
Keywords: cooperative breeding; inclusive fitness; indirect fitness; kinship..