Consequences of 'load-lightening' for future indirect fitness gains by helpers in a cooperatively breeding bird.

1. Helpers that invest energy in provisioning the offspring of related individuals stand to gain indirect fitness benefits from doing so. First, if the helper's effort is additional to that of the parents (additive) the productivity of the current breeding attempt can be increased. Secondly, if the parents reduce their workload (compensation) this can result in future indirect fitness gains to the helper via increased breeder survival; termed 'load-lightening'. 2. Long-tailed tits (Aegithalos caudatus) have a cooperative breeding system in which helpers assist kin and parents exhibit both additive and compensatory reactions in the presence of helpers. Offspring from helped nests are heavier and more likely to recruit into the breeding population, thus helpers gain indirect fitness benefits from increasing the productivity of the current breeding attempt. Despite breeders' reduction of feeding effort in the presence of helpers, previous investigations found no subsequent increase in breeder survival. 3. The aim of this study was to test the hypothesis that load-lightening resulted in indirect fitness benefits for helpers. We used data from a 14-year study to investigate the provisioning rate, survival and future fecundity of male and female long-tailed tits that did and did not receive help at the nest. 4. We found an asymmetrical response to the presence of helpers at large brood sizes. Males reduced their feeding rate more than females, and this differential response was reflected in a significant increase in male survival when provisioning large broods assisted by helpers. We found no evidence of any increase in future fecundity for helped breeders. 5. The finding that males reduce their provisioning rate in the presence of helpers (at large brood sizes) to a greater degree than females, and that this is reflected in an increase in survival rate for males only, implies that the survival increase is caused by the reduction in work-rate rather than a non-specific benefit of a larger group size. 6. The marginal benefits of help for breeder survival are likely to be more difficult to identify than the increased productivity at helped nests, but should not be overlooked when investigating the potential indirect fitness gains that supernumeraries can accrue by helping.