Abstract
Alloparental care in cooperatively breeding species may alter breeder age-specific survival and reproduction and subsequently senescence. The helping behaviour itself might also undergo age-related change, and decisions to help in facultative cooperative breeders are likely to be affected by individual condition. Helpers in long-tailed tits Aegithalos caudatus assist relatives after failing to raise their own brood, with offspring from helped nests being more likely to recruit into the breeding population. Using data collected over 25 years, we examined the age trajectories of survival and reproduction in adult long-tailed tits to determine how these were affected by the presence or absence of helpers and how helper behaviour changed with age. There was evidence for increased reproductive performance with breeder age, but no effect of age on the probability of survival. We found no evidence of significant senescent decline in survival or reproductive performance, although individuals accrued less inclusive fitness in their last year of life. Lifetime reproductive success was positively related to both reproductive life span and body mass. Within a season, breeders that were assisted by helpers enjoyed greater reproductive success through enhanced offspring recruitment in the following year. We found no evidence that age affected an individual's propensity to help, or the amount of indirect fitness accrued through helping. We found a positive correlation between life span and multiple components of reproductive success, suggesting that individual variation in quality underpins age-related variation in fitness in this species. Helping decisions are driven by condition, and lifetime inclusive fitness of immigrants was predicted by body mass. These findings further support individual heterogeneity in quality being a major driver for fitness gains across the life course of long-tailed tits.