Fitness can be profoundly influenced by the age at first reproduction (AFR), but to date the AFR-fitness relationship only has been investigated intraspecifically. Here we investigated the relationship between AFR and average lifetime reproductive success (LRS) across 34 bird species. We assessed differences in the deviation of the Optimal AFR (i.e., the species-specific AFR associated with the highest LRS) from the age at sexual maturity, considering potential effects of life-history as well as social and ecological factors. Most individuals adopted the species-specific Optimal AFR and both the mean and Optimal AFR of species correlated positively with lifespan. Interspecific deviations of the Optimal AFR were associated with indices reflecting a change in LRS or survival as a function of AFR: a delayed AFR was beneficial in species where early AFR was associated with a decrease in subsequent survival or reproductive output. Overall, our results suggest that a delayed onset of reproduction beyond maturity is an optimal strategy explained by a long lifespan and costs of early reproduction. By providing the first empirical confirmations of key predictions of life-history theory across species, this study contributes to a better understanding of life-history evolution.
Helping to rear the offspring of others may be a way for younger birds to gain access to future reproduction especially when turnover of breeding opportunities is low. However, this explanation is not applicable to cases where adults also help, or when roles shift between helpers and breeders. Over a period of 6‐yr, we studied a marked population of azure‐winged Magpie (Cyanopica cyanus) breeding in a non‐territorial, colonial system. Magpies bred in a highly flexible cooperative system, in which individuals helped at different stages of the breeding cycle, including nest building, feeding the incubating female and feeding the young and removing the faecal sacs. On average, 50% of hatched nests were assisted by helpers‐at‐the‐nest, and nest success appeared to be positively related to the presence of helpers. Helpers were predominantly males. Although juveniles were more likely to help, both juvenile and adult birds helped. Individual birds behaved as helpers either as a first‐option or after having attempted their own breeding (second‐option helpers). An individual helper may assist more than one nest during the same breeding season and in different breeding seasons. Reversals between breeder and helper roles were common in both directions, within a breeding season and between years. Helping behaviour is an option for almost any member of the colony. Therefore, hypotheses related to the enhancement of future breeding opportunities for juveniles can be discarded as general explanation of helping in this species. Although the decision to help appeared to be influenced by proximal environmental conditions hindering successful breeding, the associated benefits of helping as opposed to simply recovering for future reproduction, especially for former breeders, deserve further study.
For cooperatively breeding birds, it has been proposed that breeders should reduce their investment in eggs when they count on helpers, because this can be compensated for by helpers provisioning of nestlings. Data from some species have supported this prediction, but this is not the case in others. It has also been proposed that mothers should not reduce but rather increase investment if the presence of helpers enhances the reproductive value of offspring, a pattern that might also influence egg production as long as helpers are predictable for laying females. Here, we studied maternal expenditure in eggs and clutches in the Iberian magpie, to see whether mothers reduce their expenditure at the egg stage in the presence of helpers. Our results show that investment in clutches varied depending on the year, date in the season and age of the mother, but there were no reductions in maternal expenditure per individual egg when they counted on helpers. On the contrary, a pattern emerged in the opposite direction of more investment in eggs associated with the future presence of helpers at the nestling stage. Our data suggest that the predictability of helpers, along with the type of benefits accrued from the contribution of helpers, may be crucial to understanding the reaction of mothers at egg production.
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