Individual variability in the phenology of an asynchronous penguin species induces consequences on breeding and carry-over effects

Phenology is a major component of animals' breeding, as they need to adjust their breeding timing to match optimal environmental conditions. While the effects of shifting phenology are well-studied on populations, few studies emphasise its ecological causes and consequences at the inter-individual level. Using a 20-year monitoring of more than 2500 breeding events from similar to 500 breeding little penguins (Eudyptula minor), a very asynchronously breeding seabird, we investigated the consequences of late breeding on present and next breeding events. We found that individuals breeding later had reduced breeding success, lighter chicks at fledging, lower probability of laying a second clutch, and decreased parents' post-breeding body condition. Importantly, we found important cycling effects where delayed breeding during a given year led to significantly later laying date, lower breeding probability and lower breeding success when they breed during the next season, suggesting potential carry-over effects from one season to the next. To further understand the causes of such variability in phenology while earlier breeding is associated with better individual fitness, we aimed to assess intrinsic differences amongst individuals. We showed that the heterogeneity in breeding timing was partly fixed, the laying date being a significantly repeatable behaviour (17%), asking for more studies on heritability or early-development effects. This extensive study highlights the combined roles of carry-over effects and intrinsic differences on individual phenology, with important implications on breeding capacity through life.