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                <title xml:lang="en">Age-related breeding success in little penguins: a result of selection and ontogenetic changes in foraging and phenology</title>
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              <p>Reproductive performance typically improves with age, reaching a plateau at middle age and subsequently declining in older age classes (senescing individuals). Three potential non-exclusive mechanisms can explain the improvement in reproductive performance with age: (1) selection (poor quality individuals are removed from the population with increasing age), (2) constraint (individual efficiency increases through experience), and (3) restraint (reproductive investment increases with age as the residual reproductive value decreases). While all three mechanisms received strong empirical support, few studies have aimed at teasing apart those hypotheses and understanding their underlying functioning. In little penguins (Eudyptula minor), we used a 19-yr longitudinal dataset on breeding and foraging of more than 450 individuals to investigate the effect of age on breeding success. We separated within-individual from among-individual age effects using state-of-the-art statistical methods (within-subject centering and population change decomposition). We then assessed whether within-individual changes in breeding resulted from ontogenetic changes in foraging performances, breeding phenology or access to mates and nest sites. Fidelity and assortative pairing explained the high correlation in male and female ages within a pair. Breeding performances followed a typical bell-shaped curve with performance increasing up to 8 yr old, before reaching a plateau and subsequently declining after age 16. Both selection and within-individual processes occurred, although within-individual changes dominated differences in age-dependent breeding success. The selective appearance had almost no effect (apart from ages 2–3), and selective disappearance mostly affected changes at old ages (above 16), although they were also responsible for the slight increase in reproductive performances from ages 5–8. Focusing on within-individual changes, birds exhibited higher performances at middle ages, with birds foraging better, laying earlier and changing partner and nest less often. Their reproductive investment did not vary with age for females and slightly decreased for males. This supports the constraint hypothesis but not the restraint one. Finally, the increase in breeding performances at young ages was explained by the age-related increase in foraging performances during chick-rearing and advancement of laying. In contrast, reproductive senescence was defined by a general decrease in bird performances.</p>
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