From recruitment to senescence: food shapes the age-dependent pattern of breeding performance in a long-lived bird

We used a long-term data set (26 years) from Audouin's Gull (Larus audouinii), a long-lived seabird, to address the relationship between the age-dependent pattern of reproductive performance and environmental conditions during breeding. Although theoretical models predict that the youngest and oldest breeders (due to inexperience and senescence, respectively) will perform less well than intermediate age classes, few empirical data exist regarding how this expected pattern varies with food availability. To assess the influence of age and food availability (corrected by population size of the main consumers to take into account density dependence) on a number of breeding parameters (laying dates, egg volume, clutch size, and hatching success), we modeled mean and variances of these parameters by incorporating heterogeneity into generalized linear models. All parameters varied with age and to different degrees, depending on food availability. As expected, performance improved with increased food supply, and the observed age pattern was quadratic, with poorer breeding performances occurring in extreme ages. For most parameters (except for laying dates, for which age and food did not interact), the pattern changed with food somewhat unexpectedly; the differences in performance between age classes were higher (i.e., the quadratic pattern was more noticeable) when food was more readily available than when food availability was lower. We suggest that, under poor environmental conditions, only high-quality individuals of the younger and older birds bred and that the differences in breeding performance between age classes were smaller. Although variances for egg volume were constant, variances for laying dates were highest for the youngest breeders and tended to decrease with age, either due to the selection of higher-quality individuals or to a greater frequency of birds skipping breeding with age, especially when food was in low supply. Our results show that mean and variances of breeding parameters changed with age, but that this pattern was different for each parameter and also varied according to food availability. It is likely that, other than food, certain additional factors (e.g., sex, cohort effects, density dependence) also influence changes in breeding performance with age, and this may preclude the finding of a common pattern among traits and among studies on different taxa.