Assessment of costs of reproduction in a pelagic seabird using multistate mark-recapture models

We used a long-term population band-resight survey database, a parallel reproduction database, and multistate mark-recapture analysis to assess the costs of reproduction, a keystone concept of life-history evolution, in Nazca boobies (Sula granti) from Punta Cevallos, Isla Española, Galápagos, Ecuador. We used eight years of resight and breeding data to compare models that included sex- and state-specific survival probabilities and probabilities of transition between reproductive states using multistate mark-recapture models. Models that included state-specific effects were compared with models lacking such effects to evaluate costs of reproduction. The top model, optimizing the trade-off of model simplicity and fit to the data using the Akaike Information Criterion (AIC), showed evidence of a temporally varying survival cost of reproduction: nonbreeders showed higher annual survival than breeders did in some years. Because increasing investment among breeders showed no negative association with survival and subsequent breeding success, this evidence indicates a cost to both males and females of initiating, but not of continuing, a reproductive attempt. In some cases, breeders reaching the highest reproductive state (fledging an offspring) showed higher survival or subsequent breeding success than did failed breeders, consistent with differences in overall quality that promote both survival and reproduction. Although a male-biased adult sex ratio was observed in this population of Nazca boobies, models of state- and sex-specific survival and transition probabilities were not supported, indicating that males and females do not incur different costs of reproduction, and that the observed sex ratio bias is not due to sex-specific adult mortality.