What shapes fitness costs of reproduction in long-lived iteroparous species? A case study on the Alpine ibex.

The fitness costs of reproduction can be masked by individual differences, and may only become apparent during adverse environmental conditions. Individual differences, however, are usually assessed by reproductive success, so how fitness costs are influenced by the interplay between the environmental context and overall individual differences requires further investigation. Here, we evaluated fitness costs of reproduction based on 15 yr of monitoring of individual Alpine ibex (Capra ibex) during a period when the population was affected by a severe disease outbreak (pneumonia). We quantified fitness costs using a novel multi-event capture-mark-recapture (CMR) modeling approach that accounted for uncertainty in reproductive status to estimate the survival and reproductive success of female ibex while also accounting for overall individual heterogeneity using mixture models. Our results show that the ability of females to reproduce was highly heterogeneous. In particular, one group including 76% of females had a much higher probability of giving birth annually (between 0.66 and 0.77, depending on the previous reproductive status) than females of the second group (24% of females, between 0 and 0.05 probability of giving birth annually). Low reproductive costs in terms of future reproduction occurred and were independent of the pneumonia outbreak. There was no survival cost of reproduction either before or after the epizootic, but the cost was high during the epizootic. Our findings indicate that adverse environmental conditions, such as disease outbreaks, may lead to survival costs of reproduction in long-lived species and select against females that have a high reproductive effort. Thereby, the occurrence of adverse conditions increases the diversity of reproductive tactics within a population.