Abstract
Reproductive senescence is ubiquitous in mammals. However, patterns of senescence vary across reproductive traits, even within populations, perhaps because of differences in selection pressures, physiological constraints, and responses to environmental conditions. We investigated reproductive senescence in wild female polar bears (Ursus maritimus), using 31 years of capture-recapture data from the Svalbard area. We studied the influence of environmental conditions on age-specific litter production and litter size using generalized linear mixed models. Further, using a capture-recapture model that handles the dependency between vital rates of individuals belonging to the same family unit, we assessed maternal-age-related changes in first year cub and litter survival. We provide clear evidence for reproductive senescence in female polar bears. Litter production and litter size peaked in middle-aged females and declined sharply afterward. By contrast cub and litter survival did not decline after prime age. We found no evidence of terminal investment. The reproductive output of all females was affected by sea-ice conditions during the previous year and the Arctic Oscillation, with some effects differing greatly between age groups. Old females were affected the most by environmental conditions. Our results suggest that the decline in reproductive output is a combination of fertility and body-condition senescence, with a weak contribution of maternal-effect senescence, possibly due to benefits of experience. Further, as predicted by evolutionary theory, senescence appears to be a consequence of failures in early stages of the reproductive cycle rather than in late stages, and environmental variation affected old females more than prime-aged females. Our study emphasizes the need to study several reproductive traits and account for environmental variation when investigating reproductive senescence. Differences in senescence patterns across reproductive traits should be interpreted in light of evolutionary theory and while considering underlying physiological drivers.