Variable age structure and apparent density dependence in survival of adult ungulates

Large herbivores have strongly age-structured populations. Because recruitment often decreases as population density increases, in unexploited populations the proportion of older adults may increase with density. Because survival senescence is typical of ungulates, ignoring density-dependent changes in age structure could lead to apparent density-dependence in adult survival. To test for density dependence in adult survival, we used data from three populations that underwent considerable changes in density. Bighorn sheep (Ovis canadensis) on Ram Mountain, Alberta, ranged from 94 to 232, mountain goats (Oreamnos americanus) on Caw Ridge, Alberta, varied from 81 to 147, and estimates of roe deer (Capreolus capreolus) older than 1 year at Chizé, France, ranged from 157 to 569. We used recent developments of capture-mark-recapture modelling to assess the response of adult survival to changes in density when age structure was and was not taken into account. Survival rates were 10-15% higher during the prime-age stage than during the senescent stage for all sex-species combinations. When adults were pooled into a single age class there was an apparent negative effect of density on female survival in bighorns and roe deer, and negative trends for female mountain goats, male roe deer and male bighorn sheep. When age class was taken into account, there were no significant effects of density on adult survival. Except for male mountain goats, the strength of density dependence was lower when age was taken into account. In ungulate populations, age structure is an important determinant of adult survival. Most reports of density dependence in adult survival may have been confounded by changes in age structure.