Demographic performance of a large herbivore: effects of winter nutrition and weather

Variation among demographic rates for a population reflects the allocation of available energy by individuals to competing life-history strategies. Species exhibiting slow-paced life histories often prioritize energy allocation to adult survival over any single reproductive event, therefore maximizing future reproductive potential. Survival of adult female ungulates is generally high with little variability, whereas survival of young is lower and often highly variable. When adult survival is high with low variability, juvenile survival may have a proportionally greater effect on population growth or decline. Weather also may affect population dynamics directly by influencing survival of young or adults, or indirectly through changes in nutritional condition of adult females that influence population growth rates. We experimentally manipulated forage availability during winter, by supplementing native forage with high-energy pelleted feed ad libitum, to a subset of a population of mule deer (Odocoileus hemionus) to understand the effects of winter nutrition on survival of adult females and their young born the subsequent summer. We evaluated the effects of winter nutrition, individual-based parameters, and environmental covariates on survival of adult female mule deer from 2013 to 2018, and neonatal mule deer from 2014 to 2016. We documented a 26% decrease in annual survival of adult female mule deer in 2017 in response to increased snowpack during the preceding winter. Neonates born to females that receive enhanced nutrition during winter preceding parturition had higher survival to weaning (0.49, SE = 0.12), compared to neonates born to females that did not receive enhanced nutrition (0.29, SE = 0.07).We observed no effect of enhanced winter nutrition on survival of adult females. Our results suggested winter nutrition of maternal females may influence juvenile survival and demonstrates the importance of forage quality available to adult females during mid-pregnancy. Although we were unable to detect an effect of winter forage on survival of adults, direct effects of deep winter snow resulted in lower survival of adult females. Low survival of adult females in our study population is indicative of a declining population.