Food supplementation experiments revisited: verifying that supplemental food is used by its intended recipients

Abstract

Field-based food supplementation experiments can determine whether populations are limited by natural food availability. However, they may yield spurious results if added food is hoarded by dominants, exploited by immigrants, or delivered ineffectively. Surprisingly, population-level approaches accounting for these potential sources of bias have not been established. We explored responses of free-ranging snowshoe hares ( Lepus americanus Erxleben, 1777) to food supplementation by contrasting per capita browsing rates on three study sites with added food with those on three control sites during two winters. Food augmentation reduced foraging pressure by hares; this reduction was significant when browse species of high dietary importance were considered. By implication, hares on manipulated sites switched from favored natural foods to supplemental food, meaning that the extra food found its mark and was not heavily exploited by immigrants or hoarded by dominants. Demographic responses to food addition were not detected on the manipulated sites, indicating that the study population was not food limited. Given the success with which we detected the signal of supplementation and eliminated potential confounding factors, we suggest that future studies using food addition in the context of herbivore population dynamics would benefit from a more mechanistic approach; this may involve measurement of browsing rates on manipulated and control sites.