Foraging decisions of bison for rapid energy gains can explain the relative risk to neighboring plants in complex swards

Herbivores commonly base their foraging decisions not only on the intrinsic characteristics of plants, but also on the attributes of neighboring species. Although herbivores commonly orient their food choices toward the maximization of energy intake, the impact of such choices on neighboring plants remains largely unexplored. We evaluated whether foraging decisions by herbivores aiming at a rapid intake of digestible energy could explain multiple neighboring effects in complex swards. Specifically, we assessed how spatial patterns of occurrence of Carex atherodes, a highly profitable sedge species, could control the risk of bison (Bison bison) herbivory for seven other plant species. The foraging behavior of 70 free-ranging bison was evaluated in their natural environment during summer, and then related to plant characteristics. We used this information to estimate the instantaneous intake rate of digestible energy at individual feeding stations. We found that neighbor contrast defense and associational susceptibility can both be explained by simple foraging rules of energy maximization. Energy gains were higher when C. atherodes was consumed while avoiding the species for which we detected neighbor contrast defense. The lower intake rate associated with their consumption was due to an increase in handling time caused by their small size relative to C. atherodes. Bison also had higher energy gains by consuming instead of avoiding the plant species that experienced associational susceptibility. Because most of these plants were at least as tall as C. atherodes, their presence increased the heterogeneity of the grazed stratum. Avoiding their consumption increased handling time thereby reducing the instantaneous rate of energy intake. Overall, we found that bison adjust their fine-scale foraging decisions to vertical and horizontal sward structures in a way that maximizes their energy intake rate. Energy maximization principles thus provide a valuable framework to evaluate a broad-range of neighboring effects for prey faced with generalist consumers.