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.

Group-size-mediated habitat selection and group fusion-fission dynamics of bison under predation risk.

For gregarious animals the cost-benefit trade-offs that drive habitat selection may vary dynamically with group size, which plays an important role in foraging and predator avoidance strategies. We examined how habitat selection by bison (Bison bison) varied as a function of group size and interpreted these patterns by testing whether habitat selection was more strongly driven by the competing demands of forage intake vs. predator avoidance behavior. We developed an analytical framework that integrated group size into resource selection functions (RSFs). These group-size-dependent RSFs were based on a matched case-control design and were estimated using conditional logistic regression (mixed and population-averaged models). Fitting RSF models to bison revealed that bison groups responded to multiple aspects of landscape heterogeneity and that selection varied seasonally and as a function of group size. For example, roads were selected in summer, but not in winter. Bison groups avoided areas of high snow water equivalent in winter. They selected areas composed of a large proportion of meadow area within a 700-m radius, and within those areas, bison selected meadows. Importantly, the strength of selection for meadows varied as a function of group size, with stronger selection being observed in larger groups. Hence the bison-habitat relationship depended in part on the dynamics of group formation and division. Group formation was most likely in meadows. In contrast, risk of group fission increased when bison moved into the forest and was higher during the time of day when movements are generally longer and more variable among individuals. We also found that stronger selection for meadows by large rather than small bison groups was caused by longer residence time in individual meadows by larger groups and that departure from meadows appears unlikely to result from a depression in food intake rate. These group-size-dependent patterns were consistent with the hypothesis that avoidance of predation risk is the strongest driver of habitat selection.