The scale of the 1988 fires in Yellowstone National Park (YNP) raised numerous questions for the management of natural areas subject to large, infrequent disturbances. An important management issue in YNP involves the interaction of large-scale fire with the large assemblage of native ungulates and vegetation dynamics in the landscape. We used landscape modeling and field studies to address basic questions about the effects of fire scale and heterogeneity on resource utilization and survival of free-ranging elk (Cervus elaphus) and bison (Bison bison), and the production and regeneration of preferred forage grasses and aspen in northern Yellowstone Park. More specifically, we asked (1) how fire size interacts with winter severity to control ungulate feeding behavior and survival, both in the initial postfire winter, when fire reduces forage, and in later postfire winters, when fire augments forage; (2) how fire pattern (e.g., clumped vs. dispersed burn sites) modifies the effects of fire size; (3) which environmental factors, including fire, influence selection of feeding areas by wintering ungulates at a variety of scales, from a single feeding station to the entire northern winter range; and (4) how the size and spatial pattern of burning influence regeneration of aspen (Populus tremuloides), a preferred and heavily browsed species in YNP. We focus on elk and bison because these are by far the most numerous ungulates in the area (Houston 1982), and we have chosen to examine winter grazing and browsing for several reasons. Winter range conditions are the primary determinant of ungulate survival and reproduction in Yellowstone, and winter utilization of the vegetation by ungulates appears to be intense in some areas. Ungulates make distinct foraging choices in the winter as in the rest of the year, and burn patterns may influence those choices in ways that we represent as hypotheses described later. In addition, the activities of animals can be readily monitored in the winter, and the exact locations of feeding and bedding sites can be determined. Travel routes are easily monitored, and the ability to sight animals is high; therefore, group locations and sizes can be readily determined. This research complements ongoing studies in Yellowstone by expanding the spatial scale at which plant-herbivore dynamics are considered and by explicitly addressing the effects of spatial heterogeneity. We produced a spatially explicit simulation model of the winter range that predicts plant and ungulate dynamics under varying fire sizes, fire patterns, winter weather scenarios. The model and field studies will generate quantitative comparisons of the effects of large and small fires on ungulate survival and will thereby permit the simulation of the effects of alternative fire management scenarios.
Wolf, Canis lupus, Avoidance Behaviour of American Elk, Cervus elaphus, in Jasper National Park, Alberta