Scaling the effects of moose browsing on forage distribution, from the geometry of plant canopies to landscapes

Landscape heterogeneity influences large herbivores by altering their feeding rates, but as herbivores attempt to maximize feeding rates they also create spatial heterogeneity by altering plant growth. Herbivore feeding rates thus provide a quantitative link between the causes and consequences of spatial heterogeneity in herbivore-dominated ecosystems. The fractal geometry of plant canopies determines both the density and mass of twigs available to foraging herbivores. These properties determine a threshold distance between plants (d*) that distinguishes the mechanisms regulating herbivore intake rates. When d* is greater than the actual distance between plants (d), intake is regulated by the rate of food processing in the mouth. But when d*, d, intake is regulated by the rate at which the herbivore encounters new plants. Alterations to plant geometry due to past browsing could change the rate at which herbivores encounter and process bites of plant tissue, modify d* relative to d, and thus change intake rates and the distribution of mechanisms regulating it across landscapes. We measured changes in the geometry of aspen (Populus tremuloides) and balsam fir (Abies balsamea) saplings along gradients of moose browsing from 2001 to 2005 at Isle Royale National Park, Michigan, USA. For aspen saplings, fractal dimension of bite density, bite mass, and forage biomass responded quadratically to increasing moose browsing and were greatest at similar to 3-4 g.m(-2).yr(-1) consumption. For balsam fir, in contrast, these same measures declined steadily with increasing moose browsing. The different responses of plant canopies to increased browsing altered d* around plants. In summer, d* >= d for aspen saplings at all prior consumption levels. Food processing therefore regulated summer moose feeding rates across our landscapes. In winter, changes in bite mass due to past browsing were sufficient to cause d* < d for aspen and balsam fir. Therefore, travel velocity and food processing jointly regulated intake rate during winter. Browsing-induced changes in the small-scale geometry of plant canopies can determine intake rate at larger spatial scales by changing d* relative to d and, hence, which mechanisms determine intake rate, essentially altering how herbivores sense the distribution of their food resources.