Ungulate herbivores as drivers of aspen recruitment and understory composition throughout arid montane landscapes

Abstract

Herbivory by wild and domestic ungulates can influence tree recruitment and understory forest communities throughout the world. Herbivore-driven declines in tree recruitment have been observed for quaking aspen (Populus tremuloides), a foundation species whose health and management is recognized as a critical priority throughout much of its range. Livestock fencing is commonly used to promote aspen regeneration, but its effectiveness is rarely assessed, especially across large spatial scales. Using a livestock-reduction experiment, we evaluated the effects of ungulate herbivory on aspen in the Great Basin and southern Cascades, an expansive and environmentally heterogeneous region where aspen faces the interacting threats of climate change, conifer encroachment, and herbivory. We found that livestock fencing only reduced the intensity of herbivore browsing on aspen when wild ungulate abundance was low and did not increase stem densities of aspen recruits. Contrary to expectations, wild ungulate abundance was a strong driver of browsing intensity on juvenile aspen within fenced, but not unfenced, aspen stands, and when the abundance of these herbivores was high, browsing intensity in fenced stands exceeded that in unfenced stands. The density of aspen recruits decreased with browsing intensity on juvenile aspen and with the density of both adult aspen and conifers, suggesting that both herbivory and intraand interspecific competition are important drivers of recruitment. Fire history was also an important driver of recruitment, with stands that burned 10-20 years ago having the greatest density of aspen recruits. Finally, in the stand understory, we found that livestock fencing decreased forb cover, increased shrub species richness, and increased the cover of exotic annual grasses, a group dominated by Bromus tectorum. This latter finding suggests that livestock fencing may not be appropriate in areas where controlling the spread of this invader is a priority. In sum, our findings indicate that aspen recruitment is limited by browsing by both wild and domestic ungulates, is mediated by competition with neighboring trees and fire history, and will