Linking wolves to willows via risk-sensitive foraging by ungulates in the northern Yellowstone ecosystem

After an absence of approximately 70 years, gray wolves (Canis lupus) were re-introduced into Yellowstone National Park in the mid-1990s. We studied the potential influence of wolf/ungulate interactions upon willow (Salix spp.) growth in the valleys of the Gallatin and Lamar Rivers, as well as Slough and Soda Butte Creeks, in the northern Yellowstone ecosystem. When we compared willow heights from photographs taken prior to 1998 (willows <2 m tall) with those taken in 2004, we found an increase in willow height for 22 of 42 sites within the study area. Based on comparisons of the chronosequence photos, since wolf introduction none of the 16 upland riparian sites showed an increase in willow height, while 22 of 26 of the valley-bottom riparian sites had willow height increases. In 2004, willow height exhibited a strong inverse relationship with the percentage of browsed stems (r = 0.81, p < 0.01, n = 42). Results of regressions for valley-bottom sites indicated that view distance, impediment distance, and the number of bison (Bison bison) flops were inversely related to willow height ( p 0.02). Increased willow heights were not significantly ( p = 0.18) related to patterns of moisture availability. Willow height increases documented in this study appear to have been at least partially due to behaviorally mediated trophic cascades involving wolves and ungulates, via a mechanism of predation risk. While willow release (i.e., increased height growth) within the study area is in a very early stage, results suggest potentially important indirect effects of a top carnivore in a terrestrial food chain that may aid in the restoration of riparian species and the preservation of biodiversity. # 2006 Elsevier B.V. All rights reserved.