Wolf presence and increased willow consumption by Yellowstone elk: implications for trophic cascades.

Recent increases in the height and growth ring width of willow (Salix spp.) and other woody plants in the Greater Yellowstone Ecosystem (GYE) have been attributed to a behaviorally mediated trophic cascade from wolves (Canis lupus) to elk (Cervus elaphus) to willows. This hypothesis predicts that individual elk consume less willow in response to the presence of wolves, but this prediction has not been directly tested with data from elk. We collected 727 fecal samples from elk in the Gallatin Canyon portion of the GYE over three winters and used microhistological methods to quantify the proportion of willow in each sample. We then tested the effect of wolf presence on willow consumption by elk, controlling for the effects of snow conditions, sex, and habitat type. During the period of study, 8-17 wolves occupied the study area, and wolves were locally present on 49% of 260 sampling days, stratified at two-week intervals across three drainages. Over the three years combined, willow consumption was related to snow conditions, wolf presence, and a wolf X sex interaction. As expected, willow consumption increased with deeper and less penetrable snow, and this effect was strong. Contrary to expectation, willow consumption increased in the presence of wolves. As with other aspects of antipredator behavior, wolves had different effects on willow consumption by males and females. Finally, we aggregated the data to estimate winter-long mean willow consumption within each drainage; at this broader scale, willow consumption again increased as predation risk increased. In summary, willow consumption was more strongly affected by snow conditions than by the presence of wolves. Interactions between elk and willow were affected by wolves, but not as predicted by the hypothesis that wolf presence favors willow release through a reduction in the selection of willow by individual elk. If a trophic cascade is operating, our results suggest that a decline in the size of the elk population (to roughly one-half its size immediately prior to wolf recovery) may be more important than changes in the willow consumption of individual elk. Finally, reduced grazing of herbaceous vegetation may be equally important for vegetation dynamics.