Interaction of beaver and elk herbivory reduces standing crop of willow

Populations of beaver and willow have not thrived in riparian environments that are heavily browsed by livestock or ungulates, such as elk. The interaction of beaver and elk herbivory may be an important mechanism underlying beaver and willow declines in this competitive environment. We conducted a field experiment that compared the standing crop of willow three years after simulated beaver cutting on paired plants with and without intense elk browsing (similar to85% utilization rate). Simulated beaver cutting with intense elk browsing produced willow that was small (biomass and diameter) and short, with far fewer, but longer, shoots and a higher percentage of dead biomass. In contrast, simulated beaver cutting without elk browsing produced willow that was large, tall, and leafy, with many more, but shorter, shoots (highly branched) and a lower percentage of dead biomass. Total stem biomass after three years was 10 times greater on unbrowsed plants than on browsed plants. Unbrowsed plants recovered 84% of their pre-cut biomass after only two growing seasons, whereas browsed plants recovered only 6%. Thus, the interaction of beaver cutting and elk browsing strongly suppressed the standing crop of willow. We predict that a lack of willow suitable as winter food for beaver can cause beaver populations to decline, creating a feedback mechanism that reduces beaver and willow populations. Thus, intense herbivory by ungulates or livestock can disrupt beaver-willow mutualisms that naturally occur in less competitive environments.