Quantifying the Indirect Effect of Wolves on Aspen in Northern Yellowstone National Park: Evidence for a Trophic Cascade?

Yellowstone National Park is renowned for its incredible wildlife, and perhaps the most famous of these species is the gray wolf, which was reintroduced to the Park in the mid-1990s. After reintroduction, it was highly publicized by scientists, journalists, and environmentalists that the wolf both decreased elk density and changed elk behavior in a way that reduced elk effects on plants, a process known as a “trophic cascade.” Aspen, which is eaten by elk in winter, is one species at the forefront of Yellowstone trophic cascade research because it has been in decline across the Park for over a century. However, due to the challenges of measuring trophic cascades, there is continued uncertainty regarding the effects of wolves on aspen in northern Yellowstone. Thus, the purpose of my dissertation was to provide a comprehensive test of a trophic cascade in this system. Specifically, I used 20 years of data on aspen, elk, and wolves in Yellowstone to: 1) clarify annual trends in browsing and height of young aspen (a proxy for regeneration) after wolf reintroduction, 2) assess the influence of wolves scaring elk on aspen (“trait-mediated indirect effects”), and 3) evaluate the effect of wolves killing elk on aspen (“density-mediated indirect effects”). My research suggests that wolves indirectly contributed to increased aspen over story recruitment following their reintroduction by helping to reduce the elk population size, but elk response to the risk of wolf predation did not reduce elk foraging in a way that measurably increased aspen recruitment. Additionally, hunter harvest of elk north of the park was twice as important as wolf predation in causing increased aspen recruitment. However, despite wolves and hunters limiting elk abundance, it is still uncommon for young aspen to grow past peak browsing height (120-cm), indicating that many stands remain vulnerable to elk herbivory nearly 30 years after wolf reintroduction. These results highlight that the strength and mechanism of predator effects on plant communities are context-specific. Thus, using predator reintroduction as a tool for ecosystem restoration without considering the many factors that shape trophic cascades may result in different management and conservation outcomes than intended.