The role of predation is of major importance to
conservationists as the ranges of large carnivores continue
to collapse around the world. In North America, for example,
the gray wolf (Canis lupus) and the grizzly bear (Ursus
arctos) have respectively lost 53% and 42% of their historic
range, with nearly complete extirpation in the contiguous 48
United States (Laliberte and Ripple 2004). Reintroduction of
these and other large carnivores is the subject of intense scientific
and political debate, as growing evidence points to the
importance of conserving these animals because they have cascading
effects on lower trophic levels. Recent research has
shown how reintroduced predators such as wolves can influence
herbivore prey communities (ungulates) through direct
predation, provide a year-round source of food for
scavengers, and reduce populations of mesocarnivores such
as coyotes (Canis latrans) (Smith et al. 2003). In addition, vegetation
communities can be profoundly altered by herbivores
when top predators are removed from ecosystems, as a
result of effects that cascade through successively lower trophic
levels (Estes et al. 2001). The absence of highly interactive carnivore
species such as wolves can thus lead to simplified or
degraded ecosystems (Soulé et al. 2003). A similar point was
made more than 50 years ago by Aldo Leopold (1949):“Since
then I have lived to see state after state extirpate its wolves....
I have seen every edible bush and seedling browsed, first to
anemic desuetude, and then to death” (p. 139).
Increased ungulate herbivory can affect vegetation structure,
succession, productivity, species composition, and diversity
as well as habitat quality for other fauna. Although the
topic remains contentious, a substantial body of evidence indicates
that predation by top carnivores is pivotal in the
maintenance of biodiversity. Most studies of these carnivores
have emphasized their lethal effects (Terborgh et al. 1999).
Here our focus is on how nonlethal consequences of predation
(predation risk) affect the structure and function of
ecosystems. The objectives of this article are twofold: (1) to
provide a brief synthesis of potential ecosystem responses to
predation risk in a three-level trophic cascade involving large
carnivores (primarily wolves), ungulates, and vegetation; and
(2) to present research results that center on wolves, elk
(Cervus elaphus), and woody browse species in the northern
range of Yellowstone National Park (YNP).