Are wolves saving Yellowstone's aspen? A landscape-level test of a behaviorally mediated trophic cascade: comment.

By the early 1900s, Euro-Americans had extirpated gray wolves (Canis lupus) from most of the contiguous United States. Yellowstone National Park was not immune to wolf persecution and by the mid-1920s they were gone. After seven decades of absence in the park, gray wolves were reintroduced in 1995–1996, again completing the large predator guild (Smith et al. 2003). Yellowstone’s ‘‘experiment in time’’ thus provides a rare opportunity for studying potential cascading effects associated with the extirpation and subsequent reintroduction of an apex predator. Wolves represent a particularly important predator of large mammalian prey in northern hemisphere ecosystems by virtue of their group hunting and year-round activity (Peterson et al. 2003) and can have broad top-down effects on the structure and functioning of these systems (Miller et al. 2001, Soulé et al. 2003, Ray et al. 2005). If a tri-trophic cascade involving wolves–elk (Cervus elaphus)–plants is again underway in northern Yellowstone, theory would suggest two primary mechanisms: (1) density mediation through prey mortality and (2) trait mediation involving changes in prey vigilance, habitat use, and other behaviors (Brown et al. 1999, Berger 2010). Both predator-caused reductions in prey numbers and fear responses they elicit in prey can lead to cascading trophic-level effects across a wide range of biomes (Beschta and Ripple 2009, Laundré et al. 2010, Terborgh and Estes 2010). Thus, the occurrence of a trophic cascade could have important implications not only to the future structure and functioning of northern Yellowstone’s ecosystems but also for other portions of the western United States where wolves have been reintroduced, are expanding their range, or remain absent. However, attempting to identify the occurrence of a trophic cascade in systems with large mammalian predators, as well as the relative importance of density and behavioral mediation, represents a continuing scientific challenge. In Yellowstone today, there is an ongoing effort by various researchers to evaluate ecosystem processes in the park’s two northern ungulate winter ranges: (1) the ‘‘Northern Range’’ along the northern edge of the park (NRC 2002, Barmore 2003) and (2) the ‘‘Upper Gallatin Winter Range’’ along the northwestern corner of the park (Ripple and Beschta 2004b). Previous studies in northern Yellowstone have generally found that elk, in the absence of wolves, caused a decrease in aspen (Populus tremuloides) recruitment (i.e., the growth of seedlings or root sprouts above the browse level of elk). Within this context, Kauffman et al. (2010) initiated a study to provide additional understanding of factors such as elk density, elk behavior, and climate upon historical and contemporary patterns of aspen recruitment in the park’s Northern Range. Like previous studies, Kauffman et al. (2010) concluded that, irrespective of historical climatic conditions, elk have had a major impact on long-term aspen communities after the extirpation of wolves. But, unlike other studies that have seen improvement in the growth or recruitment of young aspen and other browse species in recent years, Kauffman et al. (2010) concluded in their Abstract: ‘‘. . . our estimates of relative survivorship of young browsable aspen indicate that aspen are not currently recovering in Yellowstone, even in the presence of a large wolf population.’’ In the interest of clarifying the potential role of wolves on woody plant community dynamics in Yellowstone’s northern winter ranges, we offer several counterpoints to the conclusions of Kauffman et al. (2010). We do so by readdressing several tasks identified in their Introduction (p. 2744): (1) the history of aspen recruitment failure, (2) contemporary aspen recruitment, and (3) aspen recruitment and predation risk. Task 1 covers the period when wolves were absent from Yellowstone and tasks 2 and 3 focus on the period when wolves were again present. We also include some closing comments regarding trophic cascades and ecosystem recovery. 1. History of aspen recruitment failure.—Although records of wolf and elk populations in northern Yellowstone are fragmentary for the early 1900s, the Northern Range elk population averaged ;10 900 animals (7.3 elk/km; Fig. 1A) as the last wolves were being removed in the mid 1920s. Soon thereafter increased browsing by elk of aspen and other woody species was noted in northern Yellowstone’s winter ranges (e.g., Rush 1932, Lovaas 1970). In an attempt to reduce the effects this large herbivore was having on vegetation, soils, and wildlife habitat in the Northern Manuscript received 13 January 2011; revised 10 June 2011; accepted 20 June 2011.  C. C. Wilmers. 1 Department of Forest Ecosystems and Society, Oregon State University, Corvallis, Oregon 97331 USA. 2 E-mail: Robert.Beschta@oregonstate.edu