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Recovering Riparian Plant Communities with Wolves in Northern Yellowstone

Recovering Riparian Plant Communities with Wolves in Northern Yellowstone

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Recovering Riparian Plant Communities withWolves in Northern Yellowstone, U.S.A.
Robert L. Beschta
1,2
and William J. Ripple
1
Abstract
Gray wolves (
Canis lupus
) were extirpated from Yellow-stone National Park in the 1920s. The ensuing seven dec-ades marked a period when wild ungulates, principallyElk (
Cervus elaphus
), extensively used woody browsespecies in the upper Gallatin and northern winter ranges,thus limiting the capability of establishing plants to growmore than 100 cm in height. Following the reintroductionof wolves in the mid-1990s, we evaluated patterns of woody browse species recovery within riparian areas of these winter ranges. Measurements indicated that cotton-wood (
 Populus
spp.) recruitment (growth of seedlings/ sprouts into tall saplings and trees) was occurring for thefirst time in several decades. A spatially patchy increasein the heights of young willow (
 Salix 
sp.) and cotton-wood in the upper Gallatin and northern winter ranges,respectively, was also found within riparian transectscomprising nearly 20 km in total length. Within sometransects, heights of woody species have begun to exceed200 cm (the approximate upper browse level of Elk). Re-sults are consistent with the reestablishment of a tri-leveltrophic cascade involving wolves, ungulates, and riparianvegetation. We additionally present conceptual modelsof vegetation recovery, illustrating differences in plantheight responses to behaviorally and density-mediatedtrophic cascades. Northern Yellowstone’s ‘‘experiment intime,’’ whereby wolves were removed and then reintro-duced, provides new insights regarding how top preda-tors can influence the structure and biodiversity of terrestrial ecosystems. Restoration ecologists and policy-makers should consider the potential benefits of largepredators as an option for helping restore degraded eco-systems.
Key words:
restoration, riparian ecosystems, trophiccascades, ungulates Yellowstone, wolves.
Introduction
Ecological restoration can include both active and passiveapproaches for reestablishing historical plant and animalcommunities, as well as the renewal of ecosystem func-tions necessary to sustain those communities (Kauffmanet al. 1997). Active restoration, for example, might rangefrom minimizing human interventions to the reintroduc-tion of previously extirpated species, thereby allowing nat-ural disturbances (passive restoration) to reorganize andrestore an ecosystem over time. Inherent in any restora-tion effort is an adequate understanding of how ecosys-tems historically functioned (reference conditions) andhow they have changed over time (Ripple & Beschta2005). Such information is usually crucial for assessing themagnitude of historical and contemporary human effects,as well as establishing recovery goals.Disturbances alter the function, species composition,and population structure of ecosystems. Wildfires andlarge floods represent ‘‘pulse’’ disturbances (Bender et al.1984) because individual events can reset plant communi-ties, often with extended time periods between occurren-ces. In contrast, annual herbivory by ungulates createsa ‘‘press’’ disturbance regime. Although the annual incre-mental cropping of vegetation may appear to be less sig-nificant than the effects of pulse disturbances, if suchherbivory comprises a sufficiently large percentage of cur-rent annual growth and is persistent (occurs year afteryear), these disturbances can have major impacts on plantcommunities and ecosystems (Ohmart 1996; Belsky et al.1999; Barmore 2003). Wolf predation throughout the yearalso represents a press disturbance and its effects on con-sumers (ungulates) may be transmitted to plant communi-ties as a trophic cascade (Estes et al. 2001). Over time,and in conjunction with other predators, this wide-rangingcarnivore may influence not only the size of prey popula-tions (density mediation) but also their patterns of herbiv-ory (behavioral mediation).During the 1800s and early 1900s, Euro-Americans erad-icated Gray wolves (
Canis lupus
) and Grizzly bears (
Ursusarctos
) from some 95 to 99% of their original range in theconterminous United States, thus allowing (1) the relaxa-tion of predation as a selective force and (2) the irruptionof herbivore populations (Berger 1999). Wolves, a largepredator capable of affecting a variety of animal species(Soule´ et al. 2005), were extirpated from northern Yellow-stone’s winter ranges in the early 1900s. During subsequentdecades, the effects of wild ungulate herbivory, principallyElk (
Cervus elaphus
), upon riparian plant communitiesremained a controversial management and scientific issue
1
College of Forestry, Oregon State University, Corvallis, OR 97331, U.S.A.
2
Address correspondence to R. L. Beschta, email robert.beschta@oregonstate.edu
Ó
 2008 Society for Ecological Restoration International 
doi: 10.1111/j.1526-100X.2008.00450.x
380
Restoration Ecology Vol. 18, No. 3, pp. 380–389
MAY 2010
 
(e.g., Chadde and Kay 1991; NRC 2002; Wagner et al.2006). However, most of this debate occurred without con-sidering the potential ecological role of wolves.The general thrust of this article embraces a majorpremise—following the extirpation of Gray wolves (anapex predator), riparian plant communities within thewinter ranges of northern Yellowstone were severelyimpacted by ungulate herbivory over a period of sevendecades (mid-1920s to mid-1990s). This premise is brieflysummarized in our description of study areas below. How-ever, the primary goal of this study was to assess theextent that trophic cascades following the reintroductionof wolves in the mid-1990s may have initiated recovery of these plant communities. If removal, by humans, of a wolf-dominated disturbance regime allowed ungulateherbivory to significantly impact winter range riparianvegetation over multiple decades, then reinstatement of that disturbance regime may represent a prerequisite forinitiating recovery of plant communities.
Study Areas
Roughly 80 km apart along the northern portion of Yel-lowstone National Park (YNP), the ‘‘upper Gallatin’’ and‘‘northern’’ winter ranges comprise approximately 200 and1,500 km
2
, respectively, of lower mountain slopes and val-ley bottom terrain in the northern Rocky Mountains(Barmore 2003; Ripple & Beschta 2004
b
). Most of theupper Gallatin winter range occurs outside the park,whereas approximately two-thirds of the northern winterrange occurs inside the park. Summers are short and cool,whereas winters are typically long and cold. Winter snow-packs are usually shallow along valley bottoms (elevationapproximately 2,000 m) and south-facing slopes, withincreasing snowpack depths at higher elevations.Low-elevation portions of these winter ranges encom-pass shrub-steppe plant communities dominated by Bigsagebrush (
 Atremisia tridentata
), with patches of inter-mixed Douglas fir (
Pseudotsuga menziesii
), Lodgepolepine (
Pinus contorta
), and Aspen (
Populus tremuloides
).Riparian areas along the upper Gallatin River are domi-nated by willows (
Salix
spp.), whereas cottonwoods (
Pop-ulus
spp.), willows, and Aspen variously occur withinriparian areas of Rose Creek, Soda Butte Creek, and theLamar River in the northern range.In addition to Gray wolves, Yellowstone’s large preda-tors include Grizzly bear, Black bear (
Ursus americanus
),and Cougar (
Felis concolor 
). Elk frequent both winterranges along with smaller populations of Mule deer (
Odo-coileus hemionus
), White-tailed deer (
Od. virginianus
),Moose (
 Alces alces
), Pronghorn (
 Antilocapra americana
),and Bighorn sheep (
Ovis canadensis
). Although Bison(
Bison bison
) have been historically absent from theupper Gallatin, a herd has been present in the northernrange throughout the 1900s. In recent years, the size of this herd has been increasing.Although YNP was established in 1872, market huntingof ungulates and predators continued until 1886 when theU.S. Army was assigned responsibility for protecting thepark’s wildlife (YNP 1997). However, persecution of wolves continued inside and outside the park through theearly 1900s. With the extirpation of wolves in the mid-1920s (Schullery & Whittlesey 1992), northern Yellow-stone’s Elk had unimpeded access to winter range plantcommunities, and browsing impacts to vegetation weresoon observed (Callahan 1923). In the upper Gallatin win-ter range, Elk numbers generally decreased in subsequentdecades due to annual harvest (by hunting) of animals thatmigrated outside the park as well as periodic die-offs asso-ciated with a degraded winter range and the occurrence of severe winters (Peek et al. 1967). Annual hunting of Elkoutside the park’s portion of the northern range alsooccurred. In addition, in the 1920s, the Park Service begancapturing Elk within the northern range and shippingthem to other locations, both to assist in the reestablish-ment of Elk herds in various parts of the western UnitedStates and Canada and to reduce browsing impacts. Even-tually, the agency resorted to killing Elk within the parkin an attempt to control their impacts upon vegetationand soils.Even though reduced Elk numbers occurred during thedecades following wolf extirpation in the upper Gallatinwinter range, Lovaas (1970) indicated that upland plantcommunities were heavily grazed, Aspen ramets (rootsprouts) were no longer able to grow above the browselevel of Elk, and hillslope erosion became a concern.Riparian willow communities were also heavily browsed(Patten 1968; Ripple & Beschta 2004
b
), eventually leadingto unstable channels and a hydrologically disconnectedfloodplain (Beschta & Ripple 2006).Following the loss of wolves in the northern range, pal-atable woody species were increasingly unable to establishand grow above the browse level of Elk (NRC 2002;Barmore 2003). Similar to the upper Gallatin, once plantcommunities had been degraded, even reduced numbersof Elk were capable of continuing to suppress vegetativegrowth. The Park Service’s Elk culling program in thenorthern range was terminated in 1968, and within twodecades, its Elk population of approximately 4,000 hadirrupted to nearly 19,000 animals. This large Elk popula-tion only served to increase the severity of impacts toplant communities and to curtail the recruitment (i.e.,growth of seedlings/sprouts into tall saplings or trees) of woody browse species (Chadde & Kay 1991; Ripple &Larsen 2000; Beschta 2005). Northern range vegetationstudies that spanned over five decades (1935–1989) foundthat average heights of young willow, Aspen, and otherwoody browse species outside ungulate exclosures neverexceeded 83 cm (
n
¼
81 species-years), whereas the samespecies inside exclosures rapidly increased in height(Chadde & Kay 1991; Singer 1996; Barmore 2003). Fruitproduction of heavily browsed berry-producing shrubsoutside exclosures was severely reduced or eliminated
Riparian Recovery with Wolves
MAY 2010
Restoration Ecology
381
 
(Kay 1995). With the deterioration of woody plant com-munities, Beaver (
Castor canadensis
) underwent decline,and by the early 1950s, only scattered colonies remained(Jonas 1955).During March 1995, 14 wolves were released into thepark with an additional 17 wolves the following winter.They were soon breeding and establishing packs acrossnorthern Yellowstone.
Methods
We summarized recent population estimates for wolves,Elk, and Bison in the upper Gallatin and northern winterranges. In the northern range study area, we determinedlong-term patterns of cottonwood recruitment. A diame-ter at breast height (dbh) of 5 cm was chosen as a thresholddiameter for measurement as it was assumed that cotton-woods of this diameter, and larger, would have attaineda height well above the browse level of Elk. We thussearched for and measured the diameters of all cotton-wood trees more than or equal to 5 cm in dbh during latesummer of 2001 with reinventories in 2002, 2003, 2004,and 2006. The establishment date of each inventoried cot-tonwood was estimated from tree age versus dbh relation-ships to establish the age structure of cottonwoods in thenorthern range study area (Beschta 2005). From thoseresults, we regressed tree frequency versus establishmentdate for the decades when wolves were present (pre-1920s). This exponential relationship provided a basis of comparison regarding cottonwood tree recruitment duringrecent decades.We further characterized the general recovery status of riparian plant communities by measuring heights of youngBooth willow (
Salix boothii
) in the upper Gallatin andyoung cottonwoods (
Populus angustifolia
and
Po. tricho-carpa
) in northern range study areas. We selected thesewoody species because they commonly occurred in theirrespective riparian areas, are highly palatable, andappeared to reflect the general pattern of height growthfor other woody browse species. Our sampling of plantheights in riparian areas was undertaken to providea ‘‘leading edge’’ indication of ecosystem recovery in thedecade following wolf reintroduction.In the upper Gallatin study area, we initially measuredBooth willow heights within a 3-km riparian transect alongthe Gallatin River in August 2003. This transect occurredalong a predation risk gradient (low predation risk at theupstream end of the transect, with increasing risk ina downstream direction; Ripple & Beschta 2004
b
). Withineach 100-m segment of the transect (30 segments in total),the three tallest Booth willow plants were selected. Weundertook plant architecture measurements (Keigley1998; Beschta & Ripple 2007) on the tallest stem (leader)of each selected plant to evaluate its browsing history andspringtime height (after winter browsing) for the previous2 years. In August 2006, we measured springtime and late-summer (after current annual growth had occurred) leaderheights of the three tallest Booth willows within each 100-m segment of the same 3-km transect. We used linearregression to illustrate trends in springtime willow heights,by year, along the 3-km transect. Using the linear relation-ship of 2001 as a reference, we used multiple regression totest for significant (
 p
0.05) slope and intercept differen-ces in subsequent years.In the northern range study area, we measured late-summer heights of young cottonwoods along ripariantransects paralleling major streams, including:(1) ‘‘Lower Lamar’’—a 9.9-km transect (measured inAugust 2002 and 2006) along the Lamar River down-stream from the Soda Butte Creek confluence.(2) ‘‘Rose Creek’’—a 2.2-km transect (measured inAugust 2002 and 2006) encompassing three distribu-taries of Rose Creek that flow across the alluvial fanat the Lamar Ranger Station (Buffalo Ranch).(3) ‘Soda Butte Creek’’—a 2.2-km transect (measured inAugust 2006) along Soda Butte Creek starting approx-imately 4-km upstream of its confluence with theLamar River.(4) ‘‘Upper Lamar’’—a 1.8-km transect (measured inAugust 2003 and 2006) along the Lamar Riverupstream of the Soda Butte Creek confluence.Within each 50-m segment of a transect, we searchedfor the three tallest cottonwood seedlings or root sproutsmore than or equal to 20 cm tall and averaged theirheights. For evaluating height changes over time, we ex-pressed segment heights of a transect as a percentile andcompared any change in median height between samplingperiods. Height measurements of woody browse species inboth the upper Gallatin and the northern winter ranges, inconjunction with other Yellowstone studies and thebroader literature, allowed us to develop conceptual mod-els that demonstrate differences in plant communityresponses from behaviorally mediated (risk based) anddensity-mediated (mortality based) components of a tro-phic cascade.
Results
Wolves and Ungulates
Following the 1995–1996 reintroduction of wolves, theirnumbers began to increase. Over the past 5 years, north-ern YNP populations have ranged between 54 and 106wolves (Smith et al. 2006). Only two Elk population esti-mates, ranging from 1,048 to 3,028 animals, were availablein the upper Gallatin study area for the 10 years (1987–1996) leading up to wolf reintroduction. After wolf rein-troduction, census estimates indicate that approximately1,000 Elk have frequented this winter range (Fig.1a). Forthe northern range, Elk populations peaked at approxi-mately 19,000 animals in the decade prior to wolf reintro-duction. Elk densities after wolf reintroduction havetrended downward, whereas Bison densities have recently
Riparian Recovery with Wolves
382
Restoration Ecology
MAY 2010

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