Aspen and Willow Restoration Using Beaver on the Northern Yellowstone Winter Range

Aspen (Populus tremuloides) on the northern Yellowstone winter range has declined over the last half‐century. Beaver (Castor canadensis) were reintroduced in Eagle Creek in 1991 in an attempt to reverse this trend. In 2005, we assessed the efficacy of this project by quantifying the long‐term effects of beaver on aspen stands and the riparian area in this drainage. Between 1990 and 2005, the canopy cover of mature aspen decreased more than 62%, whereas immature aspen cover more than tripled, resulting in a total aspen canopy cover decrease (p < 0.05) from 43 to 25% (a loss of 7.25 ha). Willow canopy cover increased from 10 to 14% during the same period. The impacts of beaver on aspen stands were estimated by comparing vegetative changes among control sites with less than 10% beaver use (n = 5), active beaver sites (n = 6), sites inactive for 1–3 years (n = 7), sites inactive for 4–6 years (n = 4), and sites inactive for 7–11 years (n = 5). Aspen sprout and sapling densities were greater (p = 0.01) on sites which were active and inactive for 1–3 years compared to the other sites. Aspen ramets were not able to grow taller than 2 m on sites without beaver activity for 4–11 years due to ungulate herbivory. Although beaver stimulated the growth of aspen sprouts and saplings, ungulate herbivory prevented successful aspen recovery in the Eagle Creek drainage of the northern Yellowstone winter range 14 years after beaver reintroduction.     

Elk browsing increases aboveground growth of water-stressed willows by modifying plant architecture

In the northern elk wintering range of Yellowstone National Park, USA, wolf (Canis lupus) removal allowed elk (Cervus elaphus) to overbrowse riparian woody plants, leading to the exclusion of beaver (Castor canadensis) and a subsequent water table decline in many small stream valleys. Reduced elk browsing following wolf reintroduction may or may not facilitate willow (Salix sp.) recovery in these areas. To determine if the effect of elk browsing on willow interacts with that of beaver abandonment, we manipulated elk browsing and the water table in a factorial experiment. Under the condition of an ambient (low) water table, elk browsing increased shoot water potential (Ψ _s), photosynthesis per unit leaf area (A), stomatal conductance per unit leaf area (g _s), and aboveground current annual growth (CAG) by 50%. Elk browsing occurred entirely during dormancy and did not affect total plant leaf area (L). Improved water balance, photosynthetic rate, and annual aboveground productivity in browsed willows appeared to be due to morphological changes, such as increased shoot diameter and decreased branching, which typically increase plant hydraulic conductivity. An elevated water table increased Ψ _s, A, g _s, CAG, and L, and eliminated or lessened the positive effect of browsing on CAG for most species. Because low water tables create conditions whereby high willow productivity depends on the morphological effects of annual elk browsing, removing elk browsing in areas of water table decline is unlikely to result in vigorous willow stands. As large willow standing crops are required by beaver, a positive feedback between water-stressed willow and beaver absence may preclude the reestablishment of historical conditions. In areas with low water table, willow restoration may depend on actions to promote the re-establishment of beaver in addition to reducing elk browsing.     

Sampling bias exaggerates a textbook example of a trophic cascade

Understanding trophic cascades in terrestrial wildlife communities is a major challenge because these systems are difficult to sample properly. We show how a tradition of non-random sampling has confounded this understanding in a textbook system (Yellowstone National Park) where carnivore [Canis lupus (wolf)] recovery is associated with a trophic cascade involving changes in herbivore [Cervus canadensis (elk)] behaviour and density that promote plant regeneration. Long-term data indicate a practice of sampling only the tallest young plants overestimated regeneration of overstory aspen (Populus tremuloides) by a factor of 4–7 compared to random sampling because it favoured plants taller than the preferred browsing height of elk and overlooked non-regenerating aspen stands. Random sampling described a trophic cascade, but it was weaker than the one that non-random sampling described. Our findings highlight the critical importance of basic sampling principles (e.g. randomisation) for achieving an accurate understanding of trophic cascades in terrestrial wildlife systems.     

Influence of harvest, climate and wolf predation on Yellowstone elk, 1961-2004

In the period following wolf ( Canis lupus ) reintroduction to Yellowstone National Park (1995–2004), the northern Yellowstone elk ( Cervus elaphus ) herd declined from ∼17 000 to ∼8000 elk (8.1% yr⁻¹). The extent to which wolf predation contributed to this decline is not obvious because the influence of other factors (human harvest and lower than average annual rainfall) on elk dynamics has not been quantified. To assess the contribution of wolf predation to this elk decline, we built and assessed models based on elk-related data prior to wolf reintroduction (1961 to 1995). We then used the best of these models to predict how elk dynamics might have been realized after wolf reintroduction (1995 to 2004) had wolves never been reintroduced. The best performing model predicted 64% of the variance in growth rate and included elk abundance, harvest rate, annual snowfall, and annual precipitation as predictor variables. The best performing models also suggest that harvest may be super-additive. That is, for every one percent increase in harvest rate, elk population growth rate declines by more than one percent. Harvest rate also accounted for ∼47% of the observed variation in elk growth rate. According to the best-performing model, which accounts for harvest rate and climate, the elk population would have been expected to decline by 7.9% per year, on average, between 1995 and 2004. Within the limits of uncertainty, which are not trivial, climate and harvest rate are justified explanations for most of the observed elk decline. To the extent that this is true, we suggest that between 1995 and 2004 wolf predation was primarily compensatory.     

Evidence for top predator control of a grazing ecosystem

The importance of top predators in controlling ecological processes in large, intact ecosystems is unclear. In grasslands that support abundant ungulates, top–down control by predators may be particularly important, because of the tight biogeochemical linkages of ungulate prey with plants and soil microbes. Here, I examined the effects of the recent reintroduction of the gray wolf Canis lupus on ecosystem processes in Yellowstone National Park, where herds of grazing ungulates previously have been shown to stimulate several processes, including soil net nitrogen (N) mineralization. Rates of ungulate grazing intensity and soil net N mineralization were compared before and after wolf reintroduction in grasslands ranging five‐fold in aboveground production. Grazing intensity and grassland net N mineralization declined after wolf reintroduction, a likely partial function of fewer ungulates; wolf predation has been one of several factors implicated in causing the decline in Yellowstone ungulates. In addition, the spatial pattern of grazing and net N mineralization changed after reintroduction. A shift in the spatial patterns of grazer‐associated processes is consistent with a growing body of work indicating that wolves have changed habitat use patterns of ungulates in Yellowstone National Park. These findings suggest widespread wolf effects on ungulate prey, plants, and microbial activity that have spatially reorganized grassland energy and nutrient dynamics in Yellowstone Park.     

Ecosystem Scale Declines in Elk Recruitment and Population Growth with Wolf Colonization: A Before-After-Control-Impact Approach

The reintroduction of wolves (Canis lupus) to Yellowstone provided the unusual opportunity for a quasi-experimental test of the effects of wolf predation on their primary prey (elk – Cervus elaphus) in a system where top-down, bottom-up, and abiotic forces on prey population dynamics were closely and consistently monitored before and after reintroduction. Here, we examined data from 33 years for 12 elk population segments spread across southwestern Montana and northwestern Wyoming in a large scale before-after-control-impact analysis of the effects of wolves on elk recruitment and population dynamics. Recruitment, as measured by the midwinter juvenile∶female ratio, was a strong determinant of elk dynamics, and declined by 35% in elk herds colonized by wolves as annual population growth shifted from increasing to decreasing. Negative effects of population density and winter severity on recruitment, long recognized as important for elk dynamics, were detected in uncolonized elk herds and in wolf-colonized elk herds prior to wolf colonization, but not after wolf colonization. Growing season precipitation and harvest had no detectable effect on recruitment in either wolf treatment or colonization period, although harvest rates of juveniles∶females declined by 37% in wolf-colonized herds. Even if it is assumed that mortality due to predation is completely additive, liberal estimates of wolf predation rates on juvenile elk could explain no more than 52% of the total decline in juvenile∶female ratios in wolf-colonized herds, after accounting for the effects of other limiting factors. Collectively, these long-term, large-scale patterns align well with prior studies that have reported substantial decrease in elk numbers immediately after wolf recolonization, relatively weak additive effects of direct wolf predation on elk survival, and decreased reproduction and recruitment with exposure to predation risk from wolves.     

Winter predation patterns of wolves in Northwestern Wyoming

Wolf (Canis lupus) diets and potential effects on prey have been a prominent subject of interest to wildlife researchers and managers since reintroduction into Yellowstone National Park, Wyoming, USA, in 1995 and 1996. Post-reintroduction, wolves expanded south and recolonized areas in the southern Yellowstone ecosystem. Elk (Cervus elaphus) in this area are supplementally fed during winter (Dec–Mar) at state-managed feedgrounds, resulting in high-density congregations of elk. From December to March 2000–2007, we determined the winter predation patterns of wolves by examining the remains of 289 wolf kills on 3 state-managed feedgrounds and adjacent winter range near Jackson, Wyoming. During winters 2002–2005, we also monitored the movements of radio-collared elk on feedgrounds to describe the response of elk to the presence of wolf kills. Thirty-seven percent (n = 106) of kills were located on elk feedgrounds where elk composition included 49% calves, 42% adult females, 5% adult males, and 5% unknown. Sixty-three percent (n = 183) of kills were located on winter range adjacent to feedgrounds and prey species consisted of 90% elk (38% calves, 35% adult females, 24% adult males, 2% unknown), 9% moose (Alces alces; 13% calves, 69% adult females, 6% adult males, 1% unknown), 1% mule deer (Odocoileus hemionus; 1 fawn, 1 adult female), and 0.5% adult female bison (Bison bison). Mean age of elk killed on feedgrounds was 4.2 years (range = 0–20) and 4.6 years (range = 0–23) on winter range. Calves were selected more than available in most years with female elk killed less than expected. Adult males were killed more than expected in 2005–2007. Eighty-eight percent (n = 198) of the time elk remained on the feedground even when wolves made a kill. Less commonly, elk left the feedground, gathered in larger herds on adjacent feedgrounds absent of wolves, and returned within a few days (6%, n = 13) or left the feedground for another feedground and did not return for the rest of the winter (6%; n = 14). Elk were less likely to leave feedgrounds in the presence of a wolf kill when there were more elk on that feedground. Elk left feedgrounds with greater topography and tree cover (Alkali and Fish Creek) and gathered on the flat, open feedgrounds (Patrol Cabin) more frequently than they left flat, open feedgrounds for feedgrounds with greater topography and tree cover. Our results indicate wolves in our study area primarily preyed on elk and exhibited a strong preference for elk calves. High-density concentrations of elk on feedgrounds will continue to be an attractant for wolves. Although elk leave feedgrounds for reasons other than wolf presence, any displacement of elk from feedgrounds due to wolves will be temporary. State managers have the ability to alter management strategies (e.g., increasing wolf harvest, phasing out elk feeding, increasing the intensity of elk feeding) in an effort to affect predator-prey relationships. © 2019 The Wildlife Society.     

The forgotten prey of an iconic predator: a review of interactions between grey wolves Canis lupus and beavers Castor spp.

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Ungulate Browsing Maintains Shrub Diversity in the Absence of Episodic Disturbance in Seasonally-Arid Conifer Forest

Ungulates exert a strong influence on the composition and diversity of vegetation communities. However, little is known about how ungulate browsing pressure interacts with episodic disturbances such as fire and stand thinning. We assessed shrub responses to variable browsing pressure by cattle and elk in fuels treated (mechanical removal of fuels followed by prescribed burning) and non-fuels treated forest sites in northeastern Oregon, US. Seven treatment paddocks were established at each site; three with cattle exclusion and low, moderate and high elk browsing pressure, three with elk exclusion and low, moderate and high cattle browsing pressure, and one with both cattle and elk exclusion. The height, cover and number of stems of each shrub species were recorded at multiple plots within each paddock at the time of establishment and six years later. Changes in shrub species composition over the six year period were explored using multivariate analyses. Generalized Linear Mixed Models were used to determine the effect of browsing pressure on the change in shrub diversity and evenness. Vegetation composition in un-browsed paddocks changed more strongly and in different trajectories than in browsed paddocks at sites that were not fuels treated. In fuels treated sites, changes in composition were minimal for un-browsed paddocks. Shrub diversity and evenness decreased strongly in un-browsed paddocks relative to paddocks with low, moderate and high browsing pressure at non-fuels treated sites, but not at fuels treated sites. These results suggest that in the combined absence of fire, mechanical thinning and ungulate browsing, shrub diversity is reduced due to increased dominance by certain shrub species which are otherwise suppressed by ungulates and/or fuels removal. Accordingly, ungulate browsing, even at low intensities, can be used to suppress dominant shrub species and maintain diversity in the absence of episodic disturbance events.     

The effects of elk on aspen in the winter range in Rocky Mountain National Park

National parks face problems in managing wild ungulates and their forage resources, including small park sizes and incomplete protection of winter ranges, absence of major predators, and influences from exterior management activities Our study focuses on the effects of elk Cervus elaphus browsing on aspen Populus tremuloides in the elk winter range of Rocky Mountain National Park Elk can prevent successful regeneration of aspen by suckers, and increase the mortality of established trees Here we quantify the effect of elk on aspen in the elk winter range of the park particularly since the adoption of a policy of “natural regulation” in 1968 We collected a stratified-random sample of aspen cohorts in the elk winter range, and also inside two long-term exclosures In each cohort we tallied live and dead trees by diameter, recorded the height and percentage of stem surface covered by bark-stripping, and estimated the date of regeneration of the cohort using increment cores We also tallied live and dead aspen suckers (young trees < 2 5 in tall) by height and the number of live and dead branches on each sucker Aspen outside the exclosures are declining as evidenced by 1) high mortality among established trees, including entire stands that are dead, 2) low density of live suckers, insufficient to regenerate the stand, 3) suckers that are excessively branched and seldom surviving to reach tree height, and 4) almost no cohort regeneration since adoption of natural regulation Exclosures and early photos of the elk winter range suggest these are anomalous conditions Over the last century, aspen cohorts regenerated only when there were fewer than ca 600 elk on the park's elk winter range, far fewer than the present estimated 1600 head Fire suppression, natural succession, fluctuations in beaver populations, climatic fluctuations, and other factors are of no or lesser importance than elk in the regeneration and survival of aspen cohorts on the elk winter range The decline of aspen is a concern when management is focused on the ecosystem scale rather than simply the scale of wildlife and their primary forage resources     

Black bear (<Emphasis Type="Italic">Ursus americanus</Emphasis>) and wolf (<Emphasis Type="Italic">Canis</Emphasis> spp.) summer diet composition and ungulate prey selectivity in Ontario,Canada

Understanding predator-prey dynamics is an important component of management strategy development for wildlife populations that are directly affected by predation. Ungulates often serve as a significant source of prey for many large mammal predators, and patterns of predation are known to influence population dynamics. Although black bear and wolf diets have been investigated extensively, prey preference has been less commonly examined, especially in analyses that take into account age class (i.e., juvenile and adult) of the ungulate prey. We examined black bear (Ursus americanus), wolf (Canis lupus), and hybrids (Canis spp.) prey preference in Ontario based on the availability of three ungulate species—elk (Cervus elaphus), moose (Alces alces), and white-tailed deer (Odocoileus virginianus). We analyzed the presence of prey items in black bear and wolf scats collected over 3 years by examining prey hair cuticular scale patterns. We applied correction factors to frequencies of occurrence of prey items found in predator scat and related diet composition to the availability of ungulates, determined by fecal pellet transect surveys. In addition, non-ungulate diet items were identified to obtain full diet composition profiles. We found that black bear diet consisted of more than 87% vegetation, and they were opportunistic, not selecting for any particular ungulate species in either adult or juvenile age class. Wolf diet was comprised mainly of ungulates (~?73.2%), muskrat (Ondatra zibethica; ~?8.5%), and beaver (Castor canadensis; ~?14.6%), and although moose were at least 1.5 times more abundant then each of the other ungulate prey species in the study area, wolves preferred elk, using moose less than expected. Although we found black bear diet to be opportunistic during the summer, wolves in our study heavily utilized both juvenile and adult ungulates, however, among ungulate species, displayed preference for elk. The preference displayed by wolves provides insight that wildlife managers can use to guide further investigation and assist with the development of strategies to ensure continued elk reintroduction success, and moose and white-tailed deer population sustainability.     

Cougar survival and source-sink structure on Greater Yellowstone's Northern Range

We studied survival and causes of mortality of radiocollared cougars (Puma concolor) on the Greater Yellowstone Northern Range (GYNR) prior to (1987–1994) and after wolf (Canis lupus) reintroduction (1998–2005) and evaluated temporal, spatial, and environmental factors that explain variation in adult, subadult, and kitten survival. Using Program MARK and multimodel inference, we modeled cougar survival based on demographic status, season, and landscape attributes. Our best models for adult and independent subadults indicated that females survived better than males and survival increased with age until cougars reached older ages. Lower elevations and increasing density of roads, particularly in areas open to cougar hunting north of Yellowstone National Park (YNP), increased mortality risks for cougars on the GYNR. Indices of ungulate biomass, cougar and wolf population size, winter severity, rainfall, and individual characteristics such as the presence of dependent young, age class, and use of Park or Wilderness were not important predictors of survival. Kitten survival increased with age, was lower during winter, increased with increasing minimum estimates of elk calf biomass, and increased with increasing density of adult male cougars. Using our best model, we mapped adult cougar survival on the GYNR landscape. Results of receiver operating characteristic (ROC) analysis indicated a good model fit for both female (area under the curve [AUC] = 0.81, 95%CI = 0.70–0.92, n = 35 locations) and male cougars (AUC = 0.84, 95%CI = 0.74–0.94, n = 49 locations) relative to hunter harvest locations in our study area. Using minimum estimates of survival necessary to sustain the study population, we developed a source-sink surface and we identify several measures that resource management agencies can take to enhance cougar population management based on a source-sink strategy. © 2011 The Wildlife Society.     

What best explains vigilance in elk: characteristics of prey, predators, or the environment?

We quantified the vigilance levels of elk (Cervus elaphus) preyedon by wolves (Canis lupus) in Yellowstone National Park, betweenJanuary and May in 2005 and 2006, and used Akaike's informationcriterion to compare a set of 38 regression models for vigilancelevels. These models combined up to 9 predictor variables of3 types: characteristics of the prey group (herd size and composition),characteristics of the predator (wolf pack size, distance away,and the presence/absence of a kill), and characteristics ofthe local environment (distance to woodland edges, snow depth,and snow cover). The set of models spanned a range of complexityfrom simple univariate models to complex combinations with upto 3 variables of each type. Complex models incorporating thecharacteristics of the wolf pack, the structure of the elk herd,and the environmental conditions had higher information contentthan simple models. Although univariate models of vigilancedetect significant relationships, they have low informationcontent relative to multivariate models. These results showthat elk assesses factors of several types when assessing riskand deciding how much time to allocate to vigilance. In particular,we found that all well-supported models of vigilance includedseveral "prey" variables and several "predator" variables. Thisresult highlights the need to consider information about predatorswhen trying to explain variation in vigilance levels in prey.     

Effect of selective logging on tree and understory regeneration in miombo woodland in western Tanzania

Tree regeneration and understory response to selective cutting of Pterocarpus angolensis DC, and Sterculia quinqueloba Sim, was measured outside Katavi National Park, Tanzania. Contrary to expectations, a selective harvest had no effect on tree recruitment for either species in this miombo woodland. In unlogged plots along transects taken at increasing distance from a main road, the stand densities of young trees declined away from the road. Since anthropogenic activity along the road results in a high fire frequency but low native ungulate densities, the increased rate of tree regeneration near the road may result from either reduced grazing and browsing pressure by native ungulates, or reduced fire intensity and severity along roads. These effects appear to mask any possible effect of selective harvest on tree recruitment. Currently, there is no evidence of compensatory recruitment of trees of any species into the canopy to replace logged trees and this will lead to a gradual thinning in overstory stand density.     

Climate change intensification of herbivore impacts on tree recruitment

Altered species interactions are difficult to predict and yet may drive the response of ecological communities to climate change. We show that declining snowpack strengthens the impacts of a generalist herbivore, elk (Cervus elaphus), on a common tree species. Thick snowpack substantially reduces elk visitation to sites; aspen (Populus tremuloides) shoots in these areas experience lower browsing rates, higher survival and enhanced recruitment. Aspen inside herbivore exclosures have greatly increased recruitment, particularly at sites with thick snowpack. We suggest that long-term decreases in snowpack could help explain a widespread decline of aspen through previously unconsidered relationships. More generally, reduced snowpack across the Rocky Mountains, combined with rising elk populations, may remove the conditions needed for recruitment of this ecologically important tree species. These results highlight that herbivore behavioural responses to altered abiotic conditions are critical determinants of plant persistence. Predictions of climate change impacts must not overlook the crucial importance of species interactions.     

Genetic variation in postfire aspen seedlings in yellowstone national park

A rare episode of regeneration of aspen (Populus tremuloides Michx.) by seeds occurred in Yellowstone National Park (YNP), Wyoming, USA, following extensive fires that occurred in 1988. In 1997, we sampled 410 aspen seedlings from 23 local populations distributed widely across YNP to determine how genetic diversity varies with elevation, substrate, plant competition, ungulate browsing, and geographical location. We employed 132 randomly amplified polymorphic DNA (RAPD) markers based on six primers to show genetic relationships within and among the postfire aspen seedling populations. Measures of genetic variation, including estimates of percentage polymorphic loci, expected heterozygosity, and Nei's FST, indicated that most of the variation occurred within rather than among local populations. There was no indication of geographical differentiation among sampled populations based on hierarchal estimates of Nei's FST, neighbour-joining, or correlations between genetic distance and geographical distance. Even genetically distant populations shared nearly 90% of the same markers. Within plots, the amount of genetic variation decreased slightly in response to increased percentage vegetative cover, mean seedling basal diameter, and mean seedling height. Geological substrate, density of lodgepole pine (Pinus contorta var. latifolia Dougl.) seedlings, browsing intensity, and elevation were not significantly related to levels of genetic variation within the seedling plots. These data suggest that genetic variation and geographical structure among seedling populations may occur over time as the transition from seedling-dominated stands to clone-dominated stands occurs.     

Responses of wolf feeding habits after adverse climatic events in central-western Belarus

Many studies have investigated the ecology of wolf populations of Eurasia, showing that although wolves are mostly opportunistic in seeking meso-large enough mammalian prey, they can also be selective, depending on local availability of prey and their population biomass. Yet prey preferences of the wolf have been poorly evaluated in situations of complex predator/prey systems because such ecological situations are extremely rare in Europe. In particular, the role of beaver is poorly known due to the extreme decline in its range and population over the last few centuries.We conducted a 15-year study (1999–2014) of wolf Canis lupus diet in the Naliboki forest of central-western Belarus to determine the dietary responses of the wolf population in a context of a rich prey supply (beaver 650 inds/100 km², elk 47 inds/100 km², red deer 98 inds/100 km, roe deer 398 inds/100 km², wild boar 234 inds/100 km²). The bison, although present, is not preyed on. We compared the seasonal and annual diet variations of both wolf adults and pups, by scat analysis and hair identification. In winter 2012–2013, the winter was quite harsh with a long period of snow, which severely affected the roe deer and wild boar populations. Five severe summer droughts also occurred (1999, 2001, 2002, 2004 and 2013), greatly decreasing the water level in rivers and canals. We took advantage of these stressful events to evaluate the diet responses of the wolves.In “normal” years, we identified 11 food categories, essentially beaver and medium-sized ungulates (66%), and large ungulates to a lesser extent (9% in summer, 20% in winter). The adults were found to selectively supply pups with beaver, probably because of its easy transportability. Beaver consumption also increased during summer droughts when water levels were very low. After the harsh winter of 2012–2013, which was followed by a sharp decline in medium-sized prey, we observed a shift in the winter diet breadth of the wolves towards greater consumption of both large wild ungulates and small carnivores. We concluded that:1. Beaver is a functional element in wolf ecology, as a primary food for adults and pups;2. A large range of available prey species is important to maintaining a viable wolf population in cases of extreme climatic events.     

Disentangling herbivore impacts on Populus tremuloides: a comparison of native ungulates and cattle in Canada’s Aspen Parkland

Ungulates impact woody species’ growth and abundance but little is understood about the comparative impacts of different ungulate species on forest expansion in savanna environments. Replacement of native herbivore guilds with livestock [i.e., beef cattle (Bos taurus)] has been hypothesized as a factor facilitating trembling aspen (Populus tremuloides Michx.) encroachment into grasslands of the Northern Great Plains. We used a controlled herbivory study in the Parklands of western Canada to compare the impact of native ungulates and cattle on aspen saplings. Native ungulate treatments included a mixed species guild and sequences of herbivory by different ungulates [bison (Bison bison subsp. bison), elk (Cervus elaphus) then deer (Odocoileus hemionus); or deer, elk, then bison]. Herbivory treatments were replicated in three pastures, within which sets of 40 marked aspen saplings (<1.8 m) were tracked along permanent transects at 2-week intervals, and compared to a non-grazed aspen stand. Stems were assessed for mortality and incremental damage (herbivory, leader breakage, stem abrasion and trampling). Final mortality was greater with exposure to any type of herbivore, but remained similar between ungulate treatments. However, among all treatments, the growth of aspen was highest with exposure only to cattle. Herbivory of aspen was attributed primarily to elk within the native ungulate treatments, with other forms of physical damage, and ultimately sapling mortality, associated with exposure to bison. Overall, these results indicate that native ungulates, specifically elk and bison, have more negative impacts on aspen saplings and provide evidence that native and domestic ungulates can have different functional effects on woody plant dynamics in savanna ecosystems.     

Planning and Implementing a Reintroduction of Wolves to Yellowstone National Park and Central Idaho

The Northern Rocky Mountain Wolf Recovery plan proposed reintroduction of Canis lupus (gray wolf) to Yellowstone National Park and central Idaho as part of a wolf restoration plan for the northern Rocky Mountains of the United States. Strong opposition from some factions within the region forestalled the action for two decades. An environmental impact statement, conducted in 1992–1994 with extensive public input, culminated in a proposal to reintroduce wolves designated as “non-essential—experimental” under Section 10 (j) of the federal Endangered Species Act. This approach, approved by the Secretary of the Interior in 1994, provided for wolf restoration while allowing management flexibility to deal with concerns of the local public. A reintroduction plan was developed in the summer and fall of 1994. Acquiring, holding, transporting, and releasing suitable wolves for reintroduction presented a myriad of technical and logistical challenges that required effective planning and coordination by all participants. In January 1995, 29 wolves were captured in Alberta and transported to Yellowstone National Park (14) and central Idaho (15). Idaho wolves were freed immediately upon arrival; Yellowstone wolves (three family groups) were held in acclimation pens in the park until late March. Most Idaho wolves traveled extensively within the area intended for them, averaging 82 km net distance away from release sites after 5 months (range = 30–220 km), and three male-female pairs formed by July. After 5 months in the wild, at least 13 of 15 Idaho-released wolves were alive within the intended area, as were 13 of 14 Yellowstone wolves; one wolf was known to have been illegally killed in each area. No livestock were killed. Wolves released into Yellowstone Park continued to live as packs, stayed closer to their release sites (x = 22 km at end of June), and settled into home ranges; two packs produced a total of nine pups. The progress of the reintroduction program in its first year far exceeded expectations. Reintroductions of about 15 wolves to each area for 2–4 more years are scheduled, but the project may be shortened because of early successes. Future reintroduction planners can expect sociocultural issues to pervade the effort, but they can be optimistic that, from a biological standpoint, reintroduction of wolves has strong potential as a restoration technique.     

Regeneration response and seedling bank dynamics on a Dendroctonus rufipennis‐killed Picea engelmannii landscape

Question: How does regeneration response to a host‐specific, high‐severity, infrequent Dendroctonus rufipennis outbreak differ from our conceptualization of high‐severity, infrequent/low‐severity, frequent disturbance regimes in Picea engelmannii‐Abies lasiocarpa communities? Location: Southern Utah, USA. Methods: One hundred and seven plots across a high‐elevation P. engelmannii forest were sampled to reconstruct pre‐outbreak overstory and seedling bank densities, and calculate their associated metrics of diversity. Decade of establishment by seedling bank trees indicated “chronic” and “pulse” regenerators. Results: The post‐outbreak overstory and seedling bank were dominated by A. lasiocarpa. Although Pinus flexilis, Pinus ponderosa, Picea pungens, and Psuedotsuga menziesii were present in the overstory, they were virtually absent in the seedling bank. Seedling bank recruitment of A. lasiocarpa and P. engelmannii has been occurring chronically for at least the last ～205 and ～152 years, respectively. A pulse response of seedling bank Populus tremuloides was apparent; however, results were complicated by intense ungulate browsing. Conclusions: Despite some similarities to the high‐severity, infrequent/low‐severity, frequent conceptualization of regeneration response to disturbance, the high‐severity D. rufipennis outbreak is best described by explicitly considering host specificity and severity. Although, the outbreak simultaneously promoted both a pulse of P. tremuloides and a release of chronically regenerated A. lasiocarpa, the P. tremuloides response was generally masked by ungulate browsing, and the regeneration response came overwhelmingly from the A. lasiocarpa seedling bank. In this landscape, once dominated by P. engelmannii, the chronically regenerating seedling bank, typically thought to take advantage of canopy gaps associated with low‐severity disturbances, is poised to dominate forest reorganization in response to the host‐specific outbreak.