Kerry A. Gunther

Distribution of grizzly bears in the Greater Yellowstone Ecosystem in 2004

Abstract The US Fish and Wildlife Service (USFWS) proposed delisting the Yellowstone grizzly bear (Ursus arctos horribilis) in November 2005. Part of that process required knowledge of the most current distribution of the species. Here, we update an earlier estimate of occupied range (1990–2000) with data through 2004. We used kernel estimators to develop distribution maps of occupied habitats based on initial sightings of unduplicated females (n = 481) with cubs of the year, locations of radiomarked bears (n = 170), and spatially unique locations of conflicts, confrontations, and mortalities (n = 1,075). Although each data set was constrained by potential sampling bias, together they provided insight into areas in the Greater Yellowstone Ecosystem (GYE) currently occupied by grizzly bears. The current distribution of 37,258 km2 (1990–2004) extends beyond the distribution map generated with data from 1990–2000 (34,416 km2). Range expansion is particularly evident in parts of the Caribou–Targhee National Forest in Idaho and north of Spanish Peaks on the Gallatin National Forest in Montana.

A proposed lexicon of terms and concepts for human-bear management in North America

Abstract We believe that communication within and among agency personnel in the United States and Canada about the successes and failures of their human–bear (Ursidae) management programs will increase the effectiveness of these programs and of bear research. To communicate more effectively, we suggest agencies clearly define terms and concepts used in human–bear management and use them in a consistent manner. We constructed a human–bear management lexicon of terms and concepts using a modified Delphi method to provide a resource that facilitates more effective communication among human–bear management agencies. Specifically, we defined 40 terms and concepts in human–bear management and suggest definitions based on discussions with 13 other professionals from the United States and Canada. Although new terms and concepts will emerge in the future and definitions will evolve as we learn more about bear behavior and ecology, our purpose is to suggest working definitions for terms and concepts to help guide human–bear management and research activities in North America. Applications or revisions of these definitions may be useful outside of North America.

Changing numbers of spawning cutthroat trout in tributary streams of Yellowstone Lake and estimates of grizzly bears visiting streams from DNA

Abstract Spawning Yellowstone cutthroat trout (Oncorhynchus clarki) provide a source of highly digestible energy for grizzly bears (Ursus arctos) that visit tributary streams to Yellowstone Lake during the spring and early summer. During 1985–87, research documented grizzly bears fishing on 61% of the 124 tributary streams to the lake. Using track measurements, it was estimated that a minimum of 44 grizzly bears fished those streams annually. During 1994, non-native lake trout (Salvelinus namaycush) were discovered in Yellowstone Lake. Lake trout are efficient predators and have the potential to reduce the native cutthroat population and negatively impact terrestrial predators that use cutthroat trout as a food resource. In 1997, we began sampling a subset of streams (n = 25) from areas of Yellowstone Lake surveyed during the previous study to determine if changes in spawner numbers or bear use had occurred. Comparisons of peak numbers and duration suggested a considerable decline between study periods in streams in the West Thumb area of the lake. The apparent decline may be due to predation by lake trout. Indices of bear use also declined on West Thumb area streams. We used DNA from hair collected near spawning streams to estimate the minimum number of bears visiting the vicinity of spawning streams. Seventy-four individual bears were identified from 429 hair samples. The annual number of individuals detected ranged from 15 in 1997 to 33 in 2000. Seventy percent of genotypes identified were represented by more than 1 sample, but only 31% of bears were documented more than 1 year of the study. Sixty-two (84%) bears were only documented in 1 segment of the lake, whereas 12 (16%) were found in 2–3 lake segments. Twenty-seven bears were identified from hair collected at multiple streams. One bear was identified on 6 streams in 2 segments of the lake and during 3 years of the study. We used encounter histories derived from DNA and the Jolly-Seber procedure in Program MARK to produce annual estimates of grizzly bears visiting streams. Approximately 68 grizzly bears visited the vicinity of cutthroat trout spawning streams annually. Thus, approximately 14–21% of grizzly bears in the Greater Yellowstone Ecosystem (GYE) may have used this threatened food resource annually. Yellowstone National Park (YNP) is attempting to control the lake trout population in Yellowstone Lake; our results underscore the importance of that effort to grizzly bears.

Interactions between wolves and female grizzly bears with cubs in Yellowstone National Park

Gray wolves (Canis lupus) were extirpated from Yellowstone National Park (YNP) by the 1920s through predator control actions (Murie 1940,Young and Goldman 1944, Weaver 1978), then reintroduced into the park from 1995 to 1996 to restore ecological integrity and adhere to legal mandates (Bangs and Fritts 1996, Phillips and Smith 1996, Smith et al. 2000). Prior to reintroduction, the potential effects of wolves on the region’s threatened grizzly bear (Ursus arctos) population were evaluated (Servheen and Knight 1993). In areas where wolves and grizzly bears are sympatric, interspecific killing by both species occasionally occurs (Ballard 1980, 1982; Hayes and Baer 1992). Most agonistic interactions between wolves and grizzly bears involve defense of young or competition for carcasses (Murie 1944, 1981; Ballard 1982; Hornbeck and Horejsi 1986; Hayes and Mossop 1987; Kehoe 1995; McNulty et al. 2001). Servheen and Knight (1993) predicted that reintroduced wolves could reduce the frequency of winter-killed and disease-killed ungulates available for bears to scavenge, and that grizzly bears would occasionally usurp wolf-killed ungulate carcasses. Servheen and Knight (1993) hypothesized that interspecific killing and competition for carcasses would have little or no population level effect on either species. As a component of post-reintroduction wolf and grizzly bear monitoring programs, interspecific interactions between the species were recorded. We expected reintroduced wolves to occasionally kill grizzly bears, especially cubs-of-the-year (cubs). We also predicted that adult males, solitary adult females, and female grizzly bears accompanied by yearling or 2-year-old offspring would occasionally usurp wolf-killed ungulates and scavenge the remains. We hypothesized that these cohorts of grizzly bears would be more successful than subadults at usurping wolf-kills. We further hypothesized that due to potential danger to cubs, females with cubs would not attempt to displace wolf packs from their kills. Our monitoring of interspecific interactions between wolves and grizzly bears is ongoing. From wolf reintroduction in 1995 until January of 2003, 96 wolf– grizzly bear interactions have been recorded (Ballard et al. 2003; D. Smith, National Park Service, Yellowstone National Park, Wyoming, USA, unpublished data). Here we report observations of interactions between wolves and female grizzly bears with cubs and evidence of wolf packs killing grizzly bear cubs near carcasses. Due to grizzly bears’ low reproductive rate (Schwartz et al. 2003) and status as a threatened species (USFWS 1993), the effects of wolves on carcass availability and cub survival is an important consideration for wolf reintroduction and grizzly bear conservation efforts. YNP encompasses 891,000 ha in the states of Wyoming, Montana, and Idaho, USA. The park contains a variety of habitats from high elevation alpine to low elevation sagebrush grasslands (Despain 1990). YNP and the surrounding area (Greater Yellowstone Ecosystem, GYE) support an estimated 56,100 elk (Cervus elaphus), 29,500 mule deer (Odocoileus hemionus), 5,800 moose (Alces alces), 3,900 bighorn sheep (Ovis canadensis), 3,600 bison (Bison bison), and smaller numbers of whitetail deer (Odocoileus virginianus), mountain goat (Oreamnos americanus), and pronghorn antelope (Antilocapra americana) (U.S. Fish and Wildlife Service 1994). Large carnivores in the GYE include grizzly bears, American black bears (U. americanus), wolves, and mountain lions (Felis concolor). In 2002, the reintroduced wolf population in the GYE was estimated at 273 wolves in 31 packs (Smith et al. 2003a). More than 90% of the prey killed by wolves in the GYE is elk (Smith et al. 2003b). Other prey species killed by wolves include deer, bison, and moose, but individually none of these prey comprise .2% of GYE wolves’ diet. The GYE grizzly bear population is estimated at 280–610 bears (Eberhardt and Knight 1996). The GYE is unique among areas inhabited by grizzly bears in North America because kerry_gunther@nps.gov doug_smith@nps.gov

Dietary breadth of grizzly bears in the Greater Yellowstone Ecosystem

Abstract Grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem (GYE) are opportunistic omnivores that eat a great diversity of plant and animal species. Changes in climate may affect regional vegetation, hydrology, insects, and fire regimes, likely influencing the abundance, range, and elevational distribution of the plants and animals consumed by GYE grizzly bears. Determining the dietary breadth of grizzly bears is important to document future changes in food resources and how those changes may affect the nutritional ecology of grizzlies. However, no synthesis exists of all foods consumed by grizzly bears in the GYE. We conducted a review of available literature and compiled a list of species consumed by grizzly bears in the GYE. We documented ≥266 species within 200 genera from 4 kingdoms, including 175 plant, 37 invertebrate, 34 mammal, 7 fungi, 7 bird, 4 fish, 1 amphibian, and 1 algae species as well as 1 soil type consumed by grizzly bears. The average energy values of the ungulates (6.8 kcal/g), trout (Oncorhynchus spp., 6.1 kcal/g), and small mammals (4.5 kcal/g) eaten by grizzlies were higher than those of the plants (3.0 kcal/g) and invertebrates (2.7 kcal/g) they consumed. The most frequently detected diet items were graminoids, ants (Formicidae), whitebark pine seeds (Pinus albicaulis), clover (Trifolium spp.), and dandelion (Taraxacum spp.). The most consistently used foods on a temporal basis were graminoids, ants, whitebark pine seeds, clover, elk (Cervus elaphus), thistle (Cirsium spp.), and horsetail (Equisetum spp.). Historically, garbage was a significant diet item for grizzlies until refuse dumps were closed. Use of forbs increased after garbage was no longer readily available. The list of foods we compiled will help managers of grizzly bears and their habitat document future changes in grizzly bear food habits and how bears respond to changing food resources.

Influence of whitebark pine decline on fall habitat use and movements of grizzly bears in the Greater Yellowstone Ecosystem.

When abundant, seeds of the high-elevation whitebark pine (WBP; Pinus albicaulis) are an important fall food for grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem. Rates of bear mortality and bear/human conflicts have been inversely associated with WBP productivity. Recently, mountain pine beetles (Dendroctonus ponderosae) have killed many cone-producing WBP trees. We used fall (15 August–30 September) Global Positioning System locations from 89 bear years to investigate temporal changes in habitat use and movements during 2000–2011. We calculated Manly–Chesson (MC) indices for selectivity of WBP habitat and secure habitat (≥500 m from roads and human developments), determined dates of WBP use, and documented net daily movement distances and activity radii. To evaluate temporal trends, we used regression, model selection, and candidate model sets consisting of annual WBP production, sex, and year. One-third of sampled grizzly bears had fall ranges with little or no mapped WBP habitat. Most other bears (72%) had a MC index above 0.5, indicating selection for WBP habitats. From 2000 to 2011, mean MC index decreased and median date of WBP use shifted about 1 week later. We detected no trends in movement indices over time. Outside of national parks, there was no correlation between the MC indices for WBP habitat and secure habitat, and most bears (78%) selected for secure habitat. Nonetheless, mean MC index for secure habitat decreased over the study period during years of good WBP productivity. The wide diet breadth and foraging plasticity of grizzly bears likely allowed them to adjust to declining WBP. Bears reduced use of WBP stands without increasing movement rates, suggesting they obtained alternative fall foods within their local surroundings. However, the reduction in mortality risk historically associated with use of secure, high-elevation WBP habitat may be diminishing for bears residing in multiple-use areas.

Genetic analysis of individual origins supports isolation of grizzly bears in the Greater Yellowstone Ecosystem

Abstract The Greater Yellowstone Ecosystem (GYE) supports the southernmost of the 2 largest remaining grizzly bear (Ursus arctos) populations in the contiguous United States. Since the mid-1980s, this population has increased in numbers and expanded in range. However, concerns for its long-term genetic health remain because of its presumed continued isolation. To test the power of genetic methods for detecting immigrants, we generated 16-locus microsatellite genotypes for 424 individual grizzly bears sampled in the GYE during 1983–2007. Genotyping success was high (90%) and varied by sample type, with poorest success (40%) for hair collected from mortalities found ≥1 day after death. Years of storage did not affect genotyping success. Observed heterozygosity was 0.60, with a mean of 5.2 alleles/marker. We used factorial correspondence analysis (Program GENETIX) and Bayesian clustering (Program STRUCTURE) to compare 424 GYE genotypes with 601 existing genotypes from grizzly bears sampled in the Northern Continental Divide Ecosystem (NCDE) (FST  =  0.096 between GYE and NCDE). These methods correctly classified all sampled individuals to their population of origin, providing no evidence of natural movement between the GYE and NCDE. Analysis of 500 simulated first-generation crosses suggested that over 95% of such bears would also be detectable using our 16-locus data set. Our approach provides a practical method for detecting immigration in the GYE grizzly population. We discuss estimates for the proportion of the GYE population sampled and prospects for natural immigration into the GYE.

The economics of roadside bear viewing

Viewing bears along roadside habitats is a popular recreational activity in certain national parks throughout the United States. However, safely managing visitors during traffic jams that result from this activity often requires the use of limited park resources. Using unique visitor survey data, this study quantifies economic values associated with roadside bear viewing in Yellowstone National Park, monetary values that could be used to determine whether this continued use of park resources is warranted on economic grounds. Based on visitor expenditure data and results of a contingent visitation question, it is estimated that summer Park visitation would decrease if bears were no longer allowed to stay along roadside habitats, resulting in a loss of 155 jobs in the local economy. Results from a nonmarket valuation survey question indicate that on average, visitors to Yellowstone National Park are willing to pay around

Influence of overnight recreation on grizzly bear movement and behavior in Yellowstone National Park

41 more in Park entrance fees to ensure that bears are allowed to remain along roads within the Park. Generalizing this value to the relevant population of visitors indicates that the economic benefits of allowing this wildlife viewing opportunity to continue could outweigh the costs of using additional resources to effectively manage these traffic jams.

Roadside bear viewing opportunities in Yellowstone National Park: characteristics, trends, and influence of whitebark pine

Abstract Interactions among recreational users and grizzly bears (Ursus arctos) are a continuous challenge for bear managers. Yellowstone National Park, Wyoming, USA uses a system of designated backcountry campsites to manage overnight use and provides bear-resistant food-storage devices for recreational users. Few studies have evaluated how this type of management and recreation influences grizzly bear behavior. We used global positioning system (GPS) data for humans and bears to determine how overnight use influenced grizzly bear movement behavior. We determined times of day campsites were occupied and contrasted grizzly bear locations to random locations near occupied campsites. We conducted a similar analysis ignoring campsite occupancy to assess the utility of including a temporal variable. Grizzly bears were 0.35 times as likely as random locations to be ≤200 m from occupied campsites (95% CI  =  0.19–0.62, P ≤ 0.001). Conversely, when human occupancy was ignored, bears were 2.11 times more likely than random locations to be ≤200 m from campsites (95% CI  =  1.85–2.41, P ≤ 0.001). We conclude that overnight backcountry camping can displace grizzly bears within 200 m of campgrounds. To avoid confounding results, we suggest considering use of a temporal variable in studies of human–bear interactions.