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Symposium 9: Rocky Mountain futures: preserving, utilizing, and sustaining Rocky Mountain ecosystems

In 2002 we published Rocky Mountain Futures, an Ecological Perspective (Island Press) to examine the cumulative ecological effects of human activity in the Rocky Mountains. We concluded that multiple local activities concerning land use, hydrologic manipulation, and resource extraction have altered ecosystems, although there were examples where the “tyranny of small decisions” worked in a positive way toward more sustainable coupled human/environment interactions. Superimposed on local change was climate change, atmospheric deposition of nitrogen and other pollutants, regional population growth, and some national management policies such as fire suppression.

Bulletin of the Ecological Society of America↗

Bromus tectorum invasion alters nitrogen dynamics in an undisturbed arid grassland ecosystem

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C 3 and C 4 ). The response of these communities was compared to an invaded/ disturbed grassland. The invaded/disturbed communities had higher surface NH 4 + input in spring, whereas there were no differences for surface input of NO 3 − . Soil inorganic N was dominated by NH 4 + , but invaded sites had greater subsurface soil NO 3 − . Invaded sites had greater total soil N at the surface four years post‐invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil δ 15 N increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar δ 15 N values suggest that Bromus assimilated subsurface NO 3 − , whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil δ 15 N; however, the invaded/ disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO 3 − concentrations were greater in the invaded communities during periods of ample precipitation, and NO 3 − leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO 3 − from decomposing Bromus litter to subsurface soil layers, accessing that deep‐soil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.

Ecology↗

Effects of management and climate on elk brucellosis in the Greater Yellowstone Ecosystem

Every winter, government agencies feed ∼6000 metric tons (6 × 10 6 kg) of hay to elk in the southern Greater Yellowstone Ecosystem (GYE) to limit transmission of Brucella abortus, the causative agent of brucellosis, from elk to cattle. Supplemental feeding, however, is likely to increase the transmission of brucellosis in elk, and may be affected by climatic factors, such as snowpack. We assessed these possibilities using snowpack and feeding data from 1952 to 2006 and disease testing data from 1993 to 2006. Brucellosis seroprevalence was strongly correlated with the timing of the feeding season. Longer feeding seasons were associated with higher seroprevalence, but elk population size and density had only minor effects. In other words, the duration of host aggregation and whether it coincided with peak transmission periods was more important than just the host population size. Accurate modeling of disease transmission depends upon incorporating information on how host contact rates fluctuate over time relative to peak transmission periods. We also found that supplemental feeding seasons lasted longer during years with deeper snowpack. Therefore, milder winters and/or management strategies that reduce the length of the feeding season may reduce the seroprevalence of brucellosis in the elk populations of the southern GYE.

Wyoming↗

Probable causes of increasing brucellosis in free-ranging elk of the Greater Yellowstone Ecosystem

While many wildlife species are threatened, some populations have recovered from previous overexploitation, and data linking these population increases with disease dynamics are limited. We present data suggesting that free-ranging elk (Cervus elaphus) are a maintenance host for Brucella abortus in new areas of the Greater Yellowstone Ecosystem (GYE). Brucellosis seroprevalence in free-ranging elk increased from 0-7% in 1991-1992 to 8-20% in 2006-2007 in four of six herd units around the GYE. These levels of brucellosis are comparable to some herd units where elk are artificially aggregated on supplemental feeding grounds. There are several possible mechanisms for this increase that we evaluated using statistical and population modeling approaches. Simulations of an age-structured population model suggest that the observed levels of seroprevalence are unlikely to be sustained by dispersal from supplemental feeding areas with relatively high seroprevalence or an older age structure. Increases in brucellosis seroprevalence and the total elk population size in areas with feeding grounds have not been statistically detectable. Meanwhile, the rate of seroprevalence increase outside the feeding grounds was related to the population size and density of each herd unit. Therefore, the data suggest that enhanced elk-to-elk transmission in free-ranging populations may be occurring due to larger winter elk aggregations. Elk populations inside and outside of the GYE that traditionally did not maintain brucellosis may now be at risk due to recent population increases. In particular, some neighboring populations of Montana elk were 5-9 times larger in 2007 than in the 1970s, with some aggregations comparable to the Wyoming feeding-ground populations. Addressing the unintended consequences of these increasing populations is complicated by limited hunter access to private lands, which places many ungulate populations out of administrative control. Agency-landowner hunting access partnerships and the protection of large predators are two management strategies that may be used to target high ungulate densities in private refuges and reduce the current and future burden of disease.

Ecological Applications↗

Persistence of canine distemper virus in the Greater Yellowstone Ecosystem's carnivore community

Canine distemper virus (CDV) is an acute, highly immunizing pathogen that should require high densities and large populations of hosts for long-term persistence, yet CDV persists among terrestrial carnivores with small, patchily distributed groups. We used CDV in the Greater Yellowstone ecosystem's (GYE) wolves (Canis lupus) and coyotes (Canis latrans) as a case study for exploring how metapopulation structure, host demographics, and multi-host transmission affect the critical community size and spatial scale required for CDV persistence. We illustrate how host spatial connectivity and demographic turnover interact to affect both local epidemic dynamics, such as the length and variation in inter-epidemic periods, and pathogen persistence using stochastic, spatially explicit susceptible-exposed-infectious-recovered simulation models. Given the apparent absence of other known persistence mechanisms (e.g., a carrier or environmental state, densely populated host, chronic infection, or a vector), we suggest that CDV requires either large spatial scales or multi-host transmission for persistence. Current GYE wolf populations are probably too small to support endemic CDV. Coyotes are a plausible reservoir host, but CDV would still require 50 000-100 000 individuals for moderate persistence (>50% over 10 years), which would equate to an area of 1-3 times the size of the GYE (60000-200000 km2). Coyotes, and carnivores in general, are not uniformly distributed; therefore, this is probably a gross underestimate of the spatial scale of CDV persistence. However, the presence of a second competent host species can greatly increase the probability of long-term CDV persistence at much smaller spatial scales. Although no management of CDV is currently recommended for the GYE, wolf managers in the region should expect periodic but unpredictable CDV-related population declines as often as every 2-5 years. Awareness and monitoring of such outbreaks will allow corresponding adjustments in management activities such as regulated public harvest, creating a smooth transition to state wolf management and conservation after >30 years of being protected by the Endangered Species Act.

Ecological Applications↗

Ecosystem ecology meets adaptive management: food web response to a controlled flood on the Colorado River, Glen Canyon

Large dams have been constructed on rivers to meet human demands for water, electricity, navigation, and recreation. As a consequence, flow and temperature regimes have been altered, strongly affecting river food webs and ecosystem processes. Experimental high‐flow dam releases, i.e., controlled floods, have been implemented on the Colorado River, USA, in an effort to reestablish pulsed flood events, redistribute sediments, improve conditions for native fishes, and increase understanding of how dam operations affect physical and biological processes. We quantified secondary production and organic matter flows in the food web below Glen Canyon dam for two years prior and one year after an experimental controlled flood in March 2008. Invertebrate biomass and secondary production declined significantly following the flood (total biomass, 55% decline; total production, 56% decline), with most of the decline driven by reductions in two nonnative invertebrate taxa, Potamopyrgus antipodarum and Gammarus lacustris . Diatoms dominated the trophic basis of invertebrate production before and after the controlled flood, and the largest organic matter flows were from diatoms to the three most productive invertebrate taxa ( P. antipodarum , G. lacustris , and Tubificida). In contrast to invertebrates, production of rainbow trout ( Oncorhynchus mykiss ) increased substantially (194%) following the flood, despite the large decline in total secondary production of the invertebrate assemblage. This counterintuitive result is reconciled by a post‐flood increase in production and drift concentrations of select invertebrate prey (i.e., Chironomidae and Simuliidae) that supported a large proportion of trout production but had relatively low secondary production. In addition, interaction strengths, measured as species impact values, were strongest between rainbow trout and these two taxa before and after the flood, demonstrating that the dominant consumer–resource interactions were not necessarily congruent with the dominant organic matter flows. Our study illustrates the value of detailed food web analysis for elucidating pathways by which dam management may alter production and strengths of species interactions in river food webs. We suggest that controlled floods may increase production of nonnative rainbow trout, and this information can be used to help guide future dam management decisions.

Arizona↗

Effects of landcover, water redistribution, and temperature on ecosystem processes in the South Plate Basin

Over one-third of the land area in the South Platte Basin of Colorado, Nebraska, and Wyoming, has been converted to croplands. Irrigated cropland now comprises 8% of the basin, while dry croplands make up 31%. We used the RHESSys model to compare the changes in plant productivity and vegetation-related hydrological processes that occurred as a result of either land cover alteration or directional temperature changes (&minus;2&deg;C, +4&deg;C). Land cover change exerted more control over annual plant productivity and water fluxes for converted grasslands, while the effect of temperature changes on productivity and water fluxes was stronger in the mountain vegetation. Throughout the basin, land cover change increased the annual loss of water to the atmosphere by 114 mm via evaporation and transpiration, an increase of 37%. Both irrigated and nonirrigated grains became active earlier in the year than shortgrass steppe, leading to a seasonal shift in water losses to the atmosphere. Basin-wide photosynthesis increased by 80% due to grain production. In contrast, a 4&deg;C warming scenario caused annual transpiration to increase by only 3% and annual evaporation to increase by 28%, for a total increase of 71 mm. Warming decreased basin-wide photosynthesis by 16%. There is a large elevational range from east to west in the South Platte Basin, which encompasses the western edge of the Great Plains and the eastern front of the Rocky Mountains. This elevational gain is accompanied by great changes in topographic complexity, vegetation type, and climate. Shortgrass steppe and crops found at elevations between 850 and 1800 m give way to coniferous forests and tundra between 1800 and 4000 m. Climate is increasingly dominated by winter snow precipitation with increasing elevation, and the timing of snowmelt influences tundra and forest ecosystem productivity, soil moisture, and downstream discharge. Mean annual precipitation of <500 mm on the plains below 1800 m is far less than potential evapotranspiration of 1000&ndash;1500 mm and is insufficient for optimum plant productivity. The changes in water flux and photosynthesis from conversion of steppe to cropland are the result of redistribution of snowmelt water from the mountains and groundwater pumping through irrigation projects.

Ecological Applications↗

Elk migration patterns and human activity influence wolf habitat use in the Greater Yellowstone Ecosystem

Identifying the ecological dynamics underlying human&ndash;wildlife conflicts is important for the management and conservation of wildlife populations. In landscapes still occupied by large carnivores, many ungulate prey species migrate seasonally, yet little empirical research has explored the relationship between carnivore distribution and ungulate migration strategy. In this study, we evaluate the influence of elk ( Cervus elaphus ) distribution and other landscape features on wolf ( Canis lupus ) habitat use in an area of chronic wolf&ndash;livestock conflict in the Greater Yellowstone Ecosystem, USA. Using three years of fine-scale wolf ( n = 14) and elk ( n = 81) movement data, we compared the seasonal habitat use of wolves in an area dominated by migratory elk with that of wolves in an adjacent area dominated by resident elk. Most migratory elk vacate the associated winter wolf territories each summer via a 40&ndash;60 km migration, whereas resident elk remain accessible to wolves year-round. We used a generalized linear model to compare the relative probability of wolf use as a function of GIS-based habitat covariates in the migratory and resident elk areas. Although wolves in both areas used elk-rich habitat all year, elk density in summer had a weaker influence on the habitat use of wolves in the migratory elk area than the resident elk area. Wolves employed a number of alternative strategies to cope with the departure of migratory elk. Wolves in the two areas also differed in their disposition toward roads. In winter, wolves in the migratory elk area used habitat close to roads, while wolves in the resident elk area avoided roads. In summer, wolves in the migratory elk area were indifferent to roads, while wolves in resident elk areas strongly avoided roads, presumably due to the location of dens and summering elk combined with different traffic levels. Study results can help wildlife managers to anticipate the movements and establishment of wolf packs as they expand into areas with migratory or resident prey populations, varying levels of human activity, and front-country rangelands with potential for conflicts with livestock.

Wyoming↗

Regional signatures of plant response to drought and elevated temperature across a desert ecosystem

The performance of many desert plant species in North America may decline with the warmer and drier conditions predicted by climate change models, thereby accelerating land degradation and reducing ecosystem productivity. We paired repeat measurements of plant canopy cover with climate at multiple sites across the Chihuahuan Desert over the last century to determine which plant species and functional types may be the most sensitive to climate change. We found that the dominant perennial grass, Bouteloua eriopoda, and species richness had nonlinear responses to summer precipitation, decreasing more in dry summers than increasing with wet summers. Dominant shrub species responded differently to the seasonality of precipitation and drought, but winter precipitation best explained changes in the cover of woody vegetation in upland grasslands and may contribute to woody-plant encroachment that is widespread throughout the southwestern United States and northern Mexico. Temperature explained additional variability of changes in cover of dominant and subdominant plant species. Using a novel empirically based approach we identified ‘‘climate pivot points’’ that were indicative of shifts from increasing to decreasing plant cover over a range of climatic conditions. Reductions in cover of annual and several perennial plant species, in addition to declines in species richness below the long-term summer precipitation mean across plant communities, indicate a decrease in the productivity for all but the most drought-tolerant perennial grasses and shrubs in the Chihuahuan Desert. Overall, our regional synthesis of long-term data provides a robust foundation for forecasting future shifts in the composition and structure of plant assemblages in the largest North American warm desert.

Ecology↗

Cross-ecosystem impacts of stream pollution reduce resource and contaminant flux to riparian food webs

The effects of aquatic contaminants are propagated across ecosystem boundaries by aquatic insects that export resources and contaminants to terrestrial food webs; however, the mechanisms driving these effects are poorly understood. We examined how emergence, contaminant concentration, and total contaminant flux by adult aquatic insects changed over a gradient of bioavailable metals in streams and how these changes affected riparian web-building spiders. Insect emergence decreased 97% over the metal gradient, whereas metal concentrations in adult insects changed relatively little. As a result, total metal exported by insects (flux) was lowest at the most contaminated streams, declining 96% among sites. Spiders were affected by the decrease in prey biomass, but not by metal exposure or metal flux to land in aquatic prey. Aquatic insects are increasingly thought to increase exposure of terrestrial consumers to aquatic contaminants, but stream metals reduce contaminant flux to riparian consumers by strongly impacting the resource linkage. Our results demonstrate the importance of understanding the contaminant-specific effects of aquatic pollutants on adult insect emergence and contaminant accumulation in adults to predict impacts on terrestrial food webs.

Ecological Applications↗

Multi-trophic resilience of boreal lake ecosystems to forest fires

Fires are the major natural disturbance in the boreal forest, and their frequency and intensity will likely increase as the climate warms. Terrestrial nutrients released by fires may be transported to boreal lakes, stimulating increased primary productivity, which may radiate through multiple trophic levels. Using a before‐after‐control‐impact (BACI) design, with pre‐ and postfire data from burned and unburned areas, we examined effects of a natural fire across several trophic levels of boreal lakes, from nutrient and chlorophyll levels, to macroinvertebrates, to waterbirds. Concentrations of total nitrogen and phosphorus were not affected by the fire. Chlorophyll a levels were also unaffected, likely reflecting the stable nutrient concentrations. For aquatic invertebrates, we found that densities of three functional feeding groups did not respond to the fire (filterers, gatherers, scrapers), while two groups increased (shredders, predators). Amphipods accounted for 98% of shredder numbers, and we hypothesize that fire‐mediated habitat changes may have favored their generalist feeding and habitat ecology. This increase in amphipods may, in turn, have driven increased predator densities, as amphipods were the most numerous invertebrate in our lakes and are commonly taken as prey. Finally, abundance of waterbird young, which feed primarily on aquatic invertebrates, was not affected by the fire. Overall, ecosystems of our study lakes were largely resilient to forest fires, likely due to their high initial nutrient concentrations and small catchment sizes. Moreover, this resilience spanned multiple trophic levels, a significant result for ecologically similar boreal regions, especially given the high potential for increased fires with future climate change.

Alaska↗

Amphibians in the climate vise: loss and restoration of resilience of montane wetland ecosystems in the western US

Wetlands in the remote mountains of the western US have undergone two massive ecological “experiments” spanning the 20th century. Beginning in the late 1800s and expanding after World War II, fish and wildlife managers intentionally introduced millions of predatory trout (primarily Oncorhynchus spp) into fishless mountain ponds and lakes across the western states. These new top predators, which now occupy 95% of large mountain lakes, have limited the habitat distributions of native frogs, salamanders, and wetland invertebrates to smaller, more ephemeral ponds where trout do not survive. Now a second “experiment” – anthropogenic climate change – threatens to eliminate many of these ephemeral habitats and shorten wetland hydroperiods. Caught between climate-induced habitat loss and predation from introduced fish, native mountain lake fauna of the western US – especially amphibians – are at risk of extirpation. Targeted fish removals, guided by models of how wetlands will change under future climate scenarios, provide innovative strategies for restoring resilience of wetland ecosystems to climate change.

Western United States↗

Rapid toxicity assessment of sediments from estuarine ecosystems: A new tandem in vitro testing approach

Microtox® and Mutatox® were used to evaluate the acute toxicity and genotoxicity, respectively, of organic sediment extracts from Pensacola Bay and St. Andrew Bay, two estuaries that cover about 273 and 127 km 2 , respectively, along the Gulf coast of Florida, USA. The sensitivity and selectivity of these two bioluminescent toxicity assays were demonstrated in validation studies with over 50 pesticides, genotoxins, and industrial pollutants, both as single compounds and in complex mixtures. The 50% effective concentration (EC50) values of insecticides, petroleum products, and polychlorinated biphenyls determined by Microtox all tended to group around the mean EC50 value of 1.2 (0.8) mg/L. The polycyclic aromatic hydrocarbon sensitivity of Mutatox was in general similar to that reported in the Ames test. Surficial sediment samples were collected, extracted with dichloromethane, evaporated and concentrated under nitrogen, dissolved in dimethyl sulfoxide, assayed for acute toxicity and genotoxicity, and compared with reference sediments. Samples with low EC50 values, and determined to be genotoxic, were detected in Massalina Bayou, Watson Bayou, East Bay, and St. Andrew Bay–East in St. Andrew Bay as well as Bayou Grande, Bayou Chico, and Bayou Texar in Pensacola Bay. An overview of these data sets analyzed by Spearman rank correlation showed a significant correlation between acute toxicity and genotoxicity ( p < 0.05). Microtox and Mutatox in tandem was a sensitive, cost‐effective, and rapid (<24 h) screening tool that identified troublesome areas of pollution and assessed the potential sediment toxicity of lipophilic contaminants in aquatic ecosystems.

Environmental Toxicology and Chemistry↗

Catchment disturbance and stream metabolism: Patterns in ecosystem respiration and gross primary production along a gradient of upland soil and vegetation disturbance

Catchment characteristics determine the inputs of sediments and nutrients to streams. As a result, natural or anthropogenic disturbance of upland soil and vegetation can affect instream processes. The Fort Benning Military Installation (near Columbus, Georgia) exhibits a wide range of upland disturbance levels because of spatial variability in the intensity of military training. This gradient of disturbance was used to investigate the effect of upland soil and vegetation disturbance on rates of stream metabolism (ecosystem respiration rate [ER] and gross primary production rate [GPP]). Stream metabolism was measured using an open-system, single-station approach. All streams were net heterotrophic during all seasons. ER was highest in winter and spring and lowest in summer and autumn. ER was negatively correlated with catchment disturbance level in winter, spring, and summer, but not in autumn. ER was positively correlated with abundance of coarse woody debris, but not significantly related to % benthic organic matter. GPP was low in all streams and generally not significantly correlated with disturbance level. Our results suggest that the generally intact riparian zones of these streams were not sufficient to protect them from the effect of upland disturbance, and they emphasize the role of the entire catchment in determining stream structure and function. ?? 2005 by The North American Benthological Society.

Journal of the North American Benthological Societ↗

Importance of storm events in controlling ecosystem structure and function in a Florida Gulf Coast estuary

From 8/95 to 2/01, we investigated the ecological effects of intra- and inter-annual variability in freshwater flow through Taylor Creek in southeastern Everglades National Park. Continuous monitoring and intensive sampling studies overlapped with an array of pulsed weather events that impacted physical, chemical, and biological attributes of this region. We quantified the effects of three events representing a range of characteristics (duration, amount of precipitation, storm intensity, wind direction) on the hydraulic connectivity, nutrient and sediment dynamics, and vegetation structure of the SE Everglades estuarine ecotone. These events included a strong winter storm in November 1996, Tropical Storm Harvey in September 1999, and Hurricane Irene in October 1999. Continuous hydrologic and daily water sample data were used to examine the effects of these events on the physical forcing and quality of water in Taylor Creek. A high resolution, flow-through sampling and mapping approach was used to characterize water quality in the adjacent bay. To understand the effects of these events on vegetation communities, we measured mangrove litter production and estimated seagrass cover in the bay at monthly intervals. We also quantified sediment deposition associated with Hurricane Irene's flood surge along the Buttonwood Ridge. These three events resulted in dramatic changes in surface water movement and chemistry in Taylor Creek and adjacent regions of Florida Bay as well as increased mangrove litterfall and flood surge scouring of seagrass beds. Up to 5 cm of bay-derived mud was deposited along the ridge adjacent to the creek in this single pulsed event. These short-term events can account for a substantial proportion of the annual flux of freshwater and materials between the mangrove zone and Florida Bay. Our findings shed light on the capacity of these storm events, especially when in succession, to have far reaching and long lasting effects on coastal ecosystems such as the estuarine ecotone of the SE Everglades.

Journal of Coastal Research↗

Measuring environmental change in forest ecosystems by repeated soil sampling: a North American perspective

Environmental change is monitored in North America through repeated measurements of weather, stream and river flow, air and water quality, and most recently, soil properties. Some skepticism remains, however, about whether repeated soil sampling can effectively distinguish between temporal and spatial variability, and efforts to document soil change in forest ecosystems through repeated measurements are largely nascent and uncoordinated. In eastern North America, repeated soil sampling has begun to provide valuable information on environmental problems such as air pollution. This review synthesizes the current state of the science to further the development and use of soil resampling as an integral method for recording and understanding environmental change in forested settings. The origins of soil resampling reach back to the 19th century in England and Russia. The concepts and methodologies involved in forest soil resampling are reviewed and evaluated through a discussion of how temporal and spatial variability can be addressed with a variety of sampling approaches. Key resampling studies demonstrate the type of results that can be obtained through differing approaches. Ongoing, large-scale issues such as recovery from acidification, long-term N deposition, C sequestration, effects of climate change, impacts from invasive species, and the increasing intensification of soil management all warrant the use of soil resampling as an essential tool for environmental monitoring and assessment. Furthermore, with better awareness of the value of soil resampling, studies can be designed with a long-term perspective so that information can be efficiently obtained well into the future to address problems that have not yet surfaced.

Journal of Environmental Quality↗

Dietary breadth of grizzly bears in the Greater Yellowstone Ecosystem

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.

Idaho, Montana, Wyoming↗