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Gavin G. Cotterill

Publications and source records attributed to Gavin G. Cotterill.

16 recordsLinked to original sources

Estimating GPS-based social aggregation metrics using collar data

Understanding social aggregation patterns in ungulate herds is essential for gaining behavioral insights, optimizing resource use, reducing human-wildlife conflict, and managing disease risk. As chronic wasting disease is the preeminent disease-related threat to cervid populations in North America, knowledge of contact between individuals and spatiotemporal patterns of aggregation provides opportunity to understand and potentially reduce disease risk while supporting sustainable population sizes. Herd density metrics, derived from global positioning system (GPS) data, can be used to inform management decisions. To effectively compare aggregation behavior within and between herds, aggregation metrics must be accurate. However, the consistency of metrics across different GPS collar sample sizes remains unclear and robust studies of big game require understanding how these factors may vary in different contexts. We examined the minimum sample size necessary for reliable calculations of three aggregation metrics: pairwise inter-animal distances, daily proximity rates, and kernel density estimate (KDE) areas. We used GPS collar data from the Jackson and West Green River elk herds ( Cervus canadensis ) in western Wyoming, USA, that differ in herd size and group structure (single versus multiple sub-groups), representing common practical contexts. Elk locations were acquired for the Jackson herd between 2016 and 2019 and from 2005 to 2010 for the West Green River herd. Herd-specific characteristics substantially influence the sample size necessary for accurate density measurements. As predicted, larger herds with many groups require more GPS collars than small herds with fewer groups. The sample size needed to accurately estimate aggregation varies by metric, with KDE areas, useful for indexing environmentally transmitted disease risk, generally requiring fewer samples, especially in high-density contexts. The required sample size also varies with seasonal changes in density. During periods of highest density, similar sample sizes are required to estimate inter-animal distances and proximity rates regardless of herd characteristics. Our results have implications for costs associated with studying big game herds, indicating fewer collars may be sufficient in some cases. These insights can aid researchers and managers in determining the appropriate number of GPS collars required for effective herd monitoring and informing relevant aggregation metrics for their management goals.

Wyoming

When do single-species occupancy models outperform multispecies models?

Occupancy models have become increasingly popular for species monitoring and assessment, in part, because detection/non-detection data are readily obtained using a variety of methods. Multispecies occupancy models (MSOMs) can yield more accurate parameter estimates than single-species models (SSOMs) with less data through their hierarchical structure, making MSOMs an attractive option when species are hard to detect or when data collection is constrained, leading to sparse datasets. Such constraints may arise from limited sampling resources, but also occur in rare species monitoring or where preliminary results are desired to inform adaptive management. Further, experimental habitat treatments often impose spatial constraints on sampling based on the scale of their implementation. Whether a MSOM outperforms SSOMs depends on the volume of data, characteristics of the ecological community, research goals of a study and how these factors align with modeling assumptions. We performed a simulation study of hypothetical pollinator communities under varying sampling intensities for scenarios in which experimental habitat treatments produced different community-level effects. We fit occupancy models to simulated datasets and assessed model performance. At lower sampling intensities (< 20 spatial replicates and < 4 temporal replicates), MSOM community-level treatment effect estimates were biased. Even at twice this sampling intensity, SSOMs yielded more accurate species-specific effect estimates in treatment effect scenarios with high variance. In some cases, MSOMs can pull species in the tails of distributions too far toward the community mean effect, which risks incorrect conclusions concerning whether treatments help or harm individual species. When quantifying species-specific effects is the main objective, particularly for rarely observed species, SSOMs are more robust to outliers across a range of community response scenarios. Researchers can use this information to inform study design, guide simulation studies and decide whether the higher precision of MSOMs outweighs risks of improperly estimated effects for some species.

Ecology and Evolution

Decision framing overview and performance of management alternatives for bison and elk feedground management at the National Elk Refuge in Jackson, Wyoming

This report was developed to evaluate the performance of a set of proposed alternatives for Cervus elaphus canadensis (elk) and Bison bison (bison) management at the National Elk Refuge in Wyoming, U.S.A., and to inform a National Environmental Policy Act Environmental Impact Statement focused on developing the next “Bison and Elk Management Plan” (BEMP). The U.S. Geological Survey facilitated a structured decision-making process for the U.S. Fish and Wildlife Service to develop the alternatives and the criteria (performance metrics) for evaluating the alternatives. Chapter A (this chapter) provides scoping details of the “BEMP,” a summary of the 19 metrics used to evaluate the performance of each of the 6 alternatives, and methodological details of 2 performance metrics that were not covered in other technical chapters. Additional technical details, results, and interpretations are briefly covered in this chapter but are mostly contained in chapters B–F.

Wyoming

Evaluating elk distribution and conflict under proposed management alternatives at the National Elk Refuge in Jackson, Wyoming

We evaluated measurable attributes describing the current and future distribution of Cervus elaphus canadensis (elk) across a region surrounding Jackson, Wyoming, for five feedground management alternatives proposed by the U.S. Fish and Wildlife Service as a revision to the 2007 “Bison and Elk Management Plan” of the National Elk Refuge. A resource selection function evaluated measurable attributes of interest to managers, including elk use of private property and sensitive habitat types at monthly timesteps and varying winter conditions. The study area boundaries were created through an expert elicitation process and consist of the Jackson Elk Herd Unit, Grand Teton National Park, the National Elk Refuge, and the northern third of the Fall Creek Elk Herd Unit. For each of the five alternatives, we distributed monthly elk numbers calculated in a concurrent analysis that simulated chronic wasting disease dynamics in this system for 20 years. Measurable attributes representing potential elk use of (1) private property, (2) cattle properties as an index of Brucella abortus risk, and sensitive habitats consisting of (3) Populus tremuloides Michx. (quaking aspen), (4) Populus angustifolia E. James (narrowleaf cottonwood), and (5) Salix L. (willow) in core winter use areas all closely followed the declines of elk abundance projected by the elk chronic wasting disease model. After 20 years, the continue feeding alternative ranked most favorably in terms of limiting elk days on private property and reducing brucellosis risk from elk to cattle because this alternative concentrated elk on the National Elk Refuge and resulted in the lowest elk population sizes. However, other management alternatives, including increase harvest and reduce feeding, tended to limit elk use of sensitive quaking aspen, narrowleaf cottonwood, and willow habitats during winter (December–April).

Wyoming

Bison population dynamics, harvest, and human conflict potential under feedground management alternatives at the National Elk Refuge in Jackson, Wyoming

Bison bison were once abundant across North America but declined due to overharvesting in the late 1800s. The reintroduced population in and around Jackson, Wyoming has averaged 485 individuals between 2018–2023 and is the subject of a planning process to inform management strategies that will guide the U.S. Fish and Wildlife’s next “Bison and Elk Management Plan” for the National Elk Refuge. This small population may benefit from historical winter-feeding operations on the National Elk Refuge because those operations may increase overwinter survival and limit human-bison conflicts, which are the number of individual bison that engage in nuisance, damaging, or otherwise aggressive behaviors with humans and livestock, that may lead to culling and other sources of mortality (for example, vehicle collisions). To inform the next “Bison and Elk Management Plan,” the U.S. Geological Survey used a population model to evaluate five management alternatives for bison and Cervus elaphus canadensis feedground operations that included continuing the elk and bison feeding program, immediately stopping the feeding program, and three other alternatives that would phase out the feeding program after a period of time. The results indicate that the bison population would be expected to decline over the next 20 years under all alternatives that stop feeding bison on the refuge. Further, this decline would lead to an associated reduction in bison harvest opportunities for resident, nonresident, and Tribal hunters. Finally, human-bison conflicts would also be expected to increase under the no feeding alternatives because bison may venture onto private lands in greater numbers if feed is not provisioned during winter months. In combination, these results suggest that feeding may lead to better outcomes for bison over the next 20 years; however, these effects may be traded off against other downsides of the feedground program, such as increased rates of animal-to-animal contact on feedgrounds that can lead to disease transmission.

Wyoming

Estimating the social and economic consequences of proposed management alternatives at the National Elk Refuge in Jackson, Wyoming

The National Elk Refuge (Refuge) is managed by the U.S. Fish and Wildlife Service and includes habitats for bison and elk. Bison and elk provide opportunities for wildlife-related recreation and contribute to the tourism industry in and around Jackson, Wyoming. Over the last century, the Refuge has provisioned supplemental feed to elk and, more recently, bison during winter months to ensure adequate forage and prevent starvation and conflict with private landowners. However, supplemental feeding artificially aggregates animals and can increase rates of disease transmission and localized damage to sensitive habitats near the feeding areas. This report presents analyses and results to support two of the nine management objectives in the next “Bison and Elk Management Plan,” with a particular focus on the social and economic consequences of five management alternatives considered in this study. The alternatives are to continue feeding bison and elk during winter months on the Refuge, stop feeding after CWD is measured at 3 percent prevalence or above in the Jackson elk herd, stop feeding immediately, reduce feeding for five years and then stop feeding, and increase elk harvest for five years and then stop feeding. These alternatives are anticipated to alter bison and elk population and space-use dynamics, with corresponding effects on wildlife-related recreation and tourism, including the number of visitors and sleigh-ride participants on the Refuge, and hunters and outfitters within the Jackson Elk Herd Unit. The performance of each of this study’s alternatives was variable, resulting in overlap in the performance of alternatives on the select objectives over the next 20 years. Generally, visitation-related objectives performed better under the continue feeding alternative, whereas hunting-related objectives performed better under the increase harvest alternative. The results presented here may assist U.S. Fish and Wildlife Service decision makers in balancing social and economic benefits identified in the decision-making process for the “Bison and Elk Management Plan” with other objectives evaluated in this report.

Wyoming

Elk personality and anthropogenic food subsidy: Managing conflict and migration loss

The continued decline of long-distance ungulate migrations threatens to decouple important ecological processes that increase biodiversity and wildlife abundance. Past research has focused on preserving migration paths where habitat fragmentation and loss disrupt movement corridors. However, shifting residency-migration trade-offs are the stronger driver of migration loss in some populations. Suburban residential developments may provide ungulates with anthropogenic food sources and refuge from predators, which can increase population growth among short-distance migrants relative to long-distance migrants. This trend can increase wildlife vehicle collisions and other human–wildlife conflicts while simultaneously reducing hunting opportunities. Yet, individual animals vary in their tolerance of human disturbance. We investigated how interindividual variation relative to conflict and human habituation influences elk migration and space use on shared winter range. We used a clustering algorithm applied to GPS collar data to identify elk use of anthropogenic food resources in suburban habitat. Cluster locations identified all known anthropogenic subsidy locations during the study period. Elk that used suburban anthropogenic food sources also migrated 60% shorter distances between summer and winter ranges than elk with no known use of these food subsidies. Elk use of protected wintering grounds was spatially structured such that conflict-prone, short-distance migrants disproportionately used areas with more human activity. Clustering algorithms applied to GPS collar data may allow managers to identify foci of concentrated use that generates human–wildlife conflict, and where prion deposition and environmental contamination facilitate the spread of chronic wasting disease, particularly in suburban areas with anthropogenic food subsidies. The apparent spatial structuring of shared winter range according to the conflict potential and migration strategy of individual elk may also permit managers to assess relative recruitment among cryptic population segments using different migration strategies and facilitate targeted, adaptive management actions. These associations between conflict, human habituation, and migration shed light on the urbanization of wildlife species, inform efforts to manage human–wildlife conflict and disease spread, and emphasize that a multipronged approach beyond maintaining habitat corridors may be necessary to conserve long-distance migrations for species that can become human-habituated.

Wyoming

Failure to meet the exchangeability assumption in Bayesian multispecies occupancy models: Implications for study design

Bayesian hierarchical models are ubiquitous in ecology. Random effect model structures are often employed that treat individual effects as deviations from larger population-level effects. In this way individuals are assumed to be "exchangeable" samples. Ecologists may address this exchangeability assumption intuitively, but might in certain modeling contexts ignore it altogether, including in situations where it may have large implications for study design. Multispecies occupancy models based on detection/non-detection data are an approach that can be utilized by those tasked with monitoring rare and endangered species because most literature suggests that, compared to single species occupancy models, improved parameter estimates are assured. Yet, we illustrate through a power analysis how sampling requirements to detect experimental treatment effects vary tremendously depending on whether the species exchangeability assumption is met. The degree to which species in a community respond similarly to covariates governs the ability to accurately estimate parameters using multispecies occupancy models. Detecting small or moderate changes in occupancy resulting from habitat restoration treatments may be impossible for small datasets (e.g., < 36 sampling locations, each surveyed < 8 times) even with a paired treatment-control design if the exchangeability assumption is violated. By contrast, when the assumption is met, small effects may be confidently estimated with as few as 12 sampling locations (6 pairs) and 6-8 survey events. Often, it may be impossible to know whether the exchangeability assumption is met. The statistical power needed to accurately estimate species-specific effects using detection/non-detection multispecies occupancy models depends on the unknown values of treatment effects and whether responses by species in the community diverge. When the species exchangeability assumption is violated, and at lower levels of sampling effort, multispecies occupancy models may provide worse inference than single species occupancy models.

BioRxiv

Decision framing overview and performance of management alternatives for bison and elk feedground management at the National Elk Refuge in Jackson, Wyoming

This report was developed to evaluate the performance of a set of proposed alternatives for Cervus elaphus canadensis (elk) and Bison bison (bison) management at the National Elk Refuge in Wyoming, U.S.A., and to inform a National Environmental Policy Act Environmental Impact Statement focused on developing the next “Bison and Elk Management Plan” (BEMP). The U.S. Geological Survey facilitated a structured decision-making process for the U.S. Fish and Wildlife Service to develop the alternatives and the criteria (performance metrics) for evaluating the alternatives. Chapter A (this chapter) provides scoping details of the “BEMP”, a summary of the 19 metrics that are used to evaluate the performance of each of 5 alternatives, and methodological details of 2 performance metrics that were not covered in other technical chapters. Additional technical details, results, and interpretations are briefly covered in this chapter, but are mostly contained in chapters B–E.

Wyoming

Evaluating elk distribution and conflict under proposed management alternatives at the National Elk Refuge in Jackson, Wyoming

We evaluated measurable attributes describing the current and future distribution of Cervus elaphus canadensis (elk) across a region surrounding Jackson, Wyoming, for five feedground management alternatives proposed by the U.S. Fish and Wildlife Service as a revision to the 2007 “Bison and Elk Management Plan” of the National Elk Refuge. A resource selection function evaluated measurable attributes of interest to managers, including elk use of private property and sensitive habitat types at monthly timesteps and varying winter conditions. The study area boundaries were created through an expert elicitation process and consist of the Jackson Elk Herd Unit, Grand Teton National Park, the National Elk Refuge, and the northern third of the Fall Creek Elk Herd Unit. For each of the five alternatives, we distributed monthly elk numbers calculated in a concurrent analysis that simulated chronic wasting disease dynamics in this system for 20 years. Measurable attributes representing potential elk use of (1) private property, (2) cattle properties as an index of Brucella abortus risk, and sensitive habitats consisting of (3) Populus tremuloides Michx. (quaking aspen), (4) Populus angustifolia E. James (narrowleaf cottonwood), and (5) Salix L. (willow) in core winter use areas all closely followed the declines of elk abundance projected by the elk chronic wasting disease model. After 20 years, the continue feeding alternative ranked most favorably in terms of limiting elk days on private property and reducing brucellosis risk from elk to cattle because this alternative concentrated elk on the National Elk Refuge and resulted in the lowest elk population sizes. However, other management alternatives, including increase harvest and reduce feeding, tended to limit elk use of sensitive quaking aspen, narrowleaf cottonwood, and willow habitats during winter (December–April).

Wyoming

Bison population dynamics, harvest, and conflict potential under feedground management alternatives at the National Elk Refuge in Jackson, Wyoming

Bison bison (bison) were once abundant across North America but declined due to overharvesting in the late 1800s. The reintroduced population in and around Jackson, Wyoming has averaged 485 individuals between 2018–2023 and is the subject of a planning process to inform management strategies that will guide the U.S. Fish and Wildlife’s next “Bison and Elk Management Plan” for the National Elk Refuge. This small population may benefit from historical winter-feeding operations on the National Elk Refuge because those operations may increase overwinter survival and limit human-bison conflicts, which are the number of individual bison that engage in nuisance, damaging, or otherwise aggressive behaviors with humans and livestock, that may lead to culling and other sources of mortality (for example, vehicle collisions). To inform the next “Bison and Elk Management Plan,” the U.S. Geological Survey used a population model to evaluate five management alternatives for bison and Cervus elaphus canadensis (elk) feedground operations that included continuing the elk and bison feeding program, immediately stopping the feeding program, and three other alternatives that would phase out the feeding program after a period of time. The results indicate that the bison population would be expected to decline over the next 20 years under all alternatives that stop feeding bison on the refuge. Further, this decline would lead to an associated reduction in bison harvest opportunities for resident, nonresident, and Tribal hunters. Finally, human-bison conflicts would also be expected to increase under the no feeding alternatives because bison may venture onto private lands in greater numbers if feed is not provisioned during winter months. In combination, these effects suggest that feeding may lead to better outcomes for bison over the next 20 years; however, these effects may be traded off against other downsides of the feedground program, such as increased rates of animal-to-animal contact on feedgrounds that can lead to disease transmission.

Wyoming

Estimating the social and economic consequences of proposed management alternatives at the National Elk Refuge

The National Elk Refuge (Refuge) is managed by the U.S. Fish and Wildlife Service and includes habitats for bison and elk. Bison and elk provide opportunities for wildlife-related recreation and contribute to the tourism industry in and around Jackson, Wyoming. Over the last century, the Refuge has provisioned supplemental feed to elk and, more recently, bison during winter months to ensure adequate forage and prevent starvation and conflict with private landowners. However, supplemental feeding artificially aggregates animals and can increase rates of disease transmission and localized damage to sensitive habitats near the feeding areas. This report presents analyses and results to support two of the nine management objectives in the next “Bison and Elk Management Plan,” with a particular focus on the social and economic consequences of five management alternatives considered in this study. The alternatives are to continue feeding bison and elk during winter months on the Refuge, stop feeding after CWD is measured at 3 percent prevalence or above in the Jackson elk herd, stop feeding immediately, reduce feeding for five years and then stop feeding, and increase elk harvest for five years and then stop feeding. These alternatives are anticipated to alter bison and elk population and space-use dynamics, with corresponding effects on wildlife-related recreation and tourism, including the number of visitors and sleigh-ride participants on the Refuge, and hunters and outfitters within the Jackson Elk Herd Unit. The performance of each of this study’s alternatives was variable, resulting in overlap in the performance of alternatives on the select objectives over the next 20 years. Generally, visitation-related objectives performed better under the continue feeding alternative, whereas hunting-related objectives performed better under the increase harvest alternative. The results presented here may assist U.S. Fish and Wildlife Service decision makers in balancing social and economic benefits identified in the decision-making process for the “Bison and Elk Management Plan” with other objectives evaluated in this report.

Wyoming

Ungulate personality and the human shield contribute to long-distance migration loss

Long-distance ungulate migrations are declining and past research has focused on preserving migration paths where habitat fragmentation and loss disrupts movement corridors. However, changing residency-migration tradeoffs are the stronger driver of long-distance migration loss in some populations. The human shield effect relative to predation risk and anthropogenic food resources likely shapes these tradeoffs, but individual animals also vary in their propensity to tolerate proximity to humans and developed areas. We investigated how personality relative to human-habituation affects migration behavior. We categorized elk as bold or shy based on use of anthropogenic food resources identified through a clustering algorithm applied to GPS collar data. Bold elk were 4 times more likely to select wintering areas close to human activity and migrated 60% shorter distances compared to shy elk. As a result, elk wintering grounds were spatially structured such that conflict- and disease-prone individuals selected areas adjacent to human activity. Our results suggest that bold personality traits act as a precursor to human-habituation, which permits bold elk to reap the forage and predation rewards that occur in suburban landscapes. A multi-pronged approach beyond just maintaining habitat corridors may be necessary to conserve long-distance migrations for species that can become human-habituated.

BioRxiv

Parsing the effects of demography, climate, and management on recurrent brucellosis outbreaks in elk

1. Zoonotic pathogens can harm human health and well-being directly or by impacting livestock. Pathogens that spillover from wildlife can also impair conservation efforts if humans perceive wildlife as pests. Brucellosis, caused by the bacterium Brucella abortus, circulates in elk and bison herds of the Greater Yellowstone Ecosystem and poses a risk to cattle and humans. Our goal was to understand the relative effects of climatic drivers, host demography, and management control programs on disease dynamics. 2. Using >20 years of serologic, demographic, and environmental data on brucellosis in elk, we built stochastic compartmental models to assess the influences of climate forcing, herd immunity, population turnover, and management interventions on pathogen transmission. Data were collected at feedgrounds visited in winter by free-ranging elk in Wyoming, USA. 3. Snowpack, hypothesized as a driver of elk aggregation and thus brucellosis transmission, was strongly correlated across feedgrounds. We expected this variable to drive synchronized disease dynamics across herds. Instead, we demonstrate asynchronous epizootics driven by variation in demographic rates. 4. We evaluated the effectiveness of test-and-slaughter of seropositive female elk at two feedgrounds. Test-and-slaughter temporarily reduced herd-level seroprevalence but likely reduced herd immunity while removing few infectious individuals, resulting in subsequent outbreaks once the intervention ceased. We simulated an alternative strategy of removing seronegative female elk and found it would increase herd immunity, yielding fewer infections. We evaluated a second experimental treatment wherein feeding density was reduced at one feedground, but we found no evidence for an effect despite a decade of implementation. 5. Synthesis and applications: Positive serostatus is often weakly correlated with infectiousness but is nevertheless used to make management decisions including lethal removal in wildlife disease systems. We show how this can have adverse consequences whereas efforts that maintain herd immunity can have longer-term protective effects. Climatic drivers may not result in synchronous disease dynamics across populations unless vital rates are also similar because demographic factors have a large influence on disease patterns.

Idaho, Montana, Wyoming

Hidden cost of disease in a free‐ranging ungulate: brucellosis reduces mid‐winter pregnancy in elk

Demonstrating disease impacts on the vital rates of free‐ranging mammalian hosts typically requires intensive, long‐term study. Evidence for chronic pathogens affecting reproduction but not survival is rare, but has the potential for wide‐ranging effects. Accurately quantifying disease‐associated reductions in fecundity is important for advancing theory, generating accurate predictive models, and achieving effective management. We investigated the impacts of brucellosis ( Brucella abortus ) on elk ( Cervus canadensis ) productivity using serological data from over 6,000 captures since 1990 in the Greater Yellowstone Ecosystem, USA. Over 1,000 of these records included known age and pregnancy status. Using Bayesian multilevel models, we estimated the age‐specific pregnancy probabilities of exposed and naïve elk. We then used repeat‐capture data to investigate the full effects of the disease on life history. Brucellosis exposure reduced pregnancy rates of elk captured in mid‐ and late‐winter. In an average year, we found 60% of exposed 2‐year‐old elk were pregnant compared to 91% of their naïve counterparts (a 31 percentage point reduction, 89% HPDI = 20%–42%), whereas exposed 3‐ to 9‐year‐olds were 7 percentage points less likely to be pregnant than naïve elk of their same age (89% HPDI = 2%–11%). We found these reduced rates of pregnancy to be independent from disease‐induced abortions, which afflict a portion of exposed elk. We estimate that the combination of reduced pregnancy by mid‐winter and the abortions following mid‐winter reduces the reproductive output of exposed female elk by 24%, which affects population dynamics to a similar extent as severe winters or droughts. Exposing hidden reproductive costs of disease is essential to avoid conflating them with the effects of climate and predation. Such reproductive costs cause complex population dynamics, and the magnitude of the effect we found should drive a strong selection gradient if there is heritable resistance.

Ecology and Evolution

Winter feeding of elk in the Greater Yellowstone Ecosystem and its effects on disease dynamics

Providing food to wildlife during periods when natural food is limited results in aggregations that may facilitate disease transmission. This is exemplified in western Wyoming where institutional feeding over the past century has aimed to mitigate wildlife–livestock conflict and minimize winter mortality of elk ( Cervus canadensis ). Here we review research across 23 winter feedgrounds where the most studied disease is brucellosis, caused by the bacterium Brucella abortus . Traditional veterinary practices (vaccination, test-and-slaughter) have thus far been unable to control this disease in elk, which can spill over to cattle. Current disease-reduction efforts are being guided by ecological research on elk movement and density, reproduction, stress, co-infections and scavengers. Given the right tools, feedgrounds could provide opportunities for adaptive management of brucellosis through regular animal testing and population-level manipulations. Our analyses of several such manipulations highlight the value of a research–management partnership guided by hypothesis testing, despite the constraints of the sociopolitical environment. However, brucellosis is now spreading in unfed elk herds, while other diseases (e.g. chronic wasting disease) are of increasing concern at feedgrounds. Therefore experimental closures of feedgrounds, reduced feeding and lower elk populations merit consideration.

Wyoming