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Evan H. Campbell Grant

Publications and source records attributed to Evan H. Campbell Grant.

At least 73 records · Page 4Linked to original sources
USGS2020

Assessing the risks posed by SARS-CoV-2 in and via North American bats — Decision framing and rapid risk assessment

The novel β-coronavirus, SARS-CoV-2, may pose a threat to North American bat populations if bats are exposed to the virus through interaction with humans, if the virus can subsequently infect bats and be transmitted among them, and if the virus causes morbidity or mortality in bats. Further, if SARS-CoV-2 became established in bat populations, it could possibly serve as a source for new infection in humans, domesticated animals, or other wild animals. Wildlife management agencies in the United States are concerned about these potential risks and have begun to issue guidance regarding work that brings humans into contact with bats, but decision making is difficult because of the high degree of uncertainty about many of the relevant processes that could lead to virus transmission and establishment. The risk assessment described in this report was undertaken to provide management agencies with an understanding of the likelihood that the various steps in the causal pathways would lead to SARS-CoV-2 infection of North American bats from people. This assessment focused on the active season for bats in the temperate zone of North America (April 15 through November 15), and used Myotis lucifugus (little brown bats) as a surrogate species. At the time of this work (April 2020), no empirical data about the effects of SARS-CoV-2 on North American bats were available, so a formal process of expert judgment was used to elicit estimates of the underlying parameters. Twelve experts in bat ecology, epidemiology, virology, and wildlife disease from the United States, United Kingdom, and Australia participated in the elicitation. A Monte Carlo simulation model was used to integrate the parameter estimates elicited from the experts and to predict the likelihood of exposure and infection in bats through a series of transmission pathways, with particular attention to capturing uncertainty in the predictions. Given the current state of knowledge as expressed by the expert panel, the results of this assessment indicate that there is a non-negligible risk of transmission of SARS-CoV-2 from humans to bats. For example, if a research scientist were shedding SARS-CoV-2 virus while handling bats under the field protocols used in North America prior to the COVID-19 pandemic, the risk model indicates that 50 percent (uncertainty, 15–84 percent) of those bats could be exposed to virus, and 17 percent (uncertainty, 3–51 percent) could become infected. Use of personal protective equipment, especially a respirator, is expected to reduce the exposure risk. The expert panel estimated that exposure risk from research scientists could be reduced 94–96 percent (uncertainty, 86–99 percent) through proper use of appropriate N95 respirators (a type of mechanical filter worn over the nose and mouth), dedicated clothing (such as Tyvek coveralls), and gloves. Should any North American bats become infected with SARS-CoV-2, the expert panel estimated that there is an approximately 33-percent chance the virus could spread within a bat population. This study, conducted by the U.S. Geological Survey in cooperation with the U.S. Fish and Wildlife Service, identified several critical uncertainties that could affect the estimate of risks associated with SARS-CoV-2 entering bat populations—notably, the underlying probability that a human would be shedding virus while working with bats, the likelihood of the virus replicating in bat tissue, and the likelihood of transmission of the virus within bat populations. Ongoing empirical work during May–October 2020 may shed light on these issues. Follow-up work is needed to better understand the probability of transmission of SARS-CoV-2 to bats from the general public; the manner in which the probabilities of exposure, infection, and transmission would differ during hibernation compared to the breeding season; and the likelihood of important effects, like morbidity and mortality in bats, the possibility of zoonosis from a North American bat reservoir, and effects of and on other wildlife.

North America↗
USGS2020

Identifying research needs to inform white-nose syndrome management decisions

Ecological understanding of host–pathogen dynamics is the basis for managing wildlife diseases. Since 2008, federal, state, and provincial agencies and tribal and private organizations have collaborated on bat and white‐nose syndrome (WNS) surveillance and monitoring, research, and management programs. Accordingly, scientists and managers have learned a lot about the hosts, pathogen, and dynamics of WNS. However, effective mitigation measures to combat WNS remain elusive. Host–pathogen systems are complex, and identifying ecological research priorities to improve management, choosing among various actions, and deciding when to implement those actions can be challenging. Through a cross‐disciplinary approach, a group of diverse subject matter experts created an influence diagram used to identify uncertainties and prioritize research needs for WNS management. Critical knowledge gaps were identified, particularly with respect to how WNS dynamics and impacts may differ among bat species. We highlight critical uncertainties and identify targets for WNS research. This tool can be used to maximize the likelihood of achieving bat conservation goals within the context and limitations of specific real‐world scenarios.

Conservation Biology↗
USGS2020

A hierarchical analysis of habitat area, connectivity, and quality on amphibian diversity across spatial scales

Habitat fragmentation can alter species distributions and lead to reduced diversity at multiple scales. Yet, the literature describing fragmentation effects on biodiversity patterns is contradictory and inconclusive, possibly because most studies fail to integrate spatial scale into experimental designs and statistical analyses. As a result, it is difficult to extrapolate the effects of fragmentation to large-scaled systems in which conservation management is of immediate importance. Objectives To explore the influence of fragmentation on biodiversity across scales, we (1) estimated the effects of habitat area, connectivity, and quality at both local (i.e. community) and regional (i.e. metacommunity) scales; and (2) evaluated the direction, magnitude, and precision of these effect estimates at both spatial scales. Methods We developed a multi-region community occupancy model to analyze 13 years (2005-2017) of amphibian monitoring data within the National Capital Region, a network of U.S. National Parks. Results Overall, we found a positive effect of park size and a negative effect of isolation on species richness at the park-level (i.e. metacommunity), and generally positive effects of wetland area, connectivity, and quality on species richness at the wetland-level (i.e. community), although parameter estimates varied among species. Covariate effects were less precise, but effects sizes were larger, at the local wetland-level as compared to the larger park-level scale. Conclusions Our analysis reveals how scale can mediate interpretation of results from scientific studies, which might help explain conflicting narratives concerning the impacts of fragmentation in the published literature. Our hierarchical framework can help managers and policymakers elucidate the relevant spatial scale(s) to target conservation efforts.

Landscape Ecology↗
USGS2020

Moving from decision to action in conservation science

Biodiversity loss is a major threat to the integrity of ecosystems and is projected to worsen, yet the path to successful conservation remains elusive. Decision support frameworks (DSFs) are increasingly applied by resource managers to navigate the complexity, uncertainty, and differing socio-ecological objectives inherent to conservation problems. Most published conservation research that uses DSFs focuses on analytical stages (e.g., identifying an optimal decision), making it difficult to assess and learn from previous examples in a conservation practice context. Here, we (1) evaluate the relationship between the application of decision science and the resulting conservation outcomes, and (2) identify and address existing barriers to the application of DSFs to conservation practice. To do this, we develop a framework for evaluating conservation initiatives using decision science that emphasizes setting attainable goals, building momentum, and obtaining partner buy-in. We apply this framework to a systematic review of amphibian conservation decision support projects, including a follow-up survey of the pertinent conservation practitioners, stakeholders, and scientists. We found that all projects identified optimal solutions to reach stated objectives, but positive conservation outcomes were limited when implementation challenges arose. Further, we identified multiple barriers (e.g., dynamic and hierarchical leadership, scale complexity, limited resource availability) that can inhibit the progression from decision identification to action implementation (i.e., ‘decision-implementation gap’), and to successful conservation outcomes. Based on these results, we provide potential actionable steps and avenues for future development of DSFs to facilitate the transition from decision to action and the realization of conservation successes.

Biological Conservation↗
USGS2020

Factors facilitating co-occurrence at the Range Boundary of Shenandoah and Red-backed Salamanders

The transition from species in allopatry to sympatry, i.e., the co-occurrence zone of competing species, allows for investigation of forces structuring range limits and provides evidence of the evolutionary and population responses of competing species, including mechanisms facilitating co-occurrence (e.g., character displacement). The Shenandoah Salamander (Plethodon shenandoah), an endangered plethodontid, is limited to three mountaintops in Shenandoah National Park, Virginia, USA. This species’ distributional limits are attributed to competitive exclusion by the Red-backed Salamander (P. cinereus). Recent work showed range overlap between these species is greater than previously thought, requiring investigation of species morphology, behavior, and demographic measures in single-species and co-occurrence zones that might facilitate such overlap. We analyzed individual characteristics (e.g., life stage, size, color, and microhabitat-use) from two years of transect surveys to see if traits differed within and outside co-occurrence zones. Measures showed species- and zonal-specific differences, but we found limited support for character displacement. Both species were larger when co-occurringin the co-occurrence zone, indicating larger animals might better compete for resources or that symmetric competition restricts dispersal or recruitment processes at the co-occurrence zone. Microhabitat use also differed by species across transects, with Red-backed Salamanders using more rock microhabitats in the co-occurrence zone, potentially due to competition for microclimates that minimize physiological stress. The lack of strong evidence for morphologic, behavioral, or demographic differentiation in situ at the range edge suggests competition may be weaker than previously thought with other factors contributing to the range limits of Shenandoah Salamanders.

Journal of Herpetology↗
USGS2020

Mapping climate change resistant vernal pools in the northeastern U.S.

Vernal pools are seasonal wetlands that provide important breeding habitat for a variety of amphibian species. As future climate projections indicate warmer growing seasons and earlier seasonal increases in evapotranspiration, some managers of vernal pools have expressed concern that pools may dry earlier in the season, potentially interfering with completion of amphibian life cycles. In this context, a subset of pools might function as hydrologic refugia by providing wetland habitat later into the year under relatively dry conditions, thus supporting species persistence even as summer conditions become warmer and droughts more frequent. This study used approximately 3,000 field observations of inundation from 450 pools in the northeastern United States—located from West Virginia to Maine—to train machine-learning models for predicting the likelihood of pool inundation. Inputs to these models included pool size, day of the year, climate conditions, short-term weather patterns, and attributes of the landscapes in which pools were embedded. Predictions of pool wetness were generated on a daily time step from late April through late July using three short-term weather scenarios (dry, wet, and average) under historical climate conditions and four sets of downscaled climate projections (2050s and 2080s under Representative Concentration Pathways 4.5 and 8.5). The modeling and inundation prediction process was replicated using four inundation thresholds on wetted area and depth. Model outputs can enable users to examine the inundation thresholds, time points, weather scenarios, and future climate projections most relevant to their management needs. Together with long-term monitoring of individual pools at the site scale, this regional-scale study can support amphibian conservation by helping to identify subsets of pools that may be most likely to function as hydrologic refugia from changing climate conditions.

Connecticut, Delaware, Maine, Maryland, Massachuse↗
USGS2019

Successful molecular detection studies require clear communication among diverse research partners

Molecular detection techniques are powerful tools used in ecological applications ranging from diet analyses to pathogen surveillance. Research partnerships that use these tools often involve collaboration among professionals with expertise in field biology, laboratory techniques, quantitative modeling, wildlife disease, and natural resource management. However, in many cases, each of these collaborators lacks specific knowledge about the approaches, decisions, methods, and terminology used by their research partners, which can impede effective communication and act as a barrier to the efficient use of molecular data for ecological inferences and subsequent conservation decision making. We outline a collaborative framework to assist colleagues with diverse types of expertise to effectively translate their scientific and management needs to research partners from other specialties. The molecular techniques used to detect organisms will continue to advance both in sophistication and in the breadth of ecological applications. Our objective is to enable ecologists to harness the full utility of these methods by developing effective collaborative partnerships.

Frontiers in Ecology and the Environment↗
USGS2019

Different management strategies are optimal for combating disease in East Texas cave versus culvert hibernating bat populations

Management decisions for species impacted by emerging infectious diseases are challenging when there are uncertainties in the effectiveness of management actions. Wildlife managers must balance trade‐offs between mitigating the effects of the disease and the associated consequences on other aspects of the managed system. An example of this challenge is exemplified in the response to white‐nose syndrome (WNS), a disease of hibernating bats. The fungal pathogen that causes WNS, Pseudogymnoascus destructans , continues to spread throughout North America. Texas, recently confirmed positive for the fungus, has documented 33 bat species in the state, with nearly half of those species naïve to the pathogen. We explicitly incorporated multiple management objectives, uncertainty, and risk in the Texas Parks and Wildlife Department decision to manage East Texas populations of the tri‐colored bat ( Perimyotis subflavus ), a species highly susceptible to WNS. Alternatives included individual actions that act against P . destructans or benefit bats, a no active management option, and combinations of actions. Although our main objective was to identify WNS mitigation measures for tri‐colored bats in culverts, we also considered the transferability of the decision for natural caves. In this scenario, the optimal decision differed for culverts and caves, with a “portfolio” combination of actions ranking as the best alternative for culverts and a single vaccine alternative for caves. Because the top management alternatives differed markedly between these two systems, finding treatments that have broad application is likely infeasible, given that each management decision is characterized by different mixtures of competing objectives.

Texas↗
USGS2019

Identifying common decision problem elements for the management of emerging fungal diseases of wildlife

Emerging infectious diseases (EIDs) of wildlife have characteristics that make them difficult to manage, leading to reactive and often ineffective management strategies. Currently, two fungal pathogens, Pseudogymnoascus destructans (Pd) and Batrachochytrium salamandrivorans (Bsal), are causing declines in novel host species. To improve the application of management strategies addressing the risk of these pathogens to North American wildlife, we queried wildlife managers about their concerns regarding managing populations of bats and amphibians potentially impacted by Pd and Bsal. Using these responses, we identified aspects of each decision problem that were shared across pathogens, regions and agencies – and found similarities in decision-problem elements for disease management. Reframing management problems as decisions can enable managers to identify similarities across EIDs, i.e. uncertainties within management actions, and improve reactive responses if proactive management is not possible. Such an approach recognizes context-specific constraints and identifies relevant uncertainties that must be reduced in developing a response.

Society and Natural Resources↗
USGS2019

Knowing your limits: Estimating range boundaries and co-occurrence zones for two competing plethodontid salamanders

Understanding threats to species persistence requires knowledge of where species currently occur. We explore methods for estimating two important facets of species distributions, namely where the range limit occurs and how species interactions structure distributions. Accurate understanding of range limits is crucial for predicting range dynamics and shifts in response to interspecific interactions and climate change. Additionally, species interactions are increasingly recognized as an important but not well‐understood predictor of range shifts. Our objective was to predict range limits and contact zones for two plethodontid salamanders, the highly range‐restricted Shenandoah salamander ( Plethodon shenandoah ) and the wide‐ranging red‐backed salamander ( Plethodon cinereus ). Using detection/non‐detection data, we assess four methodological decisions when estimating species’ distributions: (1) accounting for imperfect detection, (2) covariates to predict species occurrences, (3) accounting for species interactions, and (4) the inclusion of spatial autocorrelation. We found that Shenandoah salamander and red‐backed salamander co‐occurrence would have been underestimated and the range edge misidentified had we not accounted for incomplete detection. Covariates related to habitat were not sufficient to explain species’ range boundaries. Models that included spatial autocorrelation (i.e., a conditional autoregressive random effect) performed better than models that included just species interactions (i.e., detection and occurrence were conditional on the other species being present) and models that included both spatial autocorrelation and species interactions. Further, we found that the breadth of primary contact zones was typically 60–170 m, which is greater on average than previous estimates. In addition, we frequently observed secondary, disjunct contact zones along the range boundary. Understanding the extent to which species co‐occur and how the range boundaries are shaped is crucial to conservation efforts. Our work indicates that accounting for detection is crucial for accurately characterizing range edges and that spatial models may be especially effective in modeling distributions at the boundary.

Ecosphere↗
USGS2019

North-facing slopes and elevation shape asymmetric genetic structure in the range-restricted salamander Plethodon shenandoah

Species with narrow environmental preferences are often distributed across fragmented patches of suitable habitat, and dispersal among subpopulations can be difficult to directly observe. Genetic data collected at population centers can help quantify gene flow, which is especially important for vulnerable species with a disjunct range. Plethodon shenandoah is a Federally Endangered salamander known only from three mountaintops in Virginia, USA. To reconstruct the evolutionary history and population connectivity of this species, we generated both mitochondrial and nuclear data using sequence capture for all three populations and found strong population structure that was independent of geographic distance. Both the nuclear markers and mitochondrial genome indicated a deep split between the most southern population and the combined central and northern population. Although there was some mitochondrial haplotype-splitting between the central and northern populations, there was complete admixture in nuclear markers. This is indicative of either a recent split or current male-biased dispersal among mountain isolates. Models of landscape resistance found that dispersal across north-facing slopes at mid-elevation levels best explain the observed genetic structure among populations. These unexpected results highlight the importance of landscape features in understanding and predicting movement and fragmentation of salamanders across space.

Ecology and Evolution↗
USGS2019

Functional variation at an expressed MHC class IIß locus associates with Ranavirus infection intensity in larval anuran populations

Infectious diseases are causing catastrophic losses to biodiversity globally. Iridoviruses in the genus Ranavirus are among the leading causes of amphibian disease-related mortality. Polymorphisms in major histocompatibility complex (MHC) genes are significantly associated with variation in amphibian susceptibility to pathogens. MHC genes encode diverse cell-surface molecules that can recognize and bind to a wide array of pathogen peptides, and are divided into two classes. While MHC class I genes are the classic mediators of viral acquired immunity, larval amphibians do not express them. However, MHC class II gene diversity may be an important predictor of Ranavirus susceptibility in larval amphibians, the life stage most susceptible to Ranavirus. We surveyed natural populations of larval wood frogs (Lithobates sylvaticus), which are highly susceptible to Ranavirus, across 17 ponds and two years in Maryland, USA. We sequenced the peptide-binding region of an expressed MHC class IIß locus and assessed allelic and genetic diversity. We converted alleles to functional supertypes and determined if physiochemical properties of peptide-binding regions influenced host responses to Ranavirus. Among 334 sampled individuals, 26% were infected with Ranavirus, and among infected individuals the average intensity was 7.12 x 107 virus copies. We recovered 20 unique MHC class IIß alleles that fell into two deeply diverged clades and seven functional supertypes. Variation in MHC supertypes were associated with Ranavirus infection intensity, but not prevalence. MHC supertype heterozygotes and individuals with the MHC supertype genotype ST1/ST7 had significantly lower Ranavirus infection intensity compared to homozygotes and all other genotypes. We conclude that MHC class IIß functional genetic variation is an important component of Ranavirus susceptibility. Identifying immune system gene signatures linked to variation in disease susceptibility can inform mitigation strategies for combatting global amphibian declines.

Immunogenetics↗
USGS2019

Disease‐structured N‐mixture models: A practical guide to model disease dynamics using count data

Obtaining inferences on disease dynamics (e.g., host population size, pathogen prevalence, transmission rate, host survival probability) typically requires marking and tracking individuals over time. While multistate mark–recapture models can produce high‐quality inference, these techniques are difficult to employ at large spatial and long temporal scales or in small remnant host populations decimated by virulent pathogens, where low recapture rates may preclude the use of mark–recapture techniques. Recently developed N ‐mixture models offer a statistical framework for estimating wildlife disease dynamics from count data. N ‐mixture models are a type of state‐space model in which observation error is attributed to failing to detect some individuals when they are present (i.e., false negatives). The analysis approach uses repeated surveys of sites over a period of population closure to estimate detection probability. We review the challenges of modeling disease dynamics and describe how N ‐mixture models can be used to estimate common metrics, including pathogen prevalence, transmission, and recovery rates while accounting for imperfect host and pathogen detection. We also offer a perspective on future research directions at the intersection of quantitative and disease ecology, including the estimation of false positives in pathogen presence, spatially explicit disease‐structured N ‐mixture models, and the integration of other data types with count data to inform disease dynamics. Managers rely on accurate and precise estimates of disease dynamics to develop strategies to mitigate pathogen impacts on host populations. At a time when pathogens pose one of the greatest threats to biodiversity, statistical methods that lead to robust inferences on host populations are critically needed for rapid, rather than incremental, assessments of the impacts of emerging infectious diseases.

Ecology and Evolution↗
USGS2019

The contribution of road-based citizen science to the conservation of pond-breeding amphibians

Roadside amphibian citizen science (CS) programmes bring together volunteers focused on collecting scientific data while working to mitigate population declines by reducing road mortality of pond‐breeding amphibians. Despite the international popularity of these movement‐based, roadside conservation efforts (i.e. “big nights,” “bucket brigades” and “toad patrols”), direct benefits to conservation have rarely been quantified or evaluated. As a case study, we used a population simulation approach to evaluate how volunteer intensity, frequency and distribution influence three conservation outcomes (minimum population size, population growth rate and years to extinction) of the spotted salamander ( Ambystoma maculatum ), often a focal pond‐breeding amphibian of CS and conservation programmes in the United States. Sensitivity analysis supported the expectation that spotted salamander populations were primarily recruitment‐driven. Thus, conservation outcomes were highest when volunteers focused on metamorph outmigration as opposed to adult in‐migration—contrary to the typical timing of such volunteer events. Almost every volunteer strategy resulted in increased conservation outcomes compared to a no‐volunteer strategy. Specifically, volunteer frequency during metamorph migration increased outcomes more than the same increases in volunteer effort during adult migration. Small population sizes resulted in a negligible effect of volunteer intensity. Volunteers during the first adult in‐migration had a relatively small effect compared to most other strategies. Synthesis and applications . Although citizen science (CS)‐focused conservation actions could directly benefit declining populations, additional conservation measures are needed to halt or reverse local amphibian declines. This study demonstrates a need to evaluate the effectiveness of focusing CS mitigation efforts on the metamorph stage, as opposed to the adult stage. This may be challenging, compared to other management actions such as road‐crossing infrastructure. Current amphibian CS programmes will be challenged to balance implementing evidence‐based conservation measures on the most limiting life stage, while retaining social and community benefits for volunteers.

Journal of Applied Ecology↗
USGS2019

Principles of translational science education

In a recent special issue in Frontiers in Ecology and the Environment, Enquist et al. (2017) present a welcome streamlining of modern applied ecology emphasizing a collaborative approach to applied ecological research involving resource-managers and scientists to produce actionable science: translational ecology (TE). The authors, including ecologists, social scientists, and conservation professionals, identified six principles defining the practice of translational ecology: collaboration, engagement, commitment, communication, process, and framing (Figure 1). In thinking about how science education can be enhanced by directly involving researchers in science education, we recognized remarkable overlap between the principles of translational ecology (Enquist et al. 2017) and a framework for developing mutually beneficial integrative partnerships between scientists and educators. Here, we describe scientist-educator analogies of the six principles of translational ecology: translational science education (TSE).

Frontiers in Ecology and Evolution↗
USGS2019

Amphibian conservation in the Anthropocene

Research is necessary to identify patterns in nature, to understand how a system functions, and to make predictions about the future state of an ecosystem. Applied research in conservation biology can identify effective strategies to maintain biodiversity, though many papers end with the conclusion that more research is needed. However, more research does not necessarily lead to solutions. We use the ongoing global decline of amphibians as a salient example to highlight limitations in current conservation research, and to focus on finding solutions which are directly relevant for conservation. While research has been conducted since declines were first detected in the 1990s, outside a few specific examples, little progress in conservation has been achieved. We suggest that the case of amphibian declines is relevant to conservation science in general, as the current paradigm for conservation is that management is planned after research is completed; research and management are not effectively (and not directly) connected. This disconnect illustrates the knowledge-action divide which has been identified as a serious deficiency in conservation. Accordingly, we use this introductory paper to the Special Issue (Amphibian conservation in the Anthropocene: Progress and challenges) to describe amphibians as a conservation dilemma, and to make the case for a different, more pragmatic, and more solutions-focused view of conservation research.

Kansas, Oklahoma↗
USGS2019

Effect of amphibian chytrid fungus (Batrachochytrium dendrobatidis) on apparent survival of frogs and toads in the western USA

Despite increasing interest in determining the population-level effects of emerging infectious diseases on wildlife, estimating effects of disease on survival rates remains difficult. Even for a well-studied disease such as amphibian chytridiomycosis (caused by the fungus Batrachochytrium dendrobatidis [Bd]), there are few estimates of how survival of wild hosts is affected. We applied hierarchical models to long-term capture-mark-recapture data (mean = 10.6 yrs, range = 6–15 yrs) from >5500 uniquely-marked individuals to estimate the effect of Bd on apparent survival of four threatened or endangered ranid frog species ( Rana draytonii , R. muscosa , R. pretiosa , R. sierrae ) at 14 study sites in California and Oregon (USA) and one bufonid toad ( Anaxyrus boreas ) at two study sites in Wyoming and Montana. Our models indicated that the presence of Bd on an individual reduced apparent survival of ranid frogs by ~6–15% depending on species and sex. The estimated difference between toads with and without Bd was 19% for the Montana population and 55% for the Wyoming population; however, the 95% Credible Interval of these estimates included zero. These results provide evidence for negative effects of Bd on survival in wild populations even in the absence of obvious die-offs. Determining what factors influence the magnitude of the effects of Bd on wildlife populations is an important next step toward identifying management actions. These estimates of Bd effects are important for understanding the extent and severity of disease, whether disease effects have changed over time, and for informing management actions.

California, Montana, Oregon, Wyoming↗