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The role of genetics in chronic wasting disease of North American cervids

Chronic wasting disease (CWD) is a major concern for the management of North American cervid populations. This fatal prion disease has led to declines in populations which have high CWD prevalence and areas with both high and low infection rates have experienced economic losses in wildlife recreation and fears of potential spill-over into livestock or humans. Research from human and veterinary medicine has established that the prion protein gene ( Prnp ) encodes the protein responsible for transmissible spongiform encephalopathies (TSEs). Polymorphisms in the Prnp gene can lead to different prion forms that moderate individual susceptibility to and progression of TSE infection. Prnp genes have been sequenced in a number of cervid species including those currently infected by CWD (elk, mule deer, white-tailed deer, moose) and those for which susceptibility is not yet determined (caribou, fallow deer, sika deer). Over thousands of sequences examined, the Prnp gene is remarkably conserved within the family Cervidae; only 16 amino acid polymorphisms have been reported within the 256 amino acid open reading frame in the third exon of the Prnp gene. Some of these polymorphisms have been associated with lower rates of CWD infection and slower progression of clinical CWD. Here we review the body of research on Prnp genetics of North American cervids. Specifically, we focus on known polymorphisms in the Prnp gene, observed genotypic differences in CWD infection rates and clinical progression, mechanisms for genetic TSE resistance related to both the cervid host and the prion agent and potential for natural selection for CWD-resistance. We also identify gaps in our knowledge that require future research.

Prion

The ecology of chronic wasting disease in wildlife

Prions are misfolded infectious proteins responsible for a group of fatal neurodegenerative diseases termed transmissible spongiform encephalopathy or prion diseases. Chronic Wasting Disease (CWD) is the prion disease with the highest spillover potential, affecting at least seven Cervidae (deer) species. The zoonotic potential of CWD is inconclusive and cannot be ruled out. A risk of infection for other domestic and wildlife species is also plausible. Here, we review the current status of the knowledge with respect to CWD ecology in wildlife. Our current understanding of the geographic distribution of CWD lacks spatial and temporal detail, does not consider the biogeography of infectious diseases, and is largely biased by sampling based on hunters' cooperation and funding available for each region. Limitations of the methods used for data collection suggest that the extent and prevalence of CWD in wildlife is underestimated. If the zoonotic potential of CWD is confirmed in the short term, as suggested by recent results obtained in experimental animal models, there will be limited accurate epidemiological data to inform public health. Research gaps in CWD prion ecology include the need to identify specific biological characteristics of potential CWD reservoir species that better explain susceptibility to spillover, landscape and climate configurations that are suitable for CWD transmission, and the magnitude of sampling bias in our current understanding of CWD distribution and risk. Addressing these research gaps will help anticipate novel areas and species where CWD spillover is expected, which will inform control strategies. From an ecological perspective, control strategies could include assessing restoration of natural predators of CWD reservoirs, ultrasensitive CWD detection in biotic and abiotic reservoirs, and deer density and landscape modification to reduce CWD spread and prevalence.

Biological Reviews

Significance of stomach oil for reproduction in seabirds: An interspecies cross-fostering experiment

Stomach oil, a complex mixture of neutral dietary lipids, is a unique attribute of seabirds in the order Procellariiformes. With the exception of diving-petrels, all procellariiforms produce stomach oil and feed it to their young. We conducted an interspecies cross-fostering experiment on Bird Island, South Georgia, that was designed to reveal how the presence or absence of stomach oil in meals fed to young seabirds influences their growth, development, and survival. Hatchling South Georgia Diving-Petrels ( Pelecanoides georgicus ), a species that lacks stomach oil, were switched with hatchling Antarctic Prions ( Pachyptila desolata ), a species that feeds its young stomach oil. Diving-petrel foster parents did not successfully raise prion nestlings, presumably due to the absence of stomach oil in meals fed to nestlings. Prion foster parents successfully raised diving-petrel nestlings to fledging, but growth rates were lower, nestling fat reserves were lower, and fledging was delayed compared with controls. These results suggest that stomach oil is an essential dietary component for prion nestlings to meet their energy requirements, but diving-petrel nestlings apparently cannot efficiently assimilate stomach oil. This experiment supports the hypothesis that the production of stomach oil is an adaptation that allows breeding seabirds to enhance provisioning rates of energy to the nest, while foraging on a distant and dispersed food supply.

Georgia

Chronic wasting disease (CWD) susceptibility of several North American rodents that are sympatric with cervid CWD epidemics

Chronic wasting disease (CWD) is a highly contagious always fatal neurodegenerative disease that is currently known to naturally infect only species of the deer family, Cervidae. CWD epidemics are occurring in free-ranging cervids at several locations in North America, and other wildlife species are certainly being exposed to infectious material. To assess the potential for transmission, we intracerebrally inoculated four species of epidemic-sympatric rodents with CWD. Transmission was efficient in all species; the onset of disease was faster in the two vole species than the two Peromyscus spp. The results for inocula prepared from CWD-positive deer with or without CWD-resistant genotypes were similar. Survival times were substantially shortened upon second passage, demonstrating adaptation. Unlike all other known prion protein sequences for cricetid rodents that possess asparagine at position 170, our red-backed voles expressed serine and refute previous suggestions that a serine in this position substantially reduces susceptibility to CWD. Given the scavenging habits of these rodent species, the apparent persistence of CWD prions in the environment, and the inevitable exposure of these rodents to CWD prions, our intracerebral challenge results indicate that further investigation of the possibility of natural transmission is warranted. Copyright ?? 2010, American Society for Microbiology. All Rights Reserved.

Journal of Virology

Chronic wasting disease—Research by the U.S. Geological Survey and partners

Introduction Chronic wasting disease (CWD) is the only transmissible spongiform encephalopathy, a class of invariably fatal neurodegenerative mammalian diseases associated with a misfolded cellular prion protein found in wild free-ranging animals. Because it has a long incubation period, affected animals in Cervidae (the deer family; referred to as “cervids”) may not show signs of disease for several years. While signs are not specific to CWD, affected cervids (deer, elk, moose, and reindeer) show changes in appearance (such as progressive weight loss) and changes in behavior such as stumbling, tremors, and teeth grinding. CWD can be transmitted by direct contact or through a contaminated environment. The causative prion agent is highly resistant to degradation. In recent decades, CWD has transitioned from a novel, obscure prion disease of cervids with limited geographical distribution, to a disease that poses substantial ecological, agricultural, and economic risks across large regions of North America. Since its discovery in free-ranging elk and deer populations in the western United States in the 1980s, CWD has been reported in captive or free-ranging cervid populations in 26 States, 3 Canadian Provinces, the Republic of South Korea, Finland, Sweden, and Norway. In addition, the proportion of CWD-infected animals is increasing in many areas where the disease is already established. In some heavily affected areas, total cervid numbers have decreased over time due to CWD, which suggests that these cervid populations may not be sustainable in the long-term. The U.S. Geological Survey (USGS) conducts wildlife disease surveillance and research to support management of CWD-affected species and their habitats. The scientific information is relevant to governmental agencies that manage wildlife and their habitats including the U.S. Fish and Wildlife Service, the National Park Service, the U.S. Department of Agriculture, and other Federal, State, and Tribal agencies as well as conservation partners (non-governmental organizations, businesses, and private landowners). Each project description in this report (1–30) includes the non-USGS collaborators (Federal, State, Tribal agencies, universities) and a USGS point of contact (principal investigator). If there are USGS publications associated with the project, a publication list is provided at the end of each project description.

Alberta, California, Colorado, Idaho, Iowa, Kansas

Diagnostic testing of chronic wasting disease in white-tailed deer (Odocoileus virginianus) by RT-QuIC using multiple tissues

Chronic wasting disease (CWD) is a fatal prion disease affecting cervids (deer, elk, moose). Current methods to monitor individual disease state include highly invasive antemortem rectal biopsy or postmortem brain biopsy. Efficient, sensitive, and selective antemortem and postmortem testing of populations would increase knowledge of the dynamics of CWD epizootics as well as provide a means to track CWD progression into previously unaffected areas. Here, we analyzed the presence of CWD prions in skin samples from two easily accessed locations (ear and belly) from 30 deceased white-tailed deer ( Odocoileus viginianus ). The skin samples were enzymatically digested and analyzed by real-time quaking-induced conversion (RT-QuIC). The diagnostic sensitivity of the ear and belly skin samples were both 95%, and the diagnostic specificity of the ear and belly skin were both 100%. Additionally, the location of the skin biopsy on the ear does not affect specificity or sensitivity. These results demonstrate the efficacy of CWD diagnosis with skin biopsies using RT-QuIC. This method could be useful for large scale antemortem population testing.

Wisconsin

Chronic wasting disease: State of the science

Chronic wasting disease (CWD) is a prion disease affecting cervid species, both free-ranging and captive populations. As the geographic range continues to expand and disease prevalence continues to increase, CWD will have an impact on cervid populations, local economies, and ecosystem health. Mitigation of this “wicked” disease will require input from many different stakeholders including hunters, landowners, research biologists, wildlife managers, and others, working together. The NC1209 (North American interdisciplinary chronic wasting disease research consortium) is composed of scientists from different disciplines involved with investigating and managing CWD. Leveraging this broad breadth of expertise, the Consortium has created a state-of-the-science review of five key aspects of CWD, including current diagnostic capabilities for detecting prions, requirements for validating these diagnostics, the role of environmental transmission in CWD dynamics, and potential zoonotic risks associated with CWD. The goal of this review is to increase stakeholders’, managers’, and decision-makers’ understanding of this disease informed by current scientific knowledge.

Pathogens

Experimental infection of meadow voles ( Microtus pennsylvanicus ) with sheep scrapie

Meadow voles (Microtus pennsylvanicus) are permissive to chronic wasting disease (CWD) infection, but their susceptibility to other transmissible spongiform encephalopathies (TSEs) is poorly characterized. In this initial study, we intracerebrally challenged 6 meadow voles with 2 isolates of sheep scrapie. Three meadow voles acquired a TSE after the scrapie challenge and an extended incubation period. The glycoform profile of proteinase K-resistant prion protein (PrP(res)) in scrapie-sick voles remained similar to the sheep inocula, but differed from that of voles clinically affected by CWD. Vacuolization patterns and disease-associated prion protein (PrP(Sc)) deposition were generally similar in all scrapie-affected voles, except in the hippocampus, where PrP(Sc) staining varied markedly among the animals. Our results demonstrate that meadow voles can acquire a TSE after intracerebral scrapie challenge and that this species could therefore prove useful for characterizing scrapie isolates.

Canadian Journal of Veterinary Research

The effectiveness of harvest for limiting wildlife disease: Insights from 20 years of chronic wasting disease in Wyoming

Effective, practical options for managing disease in wildlife populations are limited, especially after diseases become established. Removal strategies (e.g., hunting or culling) are used to control wildlife diseases across a wide range of systems, despite conflicting evidence of their effectiveness. This is especially true for chronic wasting disease (CWD), an untreatable, fatal prion disease threatening cervid populations across multiple countries, for which recreational harvest has been suggested as an important disease control strategy. Using observational data to evaluate whether harvest effectively limits CWD prevalence has been difficult because statistical relationships between harvest and disease prevalence can arise from a causal effect of harvest (i.e., harvest's impacts on prevalence via changes in transmission or demographic structure) or from a number of alternative mechanisms. For instance, correlations between harvest and disease prevalence can also be driven by disease's impacts on population size and harvest (i.e., reverse causality) or from confounding variables (e.g., habitat or geographic location) that impact both harvest rates and disease prevalence. We analyzed two decades of surveillance data (2000–2021) from 10 mule deer herds in Wyoming, using statistical approaches informed by causal inference theory, to test for the effects of harvest on CWD prevalence. Herds with consistently high harvest pressure across 20 years had significantly lower prevalence. Our models predicted that harvesting 40% of adult males per year across 20 years would maintain prevalence below 5% on average, whereas if only 20% of males were harvested in each year, prevalence would increase to >30% by year 20. Moreover, shifting the relative harvest pressure within a herd over a shorter period (3 years) reduced subsequent prevalence, albeit to a smaller degree. Although high harvest is unlikely to completely eradicate CWD, our analysis suggests that maintaining hunting pressure on adult males is an important tactic for slowing CWD epidemics within mule deer herds. Our study also provides guidance for future analyses of longitudinal surveillance data, including the importance of demographic data and appropriate time lags.

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

Landscape features fail to explain spatial genetic structure in white-tailed deer across Ohio, USA

Landscape features influence wildlife movements across spatial scales and have the potential to influence the spread of disease. Chronic wasting disease (CWD) is a fatal prion disease affecting members of the family Cervidae, particularly white-tailed deer ( Odocoileus virginianus ), and the first positive CWD case in a wild deer in Ohio, USA, was recorded in 2020. Landscape genetics approaches are increasingly used to better understand potential pathways for CWD spread in white-tailed deer, but little is known about genetic structure of white-tailed deer in Ohio. The objectives of our study were to evaluate spatial genetic structure in white-tailed deer across Ohio and compare the support for isolation by distance (IBD) and isolation by landscape resistance (IBR) models in explaining this structure. We collected genetic data from 619 individual deer from 24 counties across Ohio during 2007–2009. We used microsatellite genotypes from 619 individuals genotyped at 11 loci and haplotypes from a 547-base pair fragment of the mitochondrial DNA control region. We used spatial and non-spatial genetic clustering tests to evaluate genetic structure in both types of genetic data and empirically optimized landscape resistance surfaces to compare IBD and IBR using microsatellite data. Non-spatial genetic clustering tests failed to detect spatial genetic structure, whereas spatial genetic clustering tests indicated subtle spatial genetic structure. The IBD model consistently outperformed IBR models that included land cover, traffic volume, and streams. Our results indicated widespread genetic connectivity of white-tailed deer across Ohio and negligible effects of landscape features. These patterns likely reflect some combination of minimal resistive effects of landscape features on white-tail deer movement in Ohio and the effects of regional recolonization or translocation. We encourage continued CWD surveillance in Ohio, particularly in the proximity of confirmed cases.

Ohio

Identifying priority chronic wasting disease surveillance areas for mule deer in Montana

Chronic wasting disease (CWD) is a fatal prion disease that affects a variety of ungulate species including mule deer ( Odocoileus hemionus ). As of 2014, no CWD cases had been reported in free-ranging ungulates in Montana. However, nearby cases in Canada, Wyoming, and the Dakotas indicated that the disease was encroaching on Montana's borders. Mule deer are native and common throughout Montana, and they represent a significant portion of the total hunter-harvested cervids in the state. The arrival of CWD in Montana may have significant ecosystem and socioeconomic impacts as well as potential consequences for wildlife management. We used 18,879 mule deer locations from 892 individual deer collected during 1975–2011 and modeled habitat selection for 7 herds in 5 of the 7 wildlife management regions in Montana. We estimated resource selection functions (RSF) in a Bayesian framework to predict summer and winter habitat preferences for mule deer. We estimated deer abundance from flyover counts for each region, and used the RSF predictions as weights to distribute the deer across the region. We then calculated the distance to the nearest known infected herds. We predicted areas of high risk of CWD infection in mule deer as areas with densities above the median density estimate and within the lowest quartile of distances to known infected herds. We identified these areas, the southeast corner of Montana and the north-central border near Alberta and Saskatchewan, as priority areas for CWD surveillance and management efforts.

Montana

White‐tailed deer habitat use and implications for chronic wasting disease transmission

Animal space use, activity patterns, and habitat selection—and heterogeneity in these patterns—have important implications for where and when infectious diseases are transmitted. White-tailed deer ( Odocoileus virginianus ) are habitat generalists, with a high degree of heterogeneity in their movement ecology based on sex, age, season, and region. These heterogeneities have important implications for the transmission and management of chronic wasting disease (CWD), which is a deadly prion disease transmitted both directly and indirectly through the environment. As such, favored deer habitats may promote direct interactions between conspecifics or indirect spatial overlap and subsequent environmental transmission. However, little is known about how individual animal space use translates to actual spatial overlap between individuals, leaving uncertainty in how habitat shapes the risk of direct or environmental CWD transmission. In this study, we evaluated seasonal activity patterns, home ranges, and habitat selection for 596 white-tailed deer in southwest Wisconsin, USA, from 2017-2022. We also estimated seasonal encounter distributions—regions where a pair of deer were most likely to encounter each other—for all pairs of deer putatively in different social groups (between-group) in our study, and quantified seasonal variation in the habitat composition of these areas. We found that deer selection for crops, pasture, or grasslands was generally low, relative to forest, but was highest in the post-fawning (summer) and non-breeding (winter) seasons. We observed similar patterns for the habitat composition of encounter distributions, suggesting that crops, pasture, and grasslands may be attractive resources that facilitate between-group transmission. Site fidelity between years was generally high; combined with small female home ranges in the fawning season, this implies that females likely re-use the same small, high-quality fawning habitats from year to year. We found that attraction toward between-group individuals was low during the post-fawning season but high during the breeding (fall) and non-breeding seasons. These results suggest that space use and habitat selection could shape the risk of environmental transmission in the fawning and post-fawning seasons, social selection could favor direct transmission risk in the breeding season, and combined social and habitat selection may shape risk of both direct and environmental transmission during the non-breeding season. We provide a detailed picture of the physiological and social drivers of deer movement through the year, with implications for CWD transmission and management.

Iowa, Wisconsin

Water masses, ocean fronts, and the structure of Antarctic seabird communities: putting the eastern Bellingshausen Sea in perspective

Waters off the western Antarctic Peninsula (i.e., the eastern Bellingshausen Sea) are unusually complex owing to the convergence of several major fronts. Determining the relative influence of fronts on occurrence patterns of top-trophic species in that area, therefore, has been challenging. In one of the few ocean-wide seabird data syntheses, in this case for the Southern Ocean, we analyzed ample, previously collected cruise data, Antarctic-wide, to determine seabird species assemblages and quantitative relationships to fronts as a way to provide context to the long-term Palmer LTER and the winter Southern Ocean GLOBEC studies in the eastern Bellingshausen Sea. Fronts investigated during both winter (April–September) and summer (October–March) were the southern boundary of the Antarctic Circumpolar Current (ACC), which separates the High Antarctic from the Low Antarctic water mass, and within which are embedded the marginal ice zone and Antarctic Shelf Break Front; and the Antarctic Polar Front, which separates the Low Antarctic and the Subantarctic water masses. We used clustering to determine species' groupings with water masses, and generalized additive models to relate species' densities, biomass and diversity to distance to respective fronts. Antarctic-wide, in both periods, highest seabird densities and lowest species diversity were found in the High Antarctic water mass. In the eastern Bellingshausen, seabird density in the High Antarctic water mass was lower (as low as half that of winter) than found in other Antarctic regions. During winter, Antarctic-wide, two significant species groups were evident: one dominated by Adélie penguins ( Pygoscelis adeliae ) (High Antarctic water mass) and the other by petrels and prions (no differentiation among water masses); in eastern Bellingshausen waters during winter, the one significant species group was composed of species from both Antarctic-wide groups. In summer, Antarctic-wide, a High Antarctic group dominated by Adélie penguins, a Low Antarctic group dominated by petrels, and a Subantarctic group dominated by albatross were evident. In eastern Bellingshausen waters during summer, groups were inconsistent. With regard to frontal features, Antarctic-wide in winter, distance to the ice edge was an important explanatory factor for nine of 14 species, distance to the Antarctic Polar Front for six species and distance to the Shelf Break Front for six species; however, these Antarctic-wide models could not successfully predict spatial relationships of winter seabird density (individual species or total) and biomass in the eastern Bellingshausen. Antarctic-wide in summer, distance to land/Antarctic continent was important for 10 of 18 species, not a surprising result for these summer-time Antarctic breeders, as colonies are associated with ice-free areas of coastal land. Distance to the Shelf Break Front was important for 8 and distance to the southern boundary of the ACC was important for 7 species. These summer models were more successful in predicting eastern Bellingshausen species density and species diversity but failed to predict total seabird density or biomass. Antarctic seabirds appear to respond to fronts in a way similar to that observed along the well-studied upwelling front of the California Current. To understand fully the seabird patterns found in this synthesis, multi-disciplinary at-sea investigations, including a quantified prey field, are needed.

Deep-Sea Research Part II: Topical Studies in Ocea

Genetic susceptibility to chronic wasting disease in free-ranging white-tailed deer: complement component C1q and Prnp polymorphisms

The genetic basis of susceptibility to chronic wasting disease (CWD) in free-ranging cervids is of great interest. Association studies of disease susceptibility in free-ranging populations, however, face considerable challenges including: the need for large sample sizes when disease is rare, animals of unknown pedigree create a risk of spurious results due to population admixture, and the inability to control disease exposure or dose. We used an innovative matched case&ndash;control design and conditional logistic regression to evaluate associations between polymorphisms of complement C1q and prion protein (Prnp) genes and CWD infection in white-tailed deer from the CWD endemic area in south-central Wisconsin. To reduce problems due to admixture or disease-risk confounding, we used neutral genetic (microsatellite) data to identify closely related CWD-positive ( n = 68) and CWD-negative ( n = 91) female deer to serve as matched cases and controls. Cases and controls were also matched on factors (sex, location, age) previously demonstrated to affect CWD infection risk. For Prnp, deer with at least one Serine (S) at amino acid 96 were significantly less likely to be CWD-positive relative to deer homozygous for Glycine (G). This is the first characterization of genes associated with the complement system in white-tailed deer. No tests for association between any C1q polymorphism and CWD infection were significant at p < 0.05. After controlling for Prnp, we found weak support for an elevated risk of CWD infection in deer with at least one Glycine (G) at amino acid 56 of the C1qC gene. While we documented numerous amino acid polymorphisms in C1q genes none appear to be strongly associated with CWD susceptibility.

Wisconsin

Association mapping of genetic risk factors for chronic wasting disease in wild deer

Chronic wasting disease (CWD) is a fatal transmissible spongiform encephalopathy affecting North American cervids. We assessed the feasibility of association mapping CWD genetic risk factors in wild white-tailed deer ( Odocoileus virginianus ) and mule deer ( Odocoileus hemionus ) using a panel of bovine microsatellite markers from three homologous deer linkage groups predicted to contain candidate genes. These markers had a low cross-species amplification rate (27.9%) and showed weak linkage disequilibrium (<1 cM). Markers near the prion protein and the neurofibromin 1 ( NF1 ) genes were suggestively associated with CWD status in white-tailed deer ( P = 0.006) and mule deer ( P = 0.02), respectively. This is the first time an association between the NF1 region and CWD has been reported.

Evolutionary Applications

Demographic patterns and harvest vulnerability of chronic wasting disease infected white-tailed deer in Wisconsin

Chronic wasting disease (CWD) is a fatal disease of white-tailed deer (Odocoileus virginianus) caused by transmissible protease-resistant prions. Since the discovery of CWD in southern Wisconsin in 2001, more than 20,000 deer have been removed from a >2,500-km2 disease eradication zone surrounding the three initial cases. Nearly all deer removed were tested for CWD infection and sex, age, and harvest location were recorded. Our analysis used data from a 310-km2 core study area where disease prevalence was higher than surrounding areas. We found no difference in harvest rates between CWD infected and noninfected deer. Our results show that the probability of infection increased with age and that adult males were more likely to be infected than adult females. Six fawns tested positive for CWD, five fawns from the core study area, including the youngest (5 months) free-ranging cervid to test positive. The increase in male prevalence with age is nearly twice the increase found in females. We concluded that CWD is not randomly distributed among deer and that differential transmission among sex and age classes is likely driving the observed patterns in disease prevalence. We discuss alternative hypotheses for CWD transmission and spread and, in addition, discuss several possible nonlinear relationships between prevalence and age. Understanding CWD transmission in free-ranging cervid populations will be essential to the development of strategies to manage this disease in areas where CWD is found, as well as for surveillance strategies in areas where CWD threatens to spread.

Journal of Wildlife Management

Deer carcass decomposition and potential scavenger exposure to chronic wasting disease

Chronic wasting disease (CWD) is a transmissible spongiform encephalopathy afflicting the Cervidae family in North America, causing neurodegeneration and ultimately death. Although there are no reports of natural cross-species transmission of CWD to noncervids, infected deer carcasses pose a potential risk of CWD exposure for other animals. We placed 40 disease-free white-tailed deer (Odocoileus virginianus) carcasses and 10 gut piles in the CWD-affected area of Wisconsin (USA) from September to April in 2003 through 2005. We used photos from remotely operated cameras to characterize scavenger visitation and relative activity. To evaluate factors driving the rate of carcass removal (decomposition), we used KaplanMeier survival analysis and a generalized linear mixed model. We recorded 14 species of scavenging mammals (6 visiting species) and 14 species of scavenging birds (8 visiting species). Prominent scavengers included American crows (Corvus brachyrhynchos), raccoons (Procyon lotor), and Virginia opossums (Didelphis virginiana). We found no evidence that deer consumed conspecific remains, although they visited gut piles more often than carcasses relative to temporal availability in the environment. Domestic dogs, cats, and cows either scavenged or visited carcass sites, which could lead to human exposure to CWD. Deer carcasses persisted for 18 days to 101 days depending on the season and year, whereas gut piles lasted for 3 days. Habitat did not influence carcass decomposition, but mammalian and avian scavenger activity and higher temperatures were positively associated with faster removal. Infected deer carcasses or gut piles can serve as potential sources of CWD prions to a variety of scavengers. In areas where surveillance for CWD exposure is practical, management agencies should consider strategies for testing primary scavengers of deer carcass material.

Journal of Wildlife Management