Steven N. Winter
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Previous Organizations / Institutes
The organizations / institutes from the scholar's other articles in the past ten years
Steven N. Winter has a total of 34 co-authors. Here are the top co-authors by number of shared publications:
- Margaret A. Wild
Department of Veterinary Microbiology and Pathology, College of Veterinary Medicine, Washington State University
4 shared publications - Erin Clancey
Paul G. Allen School for Global Health, Washington State University
2 shared publications - Luis Emilio Escobar
Department of Fish and Wildlife Conservation Virginia Tech Blacksburg Virginia USA
2 shared publications - María del Pilar Fernández
Paul G. Allen School for Global Health, Washington State University, Pullman, USA
2 shared publications A. Townsend Peterson
Biodiversity Institute and Department of Ecology and Evolutionary Biology The University of Kansas Lawrence KS, 66045 U.S.A
1 shared publicationCarrington Hilson
California Department of Fish and Wildlife, Northern Region
1 shared publication
Latest Publications
- Open Access
Use of Leg-Mounted Monitors to Assess the Effects of Treponeme-Associated Hoof Disease on Elk (Cervus canadensis) Activity
Treponeme-associated hoof disease (TAHD) is an emerging disease of free-ranging elk (Cervus canadensis) in the northwestern United States. Affected elk develop chronic foot lesions, lameness, debilitation, and an apparent increase in mortality, but the onset of lameness and associated changes in activity are not fully understood. We evaluated the accuracy of a newly developed leg-mounted tri-axial accelerometer monitor (Advanced Telemetry Systems) on captive elk and collected monitor-derived data to assess activity before and during an experimental TAHD challenge. Monitors provided reliable data with 85% overall accuracy of the continuous onboard classification of activity as standing, moving, or bedded against direct visual observation using seven healthy elk. Further, following TAHD challenge, monitor-derived data were able to detect that treatment elk exhibiting abnormal locomotion spent more time bedded and less time moving or standing. During the challenge period, treatment elk spent roughly 10% more of the day bedded than control elk. These findings suggest that leg-mounted activity monitors can detect changes in elk activity and may serve as a useful tool for future wildlife disease monitoring efforts.
Open AccessUse of Leg-Mounted Monitors to Assess the Effects of Treponeme-Associated Hoof Disease on Elk (Cervus canadensis) Activity
Treponeme-associated hoof disease (TAHD) is an emerging disease of free-ranging elk (Cervus canadensis) in the northwestern United States. Affected elk develop chronic foot lesions, lameness, debilitation, and an apparent increase in mortality, but the onset of lameness and associated changes in activity are not fully understood. We evaluated the accuracy of a newly developed leg-mounted tri-axial accelerometer monitor (Advanced Telemetry Systems) on captive elk and collected monitor-derived data to assess activity before and during an experimental TAHD challenge. Monitors provided reliable data with 85% overall accuracy of the continuous onboard classification of activity as standing, moving, or bedded against direct visual observation using seven healthy elk. Further, following TAHD challenge, monitor-derived data were able to detect that treatment elk exhibiting abnormal locomotion spent more time bedded and less time moving or standing. During the challenge period, treatment elk spent roughly 10% more of the day bedded than control elk. These findings suggest that leg-mounted activity monitors can detect changes in elk activity and may serve as a useful tool for future wildlife disease monitoring efforts.
- Open Access
Land use and soil characteristics are associated with increased risk of treponeme‐associated hoof disease in elk
Environments can shape the occurrence and extent of disease outbreaks in wildlife. We studied the effects of environmental features on the occurrence of treponeme‐associated hoof disease (TAHD), an emerging infectious disease of free‐ranging elk ( Cervus canadensis ), in southwestern Washington, USA. During the 2016–2022 harvest seasons, successful elk hunters returned mandatory harvest reports and noted the presence or absence of hoof abnormalities indicative of TAHD. We used generalized linear models and an information‐theoretic approach to model selection to relate (1) the spatial distribution of hoof abnormalities to features of landscapes (land cover, topography, and soil characteristics) and (2) the temporal distribution of hoof abnormalities to precipitation during the year preceding the harvest season. The probability of hoof disease increased with soil clay content and proportion of agricultural land (88% of model weight). We found no conclusive evidence for an effect of precipitation on the occurrence of TAHD, but this could relate to relatively high annual precipitation (>140 cm) in the study area. Nevertheless, disease cases may have been negatively associated with precipitation during February–June (55% of model weight). Soils and land management practices may increase the risk of hoof disease by promoting the survival of pathogens that cause TAHD, the susceptibility of elk to infection, or the intensity of pathogen transmission among elk when congregated. Focusing on areas where the risk of disease is greatest may facilitate the detection of TAHD during surveillance. Likewise, removing infected elk and dispersing uninfected elk from areas with the greatest risk of disease may enhance the effectiveness of efforts to reduce transmission. Basing this work on the knowledge that disease risk is modified by factors of hosts, pathogens, and environments, this study serves as an application of the epidemiological triad framework to better understand the ecology and epidemiology of an emerging infectious disease in wildlife.
Open AccessLand use and soil characteristics are associated with increased risk of treponeme‐associated hoof disease in elk
Environments can shape the occurrence and extent of disease outbreaks in wildlife. We studied the effects of environmental features on the occurrence of treponeme‐associated hoof disease (TAHD), an emerging infectious disease of free‐ranging elk ( Cervus canadensis ), in southwestern Washington, USA. During the 2016–2022 harvest seasons, successful elk hunters returned mandatory harvest reports and noted the presence or absence of hoof abnormalities indicative of TAHD. We used generalized linear models and an information‐theoretic approach to model selection to relate (1) the spatial distribution of hoof abnormalities to features of landscapes (land cover, topography, and soil characteristics) and (2) the temporal distribution of hoof abnormalities to precipitation during the year preceding the harvest season. The probability of hoof disease increased with soil clay content and proportion of agricultural land (88% of model weight). We found no conclusive evidence for an effect of precipitation on the occurrence of TAHD, but this could relate to relatively high annual precipitation (>140 cm) in the study area. Nevertheless, disease cases may have been negatively associated with precipitation during February–June (55% of model weight). Soils and land management practices may increase the risk of hoof disease by promoting the survival of pathogens that cause TAHD, the susceptibility of elk to infection, or the intensity of pathogen transmission among elk when congregated. Focusing on areas where the risk of disease is greatest may facilitate the detection of TAHD during surveillance. Likewise, removing infected elk and dispersing uninfected elk from areas with the greatest risk of disease may enhance the effectiveness of efforts to reduce transmission. Basing this work on the knowledge that disease risk is modified by factors of hosts, pathogens, and environments, this study serves as an application of the epidemiological triad framework to better understand the ecology and epidemiology of an emerging infectious disease in wildlife.
- Open Access
- 19 November 2025
- Published by American Society for Microbiology in Applied and Environmental Microbiology
- Vol. 91 (11) , e0088825
Bacterial community analysis of treponeme-associated hoof disease in free-ranging elk ( Cervus canadensis ): evidence for a polybacterial etiology with geographic consistency
Treponeme-associated hoof disease (TAHD) is an emerging infectious disease in free-ranging elk ( Cervus canadensis ) characterized by ulcerative and necrosuppurative pododermatitis with spirochete bacteria that leads to lameness and apparent increased mortality. While TAHD is hypothesized to have a polybacterial etiology, the causative agents remain poorly characterized, particularly across its geographic range. In this study, we developed a histologic categorization system for hoof lesions and employed 16S rRNA gene amplicon sequencing to characterize bacterial communities in samples from 129 free-ranging elk across regions with endemic or sporadic TAHD and where TAHD remains undetected. Differential abundance analysis revealed strong associations between TAHD-positive lesions and the bacterial genera Treponema , unidentified Spirochaetaceae , Mycoplasma , and Fusobacterium , along with their respective families and amplicon sequence variants. Many of these TAHD-associated operational taxonomic units (OTUs) were also more frequently detected at increased abundance in more severe, histologic lesions of pododermatitis. Correlation analysis demonstrated a strong positive association between Treponema and Mycoplasma in TAHD lesions, suggesting a more significant role of Mycoplasma in TAHD’s etiology than previously recognized. Additionally, we identified novel TAHD-associated OTUs, including Corynebacterium freneyi-xerosis , that warrant further investigation. Comparative analysis of TAHD-positive lesions from endemic and sporadic areas revealed minimal differences in the microbial community. These findings advance our understanding of the bacterial contributors to TAHD, highlighting putative pathogens for further investigation and as potential targets for diagnostic development. IMPORTANCE: While detection of Treponema is a hallmark of treponeme-associated hoof disease (TAHD), a comprehensive understanding of other bacterial contributors is necessary to improve diagnostic testing and inform control measures. Our study confirmed strong associations between Spirochaetaceae and TAHD lesions and revealed a previously underappreciated role of Mycoplasma in TAHD’s etiology. Treponema and Mycoplasma were significantly enriched in TAHD-positive lesions, absent from TAHD-negative tissues, and strongly and positively correlated with each other, suggesting a potential synergistic relationship. By developing and applying a histologic categorization system, we characterized shifts in bacterial communities as lesion severity progressed. Comparisons of TAHD-positive lesions from endemic and sporadic regions revealed minimal differences in the microbial composition, indicating strong geographic consistency. These findings enhance our understanding of TAHD’s etiology and provide a foundation for future research, including the development of improved diagnostic tests and targeted disease management strategies.
- 19 November 2025
- Published by American Society for Microbiology in Applied and Environmental Microbiology
- Vol. 91 (11) , e0088825
Open AccessBacterial community analysis of treponeme-associated hoof disease in free-ranging elk ( Cervus canadensis ): evidence for a polybacterial etiology with geographic consistency
Treponeme-associated hoof disease (TAHD) is an emerging infectious disease in free-ranging elk ( Cervus canadensis ) characterized by ulcerative and necrosuppurative pododermatitis with spirochete bacteria that leads to lameness and apparent increased mortality. While TAHD is hypothesized to have a polybacterial etiology, the causative agents remain poorly characterized, particularly across its geographic range. In this study, we developed a histologic categorization system for hoof lesions and employed 16S rRNA gene amplicon sequencing to characterize bacterial communities in samples from 129 free-ranging elk across regions with endemic or sporadic TAHD and where TAHD remains undetected. Differential abundance analysis revealed strong associations between TAHD-positive lesions and the bacterial genera Treponema , unidentified Spirochaetaceae , Mycoplasma , and Fusobacterium , along with their respective families and amplicon sequence variants. Many of these TAHD-associated operational taxonomic units (OTUs) were also more frequently detected at increased abundance in more severe, histologic lesions of pododermatitis. Correlation analysis demonstrated a strong positive association between Treponema and Mycoplasma in TAHD lesions, suggesting a more significant role of Mycoplasma in TAHD’s etiology than previously recognized. Additionally, we identified novel TAHD-associated OTUs, including Corynebacterium freneyi-xerosis , that warrant further investigation. Comparative analysis of TAHD-positive lesions from endemic and sporadic areas revealed minimal differences in the microbial community. These findings advance our understanding of the bacterial contributors to TAHD, highlighting putative pathogens for further investigation and as potential targets for diagnostic development. IMPORTANCE: While detection of Treponema is a hallmark of treponeme-associated hoof disease (TAHD), a comprehensive understanding of other bacterial contributors is necessary to improve diagnostic testing and inform control measures. Our study confirmed strong associations between Spirochaetaceae and TAHD lesions and revealed a previously underappreciated role of Mycoplasma in TAHD’s etiology. Treponema and Mycoplasma were significantly enriched in TAHD-positive lesions, absent from TAHD-negative tissues, and strongly and positively correlated with each other, suggesting a potential synergistic relationship. By developing and applying a histologic categorization system, we characterized shifts in bacterial communities as lesion severity progressed. Comparisons of TAHD-positive lesions from endemic and sporadic regions revealed minimal differences in the microbial composition, indicating strong geographic consistency. These findings enhance our understanding of TAHD’s etiology and provide a foundation for future research, including the development of improved diagnostic tests and targeted disease management strategies.
- Open Access
- 14 December 2023
- Published by Wiley in Transboundary and Emerging Diseases
- Vol. 2023, 1-14
Community Science Strategies Reveal Distributional Patterns of Treponeme-Associated Hoof Disease in Washington Elk (Cervus canadensis)
Treponeme-associated hoof disease (TAHD) is an emerging disease of conservation concern in elk (Cervus canadensis) in the Northwest USA. Elk with TAHD exhibit characteristic hoof lesions that are often accompanied by lameness and limping. The gold standard approach traditionally used for infectious disease surveillance is laboratory confirmation, which for TAHD is a histologic examination of abnormal elk hooves submitted by wildlife agencies. Diagnostic evaluation affords certainty in confirming TAHD; however, these examinations are also labor and resource intensive, and therefore, not conducive to the collection of sufficient data for epidemiologic investigations. In response, two community science (CS) surveillance strategies have been implemented in Washington State: public observations of limping elk from a web-based reporting tool and hunter reports of hoof abnormalities on harvested elk. Surveillance using CS strategies can be implemented widely and may be useful for describing broad distributional patterns of TAHD, despite their unknown relationship to laboratory-confirmed cases. We described and compared the spatialtemporal distribution of TAHD in western Washington game management units (GMU) using the two CS strategies to assess congruences and discrepancies between observed patterns. We used spatial scan statistics to identify possible core-affected and newly emerging areas at the GMU level. Lastly, we contrasted CS observations against confirmed case data to examine possible delays in TAHD detection and co-occurrence among surveillance strategies. We found public observations of limping elk often predated TAHD confirmations in GMUs by several years, while hunter-reported abnormalities predated confirmations in GMUs by several months. High co-occurrence between the presence of apparent and confirmed cases under different surveillance strategies further supports the use of CS sources. This study capitalizes on wide-reaching CS data to provide new and complementary epidemiological information that can help guide future surveillance, management, and research efforts for this novel elk hoof disease.
- 14 December 2023
- Published by Wiley in Transboundary and Emerging Diseases
- Vol. 2023, 1-14
Open AccessCommunity Science Strategies Reveal Distributional Patterns of Treponeme-Associated Hoof Disease in Washington Elk (Cervus canadensis)
Treponeme-associated hoof disease (TAHD) is an emerging disease of conservation concern in elk (Cervus canadensis) in the Northwest USA. Elk with TAHD exhibit characteristic hoof lesions that are often accompanied by lameness and limping. The gold standard approach traditionally used for infectious disease surveillance is laboratory confirmation, which for TAHD is a histologic examination of abnormal elk hooves submitted by wildlife agencies. Diagnostic evaluation affords certainty in confirming TAHD; however, these examinations are also labor and resource intensive, and therefore, not conducive to the collection of sufficient data for epidemiologic investigations. In response, two community science (CS) surveillance strategies have been implemented in Washington State: public observations of limping elk from a web-based reporting tool and hunter reports of hoof abnormalities on harvested elk. Surveillance using CS strategies can be implemented widely and may be useful for describing broad distributional patterns of TAHD, despite their unknown relationship to laboratory-confirmed cases. We described and compared the spatialtemporal distribution of TAHD in western Washington game management units (GMU) using the two CS strategies to assess congruences and discrepancies between observed patterns. We used spatial scan statistics to identify possible core-affected and newly emerging areas at the GMU level. Lastly, we contrasted CS observations against confirmed case data to examine possible delays in TAHD detection and co-occurrence among surveillance strategies. We found public observations of limping elk often predated TAHD confirmations in GMUs by several years, while hunter-reported abnormalities predated confirmations in GMUs by several months. High co-occurrence between the presence of apparent and confirmed cases under different surveillance strategies further supports the use of CS sources. This study capitalizes on wide-reaching CS data to provide new and complementary epidemiological information that can help guide future surveillance, management, and research efforts for this novel elk hoof disease.
- Open Access
- 18 January 2023
- Published by IntechOpen
Drugs and Biodiversity Loss: Narcotraffic-Linked Landscape Change in Guatemala
Characteristic of the Anthropocene, human impacts have resulted in worldwide losses in forested land cover, which can directly and indirectly drive biodiversity loss. The global illicit drug trade is one source of deforestation directly implicated with habitat loss in Central America, typically for drug trafficking and livestock production for money laundering. Given reports of deforestation in Central America linked to narcotraffic, we explored vegetation changes within Guatemala’s highly biodiverse Maya Biosphere Reserve by examining trends suggestive of deforestation in a protected area. As such, we collected satellite-derived data in the form of enhanced vegetation index (EVI), as well as history of burned areas, published human-“footprint” data, official population density, and artificial light activity in Laguna del Tigre National Park from 2002 to 2020 for descriptive analysis. We found consistent reductions in EVI and trends of anomalous losses of vegetation despite a baseline accounting for variation within the park. Analyses revealed weak correlations (R2 ≤ 0.26) between EVI losses and official sources of anthropogenic data, which may be attributable to the data’s limited spatial and temporal resolution. Alarmingly, simple analyses identified vegetation losses within a protected area, thus emphasizing the need for additional monitoring and science-based, but interdisciplinary policies to protect this biodiversity hotspot.
- 18 January 2023
- Published by IntechOpen
Open AccessDrugs and Biodiversity Loss: Narcotraffic-Linked Landscape Change in Guatemala
Characteristic of the Anthropocene, human impacts have resulted in worldwide losses in forested land cover, which can directly and indirectly drive biodiversity loss. The global illicit drug trade is one source of deforestation directly implicated with habitat loss in Central America, typically for drug trafficking and livestock production for money laundering. Given reports of deforestation in Central America linked to narcotraffic, we explored vegetation changes within Guatemala’s highly biodiverse Maya Biosphere Reserve by examining trends suggestive of deforestation in a protected area. As such, we collected satellite-derived data in the form of enhanced vegetation index (EVI), as well as history of burned areas, published human-“footprint” data, official population density, and artificial light activity in Laguna del Tigre National Park from 2002 to 2020 for descriptive analysis. We found consistent reductions in EVI and trends of anomalous losses of vegetation despite a baseline accounting for variation within the park. Analyses revealed weak correlations (R2 ≤ 0.26) between EVI losses and official sources of anthropogenic data, which may be attributable to the data’s limited spatial and temporal resolution. Alarmingly, simple analyses identified vegetation losses within a protected area, thus emphasizing the need for additional monitoring and science-based, but interdisciplinary policies to protect this biodiversity hotspot.
- 1 October 2020
- Published by Wildlife Disease Association in Journal of Wildlife Diseases
- Vol. 56 (4) , 741-758
CHRONIC WASTING DISEASE MODELING: AN OVERVIEW
Chronic wasting disease (CWD) is an infectious and fatal prion disease occurring in the family Cervidae. To update the research community regarding the status quo of CWD epidemic models, we conducted a meta-analysis on CWD research. We collected data from peer-reviewed articles published since 1980, when CWD was first diagnosed, until December 2018. We explored the analytical methods used historically to understand CWD. We used 14 standardized variables to assess overall analytical approaches of CWD research communities, data used, and the modeling methods used. We found that CWD modeling initiated in the early 2000s and has increased since then. Connectivity of the research community was heavily reliant on a cluster of CWD researchers. Studies focused primarily on regression and compartment-based models, population-level approaches, and host species of game management concern. Similarly, CWD research focused on single populations, species, and locations, neglecting modeling using community ecology and biogeographic approaches. Chronic wasting disease detection relied on classic diagnostic methods with limited sensitivity for most stages of infection. Overall, we found that past modeling efforts generated a solid baseline for understanding CWD in wildlife and increased our knowledge on infectious prion ecology. Future analytical efforts should consider more sensitive diagnostic methods to quantify uncertainty and broader scale studies to elucidate CWD transmission beyond population-level approaches. Considering that infectious prions may not follow biological rules of well-known wildlife pathogens (i.e., viruses, bacteria, fungi), assumptions used when modeling other infectious disease may not apply for CWD. Chronic wasting disease is a new challenge in wildlife epidemiology.
- 1 October 2020
- Published by Wildlife Disease Association in Journal of Wildlife Diseases
- Vol. 56 (4) , 741-758
CHRONIC WASTING DISEASE MODELING: AN OVERVIEW
Chronic wasting disease (CWD) is an infectious and fatal prion disease occurring in the family Cervidae. To update the research community regarding the status quo of CWD epidemic models, we conducted a meta-analysis on CWD research. We collected data from peer-reviewed articles published since 1980, when CWD was first diagnosed, until December 2018. We explored the analytical methods used historically to understand CWD. We used 14 standardized variables to assess overall analytical approaches of CWD research communities, data used, and the modeling methods used. We found that CWD modeling initiated in the early 2000s and has increased since then. Connectivity of the research community was heavily reliant on a cluster of CWD researchers. Studies focused primarily on regression and compartment-based models, population-level approaches, and host species of game management concern. Similarly, CWD research focused on single populations, species, and locations, neglecting modeling using community ecology and biogeographic approaches. Chronic wasting disease detection relied on classic diagnostic methods with limited sensitivity for most stages of infection. Overall, we found that past modeling efforts generated a solid baseline for understanding CWD in wildlife and increased our knowledge on infectious prion ecology. Future analytical efforts should consider more sensitive diagnostic methods to quantify uncertainty and broader scale studies to elucidate CWD transmission beyond population-level approaches. Considering that infectious prions may not follow biological rules of well-known wildlife pathogens (i.e., viruses, bacteria, fungi), assumptions used when modeling other infectious disease may not apply for CWD. Chronic wasting disease is a new challenge in wildlife epidemiology.
- 21 November 2019
- Published by Wiley in Biological Reviews
- Vol. 95 (2) , 393-408
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.
- 21 November 2019
- Published by Wiley in Biological Reviews
- Vol. 95 (2) , 393-408
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.