Evaluating Factors Influencing Elk Recruitment in Colorado
Led By: Nathaniel Rayl, Mat Alldredge, and Chuck Anderson Study Areas Bear’s Ears elk herd (DAU E-2) Trinchera elk herd (DAU E-33) Uncompahgre Plateau elk herd (DAU E-20) Project Status Ongoing Research Objectives To estimate calf and adult female survival and cause-specific mortality rates To quantify pregnancy rates and nutritional condition To evaluate factors influencing survival, pregnancy rates, and nutritional condition Project Description In Colorado, elk are an important natural resource valued for ecological, consumptive, aesthetic, and economic reasons. In 1910, Colorado had less than 1,000 elk; today, the state population is estimated to be the largest in the country, with more than 290,000 elk. Over the last two decades, however, Colorado Parks and Wildlife has become increasingly concerned about declining calf recruitment (the number of calves making it to adulthood). In 2017, researchers began a 2-year pilot study to investigate factors influencing elk recruitment in two study areas in the state. The initial pilot project was expanded into a 3rd study area in 2019 and the research continues to better understand elk population dynamics in Colorado and determine how predators, habitat, and weather conditions are impacting elk recruitment in Colorado. Reports: Pilot Study: Elk Recruitment and Habitat Use in Colorado (2017-2018) Evaluating Factors Influencing Elk Recruitment in Colorado (2019-current) Publications: Crews, S., N.D. Rayl, M.W. Alldredge, E.J. Bergman, C.R. Anderson Jr., E.H. VanNatta, J.D. Holbrook, and G. Bastille-Rousseau. 2025. Hierarchy in structuring of resource selection: understanding elk selection across space, time, and movement strategies. Ecology and Evolution 15:e71097. Crews, S., N.D. Rayl, M.W. Alldredge, E.J. Bergman, C.R. Anderson Jr., and G. Bastille-Rousseau. 2025. Drivers of spring migration phenology in Rocky Mountain elk. Scientific Reports 15:7807.
Rayl, Nathaniel
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Response of Elk to Human Recreation at Multiple Scales
Led By Eric Bergman, Nathaniel Rayl Study Areas Avalanche Creek elk herd (DAU E-15) Bear’s Ears elk herd (DAU E-2) Project Status: Ongoing Research Objectives To assess how elk respond to human recreation To evaluate the influence of human recreation on elk distribution To estimate calf and adult female survival and cause-specific mortality rates To quantify pregnancy rates and nutritional condition To evaluate factors influencing survival, pregnancy rates, and nutritional condition Project Description In Colorado, outdoor recreation contributes 511,000 jobs, $62.5 billion in economic output, and $9.4 billion in local, state, and federal tax revenue. Thus, the economies of Colorado, its counties, and its communities, rely on managing the landscape for a multitude of outdoor recreational opportunities. The sensitivity of elk to human presence and human activity has been a topic of interest for many decades. In Colorado, increasing public concerns over human recreational use have coincided with declines in elk productivity, but a direct relationship to this activity in Colorado remains unaddressed. This research aims to better understand the influence of human recreation on elk behavior and distribution. Reports Response of Elk To Human Recreation at Multiple Scales: Demographic Shifts and Behaviorally Mediated Fluctuations in Local Abundance (2019-Current) Publications Crews, S., N.D. Rayl, M.W. Alldredge, E.J. Bergman, C.R. Anderson Jr., E.H. VanNatta, J.D. Holbrook, and G. Bastille-Rousseau. 2025. Hierarchy in structuring of resource selection: understanding elk selection across space, time, and movement strategies. Ecology and Evolution 15:e71097. Crews, S., N. D. Rayl, M. W. Alldredge, E. J. Bergman, C. R. Anderson Jr., and G. Bastille-Rousseau. 2025. Drivers of spring migration phenology in Rocky Mountain elk. Scientific Reports 15:7807 Bastille-Rousseau, G., S. A. Crews, E. B. Donovan, M. E. Egan, N. T. Gorman, J. B. Pitman, A. M. Weber, E. M. Audia, M. R. Larreur, H. Manninen, S. Blake, M. W. Eihholz, E. Bergman, and N. D. Rayl. 2024. A multi‐property assessment of intensity of use provides a functional understanding of animal movement. Methods in Ecology and Evolution 15:345-357. Egan, M. E., N. T. Gorman, S. Crews, M. W. Eichholz, D. Skinner, P. E. Schlichting, N. D. Rayl, E. J. Bergman, E. H. Ellington, and G. Bastille-Rousseau. 2024. Estimating encounter-habitat relationships with scale-integrated resource selection functions. Journal of Animal Ecology 93:1036-1048.
Spatiotemporal Effects of Human Recreation on Elk Behavior
Led by Eric VanNatta, Nathaniel Rayl, Eric Bergman, Joe Holbrook Study Areas Bear’s Ears elk herd (DAU E-2) Project Status Ongoing Research Objectives To estimate the distribution of human recreation during four time periods relevant to adult female elk: calving, calf-rearing, hunting, and post-hunting To evaluate whether adult female elk alter their movements and habitat selection when exposed to different intensities of human recreation. Project Description The influence of recreational disturbance on ungulate populations is of particular interest to wildlife managers in Colorado, as there is growing concern about its potential impacts within the state. Currently, the western United States is experiencing some of the highest rates of human population growth in the country, with growth in rural and exurban areas frequently outpacing growth in urban areas. Understanding potential impacts of recreational activity on elk spatial ecology in Colorado is critical for guiding management actions, as altered movements may result in reduced foraging time and higher energetic costs, which may decrease fitness. This research will evaluate potential impacts of recreational activities on elk from the resident portion of the Bear’s Ears elk herd in Colorado. Results from this research will be used to develop management recommendations to mitigate any negative effects of recreational activity on elk. Reports Spatiotemporal Effects of Human Recreation on Elk Behavior: an Assessment Within Critical Time Stages (2019-current) Publications Crews, S., N.D. Rayl, M.W. Alldredge, E.J. Bergman, C.R. Anderson Jr., E.H. VanNatta, J.D. Holbrook, and G. Bastille-Rousseau. 2025. Hierarchy in structuring of resource selection: understanding elk selection across space, time, and movement strategies. Ecology and Evolution 15:e71097.
Effects of Habitat Treatments on Mule Deer Survival and Health During the Winter
Led By: Eric Bergman Study Area: Uncompahgre Plateau and adjacent valleys in southwest Colorado Project Status: Completed Research Objectives: To evaluate the impacts of large-scale vegetation habitat treatment on mule deer survival and health during the winter. To develop habitat management strategies for the benefit of deer To put research results into context with historical records and recent declines in mule deer herd populations. Project Description: Mule deer populations in Colorado have shown dramatic fluctuations since the 1960s. Beginning in the 1990s, some western Colorado mule deer herds demonstrated the most recent decline. For many decades, wildlife managers have used habitat management as a way to increase the health and survival of mule deer, but the effect of these efforts had not yet been evaluated. Evaluating these efforts was the focus of this 4-year study to measure the health and survival of mule deer herds during the winter months in areas treated with different levels of habitat management. In the course of this study, researchers monitored fawn survival, mule deer body conditions, and population densities in areas with different levels of habitat management. Habitat treatment varied from mechanical disturbance in the form of hydro-axe clearing and roller-chopping of late seral stage pinyon-juniper forests, chemical control of weeds, and reseeding with desirable mule deer browse species. The results of this study will help managers select the most effective habitat treatments to apply to mule deer winter ranges. Associated Publications Bergman, E.J., C.J. Bishop, D.J. Freddy, G.C. White and P.F. Doherty. 2014. Habitat management influence overwinter survival of mule deer fawns in Colorado. Journal of Wildlife Management 78(3):448–455. Bergman, E.J. 2013. Evaluation of winter range habitat treatments on overwinter survival, density, and body condition of mule deer. Dissertation, Colorado State University, Fort Collins, Colorado, USA, 137pp. Watkins, B.E., C.J. Bishop, E.J. Bergman, B. Hale, B.F. Wakeling, A. Bronson, L.H. Carpenter, and D.W. Lutz. 2007. Habitat guidelines for mule deer: Colorado Plateau shrubland and forest ecoregion. Mule Deer Working Group, Western Association of Fish and Wildlife Agencies, Cheyenne, Wyoming, USA. 72pp. Bergman, E.J., P.F. Doherty, G.C. White, and D.J. Freddy. 2014. Habitat and herbivore density: response of mule deer to habitat management. The Journal of Wildlife Management 79:60–68. Bergman, E.J., P.F. Doherty, G.C. White, and A.A. Holland. 2015. Density dependence in mule deer: a review of evidence. Wildlife Biology 21:18–29. Bergman, E.J., P.F. Doherty, C.J. Bishop, L.L. Wolfe, and B. Banulis. 2014. Herbivore body condition response in altered environments: mule deer and habitat management. PloS one 9: e106374. Resources West Slope Mule Deer Strategy
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Ellen Brandell, Wildlife Research Scientist
Current or Recent Research Projects Wolf ecology Wolf habitat selection and movement during re establishment in Colorado Wildlife management Statistical estimation of wolf abundance Quantifying the effects of management practices and disease on wildlife harvest Wolf-livestock dynamics Identifying and examining potential nonconsumptive effects of wolves on cattle Predator-prey dynamics Examining wolf and hunter effects on elk movement, space use, and aggregation patterns Quantifying conditions under which predators influence prey diseases​ Areas of Interest and Expertise​ My background is in terrestrial wildlife ecology with a focus on carnivores (primarily gray wolves) and infectious diseases. I am interested in many facets of wildlife ecology, especially social behaviors, predator-prey interactions, population dynamics, and disease dynamics. My work intersects wildlife ecology, wildlife management, statistics, and disease ecology.​ Select Publications Draper, J. P., E. E. Brandell, J. Isabelle, C. Jacques, C. McCoy, E. Michel, D. J. Storm, C. Ott-Conn, B. Wojcik, W. C. Turner, and D. P. Walsh. 2025. Doe diligence: a regional analysis of antlerless deer harvest regulations in the Midwestern United States of America. PLoS One, 20(6): p.e0324708. https://doi.org/10.1371/journal.pone.0324708 Inzalaco, H. N. t., E. E. Brandell, S. P. Wilson, M. Hunsaker, D. R. Stahler, K. Woelfel, D. P. Walsh, T. Nordeen, D. J. Storm, and S. S. Lichtenberg. 2024. Detection of prions from spiked and free-ranging carnivore feces. Scientific Reports 14:3804. doi.org/10.1038/s41598-023-44167-7 Cubaynes, S., E. E. Brandell, D. R. Stahler, D. W. Smith, E. S. Almberg, S. Schindler, R. K. Wayne, A. P. Dobson, B. M. vonHoldt, D. R. MacNulty, P. D. Cross, P. J. Hudson, and T. Coulson. 2022. Disease outbreaks select for mate choice and coat color in wolves. Science 378:300–303; DOI: 10.1126/science.abi8745 Meyer, C. J., K. A. Cassidy, E. E. Stahler, E. E. Brandell, C. B. Anton, D. R. Stahler, and D. W. Smith 2022. Parasitic infection increases risk-taking in a social, intermediate host carnivore. Communications Biology 5:1–10; doi.org/10.1038/s42003-022-04122-0 Brandell, E. E., M. K. Jackson, P. C. Cross, A. J. Piaggio, D. R. Taylor, D. W. Smith, B. Boufana, D. R. Stahler, and P. J. Hudson. 2022. Evaluating noninvasive methods for estimating cestode prevalence in a wild carnivore population. PLoS ONE 17(11): e0277420; doi.org/10.1371/journal.pone.0277420 Brandell, E. E., D. J. Storm, T. R. Van Deelen, D. P. Walsh, and W. C. Turner. 2022. A call to action: standardizing white-tailed deer harvest data in the Midwestern United States and implicationsfor quantitative analysis and disease management. Frontiers in Ecology and Evolution; https://doi.org/10.3389/fevo.2022.943411 Gilbertson, M. L., E. E. Brandell, M. E. Pinkerton, N. M. Meaux, M. Hunsaker, D. Jarosinski, W. Ellarson, D. P. Walsh, D. J. Storm, and W. C. Turner. 2022. Cause of death, pathology, and chronic wasting disease status of white-tailed deer (Odocoileus virginianus) mortalities in Wisconsin, USA. Journal of Wildlife Diseases 54:803–815; DOI: 10.7589/JWD-D-21-00202​​ ​Brandell, E. E., P. C. Cross, D. W. Smith, W. Rogers, N. L. Galloway, D. MacNulty, D. R. Stahler, J. Treanor, and P. J. Hudson. 2022. Examination of the interaction between age‐specific predation and chronic disease in the Greater Yellowstone Ecosystem​. Journal of Animal Ecology. 91:1373-1384, https://doi.org/10.1111/1365-2656.13661​ Rogers, W., E. E. Brandell, and P. C. Cross. 2022. Epidemiological differences between sexes affect management efficacy in simulated chronic wasting disease systems. Journal of Applied Ecology 59:1122-1133. https://doi.org/10.1111/1365-2664.14125​ Brandell, E.E., D. J. Becker, L. Sampson, and K. M. Forbes. 2021. Demography, education, and research trends in the interdisciplinary field of disease ecology. Ecology and evolution 11:17581-17592, https://doi.org/10.1002/ece3.8466​ Brandell, E. E., A. P. Dobson, P. J. Hudson, P. C. Cross, and D. W. Smith. 2021. A metapopulation model of social group dynamics and disease applied to Yellowstone wolves. Proceedings of the National Academy of Sciences, 118(10), https://doi.org/10.1073/pnas.2020023118 Brandell, E.E., P. C. Cross, M. E. Craft, D. W. Smith, E. J. Dubovi, M. L. Gilbertson, T. Wheeldon, J. A. Stephenson, S. Barber-Meyer, B. L. Borg, and M. Sorum. 2021. Patterns and processes of pathogen exposure in gray wolves across North America​​. Scientific reports 11(1):1-14.​ DeCandia, A. L., E. C. Schrom, E. E. Brandell, D. R. Stahler, and B. M. vonHoldt. 2021. Sarcoptic mange severity is associated with reduced genomic variation and evidence of selection in Yellowstone National Park wolves (Canis lupus). Evolutionary applications, 14(2):429-445, https://doi.org/10.1111/eva.13127 Brandell, E. E., N. M. Fountain‐Jones, M. L. Gilbertson, P. C. Cross, P. J. Hudson, D. W. Smith, D. R. Stahler, C. Packer, and M. E. Craft. 2021. Group density, disease, and season shape territory size and overlap of social carnivores. Journal of Animal Ecology, 90(1):87-101. Education Ph.D., Ecology – Pennsylvania State University, 2021 B.S., Wildlife Biology –​ University of Montana, 2015 Current or Recent Positions Wildlife Research Scientist – Colorado Parks and Wildlife, 2022-Present Postdoctoral Associate –​ Wisconsin Cooperative Wildlife Research Unit, Dept. of Forest and Wildlife Ecology, University of Wisconsin-Madison, 2020-2022 Contact Information 317 W. Prospect Rd.​ Fort Collins, CO 80526 Email: ellen.brandell@stat​e.co.us​ Phone: ​970-698-0252
Shane Frank, Wildlife Research Scientist
Current or Recent Research Projects Bobcat Ecology and Management Bobcat ecology Where do bobcats occur? Determining where bobcats select habitats and resources What do bobcats eat? Spatial and temporal variation in prey availability Determining dietary composition and prey use/selection Bobcat monitoring and management How many bobcats are out there? Evaluating statistical methods for estimating bobcat abundance/density What are causes of bobcat mortality? Estimating the relative magnitude of mortality sources Bobcat reproduction Spatial and temporal variation in bobcat family group occurrence Understanding changes in bobcat population size Determining potential important factors affecting bobcat population density and dynamics (e.g. habitat, prey availability, and harvest rate) Improving bobcat data collection methods and population inference Enhancing age classification through dentition and teeth measurements Exploring cutting-edge methods in population size/density estimation Areas of Interest and Expertise​ My background is in terrestrial wildlife ecology with a focus on carnivores (primarily bears and bobcats) and harvest, though my interests vary widely. I have interest in exploring both the lives and deaths of wildlife and how individual contexts scale up to have population level effects. This encapsulates both the contextual decisions of individual animals and how we manage and conserve their populations. I have researched several themes, including animal movement, connectivity, sociality, carcass ecology, and the effects of harvest. Moreover, I have utilized a multitude of data collection methods (e.g. location data, genetics, isotopes) and statistical approaches to answer research questions under these themes. My overarching aim is help provide pragmatic answers to (1) applied questions and (2) contribute to the ever-advancing suite of data collection and analytical tools available to practitioners. Moreover, I hope to (3) help bridge the knowledge gap between wildlife science, research findings, and people. Select Publications Heeres, R. W., Leclerc, M., Frank, S., Kopatz, A., Pelletier, F., & Zedrosser, A. (2025). Flexibility in female spatiotemporal behavioral tactics to counter infanticide risk during the mating season. Movement Ecology, 13(1), 35. Hertel, A. G., Parres, A., Frank, S. C., Renaud, J., Selva, N., Zedrosser, A., Balkenhol, N., Maiorano, L., Fedorca, A., Dutta, T., Bogdanovic, N., Bragalanti, N., Chiriac, S., Cirovic, D., Ciucci, P., Domokos, C., Fedorca, M., Filacorda, S., Findo, S., . . . De Barba, M. (2025). Human Footprint and Forest Disturbance Reduce Space Use of Brown Bears (Ursus arctos) Across Europe. Glob Chang Biol, 31(1), e70011. https://doi.org/10.1111/gcb.70011 Lucas, P. M., Thuiller, W., Talluto, L., Polaina, E., Albrecht, J., Selva, N., De Barba, M., Penteriani, V., Gueguen, M., Balkenhol, N., Dutta, T., Fedorca, A., Frank, S. C., Zedrosser, A., Afonso-Jordana, I., Ambarli, H., Ballesteros, F., Bashta, A. T., Bilgin, C. C., . . . Pollock, L. J. (2025). Trophic Interactions Are Key to Understanding the Effects of Global Change on the Distribution and Functional Role of the Brown Bear. Glob Chang Biol, 31(6), e70252. https://doi.org/10.1111/gcb.70252 Zarzo-Arias, A., Heeres, R. W., Hertel, A. G., Leclerc, M., Frank, S., Steyaert, S. M., ... & Zedrosser, A. (2025). “The landscape of love”: sex-specific habitat-use during the mating season in a solitary large carnivore. Landscape Ecology, 40(12), 222. Hertel, A. G., Albrecht, J., Selva, N., Sergiel, A., Hobson, K. A., Janz, D. M., Mulch, A., Kindberg, J., Hansen, J. E., Frank, S. C., Zedrosser, A., & Mueller, T. (2024). Ontogeny shapes individual dietary specialization in female European brown bears (Ursus arctos). Nat Commun, 15(1), 10406. https://doi.org/10.1038/s41467-024-54722-z Heeres, R. W., Leclerc, M., Frank, S., Kopatz, A., Pelletier, F., & Zedrosser, A. (2024). Are nonsocial species more social than we think? Seasonal patterns in sociality in a solitary terrestrial carnivore. Animal Behaviour, 216, 107-130. https://doi.org/10.1016/j.anbehav.2024.07.022 Zedrosser, A., Frank, S. C., Hansen, J. E., Steyaert, S. M. J. G., & Swenson, J. E. (2023). The social system of a "nonsocial" species, the brown bear. In Bear and Human (Vol. 3, pp. 121-128). Brepols Publishers. https://doi.org/doi:10.1484/M.TANE-EB.5.134329 Hansen, J. E., Hertel, A. G., Frank, S. C., Kindberg, J., & Zedrosser, A. (2023). The role of familial conflict in home range settlement and fitness of a solitary mammal. Animal Behaviour, 202, 39-50. https://doi.org/10.1016/j.anbehav.2023.05.006 Dutta, T., De Barba, M., Selva, N., Fedorca, A. C., Maiorano, L., Thuiller, W., Zedrosser, A., Signer, J., Pflüger, F., Frank, S., Lucas, P. M., & Balkenhol, N. (2023). An objective approach to select surrogate species for connectivity conservation. Frontiers in Ecology and Evolution, 11. https://doi.org/10.3389/fevo.2023.1078649 Thorsen, N. H., Hansen, J. E., Stoen, O. G., Kindberg, J., Zedrosser, A., & Frank, S. C. (2022). Movement and habitat selection of a large carnivore in response to human infrastructure differs by life stage. Mov Ecol, 10(1), 52. https://doi.org/10.1186/s40462-022-00349-y Hansen, J. E., Hertel, A. G., Frank, S. C., Kindberg, J., & Zedrosser, A. (2022). Social environment shapes female settlement decisions in a solitary carnivore. Behav Ecol, 33(1), 137-146. https://doi.org/10.1093/beheco/arab118 Arnberg, M. P., Frank, S. C., Blaalid, R., Davey, M. L., Eycott, A. E., & Steyaert, S. (2022). Directed endozoochorous dispersal by scavengers facilitate sexual reproduction in otherwise clonal plants at cadaver sites. Ecol Evol, 12(1), e8503. https://doi.org/10.1002/ece3.8503 Frank, S. C., Pelletier, F., Kopatz, A., Bourret, A., Garant, D., Swenson, J. E., Eiken, H. G., Hagen, S. B., & Zedrosser, A. (2021). Harvest is associated with the disruption of social and fine-scale genetic structure among matrilines of a solitary large carnivore. Evol Appl, 14(4), 1023-1035. https://doi.org/10.1111/eva.13178 Frank, S. C., Blaalid, R., Mayer, M., Zedrosser, A., & Steyaert, S. (2020). Fear the reaper: ungulate carcasses may generate an ephemeral landscape of fear for rodents. R Soc Open Sci, 7(6), 191644. https://doi.org/10.1098/rsos.191644 Rivrud, I. M., Frank, S. C., Bischof, R., Mysterud, A., Steyaert, S., Hertel, A. G., Hagen, S. B., Eiken, H. G., Swenson, J. E., & Zedrosser, A. (2019). Heritability of head size in a hunted large carnivore, the brown bear (Ursus arctos). Evol Appl, 12(6), 1124-1135. https://doi.org/10.1111/eva.12786 Badia, R., Blaalid, R., Puliti, S., Steyaert, S. M. J. G., & Frank, S. C. (2019). Reindeer carcasses provide foraging habitat for insectivorous birds of the alpine tundra. Ornis Norvegica, 42, 36-40. https://doi.org/10.15845/on.v42i0.2639 Steyaert, S., Frank, S. C., Puliti, S., Badia, R., Arnberg, M. P., Beardsley, J., Okelsrud, A., & Blaalid, R. (2018). Special delivery: scavengers direct seed dispersal towards ungulate carcasses. Biol Lett, 14(8). https://doi.org/10.1098/rsbl.2018.0388 Frank, S. C., Leclerc, M., Pelletier, F., Rosell, F., Swenson, J. E., Bischof, R., Kindberg, J., Eiken, H. G., Hagen, S. B., & Zedrosser, A. (2018). Sociodemographic factors modulate the spatial response of brown bears to vacancies created by hunting. J Anim Ecol, 87(1), 247-258. https://doi.org/10.1111/1365-2656.12767 Leclerc, M., Frank, S. C., Zedrosser, A., Swenson, J. E., & Pelletier, F. (2017). Hunting promotes spatial reorganization and sexually selected infanticide. Sci Rep, 7, 45222. https://doi.org/10.1038/srep45222 Frank, S. C., Ordiz, A., Gosselin, J., Hertel, A., Kindberg, J., Leclerc, M., Pelletier, F., Steyaert, S. M. J. G., Støen, O.-G., Van de Walle, J., Zedrosser, A., & Swenson, J. E. (2017). Indirect effects of bear hunting: a review from Scandinavia. Ursus, 28(2), 150-164. https://doi.org/10.2192/ursu-d-16-00028.1 Steyaert, S. M. J. G., Zedrosser, A., Elfström, M., Ordiz, A., Leclerc, M., Frank, S. C., Kindberg, J., St⊘en, O. G., Brunberg, S., & Swenson, J. E. (2016). Ecological implications from spatial patterns in human‐caused brown bear mortality. Wildlife Biology, 22(4), 144-152. https://doi.org/10.2981/wlb.00165 Frank, S. C., Steyaert, S. M., Swenson, J. E., Storch, I., Kindberg, J., Barck, H., & Zedrosser, A. (2015). A "clearcut" case? Brown bear selection of coarse woody debris and carpenter ants on clearcuts. For Ecol Manage, 348, 164-173. https://doi.org/10.1016/j.foreco.2015.03.051 Education Ph.D., Ecology – University of South-Eastern Norway (formerly Telemark University College), 2017 M.S., Forest Ecology and Management – University of Freiburg, 2011 B.S., Wildlife Biology –​ Colorado State University, 2003 Current or Recent Positions Wildlife Research Scientist – Colorado Parks and Wildlife, State of Colorado, 2022-Present Statistical Analyst / Quality Specialist Colorado Department of Public Health and Environment, 2020-2022, State of Colorado Postdoctoral Associate –​ “BearConnect”, Department of Nature, Health, and Environment, University of South-Eastern Norway, 2017-2020 Contact Information 346 Co Rd 362 Hot Sulphur Springs, CO 80451 Email: shane.frank@state.co.us Phone: ​970-646-2961
Chuck Anderson, Head of Mammals Research
Chuck Anderson Mammals Research Section Leader Current or Recent Research Projects Mitigating Impacts of Energy Development on Piceance Basin Mule Deer Populations Areas of Interest and Expertise My research focuses on large mammal ecology and management, with an emphasis on population dynamics, population monitoring, and population genetics. Streaming Videos Wild Science: Mule Deer in the Piceance Basin Select Publications Crews, S., N. D. Rayl, M. W. Alldredge, E. J. Bergman, C. R. Anderson Jr., E. H. VanNatta, J. D. Holbrook, and G. Bastille-Rousseau. 2025. Hierarchy in structuring of resource selection: understanding elk selection across space, time, and movement strategies. Ecology and Evolution 15:e71097. https://doi.org/10.1002/ece3.71097 Crews, S., N. D. Rayl, M. W. Alldredge, E. J. Bergman, C. R. Anderson Jr., and G. Bastille-Rousseau. 2025. Drivers of spring migration phenology in Rocky Mountain elk. Scientific Reports 15:7807. https://doi.org/10.1038/s41598-025-91947-4 Hurley, M. A., C. R. Anderson Jr., T. D. Forrester, and J. A. Gude. 2023. Carnivore-prey relationships. Pages 125-139 in J. R. Heffelfinger and P. R. Krausman, editors. Ecology and management of black-tailed and mule deer of North America. CRC Press, Boca Raton, Florida, USA.​ ​Johnston, D. H., and C. R. Anderson Jr. 2023. Plant and mule deer responses to pinyon-juiper removal by three mechanical methods. Wildlife Society Bulletin 47:e1421​. https://doi.org/10.1002/jwmg.22110​ Bonar, M., S. J. Anderson, C. R. Anderson Jr, G. Wittemyer, J. M. Northrup, and A. B. Shafer. 2022. Genomic correlates for migratory direction in a free-ranging cervid. Proceedings of the Royal Society B 289:20221969. https://doi.org/10.1098/rspb.2022.1969​ Johnston, D. H., C. R. Anderson Jr., R. D. Applegate, L. Bailey, E. Cooch, J. Feiberg, A. B. Franklin, R. J. Gutiérrez, K. V. Miller, J. D. Nichols, N. D. Niemuth, D. Otis, C. A. Ribic, M. M. Rowland, and T. L. Shaffer. 2021. Editor's Message: A perspective on the Journal of Wildlife Management. Journal of Wildlife Management 87:1305-1308. https://doi.org/10.1002/jwmg.22110 Swearingen, T. C., R. W. Klaver, C. R. Anderson Jr., and C. N. Jacques. 2023. Influence of camera model and alignment on the performance of paired camera stations. Wildlife Society Bulletin 47:e1422. doi.org/ 10.1002/wsb.1422 Rheault H., C. R. Anderson Jr, M. Bonar, R. R. Marrotte, T. R. Ross, G. Wittem​yer and J. M. Northrup. 2021. Some memories never fade: inferring multi-scale memory effects on habitat selection of a migratory ungulate using step-selection functions. Frontiers in Ecology and Evolution 9:702818. https://doi.org/10.3389/fevo.2021.702818 Northrup, J. M., ​C. R. Anderson Jr., B. D. Gerber, and G. Wittemyer. 2021. Behavioral and demographic responses of mule deer to energy development on winter range. Wildlife Monographs: 208:1–37. https://doi.org/10.1002/wmon.1060 Bishop, C. J., M. W. Alldredge, D. P. Walsh, E. J. Bergman, C. R. Anderson Jr., D. Kilpatrick, J. Bakel, and C. Febvre. 2019. A noninvasive automated device for remotely collaring and weighing mule deer. Wildlife Society Bulletin 43:717-725. https://doi.org/10.1002/wsb.1034 Northrup, J. M., A. Arvin, C. R. Anderson Jr., E. Brown, and G. Whittemyer. 2019. On-animal acoustic monitoring provides insight to ungulate foraging behavior. Journal of Mammalogy 100:1479–1489. https://doi.org/10.1093/jmammal/gyz124 Jacques, C. N., R. W. Klaver, T. S. Swearingen, E. D. Davis, C. R. Anderson Jr., J. A. Jenks, C. S. DePerno, and R. D. Bluett. 2019. Estimating density and detection of bobcats in a fragmented Midwestern landscapes using spatial capture-recapture data from camera traps. Wildlife Society Bulletin 43:256–264. https://doi.org/10.1002/wsb.968 Peterson, M. E., C. R. Anderson Jr., M. W. Alldredge, and P. F. Doherty Jr. 2018. Using maternal mule deer movements to estimate timing of parturition and assist fawn captures. Wildlife Society Bulletin 42:616–621. https://doi.org/10.1002/wsb.935 Peterson, M.E., C. R. Anderson Jr., J. M. Northrup, and P. F. Doherty Jr. 2018. Mortality of mule deer fawns in a natural gas development area. The Journal of Wildlife Management 82:1135–1148. https://doi.org/10.1002/jwmg.21476 Bergman, E. J., C. R. Anderson Jr., C. J. Bishop, A. A. Holland, and J. M. Northrup. 2017. Variation in ungulate body fat: individual versus temporal effects. The Journal of Wildlife Management 82:130–137. https://doi.org/10.1002/jwmg.21334 Lendrum, P. E., J. M. Northrup, C. R. Anderson Jr.​, G. E. Liston, C. L. Aldridge, K. R. Crooks, and G. Wittemyer. 2017. Predation risk across a dynamic landscape: effects of anthropogenic land use, natural landscape features, and prey distribution. Landscape Ecology 33:151-170. https://doi.org/10.1007/s10980-017-0590-z Peterson, M. E., C. R. Anderson Jr., J. M. Northrup, and P. F. Doherty Jr. 2017. Reproductive success of mule deer in a natural gas development area. Wildlife Biology 4:1-5. https://doi.org/10.2981/wlb.00341 Northrup, J. M., C. R. Anderson Jr., and G. Wittemyer. 2015. Quantifying spatial habitat loss from hydrocarbon development through assessing habitat selection patterns of mule deer. Global Change Biology 21:3961–3970. https://doi.org/10.1111/gcb.13037 Lendrum, P. E., C. R. Anderson Jr., K. L. Monteith, J. A. Jenks, R. T. Bowyer. 2014. Relating the movement of a rapidly migrating ungulate to spatiotemporal patterns of forage quality. Mammalian Biology 79:369–375. https://doi.org/10.1016/j.mambio.2014.05.005 Freeman, E. D., R. T. Larsen, M. E. Peterson, C. R. Anderson Jr., K. R. Hersey, B. R. McMillan. 2014. Effects of male‐biased harvest on mule deer: implications for rates of pregnancy, synchrony, and timing of parturition. Wildlife Society Bulletin 38:806–811. https://doi.org/10.1002/wsb.450 Northrup, J. M., M. B. Hooten, C. R. Anderson Jr., and G. Wittemyer. 2013. Practical guidance on characterizing availability in resource selection functions under a use–availability design. Ecology 94:1456‐1463. https://doi.org/10.1890/12–1688.1 Lendrum, P. E., C. R. Anderson Jr., K. L. Monteith, J. A. Jenks and R. T. Bowyer. 2013. Mig​rating mule deer: effects of anthropogenically altered landscapes. PLoS ONE 8(5): e64548. https://doi.org/10.1371/journal.pone.0064548 Lendrum, P. E., C. R. Anderson Jr., R. A. Long, J. G. Kie, and R. T. Bowyer. 2012. Habitat se​lection by mule deer during migration: effects of landscape structure and natural–gas development. Ecosphere 3:1–19. https://doi.org/10.1890/ES12–00165.1 Anderson, C. R. Jr., F. G. Lindzey, K. H. Knopff, M. G. Jalkotzy, and M. S. Boyce. 2010. Cougar management in North America. Pages 41-54 in M. Hornocker and S. Negri, Editors. Cougar: Ecology and Conservation. University of Chicago Press, Chicago, IL, USA. Anderson, C. R. Jr., and F. G. Lindzey. 2005. Expe​rimental evaluation of population trend and harvest composition in a Wyoming cougar population. Wildlife Society Bulletin 33:179-188. https://doi.org/10.2193/0091-7648(2005)33[179:EEOPTA]2.0.CO;2 Anderson, C. R. Jr., and F. G. Lindzey. 2003. Estimating cougar predation rates from GPS location clusters. The Journal of Wildlife Management 67:307-316. https://www.jstor.org/stable/3802772 Anderson, C. R. Jr., M. A. Ternent, and D. S. Moody. 2002. Grizzly bear-cattle interactions on two cattle allotments in northwest Wyoming. Ursus 13:247-256. https://www.jstor.org/stable/3873205 Education Ph.D., Zoology and Physiology — University of Wyoming, 2003 M.S., Zoology and Physiology — University of Wyoming, 1994 B.S., Wildlife Biology — Colorado State University, 1990 Current or Recent Positions Mammals Researcher Section Leader — Colorado Parks and Wildlife, 2013-Present Wildlife Researcher — Colorado Parks and Wildlife, 2006-2013 Affiliate Faculty — Idaho State University, 2009-Present Affiliate Faculty — Colorado State University, 2012-Present
Jake Ivan, Senior Wildlife Research Scientist
Jake Ivan Senior Wildlife Research Scientist, Species of Conservation Concern Current or Recent Research Projects Snowshoe Hare Response to Silvicultural Treatments Monitoring Canada Lynx in Colorado Mammal and Breeding Bird Response to Bark Beetle Outbreaks Areas of Interest and Expertise My research focuses on estimating wildlife population parameters and forest carnivore ecology, conservation and monitoring. Streaming Videos Catching pine Lynx: shadows of the forest Snowshoe hare Snowshoe hare releases Snowshoe hare trapping, collaring and release Select Publications Bosco-Lauth, A. M., S. M. Porter, R. M. Maison, K. A. Fox, J. M. Marano, J. S. Ivan, K. Griffin, M. Quilici, and J. J. Root. 2025. Rabbit Hemorrhagic Disease Virus 2 experimental infection in snowshoe hares (Lepus americanus). Journal of Wildlife Diseases 61:674-684. doi: 10.7589/JWD-D-24-00141 Devarajan, K. [numerous authors], J. S. Ivan, [numerous authors]. 2025. When the wild things are: defining mammalian diel activity and plasticity. Science Advances 11:eado3843. DOI: 10.1126/sciadv.ado3843 Ivan, J. S. 2024. Canada Lynx (lynx Canadensis). Pages 129-152 in J. E. Cartron and J. K. Frey, Eds. Wild Carnivores of New Mexico. University of New Mexico Press. Hooten, M. B., M. R. Schwob, D. S. Johnson and J. S. Ivan. 2024. Geostatistical capture-recapture models​​. Spatial Statistics 59:100817. doi.org/10.1016/j.spasta.2024.100817​ Squires, J. R., Olson, L. E., Ivan, J. S., McDonald, P. M., and Holbrook, J. D. 2024. Anthropogenically protected but naturally disturbed: a specialist carnivore at its southern range periphery. Biodiversity and Conservation. https://doi.org/10.1007/s10531-024-02978-8 Hooten, M. B., M. R. Schwob, D. S. Johnson, and J. S. Ivan. 2023. Multistage hierarchical capture-recapture models. Environmetrics 34:e2799. doi.org/10.1002/env.2799 Ivan, J. S., E. S. Newkirk, and B. D. Gerber. 2023. Differential impacts of spruce beetle outbreaks on snowshoe hares and red squirrels in the southern Rocky Mountains​. Forest Ecology and Management 544:121147. doi.org/10.1016/j.foreco.2023.121147 Squire​s, J. R., J. S. Ivan, K. E. Paolini, L. E. Olson, G. M. Jones, and J. D. Holbrook​​. 2022. Keystone structures maintain forest function for Canada lynx after large-scale spruce beetle outbreak​​. Environmental Research: Ecology 2:011001. https://doi.org/10.1088/2752-664X/ac8eb7 Van Ee, J. J., J. S. Ivan, and M. B. Hooten. 2022. Community confounding in joint species distribution models. Scientific Reports 12:1–14. https://doi.org/10.1038/s41598-022-15694-6​ Kraberger, S., L. E. K. Serieys, C. Richet, N. M. Fountain-Jones, G. Baele, J. M. Bishop, M. Nehring, J. S. Ivan, E. S. Newkirk, J. R. Squires, M. C. Lund, S. P. D. Riley, C. C. Wilmers, P. D. van Helden, K. Van Doorslaer, M. Culver, S. VandeWoude, D. P. Martin, and A. Varsani. 2021. Complex evolutionary history of felid anelloviruses. Virology 562:176–189. https://doi.org/10.1016/j.virol.2021.07.013 Olson, L. E., N. Bjornlie, G. Hanvey, J. D. Holbrook, J. S. Ivan, S. Jackson, B. Kertson, T. King, M. Lucid, D. Murray, R. Naney, J. Rohrer, A. Scully, D. Thornton, Z. Walker, and J. R. Squires. 2021. Improved prediction of Canada lynx distribution through regional model transferability and data efficiency. Ecology and Evolution 11:1667–1690. https://doi.org/10.1002/ece3.7157 Squires, J. R., J. D. Holbrook, L. E. Olson, J. S. Ivan, R. W. Ghormley, and R. L. Lawrence. 2020. A specialized forest carnivore navigates landscape-level disturbance: Canada lynx in spruce-beetle impacted forests. Forest Ecology and Management 475:118400. https://doi.org/10.1016/j.foreco.2020.118400 Tabak, M. A., M. S. Norouzzadeh, D. W. Wolfson, E. J. Newton, R. K. Boughton, J. S. Ivan, E. A. Odell, E. S. Newkirk, R. Y. Conrey, J. Stenglein, F. Iannarilli, J. Erb, R. K. Brook, A. J. Davis, J. Lewis, D. P. Walsh, J. C. Beasley, K. C. VerCauteren, J. Clune, and R. S. Miller. 2020. Improving the accessibility and transferability of machine learning algorithms for identification of animals in camera trap images: MLWIC2. Ecology and Evolution 10:10374-10383. https://doi.org/10.1002/ece3.6692 Latif, Q. S., J. S. Ivan, A. E. Seglund, D. L. Pavlacky, and R. L. Truex. 2020. Avian relationships with bark beetle outbreaks and underlying mechanisms in lodgepole pine and spruce-fir forests of Colorado. Forest Ecology and Management 464:118043. https://doi.org/10.1016/j.foreco.2020.118043 Lukacs, P. M., D. Evans Mack, R. Inman, J. A. Gude, J. S. Ivan, R. P. Lanka, J. C. Lewis, R. A. Long, R. Sallabanks, Z. Walker, S. Courville, S. Jackson, R. Kahn, M. K. Schwartz, S. C. Torbit, J. S. Waller, and K. Carroll. 2020. Wolverine occupancy, spatial distribution, and monitoring design. The Journal of Wildlife Management 84:841–851. https://doi.org/10.1002/jwmg.21856 Zimova, M., A. P. Sirén, J. J. Nowak, A. M. Bryan, J. S. Ivan, T. L. Morelli, S. L. Suhrer, J. Whittington, and L. S. Mills. 2019. Local climate determines vulnerability to camouflage mismatch in snowshoe hares. Global Ecology and Biogeography 29:503–515. https://doi.org/10.1111/geb.13049 Gigliotti, L. C., N. D. Berg, R. Boonstra, S. M. Cleveland, D. R. Diefenbach, E. M. Gese, J. S. Ivan, K. Kielland, C. J. Krebs, A. V. Kumar, L. S. Mills, J. N. Pauli, H. B. Underwood, E. C. Wilson, and M. J. Sheriff. 2019. Latitudinal variation in snowshoe hare (Lepus americanus) body mass: a test of Bergmann’s rule. Canadian Journal of Zoology 98:88–95. https://doi.org/10.1139/cjz-2019-0184 Squires, J. R., L. E. Olson, E. K. Roberts, J. S. Ivan, and M. Hebblewhite. 2019. Winter recreation and Canada lynx: reducing conflict through niche partitioning. Ecosphere 10:e02876. https://doi.org/10.1002/ecs2.2876 Ivan, J. S. 2019. Density estimation. Pages 20.1–20.19 in E. G. Cooch and G. C. White, editors. Program MARK - 'a gentle introduction'. Ayers, C. R., K. C. Hanson-Dorr, K. Stromborg, T. W. Arnold, J. S. Ivan, and B. S. Dorr. 2019. Survival, fidelity, and dispersal of double-crested cormorants on two Lake Michigan Islands. The Auk 136:1-10. https://doi.org/10.1093/auk/ukz040​ Magee, P. A., J. D. Coop, and J. S. Ivan. 2019. Thinning alters avian occupancy in piñon–juniper woodlands. The Condor 121:1-17. https://doi.org/10.1093/condor/duy008 Tabak, M. A., M. S. Norouzzadeh, D. W. Wolfson, S. J. Sweeney, K. C. Vercauteren, N. P. Snow, J. M. Halseth, P. A. Di Salvo, J. S. Lewis, M. D. White, B. Teton, J. C. Beasley, P. E. Schlichting, R. K. Boughton, B. Wight, E. S. Newkirk, J. S. Ivan, E. A. Odell, R. K. Brook, P. M. Lukacs, A. K. Moeller, E. G. Mandeville, J. Clune, and R. S. Miller. 2018. Machine learning to classify animal species in camera trap images: applications in ecology. Methods in Ecology and Evolution 10:585–590. https://doi.org/10.1111/2041-210X.13120 Buderman, F. E., M. B. Hooten, M. W. Alldredge, E. M. Hanks, and J. S. Ivan. 2018. Time-varying predatory behavior is primary predictor of fine-scale movement of wildland-urban cougars. Movement Ecology 6:22. https://doi.org/10.1186/s40462-018-0140-6 Ivan, J. S., A. E. Seglund, R. L. Truex, and E. S. Newkirk. 2018. Mammalian responses to changed forest conditions resulting from bark beetle outbreaks in the southern Rocky Mountains. Ecosphere 9:e02369. https://doi.org/10.1002/ecs2.2369 Olson, L. E., J. R. Squires, E. K. Roberts, J. S. Ivan, and M. Hebblewhite. 2018. Sharing the same slope: behavioral responses of a threatened mesocarnivore to motorized and nonmotorized winter recreation. Ecology and Evolution 8:8555-8572. https://doi.org/10.1002/ece3.4382 Olson, L. E., J. R. Squires, E. K. Roberts, A. D. Miller, J. S. Ivan, and M. Hebblewhite. 2017. Modeling large-scale winter recreation terrain selection with implications for recreation management and wildlife. Applied Geography 86:66-91. https://doi.org/10.1016/j.apgeog.2017.06.023 Buderman, F. E., M. B. Hooten, J. S. Ivan, and T. M. Shenk. 2018. Large-scale movement behavior in a reintroduced predator population. Ecography 41:126-139. https://onlinelibrary.wiley.com/doi/ftr/10.1111/ecog.03030 Baigas, P. E., J. R. Squires, L. E. Olsen, J. S. Ivan, and E. K. Roberts. 2017. Using environmental features to model highway crossing behavior of Canada lynx in the Southern Rocky Mountains. Landscape and Urban Planning 157:200-213. https://doi.org/10.1016/j.landurbplan.2016.06.007 Hooten, M. B., F. E. Buderman, B. M. Brost, E. M. Hanks, and J. S. Ivan. 2016. Hierarchical animal movement models for population-level inference. Environmetrics 27:322-333. https://doi.org/10.1002/env.2402 Ivan, J. S., and T. M. Shenk. 2016. Winter diet and hunting success of Canada lynx in Colorado. The Journal of Wildlife Management 80:1049-1058. https://doi.org/10.1002/jwmg.21101 Ivan, J. S., and E. S. Newkirk. 2016. CPW Photo Warehouse: a custom database to facilitate archiving, identifying, summarizing, and managing photo data collected from camera traps. Methods in Ecology and Evolution 7:499-504. https://doi.org/10.1111/2041-210X.12503 Buderman, F. E., M. B. Hooten, J. S. Ivan, and T. M. Shenk. 2016. A functional model for characterizing long-distance movement behavior. Methods in Ecology and Evolution 7:264-273. https://doi.org/10.1111/2041-210X.12465 Ellis, M. M., J. S. Ivan, J. M. Tucker, and M. K. Schwartz. 2015. rSPACE: Spatially based power analysis for conservation and ecology. Methods in Ecology and Evolution 6:621-625. https://doi.org/10.1111/2041-210X.12369 Ivan, J. S., G. C. White, and T. M. Shenk. 2014. Density and demography of snowshoe hares in central Colorado. The Journal of Wildlife Management 78:580–594. https://doi.org/10.1002/jwmg.695 Gerber, B. D., J. S. Ivan, and K. P. Burnham. 2014. Estimating the abundance of rare and elusive carnivores from photographic-sampling data when the population size is very small. Population Ecology 56:463-470. https://doi.org/10.1007/s10144-014-0431-8 Trainor, A. M., O. J. Schmitz, J. S. Ivan, and T. M. Shenk. 2014. Enhancing species distribution modeling by characterizing predator–prey interactions. Ecological Applications 24:204–216. https://doi.org/10.1890/13-0336.1 Ellis, M. M., J. S. Ivan, and M. K. Schwartz. 2014. Spatially explicit power analyses for occupancy-based monitoring of wolverine in the U.S. Rocky Mountains. Conservation Biology 28:52–62. https://doi.org/10.1111/cobi.12139 Ivan J. S., G. C. White, and T. M. Shenk. 2013. Using simulation to compare methods for estimating density from capture-recapture data. Ecology 94:817–826. https://doi.org/10.1890/12-0102.1 Ivan, J. S., G. C. White, and T. M. Shenk. 2013. Using auxiliary telemetry information to estimate animal density from capture-recapture data. Ecology 94:809–816. https://doi.org/10.1890/12-0101.1 Stromborg, K. L., J. S. Ivan, J. K. Netto, and C. R. Courtney. 2012. Survivorship and mortality patterns of double-crested cormorants at Spider Island, Wisconsin, 1988–2006. Waterbirds 35:31–39. https://doi.org/10.1675/063.035.sp105 Education Ph.D., Fish, Wildlife and Conservation Biology — Colorado State University, 2011 M.S., Wildlife Biology — University of Montana, 2000 B.S., Wildlife Science — Purdue University, 1997 Current or Recent Positions Wildlife Researcher — Colorado Parks and Wildlife, 2010–Present Refuge Biologist — U.S. Fish and Wildlife Service, 2003-2005 Piping Plover Recovery Biologist — The Nature Conservancy, 2000-2003
Eric Bergman, Mammals Research Scientist
Mat Alldredge, Senior Wildlife Research Scientist
Senior Wildlife Research Scientist, Carnivores ​ Current or Recent Research Projects Using Non-Invasive Genetic Sampling to Estimate Mountain Lion and Bobcat Abundance, Age Structure and Diet Mountain Lion Demographics and Human Interactions Along the Urban-Exurban Front Range Areas of Interest and Expertise My research interest focuses on large carnivore and ungulate ecology, population parameter estimation, habitat use, avian ecology and population sampling. Streaming Videos Mountain lions in Colorado – a 4 part series Series Trailer Part 1: Mountain Lion Biology & Historical Perspective Part 2: Mountain Lion Habitat & Human Expansion Part 3: Hunting Part 4: What to do if you encounter a mountain lion Colorado Mountain Lions (all 4 videos combined) Select Publications Crews, S., N. D. Rayl, M. W. Alldredge, E. J. Bergman, C. R. Anderson Jr., E. H. VanNatta, J. D. Holbrook, and G. Bastille-Rousseau. 2025. Hierarchy in structuring of resource selection: understanding elk selection across space, time, and movement strategies. Ecology and Evolution 15:e71097. https://doi.org/10.1002/ece3.71097 Crews, S., N. D. Rayl, M. W. Alldredge, E. J. Bergman, C. R. Anderson Jr., and G. Bastille-Rousseau. 2025. Drivers of spring migration phenology in Rocky Mountain elk. Scientific Reports 15:7807. https://doi.org/10.1038/s41598-025-91947-4 Venumière-Lefebvre, C. C., H. E. Johnson, S. W. Breck, M. W. Alldredge, and K. R. Crooks. 2025. Bears avoid residential neighborhoods in response to the experimental reduction of anthropogenic attractants. Frontiers in Ecology and Evolution 13:1657106. doi: 10.3389/fevo.2025.1657106 Fountain-Jones, N. M., S. Kraberger, R. B. Gagne, D. R. Trumbo, P. E. Salerno, W. C. Funk, K. Crooks, R. Biek, M. W. Alldredge, K. Logan, G. Baele, S. Dellicour, H. B. Ernest, S. VandeWoude, S. Carver, and M. E. Craft. 2021. Host relatedness and landscape connectivity shape pathogen spread in the puma, a large secretive carnivore​. Communications Biology 4:12. doi: 10.1038/s42003-020-01548-2. Gagne, R. B., S., Kraberger, R. McMinn, D. R. Trumbo, C. R. Anderson Jr, K. A. Logan, M. W. Alldredge, K. Griffin and S. VandeWoude. 2021. Viral sequen​ces recovered from puma tooth DNA reconstruct statewide viral phylogenies. Frontiers in Ecology and Evolution 9:734462. doi: 10.3389/fevo.2021.734462​ Kraberger, S., N. M. Fountain-Jones, R. B. Gagne, J. Malmberg, N. G. Dannemiller, K. Logan, M. Alldredge, A. Varsani, K. R. Crooks, M. Craft, S. Carver, and S. VandeWoude. 2020. Frequent cross-species transmissions of foamy virus between domestic and wild felids. Virus Evolution 6(1):vez058. https://doi.org/10.1093/ve/vez058 Bishop, C. J., M. W. Alldredge, D. P. Walsh, E. J. Bergman, C. R. Anderson Jr, D. Kilpatrick, J. Bakel, and C. Febvre. 2019. A noninvasive automated device for remotely collaring and weighing mule deer. Wildlife Society Bulletin 43:717–725. https://doi.org/10.1002/wsb.1034 Trumbo, D. R., P. E. Salerno, K. A. Logan, M. W. Alldredge, R. B. Gagne, C. P. Kozakiewicz, S. Kraberger, N. M. Fountain‐Jones, M. E. Craft, S. Carver, H. B. Ernest, K. R. Crooks, S. VandeWoude, and W. C. Funk. 2019. Urbanization impacts apex predator gene flow but not genetic diversity across an urban‐rural divide. Molecular Ecology 28:4926–4940. https://doi.org/10.1111/mec.15261 Alldredge, M. W., F. E. Buderman, and K. A. Blecha. 2019. Human–Cougar interactions in the wildland–urban interface of Colorado's front range. Ecology and Evolution 9:10415–10431. https://doi.org/10.1002/ece3.5559 Alldredge, M. W., T. Blecha, and J. H. Lewis. 2019. Less invasive monitoring of cougars in Colorado's Front Range. Wildlife Society Bulletin 43:222–230. https://doi.org/10.1002/wsb.971 Kechejian, S. R., N. Dannemiller, S. Kraberger, C. Ledesma-Feliciano, J. Malmberg, M. Roelke Parker, M. Cunningham, R. McBride, S. P. Riley, W. T. Vickers, K. Logan, M. Alldredge, K. Crooks, M. Löchelt, S. Carver, and S. VandeWoude. 2019. Feline foamy virus is highly prevalent in free-ranging puma concolor from Colorado, Florida and Southern California. Viruses 11:359. https://doi.org/10.3390/v11040359 Kirby, R., H. E. Johnson, M. W. Alldredge, and J. N. Pauli. 2019. The cascading effects of human food on hibernation and cellular aging in free-ranging black bears. Scientific Reports 9:2197. https://doi.org/10.1038/s41598-019-38937-5 Peterson, M. E., C. R. Anderson Jr, M. W. Alldredge, and P. F. Doherty Jr. 2018. Using maternal mule deer movements to estimate timing of parturition and assist fawn captures. Wildlife Society Bulletin 42:616–621. https://doi.org/10.1002/wsb.935 Buderman, F. E., M. B. Hooten, M. W. Alldredge, E. M. Hanks, and J. S. Ivan. 2018. Time-varying predatory behavior is primary predictor of fine-scale movement of wildland-urban cougars. Movement Ecology 6:22. https://doi.org/10.1186/s40462-018-0140-6 Blecha, K. A., R. B. Boone, and M. W. Alldredge. 2018. Hunger mediates apex predator's risk avoidance response in wildland-urban interface. Journal of Animal Ecology 87:609–622. https://doi.org/10.1111/1365-2656.12801 Lewis, J. S., K. A. Logan, M. W. Alldredge, S. Carver, S. N. Bevins, M. Lappin, S. VandeWoude, and K. R. Crooks. 2017. The effects of demographic, social, and environmental characteristics on pathogen prevalence in wild felids across a gradient of urbanization. PLoS One 12(11):e0187035. https://doi.org/10.1371/journal.pone.0187035 Lewis, J. S., K. A. Logan, M. W. Alldredge, D. M. Theobald, S. VandeWoude, and K. R. Crooks. 2017. Contact networks reveal potential for interspecific interactions of sympatric wild felids driven by space use. Ecosphere 8(3):e01707. https://doi.org/10.1002/ecs2.1707 Kirby, R., M. 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