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Mammals Wildlife Research Reports

Federal Aid Progress Report​​​​s

Technical Publications

Holmes, B. E. 2023. Bird and mammal inventory of the Rio Blanco Lake State Wildlife Area, Rio Blanco County, Colorado​. Colorado Parks and Wildlife Technical Publication No. 63. Denver, CO, USA

Quartuch, M. R., M. Vornholt, K. Dorman, and M. Eckert. 2023. Linking hunting access and chronic wasting disease: a case study from Northeastern Colorado​. Colorado Parks and Wildlife Technical Publication No. 62. Denver, CO, USA

Johnston, D. B. 2020. Piceance Basin restoration for wildlife​. Colorado Parks and Wildlife Technical Publication No. 57​. Denver, CO, USA​

Logan, K. A., and J. P. Runge. 2020. Effects of hunting on a puma population in Colorado. Colorado Parks and Wildlife Technical Pu​blication No. 54. Denver, CO, USA

Stonehouse, K. F., C. R. Anderson Jr, Peterson, M.E., and D. R. Collins. 2016. Approaches to field investigations of cause-specific mortality in mule deer. Colorado Parks and Wildlife Technical Publication No. 48. Denver, CO, USA

Journal Articles

Black Bear

Johnson, H. E., D. L. Lewis, and S. W. Breck. 2020. Ind​ividual and population fitness consequences associated with large carnivore use of residential development. Ecosphere 11: e03098.​

Lewis, J. H., M. W. Alldredge, B. P. Dreher, J. L. George, S. Wait, B. Petch, and J. P. Runge. 2019. Summarizing Colorado’s black bear two–strike directive 30 years after inception. Wildlife Society Bulletin 43:599-607.​

Lischka, S. A., T. L. Teel, H. E. Johnson, and K. R. Crooks. 2019. Understanding and managing human tolerance for a large carnivore in a residential sys​tem​. Biological Conservation 238:1081–1089.

Laufenberg, J. S., H. E. Johnson, P. F. Doherty, and ​​S. W. Breck. 2018. Compounding effects of human development and a natural food shortage on a black bear population along a human development–wildland interface. Biological Conservation 224:188–198. ​

Johnson, H. E., D. L. Lewis, S. A. Lischka, and S. W. Breck. 2018. Assessing ecological and social outcomes of a bear‐proofing experiment. The Journal of Wildlife Management 82:1102–1114.

Wilbur, R. C., S. A. Lischka, J. R. Young, and H. E. Johnson. 2018. Experience, attitudes, and demographic factors influence the probability of reporting human–black bear interactions. Wildlife Society Bulletin 42:22–31.

Johnson, H. E., D. L. Lewis, T. L. Verzuh, C. F. Wallace, R. M. Much, L. K. Willmarth, and S. W. Breck. 2017. Human development and climate affect hibernation in a large carnivore with implications for human–carnivore conflicts. Journal of Applied Ecology 55:663–672.–2664.13021 ​

Kirby, R., M. W. Alldredge, and J. N. Pauli. 2017. Environmental, not individual, factors drive markers of biological aging in black bears. Evolutionary Ecology 31:571–581.​ 

Kirby, R., M. W. Alldredge, and J. N. Pauli. 2016. The diet of black bears tracks the human footprint across a rapidly developing landscape. Biological Conservation 200:51–59. 

Wolfe, L. L., H. E. Johnson, M. C. Fisher, W. R. Lance, D. K. Smith, and M. W. Miller. 2016. Chemical immobilization in American black bears using a combination of nalbuphine, medetomidine, and azaperone. Ursus 27:1–4.–D–15–00018.1


Jacques, C. N., R. W. Klaver, T. S. Swearingen, E. D. Davis, C. R. Anderson, 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.


Bergman, E. J., and C. J. Bishop. 2023. Colorado Plateau ecoregion. Pages 275-287 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.

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. B., and C. R. Anderson Jr. 2023. Plant and mule deer responses to pinyon-juniper removal by three mechanical treatment methods​. Wildlife Society Bulletin 47:e1421.​

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.

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 10:943411.

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.​​​

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.

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.

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.​

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.

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.​

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.​

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.​

Lendrum, P. E., J. M. Northrup, C. R. Anderson, 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.​

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.​

Johnson,/ H. E., J. R. Sushinsky, A. Holland, E. J. Bergman, T. Balzer, J. Garner and S. E. Reed. 2016. Increases in residential and energy development are associated with reductions in recruitment for a large ungulate. Global Change Biology 23:578–591.

Northrup, J. M., C. R. Anderson, Jr., M. B. Hooten, and G. Wittemyer. 2016. Movement reveals scale dependence in habitat selection of a large ungulate. Ecological Applications 26:2746–2757.​

Northrup, J. M., C. R. Anderson, Jr., and G. Wittemyer. 2016. Environmental dynamics and anthropogenic development alter philopatry and space–use in a North American cervid. Diversity and Distributions 22:547–557.


Watkins, B. E., E. J. Bergman, L. C. Dhaseleer, and L. J. Bernal. 2022. Factors influencing productivity and recruitment of elk in northern New Mexico. The Journal of Wildlife Management: e22348.

Rayl, N. D., J. A. Merkle, K. M. Profitt, E. S. Almberg, J. D. Jones, J. A. Gude, and P. C. Cross. 2021. Elk migration influences the risk of disease spillover in the Greater Yellowstone Ecosystem​. Journal of Animal Ecology 90:1264-1275.

Rayl, N. D., K. M. Proffitt, E. S. Almberg, J. D. Jones, J. A. Merkle, J. A. Gude, and P. C. Cross. 2019. Modeling elk-to-livestock transmission risk to identify hotspots of brucellosis spillover​. The Journal of Wildlife Management 83:817-829.

Lukacs, P. M., M. S. Mitchell, M. Hebblewhite, B. K. Johnson, H. Johnson, M. Kauffman, K. M. Proffitt, P. Zager, J. Brodie, K. Hersey, and A. A. Holland. 2018. Factors influencing elk recruitment across ecotypes in the Western United States. The Journal of Wildlife Management 82:698–710.

Rayl, N. D., K. M. Proffitt, E. S. Almberg, J. D. Jones, J. A. Merkle, and P. C. Cross. 2018. Estimating the risk of elk-to-livestock brucellosis transmission in Montana. Montana Fish, Wildlife and Parks, Helena, Montana, USA


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.

Van Ee, J. J., J. S. Ivan, and M. B. Hooten. 2022. Community confounding in joint species distribution models. Scientific Reports 12:1–14.​

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.

​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.

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.​​​

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.​

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.​

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.​


DeCesare, N. J., R. B. Harris, M. P. Atwood, E. J. Bergman, A. B. Courtemanch, P. C. Cross, G. L. Fralick, K. R. Hersey, M. A. Hurley, and T. M. Koser. 2024. Warm places, warm years, and warm seasons increase parasitizing of moose by winter ticks. Ecosphere 15:e4799.

Hayes, F. P., J. J. Millspaugh, E. J. Bergman, R. M. Callaway, and C. J. Bishop. 2022. Effects of willow nutrition and morphology on calving success of moose. The Journal of Wildlife Management 202286:e22175.​

Bergman, E. J., F. P. Hayes, and K Aagaard. 2020. Estimation of moose parturition dates in Colorado: incorporating imperfect detections. Alces 56:127–135.​

Bergman, E. J., F. P. Hayes, P. M. Lukacs, and C. J. Bishop. 2020. Moose calf detection probabilities: quantification and evaluation of a ground-based survey technique​. Wildlife Biology 2020:1-9.

Nadeau, M. S., N. J. DeCesare, D. G. Brimeyer, E. J. Bergman, R. B. Harris, K. R. Hersey, K. K. Huebner, P. E. Mathews, and T. P. Thomas. 2017. Status and trends of moose populations and hunting opportunity in the western United States. Alces 53:99–112.

Mountain Lion

Human-Cougar Interactions Science Review Team. 2022. Human-Cougar Interactio​ns: A Literature Review Related to Common Management Questions. Washington Department of Fish and Wildlife, Olympia, Washington, USA. 78 pp.

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., L. E. 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. 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. doi:10.1016/j.virol.2021.07.013

Logan, K. A., and J. P. Runge. 2021. Effects of hunting on a puma population in Colorado. Wildlife Monographs 209:1-35.

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.

Logan, K. A. 2019. Puma population limitation and regulation: What matters in puma management? The Journal of Wildlife Management 83:1652-1666.​002/jwmg.21753

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.

Blecha, K. A., R. B. Boone, and M. W. Alldredge. 2018. Hung​er​ mediates a​pex predator's risk avoidance response in wildland–urban interface​. Journal of Animal Ecology 87:609–622.​​

Moss, W. E., M. W. Alldredge, and J. N. Pauli. 2016. Quantifying risk and resource use of a large carnivore in an expanding urban–wildland interface. Journal of Applied Ecology 53:371–378.–2664.12563

Moss, W. E., M. W. Alldredge, K. A. Logan, and J. N. Pauli. 2016. Human expansion precipitates niche expansion for an opportunistic apex predator (Puma concolor). Scientific Reports 6:39639. ​​

Snowshoe Hare

​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 a​mericanus) body mass: a test of Bergmann’s rule. Canadian Journal of Zoology 98:88–95.

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.


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 esti​mating cestode prevalence in a wild carnivore population​. PLoS ONE 17: e0277420.

​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.​​

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.


​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.


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.

Hooten, M. B., M. R. Schwob, D. S. Johnson and J. S. Ivan. 2024. Geostatistical capture-recapture models​. Spatial Statistics 59:100817.

Dutta, T., M. De Barba, N Selva, A. C. Fedorca, L Maiorano, W. Thuiller, A. Zedresser, J. Signer, F. Pfluger, S. Frank, P M. Lucas, and N. Balkenholl. 2023. An objective approach to select surrogate species for connectivity conservation. Frontiers in Ecology and Evolution 11:1078649. doi: 10.3389/fevo.2023.1078649

Hooten, M. B., M. R. Schwob, D. S. Johnson, and J. S. Ivan. 2023. Multistage hierarchical capture-recapture models. Environmetrics 34:e2799.

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.​

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. 10.1002/wsb.1422

Thorsen, N. H., J. E. Hansen, O. G. Stoen, J. Kindberg, A. Zedrosser, and S. C. Frank. 2022. Movement and habitat selection of a large carnivore in response to human infrastructure differs by life stage. Movement Ecology 10:52.​186/s40462-022-00349-y​​

​Van Ee, J. J., J. S. Ivan, and M. B. Hooten. 2022. Community confounding in joint sp​ecies distribution models​​. Scientific Reports 12:1–14.​

Johnson, D. H., C. 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. The Journal of Wildlife Management 87:1305-1308.

Carver, S, S. N. Bevins, M. R. Lappin, E. E. Boydston, L. M. Lyren, M. Alldredge, K. A. Logan, L. L. Sweanor, S. P. D. Riley, L. E. K. Serieys, R. N. Fisher, T. W. Vickers, W. Boyce, R. McBride, M. C. Cunningham, M. Jennings, J. Lewis, T. Lunn, K. R. Crooks, S.VandeWoude. 2016. Pathogen exposure varies widely among sympatric populations of wild and domestic felids across the United States. Ecological Applications 26:367–381.–0445

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.

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.