Large carnivores avoid humans while prioritizing prey acquisition in anthropogenic areas.
Anthropocene
human influence
kill sites
linear features
predation risk
predator
prey
wildland-urban interface
Journal
The Journal of animal ecology
ISSN: 1365-2656
Titre abrégé: J Anim Ecol
Pays: England
ID NLM: 0376574
Informations de publication
Date de publication:
04 2023
04 2023
Historique:
received:
12
07
2022
accepted:
30
01
2023
medline:
6
4
2023
pubmed:
10
2
2023
entrez:
9
2
2023
Statut:
ppublish
Résumé
Large carnivores are recovering in many landscapes where the human footprint is simultaneously growing. When carnivores encounter humans, the way they behave often changes, which may subsequently influence how they affect their prey. However, little research investigates the behavioural mechanisms underpinning carnivore response to humans. As a result, it is not clear how predator-prey interactions and their associated ecosystem processes will play out in the human-dominated areas into which carnivore populations are increasingly expanding. We hypothesized that humans would reduce predation risk for prey by disturbing carnivores or threatening their survival. Alternatively, or additionally, we hypothesized that humans would increase predation risk by providing forage resources that congregate herbivorous prey in predictable places and times. Using grey wolves Canis lupus in Jackson Hole, Wyoming, USA as a study species, we investigated 170 kill sites across a spectrum of human influences ranging from heavily restricted human activities on protected federal lands to largely unregulated activities on private lands. Then, we used conditional logistic regression to quantify how the probability of predation changed across varied types and amounts of human influences, while controlling for environmental characteristics and prey availability. Wolves primarily made kills in environmental terrain traps and where prey availability was high, but predation risk was significantly better explained with the inclusion of human influences than by environmental characteristics alone. Different human influences had different, and even converse, effects on the risk of wolf predation. For example, where prey were readily available, wolves preferentially killed animals far from motorized roads but close to unpaved trails. However, wolves responded less strongly to humans, if at all, where prey were scarce, suggesting they prioritized acquiring prey over avoiding human interactions. Overall, our work reveals that the effects of large carnivores on prey populations can vary considerably among different types of human influences, yet carnivores may not appreciably alter predatory behaviour in response to humans if prey are difficult to obtain. These results shed new light on the drivers of large carnivore behaviour in anthropogenic areas while improving understanding of predator-prey dynamics in and around the wildland-urban interface.
Identifiants
pubmed: 36757108
doi: 10.1111/1365-2656.13900
doi:
Banques de données
Dryad
['10.5061/dryad.44j0zpcj7']
Types de publication
Journal Article
Langues
eng
Sous-ensembles de citation
IM
Pagination
889-900Informations de copyright
© 2023 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.
Références
Acorn, R. C., & Dorrance, M. J. (1990). Methods of investigating predation of livestock. Alberta Agriculture and Rural Development.
Barker, K. J., Cole, E., Courtemanch, A., Dewey, S., Gustine, D., Mills, K., Stephenson, J., Wise, B., & Middleton, A. D. (2023). Data from: Large carnivores avoid humans while prioritizing prey acquisition in anthropogenic areas. Dryad Digital Repository. https://doi.org/10.5061/dryad.44j0zpcj7
Basille, M., Herfindal, I., Santin-Janin, H., Linnell, J. D. C., Odden, J., Andersen, R., Høgda, K. A., & Gaillard, J. (2009). What shapes Eurasian lynx distribution in human dominated landscapes: Selecting prey or avoiding people? Ecography, 32(4), 683-691. https://doi.org/10.1111/j.1600-0587.2009.05712.x
Behdarvand, N., Kaboli, M., Ahmadi, M., Nourani, E., Mahini, A. S., & Aghbolaghi, M. A. (2014). Spatial risk model and mitigation implications for wolf-human conflict in a highly modified agroecosystem in western Iran. Biological Conservation, 177, 156-164. https://doi.org/10.1016/j.biocon.2014.06.024
Boyce, M. S., Mao, J. S., Merrill, E. H., Fortin, D., Turner, M. G., Fryxell, J., & Turchin, P. (2003). Scale and heterogeneity in habitat selection by elk in Yellowstone National Park. Écoscience, 10(4), 421-431. https://doi.org/10.1080/11956860.2003.11682790
Bradley, E. H., Robinson, H. S., Bangs, E. E., Kunkel, K., Jimenez, M. D., Gude, J. A., & Grimm, T. (2015). Effects of wolf removal on livestock depredation recurrence and wolf recovery in Montana, Idaho, and Wyoming. The Journal of Wildlife Management, 79(8), 1337-1346. https://doi.org/10.1002/jwmg.948
Brodie, J., Johnson, H., Mitchell, M., Zager, P., Proffitt, K., Hebblewhite, M., Kauffman, M., Johnson, B., Bissonette, J., Bishop, C., Gude, J., Herbert, J., Hersey, K., Hurley, M., Lukacs, P. M., McCorquodale, S., McIntire, E., Nowak, J., Sawyer, H., … White, P. J. (2013). Relative influence of human harvest, carnivores, and weather on adult female elk survival across western North America. Journal of Applied Ecology, 50(2), 295-305. https://doi.org/10.1111/1365-2664.12044
Brooks, M. E., Kristensen, K., Benthem, K. J., van Magnusson, A., Berg, C. W., Nielsen, A., Skaug, H. J., Mächler, M., & Bolker, B. M. (2017). glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. The R Journal, 9(2), 378. https://doi.org/10.32614/rj-2017-066
Bunnefeld, N., Linnell, J. D. C., Odden, J., Duijn, M. A. J. V., & Andersen, R. (2006). Risk taking by Eurasian lynx (Lynx lynx) in a human-dominated landscape: Effects of sex and reproductive status. Journal of Zoology, 270(1), 31-39. https://doi.org/10.1111/j.1469-7998.2006.00107.x
Burnham, K. P., & Anderson, D. R. (2004). Multimodel Inference. Sociological Methods & Research, 33(2), 261-304. https://doi.org/10.1177/0049124104268644
Calenge, C. (2006). The package “adehabitat” for the R software: A tool for the analysis of space and habitat use by animals. Ecological Modelling, 197(3-4), 516-519. https://doi.org/10.1016/j.ecolmodel.2006.03.017
Caroline, v. D., Ras, E., de Vos, A. E. W., de Boer, W. F., Henkens, R. J. H. G., & Usukhjargal, D. (2009). Wolf predation among reintroduced Przewalski horses in Hustai National Park, Mongolia. Journal of Wildlife Management, 73(6), 836-843. https://doi.org/10.2193/2008-027
Carricondo-Sanchez, D., Zimmermann, B., Wabakken, P., Eriksen, A., Milleret, C., Ordiz, A., Sanz-Perez, A., & Wikenros, C. (2020). Wolves at the door? Factors influencing the individual behavior of wolves in relation to anthropogenic features. Biological Conservation, 244, 108514. https://doi.org/10.1016/j.biocon.2020.108514
Ceballos, G., & Ehrlich, P. R. (2002). Mammal population losses and the extinction crisis. Science, 296(5569), 904-907. https://doi.org/10.1126/science.1069349
Chapron, G., Kaczensky, P., Linnell, J. D. C., von Arx, M., Huber, D., Andrén, H., López-Bao, J. V., Adamec, M., Álvares, F., Anders, O., Balčiauskas, L., Balys, V., Bedő, P., Bego, F., Blanco, J. C., Breitenmoser, U., Brøseth, H., Bufka, L., Bunikyte, R., … Boitani, L. (2014). Recovery of large carnivores in Europe's modern human-dominated landscapes. Science, 346(6216), 1517-1519. https://doi.org/10.1126/science.1257553
Cooper, A. B., & Millspaugh, J. J. (1999). The application of discrete choice models to wildlife resource selection studies. Ecology, 80(2), 566-575. https://doi.org/10.1890/0012-9658(1999)080[0566:taodcm]2.0.co;2
Cotterill, G. G., Cross, P. C., Cole, E. K., Fuda, R. K., Rogerson, J. D., Scurlock, B. M., & du Toit, J. T. (2018). Winter feeding of elk in the greater Yellowstone ecosystem and its effects on disease dynamics. Philosophical Transactions of the Royal Society B: Biological Sciences, 373(1745), 20170093. https://doi.org/10.1098/rstb.2017.0093
DeCesare, N. J., Hebblewhite, M., Schmiegelow, F., Hervieux, D., McDermid, G. J., Neufeld, L., Bradley, M., Whittington, J., Smith, K. G., Morgantini, L. E., Wheatley, M., & Musiani, M. (2012). Transcending scale dependence in identifying habitat with resource selection functions. Ecological Applications, 22(4), 1068-1083. https://doi.org/10.1890/11-1610.1
DeMars, C. A., & Boutin, S. (2018). Nowhere to hide: Effects of linear features on predator-prey dynamics in a large mammal system. Journal of Animal Ecology, 87(1), 274-284. https://doi.org/10.1111/1365-2656.12760
Dickie, M., McNay, S. R., Sutherland, G. D., Cody, M., & Avgar, T. (2020). Corridors or risk? Movement along, and use of, linear features varies predictably among large mammal predator and prey species. Journal of Animal Ecology, 89(2), 623-634. https://doi.org/10.1111/1365-2656.13130
Elbroch, L. M., Lowrey, B., & Wittmer, H. U. (2018). The importance of fieldwork over predictive modeling in quantifying predation events of carnivores marked with GPS technology. Journal of Mammalogy, 99(1), 223-232. https://doi.org/10.1093/jmammal/gyx176
Estes, J. A., Terborgh, J., Brashares, J. S., Power, M. E., Berger, J., Bond, W. J., Carpenter, S. R., Essington, T. E., Holt, R. D., Jackson, J. B. C., Marquis, R. J., Oksanen, L., Oksanen, T., Paine, R. T., Pikitch, E. K., Ripple, W. J., Sandin, S. A., Scheffer, M., Schoener, T. W., … Wardle, D. A. (2011). Trophic downgrading of planet earth. Science, 333(6040), 301-306. https://doi.org/10.1126/science.1205106
Fortin, D., Fortin, M.-E., Beyer, H. L., Duchesne, T., Courant, S., & Dancose, K. (2009). Group-size-mediated habitat selection and group fusion-fission dynamics of bison under predation risk. Ecology, 90(9), 2480-2490. https://doi.org/10.1890/08-0345.1
Frey, S., Volpe, J. P., Heim, N. A., Paczkowski, J., & Fisher, J. T. (2020). Move to nocturnality not a universal trend in carnivore species on disturbed landscapes. Oikos, 129(8), 1128-1140. https://doi.org/10.1111/oik.07251
Frid, A., & Dill, L. M. (2002). Human-caused disturbance stimuli as a form of predation risk. Conservation Ecology, 6(1), 11. [online]. https://doi.org/10.5751/es-00404-060111
Gervasi, V., Sand, H., Zimmermann, B., Mattisson, J., Wabakken, P., & Linnell, J. D. C. (2013). Decomposing risk: Landscape structure and wolf behavior generate different predation patterns in two sympatric ungulates. Ecological Applications, 23(7), 1722-1734. https://doi.org/10.1890/12-1615.1
Gompper, M. E., Belant, J. L., & Kays, R. (2015). Carnivore coexistence: America's recovery. Science, 347(6220), 382-383. https://doi.org/10.1126/science.347.6220.382-b
Gula, R. (2004). Influence of snow cover on wolf Canis lupus predation patterns in Bieszczady Mountains, Poland. Wildlife Biology, 10(1), 17-23. https://doi.org/10.2981/wlb.2004.005
Haggerty, J. H., Epstein, K., Stone, M., & Cross, P. C. (2018). Land use diversification and intensification on elk winter range in greater Yellowstone: Framework and agenda for social-ecological research. Rangeland Ecology & Management, 71(2), 171-174. https://doi.org/10.1016/j.rama.2017.11.002
Haswell, P. M., Kusak, J., & Hayward, M. W. (2017). Large carnivore impacts are context-dependent. Food Webs, 12, 3-13. https://doi.org/10.1016/j.fooweb.2016.02.005
Hebblewhite, M., Merrill, E. H., & McDonald, T. L. (2005). Spatial decomposition of predation risk using resource selection functions: An example in a wolf-elk predator-prey system. Oikos, 111(1), 101-111. https://doi.org/10.1111/j.0030-1299.2005.13858.x
Hebblewhite, M., & Pletscher, D. H. (2002). Effects of elk group size on predation by wolves. Canadian Journal of Zoology, 80(5), 800-809. https://doi.org/10.1139/z02-059
Hijmans, R. J. (2022). raster: Geographic Data Analysis and Modeling (R package version 3.5-15) [Computer software]. https://CRAN.R-project.org/package=raster
Hosmer, D. W., Lemeshow, S., & Sturdivant, R. X. (2013). Chapter 4. Model-building strategies and methods for logistic regression. In Applied logistic regression (3rd ed.). John Wiley& Sons, Inc.
Kauffman, M. J., Varley, N., Smith, D. W., Stahler, D. R., MacNulty, D. R., & Boyce, M. S. (2007). Landscape heterogeneity shapes predation in a newly restored predator-prey system. Ecology Letters, 10(8), 690-700. https://doi.org/10.1111/j.1461-0248.2007.01059.x
Kertson, B. N., & Marzluff, J. M. (2011). Improving studies of resource selection by understanding resource use. Environmental Conservation, 38(1), 18-27. https://doi.org/10.1017/s0376892910000706
Khorozyan, I., Ghoddousi, A., Soofi, M., & Waltert, M. (2015). Big cats kill more livestock when wild prey reaches a minimum threshold. Biological Conservation, 192, 268-275. https://doi.org/10.1016/j.biocon.2015.09.031
Kunkel, K., & Pletscher, D. H. (2001). Winter hunting patterns of wolves in and near glacier National Park, Montana. The Journal of Wildlife Management, 65(3), 520. https://doi.org/10.2307/3803105
Leblond, M., Dussault, C., & Ouellet, J.-P. (2013). Impacts of human disturbance on large prey species: Do behavioral reactions translate to fitness consequences? PLoS ONE, 8(9), e73695. https://doi.org/10.1371/journal.pone.0073695
Lesmerises, F., Dussault, C., & St-Laurent, M.-H. (2012). Wolf habitat selection is shaped by human activities in a highly managed boreal forest. Forest Ecology and Management, 276, 125-131. https://doi.org/10.1016/j.foreco.2012.03.025
Linnell, J. D. C., Kovtun, E., & Rouart, I. (2021). Wolf attacks on humans: An update for 2002-2020 (NINA Report 1944).
Linnell, J. D. C., Swenson, J. E., & Anderson, R. (2001). Predators and people: Conservation of large carnivores is possible at high human densities if management policy is favourable. Animal Conservation, 4(4), 345-349. https://doi.org/10.1017/s1367943001001408
Lute, M. L., Carter, N. H., López-Bao, J. V., & Linnell, J. D. C. (2018). Conservation professionals agree on challenges to coexisting with large carnivores but not on solutions. Biological Conservation, 218, 223-232. https://doi.org/10.1016/j.biocon.2017.12.035
Mattson, D. J., & Merrill, T. (2002). Extirpations of grizzly bears in the contiguous United States, 1850 -2000. Conservation Biology, 16(4), 1123-1136. https://doi.org/10.1046/j.1523-1739.2002.00414.x
McCorquodale, S. M. (2001). Sex-specific bias in helicopter surveys of elk: Sightability and dispersion effects. The Journal of Wildlife Management, 65(2), 216. https://doi.org/10.2307/3802900
McKay, T. L., Pigeon, K. E., Larsen, T. A., & Finnegan, L. A. (2021). Close encounters of the fatal kind: Landscape features associated with central mountain caribou mortalities. Ecology and Evolution, 11(5), 2234-2248. https://doi.org/10.1002/ece3.7190
McPhee, H. M., Webb, N. F., & Merrill, E. H. (2012). Hierarchical predation: Wolf (Canis lupus) selection along hunt paths and at kill sites. Canadian Journal of Zoology, 90(5), 555-563. https://doi.org/10.1139/z2012-021
Miller, J. R. B. (2015). Mapping attack hotspots to mitigate human-carnivore conflict: Approaches and applications of spatial predation risk modeling. Biodiversity and Conservation, 24(12), 2887-2911. https://doi.org/10.1007/s10531-015-0993-6
Minin, E. D., Slotow, R., Hunter, L. T. B., Pouzols, F. M., Toivonen, T., Verburg, P. H., Leader-Williams, N., Petracca, L., & Moilanen, A. (2016). Global priorities for national carnivore conservation under land use change. Scientific Reports, 6(1), 23814. https://doi.org/10.1038/srep23814
Mumma, M. A., Gillingham, M. P., Johnson, C. J., & Parker, K. L. (2017). Understanding predation risk and individual variation in risk avoidance for threatened boreal caribou. Ecology and Evolution, 7(23), 10266-10277. https://doi.org/10.1002/ece3.3563
Oakleaf, J. K., Murray, D. L., Oakleaf, J. R., Bangs, E. E., Mack, C. M., Smith, D. W., Fontaine, J. A., Jimenez, M. D., Meier, T. J., & Niemeyer, C. C. (2006). Habitat selection by recolonizing wolves in the northern Rocky Mountains of the United States. Journal of Wildlife Management, 70(2), 554-563. https://doi.org/10.2193/0022-541x(2006)70[554:hsbrwi]2.0.co;2
Ordiz, A., Støen, O.-G., Delibes, M., & Swenson, J. E. (2011). Predators or prey? Spatio-temporal discrimination of human-derived risk by brown bears. Oecologia, 166(1), 59-67. https://doi.org/10.1007/s00442-011-1920-5
Oriol-Cotterill, A., Valeix, M., Frank, L. G., Riginos, C., & Macdonald, D. W. (2015). Landscapes of coexistence for terrestrial carnivores: The ecological consequences of being downgraded from ultimate to penultimate predator by humans. Oikos, 124(10), 1263-1273. https://doi.org/10.1111/oik.02224
Paton, D. G., Ciuti, S., Quinn, M., & Boyce, M. S. (2017). Hunting exacerbates the response to human disturbance in large herbivores while migrating through a road network. Ecosphere, 8(6), e01841. https://doi.org/10.1002/ecs2.1841
R Core Team. (2021). R: A language and environment for statistical computing. R Foundation for Statistical Computing. https://www.R-project.org/
Rogala, J., Hebblewhite, M., Whittington, J., White, C., Coleshill, J., & Musiani, M. (2011). Human activity differentially redistributes large mammals in the Canadian Rockies National Parks. Ecology and Society, 16(3), 16. https://doi.org/10.5751/es-04251-160316
Sikes, R. S., Gannon, W. L., & the Animal Care and Use Committee of the American Society of Mammalogists. (2016). Guidelines of the American Society of Mammalogists for the use of wild mammals in research. Journal of Mammalogy, 97, 663-688.
Skovlin, J. M., Bryant, L. D., & Edgerton, P. J. (1989). Effects of timber harvest on elk distribution in the Blue Mountains of Oregon. In Res. Pap. PNW-RP-415. U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station 10 p, 415. https://doi.org/10.2737/pnw-rp-415
Smith, J. A., Suraci, J. P., Clinchy, M., Crawford, A., Roberts, D., Zanette, L. Y., & Wilmers, C. C. (2017). Fear of the human ‘super predator’ reduces feeding time in large carnivores. Proceedings of the Royal Society B: Biological Sciences, 284(1857), 20170433. https://doi.org/10.1098/rspb.2017.0433
Smith, J. A., Wang, Y., & Wilmers, C. C. (2015). Top carnivores increase their kill rates on prey as a response to human-induced fear. Proceedings of the Royal Society B: Biological Sciences, 282(1802), 20142711. https://doi.org/10.1098/rspb.2014.2711
Stankowich, T. (2008). Ungulate flight responses to human disturbance: A review and meta-analysis. Biological Conservation, 141(9), 2159-2173. https://doi.org/10.1016/j.biocon.2008.06.026
Sublette County, Wyoming. (2019). Mapserver. https://greenwoodmap.com/sublette/mapserver/download/
Suraci, J. P., Clinchy, M., Zanette, L. Y., & Wilmers, C. C. (2019). Fear of humans as apex predators has landscape-scale impacts from mountain lions to mice. Ecology Letters, 22(10), 1578-1586. https://doi.org/10.1111/ele.13344
Suraci, J. P., Smith, J. A., Chamaillé-Jammes, S., Gaynor, K. M., Jones, M., Luttbeg, B., Ritchie, E. G., Sheriff, M. J., & Sih, A. (2022). Beyond spatial overlap: Harnessing new technologies to resolve the complexities of predator-prey interactions. Oikos, 2022 (8), e09004. https://doi.org/10.1111/oik.09004
Teton County, Wyoming. (2019). GIS file archive. https://www.greenwoodmap.com/tetonwy/mapserver/download/download.html
Therneau, T. M. (2022). survival: A Package for Survival Analysis in R (R package version 3.3-1) [Computer software]. https://CRAN.R-project.org/package=survival
Thieurmel, B., & Elmarhraoui, A. (2019). Suncalc: Compute sun position, sunlight phases, moon position and lunar phase (R package version 0.5.0) [computer software]. https://CRAN.R-project.org/package=suncalc
Torretta, E., Caviglia, L., Serafini, M., & Meriggi, A. (2017). Wolf predation on wild ungulates: How slope and habitat cover influence the localization of kill sites. Current Zoology, 64(3), zox031. https://doi.org/10.1093/cz/zox031
USFS, United States Forest Service. (2019). Bridger-Teton National Forest GIS Data. https://www.fs.usda.gov/main/btnf/landmanagement/gis
Valeix, M., Hemson, G., Loveridge, A. J., Mills, G., & Macdonald, D. W. (2012). Behavioural adjustments of a large carnivore to access secondary prey in a human-dominated landscape. Journal of Applied Ecology, 49(1), 73-81. https://doi.org/10.1111/j.1365-2664.2011.02099.x
Visscher, D. R., Macleod, I., Vujnovic, K., Vujnovic, D., & Dewitt, P. D. (2017). Human risk induced behavioral shifts in refuge use by elk in an agricultural matrix. Wildlife Society Bulletin, 41(1), 162-169. https://doi.org/10.1002/wsb.741
Wang, Y., Smith, J. A., & Wilmers, C. C. (2017). Residential development alters behavior, movement, and energetics in an apex predator, the puma. PLoS ONE, 12(10), e0184687. https://doi.org/10.1371/journal.pone.0184687
Webb, N. F., Hebblewhite, M., & Merrill, E. H. (2008). Statistical methods for identifying wolf kill sites using global positioning system locations. Journal of Wildlife Management, 72(3), 798-807. https://doi.org/10.2193/2006-566
WGFD, Wyoming Game and Fish Department, U.S. Fish and Wildlife Service, National Park Service, USDA-APHIS-Wildlife Services, & Department, Eastern Shoshone and Northern Arapahoe Tribal Fish and Game Department. (2020). Wyoming gray wolf monitoring and management 2019 annual report.
WGFD, Wyoming Game and Fish Department, U.S. Fish and Wildlife Service, National Park Service, USDA-APHIS-Wildlife Services, & Department, Eastern Shoshone and Northern Arapahoe Tribal Fish and Game Department. (2021). Wyoming gray wolf monitoring and management 2020 annual report.
WGFD, Wyoming Game and Fish Department, U.S. Fish and Wildlife Service, National Park Service, USDA-APHIS-Wildlife Services, & Department, Eastern Shoshone and Northern Arapahoe Tribal Fish and Game Department. (2022). Wyoming gray wolf monitoring and management 2021 annual report.
Whittington, J., Hebblewhite, M., DeCesare, N. J., Neufeld, L., Bradley, M., Wilmshurst, J., & Musiani, M. (2011). Caribou encounters with wolves increase near roads and trails: A time-to-event approach. Journal of Applied Ecology, 48(6), 1535-1542. https://doi.org/10.1111/j.1365-2664.2011.02043.x
Wilmers, C. C., Wang, Y., Nickel, B., Houghtaling, P., Shakeri, Y., Allen, M. L., Kermish-Wells, J., Yovovich, V., & Williams, T. (2013). Scale dependent behavioral responses to human development by a large predator, the puma. PLoS ONE, 8(4), e60590. https://doi.org/10.1371/journal.pone.0060590
Woodruff, S. P., & Jimenez, M. D. (2019). Winter predation patterns of wolves in Northwestern Wyoming. The Journal of Wildlife Management, 83(6), 1352-1367. https://doi.org/10.1002/jwmg.21710
Woodruff, S. P., Jimenez, M., & Johnson, T. R. (2018). Characteristics of winter wolf kill sites in the southern Yellowstone ecosystem in the presence of elk feedgrounds. Journal of Fish and Wildlife Management, 9(1), 155-167. https://doi.org/10.3996/032016-jfwm-024
WYDOT, Wyoming Department of Transportation. (2019). Roadway names SHP. https://gis.wyoroad.info/data/