Relative Habitat Suitability of the Gray Fox (Urocyon cinereoargenteus) in the Central Great Plains

Abstract

Although once abundant, gray foxes (Urocyon cinereoargenteus) are purported to be experiencing a population decline in the central Great Plains, calling for the need for spatially explicit maps for targeted sampling and conservation action. Using public and state data sources, we developed a habitat suitability model across six central US states to examine distributions of relative habitat suitability and land cover associations for the gray fox. Suitable habitat was discontinuous, with numerous small patches predicted across our study area. Gray fox relative habitat suitability increased with Simpson's diversity of land cover composition and decreased with row crop cover. Our model forecasted an apparent range restriction of the gray fox in comparison to historical maps, with suitable habitat only representing 2% of its historical distribution. Our findings add to the growing body of evidence regarding the purported regional declines of gray foxes.

Keywords

central Great Plains / conservation / ensemble model / Habitat Suitability Model / landscape composition / mesocarnivore

References

[1] Allen, M. L., A. C. Avrin, M. J. Farmer, et al. 2021. “Limitations of Current Knowledge about the Ecology of Grey Foxes Hamper Conservation Efforts.” Journal of Threatened Taxa 13, no. 8: 19079–19092. https://doi.org/10.11609/jott.7102.13.8.19079-19092.

[2] Allen, M. L., A. M. Green, and R. J. Moll. 2022. “Modelling the Distribution and Intraguild Associations of an Understudied Mesocarnivore Across the Contiguous United States.” Diversity and Distributions 28, no. 5: 1022–1033. https://doi.org/10.1111/ddi.13502.

[3] Antwi, E. K., R. Krawczynski, and G. Wiegleb. 2008. “Detecting the Effect of Disturbance on Habitat Diversity and Land Cover Change in a Post-Mining Area Using GIS.” Landscape and Urban Planning 87, no. 1: 22–32. https://doi.org/10.1016/j.landurbplan.2008.03.009.

[4] Aubry, K. B., C. M. Raley, and K. S. McKelvey. 2017. “The Importance of Data Quality for Generating Reliable Distribution Models for Rare, Elusive, and Cryptic Species.” PLoS One 12, no. 6: e0179152. https://doi.org/10.1371/journal.pone.0179152.

[5] Barbet-Massin, M., F. Jiguet, C. H. Albert, and W. Thuiller. 2012. “Selecting Pseudo-Absences for Species Distribution Models: How, Where and How Many?” Methods in Ecology and Evolution 3, no. 2: 327–338. https://doi.org/10.1111/j.2041-210X.2011.00172.x.

[6] Bauder, J. M., M. L. Allen, T. J. Benson, C. A. Miller, and K. W. Stodola. 2021. “An Approach for Using Multiple Indices for Monitoring Long-Term Trends of Mesopredators at Broad Spatial Scales.” Biodiversity and Conservation 30, no. 12: 3529–3547. https://doi.org/10.1007/s10531-021-02259-8.

[7] Belant, J. L., J. Schipper, and J. Conroy. 2009. “The Conservation Status of Small Carnivores in the Americas.” Small Carnivore Conservation 41: 3–8.

[8] Benedict, R. A., H. H. Genoways, and P. W. Freeman. 2000. “Shifting Distributional Patterns of Mammals in Nebraska.” Transactions of the Nebraska Academy of Sciences and Affiliated Societies 26: 55–84.

[9] Benson, D. J., J. D. La Peña, A. D. George, and C. C. Rega-Brodsky. 2024. “County Records and Noteworthy Detections of Gray Fox (Urocyon cinereoargenteus) in Southeast Kansas.” Transactions of the Kansas Academy of Science 127, no. 1–2: 1–8. https://doi.org/10.1660/062.127.0101.

[10] Brown, C. H., and H. P. Griscom. 2022. “Differentiating Between Distribution and Suitable Habitat in Ecological Niche Models: A Red Spruce (Picea rubens) Case Study.” Ecological Modelling 472: 110102. https://doi.org/10.1016/j.ecolmodel.2022.110102.

[11] Cheeseman, A. E., D. S. Jachowski, and R. Kays. 2024. “From Past Habitats to Present Threats: Tracing North American Weasel Distributions Through a Century of Climate and Land Use Change.” Landscape Ecology 39, no. 5: 104. https://doi.org/10.1007/s10980-024-01902-3.

[12] Chen, C., A. Liaw, and L. Breiman. 2004. “Using Random Forest to Learn Imbalanced Data.” Technical Report, Department of Statistics, University of California, Berkeley 110, no. 1–12: 24.

[13] Choate, J. R., and J. E. Krause. 1974. “Historical Biogeography of the Gray Fox (Urocyon cinereoargenteus) in Kansas.” Transactions of the Kansas Academy of Science (1903-) 77, no. 4: 231–236. https://doi.org/10.2307/362709.

[14] Cooper, S. E., C. K. Nielsen, and P. T. McDonald. 2012. “Landscape Factors Affecting Relative Abundance of Gray Foxes Urocyon cinereoargenteus at Large Scales in Illinois, USA.” Wildlife Biology 18, no. 4: 366–373. https://doi.org/10.2981/11-093.

[15] Cove, M. V., R. Kays, H. Bontrager, et al. 2021. “SNAPSHOT USA 2019: A Coordinated National Camera Trap Survey of the United States.” Ecology 102, no. 6: e03353. https://doi.org/10.1002/ecy.3353.

[16] Crooks, K. R., C. L. Burdett, D. M. Theobald, C. Rondinini, and L. Boitani. 2011. “Global Patterns of Fragmentation and Connectivity of Mammalian Carnivore Habitat.” Philosophical Transactions of the Royal Society, B: Biological Sciences 366, no. 1578: 2642–2651. https://doi.org/10.1098/rstb.2011.0120.

[17] Deuel, N. R., L. M. Conner, K. V. Miller, M. J. Chamberlain, M. J. Cherry, and L. V. Tannenbaum. 2017. “Habitat Selection and Diurnal Refugia of Gray Foxes in Southwestern Georgia, USA.” PLoS One 12, no. 10: e0186402. https://doi.org/10.1371/journal.pone.0186402.

[18] Egan, M. E., C. C. Day, T. E. Katzner, and P. A. Zollner. 2021. “Relative Abundance of Coyotes (Canis latrans) Influences Gray Fox (Urocyon cinereoargenteus) Occupancy Across the Eastern United States.” Canadian Journal of Zoology 99, no. 2: 63–72. https://doi.org/10.1139/cjz-2019-0246.

[19] Fahrig, L., J. Baudry, L. Brotons, et al. 2011. “Functional Landscape Heterogeneity and Animal Biodiversity in Agricultural Landscapes.” Ecology Letters 14, no. 2: 101–112. https://doi.org/10.1111/j.1461-0248.2010.01559.x.

[20] Fielding, A. H., and J. F. Bell. 1997. “A Review of Methods for the Assessment of Prediction Errors in Conservation Presence/Absence Models.” Environmental Conservation 24, no. 1: 38–49. https://doi.org/10.1017/S0376892997000088.

[21] Fraisl, D., G. Hager, B. Bedessem, et al. 2022. “Citizen Science in Environmental and Ecological Sciences.” Nature Reviews Methods Primers 2, no. 1: 64. https://doi.org/10.1038/s43586-022-00144-4.

[22] Friedman, J. H.2001. “Greedy Function Approximation: A Gradient Boosting Machine.” Annals of Statistics 29, no. 5: 1189–1232. https://doi.org/10.1214/aos/1013203451.

[23] Fritzell, E. K., and K. J. Haroldson. 1982. “Urocyon cinereoargenteus.” Mammalian Species 189: 1–8.

[24] GBIF.org. 2025. [Dataset]. GBIF Occurrence Download; 04/05/2025.

[25] Gorelick, N., M. Hancher, M. Dixon, S. Ilyushchenko, D. Thau, and R. Moore. 2017. “Google Earth Engine: Planetary-Scale Geospatial Analysis for Everyone.” Remote Sensing of Environment 202: 18–27. https://doi.org/10.1016/j.rse.2017.06.031.

[26] Guisan, A., and W. Thuiller. 2005. “Predicting Species Distribution: Offering More Than Simple Habitat Models.” Ecology Letters 8, no. 9: 993–1009. https://doi.org/10.1111/j.1461-0248.2005.00792.x.

[27] Hastie, T., R. Tibshirani, T. Hastie, and R. Tibshirani. 1986. “Generalized Additive Models.” Statistical Science 1, no. 3: 297–318.

[28] Hesselbarth, M. H. K., M. Sciaini, K. A. With, K. Wiegand, and J. Nowosad. 2019. “Landscapemetrics: An Open-Source R Tool to Calculate Landscape Metrics.” Ecography 42: 1648–1657. https://doi.org/10.1111/ecog.04617.

[29] Hirzel, A. H., G. Le Lay, V. Helfer, C. Randin, and A. Guisan. 2006. “Evaluating the Ability of Habitat Suitability Models to Predict Species Presences.” Ecological Modelling 199, no. 2: 142–152. https://doi.org/10.1016/j.ecolmodel.2006.05.017.

[30] iNaturalist. 2025. https://www.inaturalist.org.

[31] Jachowski, D. S., C. J. Marneweck, C. Olfenbuttel, and S. N. Harris. 2024. “Support for the Size-Mediated Sensitivity Hypothesis Within a Diverse Carnivore Community.” Journal of Animal Ecology 93, no. 1: 109–122. https://doi.org/10.1111/1365-2656.13916.

[32] Johnston, A., N. Moran, A. Musgrove, D. Fink, and S. R. Baillie. 2020. “Estimating Species Distributions From Spatially Biased Citizen Science Data.” Ecological Modelling 422: 108927. https://doi.org/10.1016/j.ecolmodel.2019.108927.

[33] Landholt, L. M., and H. H. Genoways. 2000. “Population Trends in Furbearers in Nebraska.” Transactions of the Nebraska Academy of Sciences and Affiliated Societies 26: 97–110.

[34] Larreur, M. R., C. K. Nielsen, D. B. Lesmeister, and G. Bastille-Rousseau. 2025. “The Precipitous Decline of a Gray Fox Population.” Global Ecology and Conservation 58: e03441. https://doi.org/10.1016/j.gecco.2025.e03441.

[35] Luoto, M., R. Virkkala, and R. K. Heikkinen. 2007. “The Role of Land Cover in Bioclimatic Models Depends on Spatial Resolution.” Global Ecology and Biogeography 16: 34–42. https://doi.org/10.1111/j.1466-822x.2006.00262.x.

[36] MacKenzie, D. I., J. D. Nichols, J. E. Hines, M. G. Knutson, and A. B. Franklin. 2003. “Estimating Site Occupancy, Colonization, and Local Extinction When a Species Is Detected Imperfectly.” Ecology 84, no. 8: 2200–2207. https://doi.org/10.1890/02-3090.

[37] Marmion, M., M. Parviainen, M. Luoto, R. K. Heikkinen, and W. Thuiller. 2009. “Evaluation of Consensus Methods in Predictive Species Distribution Modelling.” Diversity and Distributions 15, no. 1: 59–69. https://doi.org/10.1111/j.1472-4642.2008.00491.x.

[38] Monsarrat, S., A. F. Boshoff, and G. I. H. Kerley. 2019. “Accessibility Maps as a Tool to Predict Sampling Bias in Historical Biodiversity Occurrence Records.” Ecography 42, no. 1: 125–136. https://doi.org/10.1111/ecog.03944.

[39] Morin, D. J., D. B. Lesmeister, C. K. Nielsen, and E. M. Schauber. 2018. “The Truth about Cats and Dogs: Landscape Composition and Human Occupation Mediate the Distribution and Potential Impact of Non-Native Carnivores.” Global Ecology and Conservation 15: e00413. https://doi.org/10.1016/j.gecco.2018.e00413.

[40] Morin, D. J., D. B. Lesmeister, C. K. Nielsen, and E. M. Schauber. 2022. “Asymmetrical Intraguild Interactions With Coyotes, Red Foxes, and Domestic Dogs May Contribute to Competitive Exclusion of Declining Gray Foxes.” Ecology and Evolution 12, no. 7: e9074. https://doi.org/10.1002/ece3.9074.

[41] Oldfather, M. F., M. M. Kling, S. N. Sheth, N. C. Emery, and D. D. Ackerly. 2020. “Range Edges in Heterogeneous Landscapes: Integrating Geographic Scale and Climate Complexity Into Range Dynamics.” Global Change Biology 26, no. 3: 1055–1067. https://doi.org/10.1111/gcb.14897.

[42] Palomo-Munoz, G., M. Fidino, T. J. Werdel, et al. 2024. “Mesopredators Have Differing Influences on Prey Habitat Use and Diel Activity in a Multipredator Landscape.” Ecosphere 15, no. 9: e70006. https://doi.org/10.1002/ecs2.70006.

[43] Pearson, R. G., T. P. Dawson, and C. Liu. 2004. “Modelling Species Distributions in Britain: A Hierarchical Integration of Climate and Land-Cover Data.” Ecography 27, no. 3: 285–298. https://doi.org/10.1111/j.0906-7590.2004.03740.

[44] Perkins, J. C., A. A. Gibson, B. D. Wolaver, B. J. Labay, J. P. Pierre, and R. C. Dowler. 2022. “An Evaluation of Detection Methods for the Plains Spotted Skunk.” Wildlife Society Bulletin 46, no. 5: e1376. https://doi.org/10.1002/wsb.1376.

[45] Phillips, S. J., R. P. Anderson, M. Dudík, R. E. Schapire, and M. E. Blair. 2017. “Opening the Black Box: An Open-Source Release of Maxent.” Ecography 40, no. 7: 887–893. https://doi.org/10.1111/ecog.03049.

[46] Polder, E.1958. “Recent Locality Records for Some Iowa Mammals.” Proceedings of the Iowa Academy of Science 65, no. 1: 559–564.

[47] R Core Team. 2024. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing. https://www.R-project.org/.

[48] Radosavljevic, A., and R. P. Anderson. 2014. “Making Better Maxent Models of Species Distributions: Complexity, Overfitting, and Evaluation.” Journal of Biogeography 41, no. 4: 629–643. https://doi.org/10.1111/jbi.12227.

[49] Roberts, D. R., V. Bahn, S. Ciuti, et al. 2017. “Cross-Validation Strategies for Data With Temporal, Spatial, Hierarchical, or Phylogenetic Structure.” Ecography 40: 913–929. https://doi.org/10.1111/ecog.02881.

[50] Samson, F. B., F. L. Knopf, and W. R. Ostlie. 2004. “Great Plains Ecosystems: Past, Present, and Future.” Wildlife Society Bulletin 32, no. 1: 6–15. https://doi.org/10.2193/0091-7648(2004)32[6:GPEPPA]2.0.CO;2.

[51] Schipper, J., M. Hoffmann, J. W. Duckworth, and J. Conroy. 2008. “The 2008 IUCN Red Listings of the World's Small Carnivores.” Small Carnivore Conservation 39: 29–34.

[52] Schmidt, C. J., M. Peek, and G. A. Kauffman, et al. 2021. Kansas Mammal Atlas: An On-line Reference. Electronic Database. https://webapps.fhsu.edu/ksmammal.

[53] Schnell, G. D., J. A. Grzybowski, D. J. Hough, and T. M. McKenna. 1985. “Evaluation of Spatial Patterning in Oklahoma Furbearer Populations.” Southwestern Naturalist 30, no. 2: 225. https://doi.org/10.2307/3670736.

[54] Taraporevala, N. F., J. P. Beckmann, and J. K. Young. 2025. “Citizen Science Project on Urban Canids Provides Different Results From Camera Traps but Generates Interest and Revenue.” Wildlife Biology Early View.−/+ e01382. https://doi.org/10.1002/wlb3.01382.

[55] Thuiller, W., D. Georges, M. Gueguen, et al. 2025. biomod2: Ensemble platform for Species Distribution Modeling. R Package Version 4.2–6–2. https://CRAN.R-project.org/package=biomod2.

[56] Tudor-Locke, C., W. D. Johnson, and P. T. Katzmarzyk. 2009. “Accelerometer-Determined Steps Per Day in US Adults.” Medicine and Science in Sports and Exercise 41, no. 7: 1384–1391. https://doi.org/10.1249/MSS.0b013e318199885c.

[57] U.S. Geological Survey (USGS). 2024. Annual NLCD Collection 1 Science Products (ver. 1.1, June 2025): U.S. Geological Survey Data Release. https://doi.org/10.5066/P94UXNTS.

[58] Valavi, R., G. Guillera-Arroita, J. J. Lahoz-Monfort, and J. Elith. 2022. “Predictive Performance of Presence-Only Species Distribution Models: A Benchmark Study With Reproducible Code.” Ecological Monographs 92, no. 1: e01486. https://doi.org/10.1002/ecm.1486.

[59] Werdel, T. J., C. W. Piper, A. M. Ricketts, A. A. Ahlers, and A. A. Ahlers. 2022. “Scale-Specific Landscape Effects Impose Range-Limiting Constraints on the Distribution of a Prairie-Obligate Carnivore.” Landscape Ecology 37, no. 8: 2065–2079. https://doi.org/10.1007/s10980-022-01466-0.

[60] Werdel, T. J., C. W. Piper, A. M. Ricketts, M. S. Peek, and A. A. Ahlers. 2023. “Direct and Indirect Landscape-Scale Pathways Structure Carnivore Communities in an Agro-Prairie Ecosystem.” Journal of Mammalogy 104, no. 6: 1279–1289. https://doi.org/10.1093/jmammal/gyad065.

[61] Zumbaugh, D. M., and J. R. Choate. 1985. “Historical Biogeography of Foxes in Kansas.” Transactions of the Kansas Academy of Science (1903-) 88, no. 1/2: 1–13. https://doi.org/10.2307/3628361.

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