Distribution and Relative Abundance of Eastern Spotted Skunk Records Across Their Range
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Ecology and Conservation of the Eastern Spotted Skunk 2021 Southeastern Naturalist 20(Special Issue 11):13–23 Distribution and Relative Abundance of Eastern Spotted Skunk Records Across Their Range Roger W. Perry1,*, D. Blake Sasse2, J. Clint Perkins3, and Nicholas W. Sharp4 Abstract - Evidence suggests the range of Spilogale putorius (Eastern Spotted Skunk) has contracted and its abundance has declined in the past 70 years, leading to conserva- tion concerns. We summarized county records of Eastern Spotted Skunks collected during 2000–2020 to determine the current range and relative abundance of the species. We ac- cumulated 1174 records from 257 counties across its historic range in the United States, with 901 records from 197 counties considered verified. Verified records included museum specimens, photo-documented occurrences, and captures by researchers. We created 2 dis- tribution maps: one of their current range based on all occurrence records and another from only verified records. Records indicated the Eastern Spotted Skunk persisted across a large portion of its historic range, and is relatively abundant in the Interior Highlands, Appala- chian Mountains, central Texas, central South Dakota, and south Florida. Our results also suggest that the species’ overall range has contracted since 1959. Regions with a relatively high abundance of current records covered a variety of ecosystems, including agricultural areas, grasslands, woodlands, and forests. These data provide managers with information concerning where research and conservation efforts can be focused for this potentially de- clining species. Introduction Found across much of the United States (US), Spilogale putorius (L.) (Eastern Spotted Skunk) is a small mesocarnivore that ranges from Florida to Texas and west to the Rocky Mountains, and north to Minnesota (Kinlaw 1995). Van Gelder (1959) presented one of the first estimates of the geographic range of the Eastern Spotted Skunk. However, using records from fur-trapping magazines and museum specimens, Sasse (2017) estimated the species’ range in 1900−1920 covered ~60% of the area estimated in 1959 and suggested the Eastern Spotted Skunk expanded its range into additional areas in the mid-1900s, including the northern Great Plains and parts of the Mississippi Alluvial Valley of the central US. As evidenced primar- ily by fur trapping reports, the species declined in abundance in the second half of the twentieth century, and one subspecies, S.p. interrupta Rafinesque (Plains Spot- ted Skunk), has been proposed for protection under the federal Endangered Species Act (Gompper 2017, Gompper and Hackett 2005, USFWS 2012). The decline in abundance of Eastern Spotted Skunks may have coincided with range contractions in some states. Recent incidental captures by fur trappers in Arkansas have been limited to the Ozark and Ouachita Mountains, although the 1 USDA Forest Service, Southern Research Station, Hot Springs, AR 71902. 2Arkansas Game and Fish Commission, Mayflower, AR 72106. 3Department of Natural Resources Management, Texas Tech University, Lubbock, TX 79409, 4Alabama Wildlife and Freshwa- ter Fisheries, Tanner, AL 35671. *Corresponding author - roger.perry@usda.gov. Manuscript Editor: Steven Castleberry 13
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp species was once found nearly statewide (Sasse 2018, Sasse and Gompper 2006). In South Dakota, occurrence was reduced from 65 counties in 1983 to 54 counties in 1993 (Blumberg et al. 1997). Choate et al. (1973) reported possible extirpation of the species from western Kansas, and Dowler et al. (2017) concluded that its range in Texas may have contracted such that it was now only present in the central part of the state. Due to perceived range contraction and decline in numbers, land managers need current information on the distribution of Eastern Spotted Skunks so that research and conservation efforts can be concentrated in areas with the greatest probability of encountering the species. We used county records accumulated during 2000– 2020 to update the distribution and relative abundance of Eastern Spotted Skunks. Methods We accumulated locality records of Eastern Spotted Skunks using county as our datum. We requested locality information from state and federal biologists, researchers, and others across the current and historic range of Eastern Spotted Skunks (Appendix 1). We included data from museum collections, published re- search (Appendix 2), iNaturalist reports with photo documentation (iNaturalist 2020), state natural history databases, museum collections reported in biodiversity websites, capture and camera-trap data from government or university researchers, and public sightings reported to state wildlife agencies. We estimated the reliability of each record based on the source and documenta- tion, such as photos or collected specimens. We generally considered observational records from the public, fur trappers, and other non-wildlife professionals with- out photo documentation or validation by wildlife professionals to be unverified because of frequent confusion with Mephitis mephitis Schreber (Striped Skunk; Sasse 2018). We considered reports verified if they fell into one of the following categories: (1) specimens collected for museums; (2) photo documentation was provided, such as game-camera photos; (3) track-plate data collected by wildlife professionals; (4) captures by researchers or presented in published research; (5) tissue samples collected for genetic or other studies; (6) furs presented at sales by trappers; (7) visual sightings by professional wildlife biologists or research- ers; (8) captures by the public and seen by wildlife professionals; (9) mortalities (i.e., roadkills) documented by professional wildlife biologists; (10) observations reported by the public with specific details of behavior (i.e., arboreal behavior) that differentiated Spotted Skunks from Striped Skunks; and (11) records from wildlife rehabilitators. We considered records unverified when wildlife agencies reported “sighting” or other location data without additional information on the source or type of those records. We also considered fur-trapper reports unverified unless con- firmed by a state official or at a fur sale. We created 2 distribution maps; one using all records collected and another presenting only verified records. We arbitrarily included only records collected after 1 January 2000 so that oc- currences were considered recent. We did not include county records when the date of collection was not recorded. Additional records collected as part of on-going 14
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp research were not included so that researchers could publish their findings first. These records were minimal, and generally occurred in regions where other records occurred, such as in the central Appalachians of Virginia and West Virginia; recent records of Eastern Spotted Skunks from Wyoming were also not included because of ongoing research. We included an estimate of the relative abundance of Eastern Spotted Skunks in each county using total number of records from that county. We delineated rela- tive abundance into 4 classes: 0 records, 1 record, 2–5 records, and >5 records. We considered multiple photographs from the same site (e.g., camera-trap station) to be a single record because baited camera stations often record the same individual multiple times (Gerber et al. 2012). We considered each individual skunk a separate record when multiple captures of different individuals occurred at the same location during live trapping. Results We accumulated 1174 records of Eastern Spotted Skunks from 257 counties across their historic range (Fig. 1). Of these, 901 records from 197 counties were considered verified (Fig. 2). Forty-four percent of counties had only a single record of Eastern Spotted Skunks during 2000−2020 (38% of verified records), whereas 56% of counties had 2 or more records (62% of verified counties) (Fig. 3). The greatest number of both total and verified records were from Osceola County, FL (n = 216 total and verified), followed by Scott County, AR (n = 54 total and 47 verified), and Oconee County, SC (n = 39 total and verified). Additional notewor- thy counties with verified records included Macon County, NC (n = 35); Coryell County, TX (n = 17); Le Flore County, OK (n = 15); Harris County, TX (n = 14); and Hand County, SD (n = 14). Only single records occurred in Minnesota, Pennsyl- vania, and North Dakota and no Eastern Spotted Skunks were recorded since 2000 in Louisiana, Colorado, Maryland, or Wisconsin. Both verified and unverified records indicated Eastern Spotted Skunks per- sist from north-central North Dakota to central Texas, eastward to southwestern Pennsylvania and southern Florida. Both verified records and total records showed a similar distribution pattern. However, regions where only unverified records occurred included east-central North Carolina, western Tennessee, and northern Kansas. Verified records suggested relatively higher abundances (based on total number of verified records) occurred in the Interior Highlands of Missouri, Arkan- sas, and Oklahoma, and in the Appalachian Mountains from Virginia to northern Alabama. Other areas of high relative abundance included central and southeastern Texas, central South Dakota, and south Florida. Discussion Recent records of Eastern Spotted Skunks covered regions similar to their his- toric distribution in 1900–1920 (Sasse 2017). However, we found no recent records from eastern Iowa, southern Texas, eastern Colorado, and the Texas panhandle 15
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp where they apparently occurred in both 1900–1920 and 1959 (Van Gelder 1959). We recorded 257 counties with records, whereas Sasse (2017) suggested the range in 1900−1920 covered ~909 counties (a 72% smaller area in 2020). However, range maps are typically drawn to encompass a series of disjunct points where or- ganisms occurred and they often include areas where the species was not actually identified; thus, some counties covered by the map in Sasse (2017) may have not contained Eastern Spotted Skunks, and many of the counties in our data set may have had undetected skunks. We found abundant records from central South Dako- ta, which was outside the presumed range of Eastern Spotted Skunks in 1900–1920 (Sasse 2017) but included in the estimated range in 1959 (Van Gelder 1959). Van Gelder (1959) indicated that Louisiana was an area of integration between the subspecies S.p. putorius and S.p. interrupta, and northern Florida was an area of integration between the subspecies S.p. putorius and S. p. ambarvalis, but Eastern Spotted Skunks now appear rare or absent from both of these areas. We did not include records from Wyoming because state officials lacked con- fidence in the identifications due to their locations from across the state where Figure 1. Distribution of Eastern Spotted Skunk reports and relative abundance of records by county across their range; includes all records, 2000−2020. 16
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp Spilogale gracilis Merriam (Western Spotted Skunk) also occurs. Most records received from Wyoming were identified only as Spilogale, and only 9 were identi- fied as Eastern Spotted Skunks. Western and Eastern Spotted Skunks both occur in Wyoming but currently cannot reliably be distinguished without genetic testing (e.g., ESSCSG 2019), and the range boundary between the 2 species is unclear. Further research to distinguish the distribution of these 2 species in Wyoming was ongoing. Both species of spotted skunks also occur in Texas and have an area of potential sympatry in the central portion of the state (Dowler et al. 2008). For cen- tral Texas, we incorporated records from a recent genetic analysis (Shaffer et al. 2018), individuals with a confirmed genetic identification but not yet reported (R. Dowler, Angelo State University, San Angelo, TX, unpubl. data), individuals from on-going research, and photo records that had been vetted for species identification. Therefore, we are confident in the presented distribution of Eastern Spotted Skunks in central Texas. Even in areas with numerous records, this species is likely rare and often not easily captured or photographed with game cameras (e.g., Fino et al. 2019, Figure 2. Distribution of Eastern Spotted Skunk reports and relative abundance of records by county across their range; includes only verified records, 2000−2020. 17
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp Hackett et al. 2007, Higdon and Gompper 2020). For example, in Scott County, AR, Hackett et al. (2007) reported 0.81% capture success over 12,404 trap nights. In Oconee County, SC, Eng and Jachowski (2019) reported a 2.8% detection rate across 4689 camera-trap nights, and Fino et al. (2019) reported
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp may exist but simply lacked sufficient sampling. Nevertheless, the overall regions of occurrence generally remain valid regardless of perceived relative abundance. Counties where abundant records occurred were dominated by a wide variety of ecosystems. Abundant records occurred from counties dominated by open pasture, grasslands, and row-crop agriculture, (e.g., central South Dakota; Fino et al. 2019), dry coastal grasslands (e.g., Florida; Harris et al. 2020), woodlands (e.g., Texas; Dowler et al. 2017), and forest (e.g., Arkansas, Missouri, Virginia, West Virginia, and Alabama). In forests, there were multiple verified records from sites dominated by various forest types, including mixed pines (Pinus spp.) and oaks (Quercus spp.) in Arkansas (Perry et al. 2018) West Virginia (Thorne and Waggy 2017), and Alabama (Benson et al. 2019); oak (Quercus spp.)-hickory (Carya spp.) forests of Missouri (Hackett et al. 2007); and high-elevation forests of Picea rubens Sarg. (Red Spruce) in Virginia (Diggins et al. 2015). Consequently, they appear to be distributed among a wide variety of habitats across their range, and it is unclear if they have declined more or less in certain habitats or whether other factors such as disease may be primarily contributing to their perceived decline. Information on the ecology of this species has only recently began to appear, and recent increase in interest has spurred efforts to collect data on presence us- ing non-invasive methods such as game cameras. Additional resources that allow the quick distribution of information on skunk locations, such as iNaturalist, have also increased our knowledge of their distribution and abundance. These data pro- vide valuable information on presence and relative abundance to managers, which is critical to conservation planning and species assessments. Our results suggest that the current overall range of Eastern Spotted Skunks is similar to the range in 1900−1920 (Sasse 2017), although contractions have occurred in some areas. How- ever, larger range contractions may have occurred since their range was estimated in 1959 (Van Gelder 1959). Although our study provides general locations where Eastern Spotted Skunks can still be found, we cannot make conclusions on the cur- rent abundance of Eastern Spotted Skunks compared with past decades. Acknowledgments See Appendix 1 for a list of individuals that contributed data and information on Eastern Spotted Skunk records. We thank W.M. Ford and D.S. Jachowski for review of an earlier draft. Literature Cited Benson, I.W., T.L. Sprayberry, W.C. Cornelison, and A.J. Edelman. 2019. Rest-site activity patterns of Eastern Spotted Skunks in Alabama. Southeastern Naturalist 18:165−172. Blumberg, C.A., K.F. Higgins, and J.A. Jenks. 1997. Use of a mail survey to determine pres- ent mammal distributions in South Dakota. Proceedings of the South Dakota Academy of Science 76:75−89. Choate, J.R., E.D. Fleharty, and R.J. Little. 1973. Status of the Spotted Skunk, Spilogale putorius, in Kansas. Transactions of the Kansas Academy of Science 76:226−233. Diggins, C.D., D.S. Jachowski, J. Martin, and W.M. Ford. 2015. Incidental captures of Eastern Spotted Skunk in a high-elevation Red Spruce forest in Virginia. Northeastern Naturalist 22:N6−N10. 19
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp Dowler, R.C., C.E. Ebeling, G.I. Guerra, and A.W. Ferguson. 2008. The distribution of Spotted Skunks, Genus Spilogale, in Texas. Texas Journal of Science 60:321−326. Dowler, R.C., J.C. Perkins, A.A. Shaffer, B.D. Wolaver, B.J. Labay, J.P. Pierre, A.W. Fergu- son, M.M. McDonough, and L.K. Ammerman. 2017. Conservation status of the Plains Spotted Skunk, Spilogale putorius interrupta, in Texas, with an assessment of genetic variability in the species. Report to the Texas Comptrollers Office, Austin, TX. 143 pp. Eng, R.Y.Y., and D.S. Jachowski. 2019. Evaluating detection and occupancy probabilities of Eastern Spotted Skunks. Journal of Wildlife Management. 83:1244−1253. Eastern Spotted Skunk Cooperative Study Group (ESSCSG). 2019. Eastern Spotted Skunk conservation plan. 43 pp. Available online at https://easternspottedskunk.weebly.com/ uploads/3/9/7/0/39709790/ess_conservation_plan_dec92020.pdf. Earlier version ac- cessed 30 July 2020. Fino, S., J.D. Stafford, A.T. Pearse, and J.A. Jenks. 2019. Incidental captures of Plains Spot- ted Skunks in central South Dakota. Prairie Naturalist 51:33−36. Gerber, B.D., S.M. Karpanty, and M.J. Kelly. 2012. Evaluating the potential biases in carnivore capture–recapture studies associated with the use of lure and varying density estimation techniques using photographic-sampling data of the Malagasy civet. Popula- tion Ecology 54:43−54. Gompper, M.E. 2017. Range decline and landscape ecology of the Eastern Spotted Skunk. Pp. 478–492, In D.W. Macdonald, C. Newman, and L.A. Harrington (Eds.). Biology and Conservation of the Musteloids. Oxford University Press, Oxford, UK. 672 pp. Gompper, M.E., and H.M. Hackett. 2005. The long-term range-wide decline of a once com- mon carnivore: The Eastern Spotted Skunk (Spilogale putorius). Animal Conservation 8:195−201. Hackett, H.M., D.B. Lesmeister, J. Desanty-Combes, W.G. Montague, J.J. Millspaugh, and M.E. Gompper. 2007. Detection rates of Eastern Spotted Skunks (Spilogale putorius) in Missouri and Arkansas using live-capture and non-invasive techniques. American Midland Naturalist 158:123−131. Harris, S.N. 2018. Florida Spotted Skunk ecology in a dry prairie ecosystems. M.Sc. Thesis. Clemson University, Clemson, SC. 69 pp. Harris, S.N., T.J. Doonan, E.L. Hewett Rangheb, and D.S. Jachowski. 2020. Den-site selec- tion by the Florida Spotted Skunk. Journal of Wildlife Management 84:127−137. Higdon, S.D. and M.E. Gompper. 2020. Rest-site use and the apparent rarity of an Ozark population of Plains Spotted Skunk (Spilogale putorius interrupta). Southeastern Natu- ralist 19:73–88. Hurtado, L.C. 2018. Spatial and temporal structure of a canid community in Nebraska. Ph.D. Dissertation. University of Nebraska, Lincoln, NE. 250 pp. iNaturalist 2020. Social network for naturalists. Home page. Available from https://www. inaturalist.org. Accessed 10 June 2020. Kinlaw, A. 1995. Spilogale putorius. Mammalian Species 551:1–7. Lesmeister, D.B., M.E. Gompper, and J.J. Millspaugh. 2009. Habitat selection and home- range dynamics of Eastern Spotted Skunks in the Ouachita Mountains, Arkansas, USA. Journal of Wildlife Management 73:18–25 Perry, R.W., D.C. Rudolph, and R.E. Thill. 2018. Capture-site characteristics for Eastern Spotted Skunks in mature forests during summer. Southeastern Naturalist 17:298–308. Sasse, D.B. 2017. Distribution of the Eastern Spotted Skunk in the early twentieth century. Journal of the Arkansas Academy of Science 71:219−220. 20
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp Sasse, D.B. 2018. Incidental captures of Plains Spotted Skunks (Spilogale putorius inter- rupta) by Arkansas trappers, 2012–2017. Journal of the Arkansas Academy of Science 72:187−189. Sasse, D.B., and M.E. Gompper. 2006. Geographic distribution and harvest dynamics of the Eastern Spotted Skunk in Arkansas. Journal of the Arkansas Academy of Science 60:119−124. Shaffer, A.A., R.C. Dowler, J.C. Perkins, A.W. Ferguson, M.M. McDonough, and L.K. Ammerman. 2018. Genetic variation in the Eastern Spotted Skunk (Spilogale putoirus) with emphasis on the Plains Spotted Skunk (S. p. interrupta). Journal of Mammalogy 99:1237−1248. Thorne, E.D. 2020. Spatial ecology of a vulnerable species: Home-range dynamics, re- source use, and genetic differentiation of Eastern Spotted Skunks in central Appalachia. Ph.D. Dissertation. Virginia Polytechnic Institute and State University, Blacksburg, VA. 210 pp. Thorne, E.D., and C. Waggy. 2017. First reported observation of food provisioning to offspring by an Eastern Spotted Skunk, a small carnivore. Northeastern Naturalist 24:N1−N4. Thorne, E.D., C. Waggy, D.S. Jachowski, M.J. Kelly, and W.M. Ford. 2017. Winter habitat associations of Eastern Spotted Skunks in Virginia. Journal of Wildlife Management 81:1042–1050. United States Fish and Wildlife Service (USFWS). 2012. 90-Day finding on a petition to list the Prairie Gray Fox, the Plains Spotted Skunk, and a distinct population segment of the Mearn’s Eastern Cottontail in east central Illinois and Western Indiana as endangered or threatened species. Federal Register 77:71759−71771. Available online at https://www. federalregister.gov/documents/2012/12/04/2012-29188/endangered-and-threatened- wildlife-and-plants-90-day-finding-on-a-petition-to-list-the-prairie-gray. Accessed 12 January 2021. Van Gelder, R.G. 1959. A taxonomic revision of the spotted skunks (Genus Spilogale). Bul- letin of the American Museum of Natural History 117:229−392. White, K.M. 2019. Canid space use and temporal activity within a Black-Footed Ferret recovery site in western Kansas. M.Sc. Thesis. University of Wisconsin, Stevens Point, WI. 60 pp. 21
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp Appendix 1. State biologists, researchers, and individuals that provided data on Eastern Spotted Skunk records collected during the period 2000−2020, alphabetized by state. Person State Affiliation Eric Odell CO Colorado Parks and Wildlife Mark Danaher FL United States Fish and Wildlife Service William Freund FL fStop Foundation, Inc. Daniel Smith FL University of Central Florida Stephen Harris FL Clemson University Alexandra A. Gibson FL Angelo State University Katrina Morris GA Georgia Department of Natural Resources Vince Evelsizer IA Iowa Department of Natural Resources Zachary Cordes KS Kansas Department of Wildlife, Parks & Tourism Shelby Fulton KY Office of Kentucky Nature Preserves Jennifer Hogue-Manuel LA Louisiana Department of Wildlife and Fisheries Harry Spiker MD Maryland Department of Natural Resources Gerda Nordquist MN Minnesota Department of Natural Resources Laura Conlee MO Missouri Department of Conservation Janet Sternburg MO Missouri Department of Conservation Katelin Cross MS Mississippi Museum of Natural Science Ashley Hobbs NC North Carolina Wildlife Resources Commission Colleen Olfenbuttel NC North Carolina Wildlife Resources Commission Stephanie Tucker ND North Dakota Game and Fish Shaun Dunn NE Nebraska Game and Parks Commission Matthew Fullerton OK Oklahoma Department of Wildlife Conservation Greg Turner PA Pennsylvania Game Commission David Scott Jachowski SC Clemson University Jay Butfiloski SC South Carolina Department of Natural Resources Eileen Dowd-Stukel SD South Dakota Natural Heritage Program Roger Applegate TN Tennessee Wildlife Resources Agency Josh Campbell TN Tennessee Wildlife Resources Agency Charles Pekins TX Fort Hood Natural Resources Management Branch Robert C. Dowler TX Angelo State University Michael L. Fies VA Virginia Department of Game and Inland Fisheries W. Mark Ford VA United States Geological Service Emily Thorne VA Virginia Tech University Curtis Twellmann WI Wisconsin Department of Natural Resources Richard E. Rogers WV West Virginia Division of Natural Resources 22
2021 Southeastern Naturalist Vol. 20, Special Issue 11 R.W. Perry, D.B. Sasse, J.C. Perkins, and N.W. Sharp Appendix 2. Additional publications not cited in the literature cited that also presented county data used to create distribution maps of Eastern Spotted Skunk locations since the year 2000. Eng, R.Y.Y, and D.S. Jachowski. 2019. Summer rest-site selection by Appalachian eastern spotted skunks. Journal of Mammalogy. 100:1295–1304 Hackett, H.M. 2008. Occupancy modeling of forest carnivores in Missouri. Dissertation, University of Missouri-Columbia, Columbia, MO. Hawkins, N.R. 2012. Mesopredators in the blackland prairie of Texas: Occupancy, detec- tion probability, and diversity in relation to landcover change. M.Sc. Thesis. Stephen F. Austin State University, Nacogdoches, TX. 228 pp. Hardy, L.M. 2013. Eastern Spotted Skunk (Spilogale putorius) at the Ouachita Mountains Biological Station, Polk County, Arkansas. Journal of the Arkansas Academy of Science 67:59−65. Harris, S.N., T.J. Doonan, E.L. Hewett Rangheb, and D.S. Jachowski. 2019. Den-site selec- tion by the Florida Spotted Skunk. Journal of Wildlife Management 84:127−137. Lombardi, J.V., M.T. Mengak, and S.B. Castleberry. 2016. New records of the Eastern Spot- ted Skunk in northwestern Virginia. Banisteria 46:25−27. Sprayberry, T.R., and A.J Edelman. 2018. Den-site selection of Eastern Spotted Skunks in the southern Appalachian Mountains. Journal of Mammalogy 99:242−251. Stovall, E.A., and S.E. Hayslette. 2013. Activity patterns of Allegheny Woodrats in Tennes- see. Southeastern Naturalist 12:748−756. 23
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