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Mammals of the North-central Piedmont of South Carolina
Steven E. Fields

Southeastern Naturalist, Volume 6, Number 4 (2007): 577–596

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2007 SOUTHEASTERN NATURALIST 6(4):577–596 Mammals of the North-central Piedmont of South Carolina Steven E. Fields* Abstract - There is a paucity of data on mammal species richness in the Piedmont physiographic province of the Carolinas, especially in north-central South Carolina. Over the last 12 years, I conducted field surveys, searched the literature, and queried numerous museums and agencies to locate records of mammals from the Piedmont province of South Carolina. I recorded 43 species of mammals, four of which are listed as species of state concern, from six counties in north-central South Carolina. Baseline information on mammal species occurrence is critical to future studies and issues of management and conservation. Introduction Relatively little information has been published on the mammals of South Carolina or the Piedmont physiographic province. Previous work on the mammalian fauna of South Carolina (Coleman 1939, 1940, 1941, 1948, 1949) was limited and dealt with areas outside the Piedmont. Golley (1966) reported mammal species occurrence by county throughout South Carolina, but his report included only collections in the Charleston Museum. Mengak et al. (1987) and Mengak and Guynn (2003) investigated small mammals in the western Piedmont and mountains of South Carolina. Brown (1997) and Webster et al. (1985) presented regional range maps, but not specific locations for mammals in the southeastern United States. In South Carolina, two museums house the majority of the state’s mammal collections. The Clemson University Vertebrate Collections contain specimens from the entire state, but the mountain region, and to a lesser degree the southwestern Piedmont, are best represented. Similarly, the Charleston Museum maintains specimens from across South Carolina, but the majority were collected from the Coastal Plain. Other museums in the United States house specimens from South Carolina, but relatively few of those were collected from the South Carolina Piedmont. According to Stein, (2002) South Carolina ranked 15th in the United States for overall mammal richness at 96 species, with 13.5% of those listed at some level of risk. It is important, then, to establish a baseline of information regarding mammal species distribution in the north-central South Carolina Piedmont. My objective was to synthesize as much information as possible from field surveys, museum records, scientific publications, and unpublished notes of mammal species richness in the north-central Piedmont of South Carolina. *Culture and Heritage Museums of York County, 4621 Mount Gallant Road, Rock Hill, SC 29732, and Department of Biological Sciences, University of South Carolina, Columbia, SC 29208; stevefields@chmuseums.org. 578 Southeastern Naturalist Vol. 6, No. 4 Study Area Godfrey (1980) described the Piedmont in the eastern United States between the Appalachian Mountains and the Atlantic Coastal Plain, extending from New York to Alabama. The area is characterized by a landscape of partially eroded rolling hills, most of which is currently in some stage of postagricultural old-field succession. As a physiographic unit, the Piedmont is well-defined. The Appalachian Mountains rise to the west with the Blue Ridge escarpment forming a clear topographic boundary with the Piedmont in western South Carolina. The eastern margin of the Piedmont is also clearly marked where rivers suddenly descend the “Fall Zone” to the Sandhills of the Upper Atlantic Coastal Plain (Murphy 1995). The oak-hickory-pine forests typical of the southeastern United States cover much of the Piedmont in the Carolinas. The topography is moderately sloped and well drained by broad streams and narrow floodplains (Skeen et al. 1993). The climate is classified as humid subtropical with annual precipitation of 80–100 cm and average temperatures of 23 ºC in summer and 10 ºC in winter (Martin et al. 1993a). While various seres of classic old-field succession still exist, much of the Piedmont has been developed as residential and commercial property, representing a loss of natural habitats. Development is particularly heavy in York County and the adjacent metropolitan region that includes Charlotte, NC. In Figure 1. Counties of the north-central piedmont in South Carolina. The Piedmont physographic province is delineated by the gray lines. 2007 S.E. Fields 579 addition to York County, five other north-central Piedmont counties (Spartanburg, Union, Cherokee, Chester, and Lancaster) of South Carolina are included in this report (Fig. 1). Methods From 1994 to 2005, I conducted extensive field surveys and synthesized data from literature accounts, museum and university collections, and unpublished records and reports from individuals, state agencies, and national parks. Copies of notes, surveys, and records from other agencies, as well as voucher images of some specimens are on file at the Museum of York County in Rock Hill, SC. Representative specimens were prepared as vouchers and deposited into the mammal collection at the Museum of York County. I conducted field surveys at numerous sites in York, Cherokee, and Chester counties from January 1994 to July 1995, January through October 1996, May 1997, September 2002, and March 2004 to October 2005 (Fig. 1). I used a variety of survey methods (Wilson et al. 1996), including Sherman live traps, pitfall traps (4-liter to 19-liter), discarded bottle surveys (Benedict and Billeter 2004, Pagels and French 1987), carnivore scent-station surveys (SCDNR 2003a), remote-trip cameras (Wemmer et al. 1996), bat mist nets, and harp traps (Kunz and Kurta 1988) to assess mammal species occurrence. Survey effort varied among sites. In the species accounts, the following acronyms apply: AMNH = American Museum of Natural History, CNB = Cowpens National Battlefield, CM = Carnegie Museum, ChM = Charleston Museum, CUSC = Clemson University Vertebrate Collections, SCDNR = South Carolina Department of Natural Resources, FMNH = Field Museum of Natural History, KMP = Kings Mountain National Military Park, MYCO = Museum of York County, RM = notes from Rudy Mancke. In each account “specimens examined” were actually inspected by me, while “specimens reported” were not—the data were provided by another museum. All information under “survey records” comes from my own field work. Most locality data from survey records and specimens examined were reported as mileage and direction from Rock Hill (e.g., 2.5 mi SW Rock Hill). In each account “Rock Hill” specifically refers to downtown, with mileage and direction from City Hall at geographic coordinates 34º55.4N, 81º1.7W. Taxonomy and phylogenetic sequence follow Baker et al. (2003). Species Accounts Order Didelphimorphia Family Didelphidae Didelphis virginiana Kerr (Virginia opossum). This species is found throughout the north-central Piedmont of South Carolina. I have documented numerous occurrences of this species in York County by road-kill speci580 Southeastern Naturalist Vol. 6, No. 4 mens, and tracks. Published records for Chester, Lancaster, and York counties include Cloninger et al. (1977) and Golley (1966). Additionally, there are unpublished records from Cherokee (CNB, SCDNR) Chester, Lancaster, Union (SCDNR), and York (SCDNR, KMP) counties. Specimens examined – York County: 2 mi N Rock Hill (3 MYCO). Order Insectivora Family Soricidae Sorex longirostris Bachman (southeastern shrew). There are relatively few records of this shrew in the north-central Piedmont of South Carolina. I have collected two specimens in Chester County. Cloninger et al. (1977) reported this species in York and Lancaster counties. Specimens examined – Spartanburg County: near Bird Mountain (1 CUSC). Blarina carolinensis Bachman (southern short-tailed shrew). Two species of short-tailed shrew in the genus Blarina (B. brevicauda (Say) [northern short-tailed shrew] and B. carolinensis) are known from the Carolinas (Webster et al. 1985, Brown 1997). While these two species appear to be allopatric (Whitaker and Hamilton 1998) or partially sympatric (Brown 1997) throughout most of the region, there are some areas of syptopy in the Coastal Plain of eastern North Carolina (D. Webster, University of North Carolina, Wilmington, NC, pers. comm.). Where their ranges overlap, competition between the two species may be reduced by habitat partitioning (McCay 2001). Only B. carolinensis is known to occur within the six counties surveyed (D. Webster, pers. comm.). Golley (1966) reported B. brevicauda from Spartanburg County, but taxonomic revisions by Genoways and Choate (1972) prompted reassignment of these specimens to B. carolinensis. Cloninger et al. (1977) also identified short-tailed shrews captured in York County as B. brevicauda, and it is likely that they, too, were subscribing to the earlier taxonomy. Published records of B. carolinensis include York and Lancaster counties (Cloninger et al. 1977) and Spartanburg County (French 1981, Golley 1966). An unpublished record for York/Cherokee counties was available from KMP (the park overlaps the two counties). Survey records – Chester County: 12.1 mi SE Rock Hill: Landsford Canal State Park. York County: Rock Hill at Winthrop Coliseum; Rock Hill on Hilldale Drive; 6.7 mi N Rock Hill: Lake Wylie Dam; 3.7 mi WSW Rock Hill: Bridgewater Road; 5 mi NNW Rock Hill. Specimens examined – York County: 2.7 mi E York (2 MYCO); 3.9 mi WSW Rock Hill (2 MYCO); Rock Hill, 0.8 mi NE downtown (1 MYCO); 3 mi NNW Rock Hill (1 MYCO); 13 mi NW Rock Hill (1 MYCO). Cryptotis parva Say (least shrew). The least shrew inhabits the central and southeastern portions of the United States, including all of North and South Carolina (Webster et al. 1985). I have collected C. parva from York and Cherokee counties. Additionally, Cloninger et al. (1977) published records from York and Lancaster counties. 2007 S.E. Fields 581 Specimens examined – York County: 13 mi NW York: Kings Mountain National Park (9 MYCO); 2.7mi E York: JCT SC Hwy 161 and Fishing Creek (1 CUSC). Family Talpidae Scalopus aquaticus Linnaeus (eastern mole). The eastern mole is fairly common and appears to be the only mole species in the region. Condylura cristata (L.) (star-nosed mole) is known from the mountains and Coastal Plain with a few possible Piedmont records in North Carolina (Beane 1995). The range of Parascalops breweri (Bachman) (hairy-tailed mole) in the Carolinas is confined to the mountains (Webster et al. 1985). Literature records for S. aquaticus from York County include Golley (1966) and Penney (1950). There are unpublished records from York/Cherokee counties (KMP) and Cherokee County (CNB). Survey records – York County: 1.6 mi S Rock Hill; 10.4 mi NW York: Kings Mountain National Military Park; 4.3 mi S Rock Hill: Clearlake Plantation; 6.7 mi N Rock Hill: Lake Wylie Dam; 4.1 mi SW Rock Hill; JCT Hwy 5 and Catawba River. Chester County: 13.5 mi SE Rock Hill: Landsford Canal State Park. Specimens reported – York County: Catawba (1 FMNH). Specimens examined – York County: (1 MYCO); 13 mi NW York (1 MYCO). Order Chiroptera Family Vespertilionidae Myotis austroriparius Rhoads (southeastern myotis). The rarity of this species in South Carolina is reflected by the scant records that were located. Menzel et al. (2003) reported a single capture record from a mine in Cherokee County. An unpublished record from KMP also comes from Cherokee County. No museum specimens were located from the northcentral Piedmont of South Carolina. Lasiurus borealis Müller (red bat). This common bat species has been well documented throughout South Carolina, including all counties of the north-central Piedmont (Menzel et al. 2003). Additional published records include Spartanburg County (Golley 1966) and York County (Neuhauser and DiSalvo 1972). There are also unpublished records (RM) from York, Chester, and Lancaster counties. Specimens reported – York County: Catawba (1 FMNH). Specimens examined – Chester County: 13.5 mi SE Rock Hill: Landsford Canal State Park (1 MYCO). Lasiurus cinereus (Beauvois) (hoary bat). This large bat is widespread throughout South Carolina, but has only been recorded in Lancaster, Spartanburg, and York counties in the north-central Piedmont (Menzel et al. 2003). I also located unpublished records from York and Lancaster counties (RM). No museum specimens were found. 582 Southeastern Naturalist Vol. 6, No. 4 Lasiurus seminolus (Rhoads) (Seminole bat). The Seminole bat is fairly common throughout South Carolina, but it has been reported from only Lancaster and Spartanburg counties in the north-central Piedmont of the state (Menzel et al. 2003). An unpublished record also exists for Lancaster County (RM), but no museum specimens were found. Lasionycteris noctivagans (LeConte) (silver-haired bat). This bat occurs statewide and has been reported in Chester, Lancaster, Spartanburg, and York counties (Menzel et al. 2003). There is another published record (Neuhauser and DiSalvo 1972) for Chester County and one unpublished record (RM) for York County. Specimens reported – Lancaster County: (1 ChM). York County: Rock Hill (1 ChM). Pipistrellus subflavus (F. Cuvier) (eastern pipistrelle). This small, common species has been reported from Cherokee, Chester, Spartanburg, and York counties (Menzel et al. 2003). Additional unpublished records (SCDNR, RM) support the presence of this species in York County. Survey records – York County: Rock Hill: South Cherry Road. Specimens reported – Chester County (1 ChM). Eptesicus fuscus (Beauvois) (big brown bat). The big brown bat occurs in all six counties of the north-central Piedmont of South Carolina (Menzel et al. 2003). Unpublished records (RM) exist for Chester, Union, and York counties. Survey records – Chester County: JCT I-77 and Hwy 497 near Richburg. York County: 6.2 mi. NNW Rock Hill: Museum of York County Nature Trail; 6.5 mi. SSW Rock Hill: Strait Road; Rock Hill: Charlotte Avenue. Specimens examined – York County: 4.8 mi NW Rock Hill: Cross Creek Court (1 MYCO); York: West Madison Street (1 MYCO). Nycticeius humeralis (Rafinesque) (evening bat). This bat ranges throughout South Carolina with published records from Cherokee, Chester, Spartanburg, Union, and York counties (Menzel et al. 2003). There is another published account (Neuhauser and DiSalvo 1972) from York County and unpublished records (RM) from Cherokee, Chester, Lancaster, and York counties. Specimens reported – Union County (1 ChM). Order Lagomorpha Family Leporidae Sylvilagus floridanus (Allen) (eastern cottontail). This is the only lagomorph known to inhabit the north-central Piedmont of South Carolina. At least one record of S. aquaticus (Bachman) (swamp rabbit) was reported from York County. However, the specimen is believed to be an introduced individual as hunters have been known to release swamp rabbits in the region (S. Miller, Clemson University Vertebrate Collections, Clemson, SC, pers. comm.). The skull of the specimen is housed at CUSC (#2338). 2007 S.E. Fields 583 The eastern cottontail likely inhabits all counties of the north-central Piedmont of South Carolina, but published records of S. floridanus include York and Lancaster counties (Cloninger et al. 1977). Unpublished records include York/Cherokee counties (KMP) and Chester and Union counties (SCDNR). Survey records – Chester County: 13.5 mi SE Rock Hill: Landsford Canal State Park (2). York County: 3.9 mi WSW Rock Hill: Covenant Place; 4.3 mi SE Rock Hill: Clearlake Plantation; 4.8 mi SW Rock Hill; 6.5 mi NNW Rock Hill: Mount Gallant Elementary School; 9 mi WSW Rock Hill: Historic Brattonsville; JCT SC Hwy 49 and McKinney Road; Rock Hill: JCT Dave Lyle Boulevard and US Hwy 21 Bypass; Rock Hill: JCT India Hook Road and SC Hwy 161; JCT I-77 and Porter Road; near JCT Mount Gallant Road and SC Hwy 161 (3); JCT Museum Road and Walnut Ridge; JCT SC Hwy 51 and US Hwy 21. Specimens examined – S. aquaticus: York County: JCT Clark’s Fork and Bullocks Creek (1 CUSC). S. floridanus: Spartanburg County: (2 CUSC); Union County: (1 CUSC); York County: 3.1 mi S Rock Hill (1 MYCO). Order Rodentia Family Sciuridae Tamias striatus (Linnaeus) (eastern chipmunk). The eastern chipmunk reaches the southeastern edge of its range in South Carolina in the northcentral Piedmont (Webster et al. 1985). The species is known from York and Lancaster counties (Cloninger et al. 1977) and Spartanburg County (Golley 1966). I have also observed this animal on several occasions within the city limits of Rock Hill. Specimens examined – York County: Rock Hill: Sedgewood Drive (1 MYCO). Marmota monax (Linnaeus) (woodchuck). Woodchucks or groundhogs have steadily increased their range in response to human landscape alteration (Webster et al. 1985). Indeed, the presence of woodchucks in northwestern York County slightly extends the range depicted in Webster et al. (1985) and Whitaker and Hamilton (1998). I collected two road-kill specimens within a single year near the same site–6.2 mi NNW Rock Hill near the Lake Wylie Dam. Both specimens are awaiting preparation as voucher specimens. I subsequently observed other individuals within two miles of the same site. Sciurus carolinensis Gmelin (eastern gray squirrel). The presence and abundance of the gray squirrel in South Carolina is easily validated. No doubt it occurs in all counties of the north-central Piedmont. Still, the only official published records are from Lancaster County (Cloninger et al. 1977, Golley 1966) and York County (Cloninger et al. 1977). Unpublished records include Cherokee/York counties (KMP), Lancaster County (SCDNR), and Union County (SCDNR). I have also observed and documented this species at numerous sites throughout York County. 584 Southeastern Naturalist Vol. 6, No. 4 Specimens examined – Spartanburg County: “mid-county” (1 CUSC). York County: 6.1 mi NNW Rock Hill (2 MYCO); no specific locality (1 MYCO). Sciurus niger Linnaeus (fox squirrel). The fox squirrel is concentrated in the Sandhills and Coastal Plain of the Carolinas with a restricted range in the Piedmont (Webster et al. 1985). I have a single record of this species in the north-central Piedmont of South Carolina. Specimen examined – Chester County: Bethlehem Road (1 MYCO). Glaucomys volans (Linnaeus) (southern flying squirrel). This nocturnal rodent is rarely encountered, but is common in the region (Webster et al. 1985). The only published record from the north-central Piedmont of South Carolina is Lancaster County (Golley 1966). One unpublished record is known from Cherokee/York counties (KMP). Survey records – York County: River Hills: Hamilton’s Ferry Road; Rock Hill: Winthrop University campus; Rock Hill: Meadowbrook Drive; Rock Hill: JCT Eden Terrace and Charlotte Avenue; Rock Hill: Quail Drive. Specimens examined – York County (1 MYCO). Family Castoridae Castor canadensis Kuhl (American beaver). The beaver was extirpated from South Carolina, but reintroduced in 1940 (Shipes and Rainey 1986). Populations have apparently recovered and numbers appear to be increasing in the north-central Piedmont. Still, no literature records were found for the region. However, there are unpublished records from York/Cherokee counties (KMP) and Lancaster, Union, and York counties (SCDNR). Survey records – York County: 9.5 mi SW Rock Hill: Historic Brattonsville; off US Hwy 21 North: Anne Springs Close Greenway. Specimens examined – York County: 6 mi NW Rock Hill: Big Dutchman’s Creek (1 MYCO). Family Muridae Oryzomys palustris (Harlan) (marsh rice rat). According to Webster et al. (1985), the marsh rice rat occurs throughout South Carolina, but the range depicted was generally south of the north-central Piedmont. This species was reported from Chester County (Durden et al. 1999) and Lancaster and York counties (Cloninger et al. 1977). I collected a specimen in York County in an upland habitat, in keeping with the notation by Webster et al. (1985) that this species is sometimes “found well removed from wetland habitats.” Specimens examined – York County: Rock Hill: Winthrop Coliseum, 400 m E of building (1 MYCO); off US Hwy 21: Anne Springs Close Greenway (1 MYCO). Reithrodontomys humulis (Audubon and Bachman) (eastern harvest mouse). This diminutive native mouse resembles the introduced Mus musculus (L.) (house mouse). However, the harvest mouse has grooved upper incisors (Whitaker and Hamilton 1998) and a more placid disposition (W. Rogers, Winthrop University, Rock Hill, SC, pers. comm.). Published 2007 S.E. Fields 585 records include York and Lancaster counties (Cloninger et al. 1977) and Spartanburg County (Golley 1966). Survey records – York County: 5.4 mi NNE Rock Hill: near Catawba River. Genus Peromyscus Four species of mice in the genus Peromyscus inhabit South Carolina, but only two species, P. leucopus and P. polionotus have ranges that include the north-central Piedmont of South Carolina (Lae'rm and Boone 1994, Webster et al. 1985). Identification of species can be difficult, but determination is possible using external measurements and skull characters in a discriminant function developed by Laerm and Boone (1994). I used the two-way discriminant analysis to evaluate existing identifications of mice in the collections at CUSC and MYCO. The results of the analysis indicated that specimens in both collections were misidentified. See the accounts of P. leucopus and P. polionotus for more information and the resulting reclassifi cations. Peromyscus gossypinus (LeConte) (cotton mouse). Four specimens of cotton mouse (the identification of which seems valid based on external measurements) at the Charleston Museum were collected in Lancaster County, SC by R.H. Coleman in 1948. These are likely the same specimens described in Golley (1966) and Hall (1981). Although most of Lancaster County is within the Piedmont of South Carolina, the extreme eastern portion of the county occupies the Sandhills region of the Upper Coastal Plain. Coleman’s field catalog stated that the four P. gossypinus specimens were collected “16.5 mi. E. of Lancaster on SC Hwy 9” in “moist mixed woods on Flat Creek.” Some of the locality information is apparently erroneous because SC Hwy 9 runs far north of the indicated position, while Flat Creek runs far to the south. Coleman’s position was likely closer to SC Hwy 903. In any event, 16.5 miles east of Lancaster is a location outside of the Piedmont and more appropriate to the habitat requirements of P. gossypinus. Peromyscus leucopus (Rafinesque) (white-footed mouse). The whitefooted mouse has been reported from all counties in the north-central Piedmont of South Carolina except Union County, where it also likely occurs. Published accounts include Lancaster and York Counties (Cloninger et al. 1977), Lancaster, Spartanburg, and York counties (Golley 1966). In addition, there are unpublished records from Cherokee County (CNB, KMP). Survey records – Chester County: 13.5 mi SE Rock Hill: Landsford Canal State Park; 0.8 mi S Rock Hill; 3.68 mi ESE Rock Hill: Covenant Place; Inwood Dr, off Museum Road; 5.2 mi N Rock Hill, Mount Gallant Road; 6.8 mi E Rock Hill, Catawba Reservation at Tom Stevens Road (2); 8.4 mi NE Rock Hill, Anne Springs Close Greenway; Rock Hill: Aiken Street; Rock Hill: Marion Street. Specimens reported – Lancaster County (4 ChM). York County: Catawba (3 FMNH); 5 mi N York (1 ChM); 10 mi N York (1 ChM). Specimens examined –York County: Rock Hill (1 CUSC); 5.2 mi N Rock Hill (1 CUSC); 0.6 mi NNW Rock Hill (1 MYCO); 13 mi NW York (3 MYCO); 586 Southeastern Naturalist Vol. 6, No. 4 3.7 mi ESE Rock Hill (1 MYCO); 3.9 S Rock Hill (1 MYCO). Peromyscus polionotus (old-field mouse). Webster et al. (1985) depicted a range for this species that included the western portion of the north-central Piedmont of South Carolina. Brown (1997) and Whitaker and Hamilton (1998) showed a range from Spartanburg County extending west. Published records of P. polionotus from Spartanburg County (Biggers and Dawson 1971, Golley 1966, Hall 1981, Schwartz 1954) support this range limitation. Laerm and Boone (1995) tested identifications of P. polionotus in Georgia and western South Carolina using discriminant analysis of mensural characters and agreed that specimens reported from Spartanburg County were correctly identified. I performed discriminant analysis on museum specimens (CUSC, MYCO) collected in York County identified as P. polionotus. In each case, discriminant values (P > 1) indicated the mice to be P. leucopus. I have reclassified specimens at MYCO and suggested that specimens from York County at CUSC also be reclassified as P. leucopus. Based on the analyses and published records, it is possible that P. polionotus does not occur in York County. Perhaps with the exception of Spartanburg County, P. polionotus is absent from all counties of the northcentral Piedmont of South Carolina. More work is needed to define the northern limits of this species’ range in South Carolina. Ochrotomys nuttalli (Harlan) (golden mouse). The golden mouse likely occurs throughout the north-central Piedmont of South Carolina, but published records only exist for Lancaster County (Cloninger et al. 1977), Spartanburg County (Golley 1966), and York County (Cloninger et al. 1977 and Golley 1966). Specimens reported – York County: 10 mi NE York (1 ChM). Specimens examined – York County: 12 mi NW York: Kings Mountain National Military Park (1 MYCO). Sigmodon hispidus Say and Ord (hispid cotton rat). The cotton rat was one of the most frequently captured rodents in my field surveys. There are published records of this common species from Chester (Durden et al. 1999), Lancaster and York counties (Cloninger et al. 1977, Golley 1966), as well as Spartanburg County (Golley 1966). In addition, there are unpublished records from York/Cherokee counties (KMP). Survey records – York County: Rock Hill: Barrow Court; Rock Hill: fields behind Winthrop Coliseum (5); 3.7 mi WSW Rock Hill: Bridgewater Road (3); 5.2 mi NE Rock Hill; 5.3 mi S Clover: Fairhope Road; 5.4 mi NNE Rock Hill; 6.2 mi NNW Rock Hill: Museum of York County (2); 8 mi SE Rock Hill: JCT SC Hwy 5 and Catawba River; 8.4 mi NE Rock Hill: Anne Springs Close Greenway; 9 mi WSW Rock Hill: Historic Brattonsville (3); 6.2 mi NNW Rock Hill: JCT Mount Gallant Road and Friar Road (3). Specimens reported – Lancaster County: Lancaster (1 ChM). Spartanburg County: 4 mi E Inman (1 AMNH). York County: 5 mi NE York (1 ChM). Specimens examined – York County: 3.9 mi SW Rock Hill (2 MYCO); 1.8 mi NW Rock Hill (1 MYCO). Neotoma floridana (Ord) (eastern woodrat). A single eastern woodrat 2007 S.E. Fields 587 record from the South Carolina Piedmont was reported by Dowda et al. (1981). The rat was killed after it bit a child “in an exceptionally wellgroomed suburban area in York County, South Carolina" (Dowda et al. 1981). The rat tested positive for rabies, and no voucher material was retained. It is likely the rat was misidentified, as no other occurrences of N. floridana have been documented in the six counties surveyed of the north-central Piedmont of South Carolina. Accordingly, this record should be considered spurious. The closest records are three specimens collected in Greenville County (Coleman 1949). Rattus norvegicus (Berkenhout) (Norway rat). This introduced species ranges across the United States and is apparently more abundant and widely distributed than R. rattus (L.) (black rat) (Whitaker and Hamilton 1998). Still, I found no published records of this species in the north-central Piedmont of South Carolina, and museum records were scarce. Specimens reported – Spartanburg County: 4 mi N Boiling Springs (1 ChM). Specimens examined – York County: 2.9 mi ESE York (3 MYCO). Rattus rattus (Linnaeus) (black rat). Like the Norway rat, this species was introduced from Europe (Webster et al. 1985). The range of the black rat depicted in Whitaker and Hamilton (1998) did not include the north-central Piedmont of South Carolina, and I found only one published record of this species in the region (York County: Golley 1966). This record most likely refers to the specimens from the Charleston Museum listed below. Specimens reported – York County: York (1 ChM); 5 mi NE York (1 ChM). Mus musculus Linnaeus (house mouse). The house mouse, a diminutive rodent, is another non-native species. There are published records for Lancaster, Spartanburg, and York counties (Cloninger et al. 1977, Golley 1966) and unpublished records from Cherokee County (CNB). Survey records – York County: Rock Hill: fields behind Winthrop Coliseum (3); 0.8 mi S Rock Hill; 6.2 mi NNW Rock Hill: Museum of York County; 8.4 mi NE Rock Hill: Anne Springs Close Greenway; 9 mi WSW Rock Hill: Historic Brattonsville (3). Specimens reported – York County: York (1 ChM). Specimens examined – York County: 1.7 mi NW Rock Hill (2 MYCO); 1.7 mi NNE Rock Hill (1 MYCO); 2.9 mi ESE York (1 MYCO); 7.2 mi WSW Rock Hill (1 MYCO); 8.7 mi SW Rock Hill (1 MYCO); 9.5 mi SW Rock Hill (1 MYCO). Microtus pennsylvanicus (Ord) (meadow vole). Occurrence of the meadow vole has not been well documented in the region. Cloninger et al. (1977) reported this species from Lancaster and York counties, and Golley (1966) reported M. pennsylvanicus from Spartanburg County. Survey records – York County: 3.9 mi Rock Hill; 7.3 mi NNW Rock Hill: Covenant Place. Specimens examined – York County: 3.9 mi WSW Rock Hill (1 MYCO); 4 588 Southeastern Naturalist Vol. 6, No. 4 mi SW Rock Hill (1 MYCO); 5.6 mi NNW Rock Hill (1 MYCO). Microtus pinetorum (LeConte) (pine or woodland vole). The woodland or pine vole has been reported in Lancaster County (Cloninger et al. 1977), Spartanburg County (Golley 1966), and York County (Cloninger et al. 1977, Golley 1966). Specimens reported – York County: York (1 ChM); Catawba (1 FMNH). Specimens examined – 2.1 mi NNW Rock Hill (1 MYCO); 6.2 mi NNW Rock Hill (1 MYCO). Ondatra zibethicus (Linnaeus) (muskrat). This common semi-aquatic rodent has been documented in all counties of South Carolina’s north-central Piedmont. There are published records for Chester, Lancaster, and York counties (Golley 1966) and unpublished records for Cherokee, Chester, Lancaster, Union, and York counties (SCDNR). Survey records – York County: Clover: Faulkner Street; Rock Hill: pond behind Winthrop Coliseum. Specimens examined – Spartanburg County: Greer (1 CUSC). York County: 2.7 mi NW Rock Hill (2 MYCO). Family Dipodidae Zapus hudsonius Zimmermann (meadow jumping mouse). I located a single published record of this species for Spartanburg County (Golley 1966), in keeping with the range of the species depicted in Webster et al. (1985). Brown (1997) and Whitaker and Hamilton (1998) indicated that range limits in South Carolina were farther east and south, including all counties of the north-central Piedmont. However, each of the sources agreed that the meadow jumping mouse was sporadic in its occurrence. There were no other survey records or museum specimens of this species in the region. Order Carnivora Family Canidae Canis latrans Say (coyote). Over the last several decades, coyotes have expanded their range into the eastern United States. Some have moved naturally as forests were cleared; others were introduced by humans (Brown 1997, Choate et al. 1994, Webster et al. 1985). There were no published records for the study area, but there are unpublished SCDNR records based on fur harvests and scent-station surveys from Chester, Union, and York counties. Survey records – Chester County: 3 mi S Historic Brattonsville; 8.8 mi SE Chester. York County: 7.8 mi NW Rock Hill: SC Hwy 161; 6 mi S Rock Hill: JCT Bechtler Road and I-77. Specimens reported – Union County: “vicinity of Rose Hill near Tyger River” (1 ChM). Specimens examined – Chester County: 8.5 mi ESE Chester, Hwy 44 (1 MYCO). York County: 2.8 mi NW Rock Hill (1 MYCO). Vulpes vulpes (Linnaeus) (red fox). Some populations of red foxes in the eastern United States were brought from Europe during colonial times 2007 S.E. Fields 589 for fox hunting (Webster et al. 1985, Whitaker and Hamilton 1998). The red fox is widely distributed with published records from Lancaster and York counties (Cloninger et al. 1977). There are other unpublished accounts from York/Cherokee counties (KMP) and SCDNR records based on fur harvests and scent-station surveys from Cherokee, Chester, Lancaster, Union, and York counties. Survey records – York County: Rock Hill: JCT India Hook Road and SC Hwy 161; 4.8 mi SW Rock Hill; Mount Gallant Road at Big Dutchman Creek bridge; JCT Sutton Road and SC Hwy 160. Specimens examined – York County: 4.3 mi N Rock Hill (1 MYCO); 7 mi NE Rock Hill (1 MYCO). Urocyon cinereoargenteus Schreber (gray fox). The gray fox is relatively common throughout the region as evidenced by published and unpublished records. Cloninger et al. (1977) reported the gray fox in Lancaster and York counties, and Golley (1966) listed Chester and Spartanburg counties. There were also numerous unpublished reports from York/ Cherokee counties (KMP) and SCDNR records based on fur harvests and scent-station surveys from Cherokee, Chester, Lancaster, Union, and York counties. Survey records – York County: York: College Street; 3.9 mi N Rock Hill: Brookridge Drive; 9 mi WSW Rock Hill: Historic Brattonsville; JCT Mount Gallant Road and McClain Road; Museum Road ca. 2 mi S Museum of York County; JCT US Hwy 72 (Bypass) and SC Hwy 901; US Hwy 21 at Catawba River Bridge. Specimens examined – Union County (1 CUSC). York County: 20 mi NW Rock Hill near Clover (1 MYCO); 3.9 mi N Rock Hill (1 MYCO); 4.5 mi NW Rock Hill (1 MYCO); 4.8 mi NE Rock Hill (1 MYCO). Family Ursidae Ursus americanus Pallas (American black bear). Golley (1966) reported a black bear from Spartanburg County. There is one other record from the north-central Piedmont of South Carolina, but it is not supported by a voucher specimen or image. Nevertheless, the report was generated by an SCDNR official who reported scat and claw marks on a tree just off Chapel Road in southern York County, April 1995. The type of bear sign noted is quite distinctive (Elbroch 2003), and Ursus americanus is the only bear native to the Carolinas (Webster et al. 1985). Still, if this record is valid, then it almost certainly represents a transient individual that was outside of its normal mountain or Coastal Plain range. No populations of bears would be expected in the region (D. Webster, pers. comm.). Family Procyonidae Procyon lotor (Linnaeus) (raccoon). This common carnivore has been documented in every county within the region. Cloninger et al. (1977) listed raccoons from Lancaster and York counties, and Golley (1966) re590 Southeastern Naturalist Vol. 6, No. 4 ported specimens from Chester, Spartanburg, and York counties. There are unpublished records from Cherokee (CNB, SCDNR), Chester, Lancaster, Union, York (SCDNR), and Cherokee/York (KMP) counties. I have also documented the species at numerous sites in York and Chester counties during my field work. Specimens examined – York County: 4.3 mi N Rock Hill (1 MYCO). Family Mustelidae Mustela frenata Lichtenstein (long-tailed weasel). Although Whitaker and Hamilton (1998) indicated that M. frenata was one of the most widespread carnivores in the eastern United States, it is rarely encountered, and there are few records from the north-central Piedmont of South Carolina. Hall (1981) reported M. frenata from York County. I have encountered this animal only once. The survey record listed below is the result of an unusual circumstance. The weasel entered a residence and the owner locked the animal in her bathroom, whereupon she called me to come remove the “furry snake” from her house. Survey records – York County: 7.7 mi NNW Rock Hill: Allison Circle. Specimens reported – York County: 5 mi E York (1 ChM). Mustela vison Schreber (mink). The only literature accounts of mink in the north-central Piedmont of South Carolina are from Chester, Spartanburg, Union, and York counties (Golley 1966). Unpublished records include Cherokee and York counties (SCDNR, KMP) and Chester, and Union counties (SCDNR). Survey records – York County: 16 mi NNW Rock Hill: SC Hwy 274; 3.9 mi SSE Rock Hill; JCT SC Hwy 161 and Ebinport Road. Specimens examined – 3.1 mi NW Rock Hill (1 MYCO); 3.9 SSE Rock Hill (1 MYCO). Lontra canadensis (Schreber) (river otter). The river otter was reported only from Chester County by Golley (1966). However, unpublished SCDNR records based on fur harvests exist for Cherokee, Chester, Lancaster, Union, and York counties. Survey records – Cherokee County: JCT SC Hwy 5 and SC Hwy 55. Specimens examined – Cherokee County: 6.5 mi SE Blacksburg (1 MYCO). Family Mephitidae Mephitis mephitis (Schreber) (striped skunk). While I have encountered numerous of road-kill specimens throughout York County, I have only logged a single official survey record (see below). Cloninger et al. (1977) found the striped skunk in Lancaster and York counties. In addition, there are unpublished accounts from Cherokee, Chester, Union, and York counties (SCDNR). There are no museum specimens collected from the north-central Piedmont of South Carolina. This species should not be confused with Spilogale putorius (L.) (spotted skunk) that inhabits the western portion of the Carolinas (Webster et al. 1985, Whitaker and Hamilton 1998). A single record is known from Spartanburg County, SC (Golley 1966), but this record 2007 S.E. Fields 591 is unsubstantiated by a voucher specimen and highly suspect (D. Webster, pers. comm.). Survey records – York County: JCT SC Hwy 5 and Billy Wilson Road. Family Felidae Lynx rufus Schreber (bobcat). There were no published records of the bobcat in the counties of the north-central Piedmont of South Carolina. However, unpublished SCDNR records based on fur harvests exist for Cherokee, Chester, Lancaster, and York counties. Specimens examined – Chester County: 16 mi S Rock Hill (1 MYCO); 7 mi SSE Rock Hill (1 MYCO); 8 mi E York: Newport (1 MYCO). Order Artiodactyla Family Cervidae Odocoileus virginianus (Zimmermann) (white-tailed deer). While this species is no doubt common throughout the region, the official records remain relatively few. I have documented white-tailed deer on numerous occasions throughout Chester and York counties, and I will not list each individual record here. Other unpublished reports include Cherokee, Union, and York counties (SCDNR, KMP). Published accounts (Cloninger et al. 1977, Golley 1966) reported this species in Chester, Lancaster, Union, and York counties. Specimens examined – York County: 6.3 mi N Rock Hill (1 MYCO). Discussion Forty-three species of mammals inhabit the six counties of the northcentral Piedmont of South Carolina, at least two of which, the woodchuck and marsh rice rat, apparently extend the ranges shown in Webster et al. (1985). Two other species (the eastern woodrat and American black bear) were reported outside their normal ranges, but neither report is supported by voucher material. Because these were single, isolated reports, it is unlikely that these species are part of the typical mammal fauna of the region. While the synthesis of published and unpublished records, museum specimens, and field accounts of relatively common mammals may seem of little value, the accounts listed herein represent the baseline of such data for the north-central Piedmont of South Carolina. Compared to other regions of South Carolina, the data on mammal species occurrence for the north-central piedmont are relatively few. Of the 2345 South Carolina mammal records (excluding my own field survey records) I have compiled, only 371 (15.8%) are from the six counties of the north-central Piedmont. Clearly, any mammal species occurrence data collected from the region need to be reported. The Piedmont as a zone of transition None of the mammal species in the Piedmont of South Carolina appear to be 592 Southeastern Naturalist Vol. 6, No. 4 endemic to the province. This is not surprising, as floral and faunal distributions in the Piedmont have affinities with other provinces within the state. The Piedmont appears to be a zone of transition between the Appalachian oak forests (Stephenson et al. 1993) and the mixed hardwood-longleaf pine associations (Martin et al. 1993b) of the Coastal Plain. In fact, the Carolina Piedmont contains oak-hickory-pine forests (Skeen et al. 1993) with Pinus taeda L. (loblolly pine) and P. echinata P. Mill. (shortleaf pine) sharing the canopy with Quercus spp. (oaks) and Carya spp. (hickories). Mammal distributions in South Carolina follow a similar transitional pattern. Four species of shrews (Sorex spp.), five bats (Myotis spp.), one rabbit (Sylvilagus aquaticus), one squirrel (Tamiasciurus hudsonicus (Erxleben) [red squirrel]), one mouse (Peromyscus maniculatus (Wagner) [deer mouse]), and one weasel (Mustela nivalis L. [least weasel]) are restricted to the mountain province of South Carolina (northern Oconee County and northwestern Pickens County), whereas two bats (Lasiurus intermedius H. Allen [northern yellow bat] and Tadarida brasiliensis (I. Geoffroy) [Brazilian free-tailed bat]) and one rabbit (Sylvilagus palustris (Bachman) [marsh rabbit]) are restricted to the Coastal Plain. Other mammal species in South Carolina have ranges that include the Piedmont and one or both of the other physiographic provinces in the state. Surveys of birds in the north-central piedmont of South Carolina have shown similar trends (W. Rogers, Winthrop University, Rock Hill, SC, pers. comm.). Conservation The conservation of mammals demands data on the occurrence and relative abundance of species so that status can be determined and ranks can be assigned. The South Carolina Department of Natural Resources recently updated the listed mammal species in the state (SCDNR 2003b). At least four mammal species (southeastern myotis, hoary bat, fox squirrel, and meadow vole) in the north-central Piedmont of South Carolina officially merit some level of state concern. Menzel et al. (2003) discussed the issues of bat conservation in South Carolina, including the possible impacts of pesticides and loss of habitat and roost structures. Eight of the 15 (53%) species of bats in South Carolina are listed as state species of concern; two of those species are found in the north-central Piedmont. The southeastern myotis is listed as “rare in parts of its range” and ranks as a state endangered species. The hoary bat is listed as “demonstrably globally secure,” but it is considered a species of concern because its status in the state is unknown. While rodents represent the largest order and one of the most diverse groups of mammals (Nowak and Paradiso 1983, Wilson and Ruff 1999), certain species have limited abundance and distribution on local levels. The fox squirrel is listed as a species of state concern in South Carolina. The status of the meadow vole is unknown in South Carolina and is, therefore, of state concern. 2007 S.E. Fields 593 Although the eastern woodrat is represented by a single questionable record in the study area, I note here that it is recognized as a species of concern in the state (SCDNR 2003b). Similarly, the black bear, although represented by a single transient individual in the north-central piedmont, is also listed as a species of concern in South Carolina and is further designated as questionably “uncommon or rare” in the state (SCDNR 2003b). Several mammalian species that once inhabited the Carolina Piedmont have been extirpated. However, the natural history of these species is beyond the scope of this paper. Briefly, they are the Canis lupus rufus Audobon and Bachman (red wolf), Canis lupus L. (gray wolf), Cervus elaphus L. (elk), and Bison bison (L.) (bison) (Brown 1997, Webster et al. 1985, Whitaker and Hamilton 1998). Because members of the public still report sightings of “panthers” in the Carolinas, a slight elaboration on this species is in order. Puma concolor (L.) (cougar or mountain lion) was once distributed across the southeastern United States, and while there are sporadic reports of sightings in remote parts of the Carolinas, it is generally agreed that the cougar has been extirpated from the region since the late 1800s (Brown 1997, Webster et al. 1985). The remaining population in the eastern United States is in southwestern Florida (Whitaker and Hamilton 1998, Wilson and Ruff 1999). I found no museum records of these extirpated species from South Carolina. Acknowledgments I thank the museum curators and collection managers for access to collections and data. I also thank Cowpens National Battlefield, Kings Mountain National Military Park, R. Mancke, and the South Carolina Department of Natural Resources for providing data. Additionally, I thank the many residents and businesses that allowed access to their property for field studies, and the following individuals who helped in the field, laboratory, and office with a variety of tasks: J. Chism, J. Holsinger, J. Langley, A. Lazenby, H. Matthews, S. McAninch, J. Moretz, C. McNeilly, S. Olcott, M. Perrot, W. Rogers, A. Sorrow, J. Sorrow, J. Wray, S. Young, and G. Zakrzewski. I owe a special note of thanks to D. Webster for many helpful comments on drafts of this manuscript. Special thanks go to K. Francl and two anonymous reviewers. Their comments improved the overall quality of the manuscript. 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