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unpublished notes of mammal species richness in the north-central Piedmont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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