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Amphibians of the Central and Southwestern Piedmont Province of South Carolina
Steven E. Fields

Southeastern Naturalist, Volume 18, Issue 2 (2019): 202–223

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Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 202 2019 SOUTHEASTERN NATURALIST 18(2):202–223 Amphibians of the Central and Southwestern Piedmont Province of South Carolina Steven E. Fields* Abstract - There is a very limited amount of data on amphibian species richness in the Piedmont physiographic province of South Carolina. Over the last 14 y, I conducted field surveys, searched the literature, and queried numerous museums and agencies to locate records of amphibians from counties that lie completely within the boundaries of the Piedmont of South Carolina. I recorded 42 species from 12 Piedmont counties in South Carolina, 7 of which are listed as species of state concern. Such baseline amphibian data is crucial to continued studies and matters of conservation. Introduction Among vertebrates, reptiles and amphibians are typically regarded as less valuable components of biological communities (Todd et al. 2007). Witczak and Dorcas (2009) explored this notion from a novel viewpoint and ascribed economic value to reptiles and amphibians. Many biologists appreciate the inherent value of all species and their respective places within ecosystems (Blaustein et al. 1994). Over the decades, there has been a decline in amphibian diversity and abundance (Houlahan et al. 2000, Stuart et al. 2004), highlighting the importance of these organisms as indicators of environmental health (Blaustein and Wake 1995; see Blaustein et al. 1994 for a list of studies that monitored amphibian population stability). Although the above cited sources report several factors affecting amphibian declines, one such factor is the fungal disease chytridiomycosis (Voyles et al 2011). The disease is the result of Batrachochytrium dendrobatidis Longcore, Pessier, & D.K. Nichols (chytrid fungus), which has been responsible globally for amphibian population declines and extinctions (Skerratt et al. 2007). Landsford Canal State Park in Chester County is 1 site in the South Carolina Piedmont where the presence of chytrid fungus has been confirmed (Hoekstra 2013). The primary literature on amphibians in the South Carolina Piedmont is sparse. I noted a similar paucity of data for mammals (Fields 2007). Some secondary references (Beane et al. 2010, Dorcas and Gibbons 2008, Mitchell and Gibbons 2010, Powell et al. 2016) presented regional range maps but not specific locations for amphibians in the southeastern US. Still, such references are valuable for ascertaining the “expected” herpetofauna of the region. Even since the early European explorers, there has been a trend to investigate the Blue Ridge Mountain province and the Atlantic Coastal Plain, which overshadowed *Culture and Heritage Museums, Museum of York County, 4621 Mount Gallant Road, Rock Hill, SC 29732, and Department of Biology, Winthrop University, Rock Hill, SC 29733; stevefields@chmuseums.org. Manuscript Editor: Cathryn Greenberg Southeastern Naturalist 203 S.E. Fields 2019 Vol. 18, No. 2 Piedmont studies (Taylor 1998). My recent surveys of museum collections indicate that the trend has continued over the decades. Within the state, the primary comprehensive collection of amphibians (and other vertebrates) from South Carolina resides at Clemson University. However, most of those specimens were collected from the mountain region and western Piedmont. The Charleston Museum formerly maintained a collection of fluid-preserved amphibians and reptiles, with the majority of specimens taken from the Coastal Plain. Most of that collection was moved to the North Carolina Museum of Natural Sciences in 2014 (NC Museum of Natural Sciences 2018). Other museums in the US house specimens from South Carolina, but relatively few of those were collected from the Piedmont. According to Stein (2002), South Carolina,ranked 7th in the US for overall amphibian diversity (66 species), with 13.6% of the reported species listed at some level of risk. Dorcas and Gibbons (2008) and Mitchell and Gibbons (2010), however, reported 69 species of amphibians in South Carolina. It is important, then, to establish a baseline of information regarding amphibian species distribution. My objective was to synthesize as much information as possible from field surveys, museum records, scientific publications, and unpublished reports of amphibian species richness in the Piedmont of South Carolina. Study Area The Piedmont province of South Carolina covers >28,000 km2 and comprises about 35% of the total area of the state. Figure 1 shows the counties of South Carolina and the boundaries of the Piedmont region. I included in this report only counties that are entirely within the Piedmont province (Abbeville, Anderson, Cherokee, Chester, Fairfield, Greenwood, Laurens, McCormick, Newberry, Spartanburg, Union, and York). My primary reference for Piedmont boundaries was Griffith et al. (2002). I include here Fields’ (2007:578) description of the Piedmont to characterize the province: “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 post-agricultural 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 US 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 Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 204 cm and average temperatures of 23 °C in summer and 10 °C in winter (Martin et al. 1993). 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, North Carolina.” Methods I conducted field surveys at numerous sites in Cherokee, Chester, and York counties during 1994–1997, 2002–2005, and 2010–2017. I used a variety of survey methods (Heyer et al. 1994), including visual encounter surveys (VES) (Crump and Scott 1994), auditory observations, coverboards (Pittman and Dorcas 2006), and pitfall traps (Corn 1994). I prepared representative specimens or recorded images as vouchers, and deposited them in the herpetological collection at the Museum of York County (MYCO) in Rock Hill, SC. Figure 1. Counties of South Carolina and the Piedmont. The Piedmont physiographic province is within the bold gray lines. The boundaries are approximate but based on Griffith et al. (2002). Counties included in this study are shaded in gray. Base map provided by Texas Tech University Libraries, Lubbock, TX. Southeastern Naturalist 205 S.E. Fields 2019 Vol. 18, No. 2 I 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, records, and unpublished reports are on file at the Museum of York County. I also searched the VertNet database (VertNet 2018) for museum records of amphibians for all counties within the study area. I included only museum specimens that were identified to the species level. Within the last 20 y, several taxonomic revisions in herpetology have been proposed. Beane et al. (2010) used older names but included updated names published in the Society for the Study of Amphibians and Reptiles (SSAR) checklist of North American species (Crother 2008). In this paper, I present currently accepted common names and scientific names used in the updated SSAR checklist (Crother 2017) with notes, as appropriate, regarding previously used nomenclature. In the species accounts, the following institutional acronyms apply: ANSP = Academy of Natural Sciences at Philadelphia, AMNH = American Museum of Natural History, CM = Carnegie Museum, CUMV = Cornell University Museum of Vertebrates, CUSC = Clemson University Vertebrate Collections, FM = Field Museum of Natural History, FUR = Furman University, LACM = Los Angeles County Museum of Natural History, MPM = Milwaukee Public Museum, MYCO = Museum of York County, SC, TNHC = Texas Natural History Collections (The University of Texas at Austin), UF = Florida Museum of Natural History, UMMZ = University of Michigan Museum of Zoology, USNM = Smithsonian National Museum of Natural History, UTA = University of Texas at Arlington Amphibian and Reptile Diversity Research Center, and YPM = Yale Peabody Museum. Other acronyms include: CCARI = raw data from the Central Carolina Amphibian and Reptile Initiative (see Pittman and Dorcas 2006), SEF = my field observations and records, WU-AR = field data by Winthrop University graduate student Angel Rudert, and WU-NH = field data by Winthrop University graduate student Nicole Hoekstra. In each account, the notations “images examined” or “specimens examined” signify collections that I actually inspected, while “specimens reported” indicates those that I did not. I have not included individual specimen numbers. Results I found 685 county occurrence records representing 42 species of amphibians from the counties completely within the Piedmont physiographic province of South Carolina. Tables 1 and 2 summarize species occurrences by county and the nature of the specimen record. The species accounts that follow detail the county-occurrence records and include notes regarding taxonomy and nomenclature. Order Anura: Frogs Acris Duméril and Bibron, 1841 (cricket frogs) Acris crepitans Baird, 1854 (Eastern Cricket Frog). This species is found throughout the Piedmont counties of South Carolina. Abbeville: Metts and Nestor (2005); Anderson: specimens examined–CUSC; specimens reported–NCSM. Cherokee: CCARI; Dorcas (2009a, b, c), Kern et al. Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 206 Table 1. Records of frogs and toads from counties in the South Carolina Piedmont. Abbreviations for counties are ABB (Abbeville), AND (Anderson), CHK (Cherokee), CHS (Chester), FAI (Fairfield), GRW (Greenwood), LAU (Laurens), MCC (McCormick), NEW (Newberry), SPA (Spartanburg), UNI (Union), and YRK (York). Abbreviations for record types are: S = field surveys, observations, and raw data; U = unpublished reports and theses; L = published literature citations; I = image vouchers; M = museum specimens. County Species ABB AND CHK CHS FAI GRW LAU MCC NEW SPA UNI YRK Order Anura Acris crepitans L M L, S, U I, L, M, S, U L M, U L, M L L, S, U L, S I, L, M, S, U Acris gryllus M L M Anaxyrus americanus L,M L, M, U L, M L L L L L, S, U L,S I, L, M, U Anaxyrus fowleri M M L, M, U I, L, S, U L, M M M L, M L L, M, S, U L, M, S I, L, M, S, U Anaxyrus terrestris M M Gastrophryne carolinensis M M L I, L, S, U L U L L L, S M, S, U Hyla andersonii M Hyla chrysoscelis M L, U, S I, L, S, U L L L, S, U L I, L, S Hyla cinerea M M L I, L, S, U L M, U M L L, S L I, L, M, S, U Hyla gratiosa M M Hyla squirella I, L M Lithobates catesbianus M M L, M, U L, S, U L L, M L, M L, M, S, U L, S I, L, M, S, U Lithobates clamitans L, M M L, M, U I, L, S, U L M, U L L L L, M, S,U L, S L, M, S, U Lithobates palustris L, M, U L L L, S, U L L, U Lithobates sphenocephalus L M L, U, S I, L, M, S, U L M, U L L L L, S, U L, S L, M, S, U Lithobates sylvaticus M Pseudacris crucifer L, M M L, S, M, U I, L, S, U L M, U M L L, U L L, S Pseudacris feriarum M L, M, S, U I, L, S, U L, M M, U L L, M, U L, M, S L, M, S, U Scaphiopus holbrokii M I, U L S, U Southeastern Naturalist 207 S.E. Fields 2019 Vol. 18, No. 2 Table 2. Records of salamanders from counties in the South Carolina Piedmont. Abbreviations for counties are ABB (Abbeville), AND (Anderson), CHK (Cherokee), CHS (Chester), FAI (Fairfield), GRW (Greenwood), LAU (Laurens), MCC (McCormick), NEW (Newberry), SPA (Spartanburg), UNI (Union), YRK (York). Abbreviations for record types are: S = field surveys, observations, and raw data; U = unpublished reports and theses; L = published literature citations; I = image vouchers); and M = museum specimens. County Species ABB AND CHK CHS FAI GRW LAU MCC NEW SPA UNI YRK Order Caudata Ambystoma maculatum M L, M, U I, L, M, S, U U L, U L, M, S, U Ambystoma opacum L, M M L, S, U I, L, S, U L M, U L L L, S L, M, U Ambystoma talpoideum S Crytobranchus alleganiensis M Desmognathus auriculatus M L, M Desmognathus fuscus L, M M L, M, S, U I, L, S, U L L L, M L, M, U L, S L, M, S, U Desmognathus monticola M Desmognathus ocoee M M Desmognathus quadramaculatus M Eurycea chamberlaini M Eurycea cirrigera M M L, M, U L, M, S, U L L, M L, M L, M, S, U L, S I, L, M, S, U Eurycea guttolineata L, M M L, S, U L L L, M, U L, S U Eurycea quadridigitata M M Eurycea wilderae M Gyrinophilus porphyriticus M L, M, U L, S L, M, U Hemidactylium scutatum M U L, M Notophthalmus viridescens M M L M L L L Plethodon chlorobryonis L, M M L M, U M L, M S Plethodon cylindraceus L, M, U I, L, M, S, U U L L, U L I, M, S, U Plethodon tayahalee M M Plethodon websteri L, M Pseudotriton montanus M M M, U Pseudotriton ruber M M L, M, U I, U L M M, S, U L, S L, M, U Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 208 (2012), Reed and Gibbons (2005), Thomas (2002). Chester: CCARI, SEF, WUNH; Kern et al. (2012). Rudert (2005); specimens and images examined–MYCO; specimens reported–NCSM. Fairfield: Kern et al. (2012). Greenwood: Reed and Gibbons (2005); specimens reported–NCSM. McCormick: Metts and Nestor (2005); specimens reported–CUSC, NCSM. Newberry: Kern et al. (2012). Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005). Union: CCARI; Kern et al. (2012), Metts and Nestor (2005). York: CCARI, SEF, WU-AR; Brown (1992), Kern et al. (2012); images examined–MYCO; specimens examined–MYCO. Acris gryllus (LeConte, 1825) (Southern Cricket Frog). According to Dorcas and Gibbons (2008), the Southern Cricket Frog is primarily a coastal plain inhabitant. Powell et al. (2016) depicted a range that included much of the eastern and central South Carolina Piedmont, as well. The Southern Cricket Frog is sympatric with the Eastern Cricket Frog (Beane et al. 2010). Although the records of A. gryllus are fewer than A. crepitans, the majority come from western potions of the Piedmont. Abbeville: specimens reported–UTA. Chester: Dorcas et al. (2006). McCormick: specimens reported–USNM. Anaxyrus Tschudi, 1845 (North American toads). Some references retain the classification of the North American toads within the genus Bufo, but this genus concept was updated by Frost et al. (2006) and supported by Van Bocxlaer et al. (2010). Anaxyrus americanus (Holbrook, 1836) (American Toad). Formerly recognized as Bufo americanus, the American Toad is primarily a resident of the Blue Ridge and western Piedmont in South Carolina (Beane et al. 2010, Dorcas and Gibbons 2008). The records that I report here generally concur, but there are occurrences farther east and south. Abbeville: Metts and Nestor (2005). Anderson: Specimens examined–CUSV; specimens reported–USNM. Cherokee: Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Chester: Dorcas et al. (2006), Kern et al. (2012); specimens reported–NCSM. Fairfield: Kern et al. (2012), Metts and Nestor (2005). Laurens: Metts and Nestor (2005). McCormick: Metts and Nestor (2005). Newberry: Kern et al. (2012). Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005). Union: CCARI, Kern et al. (2012), Metts and Nestor (2005). York: Kern et al. (2012), Reed and Gibbons (2005), Rudert (2005). York: images examined–MYCO, specimens examined– MYCO; specimens reported–USNM Anaxyrus fowleri (Hinckley, 1882) (Fowler’s Toad). Some references still use Bufo fowleri or even the subspecies Bufo woodhousii fowleri with misspelled specific epithet of B. woodhousei (Crother 2017). Subspecies of B. woodhousii are currently recognized, but not within South Carolina. An earlier version of the Amphibians and Reptiles of the Carolinas and Virginia (Martof et al. 1980) listed the Fowler’s Toad as an eastern race of Bufo woodhousei, but Beane et al. (2010) did not report any race of B. woodhousii within the Carolinas or Virginia. According to Dorcas and Gibbons (2008) and Powell et al. (2016), B. woodhousii occurs west of the Mississippi River with a “hybrid zone” in western Louisiana and Arkansas. In this Southeastern Naturalist 209 S.E. Fields 2019 Vol. 18, No. 2 paper, I regard specimens identified as B. woodhousei or B. woodhousei fowleri as Anaxyrus fowleri. Abbeville: specimens reported–USNM. Anderson: specimens examined–CUSC; specimens reported–CM, NCSM, USNM. Cherokee: Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Chester: CCARI, SEF, WU-NH; Dorcas et al. (2006), Kern et al. (2012), Rudert (2005); images examined–MYCO. Fairfield: Kern et al. (2012); specimens reported– AMNH. Greenwood: specimens reported–NCSM, UMMZ. Laurens: specimens reported–NCSM, UMMZ. McCormick: Metts and Nestor (2005); specimens examined– CUSC, specimens reported–MVZ, NCSM, SDNHM, USNM. Newberry: Kern et al. (2012). Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005); specimens reported–AMNH. Union: CCARI, Kern et al. (2012), Metts and Nestor (2005); specimens reported–FM. York: CCARI, SEF, WU-AR; Kern et al. (2012), Rudert (2005); images examined–MYCO, specimens examined–MYCO, specimens reported–FUR, NCSM, USNM. Gastrophryne Fitzinger, 1843 (North American narrow-mouthed toads). Gastrophryne carolinensis (Holbrook, 1835) (Eastern Narrow-mouthed Toad). This species occurs throughout the Piedmont of South Carolina. Abbeville: specimens examined–CUSC; specimens reported–SLU, UTA. Anderson: specimens examined–CUSC; specimens reported–NCSM. Cherokee: Dorcas (2009a, b, c), Kern et al. (2012). Chester: CCARI, WU-NH; Dorcas et al. (2006), Metts and Nestor (2005), Rudert (2005). Images examined–MYCO. Fairfield: Kern et al. (2012). Greenwood: Reed and Gibbons (2005). McCormick: Metts and Nestor (2005). Newberry: Kern et al. (2012). Union: CCARI; Kern et al. (2012), Metts and Nestor (2005). York: CCARI, SEF, WU-AR; Rudert (2005); specimens examined– MYCO. Hyla Laurenti, 1768 (holarctic treefrogs). Hyla andersonii Baird, 1854 (Pine Barrens Treefrog). This species is uncommon and occurs in the Sandhills and Coastal Plain regions of the Carolinas (Beane et al. 2010). It is only known from 2 other areas, the southern Alabama/western Florida panhandle of Florida and the pine barrens of New Jersey (Dorcas and Gibbons 2008). A single preserved specimen, designated as the holotype, USNM 3600, is reported from Anderson County. However, this is a well-known subject of controversy in the literature (K. de Queiroz, Department of Vertebrate Zoology, National Museum of Natural History, Washington, DC, pers. comm.). See Amphibiaweb (2018) for a more detailed account. Hyla chrysoscelis Cope, 1880 (Cope’s Gray Treefrog). The Hyla versicolor Le Conte, 1825 (Gray Treefrog) complex represents a system of polyploid speciation that was reviewed by Holloway et al. (2006). Although H. chrysoscelis is diploid and its sibling species H. versicolor is tetraploid, the 2 species are visually indistinguishable in the field. However, vocalizations of each species are distinct. Martof et al. (1980) listed gray treefrogs as H. chrysoscelis and H. versicolor, but Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 210 in the recent version, Beane et al. (2010), and Dorcas and Gibbons (2008), reported only H. chrysoscelis in South Carolina with the range of H. versicolor only as far south as the North Carolina–Virginia border. I therefore report all species in the South Carolina Piedmont as Cope’s Gray Treefrog, H. chrysoscelis, but it should be noted that cellular analysis of individual specimens could reveal their identity as H. versicolor. Anderson: specimens reported–USNM; Cherokee: CCARI; Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002). Chester: CCARI, SEF (by vocalizations), WU-NH; Dorcas et al. (2006), Kern et al. (2012), Rudert (2005); images examined–MYCO. Fairfield: Kern et al. (2012). Newberry: Kern et al. (2012). Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005). Union: Kern et al. (2012). York: CCARI, SEF (by vocalization); Kern et al. (2012); images examined–MYCO. Hyla cinerea (Schneider, 1799) (Green Treefrog). Several authors cite a recent range extension from the Coastal Plain into the Piedmont for this species (Dorcas and Gibbons 2008). Indeed, the range for the Green Treefrog extends westward from that depicted in Martof et al. (1980) to Beane et al. (2010). Based on records found during this study, H. cinerea occurs throughout the South Carolina Piedmont, with recorded dates from 1990 through 2013. Abbeville: specimens examined–CUSC, specimens reported–UTA. Anderson: specimens examined–CUSC. Cherokee: Dorcas (2009c), Kern et al. (2012). Chester: CCARI, WU-NH; Dorcas et al. (2006), Kern et al, (2012), Rudert (2005); images examined–MYCO. Fairfield: Kern et al. (2012). Greenwood: Reed and Gibbons (2005); specimens examined–CUSC. McCormick: specimens examined–CUSC. Newberry: Kern et al. (2012). Spartanburg: CCARI; Kern et al. (2012); Union: Kern et al, (2012). York: CCARI, SEF (visual and vocalization); Kern et al. (2012), Rudert (2005); images examined–MYCO; specimens examined–MYCO. Hyla gratiosa LeConte, 1856 (Barking Treefrog). Beane et al. (2010) and Dorcas and Gibbons (2008) listed the Barking Treefrog as primarily an inhabitant of the Coastal Plain in the Carolinas. However, the relatively few records found during this study come from counties well within the South Carolina Piedmont. Abbeville: specimens reported–UTA. Greenwood: specimens examined–CUSC. McCormick: specimens reported–USNM. Hyla squirella Bosc, 1800 (Squirrel Treefrog). This is another Coastal Plain inhabitant that has likely expanded its range in the Piedmont (Beane et al. 2010, Martof et al. 1980). Chester: Dorcas et al. (2006); images examined–MYCO. McCormick: specimens reported–USNM. Lithobates Fitzinger, 1843 (American water frogs). The taxonomy of American ranids has been extensively disputed and discussed since Frost et al. (2006) moved them to the genus Lithobates, but Crother (2017) retained this move in the current Southeastern Naturalist 211 S.E. Fields 2019 Vol. 18, No. 2 nomenclature. I therefore refer to all records of Rana from the South Carolina Piedmont as Lithobates. Lithobates catesbianus (Shaw, 1802) (American Bullfrog). Some references (including Amphibiaweb 2018) retain Rana catesbiana (Shaw) as the name of this species, but this is not recognized by Crother (2017). Abbeville: specimens examined–CUSC. Anderson: specimens reported–USNM. Cherokee: Dorcas (2009a, b, c), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Chester: CCARI, SEF; Dorcas et al. (2006), Kern et al. (2012), Rudert (2005). Fairfield: Kern et al. (2012). McCormick: Metts and Nestor (2005); specimens reported–USNM. Newberry: Kern et al. (2012); specimens reported– UF. Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005); specimens reported–UF. Union: CCARI; Kern et al. (2012), Metts and Nestor (2005). York: CCARI, SEF, WU-AR; Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002), Rudert (2005); images examined–MYCO; specimens reported– NCSM, UMMZ. Lithobates clamitans (Latrielle, 1801) (Green Frog). This species is found throughout the Carolinas (Beane et al. (2010) and widespread throughout the eastern US (Powell et al. 2016). Abbeville: Metts and Nestor (2005); specimens examined–CUSC. Anderson: specimens examined–CUSC; specimens reported–CM, NCSM, UMMZ. Cherokee: Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Chester: CCARI, SEF, WU-NH; Dorcas et al. (2006), Rudert (2005); images examined–MYCO. Fairfield: Kern et al. (2012), Metts and Nestor (2005). Greenwood: Reed and Gibbons (2005); specimens reported–UF. Laurens: Metts and Nestor (2005). McCormick: Metts and Nestor (2005). Newberry: Kern et al. (2012). Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005); specimens reported–ANSP. Union: CCARI; Kern et al. (2012), Metts and Nestor (2005). York: CCARI, SEF, WU-AR; Kern et al. (2012), Reed and Gibbons (2005), Rudert (2005), Thomas (2002); specimens reported–UMMZ. Lithobates palustris (LeConte, 1825) (Pickerel Frog). Dorcas and Gibbons (2008) depicted a larger range for this species in the Carolinas than Beane et al. (2010), but both support a widespread occurrence in the South Carolina Piedmont. Cherokee: Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005); specimens reported–NCSM. Chester: Kern et al. (2012). Newberry: Kern et al. (2012). Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005). Union: Kern et al. (2012). York: Kern et al. (2012), Reed and Gibbons (2005). Lithobates sphenocephalus (Cope, 1886) (Southern Leopard Frog). This species maintains a long and complex taxonomic history as described by Crother (2017). Prior to the revision of Rana to Lithobates, Leopard Frogs were classified as Rana pipiens, R. utricularius, and R. utricularia, with subspecific designations further complicating the issue. Although further investigation is needed to resolve nominal subspecies, I follow Crother (2017) in recognizing the species L. sphenocephalus. Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 212 Abbeville: Metts and Nestor (2005). Anderson: specimens examined–CUSC; specimens reported–CM, NCSM. Cherokee: Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002). Chester: CCARI, SEF, WU-NH; Dorcas et al. (2006), Kern et al. 2012, Rudert (2005); images examined–MYCO; specimens reported–USNM. Fairfield: Kern et al. (2012). Greenwood: Reed and Gibbons (2005); specimens examined–CUSC. Laurens: Metts and Nestor (2005). McCormick: Metts and Nestor (2005). Newberry: Kern et al. (2012). Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005). Union: CCARI; Kern et al. (2012), Metts and Nestor (2005). York: CCARI, WU-AR; Brown (1992), Kern et al. (2012), Rudert (2005), Thomas (2002); specimens reported–UMMZ. Lithobates sylvaticus (LeConte, 1825) (Wood Frog). The range of the Wood Frog in the Carolinas appears to be confined to the Appalachian Mountains (Beane et al. 2010, Dorcas and Gibbons 2008, Powell et al. 2016). There is a single preserved specimen (USNM) reported from Anderson County, SC. I have not examined the specimen to verify the identification. Pseudacris Fitzinger, 1843 (chorus frogs). Pseudacris crucifer (Wied-Neuwied, 1838) (Spring Peeper). The Spring Peeper ranges across the Carolinas with widespread occurrence in the South Carolina Piedmont. Although Moriarty and Cannatella (2004) rejected subspecies (upheld in Powell et al. 2016), Dorcas and Gibbons (2008) listed the nominal subspecies P. c. crucifer Wied-Neuwied, 1838) (Northern Spring Peeper) as the inhabitant of South Carolina. Abbeville: Metts and Nestor (2005); specimens reported–USNM, UTA. Anderson: specimens reported–USNM; specimens examined–CUSC. Cherokee: SEF; Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005); specimens reported–LACM. Chester: CCARI, SEF, WU-NH; Dorcas et al. (2006), Kern et al. (2012), Rudert (2005); images examined–MYCO. Fairfield: Kern et al. (2012). Greenwood: Reed and Gibbons (2005); specimens reported–NCSM. Laurens: specimens reported–NCSM, UMMZ. Newberry: Kern et al. (2012). Spartanburg: Kern et al. (2012), Reed and Gibbons (2005). Union: Kern et al. (2012). York: CCARI, SEF, WU-AR; Kern et al. (2012). Pseudacris feriarum (Baird, 1854) (Upland Chorus Frog). Several species of chorus frogs have been reported as occurring in South Carolina (Beane et al. 2010, Dorcas and Gibbons 2008, Powell et al. 2016). However, based on the work of Lemmon et al. (2007), it appears that P. feriarum is the only species of Pseudacris that is valid for the South Carolina Piedmont. Because evidence from mitochondrial DNA appears to restrict the range of P. triseriata Wied-Neuwied (Western or Midland Chorus Frog) to Michigan, Indiana, Ohio, western Pennsylvania and southwestern New York (Lemmon et al. (2007), records of P. triseriata that were reported for South Carolina are treated here as P. feriarum. Anderson: specimens examined (reported as P. triseriata)–CUSC. Cherokee: SEF; Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002); specimens reported as P. triseriata–LACM, specimens reported–MPM, Southeastern Naturalist 213 S.E. Fields 2019 Vol. 18, No. 2 NCSM. Chester: CCARI, SEF, WU-NH; Dorcas et al. (2006), Kern et al. (2012), Rudert (2005); images examined–MYCO. Fairfield: Kern et al. (2012); specimens reported–NCSM. Greenwood: Reed and Gibbons (2005); specimens reported– NCSM, TNHC. Newberry: Kern et al. (2012). Spartanburg: Brown (1992), Kern et al. (2012), Reed and Gibbons (2005; as P. triseriata); specimens reported–NCSM. Union: CCARI; Brown (1992), Metts and Nestor (2005), Kern et al. (2012); specimens reported–NCSM. York: CCARI, SEF; Brown (1992), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002; as P. triseriata); specimens reported–NCSM. Scaphiopus Holbrook, 1836 (north american spadefoots). Scaphiopus holbrookii (Harlan, 1835) (Eastern Spadefoot). This species occurs throughout South Carolina but may go unnoticed due to its burrowing habits (Beane et al. 2010, Dorcas and Gibbons 2008), which could explain the relatively few records found during this study. Anderson: specimens reported–NCSM. Chester: WU-NH; Rudert (2005) images examined–MYCO. Newberry: Kern et al. (2012). York: CCARI, SEF, WU-AR; Rudert (2005). Order Caudata: Salamanders Ambystoma Tschudi, 1838 (mole salamanders) Ambystoma maculatum (Shaw, 1802) Spotted Salamander). The Spotted Salamander is widely distributed across South Carolina (Beane et al. 2010, Mitchell and Gibbons 2010). Abbeville: specimens reported–USNM. Cherokee: (Dorcas 2009b, c), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Chester: CCARI, SEF; Dorcas et al. (2006), Metts and Nestor (2005), Rudert (2005); images examined– MYCO; specimens examined–MYCO. Greenwood: Reed and Gibbons (2005). Spartanburg: Kern et al. (2012), Reed and Gibbons (2005). York: SEF; Brown (1992), Reed and Gibbons (2005), Thomas (2002); specimens examined– MYCO; specimens reported–FUR, NCSM. Ambystoma opacum (Gravenhorst, 1807) (Marbled Salamander). This salamander occurs throughout the southeastern US (Mitchell and Gibbons 2010), including all portions of South Carolina (Beane et al. 2010). Abbeville: Metts and Nestor (2005); specimens examined–CUSC. Anderson: specimens reported–USNM. Cherokee: CCARI; Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002). Chester: CCARI, SEF; Dorcas et al. (2006), Metts and Nestor (2005), Kern et al. (2012), Rudert (2005); images examined–MYCO. Fairfield: Metts and Nestor (2005). Greenwood: Reed and Gibbons (2005); specimens reported–NCSM. McCormick: Metts and Nestor (2005). Newberry: Kern et al. (2012). Union: CCARI; Kern et al. (2012), Metts and Nestor (2005). York: Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002); specimens examined–MYCO. Ambystoma talpoideum (Holbrook, 1838) (Mole Salamander). The range of the Mole Salamander is described as isolated and disjunct in the Carolinas, probably Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 214 because these salamanders spend most of their time under leaf litter or in burrows, and sightings are rare (Beane et al. 2010, Mitchell and Gibbons 2010). Two separate surveys (CCARI, SEF) recorded this species at Landsford Canal State Park in Chester County, SC. Cryptobranchus Leuckart, 1821 (hellbenders). Cryptobranchus alleganiensis (Daudin, 1803) (Hellbender). The range of the Hellbender is confined to the Appalachian Mountains in the Carolinas (Mitchell and Gibbons 2010). Beane et al. (2010:51) stated that “there are no recent records from South Carolina”. However, 3 preserved specimens (USNM) from Abbeville County, SC were reported to VertNet. No specific date or locality other than Abbeville was designated. I have not examined these specimens to verify their identification, but the presence of C. alleganiensis in the South Carolina Piedmont is highly suspect. Desmognathus Baird, 1850 (dusky salamanders) Desmognathus auriculatus (Holbrook, 1838) (Holbrook’s Southern Dusky Salamander). This salamander is regarded as a coastal plain inhabitant by Beane et al. (2010), Mitchell and Gibbons (2010), and Powell et al. (2016). Yet, several records exist for the Piedmont. Three preserved specimens (USNM) from McCormick County, SC were reported to VertNet. I have not examined these specimens to verify their identification. Brown (1992) reported D. auriculatus from York County, SC, as well as Piedmont counties (Iredell and Rowan) in North Carolina. Desmognathus fuscus (Rafinesque, 1820) (Northern Dusky Salamander). This is a common salamander throughout the northeastern US (Powell et al. 2016) and the Blue Ridge and Piedmont of the Carolinas (Beane et al. 2010, Mitchell and Gibbons (2010). Abbeville: Metts and Nestor (2005); specimens reported–USNM; specimens examined–CUSC. Anderson: specimens reported–NCSM, UMMZ, USNM; specimens examined–CUSC. Cherokee: CCARI; Brown (1992), Dorcas (2009a, b, c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Chester: CCARI, WU-NH; Dorcas et al. (2006), Rudert (2005); images examined–MYCO. Fairfield: Metts and Nestor (2005). Laurens: Metts and Nestor (2005). McCormick: Metts and Nestor (2005); specimens reported–NCSM, USNM. Spartanburg: Kern et al. (2012), Reed and Gibbons (2005); specimens reported– AMNH, NCSM. Union: CCARI; Metts and Nestor (2005). York: CCARI; Brown (1992), Reed and Gibbons (2005), Thomas (2002); specimens reported– NCSM, UMMZ. Desmognathus monticola Dunn, 1916 (Seal Salamander). While the range of D. monticola in the Carolinas appears to be restricted to the Blue Ridge province (Mitchell and Gibbons 2010), Beane et al. (2010:75) reported a range “in portions of the adjacent Piedmont.” Two specimens from Anderson County, SC, exist (CUSC). I have verified their identification as D. monticola. Desmognathus ochrophaeus Cope, 1859 (Allegheny Mountain Dusky Salamander) and Desmognathus ocoee Nicholls, 1949 (Ocoee Salamander). These 2 species, Southeastern Naturalist 215 S.E. Fields 2019 Vol. 18, No. 2 along with D. carolinensis and D. orestes, are part of the Mountain Dusky Salamander complex. Beane et al. (2010). Mitchell and Gibbons (2010) emphasized the importance of knowing specimen locality to make valid identifications of these cryptic species. Crother (2017) commented on these parapatric units as occupying primarily mountain provinces in the eastern US. Based on this information, I report all South Carolina Piedmont occurrences of dusky salamanders as D. ocoee. Anderson: specimens reported–USNM. Spartanburg: specimens reported– NCSM. Records reported as D. ochrophaeus include: Anderson: specimens reported– UMMZ; specimens examined–CUSC. Desmognathus quadramaculatus (Holbrook, 1840) (Black-bellied Salamander). This is another species that is reported to have a range in the Carolinas restricted to the Blue Ridge province (Beane et al. 2010, Mitchell and Gibbons 2010). Two preserved specimens (USNM) from Abbeville County with no specific locality other than Abbeville were reported to VertNet. I have not examined these specimens to verify their identification. Eurycea Rafinesque, 1822 (brook salamanders). Eurycea bislineata (Green, 1818) (Northern Two-lined Salamander). The range of this species includes northern Virginia, but not the Carolinas (Beane et al. 2010, Mitchell and Gibbons (2010). Specimens reported from the South Carolina Piedmont in this study are regarded as E. cirrigera. See species account below. Eurycea chamberlaini Harrison and Guttman, 2003 (Chamberlain’s Dwarf Salamander). This salamander has a spotty range within the Carolinas, including the South Carolina Piedmont (Beane et al. 2010, Mitchell and Gibbons 2010). Three specimens (NCSM) reported from Anderson County fall within the range depicted. Eurycea cirrigera (Green 1831) (Southern Two-lined Salamander). According to Beane et al. (2010), some researchers refer this salamander to a subspecies of E. bislineata. However, Crother (2017) recognized no such designation. Abbeville: specimens reported–NCSM, USNM; specimens reported as E. bislineata– UMMZ; specimens examined–CUSC. Anderson: specimens reported–CM, USNM; specimens reported as E. bislineata–NCSM; specimens examined–CUSC. Cherokee: Dorcas (2009a, b), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Chester: CCARI; Dorcas et al. (2006); Kern et al. (2012), Rudert (2005) (as E. bislineata); specimens reported–NCSM. Fairfield: Metts and Nestor (2005). Laurens: Metts and Nestor (2005); specimens reported– UF. McCormick: Metts and Nestor (2005); specimens reported–NCSM; specimens reported as E. bislineata complex–USNM. Spartanburg: CCARI; Kern et al. (2012), Reed and Gibbons (2005); specimens reported–NCSM. Union: CCARI; Metts and Nestor (2005). York: CCARI; Brown (1992), Reed and Gibbons (2005), Thomas (2002); York: images examined–MYCO; specimens reported–NCSM. Eurycea guttolineata (Holbrook, 1838) (Three-lined Salamander). This species was formerly regarded as a subspecies of E. longicauda (Conant and Collins 1991). Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 216 E. longicauda occurs in the Appalachian Mountains of Virginia with isolated populations in the Blue Ridge province of North Carolina, but is not known to occur in South Carolina (Beane et al. 2010). Specimens of E. longicauda reported from Abbeville and Anderson counties, SC, are regarded as E. guttolineata in this paper. Abbeville: Metts and Nestor (2005); specimens reported–NCSM; specimens reported as E. longicauda–USNM; specimens examined–CUSC. Anderson: specimens reported–NCSM; specimens examined–CUSC; specimens examined (reported as E. longicauda)–CUSC. Cherokee: CCARI; Dorcas (2009c), Kern et al. (2012), Reed and Gibbons (2005), Thomas (2002). Chester: Dorcas et al. (2006). Newberry: Kern et al. (2012). Spartanburg: Kern et al. (2012), Reed and Gibbons (2005); specimens reported–NCSM. Union: CCARI; Metts and Nestor (2005). York: Reed and Gibbons (2005), Thomas (2002). Eurycea longicauda (Green, 1818) (Long-tailed Salamander). See account of E. guttolineata. Eurycea quadridigitata (Holbrook, 1842) (Southeastern Dwarf Salamander). Beane et al. (2010) restricted the range of this species in South Carolina to the Coastal Plain, but Mitchell and Gibbons (2010) and Petranka (1998) also recognized a population in the upper Piedmont of South Carolina. The records that I located in this study support the increased range. Anderson: specimens reported–UF; specimens examined–CUSC. Spartanburg: specimens reported–UF. Eurycea wilderae Dunn, 1920 (Blue Ridge Two-lined Salamander). E. wilderae was once considered a subspecies of E. bislineata (Petranka 1998), but Crother (2017) recognized its status a separate species. The Blue Ridge Two-lined Salamander, as its common name indicates, primarily inhabits the mountain regions of the Carolinas and Virginia (Beane et al. 2010, Mitchell and Gibbons 2010). Although 10 preserved specimens (CUSC) were reported from several localities in Anderson County, SC, I have not examined the specimens to verify their identifications as E. wilderae. Gyrinophilus Cope, 1899 (spring salamanders) Gyrinophilus porphyriticus (Green, 1827) (Spring Salamander). The Spring Salamander ranges throughout the Appalachian Mountains and upper Piedmont of the southeastern US (Mitchell and Gibbons 2010) and the same provinces within South Carolina (Beane et al. 2010). Anderson: specimens reported–AMNH, UMMZ. Cherokee: Dorcas (2009b), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Spartanburg: CCARI; Kern et al. (2012). York: Brown (1992), Reed and Gibbons (2005), Thomas (2002); specimens reported–NCSM. Hemidactylium Tschudi, 1838 (four-toed salamander) Hemidactylium scutatum (Temminck and Schlegel in Von Siebold, 1838) (Fourtoed Salamander). Although the range of this species extends across the eastern Southeastern Naturalist 217 S.E. Fields 2019 Vol. 18, No. 2 US (Petranka 1998), populations in the Southeast are described as scattered and fragmented (Beane et al. 2010, Mitchell and Gibbons 2010). Anderson: specimens reported–USNM. York: Brown (1992) tentatively identified by eggs located at Kings Mountain State Park in 1975, 1979, and 1980; specimens reported–NCSM. Notophthalmus Rafinesque, 1820 (eastern newt) Notophthalmus viridescens (Eastern Newt). The Eastern Newt occupies an extensive range across the southeastern US (Mitchell and Gibbons (2010) and throughout the Carolinas (Beane et al. 2010). Abbeville: specimens reported–CUMV, USNM; specimens examined–CUSC. Anderson: specimens reported–NCSM, USNM; specimens examined–CUSC. Cherokee: Dorcas (2009a, b, c), Kern et al. (2012). Fairfield: specimens reported– UMMZ. McCormick: Metts and Nestor (2005); Spartanburg: Kern et al. (2012). Union: Kern et al. (2012). Plethodon Tschudi, 1838 (woodland salamanders) Salamanders formerly classified as Plethodon glutinosus have been reevaluated in recent decades. Highton et al. (1989) analyzed genetic variation in populations from 129 localities throughout the eastern US and recognized 15 species within the Plethodon glutinosus complex. Petranka (1998) only accepted 3 of the 15 species proposed by Highton et al. (1989). However, Crother (2000) upheld the taxonomy of Highton et al. (1989). Under the revised taxonomy, 2 species occur within the South Carolina Piedmont. Highton et al. (1989) confined the range of P. cylindraceus in South Carolina to the northern Piedmont and the range of P. chlorobryonis to the southern Piedmont and Coastal Plain. P. glutinosus was restricted to the Blue Ridge province with no occurrences in South Carolina. More recent references (Beane et al. 2010, Mitchell and Gibbons 2010, Powell et al. 2016) reflect these changes in distribution for the respective species. Plethodon chlorobryonis Mittleman, 1951 (Atlantic Coast Slimy Salamander). Abbeville: Metts and Nestor (2005); specimens reported–USNM, UTA, specimens examined (as P. glutinosus)–CUSC. Anderson: specimens reported (as P. glutinosus)–UMMZ, USNM, specimens examined (as P. glutinosus)–CUSC. Cherokee: Dorcas (2009a, b, c). Greenwood: Reed and Gibbons (2005) (as Plethodon glutinosus complex); specimens reported–NCSM. Laurens: specimens reported–USNM. McCormick: Metts and Nestor (2005); specimens reported– NCSM, USNM. Union: CCARI. Plethodon cylindraceus (Harlan, 1825) (White-spotted Slimy Salamander). Cherokee: Kern et al. (2012), Reed and Gibbons (2005) (as Plethodon glutinosus complex), Thomas (2002); specimens reported–NCSM; specimens examined (as P. glutinosus)–CUSC. Chester: CCARI, WU-NH; Dorcas et al. (2006), Rudert (2005) (as P. glutinosus); specimens reported–USNM; images examined (as P. glutinosus)–MYCO. Laurens: Metts and Nestor (2005). Spartanburg: Kern et al. Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 218 (2012), Reed and Gibbons (2005) (as Plethodon glutinosus complex). Union: Metts and Nestor (2005). York: CCARI, SEF; Reed and Gibbons (2005) (as Plethodon glutinosus complex), Thomas (2002); images examined–MYCO; specimens reported– NCSM. Plethodon teyahalee Hairston, 1950 (Southern Appalachian Salamander). This is another species whose reported range is limited to the Blue Ridge, but hybridization is known to occur with members of the slimy salamander complex (Beane et al. 2010, Mitchell and Gibbons 2010). Abbeville: specimens reported USNM. Anderson–specimens reported–NCSM, USNM; specimens examined–CUSC. Plethodon websteri Highton, 1979 (Webster’s Salamander). Beane et al. (2010) reported this species as occurring in Edgefield, Greenwood, and McCormick counties in South Carolina. I found records for Edgefield County and McCormick County: Metts and Nestor (2005); specimens reported–USNM; specimens examined CUSC. Pseudotriton Tschudi, 1838 (red and mud salamanders) Pseudotriton montanus Baird, 1850 (Mud Salamander). The Mud Salamander inhabits the Piedmont and Coastal Plain regions of the Carolinas (Beane et al. 2010, Mitchell and Gibbons 2010). Anderson: specimens reported–NCSM, USNM; specimens examined–CUSC. Cherokee: Reed and Gibbons (2005). Laurens: specimens reported–UTA. York: Reed and Gibbons (2005); specimens examined–MYCO. Pseudotriton ruber (Sonnini de Manoncourt and Latreille, 1801) (Red Salamander). This species is common across much of the southeastern US (Mitchell and Gibbons 2010), including the Blue Ridge and Piedmont of the Carolinas (Beane et al. 2010). Abbeville: specimens reported–USNM; specimens examined–CUSC. Anderson: specimens reported–AMNH, UMMZ, USNM; specimens examined–CUSC. Cherokee: Dorcas 2009a, b), Reed and Gibbons (2005); specimens reported– NCSM. Chester: Rudert (2005); images examined–MYCO. Laurens: Metts and Nestor (2005). McCormick: specimens reported–USNM. Spartanburg: CCARI; Reed and Gibbons (2005); specimens reported–NCSM. Union: CCARI; Metts and Nestor (2005). York: Brown (1992), Reed and Gibbons (2005); specimens reported– NCSM. Discussion I conclude that 42 species of amphibians are recognized within the Piedmont physiographic province of South Carolina. Other surveys could potentially suggest more species, but recent changes in taxonomy would likely be the reason for any discrepancy. As with mammals, none of the amphibians listed in this report are endemic to the Piedmont. Rather, their distributions cross into the Blue Ridge and/or Atlantic Coastal Plain (see Fields 2007 for a report on mammals of the South Carolina Piedmont and a discussion of the Piedmont as a zone of faunal transition). Southeastern Naturalist 219 S.E. Fields 2019 Vol. 18, No. 2 The lack of primary literature on amphibian occurrence in the South Carolina Piedmont necessitates reliance on secondary references. I relied heavily on Beane et al. (2010), Dorcas and Gibbons (2008), and Reed and Gibbons (2010), and to a lesser degree, Petranka (1998) for written distribution data and range maps. Compared to Petranka (1998), information presented herein suggests possible range extensions for some species: Acris gryllus–records from Abbeville, Chester, and McCormick counties extend the range north and west; Anaxyrus americanus–records from Chester, Fairfield, Union, and York counties extend the range south and east; Hyla andersonii–the single type-specimen from Anderson County, SC, is the subject of much controversy (see species account), but would represent a westward range extension if valid; Hyla versicolor is now restricted to a range far north of South Carolina (see species account), with all South Carolina reports now regarded as H. chrysoscelis, but verification of vocalizations or chromosome number is required for unequivocal identification; Hyla squirella–records from Chester and McCormick counties extend the range west; Lithobates sylvaticus–a single record from Anderson County (if valid) would extend the range eastward. Similar range extensions are possible for salamanders in the South Carolina Piedmont: Cryptobranchus alleganiensis—records from Abbeville County, if valid, would extend the range eastward; Desmognathus auriculatus—records from McCormick and York counties extend the range to the west; Desmognathus quadramaculatus—records from Abbeville County would extend the range east; Plethodon tayahalee—records from Abbeville and Anderson counties would extend the range east. Besides representing possible range extensions, amphibian locality data are important to conservation issues. The South Carolina Department of Natural Resources maintains a list of rare, threatened, and endangered species for the state (SCDNR 2015). Seven amphibian species (4 anurans and 3 caudatans) are listed. Acris crepitans is globally and state-ranked as secure. Hyla andersonii is globally ranked as uncommon with cause for long-term concern, listed as a threatened species within the state, and considered vulnerable on the state level, due to range restriction, few populations, and widespread declines. Lithobates palustris is globally ranked as secure but has not yet been assessed on the state level. Lithobates sylvaticus has a global rank of secure, but is considered vulnerable on the state level, citing range restriction, few populations, and widespread declines. Eurycea chamberlaini has a global ranking of uncommon, with cause for long-term concern; it is not yet assessed on the state level. Hemidactylum scutatum has a global rank of secure, but is not yet assessed on the state level. Plethodon websteri has a global rank of vulnerable; citing range restriction, few populations, and widespread declines, it is listed as a state endangered species with a state ranking of imperiled. These global and state rankings emphasize the need for information on amphibian species richness. This paper represents the baseline of such occurrence data for amphibians of the South Carolina Piedmont. This information is a starting point for further explorations into relative abundance and other studies to aid understanding and conservation of our state’s rich herpetofaunal diversity. Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 220 Acknowledgments I thank Melissa Fuentes, Stanlee Miller, and Travis Perry for access to museum collections and data. I also thank Mike Dorcas, Al James, Nicole Hoekstra, and Angel Rudert for providing field data on species occurrence, and the following individuals for providing voucher images of amphibians: Jim Augustin, Irene Boland, Stephen Crotts, John Garton, Nancy Leach, and Andrew Lazenby. I am grateful to 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: James Barnwell, Al James, Jim Knight, Jim Langley, Andrew Lazenby, Jason Moretz, Don Oneppo, Irvin Pitts, Mike Perrot, Bill Rogers, Andrea Sorrow, Jim Sorrow, Ken Wagner, Jeannie Wray, and Sarah Young. Field collections were authorized under permits issued by the South Carolina Department of Natural Resources. Funding was provided by the York County Culture and Heritage Commission. Finally, I thank Whit Gibbons, Jim Knight, and 1 anonymous reviewer of the manuscript for helpful comments to improve this paper. Literature Cited AmphibiaWeb. 2018. Online database of information relating to amphibian biology and conservation. University of California, Berkeley, CA. Available online at http://amphibiaweb. org. Accessed 23 May 2018. Beane, J.C., A.L Braswell, J.C. Mitchell, W.M. Palmer, and J.R. Harrison. 2010. Amphibians and Reptiles of the Carolinas and Virginia. 2nd Edition. The University of North Carolina Press, Chapel Hill, NC. 274 pp. Blaustein, A., and D. Wake. 1995. The puzzle of declining amphibian populations. Scientific American 272(4):52–57. Blaustein, A., D. Wake, and W. Sausa. 1994. Amphibian declines: Judging stability, persistence, and susceptibility of populations to local and global extinctions. Conservation Biology 8(1):60–71. Brown, E.E. 1992. Notes on amphibians and reptiles of the western Piedmont of North Carolina. Journal of the Elisha Mitchell Scientific Society 108( 1):38–54. Conant, R., and J.T. Collins. The Peterson Field Guide to Reptiles and Amphibians, 4th Edition. Houghton Mifflin Company, Boston, MA. 450 pp. Corn, P.S. 1994. Straight-line drift fences and pitfall traps. Pp. 109–117, In W.R. Heyer, M.A. Donnelly, R.W. McDiarmid, L.C. Hayek, and M.S. Foster (Eds.). Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Washington, DC. 364 pp. Crother, B.I. (Ed). 2000. Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in Our Understanding, 5th Edition. Society for the Study of Amphibians and Reptiles (SSAR) Herpetological Circular 29. 82 pp. Crother, B.I. (Ed.). 2008. Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in Our Understanding, 6th Edition. Society for the Study of Amphibians and Reptiles (SSAR) Herpetological Circular 37. 94 pp. Crother, B.I. (Ed.). 2017. Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in Our Understanding, 8th Edition. Society for the Study of Amphibians and Reptiles (SSAR) Herpetological Circular 43. 102 pp. Southeastern Naturalist 221 S.E. Fields 2019 Vol. 18, No. 2 Crump, M.L., and N.J. Scott Jr. 1994. Visual encounter surveys. Pp. 84–92, In W.R. Heyer, M.A. Donnelly, R.W. McDiarmid, L.C. Hayek, and M.S. Foster (Eds.). Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Washington, DC. 364 pp. Dorcas, M.E. 2009a. Herpetological surveys of the railroad corridor between Gaffney and the W.S. Lee III Nuclear Station, Cherokee County, SC. Final report to Duke Power Company, Charlotte, NC. Dorcas, M.E. 2009b. Herpetological survey of London Creek, Cherokee County, South Carolina and its vicinity. Final report to Duke Power Company, Charlotte, NC. Dorcas, M.E. 2009c. Herpetological survey of the W.S. Lee III Nuclear Station, South Carolina. Final report to Duke Power Company, Charlotte, NC. Dorcas, M.E., and J.W. Gibbons. 2008. Frogs and Toads of the Southeast. The University of Georgia Press, Athens, GA. 238 pp. Dorcas, M.E., S.J. Price, and G.E. Vaughan. 2006. Amphibians and reptiles of the Great Falls Bypassed Reaches in South Carolina. Journal of the North Carolina Academy of Science 122(1):1–9. Fields, S.E. 2007. Mammals of the north-central piedmont of South Carolina. Southeastern Naturalist 6(4):577–596. Frost, D.R., T. Grant, J. Faivovich, R.H. Bain, A. Haas, C.F.B. Haddad, R.O. DeSa, A. Channing, M. Wilkinson, S.C. Donnellan, C.J. Raxworthy, J.A. Campbell, B. L. Blotto, P. Moler, R.C. Drewes, R.A. Nussbaum, J.D. Lynch, D.M. Green, and W.C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297:1–370. Godfrey, M.A. 1980. A Sierra Club Naturalist’s Guide to the Piedmont. Sierra Club Books, San Francisco, CA. 499 pp. Griffith, G.E., J.M Omernik, J.A. Comstock, J.B. Glover, and V.B. Shelburne. 2002. Ecoregions of South Carolina (map scale 1:1,500,000). US Environmental Protection Agency, Corvallis, OR. Heyer, W.R., M.A. Donnelly, R.W. McDiarmid, L.C. Hayek, and M.S. Foster (Eds.). 1994. Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian institution Press, Washington, DC. 364 pp. Highton R., G.C. Maha, and L.R. Maxson. 1989. Biochemical evolution in the slimy salamanders of the Plethodon glutinosus complex in the eastern United States. Illinois Biological Monographs 57:1–151. Hoekstra, N. 2013. Studying Batrachochytrium dendrobatidis (Bd) in amphibian populations at Landsford Canal State Park. M.Sc. Thesis. Winthrop University, Rock Hill, SC. 74 pp. Holloway, A.K., D.C. Cannatella, H.C. Gerhardt, and D.M. Hillis. 2006. Polyploids with different origins and ancestors form a single sexual polyploid species. The American Naturalist 167(4):E88–E101. Houlahan, J.E., C.S. Findlay, B.R. Schmidt, A.H. Meyer, and S.L. Kuzmin. 2000. Quantitative evidence for global amphibian population declines. Nature 404:752–755. Kern, M.M., J.C. Guzy, S.J. Price, S.D. Hunt, E.A. Eskew, and M.E. Dorcas. 2012. Riparian- zone amphibians and reptiles within the Broad River Basin of South Carolina. Journal of the North Carolina Academy of Science 128(3/4):81–87. Lemmon, E.M., A.R. Lemmon, J.T. Collins, J.A. Lee–Yaw, and D.C. Cannatella. 2007. Phylogeny-based delimitation of species boundaries and contact zones in the trilling chorus frogs (Pseudacris). Molecular Genetics and Evolution 44:1068–1082. Southeastern Naturalist S.E. Fields 2019 Vol. 18, No. 2 222 Martin, W.H., S.G. Boyce, and A.C. Echternacht (Eds.). 1993. Biodiversity of the Southeastern United States: Upland Terrestrial Communities. John Wiley and Sons, Inc., New York, NY. 373 pp. Martof, B.S., W.M. Palmer, J.R. Bailey, and J.R. Harrison III. 1980. Amphibians and Reptiles of the Carolinas and Virginia. The University of North Carolina Press, Chapel Hill, NC. 264 pp. Metts, B.S., and J.P Nestor. 2005. Amphibian and reptile diversity in the Sumter National forest of South Carolina. Journal of the North Carolina Academy of Science 121(3):117–124. Mitchell, J., and J.W. Gibbons. 2010. Salamanders of the Southeast. The University of Georgia Press, Athens, GA. 324 pp. Moriarty, E.C., and D.C. Cannatella. 2004. Phylogenetic relationships of the North American chorus frogs (Pseudacris: Hylidae). Molecular Phylogenetics and Evolution 30(2):409–420. Murphy, C.H. 1995. Carolina Rocks: The Geology of South Carolina. Sandlapper Publishing, Inc., Orangeburg, SC. 8 pp. North Carolina Museum of Natural Sciences. 2018. Herpetology collection overview. Available online at http://naturalsciences.org/research–collections/herpetology–collection. Accessed 28 May 2018. Petranka, J.W, 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington, DC. 592 pp. Pittman, S.E., and M.E. Dorcas. 2006. Catawba River Corridor coverboard program: A citizen science approach to amphibian and reptile inventory. Journal of the North Carolina Academy of Sciences 122(4):142–151. Powell, R., R. Conant, and J.T. Collins. 2016. Peterson Field Guide to Reptiles and Amphibians of Eastern North America, 4th Edition. Houghton Mifflin Harcourt Publishing Company, New York, NY. 494 pp. Reed, R.N., and J.W. Gibbons. 2005. Results of a herpetofaunal survey of five national park units in South Carolina. A report prepared for the National Park Service. Savannah River Ecology Laboratory, Aiken, SC. 73 pp. Rudert, A.A. 2005. Effects of habitat fragmentation on herpetofauna in the central Piedmont of North and South Carolina. M.Sc. Thesis. Winthrop University, Rock Hill, SC. Skeen, J.N., P.D. Doerr, and D.H. Van Lear. 1993. Oak–hickory–pine forests. Pp. 1–33, In W.H. Martin, S.G. Boyce, and A.C. Echternacht (Eds.). Biodiversity of the Southeastern United States: Upland Terrestrial Communities. John Wiley and Sons, Inc., New York, NY. 373 pp. Skerratt, L.F., L. Berger, R. Speare, S. Cashins, K.R. McDonald, A.D. Phillott. H.B. Hines, and N. Kenyon. 2007. Spread of Chytridiomycosis has caused the rapid global decline and extinction of frogs. EcoHealth 4:125–134. South Carolina Department of Natural Resources (SCDNR). 2015. South Carolina rare, threatened, and endangered species inventory. Columbia, SC, USA. Available online at http://www.dnr.sc.gov/species/state.html. Accessed 28 May 2018. Stein, B.A. 2002. States of the Union: Ranking America’s biodiversity. NatureServe, Arlington, VA. Available online at http://www.natureserve.org/biodiversity-science/publications/ states-union-ranking-americas-biodiversity. Accessed 25 May 2018. Stuart, S., J. Chanson, N. Cox, B. Young, A. Rodrigues, D. Fischman, and R. Waller. 2004. Status and trends of amphibian declines and extinctions worldwide. Science 306:1783–1786. Taylor, D. 1998. South Carolina Naturalists: An Anthology, 1700–1860. University of South Carolina Press, Columbia, SC. 241 pp. Southeastern Naturalist 223 S.E. Fields 2019 Vol. 18, No. 2 Thomas, R.B. 2002. Herpetofaunal inventory of Kings Mountain National Military Park. Final Report to the National Park Service. University of South Carolina at Spartanburg, Spatanburg, SC. 15 pp. Todd, B.D., C.T. Winne, J.D. Willson, and J.W. Gibbons. 2007. Getting the drift: Examining the effects of timing, trap type, and taxon on herpetofaunal drift fence surveys. American Midland Naturalist 158:292–305. Van Bocxlaer, I., S.P. Loader, K. Roelants, S.D. Biju, M. Menegon, and F. Bossuyt. 2010. Gradual adaptation toward a range-expansion phenotype initiated by the global radiation of toads. Science 327(5966):679–682. VertNet. 2018. Online vertebrate biodiversity database. Available at http://vertnet.org/. Accessed 20–30 April 2018. Voyles, J., E. Rosenblem, and L. Berger. 2011. Interactions between Batrachochytrium dendrobatidis and its amphibian hosts: A review of pathogenesis and immunity. Microbes and Infection 13:25–32. Witczak, L.R., and M.E. Dorcas. 2009. What are frogs and snakes worth? The economic value of reptiles and amphibians inhabiting the Charlotte–Metropolitan area of North Carolina. Journal of the North Carolina Academy of Science 125(4):131–137.