6?// 7/ . /3 /Jg B8& number 3 July 1980 EDITORIAL STAFF John E. Cooper, Editor Alexa C. Williams, Managing Editor John B. Funderburg, Editor-in-Chief Alvin L. Braswell, Curator of Lower Vertebrates, N.C. State M useum John C. Clamp, Associate Curator ( Invertebrates ), JV. C. State M useum Martha R. Cooper, Associate Curator ( Crustaceans), N.C. State Museum James W. Hardin, Department of Botany, N.C. State University Board David S. Lee, Chief Curator of Birds and Mammals, N.C. State Museum William M. Palmer, Chief Curator of Lower Vertebrates, N.C. State Museum Thomas L. Quay, Department of ^oology, N.C. State University Rowland M. Shelley, Chief Curator of Invertebrates, N.C. State Museum Brimleyana, the Journal of the North Carolina State Museum of Natural His- tory, will appear at irregular intervals in consecutively numbered issues. Con- tents will emphasize zoology of the southeastern United States, especially North Carolina and adjacent areas. Geographic coverage will be limited to Alabama, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North Carolina, South Carolina, Tennessee, Virginia, and West Virginia. Subject matter will focus on taxonomy and systematics, ecology, zoo- geography, evolution, and behavior. Subdiscipline areas will include general in- vertebrate zoology, ichthyology, herpetology, ornithology, mammalogy, and paleontology. Papers will stress the results of original empirical field studies, but synthesizing reviews and papers of significant historical interest to southeastern zoology will be included. Suitability of manuscripts will be determined by the Editorial Board, and ap- propriate specialists will review each paper adjudged suitable. Final ac- ceptability will be decided by the Editor. Address manuscripts and all cor- respondence (except that relating to subscriptions and exchange) to Editor, Brimleyana , N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 27611. In citations please use the full name — Brimleyana. North Carolina State Museum of Natural History North Carolina Department of Agriculture James A. Graham, Commissioner CODN BRIMD 7 ISSN 0193-4406 A Late Pleistocene Vertebrate Assemblage from Edisto Island, South Carolina Janet A. Roth Department of Geology, University of Georgia AND Joshua Laerm1 . Museum of Natural History, University of Georgia, Athens, Georgia 30602 ABSTRACT. — Edisto Island, South Carolina, was discovered as a ver- tebrate fossil locality during the early 1900s. Fossils collected by amateurs and professionals comprise an extensive, predominantly Late Pleistocene assemblage (approximately 10,000 to 120,000 years B. P.). The 37 genera present represent 9 orders of mammals and 3 orders of reptiles. Although the fossiliferous rock unit(s) from which the ver- tebrate material is derived remains unidentified, original deposition is considered to be alluvial and extensive regionally. The fauna gives evidence of a nearshore environment, including streams and marshes, and adjacent open savannah with scattered wooded areas. Several species, including the giant tortoise, Geochelone, indicate a probable frost-free climate. In additon, the occurence of Palaeolama mirifica, Glyptotherium floridanum, and Geochelone sp. at Edisto, extends the known geographic range of these species 322 kilometers (200 miles) northward. INTRODUCTION Terrestrial Pleistocene biostratigraphic zones are most clearly defined in western North America, and the mammal ages that serve as subdivisions within the Pleistocene are based on faunas from classic western localities (Fig. 1). In addition, the geologic ranges of many Pleistocene vertebrate species have been determined predominantly on the basis of their western occurrences (Hibbard 1958). In contrast to the relatively clear picture of western North America, Pleistocene vertebrate paleontology in the southeastern United States is less well defined, and correlation with western localities has proved dif- ficult (Hay 1923; Simpson 1929a; Hibbard 1958). However, Webb (1974) argued that the western mammal ages can be extended to include Florida. In all probability, the same applies to the rest of the Southeast. Recently, interest in the Pleistocene vertebrates of the Southeast has increased, and several important contributions have been made. These include Webb’s (1974) summary of the Florida Pleistocene, a substantial 'Address for correspondence and reprint requests. Brimleyana No. 3: 1-29 July 1980. 1 2 Janet A. Roth and Joshua Laerm TIME (M.Y.) -0.01 GLACIAL STAGES Sea Level Low — High NORTH AMERICAN MAMMAL AGES Wisconsinian Sangamonian / Rancholabrean (Rancho La Brea, Los Angeles Co., California) -0.12 7 o o l / Illinoian Yarmouthi'an / Irvingtonian (Irvington, Alameda Co., California) Kansan' \ Aftonian \ / Blancan (Mt . Blanco, Crosby Co . , Texas) Nebraskan -1.60 Fig. 1. Pleistocene glacial stages and North American mammal ages. Sea levels from Webb (1974). Absolute dates from Berggren and Van Couvering (1974). Pleistocene fauna from Ladds Quarry, Bartow County, Georgia (Lipps and Ray 1967), and Pleistocene vertebrates in the Georgia piedmont (Voorhies 1974) and coastal regions (Frey et al. 1975). However, many areas of the Southeast have not yet been investigated, and distribution of certain species has not yet been adequately documented. Vertebrate fossils at Edisto Island, South Carolina, were first reviewed by Hay (1923). Subsequently, Auffenberg (1957), Ray, (1965, 1967), Ray et al. (1968), Kurten (1973), and Dobie and Jackson (1979) discussed individual species from Edisto. For closely comparable faunas in the Charleston and Ashley River areas see Hay (1923) , Allen (1926), and Hibbard et al. (1965). O’Kelley (1976) presented a preliminary faunal list from Edisto, based primarily on Holocene material. However, no comprehensive study of the Edisto fossil vertebrates has been attempted. The abundance of fossil material from the island coupled with its coastal geographic locale north of previously reported localities provides new in- formation on paleoclimatic conditions of the Edisto area and the geographic range of particular species. The purposes of this study are to: (1) report a rather extensive (36 species of reptiles and mammals), predominantly late Pleistocene fauna from Edisto Island, South Carolina; (2) consider the paleoecological and Edisto Island Fossil Vertebrates 3 paleoclimatic implications of the fauna; and (3) discuss the significance of the fauna in relation to other southeastern Pleistocene localities. GEOGRAPHIC AND GEOLOGICAL SETTING Edisto Island is a barrier island located 34 kilometers (21 miles) southwest of Charleston, South Carolina, at longitude 80°17’W and latitude 30°31’N (Fig. 2). Fossils are found in the intertidal zone along the beaches, along stream banks and estuaries, and occasionally are en- trapped in fishing nets. The fossil material is typically well mineralized and black in color. The fossils have not yet been found in situ and the fossiliferous stratum (or strata) has not been identified. There are, however, several possible source areas: (1) Holocene sediments on or near the beach, (2) the Silver Bluff formation of latest Pleistocene age which, according to Colquhoun (1969), lies immediately inland of the Holocene sediments, or (3) earlier Pleistocene strata which underlie the previously mentioned units and crop out offshore below sea level or in estuaries and streams. The fossils may be derived from any one or all of these units. Although often reported from Jeremy Inlet and Edingville Beach, the fossils are found also at Botany Bay Island, Edisto Beach, and the southern tip of the island (Fig. 2). This widespread occurrence suggests that the depositional area was equally broad or that, once eroded, fossils are easily transported by currents to the widely separated points of discovery. The large number of terrestrial species present in the Edisto Island fauna and the probability of an offshore source area suggest that sea level at the time of deposition was lower than the present level. Sea level fluc- tuated during the Pleistocene and Holocene, in response to alternating glacial and interglacial stages (Cazeau et al. 1964; Hoyt 1967; Whitmore et al. 1967; Winkler and Howard 1977). Webb (1974) related these fluc- tuations of sea level to the North America mammal ages and glacial stages within the Pleistocene (Fig. 1). With substantially lowered sea level during Pleistocene glacial stages, what is now Edisto Island would have been a nearshore but inland area possibly exhibiting a savannahlike terrain; the deposition on Edisto Island was probably alluvial. Numerous Pleistocene streams and es- tuaries might have cut into underlying Pliocene and/or Miocene strata, deposited the Pleistocene vertebrate fossils, and possibly caused some mixing. Although evidence is not conclusive, the widespread recovery of fossil vertebrates along the southern Atlantic coast (Frey et al. 1975) suggests that the fossiliferous unit or units is extensive regionally. At Edisto Island, this unit probably crops out both in streams and estuaries on the island and in nearby offshore areas. 4 Janet A. Roth and Joshua Laerm Fig. 2. Map of Edisto Island, South Carolina, and adjacent coastal area. Insert, map of eastern United States indicating location of Edisto Island. MATERIALS AND METHODS In addition to examining and identifying previously collected museum materials from Edisto Island, fossils were collected on the island from June to August, 1978. Collecting was carried out by general surface survey of beach and stream banks and by screening of beach sand through 1/1 6-inch wire mesh. Museum abbreviations are as follows: ChM, Charleston Museum USNM, National Museum of Natural History, Smithsonian Institution HS, Hampden-Sydney College collections UGV, University of Georgia vertebrate fossil collections FAUNAL LIST The following faunal list includes all known mammalian and rep- tilian taxa from Edisto Island. Other fossil vertebrates are not included in this report. Class Reptilia Order Chelonia Family Chelydridae Chelydra serpentina Edisto Island Fossil Vertebrates 5 Family Kinosternidae Kinosternon sp. Family Emydidae Chrysemys scrip ta cf. pet ro lei Chrysemys floridana Terrapene Carolina putnami Family Testudinidae Geochelone sp. Gopherus sp. Family Trionychidae Trionyx sp. Order Squamata Suborder Serpentes Family Colubridae cf. Coluber sp., or Masticophis sp. cf. Elaphe sp. Order Crocodilia Family Crocodylidae Alligator mississippiensis cf. Gavialosuchus sp. Class Mammalia Order Edentata Family Dasypodidae Dasypus bellus Holmes ina septentrionalis Family Glyptodontidae Glyptotherium floridanum Familty Megalonychidae Megalonyx cf. jeffersonii Family Megatheriidae Eremotherium cf. mirabile Family Mylodontidae Glossotherium ( Paramylodon) cf. harlani Order Lagomorpha Family Leporidae Sylvilagus sp. Order Rodentia Family Castoridae Castoroides cf. ohioensis Castor canadensis Family Hydrochoeridae Neochoerus pinckneyi 6 Janet A. Roth and Joshua Laerm Order Cetacea Family Delphinidae Tursiops cf. truncatus Family Physeteridae Physeter sp. Family Balaenopteridae Order Carnivora Family Canidae Canis dims Urocyon cf. cinereoargenteus Family Procyonidae Procyon lotor Family Ursidae Tremarctos floridanus Family Odobenidae cf. Odobenus rosmarus Family Phocidae cf. Halichoems grypus Family Felidae Felis onca augusta Order Proboscidea Family Elephantidae Mammuthus cf. columbi Family Mammutidae Mammut americanum Order Sirenia Family Trichechidae Trichechus sp. Order Perissodactyla Family Equidae Equus sp. Family Tapiridae Tapims sp. Order Artiodactyla Family Tayassuidae Mylohyus cf. fossilis Family Camelidae Palaeolama cf. mirifica Family Cervidae Odocoileus virginianus Family Bovidae Bison cf. antiquus Edisto Island Fossil Vertebrates 7 SYSTEMATIC PALEONTOLOGY Many of the species discussed below occurred widely in North America during the Late Pleistocene. A complete listing of their oc- currences and distributions is beyond the scope of this paper. We will, in general, limit our comments to known distributional records in the Southeast. Descriptions of the geological ranges and comments regarding habitat preferences for the following species are provided in Table 1. Class Reptilia Order Chelonia Family Chelydridae Chelydra serpentina (Linnaeus) — snapping turtle Material. — ChM.GPV1528, ChM.GPV1530-1531, ChM.GPV1533- 1536, seven peripheral scutes; ChM.GPV1529, ChM.GPV1532, two costal scutes. Family Kinosternidae Kinosternon (Spix) sp. indet. — mud turtles Material. — ChM.GPV 1554, pygal bone. Family Emydidae Chrysemys scripta cf. petrolei (Leidy) — pond slider Material. — ChM.GPV2093, carapacial fragment. Remarks. — Chrysemys scripta petrolei is distinguished from Recent C. scripta by more rugose surficial sculpturing of the carapace and by its larger size (Weaver 1967). Chrysemys floridana (LeConte) — Florida cooter Material. — ChM.GPV622-623, two carapacial fragments. Chrysemys floridana or Chrysemys concinna (LeConte) — cooters Material. — ChM.GPV 1461-1463, ChM.GPV 1465- 1470, nine carapacial fragments. Remarks. — Both species range from the Pliocene to the Holocene and now occur in aquatic habitats in eastern coastal states. Additional Chrysemys material from Edisto Island has recently been reported elsewhere (Dobie and Jackson 1979) and is not considered here. Terrapene Carolina putnami (Hay) — giant box turtle Material. — ChM.GPV2094-2095, two carapacial fragments. Remarks. —Terrapene Carolina putnami is distinguished from living T. c. Carolina on the basis of greater size and increased curvature of the peripheral scutes. Fossil localities occur from Florida to New Mexico and at least as far north as Kansas. 8 Janet A. Roth and Joshua Laerm Table 1. Geologic ranges and habitat preferences for species occurring in Edisto Island fauna. SPECIES GEOLOGIC RANGE Chelydra serpentina Kinosternon Chrysemys scripta petrolei Chrysemys floridana Chrysemys concinna Terrapene Carolina putnami Geochelone sp. Gopherus sp. Coluber sp. or Masticophis sp. Elaphe sp. Alligator mississippiensis Gavialosuchus sp. Dasypus bellus Holmesina septentrionalis Glyptotherium floridanum Megalonyx jeffersonii Eremotherium mirabile Glossotherium harlani Syvilagus sp. Castoroides ohioensis Castor canadensis Neochoerus pinckneyi Tursiops truncatus Physeter Canis dims Urocyon cinereoargenteus Felis onca augusta Procyon lotor T remar do s floridanus Odobenus rosmams Halich9ems grypus Mammuthus columbi Mammut americanum Trichechus Equus Tapirus Bison antiquus Palaeolama mirifica Odocoileus virginianus Mylohyus fossilis Hemphillian to Holocene (Holman 1966) Pliocene to Holocene (Holman 1972) Rancholabrean (Weaver 1967) Pliocene to Holocene Pliocene to Holocene Middle Pliocene to late Wisconsinian (Milstead 1969) Late Eocene to Holocene (Hibbard and Taylor 1960) Miocene to Holocene (Romer 1966) Pleistocene to Holocene (Holman, pers. comm.) Miocene to Holocene (Romer 1966) Oligocene to Holocene (Romer 1966) Miocene to Pleistocene (Auffenberg 1954, 1957; Tadao 1969) Blancan to Wisconsinian (Robertson 1976; Ray 1967) Irvingtonian to Rancholabrean (Webb 1974) Rancholabrean (Gillette and Ray, in press) Blancan to Holocene (Hibbard et al. 1965) Rancholabrean (Hibbard et al. 1965) Pleistocene (Stock 1925; Schultz 1965) Kansan to Holocene (Hibbard et al. 1965) Late Kansan to late Wisconsinian (Semken 1966) Illinoian to Holocene (Semken 1966) Wisconsinian (Lance 1958) Pliocene to Holocene (Romer 1966) Miocene to Holocene (Romer 1966) Sangamonian to late Wisconsinian (Hibbard and Taylor 1960) Rancholbrean to Holocene (Stevens 1965) Aftonian to late Wisconsinian (Ray 1967; Kurten 1973) Pleistocene to Holocene (Arta and Hutchison 1964) Illinoian to Holocene (Kurten 1966) Nebraskan to Holocene (Ray et al. 1968) Pleistocene (Ray et al. 1968) Yarmouthian to late Wisconsinian (Miller 1971; Webb 1974) Throughout Pleistocene (Miller 1971) Pliocene to Holocene (Romer 1966) Irvingtonian to Holocene (Martin and Webb 1974) Blancan to Holocene (Hibbard et al. 1965) Late Sangamonian to late Wisconsinian (Robertson 1974) Irvingtonian to Rancholabrean (Lundeluis 1972; Webb 1974) Late Irvingtonian to Holocene (Martin and Webb 1974) Blancan to late Wisconsinian (Semken and Griggs 1965) Edisto Island Fossil Vertebrates 9 HABITAT PREFERENCE Any permanent body of fresh water (Conant 1975) Muddy bottom aquatic habitats (Conant 1975) Probable habitat in or near stream (Holman 1972) Coastal plain ponds, lakes, swamps, marshes, and rivers (Conant 1975) Predominantly streams but also coastal marshes (Conant 1975) Coastal savannah subspecies (Auffenberg 1967; Martin 1974) Tropical to humid subtropical habitats (Hibbard 1960; Loveridge and Williams 1957) Predominantly dry sandy soils (Conant 1975) Open woods or grassland usually near permanent water (Conant 1975) Terrestrial and arboreal usually near reliable water sources (Holman 1972) Tropical to subtropical aquatic habitats (Woodburne 1958; Conant 1975) Tropical to subtropical aquatic habitats (Auffenberg 1954, 1957) Variety of subtropical habitats (Slaughter 1959) Variety of subtropical habitats (Gillette, pers. comm.) Tropical to subtropical climate, lush vegetation, standing water (Gillette and Ray, in press) Warm, dense, moist forest (Stock 1925) Warm, dense, moist forest (Edmund, pers. comm.) Open grasslands (Stock 1925) Wide variety of forested habitats (Blair 1968) Marshes, bogs, lakes, and rivers (Cahn 1932) Marshes, bogs, lakes, and rivers (Blair 1968) Moist temperature climate across southern North America (Hay 1927; Simpson 1930) Atlantic Ocean, Massachusetts south to South America (Hall and Kelson 1959) Arctic to tropical waters of Atlantic (Hall and Kelson 1959) Presumably similar to modern Canis Variety of woodland habitats (Blair 1968) Densely wooded areas, frequenting streams (Kurten 1973) Forested areas, frequenting streams and marshes (Hall and Kelson 1959) Variety of wooded habitats (Kurten 1966) Atlantic coastal area (Hall and Kelson 1959) North Atlantic coastal areas, migrating into inlets and estuaries (Hall and Kelson 1959) Open grasslands and savannah, with permanent water (Hibbard 1955) Coniferous forests, probably boreal (Martin and Guilday 1967; Dremanis 1968; Voorhies 1974) Atlantic coastal regions, frequenting bays and inlets (Hall and Kelson 1959) Open grasslands, presumably similar to modern Equus Predominantly humid tropical to subtropical forests (Simpson 1945; Gray and Cramer 1961) Open grasslands, presumably similar to modern Bison Grasslands and savannahs (Webb, pers. comm.) Woodlands and forest edges Warm moist woodlands (Martin and Guilday 1967) 10 Janet A. Roth and Joshua Laerm Family Testudinidae Geochelone sp. (Fitzinger) — giant land tortoise Material. — ChM.PV2475, ChM.PV2486-2487, three carapacial frag- ments; ChM.GPV2005, ChM.PV2467-2476, ChM.PV2476, ChM.PV2478, ChM.PV2479-2485, ChM.PV2488-2492, twenty-one plastral fragments; ChM.PV2457-2462, ChM.PV2464, ChM.PV2466, eight peripheral scutes; ChM.PV2463, ChM.PV2477, two nuchal sctues; ChM.PV2465, costal scute; ChM.PV2497, osteoderm. Remarks.— Geochelone is restricted to primarily subtropical and tropical climates (Hibbard and Taylor 1960). Previous reports of Geochelone include localities in Florida and several midwestern states (Hibbard 1960). Gopherus sp, (Rafinesque) — gopher tortoise Material. — ChM.GPV1537, carapacial fragment. Order Squamata Suborder Serpentes Family Colubridae cf. Coluber sp. Linnaeus or Masticophis sp. (Baird and Girard) — racers or whipsnakes Material. — ChM.GPV2088, vertebral fragment. Remarks. — The specimen is missing important processes, which precludes generic identification. cf. Elaphe sp. (Fitzinger) — rat snake Material. — ChM.PV2746, vertebral fragment. Remarks. — The specimen is severely worn, making determination of species impossible. Order Crocodilia Family Crocodylidae Alligator mississippiensis (Daudlin) — alligator Material. — ChM.PV2493, right mandible; ChM.PV2494, right man- dibular fragments; ChM.PV2501, mandibular fragment; ChM.PV2020, tooth fragment; ChM.PV2502, right dermal roof fragment; ChM.PV2503, left dermal roof fragment; ChM.PV2505, dermal roof fragment; ChM.PV2735, worn vertebral centrum; ChM.PV2496, ChM.PV2500, ChM.PV2752, three dermal scutes; UGV132a-c, dermal scutes; UGV136, skull fragment; UGV142, vertebra. Remarks. — All specimens are very dark and well mineralized. The der- mal scutes exhibit the diagnostic keel. cf. Gavialosuchus (Toula and Kail) — giant crocodile Material. — ChM.PV2495, right humerus; ChM.PV2504, right humerus; ChM.35.208. 176, dorsal plate. Edisto Island Fossil Vertebrates 11 Remarks. — The two humeri are large, elongate, and do not cpmpare well with available material of Alligator or Crocodylus acutus. The specimens are referred to Gavialosuchus; however, the limited knowledge of Gavialosuchus postcranial material prevents conclusive determination. Gavialosuchus has been reported from the late Miocene to early Pliocene of Florida (Sellards 1915; Mook 1921; Auffenberg 1954, 1957). Auffenberg (1957) also noted two occurrences of Gavialosuchus from South Carolina: a partial skull (ChM. 13745) from near Lambs, Charleston County, and one dorsal plate (ChM. 35. 208. 176) from Edisto Beach. He suggested that the Edisto Island specimen is probably of Miocene age. The two specimens reported here may also represent Miocene or Pliocene components of a predominantly Pleistocene assem- blage; or, it is possible the genus had a more extensive geologic range than has previously been reported. In the light of present knowledge of the geology of the region between Edisto Island and Charleston, a Miocene age for these specimens seems unlikely. Richards et al. (1971) reported Late Pleistocene mollusks from depths of 40 to 50 feet below the surface on Wadmalaw Island, John’s Island, and James Island and con- cluded that they were of Sangamonian age. The Sangamonian deposits unconformably overlie the Oligocene member of the Cooper Formation. Offshore they are overlain by sediments deposited during regression in Wisconsinian time. Thus, there is no immediate source for fossils of Miocene age in the vicinity of Edisto Island. Reworked Miocene fossils may well occur at the base of the Pleistocene in this area as they do at cer- tain places farther inland, but it is difficult to see how such material could be reworked upward through more than 50 feet of sediment. A gavialosuchid recorded from the Pleistocene of Japan (Tadao 1969) lends some credence to the possiblity that the Edisto specimen is Pleistocene in age. Class Mammalia Order Edentata Family Dasypodidae Dasypus bellus (Simpson) — beautiful armadillo Material. — ChM.PV2705, dermal scute. Remarks. — Dasypus bellus has been reported from as far west as Texas (Slaughter 1959) and north to Missouri, Tennessee and West Virginia (Guilday and McCrady 1966; Guilday et al. 1978). Localities include Ladds Quarry in northwest Georgia (Ray 1967). The earliest report of D. bellus is from the Blancan of Florida (Robertson 1976), but the species does not appear in the fossil record elsewhere until the Wisconsinian. See Table 2 for measurements. 12 Janet A. Roth and Joshua Laerm Table 2. Comparative measurements (mm) of Dasypus bellus dermal scutes from Edisto Island and elsewhere. ChM.PV2705,Edisto Island ChM.PV2092, Edisto Island D. novemcinctus, Recent (Martin 1974) D. bellus, Florida (Martin 1974) Thickness Width N 4.9 13.0 1 5.0 12.0 1 0.9 -2.4 3.5 -8.1 235 1.9 -6.1 6.2 - 16.5 194 cf. Holmesina ( Chylamytherium ) septentrionalis (Leidy) — giant armadillo Material. — ChM .GPV2032, ChM.PV2785-86, ChM.PV2416, ChM.PV2619-2620, carapacial scutes; ChM.PV2420 caudal ring scute. Remarks. — Chlamytheres apparently reached Flordia earlier than elsewhere in the U. S. (Webb 1974) and are known from several localities there (Robertson 1976). The Charleston specimens are referred to H. sep- tentrionalis, known from Irvingtonian and Rancholabrean deposits (Webb 1974). The group is presently under review by Dr. G. Edmund. Family Glypotodontidae Glyptotherium floridanum (Simpson) — North America glyptodont Material. — ChM.PV2415, cranial fragment; ChM.GPV2090, ChM.PV24 17-24 18, carapacial scutes. Remarks. — The cranial fragment was described by Ray (1965) and identified as Boreostracon floridanus (Simpson), but Gillette and Ray (in press) refer all glyptodont material from Edisto Island to G. floridanum. This species is confined to the Rancholabrean. Geographically, G. floridanum is restricted to the Gulf coastal plain and the southern Atlan- tic coast, and the South Carolina specimen provides the northernmost record of the species. Family Megalonychidae Magalonyx cf. M. jeffersonii (Desmarest) — Jefferson’s ground sloth Material.— ChM.PV2421, right lateral mandibular fragment, dentition not preserved; ChM.PV2423, caninform tooth; ChM.PV2743, caniniform tooth; ChM.PV2455 and ChM.GPV2012, two molariform teeth; ChM.PV2424, ungual phalanx; ChM.PV2428, cast of ungual phalanx; HS.A-48, HS.A-29, FIS.A-51, three caniniform teeth; HS.H-6 1 , astragalus; from the private collection of M. W. Wagner, two ungual phalanges; UGV150, RM1. Remarks.— Ray et al. (1967) noted that the number of species of Megalonyx and their validity are uncertain. The Edisto Island specimens compare closely with material presently identified as M. jeffersonii. The geographic distribution of Megalonyx seems to have been widespread Edisto Island Fossil Vertebrates 13 over North America, and in the Southeast includes previously reported occurrences in South Carolina (Hay 1923), Georgia (Ray 1967), Florida (Webb 1974) and Tennessee (Guilday et al. 1969). Family Megatheriidae Eremotherium cf. E. mirabile (Leidy) — giant ground sloth Material. — ChM.PV2426, ungual phalanx; ChM.PV2454, molar frag- ment; ChM.PV2400, lumbar vertebral fragment. Remarks. — Eremotherium ranged from southern South America to New Jersey (but see Ray 1979) and has been reported from South Carolina and Georgia (A. Sanders, pers. comm.; Frey et al. 1975), and Florida west to Texas (Webb 1974; Lundelius 1972). Family Mylodontidae Glossotherium ( Paramylodon) cf P. harlani (Owen) — Harlan’s ground sloth Material. — Ch M . PV2422, third lower molariform tooth; ChM.PV2427, molariform fragment; ChM.PV2741, distal fragment of humerus; ChM.PV2429, ungual phalanx; ChM.GPV2004, metapodial; USNM22842, ungual phalanx. Remarks.— The large ground sloth, Glossotherium, is known from the Pleistocene of North America. Its geographic distribution ranges from Central America to the Pacific and Atlantic coasts of the United States (Schultz 1965). Order Edentata, gen. et sp. indet. Two specimens, ChM.PV2739 and ChM.PV2740, both basal frag- ments of the neural spine, are comparable to ground sloth material but are otherwise indeterminate. Order Lagomorpha Family Leporidae Sylvilagus sp. (Gray) — rabbit Material. — ChM.PV2441, right mandible with P3, M,, and M2; ChM.PV2442, molar fragment; ChM.PV2498, molar fragment; ChM.PV2738, portion of right tibia. Remarks.— Sylvilagus is a relatively common and widespread con- stituent of North American fossil faunas beginning in the Late Pleistocene (Schultz 1965). Although the genus is an expected member of a Pleistocene fauna, the possibility that the specimens at Edisto Island represent a more recent occurrence of Sylvilagus on the island must be considered. Due to the apparent transport at Edisto Island the presence of Sylvilagus and other small mammals is a rare occurrence. Order Rodentia Family Castoridae Castoroides cf. C. ohioensis (Foster) — giant beaver Material.— ChM.PV2437-2438, two incisor fragments. 14 Janet A. Roth and Joshua Laerm Castoroides ohioensis localities occur throughout the eastern two thirds of North America, and are concentrated in the area south of the Great Lakes (Dallman 1969). The species has also been reported in the Southeast, including Georgia, South Carolina (Hay 1923), Tennessee (Parmalee et al. 1976) and Florida (Martin 1969). Castor canadensis (Kuhl) — beaver Material.— HS. 104, right femur. Remarks.— Although the species is a common constituent of fossil faunas, this specimen may be modern. Family Hydrochoeridae Neochoerus pinckneyi (Hay) — giant capybara Material. — Ch M . PV 2439, RM3; ChM.PV2440, RI1. Remarks.— This giant capybara was described by Hay (1923) as Hydrodchoerus pinckneyi on the basis of a left upper third molar from the Charleston area with which ChM.PV2439 closely compares. In 1927 Hay erected Neochoerus for the species. It is morphologically similar to the living Hydrochoerus, but of larger size. Capybara remains are common in Florida (Simpson 1930; Webb 1974). Order Cetacea Suborder Odontoceti Family Delphinidae Tursiops cf. T. truncatus (Montague) — Atlantic bottlenose dolphin Material.— ChM.PV2434, tooth fragment; ChM.PV2430, vertebra; HS.E-34, vertebra. Family Physeteridae Physeter sp. (Linnaeus) — sperm whale Material. — ChM.PV2436, tooth fragment. Suborder Mysticeti Family Balaenopteridae, gen. et sp. indet. A vertebral fragment (ChM.PV2431) and an auditory bulla (ChM.PV2433) represent balaenopterids of indeterminate genus or genera. Order Cetacea, gen. et sp. indet. Three vertebral fragments (ChM.PV23 18-2320) and two auditory bullae (ChM.PV2435 and HS.A-90) are of cetacean origin, but generic identifications remain to be determined. Order Carnivora Family Canidae Canis dirus (Leidy) — dire wolf Material. — ChM.PV2282, right mandibular fragment with broken Edisto Island Fossil Vertebrates 15 base of C and P2_4, M, preserved; HS.EB-22, LP4 with fragment of left maxilla attached. Remarks. — Fossil remains of Canis dims and Canis lupus are often con- fused; however, both Merriam (1912) and Martin (1974) concluded that the two species are distinct. Measurements from the Edisto specimens compare well with measurements provided by Martin (1974), Gillette (1979) and Nowak (1979), but are larger than those provided by Ander- son (1968). Canis dims was widely distributed in North America in the Pleistocene and occurred commonly in the Southeast (Guilday et al. 1969; Webb 1974; Gillette 1979). Urocyon cf. U. cinereoargenteus (Schreber) gray fox Material. — ChM.PV2736, left ulnar fragment. Remarks. — Although U. cinereoargenteus is an acceptable member of the Pleistocene fauna, this specimen could be Recent. The moderate but not strong degree of mineralization makes age determination of the specimen questionable. Reports of gray fox from the southeastern Pleistocene include several in Florida (Gut and Ray 1963; Arata 1959; Martin and Webb 1974) and one in Georgia (Ray 1967). It would be an expected member of a late Pleistocene fauna. Family Procyonidae Procyon lotor (Linnaeus) — raccoon Material. — ChM.PV2280-228 1, two mandibular fragments; ChM.PV2282, right mandibular fragment; ChM.PC2737, left man- dibular fragment; ChM.PV2747, right femur; HS.A-34, right mandibular fragment; UGV161, portion of left mandible. Remarks. — Arata and Hutchinson (1964) examined known Procyon material and determined that the Pleistocene species are distinguished only on the basis of variation in size. This variation is similar to the geographic variation in size found in modern P. lotor. They thus con- cluded that known specimens of Pleistocene Procyon are conspecific with P. lotor. The Edisto Island material compares closely with modern Procyon. Fossil P. lotor has been reported from various localities throughout eastern and central North America, including numerous records from Florida and at least one from the Ashley River, South Carolina (Hay 1923). Family Ursidae Tremarctos floridanus (Gidley) — Florida spectacled bear Material.— ChM.GPV2019, cast of right mandibular ramus with M2 (original in private collection of Charles Harshaw, Charleston). Remarks. Dental measurements of the specimen fall well within the range of those recorded by Kurten (1966) and Ray (1967). Tremarctos floridanus has been reported from several southeastern localities, in- 16 Janet A. Roth and Joshua Laerm eluding Tennessee (Guilday and Irving 1967), northwestern Georgia (Ray 1967), and Florida (Kurten 1966). Family Odobenidae cf. Odobenus rosmarus (Linnaeus) — walrus Material. —Cranial fragment (HS). Remarks. — This specimen has been tentatively referred to Odobenus rosmarus (Ray et al. 1968). Family Phocidae cf. Halichoerus grypus (Fabricius) — gray seal Material.— ChM.PV2283, partial right innominate. Remarks.— This specimen has been tentatively referred to Halichoerus grypus (Ray et al. 1968). Family Felidae Felis onca augusta (Leidy) — jaguar Material. — ChM.PV2284, right P4. Remarks.— This specimen is the first known occurrence of the species in South Carolina (Ray 1967; Kurten 1973). Fossil F. onca occurs throughout much of the conterminous United States, including localities in Florida (Webb 1974), Tennessee (Guilday et al. 1978), and one in northwestern Georgia (Ray 1967). Order Proboscidea Family Elephantidae Mammuthus cf. M. columbi (Falconer) — Columbian mammoth Material.— ChM.PV2291, right lower molar fragment; ChM.PV2287, left upper molar in two fragments; ChM.PV2288, two molar fragments; ChM.PV2289, lower right molar fragment; ChM.PV2292-2293, two molar fragments; ChM.PV2305, cuboid; ChM.PV2290, molar fragment; ChM.PV2285, right third metacarpal; ChM.PV2286, third phalanx, digit III; HS.101, right lower molar; HS.E-46, molar fragment; HS.102, deciduous molar fragment; HS.102, deciduous molar fragment; UGV149, anterior half of probable RM3; UGV135a-b, tooth fragments. Remarks.— Osborn (1942) described 10 genera and 59 species of elephants. Although Osborn’s criteria for distinguishing species are still used, most authors have identified only 2 genera and 5 to 6 species as valid. Aguirre (1969) regarded two species, M. imperator and M. columbi, as valid in North America, but Miller (1971) regarded even these as in- distinct. Because M. columbi has priority, the Edisto Island material is assigned to that species. The Columbian mammoth is known from numerous localities in the Southeast, including Georgia, South Carolina, and Florida (Hay 1923; Webb 1974). Edisto Island Fossil Vertebrates 17 Family Mammutidae Mammut americanum (Kerr) — American mastodon Material. — ChM.PV2294-2295, two RM3’s; ChM.PV2286, RM3 frag- ment; ChM.PV2297-2298, molar fragments; ChM.PV2742, unerupted deciduous cheek tooth; ChM.PV2299, right mandible; HS.E-45, deciduous cheek tooth fragment; HS.E-44, cheek tooth fragment; HS.103, molar fragment; UGV148, RM2; UGV133a-c, tooth fragments; UGV153, section of tusk. Remarks. —Although Osborn (1936) recorded numerous species of the genus Mammut, only M. americanum is now generally regarded as valid in the North American Pleistocene (Miller 1971). Order Proboscidea, gen. et sp. indet. Material. — ChM.PV2300, ChM.PV2307, four tusk fragments; ChM.PV2303, skull fragment; ChM.PV2306, scapular fragment; ChM.PV2733, vertebral fragment; HS.A-117, HS.A-121, HS.A-27, ver- tebral fragments; HS.E-26A, HS.A-86, fragmentary metapodials; HS.E- 67, phalanx. Order Sirenia Family Trichechidae Trichechus (Linnaeus) sp. indet. — manatee Material. -ChM.GPV627, rib fragment; ChM.PV2315, ChM.PV2317, ChM.PV2453, ChM.PV2321, four rib fragments; ChM.PV2748, thoracic vertebral fragment; HS.E-75, rib fragment; HS.A-33, right mandible. Remarks. — Trichechus is totally aquatic and prefers sheltered bays or sluggish rivers. They are known to wander far inland in Florida rivers. Simpson (1932) indicated that Pleistocene sirenians commonly occur in inland fresh water deposits of Florida. Pleistocene Trichechus remains are known along tthe Atlantic coast from Maryland to Florida (Simpson 1932). Order Perissodactyla Family Equidae Equus (Linnaeus) sp. indet. — horse Atoernz/.— ChM.PV2331, ChM.PV2449, two LP,; ChM.PV2332, LP2; ChM.PV2330, LP3; ChM.PV2333, LM,; ChM.PV2334-2335, two LM2; ChM . PV23 3 6-2337, two RP2; ChM.PV2338, ChM.PV2448, ChM.PV2451, three RP3; ChM.PV2339; ChM.PV2340;, ChM.PV2360- 2361, ChM.PV2447, four LI2; ChM.PV2341, LP1; ChM.PV2342-2343, two LP3; ChM.PV2446, LM1; ChM.PV2344; LM2; ChM.PV2348, LM3; ChM.PV2345-2346, two RP3; ChM.PV2347, ChM.PV2445, two RM1; ChM.PV2450, lower molar fragment; ChM.PV2353-2355, three upper molar fragments; ChM.PV2359, molar fragment; ChM.PV2006, premolar fragment; ChM.PV2349-2352, four upper cheek tooth frag- 18 Janet A. Roth and Joshua Laerm ments; ChM.PV2357-2358, ChM.PV2452, three lower cheek tooth frag- ments; HS.E-33A, HS.A-82, two right ulnae; HS.E-27, distal epiphyses of two radii, ChM.PV2321, right tibia; ChM.PV2413, left tibia and portion of shaft; ChM.PV2329, right pelvic fragment, including acetabulum; ChM.PV2322, right astragalus; ChM.PV2745, calcaneal fragment; ChM.PV2372, 2328, ChM.PV2444, three metapodial fragments; ChM.PV2326, fragment of proximal part of metapodial; ChM.GPV1978, metapodial; HS.E-25, two phalanges; ChM.PV2323-2325, ChM.PV2443, four phalanges; UGV1 15, LM1 or LM2; UGV1 16, RP3 or RP4; UGV1 17, RM1, UGV118, LP2; UGV119, RP3 or RP4; UGV124, buccal section of LM1; UGV125, RP3 or RP4; UGV126, lingual section of RM1 UGV127, LP3 or LP4; UGV134, LM1 or LM2; UGV113, RM3; UGV114, LP2; UGV120, RM, or RM,; UGV121, LM, UGV122, LM1, or LM2; UGV123, LM3; UGV122, LM, or LM2; UGV123, LM3; UGV128, LM, or LM2; UGV129, LP3 or LP4; UGV130, RM, or RM2; UGV160, glenoid region of scapula. Remarks. — The possibility that some of the equid remains are of Re- cent origin must be considered since E. caballus is a modern inhabitant of Edisto Island and permineralization is known to take place at a com- paratively rapid rate on the island (O’Kelley 1976). Although the anti- quity of the Equus material cannot be proven absolutely at this time, three factors indicate that at least most of the remains are Pleistocene. First, the very dark color and strong permineralization of some of the fossils is comparable to that of material from species that definitely became extinct during the Pleistocene. Second, the relative abundance of horse material in the fauna would suggest numerous individuals occur- ring over a long period of time. Finally, Equus is an expected member of the Pleistocene fauna at Edisto Island. Family Tapiridae Tapirus Brisson sp. indet. — tapir Material. — ChM.PV2362, symphyseal fragment of mandible with fragmentary root of RI2; ChM.PV2363, right upper molar fragment; ChM.PV2364, deciduous RM1; ChM.PV2365, upper molar fragment; HS.E-68, carpal element; HS.Q, right upper molar fragment; HS.R, right molar; UGV150, LM,; from the private collection of W. M. Wagner, left mandible with P3-M3 present. Remarks.— The incomplete nature of most of the specimens prevents specific determination. Order Artiodactyla Family Tayassuidae Mylohyus cf. M. fossilis (Leidy) — long nosed peccary Material. — ChM.PV2414, symphyseal mandibular fragment with roots of RI,, RI2, RC, LI,, IC; ChM.PV2499, LM,; from the private Edisto Island Fossil Vertebrates 19 collection of Mrs. Eddie Gue, RP3 and one molar fragment. Remarks.— The limited available Mylohyus material prevents definite specific determination. Lundelius (1960) recognized Mylohyus nasutus and M. fossilis, based on differences in size and geographic distribution. Mylohyus fossilis is characterized as a smaller species from eastern North America, while M. nasutus is a larger species generally occurring west of the Appalachian Mountains. Ray (1967) did not support Lundelius’ con- clusions but suggested instead that the variation in size between supposed species of Mylohyus is the result of sexual variation within one species. The Edisto Island material compares well with material presently iden- tified as M. fossilis. Fossil localities include the Ashley River in South Carolina (Allen 1926), Ladds Quarry, Georgia (Ray 1967), and numerous sites in Florida (Webb 1974). Family Camelidae Palaeolama cf. P. mirifica (Simpson) — Pleistocene llama Material. — ChM.PV2411, right distal fragment of humerus; ChM.PV2409, right radius-ulna and associated carpals; ChM.PV2410, metapodial shaft fragment; ChM.PV2407, phalanx. Remarks.— The metapodial fragment and phalanx conform closely in size and morphology to Palaeolama mirifica from the Coleman II locality in Florida. But the two limb elements (ChM.PV2411, ChM.PV2409) seem larger and more massive than comparative P. mirifica material. Measurements taken from the radius-ulna are comparable to those recorded by Webb (1974) for P. mirifica (Table 3). The species has been reported from Florida (Webb 1974), Texas (Lundelius 1972), and the coast of Georgia (Frey et al. 1975). The record of Palaeolama at Edisto Island extends its geographic distribution northward. Table 3. Comparative measurements (mm) of Palaeolama cf. mirifica. ChM.PV 2409 Webb (1974) Overall length 508 430-451 Length, olecranon 83 Articular length 428 Width, proximal articular surface 75 Width, midlength - transverse 52 Anteroposterior 36 Width, distal end 77 20 Janet A. Roth and Joshua Laerm Family Cervidae Odocoileus virginianus (Boddaert) — white tailed deer Material. — ChM . PV 2403-2404, antler bases; ChM.PV2405, antler fragment; ChM.PV2456, antler fragment, ChM.PV2749, RM2 fragment; ChM.PV2750, fragment of right humerus; ChM.PV2406, left radius, HS.A-56, antler tine; HS.E-29, anterior extremity of left mandible; HS.A-58, metacarpal fragment; UGV153, section of right tibia. Remarks. — All material is identical to modern Odocoileus virginianus, but the specimens are dark and well mineralized. Although O. virginianus is a common species in the middle to late Pleistocene, the possibility that this material is of Recent origin must be considered since the species is presently a common inhabitant of Edisto Island. Family Bovidae Bison cf. B. antiquus (Leidy) — Pleistocene bison Material. — ChM.PV2393, horn core, preserved from basal constric- tion to core tip; ChM.PV2394, horn core fragment; HS.E-77, horn core fragment; ChM.PV2392, horn core fragment; ChM.PV2389, thoracic vertebra. Remarks.— Edisto bison material is generally robust and strongly per- mineralized. Measurements from horn cores (Table 4) compare favorably with Bison antiquus (Skinner and Kaisen 1947; Robertson 1974). The remaining Bison material lacks diagnostic features and cannot with cer- tainty be referred to species. Bison sp. — bison Material. — Ch M . PV 2376, LP3; ChM.PV2377, LM,; ChM.PV2304, left frontal fragment; ChM.PV2408, right mandible; ChM.PV2366, right mandible with P3 and M,; ChM.PV2374, right mandible with M2 and M3; ChM.PV2375, left mandibular fragment with roots of M, and M2; ChM.PV2372, left anterior mandibular fragment; ChM.PV2372, RM2; ChM.PV2, atlas vertebra; ChM.PV2397, axis vertebra; ChM.PV2388, sacral vertebra; ChM.PVl, sacral vertebra; ChM.PV2390, cervical ver- tebra; ChM.PV2391, cervical vertebra; ChM.PV2395-2396, two left tibiae; ChM.PV2398, vertebral fragment; ChM.PV2399, vertebral cen- trum; ChM.PV2372-2373, proximal fragment of right scapula; ChM.PV2367, right scapular fragment; ChM.PV2368, left scapular frag- ment, glenoid cavity preserved; ChM.PV2369, proximal fragment of humerus; ChM.PV2370, right humerus; ChM.PV2371, distal fragment of right humerus; ChM.PV2378, shaft of right radius; ChM.PV2379, left radius in two fragments; ChM.PV2380, distal fragment of left radius, ChM.PV2401, partial right innominate; ChM.PV2381, distal fragment of left femur; ChM.PV2382, distal fragment of right femur; ChM.PV2383, proximal fragment of left tibia; ChM.PV2384, right calcaneum; ChM.PV2734, calcaneal fragment; ChM.PV2385, proximal fragment of Table 4. Comparative measurements (mm) of Bison antiquus horn cores, Edisto Island and other localities. Edisto Island Fossil Vertebrates 21 X NT) 03 ON 2 r~~ c < (N <3 °Q d O . NO cn) — rf re oo rc m r~- — no m m c-i m m m X NT) oo o3 ON tn §2 m m m 3 oi NO O 3 > oo m OO re co < ' — 1 CN) Cn) oci . e o O OO r- o 2 Cn) Cfi X r-~ r-~ UJ r-~ on) m ■ — 1 > Cu ^ _ ON 2 U o r~~ m O o- > cu U m ON m C4 Cn) O m NO NO <~C m m r— ■'1- ON v- a. u \J3 !_ a> t— £ ^ u. . i— E JZ D SZ 3 0£) -0 3 <3 S > s g 1 u- 3 2 g CD -j Ui rr o u. o ° o ° o U U U c 4) ^ s 1 73 55 ON TJ W C c Ki o 1— C/3 4) *—* 1— c 4) c JO lx C/5 o (N 22 Janet A. Roth and Joshua Laerm metapodial; ChM.PV2386, proximal fragment of right tibia; HS.A-31, posterior part of right mandible with basal fragment of RM3; HS.A-105, anterior portion of right mandibular ramus with P, , P3, M,, and M2; HS.A-30, anterior portion of left mandibular ramus without dentition; HS.E-57, proximal portion of scapula; HS.A-119, distal fragment of humerus; HS.A-108, left radius; HS.E-74, olecranon of ulna; HS.E-70, postzygapophysis of vertebra; HS.A-105, fragment of right ilium with acetabulum; HS.E-55, proximal epiphysis of tibia; HS.A-59, distal epiphysis of left tibia; HS.E-31, distal fragment of fibula; HS.A-60, left calcaneum; HS.E-60, astragalus; HS.A-78, left cubonavicular; HS.A-77, right cubonavicular; HS.E-12, fragment of proximal part of fused metatarsal; HS.A-57, metacarpal fragment; HS.A-114, right metatarsal; HS.E-64, HS.E-67, phalanges; HS.A-63, HS.A-65, HS.A-66, phalanges; UGV146, lower molars; UGV147, right astragalus; UGV151, right femur. PALEOECOLOGY In general, knowledge of the paleoecology of an area may be derived from the fossil-bearing sediments, the fossils themselves and palynological studies of the fossiliferous strata. At Edisto Island, only the vertebrates (and only their larger, more durable parts) are available for consideration. As a result data are highly limited and biased. However, some knowledge of the paleoecology pan be gained from the recovered part of the vertebrate record. The Edisto Island fauna can be divided into at least four general communities, representing broad habitats in the depositional area: Marine community.— This community consists of predominantly wide ranging marine species that frequent nearshore areas and oc- casionally occur in rivers and bays. The crocodile ( Gavialosuchus ), the cetaceans ( Tursiops , Physeter, and balaenopterids), the gray seal {Halichoerus), the walrus ( Odobenus ), and the manatee ( Trichechus ), comprise this community. Halichoerus, Trichechus, and Tursiops may migrate inland by means of estuaries and consequently be found in association with a primarily terrestrial fauna. But the large size of Physeter and the balaenopterid whales precludes any major inland migra- tions and indicates that deposition of remains of these two taxa must have occurred very near, if not in, a marine environment. Semi-aquatic terrestrial community. — This community relies upon a permanent source of fresh water such as a stream, marsh, or pond, but some members of the community may spend part or most of their lives beyond the limits of water. Included here are the beavers ( Castor and probably Castoroides ), the capybara ( Neochoerus ), the turtles ( Chelydra , Chrysemys, and Kinosternon ), Alligator, and possibly the glyptodont ( Glyptotherium ). Most of these genera also require substantial amounts of vegetation in or near the water, both for food and for shelter. In par- ticular, the beavers would depend on brush or standing timber in the Edisto Island Fossil Vertebrates 23 area, and alligators would require dense vegetation along stream banks or marsh edges for nest building. Forest community .—The, forest community includes species that de- pend on forest vegetation and food and/or refuge and spend most of their time in forested areas. The ground sloth ( Megalonyx ), mastodon {Mammut), tapir ( Tapirus ), peccary ( Mylohyus ), and sloth (. Eremotherium ) are all considered to be browsers, preferring dense vegetation. Felis, Odocoileus, and Sylvilagus also prefer the forest habitat. The turtles, Gopherus and Terrapene, might also frequent forested areas. Savannah community. — The savannah community is well represented in the fossil fauna and mostly consists of grazers, which rely on tall grasses and scrubby vegetation or open grasslands. Included are the glyp- todont ( Glyptotherium ), ground sloth {Par amyl odon), mammoth {Mammuthus), horse {Equus), buffalo {Bison), and llama ( Palaeolama ). Terrapene, Geochelone, and Gopherus also occur in savannah type vegeta- tion. Although most species have been assigned to a particular com- munity, many would regularly visit two or more of the described habitats. For example, the raccoon ( Procyon ) would be expected to fre- quent stream banks and marshes and to den in a wooded area, but an in- dividual might also wander through open grasslands. The rabbit { Sylvilagus ), wolf ( Canis ), fox ( Urocyon ), and deer ( Odocoileus ) also com- monly move from one habitat to another. The probable common ground for all species concerned is the water- ing place. Thus, the fauna gives evidence of a stream or marsh environ- ment bordered by open savannah and scattered wooded areas. The presence of certain species in the fauna indicates particular climatic restrictions. The most definite climatic limitation is imposed by the presence of Alligator and Geochelone in the fauna. These occur in tropical to subtropical climates in which temperatures rarely, if ever, fall below freezing and where precipitation is usually plentiful. Thus, at the time Alligator and Geochelone lived in the vicinity of Edisto Island the climate may have been more equable than the present climate of the area. Other genera that are considered to be restricted to warm, moist climates include Chlamytherium, Glyptotherium, Neochoerus, Dasypus, and Mylohyus. Several genera suggest a substantially cooler climate. These include; Mammut, Castoroides, Halichoerus, and Odobenus. The status of the mastodon with respect to climate is uncertain (Martin and Guilday 1967; Dreimanis 1968; Voorhies 1974); however, their ecological tolerances might reasonably be expected to be broad. Castoroides is most commonly found in the Great Lakes region, thus ranging into distinctly northern climates. But the genus also has been reported from southern localities, and no southern climate restrictions have been determined. Odobenus and Halichoerus are thought of as northern species, but their occurrence 24 Janet A. Roth and Joshua Laerm at Edisto may indicate individuals wandering beyond the normal range or a more extended range during the Pleistocene, possibly as a result of a cooler climate. In the absence of any stratigraphic control there is no evidence that the Edisto fauna is unified. On the contrary, it appears almost certainly heterochronous. Thus, inferences regarding the paleoclimatic conditions cannot be made with any degree of certainty. In general, however, the assemblage appears to reflect a predominantly warm, moist climate similar to or warmer than that of today. AGE AND RELATIONS OF THE EDISTO VERTEBRATES The age of the Edisto Island fauna may be established only on the basis of previously recorded biostratigraphic occurrence of the species present. The most definitive species are Neochoerus pinckneyi (restricted to the Wisconsin), Bison cf. antiquus, Dasypus bellus, Glyptotherium floridanus, and Canis dims, which are Sangamonian to Wisconsinian in age. Chrysemys scripta petrolei also is limited to the Sangamonian and Wisconsinian. Although the total span of deposition at Edisto Island cer- tainly may extend well beyond these temporal limits, much of the deposi- tion probably occurred during the Rancholabrean. The problem of a mixed fauna cannot wholly be resolved. Because of apparent rapid mineralization at Edisto Island, the separation of Pleistocene and Holocene material remains in question. Neill (1957) noted that rapid permineralization may create the illusion that Recent material is of an older age, and cautioned against interpreting all per- mineralized material as fossils. O’Kelley (1976) dated mammalian fossils, which were eroding out of a marsh clay deposit on the beach and ex- hibited the characteristic dark color and degree of mineralization of “fossils,” but found them to be of Holocene age. At this point only a partial solution to the problem may be offered. The Edisto Island fauna is composed primarily of species limited to the Pleistocene. Extant species in the fauna are included in the Pleistocene fauna recorded here and are acceptable Pleistocene species, but they must remain suspect as possibly representing a mixture of Holocene and Pleistocene material. Furthermore, the late Tertiary age for Gav- ialosuchus cannot be discounted. Failure to recover microvertebrates, even after extensive screening, suggests that the absence of the more fragile groups such as small mam- mals, amphibians, birds and some reptiles is due to transport or possible surf grinding. However, larger vertebrates are well represented. Of a possible 1 1 mammalian orders, 9 appear in the faunal list. With the exception of small vertebrates, which are absent from the fauna, the Edisto Island species list is similar to those of several coastal Florida sites of Rancholabrean age. In particular, Edisto Island and Edisto Island Fossil Vertebrates 25 Seminole Field (Simpson 1929b) share 19 mammalian genera; Edisto Island and Melbourne (Ray 1958) share 20. Of the 14 vertebrate species recovered from the Georgia estuaries (Frey et al. 1975), 9 appear at Edisto Island. Although the Edisto Island fauna is in some repects similar to the Ladds Quarry fauna (Ray 1967), Edisto Island lacks the species associated with a colder climate and upland terrain (e.g., Maries pen- nanti, Sylvilagus transitionalis, and large sciurids). However, Edisto Island has several savannah dwelling species (e.g., Palaeolama, Glyptotherium) that are not present at Ladds Quarry, in addition to the marine taxa. The Edisto Island records of Glyptotherium, Palaeolama, and Geochelone constitute northern range extensions for these genera in eastern North America. In fact, the Charleston region of South Carolina represents the approximate northern limit of the range of other Late Pleistocene taxa including Tapirus, Mylohyus, and Neochoerus. The taxa occurring at Edisto Island in the Late Pleistocene represent essentially a coastal plain assemblage similar to late Pleistocene faunas found throughout Florida and the Gulf coastal region at least as far west as Texas (Lundelius 1972). ACKNOWLEDGMENTS. — We extend our thanks to A. Sanders (Charleston Museum), A. E. Crawford (Hampden-Sydney College), C. E. Ray (Smithsonian Institution), and S. D. Webb (Florida State Museum) for access to the collections under their care. Their advice, criticism and comments were particularly helpful. We also thank G. Ed- mund, A. Holman and D. Gillette for their useful correspondence. We particularly thank C. E. Ray and S. D. Webb for their exceptionally helpful and critical reviews. Support for this study was provided by the Departments of Geology and Zoology, University of Georgia. Much of the work was completed by the senior author at the Charleston Museum, which is thanked for a positive, productive atmosphere in which to work. This paper is a con- tribution of the University of Georgia Museum of Natural History. LITERATURE CITED Aguirre, Emiliano. 1969. Evolutionary history of the elephant. Science 164 : 1366-1376. Allen, G. M. 1926. Fossil Mammals from South Carolina. Bull. Mus. Comp. Zool. Harv. Univ. 67:447-467. Anderson, Elaine. 1968. The Carnivora. Fauna of the Little Box Elder Cave, Converse County, Wyoming. Univ. Colo. Stud. Ser. Earth Sci. 6:1-59. Arata, A. A. 1959. Reevaluation of the Pleistocene Urocyon seminolensis from Florida. Q. J. Fla. Acad. Sci. 22:133. and J. H. Hutchinson. 1964. The raccoon ( Procyon ) in the Pleistocene 26 Janet A. 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G. 1967. A re-evaluation of fossil turtles of the Chrysemys scripta group. Tulane Stud. Geol. 5:53-66. Webb, S. D. (ed.) 1974. Pleistocene mammals of Florida. Univ. Presses Fla., Gainesville. 270 pp. Whitmore, Frank C., K. O. Emery, H. B. S. Cooke and D. J. P. Swift. 1967. Ele- phant teeth from the Atlantic continental shelf. Science 755:1477-1480. Winkler, C. D., and J. D. Howard. 1977. Correlation of tectonically deformed shorelines on the southern Atlantic coastal plain. Geology 5:123-317. Woodburne, M. O. 1958. A fossil alligator from the Lower Pliocene of Okla- homa. Pap. Mich. Acad. Sci. 44:47-49. Accepted 3 April 1980 . . The Status of Cleptoria shelfordi Loomis, with the Proposal of a New Genus in the Milliped Family Xystodesmidae (Polydesmida) Rowland M. Shelley North Carolina State Museum of Natural History, P. O. Box 27647, Raleigh, North Carolina 27611 ABSTRACT. — The new xystodesmid milliped genus Brevigonus is proposed to accommodate Cleptoria shelfordi Loomis. It is monotypic and distinguished by the brevity of the male gonopods, which in situ extend barely beyond the margin of the aperture. Two basic gonopodal variants of B. shelfordi exist, labeled A and B, but they do not appear to be reproductively isolated, and their sympatric ranges likewise rule out subspecific status. Variant A possesses an enlarged medial flange on the acropodite and a basal telopodal spine, whereas in variant B the spine is absent and the flange is reduced. Variant B, however, displays a promi- nent medial spur on the acropodite which is absent or greatly reduced in variant A. The two variants also differ in the curvature of the acropodite and its apical configuration. Brevigonus shelfordi occurs in a narrow band along the north side of the Savannah River in the Piedmont Plateau of South Carolina, from the vicinity of Clemson to McCormick. It has not been found across the river in Georgia, where the dominant xystodesmid is Cleptoria ahbotti Hoffman. In 1942 and 1943 Professor V. E. Shelford of the University of Illinois collected millipeds in the southeastern United States and sent them to H. F. Loomis, who published the results of the survey (Loomis 1944). He described three new xystodesmid species, one of them named for Prof. Shelford and assigned to the genus Cleptoria, probably because of its large size, the triangular prefemoral process of the male gonopod, and the general appearance of the telopodite. Since its description, Cleptoria shelfordi has been mentioned in only two other taxonomic publications. The first was the North American checklist (Chamberlin and Hoffman 1958), which merely cited the species and incorrectly repor- ted the type locality (the De La Howe Forest) as being in Lincoln County, Georgia. It is actually across the Savannah River in McCormick County, South Carolina, approximately 1 1.3 km (7 mi.) west of the town of McCormick. The second reference was Hoffman’s (1967) revision of Cleptoria, in which he transferred shelfordi to “another generic group so far unpublished.” Thirteen years later this genus is still unpublished, and technically shelfordi has no generic name, a most awkard nomenclatorial situation. Brimleyana No. 3: 31-42 July 1980. 31 32 Rowland M. Shelley The shelfordi problem has interested me since I examined the holotype in connection with a study of abnormal xystodesmids (Shelley 1977). The eighth segment of this milliped is without legs or a sternum (Fig. 1), a condition unreported at that time in the Diplopoda. Of further interest, however, was the contrast between the large size of the shelfordi type and the brevity of the gonopods, which were the shortest I had seen on an aphelorine xystodesmid and seemed almost aborted. Because of this specimen and the nomenclatorial problem, I visited the type locality in 1976 and found congeneric material in a 64 km (40 mi.) section along the north side of the Savannah River in Abbeville and McCormick coun- ties. I investigated habitat on both sides of the river, but found pertinent forms only in a narrow strip, 32 km (20 mi.) wide, on the South Carolina side. One male from Oconee County is present in Hoffman’s collection, but field trips to this area and Anderson County have been unproductive. Superficial examination revealed that shelfordi requires a new genus, and I propose the name Brevigonus to emphasize this feature of the male genitalia. Still unresolved is the number of species comprising the taxon, and I have conducted meticulous field surveys to try to find out. My current conception of a monotypic genus may eventually be altered, since two basic gonopodal variants are evident in the material at hand. They share more features than most aphelorine congeners, however, and thus seem not to be reproductively isolated. Moreover, they occur sym- patrically throughout the range of the genus, and I know of no aphelorine genus in which all the species occur over the entire range. Sampling to date has been thorough, and additional collecting will probably not further clarify the situation. Consequently, I recognize only one species in Brevigonus, and the present contribution is offered mainly to redescribe shelfordi and to provide it with a generic name. Brevigonus, new genus Type species. — Cleptoria shelfordi Loomis, 1944. Description. —A genus of large, robust xystodesmids with the following characteristics: Body composed of head and 20 segments in both sexes; W/L ratio 22-24%. Head of normal appearance, smooth, polished; epicranial suture distinct, not bifid; facial setae reduced, epicranial and interantennal ab- sent. Antennae relatively short, with four antennal cones. Terga smooth, polished. Collum large, broad, extending slightly beyond ends of following tergite on each side. Paranota moderately depressed, continuing slope of middorsum; peritremata flat on anterior- New Milliped Genus 33 Figs. 1-2. 1, B. shelfordi holotype, segments 8 and 9, ventral view. 2, topotype, segments 6 and 7, ventral view. Scale lines = 1.00 mm. most segments, becoming more distinct and elevated in midbody region; ozopores opening dorsad. Caudal segments normal for family. Sterna variable with large process on segment 4 and two small struc- tures between both pairs of legs on segment 5. Postgonopodal sterna a modification of bilobed variation discussed by Hoffman (1965). Female sterna similar to postgonopodal sterna of males. Gonopodal aperture subelliptical. Gonopods overlapping in situ, extending forward just beyond anterior margin of aperture. Coxae massive, without apophyses, connected by membrane only, no sternal remnant. Prefemora of normal size, usually with large, cuneate processes arising on dorsal sides. Acropodites extremely short, curving broadly dis- tad to prefemora, apically blunt, with or without a variable medial flange, spur, and basal spine. Cyphopodal aperture rounded, situated lateral to second legs. Cyphopods large, with receptacle and tips of valves visible in situ. 34 Rowland M. Shelley Receptacle large, convoluted, cupped around ventral edges of valves; valves elongate, oriented subdorsoventrally in body; operculum minute, located under dorsal end of valves. Range. —Along Savannah River in piedmont South Carolina from the vicinity of Clemson to McCormick. Species: One Relationships. — Brevigonus is most closely related to Cleptoria. The acropodite of the latter also is rather short, and this plus the common wedgeshaped prefemoral processes and the massive nature of the gonopods indicate close affinity between the two genera. I believe that Brevigonus evolved more recently than Cleptoria and that it may be derived from that genus. 3 Figs. 3-6. B. shelfordi holotype. 3, sternal process of segment 4, caudal view. 4, left gonopod, medial view, setation omitted. 5, same, lateral view. 6, topotype, distal portion of acropodite, medial view. Scale line = 1.00 mm for Figs. 4-5, 0.67 mm for Figs. 3 and 6. New Milliped Genus 35 Brevigonus shelfordi (Loomis), new combination Figs. 1-14 Cleptoria shelfordi Loomis, 1944:172-173, Fig. 4. Chamberlin and Hoffman, 1958:28. Type specimen. —Male holotype, Museum of Comparative Zoology (MCZ), collected by V.E. Shelford, 10 July 1942, from station 1 1, De La Howe Forest, McCormick Co., South Carolina. Diagnosis.— With the characters of the genus. Holotype. —Length 56.1 mm, width 13.0 mm, W/L ratio 23.2%. Paranota depressed, creating a high height/width ratio of 60.3% at mid- body. Body essentially parallel sided from segments 6-14, tapering at both ends as indicated by following segmental widths: collum 8.7 mm 12th- 13th 13.0 2nd 9.7 14th 12.8 3rd 10.8 15th 12.2 4th 11.6 16th 11.4 5th 11.9 17th 9.4 6th-9th 12.0 18th 6.9 10th- 11th 12.8 Color in life unknown (see topotype description). Head capsule smooth, polished; width across genal apices 5.4 mm; interantennal isthmus broad (1.9 mm), smooth; epicranial suture shallow but distinct. Antennae relatively short, extending only to caudal edge of second tergite, becoming progressively more hirsute distally; first antennomere subglobose, 2-6 clavate, 7 short and truncate; relative lengths of anten- nomeres as follows: 2>3>4>6>5>1 >7. Genae not margined laterally, trace of medial impression, ends broadly rounded and ex- tending slightly beyond adjacent cranial margins. Facial setae as follows: frontal 1-1 (impressions only, setae not seen), genal 1-1, clypeal about 9- 9, labral about 12-12. Dorsum typical for family. Paranotal corners rounded on segments 1-6, becoming slightly more distinct on 7 and progressively more so caudally. Peritremata virtually flat on segments 1-7, only slightly elevated above paranotal surface; higher and more distinct on remaining seg- ments. Ozopores small and indistinct, situated near middle of paranota. Sternum of segment 4 produced into large, apically divided process (Fig. 3), subequal in length to width of adjacent coxae; sternum of seg- ment 5 produced into two small digitiform lobes between fourth pair of legs and two knoblike processes between fifth pair. Postgonopodal sterna 36 Rowland M. Shelley Figs. 7-12. Gonopodal variation in B. shelf ordi, medial views of acropodites of specimens from selected localities for variants A and B, as discussed in text. 7, Oconee Co., Clemson vicinity. 8, Abbeville Co., 4.8 km sw of Calhoun Falls near Savannah River. 9, McCormick Co., Hickory Knob State Park. 10, topotype, McCormick Co., 11.6 km wnw of McCormick, De La Howe Forest. 1 1, Abbeville Co., 10 km s of Abbeville, along road to Parsons Mountain camp- ground, Sumter National Forest. 12, McCormick Co., Baker Creek State Park. Scale line = 1.00 mm for all figures. elevated behind stricture, interrupted by transverse groove behind first pair of legs on each segment, expanded into broad platelike structure beyond caudal legs with shallow medial depression, small blunt lobes subtending anterior legs of each segment, larger rounded lobes beside caudal legs. Hypoproct broadly rounded, subacute distally, two paramedial setae arising from distinct rounded tubercles. Paraprocts with margins strongly thickened. Coxae generally without spines, low blunt “teeth” present on postgonopodal legs; prefemoral spines distinct on postgonopodal legs; tarsal claws darkened distally, bisinuately curved. Segment 8 without any trace of legs or sterna, teratological, caudal edge of metazonite broadly indented along midline (see Shelley 1977). Gonopodal aperture subelliptical, about 3.9 mm long and 2.5 mm New Milliped Genus 37 wide at midpoint; margins not raised, smoothly continuing sternal slope, thickened along caudal edge. Gonopods (Figs. 4-5): prefemoral process short and subtriangular, cuneate; acropodite widest at base, expanded into flange along medial edge which terminates in short spur about midlength of acropodite; acropodite constricted and broadly curved cephalad at midlength, apically expanded and flattened, with rounded subterminal lobe on dorsal edge, subacute apically; prostatic groove crossing over from medial to lateral sides at midlength curvature, open- ing apically. Topotypes.— The topotypes are glossy black dorsally with red paranota and a suggestion of a stripe along the anterior edge of the collum. The metaterga, however, are without stripes. As shown in Figure 6, the gonopods of some topotypes differ from those of the holotype, being distinctly forked apically with a hoodlike lobe overhanging the solenomerite. There is only a trace of this indentation in the holotype. The brevity of the gonopods is most clearly revealed in situ (Fig. 2), where they overlap each other and extend barely beyond the anterior edge of the aperture. Cyphopods (Fig. 13) of female topotypes as described for genus. 13 Fig. 13. Right cyphopod of female topotype, lateral view. Scale line = 1.00 mm. 38 Rowland M. Shelley Variation. — The only color variation involves the presence or absence of a narrow red stripe along the anterior edge of the collum connecting the paranotal spots. As mentioned in the introduction, two basic gonopodal variants exist. In variant A (Figs. 7-9) the flange of the gonopod is greatly enlarged, partially or completely obscuring the stem of the acropodite in medial view. This condition is accompanied by a prominent basal spine on the ventral side of the acropodite; reduction of the medial acropodite spur, which is absent from some specimens (Figs. 7-8); and a more com- pact curvature of the acropodite in which the tip is directed dorsad or downward toward the coxa in medial view. In variant B (Figs. 10-12), which includes material from the type locality, the basal spine is absent and the flange is reduced so that the prostatic groove is visible in medial view to about the point where it crosses to the lateral side of the acropodite. The spur is distinct in all these specimens, and the acropodite extends cephalad apically with the tip directed only slightly dorsad. The prefemoral process varies more in variant A than in B, and is missing from the male from Oconee County (Fig. 7). Likewise, the apical indenta- tion of the acropodite occurs only in variant B (Figs. 10-1 1), but as shown in Figure 12 it is not present on all specimens. In both groups the gonopods are large and heavily sclerotized. These two variants are inter- mixed and occur throughout the range of shelfordi, but the presence of characters from each variant in the other (e.g., the spur of B in A, Fig. 19, and the apical configuration of A in B, Fig. 12) suggests that they are not reproductively isolated. Their sympatric ranges argue against subspecific recognition, and the only alternative is a single species with variable gonopods. Should future discoveries justify taxonomic recognition of the two groups, the name shelfordi will be available for variant B, and a new specific name will be required for variant A. Ecology. — Brevigonus shelfordi usually is found under thin layers of leaves on relatively hard substrates near water sources; the male from Oconee County was discovered under a dead pig. This region of South Carolina is predominantly pine, and hardwoods are typically found only along creeks and rivers. The type locality is an exception, however, and the species is abundant in the protected forest tract near the grave of John De La Howe. Visits to this site in August 1976 and May 1977 produced 7M, 4F, and 7M, 2F, respectively, and many times these numbers could have been taken. The area is hilly, and shelfordi occurs in thick leaf piles, in open litter, under logs, and on banks of creeks. The following descrip- tion of the De La Howe forest is excerpted from Radford and Martin (1975:182). New Milliped Genus 39 The De La Howe woods is the “progenitor” of all piedmont forests. The canopy is dominated by loblolly pines on the slopes and in the draws with shortleaf pine on the ridges with several age classes and trees 12-42” D. B. H. The subcanopy is a mosaic of oaks and cover hardwood elements . . . On the ridges and slopes are red, white, black, post and southern red oaks along with pignut and mockernut hickories, red mulberry, yellow poplar and sweet gum; in the draws and sheltered slopes are beeches and southern sugar maple. Most of the subcanopy trees vary from 12- 24” D. B. H. Beneath the subcanopy is a layer of hop hornbeam, storax and flowering dogwood along with some tall blue haws ( Viburnum prunifolium). The shrub layer is composed of leatherwood in the draws, dwarf pawpaw, aromatic sumac, and maple-leaved viburnum on the slopes and deerberry ( Vaccinium stamineum ) and New Jersey tea along the ridge. . . . the herb layer is scant with the woody layers above but it is a unique combina- tion of partridge berry, wintergreen and pennywort ( Obolaria virginica). This is a classic example of the eastern North American Oak- Pine Forest with a maximum woody plant diversity. Tree size for the age of the stand is not overly impressive but the combination of woody species over this nearly circumneutral soil derived from a basic igneous rock and the abundance of the Obolaria make this a truly outstanding site. The tract is now recognized as a Natural Area by the Society of American Foresters. Distribution. — Brevigonus shelfordi ranges about 112 km (70 mi.) along the north side of the Savannah River in piedmont South Carolina, from the vicinity of Clemson to McCormick. It occurs about 32 km (20 mi.) from the river into South Carolina, but most individuals have been collected less than 16 km (10 mi.) from the river in Abbeville and McCor- mick counties. The species has not been encountered in Anderson County and does not appear to occur in Georgia. I have collected along the Savannah River in Hart, Elbert, Lincoln, and Columbia counties, the Georgia counties bordering the range of shelfordi, without finding the milliped. The dominant xystodesmid in this area is Cleptoria abbotti Hoffman, which seems to replace shelfordi south of the Savannah River. In July 1979 an assistant and I collected shelfordi along South Carolina highway 72 in Abbeville County close to the Savannah River, but just across the river in essentially the same habitat in Elbert County, Georgia, we found only C. abbotti. I thus feel that shelfordi is absent from Georgia even though this area has not been sampled as thoroughly as have areas in South Carolina. Specimens have been examined from the following South Carolina localities. Unless otherwise indicated, all were collected by the author and deposited in the N. C. State Museum collection, the invertebrate catalogue number of which is shown in parentheses. Materials from the 40 Rowland M. Shelley Fig. 14. Distribution of B. shelfordi in South Carolina. New Milliped Genus 41 Florida State Collection of Arthropods and the private collection of Richard L. Hoffman are indicated by the acronyms FSCA and RLH, respectively, and MCZ is the Museum of Comparative Zoology. Oconee Co.— Clemson vicinity, under dead pig, 2M, F, 18 July 1962, J. A. Payne (RLH). Abbeville Co.— 5.8 km w of Due West, along SC hwy. 37 at Little River, M. F, 11 June 1978, R .M. Shelley and W. B. Jones (A2067). 10.8 km w of Abbeville, along SC hwy. 71 at Penny Creek, 2M, 6 May 1977 (A 1546). 10 km s of Abbeville, Sumter National Forest, along road to Parsons Mountain campground, M, 3F, 9 August 1976 (A 1390); Parsons Mountain campground, M, 3F, 10 August 1976 (A1392); and near Parsons Mountain Lake, M. F, 6 May 1977 (A1548). 4.8 km sw of Calhoun Falls, near Savannah River, M, 30 April 1960, L. Hubricht (RLH); 3.8 km sw of Calhoun Falls, along SC hwy. 72, 0.9 km n of Savannah River, 3M, F, 6 May 1977 (A 1543), and M, 19 July 1979, R. M. Shelley and R. K. Tardell (A2837). McCormick Co. — 6 km n of Mt. Carmel, along SC Hwy. 79 at Swaney Creek, 2M, F, 5 May 1977 (A 1540). 15 km nw of McCormick, along SC hwy. 39 at Little River, M, 5 May 1977 (A 1539). 10.6 km nw of McCormick, jet. SC hwys. 28 and 81, 2M, 5 May 1977 (A 1538). 12 km wnw of McCormick, De La Howe Forest, Station 1 1, M, 10 July 1952, V. E. Shelford (MCZ); De La Howe Forest at end SC hwy. 25, 3.5 km s of jet. SC hwy. 81, 7M, 4F, 9 August 1976 (A1388), and 7M, 2F, 5 May 1977 (A1536), TYPE LOCALITY. 8 km w of McCormick, along US hwy. 378, M, 2 July 1958, collector un- known (FSCA). Baker Creek State Park, M. F, 8 August 1976 (A 1386). Hickory Knob State Park, 4M, 3F, 8 August 1976 (A 1387). Remarks. — Brevigonus shelfordi is the largest aphelorine xystodesmid species. Many individuals are comparable in size to Pachydesmus crassicutis (Wood) in the tribe Pachydesmini, which attains a length of over 60 mm and is generally considered the largest polydesmoid species in the United States. ACKNOWLEDGMENTS. -\ am grateful to Herbert W. Levi, Museum of Comparative Zoology, for the privilege of examining the holotype of Cleptoria shelfordi. Howard V. Weems, Jr., kindly loaned material in the Florida State Collection of Arthropods, and Richard L. Hoffman, Radford College, did likewise with material in his collection. Specimens from Baker Creek and Hickory Knob State Parks were col- lected with permission of the South Carolina Department of Parks, Recreation, and Tourism. Thanks are also extended to John E. Cooper, N.C. State Museum, for prepublication review and Renaldo G. Kuhler, N.C. State Museum, for preparing Figures 1 and 2. This study was sup- 42 Rowland M. Shelley ported in part by National Science Foundation Grant Number DEB- 7702596. LITERATURE CITED Chamberlin, Ralph V., and Richard L. Hoffman. 1958. Checklist of the millipeds of North America. U. S. Natl. Mus. Bull. 272:1-236. Hoffman, Richard L. 1965. Revision of the milliped genera Boraria and Gyalostethus (Polydesmida: Xystodesmidae). Proc. U. S. Natl. Mus. 7 77:305-348. 1967. Revision of the milliped genus Cleptoria (Polydesmida: Xystodesmidae). Proc. U. S. Natl. Mus. 724:1-27. Loomis, Harold F. 1944. Millipeds principally collected by Professor V. E. Shelford in the eastern and southeastern states. Psyche 57:166-177. Radford, Albert E., and David L. Martin. 1975. Potential Ecological Natural Landmarks, Piedmont Region, Eastern United States. Privately published for National Park Service. 249 pp. Shelley, Rowland M. 1977. Appendicular abnormalities in the milliped family Xystodesmidae (Polydesmida). Can. J. Zool. 55: 1014-1018. Accepted 17 March 1980 Recent Range Expansion of the Groundhog, Marmota monax, in the Southeast (Mammalia: Rodentia) Sarah S. Robinson1 and David S. Lee North Carolina State Museum of Natural History, P.O. Box 27647, Raleigh, North Carolina 27611 ABSTRACT. — In recent years the groundhog, Marmota monax, has greatly expanded its range in the Southeast and now is found in areas of the Piedmont Plateau and Coastal Plain where it did not occur in historical times. This change in distribution is believed to correlate with changing land use practices. The groundhog, Marmota monax, is generally distributed throughout northeastern North America. South of Pennsylvania and New Jersey it was until recently known mostly from Piedmont Plateau and montane areas. Paradiso (1969) indicated its apparent absence from the Delmarva Peninsula and perhaps most of the Coastal Plain of Maryland. In Virginia, Handley and Patton (.1947) reported it to be most common in the mountains, although they had many Piedmont and several Coastal Plain records. Bailey (1946) noted that groundhogs were “scarce” in the “tidewater region” of Virginia. In the Carolinas, Marmota has been reported only from mountainous regions (Brimley 1944-46; Golley 1966; Hall and Kelson 1959). This note documents its re- cent expansion in the Piedmont Plateau of North Carolina and in the Coastal Plain province of Maryland, Delaware, and North Carolina. Between 1970 and 1977 we compiled Delmarva records for Marmota. Data for North Carolina were obtained from a 1974 question- naire on the distribution of the groundhog that went to all field personnel of the North Carolina Wildlife Resources Commission and from recent records compiled by the North Carolina State Museum (NCSM). The questionnaires are on file in the Department of Mammalogy and Ornithology, NCSM, and voucher specimens from the areas discussed are in the museum’s mammal collections. DELMARVA PENINSULA Paradiso (1969) noted that M. monax was in all sections of Maryland “except the Eastern Shore, where it has until lately been un- known. There is recent evidence, however, that the species is extending its range into that section.” Grizzell (1955) reported that it first appeared on the Delmarva at the turn of the century, spreading from Pennsylvania, and by 1955 populations had increased sufficiently to be a probelm in 'Present address: Coastal Zone Resources Division, Ocean Data Systems, Inc., 4505 Franklin Avenue, Wilmington, NC 28403 Brimleyana No. 3: 43-48 July 1980. 43 44 Sarah S. Robinson and David S. Lee some areas. However, neither author had new records for the region east of the Chesapeake Bay. Allen (1950) reported that the species had in- vaded the Delmarva in the previous 50 years, noting that it first appeared near Galena, Kent County, Maryland, around 1900. This is apparently the only published report of its occurrence east of Chesapeake Bay. Casual interviews with residents of the area confirmed to us that groundhogs had recently appeared there, although agreement on the length of time they had been established was lacking. It is interesting that we have few records from south of Talbot and Caroline counties, Maryland, and Kent County, Delaware, despite inten- sive field work. This may reflect the change in soil composition which occurs in this general area. The well-to-poorly drained sandy loam and sandy clay loam soils (Sassafras and Sassafras-Fallsington associations), which are nearly continuous in the northern portion of the Delmarva, become fragmented here. The 75- to 125-cm deep loamy soils are mostly replaced by more sandy soil associations, and much of the area is dis- sected by extensive Spartina marshes which would further limit dispersal of Marmota. The southern limits of the groundhog on the Delmarva appear to terminate at the northern limits of pure stands of Loblolly Pine, Pinus taeda (Conant 1945). This change in community character appears to have effectively limited the natural dispersal of several vertebrate species: the Whip-poor-will, Caprimulgus vociferus, and Scarlet Tanager, Piranga olivacea (Stewart and Robbins 1958); and the Red-spotted Newt, Notophthalmus viridescens viridescens, Spotted Salamander, Ambystoma maculatum, Northern Red Salamander, Pseudotriton ruber ruber, and Northern Two-lined Salamander, Eurycea bislineata bislineata (Harris 1975). Also, disjunct populations of at least six species of upland plants occurring in communities of Coastal Plain hardwoods on the peninsula are excluded from areas which support extensive stands of P. taeda (Franz and Lee 1976). Delaware Records Coastal Plain: New Castle County. —Road shoulders/fields, 3 mi. (4.8 km) s of Middletown on SR 896; 1973 (Matapeake-Sassafras soil type). Road shoulders/hardwood forest, 8 mi. (12.9 km) n of Smyrna on US 13; 1973 (Matapeake-Sassafras). Fields, Lums Pond State Park; 1974. US 301 between Summit and Middletown; 1972. Kent County.— Dikes/fields, Bombay Hook National Wildlife Refuge; 1974-1976. Fields, 7 mi. (1 1.3 km) s of Leipsic on SR 9; 1976 (NCSM 2933). Sussex County.— Fields, Prime Hook Wildlife Refuge; 1974 (sandy loam). Maryland Records Piedmont Plateau: Cecil County. — Riv er floodplain, Conowingo and Rt. 222, 5 mi. (8.1 km) s of Conowingo; 1968-73. Rising Sun; 1969. Groundhog Range Expansion 45 Coastal Plain: Cecil County. —Chesapeake City; 1941 (NCSM 1997). Kent County. Galena (Allen 1950). Fields, Rock Hall; 1973. Eastern Neck National Wildlife Refuge; 1972. Talbot County.— Fields, 3 mi. (4.8 km) east of jet. SR 404 and 309; 1973. Hardwood forest, Mill Creek Sanctuary, 8 mi. (12.9 km) east of Easton; 1973. Queen Annes County. — Centerville; 1974. Caroline County. — Fields, Denton; 1973. NORTH CAROLINA The distribution of the groundhog in the North Carolina Piedmont Plateau and Coastal Plain is undergoing a transition similar to that observed in Maryland and Delaware. In the western Piedmont of the state groundhogs are now known from Gaston, Cleveland, Catawba, Iredell, Davie, Forsyth, and all counties farther west. In the central Pied- mont they occur through Guilford, Randolph and Alamance counties, as well as in the tier of counties along the Virginia border — Surry, Stokes, Rockingham, Caswell, Person and Granville. The individuals forming the presently small, scattered Piedmont populations probably represent immigrants from western North Carolina and adjacent counties in Virginia. The construction of two large reservoirs on the North Carolina- Virginia border (Kerr Reservoir and Lake Gaston) may have facilitated movement of the groundhogs into North Carolina as massive areas were cleared prior to flooding. Populations in Orange and Durham counties appear to date from the early 1950s, when 13 animals from the western part of the state were released near Hillsborough, Orange County (NCSM files). Specimens and sight records from Wake County perhaps represent an expansion of this population. Groundhogs also have expanded into the northeastern Coastal Plain counties of North Carolina, where they are primarily distributed along rivers and larger streams— the Neuse River in Greene County and Meherrin River in Hertford County, the Roanoke River in several coun- ties, the Tar River in Nash and Edgecombe counties, and the Chowan River in Gates and Chowan counties. The Roanoke and Tar river areas appear to have been colonized first, and our earliest Coastal Plain records are from Halifax and Nash counties. There are now populations from eastern Vance County through Warren, Halifax, Northampton, Hertford, Gates, Chowan, Bertie, Martin, Edgecombe, Nash, Wilson, Johnston and Greene counties. The Coastal Plain groundhogs typically build their burrows in streambanks and along drainage ditches as well as at the edges of open fields. The levees along the Roanoke River, in par- ticular, contain a large number of burrows. North Carolina Records Piedmont Plateau: Davie County.— Mocksville; 1975. Durham County. — 10 mi. (16.1 km) n of Durham on US 501; 1979. Franklin 46 Sarah S. Robinson and David S. Lee County.— Lynch Creek, 0.24 mi. (0.39 km) from SR 1240; 1977 (Appling- Cecil sandy loam). Gaston County. — Shoulder of US 321, between Gas- tonia and Dallas; 1956 (Appling-Cecil sandy loam). Granville County.— Fields, Critcher Farm near Stovall; 1974 (Herndon-Georgeville silt loam or silty clay loam). Pasture, Butner; 1971. Iredell County.— 9 mi. (14.5 km) e of Statesville on US 40, 1979. Orange County. — 13 specimens from mountains released; 1950. Eno River State Park (NCSM 2675); 1977. Hillsborough (NCSM 2676); 1975. Wake County.— Urban Raleigh; 1939, and (NCSM 3026, 3062); 1976. 10 mi. (16.1 km) e of Raleigh; 1950 (Appling-Cecil sandy loam). Cary; 1977. Warren County.— 1-85 at SR 1210; 1977 (Granville- White Store sandy loam). 3 mi. (4.8 km) n of Wise near US 1 on highway shoulder; 1976 (Appling-Cecil sandy loam). Fig. 1. Distribution of Marmota monax in the Southeast. Dashed line indicates approximate eastern limit of historical range. Solid symbols are specific locality records; hollow symbols are county records only. Compiled from Golley 1962, 1966; Hall and Kelson 1959; Handley and Patton 1947; Paradiso 1969; and NCSM records. Groundhog Range Expansion 47 Coastal Plain: Bertie County.— Old homestead, 200 yd. (182 m) from Roanoke River near Woodville; 1975 (terrace soils— loamy sands to clay loams). Levee along Roanoke River near Kelford; 1976 (Dunbar- Lynchburg sandy loam). Edgecombe County.— Downtown Tarboro un- der building; no date. Gates County.— Peanut fields, US 13 near Storey’s crossroad; 1973 (Craven-Shubuta sandy loam). Greene County .—8 mi. (12.9 km) w of Snow Hill; 1972 (Craven-Shubuta sandy loam). Halifax County.— Levees along Roanoke River at Weldon; 1971-75 (alluvial soils). Hertford County.— Burrow under bridge abutment 0.7 mi. (1 km) s of Menola on SR 1150; 1973 (silt loam). Johnston County. — Clayton (NCSM 256); 1951. Martin County. — Swimming in Roanoke River near Jamesville; 1973 (terrace soils— loamy sands to clay loams). Nash County.— Surburban area west side of Rocky Mount; ca. 1948 (Norfolk- Ruston sandy loam). Shoulder NC 43; 1970 (Norfolk-Ruston sandy loams). Northampton County.— Levees and brushy fields Occoneechee Neck, along Roanoke River; 1971-1975 (Norfolk-Ruston and Appling- Cecil sandy loams). 3 to 5 mi. (4.8 to 8.1 km) n of Rich Square; no date (Lenior-Coxville silt loams). Clearing of forests for agricultural purposes has increased the amount of habitat suitable for groundhogs in the Piedmont Plateau and Coastal Plain provinces of the Southeast. Highway and utility rights-of- way and river levees appear to have provided paths of dispersal from other upland areas. The success of Marmota in this region may in part be a result of a prolonged growing season combined with a limited period of groundhog dormancy. In North Carolina we have reports of active groundhogs from all months of the year. ACKNOWLEDGMENTS. — We would like to thank Frederick S. Barkalow, Jr., N.C. State University; Merrill Lynch, formerly NCSM; and personnel of the N.C. Wildlife Resources Commission, for making available some of the distributional information presented here. John E. Cooper, NCSM, read and commented on the manuscript. LITERATURE CITED Allen, D. L. 1950. The fabulous whistlepig. Sports Afield 123: 28-29, 78-80. Bailey, John W. 1946. The Mammals of Virginia. Williams Printing Co., Rich- mond. 413 pp. Brimley, C. S. 1944-46. The Mammals of North Carolina. 18 installments. Carolina Tips, Carolina Biological Supply Co., Elon College. 39 pp. Conant, Roger. 1945. An annotated checklist of the amphibians and reptiles of the Del-Mar-Va Peninsula. Soc. Nat. Hist. Delaware, Wilmington. 9 pp. Franz, Richard, and D. S. Lee. 1976. A relict population of the Mottled sculpin, Cottus bairdi, from the Maryland coastal plain. Chesapeake Sci. / 7(4): 301-302 48 Sarah S. Robinson and David S. Lee Golley, Frank B. 1962. Mammals of Georgia. Univ. Georgia Press, Athens. 218 pp. 1966. South Carolina Mammals. Charleston Museum, Charleston. 181 pp. Grizzell, Roy A. 1955. A study of the southern woodchuck, Marmota monax monax. Am. Midi. Nat. 55:257-293. Hall, E. Raymond, and K. R. Kelson. 1959. The Mammals of North America. Vols. I and II. Ronald Press Co., New York. 1,083 pp. Handley, Charles O., and C. P. Patton. 1947. Wild Mammals of Virginia. Comm. Game and Inland Fisheries, Richmond. 220 pp. Harris, Herbert S., Jr. 1975. Distributional survey (Amphibia/Reptilia); Mary- land and the District of Columbia. Bull. Md. Herpetol. Soc. / 7(3):73- 1 67. Paradiso, John L. 1969. Mammals of Maryland. U. S. Dep. Inter., N. Am. Fauna 66, Washington. 193 pp. Stewart, Robert E., and C. S. Robbins. 1958. Birds of Maryland and the District of Columbia. U. S. Dep. Inter., N. Am. Fauna 62, Washington. 401 pp. Accepted 2 February 1980 Additional Records of Albinistic Amphibians and Reptiles from North Carolina William M. Palmer and Alvin L. Braswell North Carolina State Museum of Natural History, P. O. Box 27647, Raleigh, North Carolina 27611 ABSTRACT. — Albinism has been reported previously in 6 species of reptiles from North Carolina: Cemophora coccinea, Crotalus horridus, Elaphe guttata, E. obsoleta, Farancia abacura, and Sceloporus undulatus. This paper describes 16 additional albinistic specimens of 12 amphibian and reptile species from the state. Species represented include the salamanders Ambystoma talpoideum, Amphiuma means, Eurycea bislineata, and Siren intermedia; the toad Bufo woodhousei; the turtle Chelydra serpentina; and the snakes Carphophis amoenus, Diadophis punctatus, E. obsoleta, Nerodia fasciata, N. taxispilota, and Virginia striatula. A description of a snake (F. abacura ), previously reported but not described, also is included. INTRODUCTION Albinism is well known in amphibians and reptiles and records of its occurrence are prevalent in the literature. However, this phenomenon in North Carolina has been reported only in the lizard Sceloporus undulatus (Hensley 1968) and the snakes Cemophora coccinea (Brimley 1944), Elaphe obsoleta (Meacham 1946), and E. guttata, Farancia abacura and Crotalus horridus (Hensley 1959). This paper records from the state an additional 16 albinistic specimens of 12 species: 5 salamanders of 4 species, 1 toad, 1 turtle, and 9 snakes of 6 species. A description of an albinistic mud snake, F. abacura, recorded but not described by Hensley (1959), also is included. MATERIALS AND METHODS Color descriptions of a few individuals were made with the aid of color swatches, and capitalized color names with swatch numbers in parentheses are from Smithe (1975). SVL and TL indicate snout-vent length and total length, respectively. Common and scientific names follow those recommended by Collins et al. (1978). Specimens in the collection of the North Carolina State Museum of Natural History are designated by the acronym NCSM and the catalog number. All specimens were examined alive unless specified otherwise, and those for which no disposition is given are no longer extant. ANNOTATED LIST Amphibia Amphiuma means, Two-toed Amphiuma NEW HANOVER CO.: near Seabreeze, March or April 1968. An Brimleyana No. 3: 49-52 July 1980. 49 50 William M. Palmer and Alvin L. Braswell amphiuma about 230 mm TL was collected from a shallow roadside ditch and described by George Tregembo (pers. comm.). It was white with red eyes and scattered small, pinkish spots. Siren intermedia intermedia , Eastern Lesser Siren BRUNSWICK CO.: near Sunny Point, 23 April 1977. A specimen about 200 mm TL was collected with several individuals of the normal phenotype from a borrow pit pond in sandy flatwoods. Its dorsum and limbs were yellow, slightly brighter than Straw Yellow (56), and its venter was paler but near that color. Its gills were rust colored, between Cinnamon-Rufous (40) and Ferruginous (41), and its eyes were red. Ambystoma talpoideum, Mole Salamander MACON CO.: 22.5 km (14 mi.) wsw of Franklin, 13 December 1975, NCSM 15961 (21 larvae), 17872 (adult male). Two individuals among a series of 22 larvae collected from a floodplain pond were largely amelanistic in having the characteristic dark brown and black larval pat- terns replaced by pale gray and gray. One (37.0 mm SVL, 70.9 mm TL) was preserved with the typical specimens shortly after capture. The other, reared in the laboratory, was examined and preserved on 29 May 1977 as an adult (58.0 mm SVL, 100 mm TL). It was pale gray and translucent with many blood vessels and some internal organs visible through the skin. The parotid area and the dorsum of the tail were whitish due to clustered poison glands. The eyes were gray with dark pupils. Eurycea bislineata cirrigera, Southern Two-lined Salamander NEW HANOVER CO.: 5.6 km (3.5 mi.) n of Carolina Beach, March 1971, NCSM 10072. A female (40.0 mm SVL, 91.0 mm TL), collected with several individuals of the normal phenotype under leaves in a moist ditch, had a pinkish dorsum and venter with a network of un- derlying blood vessels visible through the skin. The sides of the head and upper tail had faint brown lines, and the middorsum of the tail was pale yellow. The eyes were brassy with dark pupils. Bufo woodhousei fowleri, Fowler’s Toad WAKE CO.: Raleigh, June 1965, NCSM 3884. This specimen (34.5 mm SVL) had translucent pinkish skin and dark blue eyes with reddish pupils. Color transparencies from life are in NCSM. Reptilia Chelydra serpentina serpentina, Common Snapping Turtle DAVIE CO.: 12 August 1978. The Taylorsville Times, 17 August 1978, contained a very clear photograph of a 16-pound albino caught in a farm pond by Jerry Petrea. From color photographs provided by Jerry and Riley Petrea, the carapace was pale yellowish brown with dark brown sutures. The head, limbs, and tail were slightly paler with scattered brown markings, and the eyes were yellowish with light green pupils. Albinistic Amphibians and Reptiles 51 Nerodia fasciata fasciata, Banded Water Snake SAMPSON CO.: Laurel Lake, 4.8 km (3 mi.) e of Salemburg, Sep- tember 1958. Two neonates, among a litter of 9 produced by a female of the normal phenotype, had red pupils and tongues and a white dorsal groundcolor with faint pink crossbands. The other siblings had typical patterns. Nerodia taxispilota, Brown Water Snake BLADEN-PENDER CO.: county line, along Black River, May 1961, American Museum of Natural Elistory (1 12347). An adult male had a pale yellowish dorsum with fine light brown stippling and without traces of blotches. The venter was whitish with pale brown markings, and the eyes were dark. Virginia striatula, Rough Earth Snake WAKE CO.: Cary, 4 April 1978, NCSM 20448. A female (about 200 mm TL), found under debris with several specimens of the normal phenotype, had red pupils and a pink tongue. The parietal band was pale yellow and faint. The remainder of the dorsum was pinkish tan, near Salmon (6). The venter was light gray, near Smoke Gray (44). Color transparencies from life are in NCSM. Diadophis punctatus ssp., Ringneck Snake DAVIDSON CO.: 1 1.3 km (7 mi.) s of Denton, May 1979, NCSM 20077. This female (348 mm SVL, 443 mm TL) had a pale olive brown dorsum, a light yellow neck ring bordered by brown, white labials and chin, and reddish brown eyes with dark pupils. The anterior venter was pale yellow, grading to orange yellow posteriorly, and the midventral spots were grayish brown. The snake, although not albinistic to the degree shown in most species here described, was nonetheless con- spicuously paler than typical specimens from the state and we consider it an amelanistic variant. Color transparencies from life are in NCSM. Carphophis amoenus amoenus, Eastern Worm Snake ROWAN CO.: 4.8 km (3 mi.) nw of Salisbury, 11 April 1979, NCSM 20454. A female about 240 mm TL, collected in a suburban yard and described by R. B. Julian (pers. comm.), was uniformly whitish with a dark pink tongue and reddish eyes. Farancia abacura abacura, Eastern Mud Snake HERTFORD CO.: Wiccacon River swamp, near confluence with Chowan River, 5.6 km (3.5 mi.) ne of Harrellsville, 17 July 1947, NCSM 3193. Hensley (1959) reported this adult on the basis of information received from the late Harry T. Davis, former director of NCSM, who apparently neglected to include its description. Color transparencies made of the living snake show that the dorsum was uniformly pinkish. Eye color cannot be ascertained from the photographs, but it appears to have been dark. Mounted and exhibited for more than 30 years, this 52 William M. Palmer and Alvin L. Braswell snake has now faded to a pale yellow with reddish brown mottling on the head. Elaphe obsoleta obsoleta, Black Rat Snake GASTON CO.: 5.6 km (3.5 mi.) sw of Gastonia, summer 1977, Schiele Museum at Gastonia. A captive adult female examined by us on 2 December 1979 had a red tongue, red pupils, and a pinkish white dorsal groundcolor with 35 rather prominent reddish body blotches. Color transparencies from life are in NCSM. HARNETT CO.: 2.4 km (1.5 mi.) w of Angier, 24 April 1978, NCSM 20043. An adult female, found among the rafters of an old barn, had bright red pupils and a red tongue. The dorsum was white with pink or pale red pigment forming 34 faint middorsal body blotches, a series of alternating lateral blotches weaker than those of the dorsum, and 1 1 or 12 obscure tail bands. Pattern was most evident on the anterior body. The venter was yellowish white and nacreous, with faint pink mottling along the edges. Color transparencies from life are in NCSM. NORTHAMPTON CO.: 6.4 km (4 mi.) nw of Rich Square, 3 June 1970, NCSM 9428. An adult male, examined shortly after it was killed by a logging crew, had a plain whitish groundcolor with scarcely discernible pinkish middorsal body blotches. Eye color was not determined. Two reports of albinistic black rat snakes from Stanly County (Meacham 1946, Hensley 1959) were based on the same specimen, originally in NCSM but now lost. ACKNOWLEDGMENTS. — We are grateful to the following persons who collected and donated specimens or otherwise provided information: Victor Ambellas, Larry D. Dunnagan, Carl Hiatt, Darrel Jones, Robert B. Julian, Brian Keho, Warren Kimsey, John E. Kiser, Dan F. Lockwood, Pat Mitchell, Jerry and Riley Petrea, Martin O. Shackleford, Richard Stout, George and Robert Tregembo, and Dick and Rick Wells. LITERATURE CITED Brimley, C. S. 1944. Amphibians and reptiles of North Carolina. Reprinted from Carolina Tips (1939-1943). Carolina Biological Supply Co., Elon College, North Carolina. 63 pp. Collins, Joseph T., J. E. Huheey, J. L. Knight and H. M. Smith. 1978. Standard common and current scientific names for North American amphibians and reptiles. Soc. Study Amphib. Reptiles Herpetol. Circ. No. 7. 36 pp. Hensley, Max. 1959. Albinism in North American amphibians and reptiles. Publ. Mus. Mich. State Univ. Biol. Ser. 7(4): 133-159. 1968. Another albino lizard, Sceloporus undulatus hyacinthinus (Green). J. Herpetol. /(l-4):92-93. Meacham, Frank B. 1946. An albino pilot black snake from North Carolina. Copeia 1946(2): 102. Smithe, Frank B. 1975. Naturalist’s color guide. Am. Mus. Nat. Hist., New York. Accepted 14 March 1980 A Distributional Checklist of the Fishes of Kentucky Brooks M. Burr Department of Zoology, Southern Illinois University, Carbondale, Illinois 62901 ABSTRACT. —A compilation of records of fishes from Kentucky waters based on specimens deposited in museums, personal collecting, and accepted literature reports revealed that 229 species occur or did oc- cur in the state. A substantial amount of new distributional data is presented in the form of an annotated list including records of several species of fishes previously unreported from the state. Distributional statements in the checklist are based on individual spot maps completed for all Kentucky fishes. A list of five problematical species is included at the end of the checklist. INTRODUCTION The fish fauna of Kentucky is more diverse than that of any other in- land area of comparable size in North America except Tennessee and Alabama. Presently, 229 species are known to occur or to have occurred in Kentucky waters and only 10 or 11 are the result of introduction by man. A major factor contributing to the present completeness of our knowledge of the Kentucky fish fauna has been its rich history of ichthyological investigations going back to the time of one of North America’s earliest ichthyologists, Constantine Samuel Rafinesque. Since Rafinesque’s groundbreaking work on Ohio River valley fishes (1820) there have been four other reports on Kentucky fishes (Woolman 1892, Garman 1894, Evermann 1918, Clay 1975). Woolman’s study is of im- mense historical value in documenting the distribution of many Ken- tucky fishes before most of the changes brought on by man took place. Garman’s and Evermann’s reports are mostly compilations containing little original information. The most recent work on Kentucky fishes (Clay 1975) did not include adequate distributional information and ex- cluded nearly 30 species of fishes that now occur in Kentucky. Moreover, much of Clay’s distributional information is in need of revision. This is in part due to the descriptions of new species, the resurrection of others from synonymy, recent intensive collecting in poorly worked areas, and examination of museum records of Kentucky fishes in many institutions throughout the eastern United States that apparently were not consulted by Clay. A new Fishes of Kentucky, aimed at summarizing the distribution Brimleyana No. 3: 53-84 July 1980. 53 54 Brooks M. Burr and biology of Kentucky fishes, is in preparation, but its appearance must await additional collecting and a critical compilation of Kentucky fish records from a few other institutions. MATERIALS AND METHODS Sources of Data I completed individual spot distributional maps for each species known to occur or to have occurred in Kentucky waters. The dis- tributional statements are based on those maps. During a final check of all records, those that seemed unreasonable and were not substantiated by specimens were discarded. The maps and resulting list are based on recent personal collecting, various regional surveys conducted by Kentucky Fish and Wildlife Resources Agency personnel, unquestioned literature records, and specimens that I examined in the following collections: California Academy of Sciences (CAS); Cornell University (CU); Eastern Kentucky University (EKU); Field Museum of Natural History (FMNH); Florida State University (FSU); Illinois Natural History Survey (INHS); Ken- tucky Fish and Wildlife Resources Agency (KFW); Kentucky Nature Preserves Commission (KNP); University of Kansas (KU); Harvard Museum of Comparative Zoology (MCZ); Murray State University (MSU); Northeast Louisiana University (NLU); Ohio State University (OSU); Southern Illinois University at Carbondale (SIUC); Tulane University (TU); University of Louisville (UL); University of Michigan Museum of Zoology (UMMZ); National Museum of Natural History (USNM); University of Tennessee at Knoxville (UT); University of Tulsa (UTULSAC); and Western Kentucky University (WKU). In order to show the areas of Kentucky that have been adequately sampled and those that need special attention, the locations of approx- imately 1150 stations sampled mostly since 1950 were plotted (Fig. 3). Despite the number of sites sampled in the Licking River, it is an area particularly worthy of further collecting efforts (many of the existing records are of only game fishes), as are the upper Cumberland and Big Sandy rivers. Treatment in the Annotated List The 229 species of lampreys and fishes known to occur or having oc- curred naturally or by way of repeated introduction in Kentucky waters are grouped under family names and arranged in phylogenetic sequence following, in part, Greenwood et al. (1966) and Bailey et al. (1970). Within each family, genera and species are listed in alphabetical order. Common and scientific names follow Bailey et al. (1970). In an effort to make the list complete, I included several species that are not tax- onomically described but are known to occur in Kentucky. Most of them Kentucky Fishes 55 have been recognized for more than 20 years and their distributions are accurately known. I have not departed from the 1970 list of names even though recent studies (some unpublished) reveal that the names of some Kentucky fishes are affected. Fundulus notti will probably be accepted for F. dispar (Wiley 1977); Menidia audens will probably be changed to M. beryllina. Although several studies indicate that Notropis chrysocephalus intergrades or hybridizes extensively with N. cornutus in various parts of its range, I have followed Gilbert (1961) in using the name N. chrysocephalus for the Kentucky populations. The Kentucky population hitherto known as Percina uranidea should now be called P. ouachitae (Williams and Etnier 1977). I divided Kentucky into 11 subunits, which correspond to river systems, or fish faunal blocks (Fig. 1). For each species, letters denoting the Kentucky subunits in which it is known to occur follow directly after the common name, which is followed by a concise statement regarding the species’ current or former distribution in the state. For species re- stricted to the large bordering Ohio and Mississippi rivers, designation of subunits is not applicable. My use of the terms “generally distributed,” “occasional,” or “sporadic” follow the definitions of Smith (1965). Distribution is sometimes expressed in terms of sections of the state, such as eastern two-thirds or western half. In many instances, it is ex- pressed in terms of specific drainage systems or waters such as Mis- sissippi River, lower Ohio River, and Big Sandy River drainage. For species known in Kentucky from only one or a few records, the name of the stream or major drainage basin and the county involved usually are given, as well as the acronym of the museum or university that contains specimens. The counties of Kentucky are depicted in Figure 2. In a few cases I cited recent references to uncommon species, especially if the papers pointed out more detailed information on their Kentucky ranges. A list of a few problematical species is included in this report. The annotations for these species, all hypothetical in the Kentucky fauna, are self-explanatory. Also included in the problematical list are species that were included in earlier lists of Kentucky fishes, but with the present state of our knowledge can now be deleted from the Kentucky list. ANNOTATED LIST OF SPECIES The 229 species in the following list represent 69 genera and 27 families. Twelve species (Alosa alabamae, Clinostomus elongatus, Hemitremia flammea, Hybopsis x-punctata, Notropis amnis, Lagochila lacera, Lota lota, Ammocrypta asprella, A. clara, A. vivax, Etheostoma microperca, Percina burtoni) have not been recently encountered and their current status in Kentucky is discussed. 56 Brooks M. Burr Fig. 1 . Divisions of major drainages and/or ichthyofaunal blocks as they relate to fish distribution patterns. A, lower Ohio River tributaries, Mayfield and Obion creeks and Bayou du Chien. B, Terrapin Creek and other small tributaries of Obion River. C, lower Tennessee and Clarks rivers. D, lower Cumberland River. E, Tradewater and lower Green rivers. F, upper Green and Barren rivers. G, Salt River. H, upper Cumberland River. J, Kentucky River. K, Licking River. L, Tygart’s Creek, Little Sandy and Big Sandy rivers. Kentucky Fishes 57 Petromyzontidae — lampreys Icthyomyzon bdellium (Jordan). Ohio lamprey. D,E,F,G,H,J,K,L. Occasional in the Green, Cumberland, Kentucky, Licking and Ohio rivers. The specific distinctiveness of this form deserves further study. Hubbs and Trautman (1937:14) resurrected the name bdellium and suggested that the two “geographic forms” ( bdellium and castaneus ) may intergrade in the lower Ohio River. Starrett et al. (1960) identified only /. castaneus in the Wabash River, Illinois, where Hubbs and Trautman had earlier identified specimens as I. bdellium. The characteristics of the two forms overlap greatly and specimens cannot be assigned with confidence to the form bdellium; consequently, parasitic lampreys from the middle and upper Ohio River and its major tributaries are identified arbitrarily as bdellium on the basis of geography. Ichthyomyzon castaneus Girard. Chestnut lamprey. C,D. Occasional in the lower Ohio, Mississippi, Cumberland and Tennessee rivers. See comments under I. bdellium. Ichthyomyzon fossor Reighard and Cummins. Northern brook lam- prey. J,L. Rare in the upper parts of the Kentucky (EKU, UMMZ, KNP), Big Sandy (UL) and Little Sandy River (KNP) systems. Ichthyomyzon gagei Hubbs and Trautman. Southern brook lamprey. C. A record based on one specimen from Clarks River, Calloway County (SIUC). Ichthyomyzon greeleyi Hubbs and Trautman. Allegheny brook lam- prey. F,H,J. Rare and sporadic in the upper Green (USNM), Cum- berland (REJ) and Kentucky (KFW, UL) drainages. Hoyt’s (1979) record of this species from the Ohio River at Paducah is not considered valid, and was probably based on misidentified I. castaneus. Ichthyomyzon unicuspis Hubbs and Trautman. Silver lamprey. C,E,F,H,J. Occasional in large rivers or their major tributaries throughout the state. Lampetra aepyptera (Abbott). Least brook lamprey. B,C,D,E,F, G,H,J,K,L. The most common lamprey in Kentucky, occurring in small to medium-size streams throughout the state except in the extreme west. Lampetra lamottei (Lesueur). American brook lamprey. F,J,L. Occasional in the upper reaches of the Barren, Green, Kentucky and Big Sandy drainages. Acipenseridae — sturgeons Acipenser fulvescens Rafinesque. Lake sturgeon. C,D,H. Formerly present in the main channels of the Ohio, Mississippi, Tennessee and Cumberland rivers (Call 1896, Woolman 1892, Evermann 1902). Most recent specimen from Cumberland River, McCreary County (UL). Status in Ohio and Mississippi rivers uncertain although several commer- cial fishermen told me that lake sturgeons are caught during early spring 58 Brooks M. Burr in the Mississippi River. Scaphirhynchus albus (Forbes and Richardson). Pallid sturgeon. Rare in the main channel of the Mississippi River where it is captured by commercial fishermen. No definite records, but descriptions of specimens by fishermen apply to this species. It has been taken farther south in the Mississippi River (Bailey and Cross 1954), so is clearly part of the Ken- tucky fauna. Scaphirhynchus platorynchus (Rafinesque). Shovelnose sturgeon. Occasional in the main channels of the Ohio and Mississippi rivers. Polyodontidae — paddlefishes Polyodon spathula (Walbaum). Paddlefish. A,C,D,H,J,K. Occa- sional in the Mississippi, Ohio, Cumberland, Tennessee, Kentucky and Licking rivers. Not reported from the Licking River since Barbour (1951). Lepisosteidae — gars Lepisosteus oculatus (Winchell). Spotted gar. A,D. Rare in the Mis- sissippi (SIUC), lower Ohio and Cumberland rivers (UL). Has not been taken farther east in the Ohio River than near the mouth of the Tradewater River. Lepisosteus osseus (Linnaeus). Longnose gar. C,D,E,F,G,H,J,K,L. The most common and widespread gar in Kentucky, occurring in large rivers (and lakes) and their major tributaries throughout the state. Lepisosteus platostomus Rafinesque. Shortnose gar. A,C,D. Limited to the western one-fourth of Kentucky where it is most common in Bayou du Chien, Obion Creek and Land Between the Lakes. Occa- sional to common in the lower Ohio and Mississippi rivers. Lepisosteus spatula Lacepede. Alligator gar. A,D. Four valid records presently are known, one each from the Tennessee River (Barbour 1963), mouth of the Ohio River (picture at INHS), Ohio River at Paducah (Hoyt 1979), and mouth of Bayou du Chien (EKU). Trautman (1957) reported other records (mostly anecdotal) of the species from the Ohio River as far east as Bracken County. Status uncertain, although Hoyt (1979) reported capture of 20 individuals during a two year study of fish impingement at the Shawnee Steam Plant, Paducah. Amiidae — bowfins Amia calva Linnaeus. Bowfin. A,C,D,E,G,TL. Sporadic on the Coastal Plain (INHS, MSU, SIUC, UL) and in backwater areas of the Green (SIUC, KNP), Salt (UL) and Kentucky rivers (KFW). Branson (1977) reported specimens from Tygart’s Creek, Carter County (EKU). BRACKfN Kentucky Fishes 59 o 3 3 O C/2 0> 3 3 O CJ 0> -3 f— c4 tob l z 60 Brooks M. Burr 0.7 mm) containing yolk were present in the oviducts or had recently been laid as evidenced by long, thin oviducts with clearly separated muscle bands (Gordon 1953; Ireland, 1976). Eurycea quadridigitata Biology 99 TIME (month) Fig. 1. Number of Eurycea quadridigitata captured during migration into and out of Rainbow Bay from 21 September 1978 through 30 December 1979. Number of individuals and rainfall plotted by weekly cumulative totals, air temperatures by weekly means. 100 Raymond D. Semlitsch and Michael A. McMillan RESULTS AND DISCUSSION Breeding migrations. —The peak terrestrial activity period for adult E. quadridigitata appears to be late July through October (Fig. 1). During the 1978 season, animals caught between 21 September and 14 October at Rainbow Bay (N = 122), and on the night of 30 September, at Ellenton Bay (N = 40), represented mature adults entering the bays. Likewise, the majority of E. quadridigitata caught from 14 July through 30 October 1979 (N = 164) were adults entering Rainbow Bay in breeding condition (males with enlarged cirri; gravid females). Since these incoming salamanders were mature and in breeding condition we assume they were entering the bays to breed. Breeding migrations during 1979 appeared to be more diffuse than during 1978. This could be due to rainfall pattern, since the autumn of 1978 was very dry, whereas rainfall occurred fre- quently during the autumn of 1979 (Fig. 1). The amount and timing of rainfall has been shown to be an important proximate factor that may determine the onset and duration of these breeding migrations of other salamanders (Baldauf 1952; Shoop 1960, 1965; Gill 1978). There was no correlation between total number of captures per week and cumulative rainfall for the week or mean minimum and maximum air temperatures (r = 0.16, p > 0.10; r = 0.09, p > 0.20; r = 0.14, p > 0.20; respectively). Fig. 2. Size-frequency distributions of breeding Eurycea quadridigitata entering Rainbow Bay and Ellenton Bay from 21 September through 30 November 1978. Eurycea quadridigitata Biology 101 However, it is clear from Fig. 1 that the initiation of breeding migrations is due to a drop in average air temperature over a several day period in late summer and early autumn. It should also be noted that breeding migrations of E. quadridigitata are probably adaptively adjusted for the autumn period since similar environmental conditions from January through June did not induce migration to breeding areas. Similar condi- tions were noted by Shoop (1960) for initiation of breeding migrations of Ambystoma talpoideum in Louisana. The breeding migration period on the SRP corresponds with that reported by Harrison (1973) for the Charleston, South Carolina area, but the onset of movement to the bays on the SRP was about a month earlier. Variation in the onset of breeding migrations can be expected because of local and annual variation in climatic factors. Data concerning the exiting of adults and/or metamorphosing juveniles during 1978 are not available since a large proportion of the breeding adult population was destructively sampled. Population size structure. — Population size structure of E. quadridigitata entering Rainbow Bay and Ellenton Bay is shown in Fig. 2. All 124 individuals of both sexes entering the bays at this time were mature and their sex ratio did not differ from 1:1 (X:= 0.48, df = 1, p > 0.40, Rainbow Bay; X2= 0.08, df = 1, p > 0.90, Ellenton Bay). The mean SVL of adults was 29.1 ± 0.20 mm (N = 74, range 24-33 mm SVL) at Rainbow Bay and 26.4 ± 0.24 mm (N = 50, range 23-31 mm SVL) at Ellenton Bay. Local variation was evident in that adults from Rainbow Bay were significantly larger than those from Ellenton Bay (t = 8.77, df = 122, p < 0.001). Similar results were noted for metamorphosing E. quadridigitata juveniles from two populations (Semlitsch, in press) and may be explained by differential growth rates of juveniles in response to water temperature and/or food availability (Shoop 1974; Stewart 1956; Wilbur 1972; Wilder 1924). Since Rainbow Bay is smaller and more shallow than Ellenton Bay, the water may warm more quickly in the spring. Larvae from Rainbow Bay may grow more rapidly and gain a growth advantage that might be maintained to maturity. Sexual dimorphism in body size was not present in E. quadridigitata since the SVL of males and females was not significantly different at Rainbow Bay and Ellenton Bay (t = 1.35, df = 72, p > 0.20; t = 0.29, df = 48, p > 0.50; respectively). Size classes other than the unimodal distribution of adults were not found entering the bays during breeding migrations (Fig. 2). Year classes among adults cannot be distinguished on the basis of size. This could be because no growth occurs after sexual maturity is reached or because E. quadridigitata is short-lived and few adults survive to breed more than once during their life. The evidence presented supports the latter explana- tion, since very few adults ever leave the bays after breeding (Fig. 1, see 1979 season.) 102 Raymond D. Semlitsch and Michael A. McMillan Table 1. A summary of female body size and reproductive output of Eurycea quadridigitata from two populations in South Carolina. Values repre- sent means ± 1 S.E. Measurements are in mm. Parameter Location Rainbow Bay Ellenton Bay Smallest SVL of Gravid Females 27.1 22.9 SVL of Females 29.4 ± 0.23 26.7 ± 0.38 N 31 24 Mean Number of Ovarian Eggs 32.7 ± 1.36 21.4 ± 1.88 Range 18-48 7-42 N 31 24 Diameter of Ovarian Eggs September 0.93 ± 0.04 0.86 ± 0.06 N 13 18 October 1.09 ± 0.06 1.43 ± 0.08 N 11 4 November 1.48 ± 0.05 1.48 ± 0.20 N 7 2 Reproduction.— All female E. quadridigitata collected entering the bays during the period 21 September through 30 November 1978 that were dissected contained enlarged ovarian eggs. The smallest SVL of females with enlarged ovarian eggs was 27.1 mm at Rainbow Bay and 22.9 mm at Ellenton Bay (Table 1). Gravid females are caught entering the bays in August through November. The diameter of ovarian eggs was similar between study sites and increased over time from 0.86 - 0.93 mm in September to 1.48 mm in November (Table 1). The largest egg diameter measured was 1.8 mm from a female caught on 15 November 1978 and is probably close to the size of eggs at deposition. Only two gravid females were caught after November. Therefore, time of egg deposition for the majority of dwarf salamanders is probably November or early December. Since egg deposi- tion is synchronous and gravid females are not found at other times of the year, clutch frequency is no more than one per year. The number of ovarian eggs was positively correlated with SVL (r = 0.67, p < 0.001, for Rainbow Bay and Ellenton Bay combined; Fig. 3). The number of ovarian eggs was significantly greater at Rainbow Bay than at Ellenton Bay (t = 5.10, df = 53, p < 0.001; Table 1) and is attributable to the significant local variation in body size between populations. Eurycea quadridigitata Biology 103 SNOUT -VENT LENGTH (mm) Fig 3. Relationship between number of ovarian eggs and snout-vent length of Eurycea quadridigitata from Rainbow Bay and Ellenton Bay. Data were pooled from both locations to calculate the regression equation. ACKNOWLEDGMENTS.— We thank Janalee P. Caldwell, Judith L. Greene, Gary B. Moran, Martha K. Nungesser, and Cynthia A. Shoemaker for their assistance collecting specimens and checking drift fences. We thank Sheri Belew and Tonya Willingham for their typing assistance and expertise. Julian R. Harrison, Janalee P. Caldwell, and J. Whitfield Gibbons critically read the manuscript. This research was sup- ported by U.S. Department of Energy Contract DE-AC09-76SR00819 with the University of Georgia (SREL). 104 Raymond D. Semlitsch and Michael A. McMillan LITERATURE CITED Baldauf, Richard. J. 1952. Climatic factors influencing the breeding migration of the spotted salamander, Ambystoma maculatum (Shaw). Copeia 1952(3): 178-181. Bishop, Sherman. C. 1947. Handbook of salamanders: the salamanders of the United States, of Canada, and of lower California. Comstock Publ. Co., Ithaca. 555 pp. Dunn, Emmett. R. 1926. The salamanders of the family Plethodontidae. Smith College 50th Anniversary Publ., Northampton. 441 pp. Gibbons, J. Whitfield. 1970. Terrestrial activity and the population dynamics of aquatic turtles. Am. Midi. Nat. 53(2):404-414. , and D. H. Bennett. 1974. Determination of anuran terrestrial activity patterns by a drift fence method. Copeia 1 974( 1 ): 236-243. Gill, Douglas E. 1978. The metapopulation ecology of the red-spotted newt, Notophthalmus viridescens (Rafinesque). Ecol. Monogr. 45:145-166. Gordon, Robert E. 1953. A population of Holbrook’s salamander, Eurycea longi- cauda guttolineata (Holbrook). Tulane Stud. Zool. / :(4)55-60. Harrison, Julian R. 1973. Observations on the life history and ecology of Eurycea quadridigitata (Holbrook). HISS News J. / (2): 57-58. Ireland, Patrick H. 1976. Reproduction and larval development of the gray- bellied salamander, Eurycea multiplicata griseogaster. Herpetologica 52(3): 233-238. Mittleman, M. B. 1947. American Caudata. I. Geographic variation in Manculus quadridigitatus. Herpetologica 3(6):209-224. 1967. Manculus and M. quadridigitatus. CAT AMER AMPHIB REPT: 44.1. Neill, Wilfred T. 1949. A series of Manculus from Georgia. Herpetologica 5:29-30. Noble, G. Kingsley. 1927. The value of life history data in the study of the evolu- tion of the Amphibia. Ann. N.Y. Acad. Sci. 30: 31-128. Odum, Eugene P. 1960. Organic production and turnover in old field succession. Ecology 47:34-49. Schalles, John F. 1979. Comparative limnology and ecosystem analysis of Carolina bay ponds on the upper Coastal Plain of South Carolina. Ph.D. Thesis, Emory Univ., Atlanta. 268 pp. Semlitsch, Raymond D. In press. Growth and metamorphosis of larval dwarf salamanders (Eurycea quadridigitata). Herpetologica. Sever, David M. 1975. Morphology and seasonal variation of the nasolabial glands of Eurycea quadridigitata (Holbrook). J. Herpetol. 9(4):337-348. Shoop, C. Robert. 1960. The breeding habits of the mole salamander, Ambystoma talpoideum (Holbrook), in southeastern Louisana. Tulane Stud. Zool. 5(3):65-82. 1965. Orientation in Ambystoma maculatum: Movements to and from breeding ponds. Science 749(3683): 558-559. 1974. Yearly variation in larval survival of Ambystoma maculatum. Ecology 55:440-444. Stewart, Margaret M. 1956. The separate effects of food and temperature differ- ences on the development of marbled salamander larvae. J. Elisha Mitchell Sci. Soc. 72:47-56. Eurycea quadridigitata Biology 105 Wilbur, Henry M. 1972. Competition, predation, and the structure of the Ambystoma-Rana sylvatica community. Ecology 55:3-21. Wilder, Inez W. 1924. The relation of growth to metamorphosis in Eurycea bislineata (Green). J. Exp. Zool. 40{ 1 ): 1 - 1 12. Accepted 28 March 1980 * Occurrence of the “Tramp” Terrestrial Amphipods Talitroides alluaudi (Chevreux) and T. topitotum (Burt) (Amphipoda: Talitridae) in South Carolina1 Charles K. Biernbaum Grice Marine Biological Laboratory, College of Charleston, 205 Ft. Johnson, Charleston, South Carolina 29412 ABSTRACT. — Populations of the terrestrial amphipods Talitroides alluaudi (Chevreux) and T. topitotum (Burt) have been found in Charleston, South Carolina. Both species are of Indo-Pacific origin, but have been transported throughout the world through commerce. Typical characteristics found in fully terrestrial talitrid amphipods are seen in these populations. INTRODUCTION Practically all coastal biologists are familiar with supralittoral talitrid amphipods, common under algae, wrack and sand near the high tide mark. However, few are familiar with the biology of those talitrid species that have become fully terrestrial. These species show a distribu- tion that is primarily tropical and Southern Hemisphere, with natural oc- currences in the Northern Hemisphere largely limited to Japan and the Indo-Malayan region, including Hawaii (Hurley 1968). Bousfield (pers. comm.) is currently studying extensive material of endemic terrestrial amphipods from Jamaica and Hispaniola, and parts of Mexico and Cen- tral America. Hurley (1968) believed that their absence from North America and Eurasia was most likely due to restrictions imposed by con- tinental climates. Their apparent absence from South America (except for a recent Brazilian introduction [Hurley 1975]) is, however, less easily explained. Although a few species are found in grasslands (Hurley 1968), most live in forest humus. In some areas they are a major component of this cryptozoic fauna; Birch and Clark (1953) found 4000m-2in an Australian forest. High susceptibility to desiccation has restricted most species to moist habitats, which, together with an inability to survive heavy frost winters (Bousfield 1968), probably explains in large part why they have been less successful in invading terrestrial ecosystems than the more desiccation-resistant isopods (Hurley 1959). 'Contribution Number 48 of the Grice Marine Biological Laboratory, College of Charleston. Brimleyana No. 3: 107-111 July 1980. 107 108 Charles K. Biernbaum In addition to natural talitrid dispersal by means of continental drift (Bousfield 1968, Hurley 1975), birds (Hurley 1959), and rafting, man has also played a part. Modern transplanting of exotic species of plants has resulted in the expanded distribution of many “tramp” (Bousfield and Howarth 1976) terrestrial species to other parts of the world. In these new locations they may be found in suitable habitats (where winters are mild) or greenhouses, where they have become a recognized component of a “hothouse fauna” including exotic species of insects, snails and isopods (Hurley 1959). RESULTS Populations of two species of terrestrial talitrid amphipods, Talitroides alluaudi (Chevreux) and T. topitotum (Burt), were found living in leaf litter and humus in Charleston, South Carolina. The population of T. alluaudi is located in a 2-4 m by 50 m strip of loose plant debris at the base of a northeast facing, residential stone wall. This habitat consists of leaf litter approximately 2 cm thick overlying dark, sandy loam, only the top 2-4 cm of which is humus rich. Leaves composing the leaf litter con- sist primarily of Quercus laurifolia (laurel oak), with additional leaves and plant debris from Tilia caroliniana (Carolina basswood), Carya il- linoensis (pecan), Prunus sp. (cherry), Ligustrum sp. (privit), Eriobotrya japonica (loquat), and Camellia japonica (camellia). Common faunal groups found in this same habitat are pulmonate snails and slugs, isopods (Armadillium sp. and Oniscus sp.), collembolans, dermapterans, ants, cockroaches, millipeds and occasionally the terrestrial turbellarian, Bipalium sp. I have collected specimens of T. alluaudi at this site since September, 1978. Their distribution within the habitat is clumped, with greatest den- sities occurring where the humus layer is thickest. A collection made in September, 1979, revealed an approximate density of 100 m -2 in such a humus rich area. Charleston specimens conform to Medcof’s (1940) pleopod description, with all specimens examined having one third pleopod consisting solely of the protopodite and the other having a but- tonlike, single outer ramal article. Males display the “mitten” second gnathopod morphology characteristic of female talitrids and males of most terrestrial species (Hurley 1959, 1968). Body lengths of Charleston specimens are approximately 5-7 mm, agreeing with lengths recorded by Shoemaker (1936) for those in California. Approximately 6 km from the population of T. alluaudi is a popula- tion of T. topitotum living in leaf litter and humus around and under a house and adjacent shed. Amphipods were found in plant debris under the house, within a strip not exceeding 0.5 m from its edge and in an ap- proximate area of 10 m2 under and surrounding the small shed. Leaves composing the leaf litter consisted of Carya illinoensis (pecan), Carya sp. (hickory), Liquidambar styraciflua (sweetgum) and Magnolia grandiflora Introduced Terrestrial Amphipods 109 (southern magnolia). Faunal groups other than amphipods found in this plant debris were the same as those listed above for T. alluaudi. I collec- ted specimens of T. topitotum at this site from August, 1974, to June, 1977, at which time a large population was present. A second population of T. topitotum was found in March, 1980, liv- ing in a relatively undisturbed forest located across from Harborview Shopping Center, approximately 3.5 km from the population described above. The approximately 20 ha mesic forest, which surrounds a lake, has a canopy (and leaf litter) consisting primarily of Quercus falcata (southern red oak), Pinus taeda (loblolly pine), and Quercus laurifolia (laurel oak). Talitroides topitotum is found in small numbers ( < l-50m-2) in the thick humus layer in various parts of the forest. Fligher densities (90-140 nv2) are present within 10 m of the edge of a small swamp adja- cent to the lake. Where the soil is saturated with water, however, T. topitotum is absent. Pleopods of Charleston specimens of T. topitotum conform to Burt’s (1934) description. Published body lengths of 5-13 mm (Shoemaker 1936) are not very different from the maximum recorded length in Charleston specimens of 10 mm. Voucher specimens of both Talitroides species have been deposited in the collections of the Canadian National Museum of Natural Sciences, Ottawa; the Grice Marine Biological Laboratory, Charleston; the North Carolina State Museum of Natural History, Raleigh (10 T. topitotum, NCSM C-323; 10 T. alluaudi, NCSM C-324); and my personal collection. DISCUSSION Both T. alluaudi and T. topitotum are of Indo-Pacific origin and un- doubtedly were transported to Charleston with exotic species of plants common in this coastal, subtropical city. They are two of only three species of terrestrial amphipods recorded from the continental United States, the third being Arcitalitrus sylvaticus (Haswell), found in Califor- nia (Bousfield and Carlton 1967). Talitroides alluaudi was first described by Chevreux (1901) from the Seychelles Islands in the Indian Ocean, where it lives in rotten trunks of coconut trees and forest humus (Shoemaker 1936). Figures of this species are in Chevreux (1901) and Reid (1947). Talitroides alluaudi, discovered in an Ohio greenhouse (Visscher and Heimlich 1930), was the first fully terrestrial amphipod found in the continental United States. Wild pop- ulations have been found on various Indian Ocean and Polynesian islands, the Canary Islands, the Azores (Hurley 1975), Hawaii (Bousfield and Howarth 1976), California (Bousfield 1975), Florida and Georgia (Bousfield unpublished). Greenhouse populations have been found in Ohio (Visscher and Heimlich 1930), New Jersey (Shoemaker 1936), Illinois, Ontario (Medcof 1939), and at least eleven European countries (Hurley 1959). Talitroides topitotum was first described by Burt (1934) from 110 Charles K. Biernbaum specimens found under a crate in a garage 80 km from the coast at 1311 m in Ceylon (Sri Lanka). Good figures of this species are in Burt (1934) and Shoemaker (1936, as Talitrus sylvaticus Haswell). The first published occurrence of T. topitotum in the United States was that by Shoemaker (1936, as Talitrus sylvaticus ) in California gardens, where it on occasion reached pest densities. If T. topitotum, T. pacificus Hurley and T. decoratus (Carl) are synonymous (Hurley 1975), T. topitotum has wild populations in India, Australia, many Indo-Pacific islands, Brazil, the Azores, Madeira (Hurley 1975) and, in the United States, California (Shoemaker 1936), Florida, the Gulf Coast (Bousfield, unpubl.) and Hawaii, where it has become the dominant amphipod species in leaf litter (Bousfield and Howarth 1976). ACKNOWLEDGMENTS.— I wish to thank Dr. E. L. Bousfield for providing additional distributional information and for confirming species identifications. Two anonymous reviewers provided criticisms which greatly improved the publication. LITERATURE CITED Birch, L. C., and D. P. Clark. 1953. Forest soil as an ecological community with special reference to the fauna. Q. Rev. Biol. 25:13-36. Bousfield, E. L. 1968. Transition to land. Discussion. Symposium on terrestrial adaptations in Crustacea. Am. Zool. 5(4):393-398. 1975. List of Talitridae. pp. 363-364 in Smith, R.I., and J. T. Carlton (eds.). Light’s manual: Intertidal invertebrates of the Central California coast. Univ. California Press, Berkeley. 716 pp. , and J. Carlton. 1967. New records of Talitridae (Crustacea: Amphipoda) from the central California coast. Bull. South. Calif. Acad. Sci. dd(4):277-284. , and F. G. Howarth. 1976. The cavernicolous fauna of Hawaiian lava tubes. 8. Terrestrial Amphipoda (Talitridae), including a new genus and species with notes on its biology. Pac. Insects 77(1): 144-1 54. Burt, D. R. R. 1934. On the amphipod genus Talitrus, with a description of a new species from Ceylon, Talitrus (Talitropsis) topitotum, sub-gen. et sp. nov. Ceylon J. Sci. (B) 75(2): 181-191. Chevreux, Ed. 1901. Mission scientifique de M. Ch. Alluaud aux lies Sechelles. Crustaces Amphipodes. Mem. Soc. Zool. Fr. 74:388-438. Hurley, Desmond E. 1959. Notes on the ecology and environmental adaptations of the terrestrial Amphipoda. Pac. Sci. 72:107-129. 1968. Transition from water to land in amphipod crustaceans. Am. Zool. 5(4):327-353. 1975. A possible subdivision of the terrestrial genus Talitrus (Crustacea Amphipoda: Family Talitridae). N. Z. Oceanogr. Inst. Rec. 2(14): 157-170. Introduced Terrestrial Amphipods 111 Medcof, J. C. 1939. Additional records of the terrestrial amphipod, Talitrus alluaudi Chevreux, in North America. Am. Midi. Nat. 22(1):216-217. 1940. Variations in the pleopod structure of the terrestrial amphipod Talitrus alluaudi Chevreux. Lloydia 2:79-80. Reid, D. M. 1947. Talitridae (Crustacea Amphipoda). Synopses of the British fauna. Linn. Soc. London 7:1-25. Shoemaker, Clarence R. 1936. The occurrence of the terrestrial amphipods, Talitrus alluaudi and Talitrus sylvaticus, in the United States. J. Wash. Acad. Sci. 26( 2):60-64. Visscher, J. P. and C. S. Heimlich. 1930. A terrestrial amphipod in the United States. Science 72:560. Accepted 26 September 1979 Some Historical Data Bearing on the Pine Barrens Treefrog, Hyla andersoni, in South Carolina E. E. Brown Box 43, Davidson, North Carolina 28036 ABSTRACT. — Hyla andersoni, the Pine Barrens Treefrog, was described in 1854 by Baird from a specimen sent apparently by Charlotte Paine of Anderson, South Carolina, a site some distance inland from the sandhills strip. The type locality has long been in question. The species has since been collected in South Carolina, but apparently not yet at a point close to Anderson. Charlotte Paine and/or her co-worker, Mrs. M. E. Daniel, were located in Anderson from 1848 through 1857 and sent Baird many specimens, including the single H. andersoni. Attempts to trace travels and routes of these women, activities of potential student collectors at the Ander- son school where they taught, and other historical aspects, have shed no further positive light on the source of the type specimen. Baird (1854) described Hyla andersoni from a single specimen sent to him from Anderson (Anderson County), South Carolina, apparently by a local teacher, Charlotte Paine. A perennial question has concerned the actual point of collection of the Anderson frog, thus the type locality of the species, especially in the absence of other South Carolina specimens (Gosner and Black 1967, Neill 1947, Wright 1932). Whereas H. andersoni is usually considered to be restricted to pockets in the pine barrens region, the town of Anderson is located well up in the Piedmont Plateau, some 70 airline miles inland from the Fall Line and perhaps 65 miles from the nearest margin of the sandhills strip. (Reduce either of these dis- tances by about 15 miles to reach only the lower border of Anderson County.) Furthermore, some early collectors paid little attention to exact locality records and individual specimens often were not labeled. Any specimens sent in by Charlotte Paine and her co-worker, Mrs. M. E. Daniel, probably were recorded at the destination as simply “from Anderson.” Although it is well known that certain typical Coastal Plain forms extend sparsely into the Piedmont here and there, especially farther south, this topic has not been adequately investigated. Folkerts (1971) reported Ambystoma tigrinum and Eurycea quadridigitata from Anderson County and the latter also from Pickens County. I have seen this Eurycea north of Greenville in Greenville County, and Scaphiopus holbrooki just south of Belton in Anderson County, and I believe Franklin Sherman had records of Cemophora coccinea in Pickens County. Although I did not see a specimen, I was told locally of an apparently reliable record of Farancia abacura from about nine miles down the Savannah River from a point opposite Anderson. While these remarks refer to typical Coastal Brimleyana No. 3: 113-117 July 1980. 1 1 3 114 E. E. Brown Plain forms, many workers would not consider Hyla andersoni to be a “typical” such form. The Fall Line, incidentally, is a convenient boundary but not necessarily a barrier. The 500-foot contour runs roughly along the Fall Line and, of course, turns inland up each river valley. Although the fact may not be significant, it is worth noting that this contour extends up the Savannah River to lower Anderson County and up the Saluda River to Ware Shoals, upper Greenwood County. I first gained some familiarity with Hyla andersoni at various points in the North Carolina sandhills during the late 1940s. The frog virtually had to be present in South Carolina. Hence, in 1950 I started looking for it there, and found it in mid-June just south of Patrick, Chesterfield County (EEB 2200-01). In early July of that year it was found in Kershaw County at a site that I again checked in 1951 (EEB 3124-5), 1952, 1965 (EEB 6529-30) and, less carefully, several times since. As the populations appeared to be small and there was no evident protection in sight for the species, it simply did not seem wise to advertise these locality records. My few South Carolina specimens of H. andersoni will be placed in the Charleston Museum and the U. S. National Museum. Any worker re- quiring the location of the Kershaw County collecting site should contact the Charleston Museum, Julian R. Harrison at The College of Charleston, or the South Carolina Wildlife and Marine Resources Department (Nongame and Endangered Species Section), Columbia. My limited forays did not yield the frog farther southwest along the sandhills strip, including the Aiken County portion of the area now con- trolled by the Savannah River Plant of the Atomic Energy Commission. Once, after a heavy night rain, I thought I heard it in Lexington County near US 1 on the Cayce side of the rivers opposite Columbia, but I could not verify this. Nor did I find the frog in, or toward, the Anderson area. In recent years, other workers have begun to encounter H. andersoni in South Carolina. With interested observers now nearer to the scene, it may eventually be possible to guess where the type specimen might have originated. Neill (1947) perhaps wisely suggested that the type locality of H. andersoni be considered unknown. Schmidt (1953) without comment designated it as the present Aiken County. Hoping that a different approach might yield helpful clues to answer the question of the type locality, I have devoted much time and effort to learning what I could concerning Charlotte Paine and the school at An- derson. I have not solved the problem of the source of the type specimen. The following gleanings comprise history rather than herpetology, but to prevent repetition of effort by other persons they should be on record. Some writers (Wright and Wright 1949, Neill 1947) seem to have mistakenly assumed that Charlotte Paine and Mrs. M. E. Daniel were one and the same person. Neill (1954:90) appeared to suggest that a Col. M. E. Daniels of Augusta may have been the husband of our Mrs. Hyla andersoni in South Carolina 115 Daniel, and that most of the Paine-Daniel specimens might have come from the Augusta area. I have not been able to identify Neill’s Col. Daniels, but details following will suggest that he could not have been associated with the Mrs. Daniel in question. Phoebe and Charlotte Paine, originally from the area of Portland, Maine (their father practiced medicine at Limerick, Windham and Port- land), but later long time residents of Carlisle, Pennsylvania, came to Spartanburg, South Carolina, about 1839 to teach (Paine 1883, Landrum 1900, Vandiver 1928). With these sisters came their niece, Mary Eliza Webb, who later married Dr. Robert M. Daniel of Spartanburg but became a widow within a few months. As Mrs. Mary E. (Webb) Daniel she, along with Charlotte Paine, in 1848 accepted a supervisory and teaching position in the newly organized Johnson Female Seminary at Anderson, South Carolina. Paine was present at Anderson from 1848 through the session beginning in 1852, thereafter apparently returning to Carlisle. Except for the 1852 session, Daniel was present from 1848 until her death in 1857 (a year when the school was ravaged by disease). While in Carlisle, both women were well acquainted with Spencer F. Baird and his wife. They obviously had promised to send Baird specimens if he would send them books. Limited correspondence be- tween them is preserved in the Smithsonian Institution archives, although several items have faded to illegibility. Charlotte Paine sent a container of specimens from Anderson to Baird at Carlisle in 1848, and another in 1849. Baird complained that the 1849 collection arrived with specimens in a very poor state of preservation (a common difficulty, as shipment re- quired months and the preservative often evaporated, or was drunk, prior to arrival at the destination). Baird moved to the Smithsonian In- stitution in 1850 and Mrs. Daniel sent another container of specimens in 1853. According to G. R. Zug and W. A. Deiss (pers. comm.) of the Smithsonian, no specimens were received during the 1850-52 period. The letters of Paine and Daniel infrequently refer to frogs and do not indicate that the women were aware of the specimen of H. andersoni when it passed through their hands. Allowing for some possible paraphrasing on my part in taking notes, C. Paine noted in a letter to Baird in July, 1848, that she “had sent a can of common specimens of reptiles, including one horn snake [presumably Farancia], furnished by a gentleman from his plantation. The little girls and boys brought in the lizards, small frogs and common snakes ... A servant brought in a basket of live frogs caught with hook and line.” In November, 1848 she wrote, “I took a great fancy to those bright little green frogs, with their rolling eyes and quick movements . . .,” and in May 1849, “Some gen- tlemen in Abbeville district have promised specimens of joint snake and horn snake, when they can be met with. These are rare and seldom found. Most Northerners disbelieve the existence of these two. Therefore I wish to send convincing proof of there being such reptiles. And the deadly power of the horn snake no one can doubt.” 116 E. E. Brown Writing to Mrs. Daniel in April 1853 Baird clearly referred to the specimen of H. andersoni : “I send today a copy of our serpent book which I trust you will look over with interest. You will see by the frequent reference to Anderson how valuable your contributions have been. “We are now at work on the frogs and other Batrachians and already have some things from Anderson. In particular there is one tree frog, the only one I ever saw, entirely new, but which, unfortunately, was so much rubbed as to be defaced. It appears to have been slender, and quite smooth above; perhaps green or olive, with a great number of small round yellowish spots on the legs. Please catch, and have caught, all the tree frogs in the country, so as to secure more of this.” In October 1853 Baird again inquired about the frog. Mrs. Daniel’s letters to Baird in 1853-54 do not indicate that she remembered the specific frog in question. To my knowledge, neither Paine nor Daniel left any helpful diary or catalogue of collections. The situation of Paine and Daniel at Anderson was a confining one, with virtually no opportunity for travel. In journeying from Pennsylvania to Anderson they came to Charleston by sea. The 135 mile railroad, said to have been the longest in the world when completed in 1833, came in from Charleston to Hamburg (just across the Savannah River from Augusta) (Phillips 1908). Its branch to Columbia was finished in 1842. The extension from there to Anderson was not completed until 1853, hence is not significant here. I am not certain about the late 1840s, but in the early days of the trip to Hamburg a traveller came in 90 miles to Blackville, then stopped for the night. At the edge of Blackville the railroad crossed an interesting “Carolina bay”. Its frog fauna includes Rana grylio, but I did not find Hyla andersoni there. Charlotte Paine stated that travel from the Augusta area to Ander- son was by wagon, or possibly by stage if a person were travelling light. She did not mention the route of travel, but rather hazy maps in Colton (1856) and Simms (1843) suggest that at least three were possible: 1) to Edgefield, then Abbeville and Anderson; 2) a route nearer the Savannah River, toward Calhoun’s Mill (apparently on Little River at the upper edge of present McCormick County) and then to Abbeville and Ander- son; 3) up the west side of the Savannah, crossing at then existent Petersburg or Vienna (just above the confluence with Broad River), thence to Calhoun’s Mill, Abbeville and Anderson. Mrs. Daniel occasionally visited friends or in-laws in Union district, apparently at or near Pacolet Mills in the edge of present Spartanburg County. However, this would have been only during December or January. The school at Anderson operated on two five-month terms, one from early February through June, and the other from early July through November. Commencement came in late July, followed by one week of “recreation”. December and January were the chief vacation period. Hyla andersoni in South Carolina 117 To speculate concerning the origin of the specimen of Hyla ander- soni, it might have been brought in by: 1) a more or less local student or other person; 2) a student coming from a more distant home at the begin- ning of a term; 3) a travelling acquaintance passing through Anderson; or 4) a parent or alumna returning to the school at commencement time or at the end of a term. My present guess is that the frog was collected some distance from Anderson and brought over many miles of rough roads stuffed in a small container of “spirit”. It was then simply dropped into the keg of specimens being accumulated by Paine and Daniel. Had this particular frog been brought in alive, it seems logical that these women would have remembered it. The Anderson school’s records were burned during a military raid in 1865. However, copies of some of the pertinent circulars or catalogs are in the Furman University Special Collections or in the University of South Carolina’s South Caroliniana Library at Columbia. I have examined the lists of students for appropriate years, especially 1848 and early 1849, in an attempt to guess who might have obtained the frog in question. Most of the students (63% to 72%) were from Anderson district, several from Abbeville district. Mary and Elizabeth Morris, Julia Horsey, and Cassandra Hewitt made the trip in from Charleston. Mary S. Coleman and Harriet Hibler were from Edgefield district. Elizabeth and Eugenia Higgins lived on the hill above the Saluda River (Newberry side) where the Saluda-Newberry highway (SC 121) now crosses. Their father ran the ferry there. Augusta G. Thompson and Jerusha Prince were respectively from Chickasaw and Tippah counties, Mississippi. Is there any chance of the frog’s being present in that northeastern portion of that state? In our present state of ignorance, all of the above in- dividuals would have to be considered possibilities. There was no student from Columbia (or north of there) until Fannie Caldwell in 1851. Any good South Carolina map of the time (I had Mitchell’s 1850 one) should show that Anderson district corresponded to present Ander- son County, but Abbeville district included present Abbeville and much of Greenwood and McCormick counties. Edgefield district included pre- sent Edgefield and Saluda, plus small portions of Greenwood, McCor- mick and Aiken counties. There was no Aiken County in 1850. One possibility is so remote that even to bring it up is questionable: Could the specimen of H. andersoni have been collected by Dr. John P. Barratt and become mixed with the Paine-Daniel specimens at the Smithsonian? Barratt’s specimens were sent to Washington from Ab- beville district. He lived a few miles south of the present town of Greenwood. According to information provided by E. D. Herd, Jr., about 1845 Barratt travelled slowly through parts of the sandhills strip and deeper into the lower Coastal Plain with M. Tuomey, state geologist. The trip was primarily a geologic one, but a person of Barratt’s biological interests would have been ever alert for unusual specimens. 118 E. E. Brown ACKNOWLEDGMENTS. —I am indebted to the following: the late A. L. Pickens and Doris Cochran for early suggestions; Daughters of the American Revolution Library (Washington); Smithsonian archives (Anita Lonnes); Cumberland County (Pennsylvania) Historical Society and Hamilton Library Association (Kathryn F. Brougher); Cumberland County (Pennsylvania) Records Office; Maine Historical Society (Elizabeth Ring); Furman University Library and Special Collections (especially R. S. Lillard); University of South Carolina South Caroliniana Library; E. Don Herd, Jr., for generously allowing me to ex- amine his unpublished study of the John P. Barratt papers; South Carolina State Archives; Anderson County Records Office; Spartanburg County Records Office; Spartanburg Herald Journal for use of microfilm. This work was supported in small part by Davidson College Faculty Research funds. LITERATURE CITED Baird, Spencer F. 1854. Descriptions of new genera and species of North Ameri- can frogs. Proc. Acad. Nat. Sci. Phila. 7:59-62. Colton, Joseph H. 1856. Atlas of the world. Vol. 1. J. H. Colton & Co., New York. Folkerts, George W. 1971. Notes on South Carolina salamanders. J. Elisha Mitchell Sci. Soc. S7(4):206-208. Gosner, Kenneth L., and I. H. Black. 1967. Hyla andersonii. CAT AMER AMPHIB REPT:54. 1-54.2. Landrum, John B. O. 1900. History of Spartanburg County. Franklin Printing Co., Atlanta, viii 4- 739 pp. Mitchell, Samuel A. 1852. A new map of South Carolina. Thomas, Copperthwait & Co., Philadelphia. ( Charlotte Observer reprint, 1967.) Neill, Wilfred T. 1947. Doubtful type localities in South Carolina. Herpeto- logica 4:75-76. 1954. Ranges and taxonomic allocations of amphibians and reptiles in the southeastern United States. Publ. Res. Div. Ross Allen’s Reptile Inst. 7(7):75-96. Paine, H. D. (ed.). 1883. Paine family records. Vol. 11. Privately printed. Phillips, Ulrich B. 1908. A history of transportation in the eastern cotton belt to 1860. Columbia Univ. Press, New York, xvii -I- 405 pp. Schmidt, Karl P. 1953. A check list of North American amphibians and reptiles. Sixth ed. Amer. Soc. Ichthyol. Herpetol., Chicago, viii -1- 280 pp. Simms, William G. 1843. Geography of South Carolina. Babcock & Co., Charleston, viii + 192 pp. Vandiver, Louise A. 1928. Traditions and history of Anderson County. Ruralist Press, Atlanta, viii -I- 318 pp. Wright, Albert H. 1932. Life-histories of the frogs of Okefinokee Swamp, Georgia. North American Salientia (Anura) No. 2. Macmillan Co., New York. 497 pp. , and A. A. Wright. 1949. Handbook of frogs and toads of the United States and Canada. Comstock Publ. Co., Ithaca, xii + 640 pp. Accepted 17 March 1980 Ambystoma mabeei Bishop (Caudata: Ambystomatidae): An Addition to the Salamander Fauna of Virginia Joseph C. Mitchell Graduate Program in Ecology, University of Tennessee, Knoxville, Tennessee 37916 1 AND S. Blair Hedges Department of Biology, George Mason University, Fairfax, Virginia 22030 ABSTRACT. — Ambystoma mabeei from Southampton County and Suf- folk, Virginia extend the known range of the species 46 km (28.6 mi.) northward from North Carolina. Juveniles of this uncommon species are described for the first time. An updated distribution map is included. Species with restricted ranges are of special interest to the biogeographer and ecologist. Their distributions, if accurately defined, may provide clues to their environmental tolerance limits. One such species is the salamander Ambystoma mabeei, known only from the Coastal Plain of North Carolina and South Carolina (Hardy and Ander- son 1970). This note reports specimens from two additional localities that significantly extend the known range of this species northward. All specimens are in the collection of the National Museum of Natural History (USNM). On 29 April 1979 an adult female A. mabeei (USNM 212212) and two recently metamorphosed juveniles (USNM 212213-14) were col- lected by Joseph C. Mitchell and Wendy H. Mitchell 7 km (4.3 mi.) northwest of Courtland, Southampton County, Virginia. On 17 May 1979 three other recently metamorphosed juveniles (USNM 212215-17) were collected at the same locality. On 16 June 1979 S. Blair Hedges found a juvenile (USNM 211210) 5 km (3. 1 mi.) south of Suffolk, City of Suffolk (formerly Nansemond County), Virginia. These specimens repre- sent the northernmost localities known and the first records of the species for Virginia. The previous northernmost site is 46 km (28.6 mi.) to the south in Perquimans County, North Carolina (Palmer et al. 1974). Figure 1 shows the updated range and all known localities of A. mabeei. The adult female from Southampton County, Virginia has the following characteristics: snout-vent length (SVL; to posterior margin of vent) 54.3 mm; total length (TL) 93.1 mm; costal grooves 13R/13L. The dorsum in life was black with scattered white flecks (most concentrated 'Mailing address: 1716 Rockwood Road, Richmond, Virginia 23226. Brimleyana No. 3: 119-121 July 1980. 119 120 Joseph C. Mitchell and S. Blair Hedges Fig. 1. Locality records for Ambystoma mabeei. Triangles indicate new localities herein reported; circles indicate literature and museum records. The dashed line represents the Fall Line. Ambystoma mabeei in Virginia 121 laterally), and brown after preservation. The venter was slate gray with some white flecking on the chin. All juveniles from this locality possess gill buds, indicating recent transformation from the larval stage. The five specimens in the series measured (mm):SVL= 36.7-46.0 (x = 42.0); TL= 60. 1-78.6 (x = 70.4). These individuals were uniform black dorsally with a single lateral row of flecking. Their ventral surfaces were uniform slate gray and their ventrolateral margins were marked with a row of white spots. The Suffolk specimen is an older juvenile (SVL = 39.5 mm, TL=63.9 mm) with no gill buds; costal grooves 13/13; concentrated lateral flecking; and brown dorsum in preservative. The Suffolk specimen was found under a pine log on the side of a sawdust pile. Here the surrounding vegetation consists mainly of Red Maple, Acer rubrum, with some Loblolly pine, Pinus taeda, and oaks, Quercus spp. The Southampton County locality is a cutover and burned pinewoods flatland with secondary growths of cane, Arundinaria spp., seedling Sweetgum, Liquidambar styraciflua, and Red Maple. Throughout this area are ditches and pools that contain varying amounts of water. All Ambystoma mabeei found at this locality were taken during the day under pieces of paper or small logs in sandy areas adjacent to water. Both localities are similar to A. mabeei habitats described by Hardy (1969). ACKNOWLEDGMENTS. -We thank Wendy H. Mitchell for field assistance and Ronald I. Crombie, Carl H. Ernst, and George R. Zug for reading the manuscript. E. E. Brown, William M. Palmer, Albert E. San- ders, and Gerald K. Williarhson supplied locality data. James A. Red- mond assisted in the location of some obscure localities. LITERATURE CITED Hardy, Jerry D., Jr. 1969. Reproductive activity, growth, and movements of Ambystoma mabeei Bishop in North Carolina. Bull. Md. Herpetol. Soc. 5(2):65-76. , and J. D. Anderson. 1970. Ambystoma mabeei. CAT AMER AMPHIB REPT:81. 1-81.2. Palmer, William M., A. L. Braswell and D. L. Stephan. 1974. Noteworthy herpetological records from North Carolina. Bull. Md. Herpetol. Soc. 7D(3):81-87. Accepted 5 March 1980 SUBSCRIPTIONS AND EXCHANGES The editors anticipate at least two issues of approximately 150 pages each annually. Rates for subscriptions for all issues appearing within the calendar year: Individual — United States $ 7.50 Individual — Foreign $10.00 Institution $12.50 Single issue purchase $ 4.50 All subscriptions must be paid in advance. Issues will be available on an exchange basis to organizations and institu- tions publishing general natural history and ecology journals or papers in a fairly regular schedule. Publications received on exchange will be placed in the State Museum’s H. H. Brimley Memorial Library. Address all subscriptions and requests for information on purchase and ex- change to Managing Editor, Brimleyana, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh, NC 2761 1. Back issues are available for $4.50 each. DATE OF PUBLICATION Brimleyana No. 2 was mailed on 30 November 1979. ERRATA The following errors appeared in Brimleyana No. 2: Page 31, lines 18 and 19 — T. toxawayi, T. howellae, and T. coweensis should appear as subspecies of T. aduncus instead of full species. Page 33, alternative 12 of the key, last line of second option — change 15 to 14. Page 94. The magnifications for Fig. 3 were in error; the caption should read — Photomicrographs of copulatory organs in sagittal section. A, Phagocata carolinensis , X42; B, Phagocata holleri, X90. Page 133. Fig. 2 was incorrectly cropped and the meter rule omitted. 123 MANUSCRIPT REVIEWERS The editor and editorial staff are indebted to the following biologists who kindly reviewed manuscripts published or submitted for publication in Brimleyana Nos. 1 and 2 (1979): Joseph R. Bailey, Duke University Donald Baird, Princeton University James R. Baker, N. C. State University E. L. Bousfield, National Museums of Canada Richard G. Bowker, Alma College Richard C. Bruce, Highlands Biological Station Donald G. Buth, University of California at Los Angeles Martha L. Crump, University of Florida David C. Culver, Northwestern University Carl H. Ernst, George Mason University David A. Etnier, University of Tennessee at Knoxville John B. Funderburg, N. C. State Museum Sidney A. Gauthreaux, Clemson University J. Whitfield Gibbons, Savannah River Ecology Laboratory Reinhard Harkema (deceased), N. C. State University Horton H. Hobbs, Jr., National Museum of Natural History Charles H. Hocutt, Appalachian Environmental Laboratory Richard Hoffman, Radford College Robert E. Jenkins, Roanoke College Robert A. Kuehne, University of Kentucky David S. Lee, N. C. State Museum Bernard S. Martof (deceased), N. C. State University John S. McIntosh, Wesleyan University Grover Miller, N. C. State University T. B. Mitchell, N. C. State University Robin M. Overstreet, Gulf Coast Research Laboratory William M. Palmer, N. C. State Museum John Parker, III, N. C. State University C. Richard Robins, University of Miami Roy T. Sawyer, University of California at Berkeley Royal D. Suttkus, Tulane University Charles Welby, N. C. State University Walter H. Wheeler, University of North Carolina at Chapel Hill Kenneth L. Williams, Northwestern State University of Louisiana Larry D. Wilson, Miami-Dade Community College 124 TABLE OF CONTENTS 1979 Number 1 Ashton, Ray E., Jr. and Alvin L. Braswell. Nest and larvae of the Neuse River Waterdog, Necturus lewisi (Brimley) (Amphibia: Proteidae) 15 Blaney, Richard M. The status of the Outer Banks Kingsnake, Lampro- peltis getulus sticticeps (Reptilia: Serpentes: Colubridae) 125 Braswell, Alvin L. and Nora M. Murdock. New records of the sala- manders Ambystoma talpoideum (Holbrook) and Hemidactylium scutatum (Schlegel) in North Carolina (Amphibia: Ambystomatidae and Plethodontidae) 135 Braswell, Alvin L. (see Ashton, Ray E., Jr.) 15 Braswell, Alvin L. (see Shelley, Rowland M.) 129 Brown, E. E. Some snake food records from the Carolinas 113 Cooper, John E. The brothers Brimley: North Carolina naturalists 1 Denoncourt, Robert F. (see Hocutt, Charles H.) 47 Filka, Marianne (see Shelley, Rowland M.) 147 Highton, Richard. A new cryptic species of salamander of the genus Plethodon from the southeastern United States (Amphibia: Plethodontidae) 31 Hocutt, Charles H., Robert F. Denoncourt and Jay R. Stauffer, Jr. Fishes of the Gauley River, West Virginia 47 Holt, Perry C. and Ann M. Weigh A new species of Xironodrilus Ellis 1918 from North Carolina (Clitellata: Branchiobdellida) 23 Howell, William M. (see Williams, James D.) 141 Iverson, John B. The female reproductive cycle in north Florida Kinosternon bciurii (Testudines: Kinosternidae) 37 McKinley, Daniel. Historical review of the Carolina Parakeet in the Carolinas 81 Murdock, Nora M. (see Braswell, Alvin L.) 135 Shelley, Rowland M. and Marianne Filka. Occurrence of the milliped Pachydesmus crassicutis incursus (Chamberlin) in the Kings Mountain region of North Carolina and the Coastal Plain of South Carolina (Polydesmida: Xystodesmidae) 147 Shelley, Rowland M., Alvin L. Braswell and David L. Stephan. Notes on the natural history of the terrestrial leech, Haemopis septagon Sawyer and Shelley (Gnathobdella: Hirudinidae) 129 Stauffer, Jay R., Jr. (see Hocutt, Charles H.) 47 Stephan, David L. (see Shelley, Rowland M.) 129 Tarplee, William H., Jr. Estimates of fish populations in two north- eastern North Carolina swamp streams 99 Weigl, Ann M. (see Holt, Perry C.) 23 125 Williams, James D. and William M. Howell. An albino sculpin from a cave in the New River drainage of West Virginia (Pisces: Cottidae) 141 Number 2 Avise, John C. (see Dickson, Gary W.) 119 Baird, Donald and John R. Horner. Cretaceous dinosaurs of North Carolina 1 Barr, Thomas C., Jr. Revision of Appalachian Trechus (Coleoptera: Carabidae) 29 Beets, James P. (see Bryant, Richard T.) 137 Bryant, Richard T., James P. Beets and Michael G. Ryon. Rediscovery of the Sharphead darter, Etheostoma acuticeps, in North Carolina (Pisces: Percidae) 137 Burgess, George H. (see Rohde, Fred C.) 97 Case, Gerard R. Cretaceous selachians from the Peedee Formation (Late Maestrichtian) of Duplin County, North Carolina 77 Dickson, Gary W., John C. Patton, John R. Holsinger and John C. Avise. Genetic variation in cave-dwelling and deep-sea organisms, with emphasis on Crangonyx antennatus (Crustacea: Amphipoda) in Virginia . 1 19 Dodd, C. Kenneth, Jr. A photographic technique to study tadpole populations 131 Holsinger, John R. (see Dickson, Gary W.) 119 Horner, John R. (see Baird, Donald) 1 Kenk, Roman. Freshwater triclads (Turbellaria) of North America. XII. Another new cave planarian from North Carolina, Phagocata carolinensis n. sp 91 Link, G. William, Jr. (see Rohde, Fred C.) 97 Norden, Beth B. and Aubrey G. Scarbrough. Nesting biology of Andrena (Larandrena) miserabilis Cresson and description of the prepupa (Hymenoptera: Andrenidae) 141 Patton, John C. (see Dickson, Gary W.) 141 Rohde, Fred C., George H. Burgess and G. William Link, Jr. Fresh- water fishes of Croatan National Forest, North Carolina, with comments on the zoogeography of Coastal Plain fishes 97 Ryon, Michael G. (see Bryant, Richard T.) 137 Scarbrough, Aubrey G. (see Norden, Beth B.) 141 126 INDEX TO SCIENTIFIC NAMES (New names in italics) Numbers 1: and 2: (1979) New Names Phagocata carolinensis 2:91-96 Plethodon websteri 1:31-36 Trechus aduncus coweensis 2:59-60 Trechus aduncus howellae 2:60-61 Trechus aduncus toxawayi 2:59 Trechus haoe 2:51-52 Trechus luculentus unicoi 2:68-69 Trechus luculentus wayahensis 2:70 Trechus nantahalae 2:72-75 Trechus schwarzi saludae 2:46-47 Trechus schwarzi scopulosus 2:46 Trechus stupkai 2:65-66 Trechus tusquitee 2:52-53 Trechus valent inei 2:62-64 Trechus vandykei pisgahensis 2:50-51 Xironodrilus bashaviae 1:23-29 Acantharcus pomotis 1:101; 2:104, 108, 112 Acer rubrum 2:143 Acris gryllus 1:116, 117 Agkistrodon contortrix 1:113, 121 piscivorus piscivorus 1:121 Alasmosaurus 2:16 Albertosaurus 2:6, 7, 25 libratus 2:9 sarcophagus 2:9 sp. 2:24 Alectrosaurus olseni 2:9 Alosa aestivalis 2:110 Ambloplites rupestris 1:62, 66 Amblyomma tuberculatum 1:118 Ambystoma opacum 1:115 talpoideum 1:135-139 Amia calva 1 : 1 0 1 ;2: 1 03, 104, 114 Anchoa mitchilli 2:110 Andrena miserabilis 2:141-146 Anguilla rostrata 1:57, 66, 101, 102; 2:103, 104, 112 Ankyrodrilus 1:24 Anolis carolinensis 1:117 Anomoeodus 2:24 Apeltes quadracus 2:110 Aphredoderus sayanus 1:101 sayanus 2:104, 108, 115, 116 Asellus brevicauda 2:124 Asteracanthus sp. 2:24 Aster sp. 2:131 Astyanax mexicanus 2:120, 124 Bairdiella chrysura 2:110 Bembidion carolinense 2:68 Betula nigra 2:131 Bidens sp. 2:131 Blarina brevicauda carolinensis 1:119, 121 Bottosaurus 2:24 Brachaelurus 2:87 Brachyrhizodus wichitaensis 2:24 Branchiobdella pulcherrima 1:23 Brevoortia tyrannus 2:110 127 Bufo americanus 2:131 houstonensis 2:136 quercicus 1:115 sp. 1:116 terrestris 1:114, 118, 120 woodhousei fowleri 1:114, 115, 116 Cambarus bartonii 1:24, 28 Campostoma anomalum 1:58, 66 Carassius auratus 2:104, 106 Carcharias 2:24, 81 Carphophis amoenus 1:117, 121 amoenus 1:116 Catostomus commersoni 1:66 Cemophora coccinea copei 1:120 Centrarchus macropterus 1:101; 2:104, 108, 112 Ceratophyllum demersum 2:99 Ceuthophilus gracilipes 2:124 Chologaster cornuta 1:101; 1:104, 107, 114 Chrysemys concinna 1:1 18 Citrus aurantium 2:141 Clinostomus funduloides 1:58, 66 Cnemidophorus sexlineatus 1:117, 118 Coelosaurus 2:25, 26 antiquus 2: 10 Colletes brimleyi 1:13 Coluber constrictor 1:113 constrictor 1:117 Condylura cristata 1:119 Conuropsis carolinensis 1:81-98 Cottus 1:65-74 bairdi 1:65, 66, 143-145 carolinae 1:141-146 ssp. 1:48, 65, 66 spp. 1:76 Crangonyx antennatus 2:120-125 Cretolamna appendiculata 2:87, 88 lata 2:82, 83, 84 biauriculata 2:82, 84, 87, 88 ?maroccana 2:84 serrata 2:82, 84, 87, 88 Cretorectolobus 2:87 Crocodylus rugosus 2:22 Crotalus horridus 1:122 atricaudatus 1:122 Cryptotis parva 1:122 Cyprinus carpio 2:104, 106 Daspletosaurus torosus 2:9 Dasyatis 2:87 Dasypus novemcinctus 2:146 Deinosuchus 2:7, 23 rugosus 2:22, 24 Deltotaria brimleardia 1:13 brimleii 1:13 Desmognathus auriculatus 1:114 brimleyorum 1:13 fuscus 1:114, 116, 117 Diadophis punctatus 1:116, 119, 121 Dina absoloni 1:131 Dolichopoda geniculata 2:124 Dormitator maculatus 2:110 Dorosoma cepedianum 2:103, 104, 112 Drosophila 2:120 Dryptosaurus 2:9, 25 aquilunguis 2:6, 7, 8 sp. 2:24 128 Edaphodon 2:24 Egertonia 2:24 Elaphe guttata guttata 1:118 obsoleta 1:113 obsoleta 1:119 quadrivittata 1:119 Elassoma evergladei 2:1 13, 114 zonatum 1:101; 2:104, 108, 113, 115 Enneacanthus chaetodon 1:101; 2:108, 111, 113, 115 gloriosus 1:101; 2:104, 108, 112 obesus 1:101; 2:104, 109, 114 Ephuta pauxilla brimleyi 1:13 Epidendrum 2:141 Ericymba buccata 1:58, 66 Erimyzon oblongus oblongus 1:101; 2:104, 107, 112 sucetta 2:104, 107, 114 Erpobdella punctata punctata 1:131 Esox americanus americanus 1:101; 2:104, 106, 112 masquinongy 1:65 niger 1:101; 2:104, 106, 112 Etheostoma 1:102 acuticeps 2:137-140 blennioides 1:63, 66 caeruleum 1:63, 66, 75 chlorobranchium 2:139 flabellare 1:63, 66 fusiforme 1:101 fusiforme 2:104, 109, 114 kanawhae 1:48, 64, 75 nigrum 1:63, 66 olmstedi olmstedi 2:104, 109, 113 osburni 1:48, 63, 74 perlongum 2: 1 12 serifferum 1:101; 2:104, 109, 111, 113 sp. 1:114 vitreum 2: 122, 1 15 Eucinostomus argenteus 2:110 Eumeces sp. 1:117, 118 Eunectes 1:122 Euphausia distinguenda 2:125 Eurycea bislineata 1:115, 116 wilderae 1:11' lucifuga 2:124 quadridigitata 1:116 Exoglossum laurae 1:48, 58, 66 Exogyra 2:3 ponderosa 2:6 Farancia erytrogramma erytrogramma 1:117 Fundulus diaphanus diaphanus 2:104, 108, 115, 116 escambiae 2:108 heteroclitus 2:110 lineolatus 1:101; 2:104, 108, 114 notti 2:108 sp. 1:114 waccamensis 2: 1 12 Gambusia affinis 1:101 holbrooki 2:104, 108, 112 Gammarus minus 2:124 Ginglymostoma 2:87 Glaucomys volans 1:119 Gobiosoma bosci 2:110 129 Godonia lasianthus 2:141 Gopherus polyphemus 1:118 Gorgosaurus 2:6, 7, 22 Gyrodus 2:24 Hadrosaurus 2:1 1 foulkii 2:18, 19, 21 tripos 2:3, 14, 19, 20, 21, 22 Haemopis septagon 1:129-134 terrestris 1:129-134 Halictus brimleyi 1:13 Helix aspersa 2:125 Hemidactylium scutatum 1:135-139 Heterandria formosa 2:114 Heterodon platyrhinos 1:116 Heterodontus 2:87 Homarus americanus 2:125 Hybodus montanensis 2:79 sp. 1 2:79, 82, 88 Hybognathus regius 2:104, 106, 115 Hybopsis dissimilis 1:65 n. sp. 2:1 15 Hydrocotyl sp. 2:99 Hyla chrysoscelis 1:114, 115, 117 crucifer 1:114 femoralis 1:116 gratiosa 1:114 sp. 1:116 Hypentelium nigricans 1:66, 114 Hypolophus 2:24, 87 Hypsibema 2: 14, 16, 25 crassicauda 2:3, 7, 10, 11, 12, 13, 15, 18, 20, 21, 24 missouriense 2:15 Ictalurus brunneus 2: 1 15 catus 2: 104, 107, 1 14 natalis 1:47, 61, 66, 76, 101, 145; 2:104, 107, 112 nebulosus 1:47, 61, 66, 76, 101; 2:104, 107, 114 platycephalus 2:115 punctatus 1:65; 2:104, 107, 115, 116 sp. 1:114 Ischyrhiza avonicola 2:87 mira 2:24, 85, 86, 87, 88 Isurus 2:24, 81 Juncus sp. 2:99, 131 Justicia americana 2:139 Kinosternon bauri 1:37-46 flavescens 1:37 leucostomum 1:37 scorpioides 1:37 subrubrum 1:37, 117 Kritosaurus 2:18 incurvimanus 2:19 notabilis 2:19 Labidesthes sicculus 1:65 Lagodon rhomboides 2:110 Lampetra aepyptera 1:47, 57, 66, 76, 78; 2:103, 104, 112, 115 Lampropeltis calligaster rhombomaculata 1:119, 120 getulus 1:119, 125 floridana 1:125-128 getulus 1:119 130 sticticeps 1:125-128 triangulum elapsoides 1:120 Leidyosuchus formidabilis 2:24 Leiostomus xanthurus 2:110 Lepisosteus osseus 2:103, 104, 112 Lepomis auritus 1:65, 101; 2:104, 109, 113 cyanellus 1:62, 66 gibbosus 1:62, 66, 101; 2:104, 109, 113 gulosus 1:101; 2:104, 109, 113 macrochirus 1:55, 62, 66, 101; 2:104, 109, 113, 125 marginatus 2:109, 111, 113, 115 microlophus 2:104, 109 punctatus 2: 1 1 1, 113, 115 Lespedeza sp. 2:131 Liquidamber styraciflua 2:141 Lonchidion 2:87 Lophorothon 2:25 atopus 2:17 Lucania parva 2: 1 10 Masticophis flagellum flagellum 1:118 Megaptera expansa 2:21 Membras martinica 2:110 Menidia beryllina 2: 1 10 extensa 2: 1 12 menidia 2: 1 10 Mesiteia 2:87 Meta menardi 2:124 Micropterus dolomieui 1:62, 66-74 punctulatus 1:62, 66-74 salmoides 1:55, 62, 66-74, 101, 114; 2:104, 109, 111, 113 Microtus pennsylvanicus 1:117, 118, 1 20- 1 22 pinetorum 1:117, 120, 121 sp. 1:119 Micrurus fulvius fulvius 1:121 Minetryma melanops 2:115 Mitsukurina owstoni 2:81 Morone americana 2:110 Moxostoma anisurum 2:104, 107, 115 collapsum 2: 107 erythrurum 1:47, 66-74, 76 macrolepidotum 2:107, 115 pappillosum 2:104, 107, 115 spp. 2:107 Mugil cephalus 2: 1 10 curema 2:1 10 Mus musculus 1:117, 119; 2:125 Myledaphus bipartitus 2:87 Myriophyllum brasiliense 2:99 Narceus 1:121 Najas sp. 2:99 Neaphaenops tellkampfii 2:124 Necturus lewisi 1:10, 15-22 maculosus 1:15, 20, 21 lewisi 1:10, 21 louisianensis 1:20, 21 punctatus 1:15, 21, 1 14 Neosaurus 2:10 missouriensis 2:14 Nerodia 1:122 erythrogaster erythrogaster 1:114 fasciata fasciata 1:117 sipedon sipedon 1:114 Niphargus spp. 2:124 131 Nocomis micropogon 1:58 platyrhynchus 1:48, 58, 66 Notemigonus crysoleucas 1:59, 66, 101; 2:104, 106, 112 Nothonotus rufilineatus 2:137 Notophthalmus 1:31 Notorhynchus 2:87 Notropis 1:48 albeolus 1:59, 66 altipinnis 2: 106, 111, 115 amoenus 2:104, 106, 114 analostanus 2:1 15 bifrenatus 2:104, 106, 112, 114 brimleyi 1:13 chalybaeus 1:101; 2:104, 106, 112 chrysocephalus 1:59, 66 cummingsae 2:104, 106, 113, 114 hudsonius 2:104, 107, 115 maculatus 2: 1 14 petersoni 2:1 13, 1 14 photogenis 1:59, 66 procne 2:104, 107 longiceps 2: 1 15 rubellus 1:59, 66 scabriceps 1:48, 59, 66, 76 scepticus 2: 1 15 sp. 1:101 spilopterus 1:60 stramineus 1:60, 66 telescopus 1:60, 66, 76 volucellus 1:60, 66 Noturus flavus 1:47, 61, 66, 76 furiosus 2:1 12 gyrinus 1:101; 2:104, 107, 112 insignis 2:104, 107, 1 12 n. sp. 2: 1 15 Nymphaea odorata 2:99 Nyssa aquatica 1:99 sylvatica 2:141 Ochrotomys nuttalli 1:122 Odontaspis 2:87 hardingi 2:83 holmdelensis 2:83 samhammeri 2:83 sp. 2:81, 82, 88 taurus 2:83 Onchopristis 2:85 Opheodrys aestivus 1:113, 117, 118, 119, 120 OphisauruS attenuatus 1:119 sp. 1:117, 121 Ornithomimus 2:10, 25 altus 2:9 sp. 2:24 Orotrechus 2:63 Orthomerus transylvanicus 2:15 Oryzomys palustris 1:119 Ostrea 2:5 Pachydesmus clarus 1:147 crassicutis 1:147 denticulatus 1:152, 153 incursus 1:147-153 retrorsus 1:148, 152 Paleogaleus 2:87 Paralbula casei 2:24 Paranomotodon 2:87 Parrosaurus 2:10, 15, 25 missouriensis 2:14 Pedinaspis brimleyi 1:13 Perea flavescens 1:65, 101; 2:104, 109, 114 Percina caprodes 1:47, 64, 66 crassa 2: 1 15 roanoka 2:104, 109, 115 evides 2:139 132 cf. maculata 1:64, 66 oxyrhyncha 1:48, 64, 66 peltata nevisense 2:112, 115 Peromyscus leucopus 1:117 sp. 1:118, 119, 121, 122 Persa palustris 2:141 Petromyzon marinus 2:103, 111 Phagocata angusta 2:91, 95 carolinensis 2:91-96 holleri 2:91, 94, 95 Phenacobius teretulus 1:48, 60, 66, 76 Philaemon 1:131 Phobosuchus 2:22 Phoxinus oreas 1:65 Pimephales notatus 1:60, 66 promelas 1:61, 66 Pituophis melanoleucus melanoleucus 1:119 Platecarpus sp. 2:24 Plethodon cinereus 1:116 clemsonae 1:10 dorsalis 1:31-36 glutinosus 1:1 14, 115, 116 jordani metcalfi 1:116 metcalfi 1:10 webster i 1:31-36 welleri 1:31, 32, 35 Plicatolamna 2:83 arcuata 2:84, 85 cf. arcuata 2:82, 84 sp. 2:88 Podostemum ceratophyllum 2:139 Polygonum punctatum 2:99 Polypodium 2:141 Polyptychodon rugosus 2:22 Pomoxis annularis 1:65 nigromaculatus 1:65, 101; 2:104, 109, 113, 115 Pontederia cordata 2:99 Potamogeton pectinatus 2:99 Proserpinaca sp. 2:99 Protamia 2:21, 24 Protoplatyrhina 2:87 Pseudacris brimleyi 1:13, 116 triseriata feriarum 1:116 Pseudemys concinna elonae 1:10 vioscana 1:10 Pseudocorax 2:87 Pseudotriton ruber schencki 1:10 sp. 1:114, 115 Ptomaphagus hirtus 2:124 spp. 2:124 Ptychotrygon 2:87 Pycnodus phaseolus 2:24 Pylodictus olivaris 1:62, 66 Python 1:122 Raja 2:87 Rana 1:121 catesbeiana 1:114, 116, 117 heckscheri 1:117 palustris 1:115 sphenocephala 1:114, 117 virgatipes 1:117 Rattus norvegicus 1:119 Regina septemvittata 1:113, 114 Reithrodontomys humulis 1:121 Rhadine subterranea 2:124 133 Rhinichthys atratulus 1:61, 66 cataractae 1:61, 66 Rhinobatos 2:87 Rhombodus binkhorsti 2:85, 87 cf. binkhorsti 2:85, 86 sp. 2:88 Ruppia maritima 2:99 Sabal palmetto 2:141 Sagittaria sp. 2:99 Salix sp. 2:131 Salmo gairdneri 1:57, 66, 145 trutta 1:57, 66 Salvelinus fontinalis 1:58, 66 Saururus cernuus 2:99 Scapanorhynchus owstoni 2:81 texanus 2:24, 80, 81, 82, 84, 88 Scaphiopus holbrooki 1:114 Sceloporus undulatus 1:117, 119 Scinceila lateralis 1:117, 120 Scirpus sp. 2:99 Sciurus carolinensis 1:119, 122 Sclerorhynchus 2:85, 87 Scoterpes copei 2:124 Scyliorhinus 2:87 Seminatrix pygaea paludis 1:115 Semotilus atromaculatus 1:61, 66, 114 Sigmodon hispidus 1:117, 119 Sigmoria latior hoffmani 1:148 Sistrurus 1:122 Solenopsis 2:145 Sparganium americanum 2:99 Spelerpes ruber schencki 1:10 Sphagnum sp. 2:99 Sphecodes sp. 2:145 “Spirobolus” 1:121 Squalicorax kaupi 2:79, 80, 82, 84, 88 pristodontus 2:24, 80, 82, 84, 88 Squalus 2:87 Sternotherus minor 1:44 Stizostedion vitreum 1:65-74 Storeria dekayi 1:115 occipitomaculata occipitomaculata 1:115 Sylvilagus floridanus 1:119 Syngnathus fuscus 2:110 Tamias striatus 1:119, 122 Tantilla coronata 1:117, 120 Taphrosphys sp. 2:24 sulcatus 2:23 Taricha 1:31 Taxodium distichum 1:99; 2:99 Thamnophis sauritus sauritus 1:116 sirtalis sirtalis 1:115 Thecachampsa 2:22 Tillandsia 2:141 Trachodon tripos 2:18 Trechus 2:29-75 aduncus 2:31, 33, 57, 59, 60, 61, 71 aduncus 2:31, 34, 39, 41, 51, 57-58, 59, 60 coweensis 2:34, 39, 41, 58 59-60 howellae 2:34, 40, 41, 58, 60-61, 70, 73 toxawayi 2:34, 39, 40, 58, 59 balsamensis 2:36, 40, 41, 62, 71 barberi 2:34, 38, 41, 50, 51, 52, 53, 54-55, 56, 57, 58, 59, 60, 61, 62, 68, 70, 71, 73 134 bowlingi 2:34, 38, 41, 49, 52, 53- 54, 56, 61, 66, 68 carolinae 2:32, 37, 41, 48-49, 50, 70 cumberlandus 2:29, 32, 37, 47-48 haoe 2:34, 37, 38, 41, 50, 51-52, 54, 61 hydropicus 2:29, 32, 42-43, 44, 49 avus 2:31, 32, 36, 42, 43 beutenmuelleri 2:31, 32, 36, 41, 42, 43-44, 49, 50 - canus 2:32, 36, 42, 44 hydropicus 2:32, 36, 37, 42-43, 44 luculentus 2:31, 36, 54, 57, 63, 67, 68, 70 luculentus 2:36, 39, 40, 41, 61, 64, 67-68, 70 unicoi 2:36, 39, 40, 41, 52, 61, 68-69, 70 wayahensis 2:36, 39, 40, 41, 45, 53, 69, 70, 73, 75 mitchellensis 2:33, 37, 41, 48, 49, 50 nantahalae 2:35, 39, 40, 41, 69, 70, 71, 72-75 nebulosus 2:36, 40, 41, 54, 62, 62, 66, 68 novaculosus 2:35, 40, 41, 54, 61, 63, 66-67, 68, 70, 73 roanicus 2:32, 37, 41, 49 rosenbergi 2:36, 40, 41, 58, 62, 70-71 satanicus 2:33, 38, 41, 57, 58 schwarzi 2:30, 31, 44-45, 47, 70 saludae 2:33, 36, 37, 46-47 schwarzi 2:33, 36, 37, 41, 44- 45, 46, 47, 51, 58 scopulosus 2:33, 36, 37, 41, 45, 46, 48, 49, 50 stupkai 2:35, 39, 40, 41, 64, 65-66 subtilis 2:34, 38, 41, 55, 56, 62, 71 talequah 2:33, 40, 41, 52, 57, 61 tennesseensis 2:29, 31, 35, 71, 72 tauricus 2:35, 40, 41, 71-72 tennesseensis 2:35, 40, 71 tonitru 2:34, 38, 41, 55-56 tuckaleechee 2:35, 39, 40, 41, 71 tusquitee 2:34, 37, 38, 41, 49, 51, 52-53, 54, 70, 73, 75 uncifer 2:33, 38, 41, 54, 56-57, 61, 62, 63, 67, 68 valentinei 2:36, 39, 40, 41, 54, 57, 61, 62-64, 65, 66, 67, 68 vandykei 2:30, 31, 34, 49, 50, 54, 71 pisgahensis 2:35, 37, 38, 41, 50-51, 52, 58 vandykei 2:35, 37, 38, 41, 49-50, 51 verus 2:30, 35, 40, 41, 61, 63, 64, 65 Trinectes maculatus 2:110 Trionyx 2:23 spp. 2:24 Tylosaurus 2:2 sp. 2:24 Typha sp. 2:99 Ulmus floridana 2:141 Umbra pygmaea 1:101; 2:103, 104, 112 Utricularia purpurea 2:99 Virginia striatula 1:113, 116, 117 valeriae valeriae 1:116 Xironodrilus appalachius 1:28 bashaviae 1; 23-29 135 dentatus 1:28 Zapus hudsonius 1:121 formosus 1:23, 24, 28 Zatomus 2:6 pulcherrimus 1:28 Xironogiton 1:24 136 INFORMATION FOR CONTRIBUTORS Submit original and two copies of manuscripts to Editor, Bnmleyana, North Carolina State Museum of Natural History, P. 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Those without such funds should so indicate in their correspondence with the Editor. This will not affect accept- ance for normal publication. Contributors who pay full page costs will be furnished 100 free reprints. Reprint order forms will be sent with galley proofs and are to be returned to the Managing Editor. On papers with more than one author, it will be the responsibility of the correspondent to assure that other authors have an opportunity to obtain reprints. Proofs are to be corrected, signed and returned to the Managing Editor within 48 hours. Changes in proofs other than type corrections will be charged to the author. CONTENTS A Late Pleistocene Vertebrate Assemblage from Edisto Island, South Carolina. Janet A. Roth and Joshua Laerm 1 The Status of Cleptoria shelfordi Loomis, with the Proposal of a New Genus in the Milliped Family Xystodesmidae (Polydesmida). Rowland M. Shelley 31 Recent Range Expansion of the Groundhog, Marmota monax, in the Southeast (Mammalia: Rodentia). Sarah S. Robinson and David S. Lee 43 Additional Records of Albinistic Amphibians and Reptiles from North Carolina. William M. Palmer and Alvin L. Braswell 49 A Distributional Checklist of the Fishes of Kentucky. Brooks M. Burr 53 New Species Groups of Pseudanophthalmus from the Central Basin of Tennessee (Coleoptera: Carabidae: Trechinae). Thomas C. Barr, Jr 85 Breeding Migrations, Population Size Structure, and Reproduction of the Dwarf Salamander, Eurycea quadridigitata, in South Carolina. Raymond D. Semlitsch and Michael A. McMillan 97 Occurrence of the “Tramp” Terrestrial Amphipods Talitroides alluaudi (Chevreux) and T. topitotum (Burt) (Amphipoda: Talitridae) in South Carolina. Charles K. Biernbaum 107 Some Historical Data Bearing on the Pine Barrens Treefrog, Hyla andersoni, in South Carolina. E. E. Brown 113 Ambystoma mabeei Bishop (Caudata: Ambystomatidae): An Addition to the Salamander Fauna of Virginia. Joseph C. Mitchell and S. Blair Hedges 119 Errata and Miscellany 123 Manuscript Reviewers 124 Table of Contents, Nos. 1 and 2 (1979) 125 Index to Scientific Names, Nos. 1 and 2 (1979) 127