6?//

7/

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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

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)

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.

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

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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.

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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:

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

<u

l—

o

E

Fig. 3. Locations of fish collections made in Kentucky, mostly between 1950 and the present. Each dot represents two

species from a site.

Kentucky Fishes

61

Anguillidae — freshwater eels

Anguilla rostrata (Lesueur). American eel. C,D,E,G,H,J,K,L.

Catadromous. Sporadic in large rivers throughout the state, although I

know of no valid records from the upper Green River.

Clupeidae — herrings

Alosa alabamae Jordan and Evermann. Alabama shad. Ana-

dromous. Formerly present in the Mississippi and Ohio rivers. Not

reported from Kentucky waters since just before 1900 although Pflieger

(1975) and Smith (1979) reported recent records from Missouri and

Illinois, respectively. Probably only enters the Kentucky part of the Mis-

sissippi River for spawning.

Alosa chrysochloris (Rafinesque). Skipjack herring. C,D,E,G,J.

Common in the main channels of the Mississippi and Ohio rivers and the

lower reaches of their major tributaries.

Dorosoma cepedianum (Lesueur). Gizzard shad. A,C,D,E,F,G,H,

J,K,L. Abundant and generally distributed throughout the state, and es-

pecially common in large rivers, lakes and reservoirs.

Dorosoma petenense (Gunther). Threadfin shad. C,D,E,F,H,J.

Generally distributed in the main channel of the Ohio River from

Louisville to its mouth (Minckley and Krumholz 1960), and the main

channels of the Mississippi, lower Cumberland, lower Tennessee and

lower Green rivers. Introduced into reservoirs throughout the state.

Hiodontidae — mooneyes

Hiodon alosoides (Rafinesque). Goldeye. A,C,D,E,F,H,J. Occa-

sional to common in the Ohio and Mississippi rivers and in large and

medium-size rivers throughout the state.

Hiodon tergisus Lesueur. Mooneye. C,D,E,F,G,H,J,K. Sporadic in

the Ohio and Mississippi rivers. Occasional in large and medium-size

rivers throughout the state, except the Big Sandy drainage.

Salmonidae — trouts

Salmo gairdneri Richardson. Rainbow trout. D,F,G,H,J,K,L. In-

troduced. Sporadic in the reservoirs and streams where it has been in-

troduced. Most stocking, with marginal success, has taken place in Lake

Cumberland (Axon 1974).

Salmo trutta Linnaeus. Brown trout. F,J. Introduced. Recently

(1977) stocked in Indian Creek (Powell and Menifee counties), Big Dou-

ble Creek (Clay County) and Trammel Fork (Allen County). However,

sampling in those streams by district fishery biologists indicates no sur-

vival (Peter W. Pfeiffer, pers. comm.).

Salvelinus fontinalis (Mitchill). Brook trout. H. Introduced. Pres-

ently known only from Martin’s Fork and Shillalah Creek, Bell County,

where it is sporadic in occurrence.

62

Brooks M. Burr

Osmeridae — smelts

Osmerus mordax (Mitchill). Rainbow smelt. Recently captured from

the Mississippi River, Carlisle County (SIUC), and now known from

several localities along the entire Mississippi River from the mouth of the

Missouri River to Louisiana (Burr and Mayden, in press; Royal D.

Suttkus, pers. comm.). It was collected only during December and

January in 1978 and 1979.

Umbridae — mudminnows

Umbra limi (Kirtland). Central mudminnow. A,B,C. A rare species

recently captured from Terrapin Creek, Graves County (SIUC), and

Clarks River, Marshall County (SIUC). Reported by Sisk (1973) from

southwestern Fulton County (MSU).

Esocidae — pikes

Esox americanus Lesueur. Grass pickerel. A,B,C,D,E,F,G,FI,J,K,L.

Common throughout the western half of the state and occasional in the

eastern half.

Esox lucius Linnaeus. Northern pike. C,J,L. Introduced. Rare and

infrequently reported from the reservoirs where it has been planted.

Status uncertain and perhaps should not be considered a part of the state

ichthyofauna.

Esox masquinongy Mitchill. Muskellunge. F,J,K,L. Rare and

sporadic in the Green, Kentucky, and Licking River drainages and

Tygarts Creek. Most populations probably are presently maintained by

stocking.

Esox niger Lesueur. Chain pickerel. A,C. Very fare in oxbow lakes

along the lower Ohio (INHS) and Mississippi (SIUC) rivers. Reported

from Clarks River, Calloway County, by Sisk (1969) but specimens ap-

parently are no longer extant. Recent specimens were discussed by Burr

and Mayden (1979).

Cyprinidae — minnows and carps

Campostoma anomalum (Rafinesque). Stoneroller. A,C,D,E,F,G,H,

J,K,L. Abundant throughout the state, being noticeably absent from

Bayou du Chien, Obion Creek, and region B.

Carassius auratus (Linnaeus). Goldfish. A,C,D,E,F,G,J,K. In-

troduced. Sporadic in large to medium-size rivers and reservoirs

throughout most of the state. Confirmed records from the Big Sandy

drainage are unavailable.

Clinostomus elongatus (Kirtland). Redside dace. K. Reported in a

manuscript list of the fishes of northeastern Kentucky (Clark 1940) from

Lick Fork of upper North Fork, Licking River, Rowan County. I have

been unable to locate the specimen(s) on which this record is based and

no one has since reported the species in Kentucky. The record could

Kentucky Fishes

63

possibly be based on a misidentification, or the species may be very rare

or extirpated here.

Clinostomus funduloides Girard. Rosyside dace. C,D,F,K,L.

Sporadic in small tributaries of the lower Tennessee (Burr and Mayden

1979, Miller 1978) and Cumberland rivers (Burr and Mayden 1979) and

several streams in region L (KFW, UL). More recently discovered in the

upper Barren River system, Barren County (TU), the upper Green river

system, Casey County (TU), and the Little Licking River (Branson 1977).

Ctenopharyngodon idella (Valenciennes). Grass carp. Introduced.

Presently known only from the main channel of the Mississippi River

where it is generally taken only by commercial Fishermen (Pflieger 1978).

Cyprinus carpio Linnaeus. Carp. A,C,D,E,F,G,FI,J,K,L. In-

troduced. Distributed throughout the state.

Ericymba buccata Cope. Silverjaw minnow. E,G,H,J,K,L. Common

from the lower Tradewater River system (Burr et al. 1980) eastward to

and including the Big Sandy River drainage.

Hemitremia flammea (Jordan and Gilbert). Flame chub. H. Re-

ported to be abundant in Big Laurel River, Laurel County (Jordan and

Brayton 1878) and also collected from Clear Fork and Wolf Creek near

Pleasant View, Whitley County (Jordan and Swain 1883). There are no

reports of this species in Kentucky waters since Jordan’s work, and it is

probably extirpated.

Hybognathus hayi Jordan. Cypress minnow. A. Occasional in Bayou

du Chien, (MSU, SIUC), Obion Creek (MSU), Mayfield Creek (INHS,

SIUC) and oxbow lakes of the Ohio River (INFIS). Considered to be

more common than formerly thought (Burr et al. 1980).

Hybognathus nuchalis Agassiz. Silvery minnow. A,B,C,D,E,F. Com-

mon in the main channel of the Ohio River from its mouth northeast to

Louisville. Abundant in the lower Green River drainage and other

medium-size streams, rivers and sloughs in the western half of the state.

A record from the Rockcastle River drainage, Rockcastle County (Bran-

son and Batch 1972) needs substantiation. Formerly known from the Big

Sandy River (Woolman 1892).

Hybognathus placitus Girard. Plains minnow. Known only from one

locality, near the mouth of the Ohio River (Pflieger 1975, Smith 1979).

Attempts to collect this species farther south in the Kentucky part of the

Mississippi River have been unsuccessful.

Hybopsis aestivalis (Girard). Speckled chub. F,H,J,K,L. Rare in the

main channel of the Ohio and Mississippi rivers. Occasional in the upper

Green, Barren, Cumberland, Kentucky, Licking and Big Sandy River

systems.

Hybopsis amblops (Rafinesque). Bigeye chub. D,E,F,G,H,J,K,L.

Occasional in clear streams throughout the eastern two-thirds of the

state. Apparently does not occur above the falls of the Cumberland

River.

64

Brooks M. Burr

Hybopsis dissimilis (Kirtland). Streamline chub. D,F,G,H,J,L. Oc-

casional in the high gradient parts of the Cumberland, Green, Barren,

Rolling Fork, Kentucky and Big Sandy River systems. I am unaware of

confirmed records from the Licking River drainage although the species

probably occurs there.

Hybopsis gelida (Girard). Sturgeon chub. Presently known from one

locality, near the mouth of the Ohio River (Pflieger 1975). Numerous at-

tempts to collect this species farther south in the Kentucky part of the

Mississippi River have been unsuccessful. It is clearly part of the Ken-

tucky fauna since it also occurs as far south in the Mississippi River as

Mississippi.

Hybopsis gracilis (Richardson). Flathead chub. Known from only

two localities, both in the main channel of the Mississippi River, where it

is uncommon.

Hybopsis insignis Hubbs and Crowe. Blotched chub. C,H. Unknown

from region C since its original description by Hubbs and Crowe (1956)

and the subsequent impoundment of the lower Tennessee River. Rare in

the upper Cumberland River where it has been taken at four localities.

Hybopsis meeki Jordan and Evermann. Sicklefin chub. Presently

known from only one locality, at the confluence of the Ohio River with

the Mississippi River (Pflieger 1975, Smith 1979). It has been taken

farther south in the Mississippi River (Bailey and Allum 1962), so is

clearly part of the Kentucky fauna.

Hybopsis storeriana (Kirtland). Silver chub. C,D,E,F,G,H,J,K,L.

Generally distributed in large and medium-size rivers throughout the

state, including the main channels of the Ohio and Mississippi rivers

where it is the most common Hybopsis.

Hybopsis x-punctata Hubbs and Crowe. Gravel chub. Known from

only three localities in the Ohio River in Campbell, Greenup and Boyd

counties (Trautman 1957); needs substantiation.

Nocomis biguttatus (Kirtland). Hornyhead chub. J. Known from

three localities in Franklin County (UMMZ, UL, CU). Lachner and

Jenkins (1971) suspected that the isolated Kentucky River population of

this species was perhaps the result of a bait or stocking introduction.

Nocomis effusus Lachner and Jenkins. Redtail chub. D,F,H.

Occasional in the Cumberland River between Cumberland Falls and the

Little River system. Rare in the upper Green River drainage where it is

known from two localities. Extensive collecting in the lower Green River

has not revealed the presence of this species, despite the suggestion by

Lachner and Jenkins (1967) that it might occur there.

Nocomis micropogon (Cope). River chub. H,J,K,L. Common in the

upper Cumberland, upper Kentucky and Licking River systems.

Occasional in region L.

Notemigonus crysoleucas (Mitchill). Golden shiner. A,B,C,D,E,

F,G,H,J. Generally distributed in the western two-thirds of the state. A

Kentucky Fishes

65

record from the Cumberland River above the falls is presumably based

on an introduction.

Notropis amnis Flubbs and Greene. Pallid shiner. C,F,H. Until

recently the only published record of this species was from Clarks River,

Marshall County (Hubbs 1951). Other records are available from

Jennings’s Creek, Warren County (SIUC), Otter Creek, Wayne County

(CU), and Green River, Hart County (UMMZ). A very rare species

which is probably nearing extinction in Kentucky.

Notropis ardens (Cope). Rosefin shiner. C,D,E,F,G,H,J,K,L.

Generally distributed from the lower Tennessee River system eastward,

avoiding lowland areas in region E. Apparently not present in the Big

Sandy drainage.

Notropis ariommus (Cope). Popeye shiner. F,G,H,J. Uncommon in

the upper Green, Barren, Rolling Fork, Cumberland and Kentucky

River drainages. No records are available from above Cumberland Falls.

Notropis atherinoides Rafinesque. Emerald shiner. A,C,D,E,F,G,

H,J,K,L. Generally distributed throughout the state.

Notropis blennius (Girard). River shiner. C,D. Common in the main

channels of the Mississippi and Ohio rivers. Rare in the lower Tennessee

and Cumberland rivers since their impoundment.

Notropis boops Gilbert. Bigeye shiner. C,D,E,F,G,H,J,K,L.

Somewhat sporadic in the western two-thirds of Kentucky where it

occurs in the Clarks, upper Pond, upper Barren, upper Green and upper

Cumberland (below the falls) River systems. More evenly distributed in

the Salt, lower Kentucky and lower Licking rivers and in the northern

portion of region L.

Notropis buchanani Meek. Ghost shiner. D,E,F,G,H,J,K. Occa-

sional in large rivers (including the Ohio River) and their major

tributaries throughout Kentucky, except in the extreme western counties

and the Big Sandy drainage. Probably most common in the lower Green

River.

Notropis camurus (Jordan and Meek). Bluntface shiner. A,B. Known

only from Terrapin Creek, Graves County (SIUC), where it is common,

and Obion Creek, Hickman County (FSU), where it is rare.

Notropis chrysocephalus (Rafinesque). Striped shiner. C,D,E,F,G,

H,J,K,L. Generally distributed throughout the state avoiding the Coastal

Plain and parts of the lower Green River drainage.

Notropis emiliae (Hay). Pugnose minnow. A,C,D,E,F. Sporadic

throughout the western half of the state.

Notropis fumeus Evermann. Ribbon shiner. A,B,C,D,E. Abundant

throughout the western third of the state (Burr et al. 1980).

Notropis galacturus (Cope). Whitetail shiner. D,H,L. Known in

Kentucky only from the Cumberland and Big Sandy River drainages. A

population exists in Red River, Todd and Logan counties, and the

species is evenly distributed in the upper Cumberland River below the

66

Brooks M. Burr

falls. Uncommon in the Big Sandy drainage where it is known only from

Pike County (KNP).

Notropis hudsonius (Clinton). Spottail shiner. Known only from a

single locality near the confluence of the Ohio and Mississippi rivers

(Pflieger 1975).

Notropis leuciodus (Cope). Tennessee shiner. F,H. Occasional to

common in the upper Barren, Green and Cumberland River drainages.

Notropis lutrensis (Baird and Girard). Red shiner. A,E. Common in

Bayou du Chien, Obion Creek, and small tributaries to the lower Ohio

River. A small population is present in the lower Tradewater River (Burr

et al. 1980). Occasional in the main channels of the Mississippi and lower

Ohio rivers.

Notropis maculatus (Hay). Taillight shiner. A. Occasional to

common in the oxbow lakes and ponds of the lower Ohio and Mississippi

rivers (Burr and Page 1975).

Notropis photogenis (Cope). Silver shiner. E,F,G,H,J,K,L. Generally

distributed throughout the eastern half of the state.

Notropis rubellus (Agassiz). Rosyface shiner. F,G,H,J,K,L.

Generally distributed throughout the eastern half of the state.

Notropis shumardi (Girard). Silverband shiner. D. Occasional to

common in the main channels of the lower Ohio (Union County

westward) and Mississippi rivers. A record from the lower Cumberland

River is based on a preimpoundment study and I doubt that the species is

still present there.

Notropis spilopterus (Cope). Spotfin shiner. A,C,D,E,F,G,H,J,K,L.

Generally distributed throughout the state, but sporadic on the Coastal

Plain.

Notropis stramineus (Cope). Sand shiner. A,G,J,K,L. Occasional to

common from Doe Run, Meade County, and eastward. Disjunct in

Mayfield Creek, Carlisle and Graves counties (Burr et al. 1980). Hoyt’s

(1979) record for this species from the Green River at Paradise was based

on N. volucellus.

Notropis telescopus (Cope). Telescope shiner. H. Known only from

region H of the Cumberland River below the falls.

Notropis umbratilis (Girard). Redfin shiner. A,B,C,D,E,G,J,K,L.

Common in the Coastal Plain. Generally distributed along the northern

border of the state from the lower Cumberland River to region L.

Notropis venustus (Girard). Blacktail shiner. A. Known from

comparatively few specimens from two localities each in Bayou du Chien

(MSU), the Mississippi River (SIUC), and the lower Ohio River (INHS).

This species is rare, although apparently suitable habitat is present, in

western Kentucky.

Notropis volucellus (Cope). Mimic shiner. A,C,D,E,F,G,H,J,K,L.

Sporadic in occurrence but generally distributed throughout the state,

usually in large to medium-size rivers.

Kentucky Fishes

67

Notropis whipplei (Girard). Steelcolor shiner. A,C,D,E,F,G,H,

J,K,L. Rare in extreme western Kentucky, and occasional to common

throughout the rest of the state.

Notropis species. Undescribed. Palezone shiner. H. A species or

subspecies of Notropis allied to N. procne has been identified from the

Little South Fork of the Cumberland River, Wayne County (Robert E.

Jenkins, pers. comm.).

Notropis species. Undescribed. Sawfin shiner. H. An undescribed

species related to N. spectrunculus occurs in the Little and Big South

Forks of the Cumberland River, Wayne County, and probably also

occurs in tributaries of Wolf River in Cumberland and Clinton counties.

Phenacobius mirabilis (Girard). Suckermouth minnow. A,B,C,E,

G,FI,J,K,L. Common in the Coastal Plain and lowland areas in region E.

Occasional to uncommon in the Salt, Kentucky, Licking, upper

Cumberland and Big Sandy River systems.

Phenacobius uranops (Cope). Stargazing minnow. F,FL Occasional

to common in the upper Barren and Green rivers. Uncommon in the

Rockcastle River.

Phoxinus cumberlandensis Starnes and Starnes. Blackside dace. H. A

recently described species endemic to the upper Cumberland River (TU,

UL, UMMZ, UT) where it is uncommon and considered endangered due

to strip mine pollution (Starnes and Starnes 1978). Several additional

populations were recently discovered in the Rockcastle and Laurel River

drainages (KNP).

Phoxinus erythrogaster (Rafinesque). Southern redbelly dace.

C,D,E,F,G,F1,J,K,L. A common inhabitant of small, springfed, upper

elevation streams throughout the eastern two-thirds of the state. A

population on the Coastal Plain was recently discovered (Freeze and

Rayburn 1977).

Pimephales notatus (Rafinesque). Bluntnose minnow. A,C,D,E,F,

G,FI,J,K,L. One of the most abundant and ubiquitous minnows in

Kentucky, occurring throughout the state and avoiding only the extreme

lowland areas in the west (e.g., Mayfield Creek, Bayou du Chien, Obion

Creek).

Pimephales promelas (Rafinesque). Fathead minnow. A,C,E,F,G,

FI,J,K,L. Sporadic in the Coastal Plain and entire Green River.

Occasional to common in the eastern half of Kentucky. Several records

may be the result of deliberate introductions.

Pimephales vigilax (Baird and Girard). Bullhead minnow.

C,D,E,F,G,H,J,K,L. Generally distributed throughout the state in large

to medium-size rivers, probably reaching its greatest abundance in the

lower Ohio and Green rivers.

Rhinichthys atratulus (Flermann). Blacknose dace. C,D,F,G,FI,

J,K,L. Occasional to sporadic throughout most of the state except the

extreme western part. Common above Cumberland Falls. The

68

Brooks M. Burr

distribution of this species in Kentucky is spotty, which most likely

reflects inadequate sampling of its preferred habitat.

Semotilus atromaculatus ( M itch ill). A,B,C,D,E,F,G,H,J,K,L.

Abundant and generally distributed in small to medium-size streams

throughout the state.

Catostomidae — suckers

Carpiodes carpio (Rafinesque). River carpsucker. A,C,D,E,G,H,

J,K,L. Generally distributed in large rivers and the lower reaches of their

major tributaries throughout Kentucky.

Carpiodes cyprinus (Lesueur). Quillback. C,D,E,G,H,J,K,L.

Generally distributed in large to medium-size rivers in the eastern half of

the state and occasional in the western half.

Carpiodes velifer (Rafinesque). Highfin carpsucker. D,H,J,K.

Sporadic throughout the eastern half of Kentucky (17 localities), and rare

in the western half (3 localities).

Catostomus commersoni (Lacepede). White sucker. A,C,D,E,F,

G,H,J,K,L. Common and generally distributed in small and medium-size

streams from Massac Creek, McCracken County, and eastward

throughout the state.

Cycleptus elongatus (Lesueur). Blue sucker. D,E,K. Rare in the main

channel of the Ohio River (10 records within the last 50 years; INHS,

UL, OSU). Rare in the lower Green, Kentucky and Licking rivers.

Formerly present in the lower Tennessee and Cumberland rivers

(Woolman 1892) before impoundment.

Erimyzon oblongus (Mitchill). Creek chubsucker. A,B,C,D,E,F,G.

Generally distributed from the lower Salt and Green rivers westward.

Erimyzon sucetta (Lacepede). Lake chubsucker. A,E. One of the

rarest species in Kentucky and presently known*from only five localities:

Obion Creek, Hickman County (KFW, UL, SIUC), Snapneck Creek,

Fulton County (SIUC), Long Falls Creek, McLean County (SIUC),

Ohio River at Paducah (Hoyt 1979), and Cypress Creek, Muhlenberg

County (KNP).

Hypentelium nigricans (Lesueur). Northern hog sucker. C,D,E,F,

G,H,J,K,L. Generally distributed from Clarks River to the Big Sandy

basin, avoiding lowland areas in region E.

Ictiobus bubalus (Rafinesque). Smallmouth buffalo. A,C,D,E,G,

J,K,L. Generally distributed in the lower reaches of large, and medium-

size rivers and reservoirs throughout the state, including the Ohio and

Mississippi rivers proper.

Ictiobus cyprinellus (Valenciennes). Bigmouth buffalo. A,C,D,E,

F,G,H,J,K. Occasional in large and medium-size rivers (and reservoirs)

throughout Kentucky, excluding the Big Sandy basin. Also present in the

main channels of the Ohio and Mississippi rivers.

Ictiobus niger (Rafinesque). Black buffalo. A,D,E. Sporadic in large

Kentucky Fishes

69

rivers and reservoirs throughout the western half of Kentucky.

Occasional in the main channels of the Ohio and Mississippi rivers.

Lagochila lacera Jordan and Brayton. Harelip sucker. H,J. Extinct.

Formerly known from five localities, one in the Kentucky River system,

Jessamine County (MCZ) and four in the upper Cumberland River

system (Woolman 1892, Kirsch 1893).

Minytrema melanops (Rafinesque). Spotted sucker. A,C,D,E,

F,G,H,J,K,L. Generally distributed throughout the state but rarely taken

in large numbers.

Moxostoma anisurum (Rafinesque). Silver redhorse. D,F,H,J,K,L.

Sporadic throughout the eastern half of Kentucky. Not taken from

region D since impoundment of the Cumberland River.

Moxostoma atripinne Bailey. Blackfin sucker. F. Endemic to the

Barren River system where it is known from nine localities, in Allen,

Barren and Monroe counties.

Moxostoma carinatum (Cope). River redhorse. D,F,H,J,K,L.

Sporadic in large and medium-size rivers throughout the eastern half of

the state. Rare in the main channel of the Ohio River.

Moxostoma duquesnei (Lesueur). Black redhorse. D,E,F,G,H,J,K,L.

Occasional throughout the eastern four-fifths of Kentucky, from the

Little River system to the Big Sandy system.

Moxostoma erythrurum (Rafinesque). Golden redhorse. A,C,D,E,

F,G,H,J,K,L. Generally distributed throughout the state, and the most

common Moxostoma in Kentucky. Uncommon, however, in the Coastal

Plain.

Moxostoma macrolepidotum (Lesueur). Shorthead redhorse. A,C,

D,F,G,H,J,K,L. Occasional throughout the state, in large and medium-

size rivers. Very rare in the Coastal Plain.

Ictaluridae — freshwater catfishes

Ictalurus catus (Linnaeus). White catfish. L. Introduced. The present

distribution of this species in Kentucky is difficult to ascertain. It has

been widely introduced in private commercial fishing lakes in Kentucky,

and planted once in Greenbo Lake, Greenup County (Clay 1975). A

single specimen from the Ohio River, Breckenridge County is known

(Clay 1975).

Ictalurus furcatus (Lesueur). Blue catfish. A,C,D,E,J,K. Occasional

in the Ohio and Mississippi rivers and the lower reaches of their major

tributaries.

Ictalurus melas (Rafinesque). Black bullhead. A,B,C,D,E,F,G,H,

J,K,L. Generally distributed throughout the state.

Ictalurus natalis (Lesueur). Yellow bullhead. A,B,C,D,E,F,G,H,

J,K,L. Generally distributed throughout the state and generally more

common than the black bullhead, with which it is often taken.

Ictalurus nebulosus (Lesueur). Brown bullhead. D,E,G,J,K. Sporadic

70

Brooks M. Burr

in large and medium-size rivers from the lower Cumberland to the

Licking River.

Ictalurus punctatus (Rafinesque). Channel catfish. A,C,D,E,F,G,

H,J,K,L. Generally distributed in sandy-gravelly sections of large and

medium-size rivers throughout Kentucky.

Noturus elegans Taylor. Elegant madtom. F,L. Previously known in

Kentucky only from the Barren and Green rivers, where it is common in

certain localities. Bauer and Branson (1979) reported this species from

the Little Sandy River system, Elliott County (EKU).

Noturus eleutherus Jordan. Mountain madtom. F,G,H,J,K,L.

Sporadic in the main channel of the Ohio River, occasional in the upper

Green, upper Cumberland, Salt, Kentucky, Licking and Big Sandy

rivers.

Noturus exilis Nelson. Slender madtom. D,F. Rare in the lower

Cumberland (KFW, UL), upper Green (KFW, UL) and Barren (KFW)

rivers. One collection from Eagle Creek, Grant County (UL), needs

substantiation.

Noturus flavus Rafinesque. Stonecat. D,F,G,H,J,K,L. Common in

the eastern half of Kentucky although apparently absent from the Big

Sandy basin. Older records are from the Ohio River, Livingston County,

(UL) and Little River, Trigg County (KFW).

Noturus gyrinus (Mitchill). Tadpole madtom. A,C,D,E,G,J.

Occasional in the Coastal Plain and the lower reaches of streams along

the northern border of the state, to the Kentucky River drainage.

Noturus hildebrandi (Bailey and Taylor). Least madtom. B. Known

only from Terrapin Creek, Graves County (SIUC) where it is taken most

frequently from sandy raceways and brush piles at night (Burr and

Mayden 1979).

Noturus miurus Jordan. Brindled madtom. A,C,D,E,F,G,H,J,K,L.

Generally distributed throughout the state, including the upper

Cumberland River system both above (Starnes and Starnes 1978) and

below the falls.

Noturus nocturnus Jordan and Gilbert. Freckled madtom.

A,C,D,E,F,J,L. Occasional in the Coastal Plain and the Green River

basin. Sporadic throughout the rest of the state with only one record

from the upper Kentucky River and two from region L.

Noturus phaeus Taylor. Brown madtom. B. Known only from

Terrapin and Powell creeks, Graves County (SIUC), where it is fairly

common in riffles with accumulated sticks and debris (Burr and Mayden

1979).

Noturus stigmosus Taylor. Northern madtom. F,G,J,K,L. Sporadic

in the eastern half of the state. Two records are from the main channel of

the lower Ohio River.

Pylodictis olivaris (Rafinesque). Flathead catfish. A,C,D,E,F,G,

H,J,K,L. Generally distributed throughout Kentucky in large and

Kentucky Fishes

71

medium-size rivers and reservoirs. Consistently taken from the Ohio and

Mississippi rivers by commercial fishermen.

Amblyopsidae — cavefishes

Amblyopsis spelaea DeKay. Northern cavefish. F,G. Occurs from

Mammoth Cave, Edmonson County, north to caves in Breckenridge

County and probably occurs in caves between those sites.

Chologaster agassizi Putnam. Spring cavefish. C,D,E,F. From

springfed streams near the Ohio River, Livingston County, through Land

Between the Lakes and Red River of the Cumberland, to Mammoth

Cave, Edmonson County. Recently captured in a small stream in the

Pond River drainage, Muhlenberg County (SIUC).

Typhlichthys subterraneus Girard. Southern cavefish. F. Known

from several caves in Barren, Edmonson, Hart, Pulaski and Warren

counties.

Aphredoderidae — pirate perches

Aphredoderus sayanus (Gilliams). Pirate perch. A,B,C,D,E,G.

Generally distributed throughout the western half of the state; two

localities in Jefferson County (UL).

Percopsidae — trout-perches

Percopsis omiscomaycus (Walbaum). Trout-perch. F,G,K,L.

Sporadic in the middle Licking River and the northern half of region L.

Two localities for Jefferson County (UL), one locality in the Green River

system, Casey County (KFW), and one locality in the Barren River

system, Allen County (Bauer and Branson 1979).

Gadidae — codfishes

Lota lota (Linnaeus). Burbot. J,K. Known from five localities in

Kentucky, all reported by Clay (1975). A rare and poorly known species

in the state. It is uncertain whether the records are based on escapees

from introduced populations or whether the species is native.

Cyprinodontidae — killifishes

Fundulus catenatus (Storer). Northern studfish. F,G,H,J. Common

in the upper Barren, Green, Salt and Cumberland rivers. Also known

from a tributary of the Kentucky River, Lincoln County (EKU,

UMMZ), and Dix River, Rockcastle County (WKU). A preimpound-

ment record from a tributary of the lower Cumberland River, Caldwell

County (KFW), needs substantiation.

Fundulus chrysotus (Gunther). Golden topminnow. A. Known only

from Open Pond (Reelfoot Lake drainage), Fulton County, where it is

uncommon (Sisk 1973).

Fundulus notatus (Rafinesque). Blackstripe topminnow. A,C,D,E,

72

Brooks M. Burr

F,G,J,K. Occasional to common in the lower Licking, Kentucky, Salt

and Green River systems. Uncommon in the Cumberland River below

Lake Cumberland. Usually replaces F. olivaceus in lowland swamps and

ditches of region A.

Fundulus notti (Agassiz). Starhead topminnow. A. Known from two

localities in extreme western Kentucky: Open Pond, Fulton County (Sisk

1973), and Murpheys Pond, Hickman County (Branson 1972).

Fundulus olivaceus (Storer). Blackspotted topminnow. A,B,C,D,E.

Abundant from the Tradewater River westward. Sporadic in the lower

Green River where the few specimens available are somewhat inter-

mediate in appearance between F. notatus and F. olivaceus. Specimens of

this species reported from Martin’s Fork, Harlan County (Clay 1975)

could not be located.

Poeciliidae — livebearers

Gambusia affinis (Baird and Girard). Mosquitofish. A,B,C,D,E,F,

G,H,J,K. Abundant and ubiquitous in the Coastal Plain, the lower

Green River and its tributaries, and the Salt River and other tributaries

along the Ohio River, where it is presumed to be native. Probably in-

troduced into the Kentucky, Licking and upper Cumberland Rivers for

mosquito control. Several records are now available from the presumed

areas of introduction.

Atherinidae — silversides

Labidesthes sicculus (Cope). Brook silverside. A,C,D,E,F,G,H,

J,K,L. Generally distributed throughout the state in or near large to

medium-size rivers, lakes and reservoirs, and bottomland swamps, lakes

and sloughs.

Menidia audens Hay. Mississippi silverside. A. Known only from the

main channel of the Mississippi River and two localities in extreme

southwestern Fulton County, where the species was first reported from

Kentucky by Sisk (1973).

Cottidae — sculpins

Cottus bairdi Girard. Mottled sculpin. J,K,L. Unevenly distributed

in the upper Kentucky and upper Licking rivers and region L. A record

from Clear Creek, Rockcastle County (Branson and Batch 1972) needs

verification.

Cottus carolinae (Gill). Banded sculpin. D,E,F,G,H,J. Generally dis-

tributed from the lower Cumberland River system eastward to and in-

cluding the Kentucky river system, avoiding lowland areas in Region E.

Percichthyidae — temperate basses

Morone chrysops (Rafinesque). White bass. A,C,D,E,F,G,H,J.

Occasional in the main channels of the Ohio and Mississippi rivers and

Kentucky Fishes

73

their major tributaries. I am unaware of any confirmed records from the

Licking, Big Sandy and Salt rivers, although the species probably occurs

in those drainages.

Morone mississippiensis Jordan and Eigenmann. Yellow bass.

A,C,D. Limited to the main channels of the lower Cumberland, lower

Tennessee, Mississippi and lower Ohio rivers, where it is uncommon.

Morone saxatilis (Walbaum). Striped bass. C,D,F,H,J,L. In-

troduced. Continually restocked in several of the manmade lakes

throughout Kentucky. I have captured escapees and seen specimens from

the main channels of the lower Ohio, lower Tennessee and Mississippi

rivers.

Centrarchidae — sunfishes

Ambloplites rupestris (Rafinesque). Rock bass. D,E,F,G,H,J,K,L.

Generally distributed throughout the eastern two-thirds of the state

avoiding lowland areas in region E. Stragglers are known from as far

west on the Ohio River as Union County.

Centrarchus macropterus (Lacepede). Flier. A,C,D,E. Sporadic to

occasional in extreme western Kentucky and the lower Green River and

its tributaries (over 10 recent records). One record from the lower Cum-

berland River drainage, Livingston County (INHS), and one from Clarks

River, Marshall County (SIUC).

Elassoma zonatum Jordan. Banded pygmy sunfish. A,B,C,E. For-

merly thought to be restricted to the Coastal Plain, where it is common in

swamps, ditches, and lowland streams. Recently discovered in Cypress

Creek, Muhlenberg County, (KNP, SIUC) and Terrapin Creek, Graves

County (SIUC). A record from West Fork Clarks River, Graves County

(Sisk 1969), needs substantiation.

Lepomis auritus (Linnaeus). Redbreast sunfish. D,H,J. Introduced.

Specimens from introduced populations are available from Little River,

Trigg County (MSU), upper Cumberland River, Bell County (KFW),

and Kentucky River, Franklin County (UL). If this species has suc-

cessfully become established in Kentucky waters, the fact has not been

confirmed.

Lepomis cyanellus Rafinesque. Green sunfish. A,B,C,D,E,F,G,H,

J,K,L. Generally distributed throughout the state.

Lepomis gibbosus (Linnaeus). Pumpkinseed. Known from only two

localities in the Ohio River in Jefferson and Campbell counties. Inclusion

of this species in the Kentucky fish list is probably based on escapees

from introductions to commercial fishing lakes and farm ponds.

Lepomis gulosus (Cuvier). Warmouth. A,B,C,D,E,F,G,H,J,K. Oc-

casional from the Licking River drainage westward, but most common

from the lower Green River basin westward.

Lepomis humilis (Girard). Orangespotted sunfish. A,C,D,E,F,G,H,

J,K,L. Occasional to common in the Coastal Plain, sporadic throughout

the rest of the state.

74

Brooks M. Burr

Lepomis macrochirus Rafinesque. Bluegill. A,B,C,D,E,F,G,H,

J,K,L. Abundant and generally distributed throughout the state.

Lepomis marginatus (Holbrook). Dollar sunfish. A,B. Known only

from Murpheys Pond, Hickman County (SIUC), where it is uncommon

(Burr and Mayden 1979), and Terrapin Creek, Graves County (SIUC),

where five specimens were recently taken.

Lepomis megalotis (Rafinesque). Longear sunfish. A,B,C,

D,E,F,G,H,J,K,L. Abundant throughout the state.

Lepomis microlophus (GUnther). Redear sunfish. A,C,D,E,F,G,

J,K,L. Sporadic and uncommon throughout the state, excluding the up-

per Cumberland River drainage. Several records from eastern Kentucky

probably are based on introductions.

Lepomis punctatus (Valenciennes). Spotted sunfish. A. Known from

seven localities in extreme western Kentucky (INHS, SIUC), where it is

taken from lowland streams and lakes (Burr and Mayden 1979) and

never appears to be common.

Lepomis symmetricus Forbes. Bantam sunfish. A. Known from six

localities in extreme western Kentucky (INHS, MSU, UL, SIUC), where

it is sometimes common (e.g., in Murphey’s Pond, Hickman County).

Micropterus coosae Hubbs and Bailey. Redeye bass. H. Introduced.

Known only from Martins Fork and tributaries, Bell County (UL).

Although the provenance of this population is unknown (Clay 1975) it

seems highly unlikely that the species is native to Kentucky.

Micropterus dolomieui Lacepede. Smallmouth bass. C,D,E,F,G,H,

J,K,L. Generally distributed in upland streams throughout the eastern

two-thirds of Kentucky. Occasional in the Land Between the Lakes area,

where it was more common before impoundment.

Micropterus punctulatus (Rafinesque). Spotted bass. A,C,D,E,F,

G,H,J,K,L. Generally distributed throughout the state except in the ex-

treme west, where it is uncommon.

Micropterus salmoides (Lacepede). Largemouth bass. A,C,D,E,

F,G,H,J,K. Generally distributed throughout the state, except for region

L.

Pomoxis annularis Rafinesque. White crappie. A,C,D,E,F,G,H,

J,K,L. Generally distributed in lakes and rivers throughout the state.

Pomoxis nigromaculatus (Lesueur). Black crappie. A,C,D,E,F,G,H,

J,K. Sporadic in lakes and rivers throughout the state except for region

L. Records indicate that it is less common than the white crappie.

Percidae — perches

Ammocrypta asprella (Jordan). Crystal darter. D,F. Known from

four old records: Cumberland River, Lyon County (FMNH), Green

River, Edmonson County (USNM), Green River, Green County

(UMMZ), and Ohio River, Greenup-Boyd counties (CAS-SU). Not

collected in Kentucky since 1929 and considered extirpated.

Kentucky Fishes

75

Ammocrypta clara Jordan and Meek. Western sand darter. F,H.

Known from two localities: Green River, Green County (Woolman

1892), and Cumberland River, Wayne County (UMMZ). Not collected

in Kentucky since 1925, when it was recorded from the part of the Cum-

berland River now impounded as Lake Cumberland. Probably extir-

pated.

Ammocrypta pellucida (Putnam). Eastern sand darter. E,F,J,K,L.

Recent collecting effort indicates that this species, which was formerly

reasonably common in eastern Kentucky, is rapidly declining in num-

bers. Probably still present in portions of the Green, Kentucky, Licking,

Big Sandy and Ohio rivers.

Ammocrypta vivax Hay. Scaly sand darter. C. Known only from two

specimens taken in a preimpoundment study of the lower Tennessee

River. The specimens are from Jonathon Creek, Marshall County

(UMMZ), and were reported by Starnes et al. (1977). Probably extir-

pated, since most of lower Jonathon Creek is now part of the impounded

waters of Kentucky Lake.

Etheostoma asprigene (Forbes). Mud darter. A,D,E. Most collec-

tions come from areas bordering the Mississippi and lower Ohio rivers.

Seven recent collections (KNP, SIUC) are available from the lower

Green River drainage. Common at some localities in extreme western

Kentucky.

Etheostoma atripinne (Jordan). Cumberland snubnose darter. D,H.

An uncommon species known from six localities in the Cumberland

River drainage in Todd, Logan, Wayne and Pulaski counties.

Etheostoma barbouri Kuehne and Small. Teardrop darter. E,F. En-

demic to the upper Barren and upper Green rivers where it is common in

small to medium-size streams.

Etheostoma bellum Zorach. Orangefin darter. F. Endemic to the up-

per Green and Barren rivers where it is common in fast, gravel and

boulder riffles.

Etheostoma blennioides Rafinesque. Greenside darter. D,E,F,G,H,

J,K,L. Generally distributed from the lower Cumberland River, Trigg

County, eastward, avoiding most of the lowlands in region E.

Etheostoma caeruleum Storer. Rainbow darter. C,D,E,F,G,H,J,K,L.

Generally distributed from the Land Between the Lakes area eastward,

avoiding the lowland areas in region E.

Etheostoma camurum (Cope). Bluebreast darter. H,J. Found only in

the upper Cumberland River drainage below Cumberland Falls, and the

upper Kentucky River drainage.

Etheostoma chlorosomum (Hay). Bluntnose darter. A,C,D,E. Com-

mon in the lower Cumberland River and westward. Three recent records

from the Tradewater River drainage (SIUC). A record from Caney

Creek, Grayson County (UL), needs substantiation.

Etheostoma cinereum Storer. Ashy darter. D,H. Known only from

76

Brooks M. Burr

the Cumberland River drainage where it is rare. Five records are

available: Red River, Logan County (UMMZ), Little South Fork of the

Cumberland River, Wayne County (UT), Buck Creek, Pulaski County

(UMMZ), Rockcastle River, Rockcastle County (KNP), and Big South

Fork of the Cumberland River, McCreary County (KNP).

Etheostoma flabellare Rafinesque. Fantail darter. C,D,E,F,G,FI,

J,K,L. Generally distributed and abundant from Clarks River eastward,

avoiding the lowland areas in region E.

Etheostoma fusiforme (Girard). Swamp darter. A. Known only from

two localities in the Reelfoot Lake drainage in extreme southwestern

Fulton County (Sisk 1973).

Etheostoma gracile (Girard). Slough darter. A,B,C,D,E. Generally

distributed and common from the lower Green River westward.

Etheostoma histrio Jordan and Gilbert. Harlequin darter. A,B,C,E.

Uncommon and sporadic in the lower portions of Obion Creek, Bayou

du Chien, and Mayfield Creek (Sisk and Webb 1976). Formerly known

from Blood River, Calloway County (UL), and Rough River, Ohio

County (Woolman 1892).

Etheostoma kennicotti (Putnam). Stripetail darter. C,D,E,F,H. Com-

mon in upper Clarks, Tradewater, upper Pond, upper Rough, upper

Green and upper Cumberland rivers, and a few small tributaries of the

Ohio River in Livingston and Crittenden counties. A record from the

Licking River drainage, Bath County (Bauer and Branson 1979), needs

verification.

Etheostoma maculatum Kirtland. Spotted darter. F,H,K. Uncom-

mon in the upper Green, upper Barren and upper Cumberland rivers.

Woolman (1892) reported this species from South Fork Licking River

where it has not been reported since.

Etheostoma microlepidum Raney and Zorach. Smallscale darter. D.

Known from only three localities in Kentucky, all from the lower Cum-

berland River drainage: Little River, Trigg County (UMMZ), Little

River, Trigg-Christian counties (UMMZ), and Red River, Logan County

(EKU).

Etheostoma microperca Jordan and Gilbert. Least darter. G. Known

from only one specimen collected in Beargrass Creek, Jefferson County

(UL). Other records reported by Clay (1975) were based on misidentifica-

tions of other species (Burr 1978). Woolman’s (1892) record for the Little

Barren River, Green County, is probably an error (Burr 1978). Recent

collecting in Beargrass Creek has not revealed the presence of this species

and it may be extirpated.

Etheostoma neopterum Howell and Dingerkus. Lollypop darter. C.

Found in Kentucky only in tributaries of the Tennessee River, where it is

abundant (Burr and Mayden 1979).

Etheostoma nigrum Rafinesque. Johnny darter. B,C,D,E,F,G,H,

J,K,L. Sporadic in distribution in the western half of the state but com-

Kentucky Fishes

77

mon and evenly distributed in the eastern half. Rare in the upper Cum-

berland River drainage (Starnes and Starnes 1979).

Etheostoma obeyense Kirsch. Barcheek darter. H. Known only from

the upper Cumberland River below Cumberland Falls, where it is oc-

casional to common in small to medium-size creeks (Page and Braasch

1976).

Etheostoma parvipinne Gilbert and Swain. Goldstripe darter. B,C.

Known from four localities in western Kentucky: Terrapin and Powell

creeks, Graves County (SIUC), and Sugar Creek (SIUC) and Billie

Branch, Calloway County (MSU). Occurrences are sporadic and at cer-

tain times of the year the species is very hard to find (Burr and Mayden

1979).

Etheostoma proeliare (Hay). Cypress darter. A,C,D. Rare to oc-

casional in sloughs and lakes that border the Mississippi and lower Ohio

rivers. Two records are available from the lower Cumberland River

drainage, Livingston County (INHS), and five from the lower Tennessee

River drainage (SIUC, UMMZ). Three of the latter were taken before

impoundment (Burr and Mayden 1979).

Etheostoma rufilineatum (Cope). Redline darter. C,D,H. Uncom-

mon to occasional in the Tennessee and Cumberland River drainages.

Somewhat disjunct in Clarks River where specimens display some

morphological differentiation from other populations.

Etheostoma sagitta (Jordan and Swain). Arrow darter. H,J. Oc-

casional in the upper Cumberland and upper Kentucky river drainages.

Etheostoma smithi Page and Braasch. Slabrock darter. D. Known

only from the lower Cumberland River drainage, where it is sometimes

common in small headwater creeks (Page and Braasch 1976).

Etheostoma spectabile (Agassiz). Orangethroat darter. A,C,

D,E,F,G,H,J,K,L. Sporadic in distribution but common. Populations of

this species in western Kentucky are limited to Mayfield Creek, Land

Between the Lakes, and upper Pond River. The species is more evenly

distributed from the Salt River eastward to the upper part of region L.

Etheostoma squamiceps Jordan. Spottail darter. D,E,F. An abun-

dant species in the lower Cumberland River and adjacent small creeks

tributary to the Ohio River in Livingston and Caldwell counties, and up-

per Pond, Barren, Nolin, Rough, and Green rivers. Specimens reported

from Bayou du Chien by Webb and Sisk (1975) are under study by other

workers.

Etheostoma stigmaeum (Jordan). Speckled darter. C,D,E,F,H. Oc-

casional to common in Clarks, upper Rough, upper Green, upper Barren

and upper Cumberland (below the falls) rivers.

Etheostoma swaini (Jordan). Gulf darter. B. Occurs only in

tributaries of Obion River, Graves County (SIUC), where it is occasional

to common (Burr and Mayden 1979).

Etheostoma tippecanoe Jordan and Evermann. Tippecanoe darter.

78

Brooks M. Burr

F,H,J,K. Uncommon in the Green River, Green County (UMMZ), the

upper Kentucky River drainage, Owsley County (UL, UMMZ), and Big

South Fork of the Cumberland River, McCreary County (KNP). Oc-

casional in the middle part of the Licking River (KFW, UL).

Etheostoma variatum Kirtland. Variegate darter. J,K,L. Generally

distributed throughout the upper Kentucky and Licking River systems,

and all of region L where it may be common in swift riffles.

Etheostoma virgatum (Jordan). Striped darter. D,H. Known only

from the Cumberland River drainage, where it is common in Red River,

Todd and Logan counties, and Buck Creek and Rockcastle River

tributaries below the falls in the upper Cumberland River.

Etheostoma zonale (Cope). Banded darter. B,E,F,G,FI,J,K,L. Un-

evenly distributed in the eastern two-thirds of the state. Specimens from

region B represent the distinctive subspecies E. z. lynceum.

Etheostoma species. Undescribed. Golden snubnose darter. C,D.

Generally distributed throughout the lower Cumberland River drainage

in Kentucky. There is one preimpoundment record from the lower Ten-

nessee River drainage (UMMZ).

Etheostoma species. Undescribed. Emerald darter. H ,J . Common in

parts of the upper Cumberland and Kentucky River drainages.

Etheostoma species. Undescribed. Lowland snubnose darter. C.

Abundant in Clarks and Blood River drainages, and there is one preim-

poundment record from the lower Tennessee River.

Etheostoma species. Undescribed. Red snubnose darter. B. Oc-

casional in Terrapin Creek, Graves County (SIUC).

Etheostoma species. Undescribed. Splendid darter. F. Endemic to

the upper Barren River system where it is common in pools and

raceways.

Etheostoma species. Undescribed. Green River snubnose darter. F.

Endemic to the upper Green River system where it is occasional to

sporadic.

Perea flavescens (Mitchill). Yellow perch. D. Sporadic and rare in

the main channel of the Ohio River; one record from the lower Cum-

berland River is probably an introduction. It is not certain whether the

individuals that occasionally appear in the Ohio River are stragglers from

native populations in other parts of the species’ range or escapees from

introduced populations.

Percina burtoni Fowler. Blotchside logperch. H. Known from one

record from Little South Fork of the Cumberland River, Wayne County

(Woolman 1892). Some of the specimens that Woolman called P.

caprodes were later reidentified as P. burtoni by Robert E. Jenkins

(Comiskey and Etnier 1972). Probably extirpated in Kentucky, although

Comiskey nearly missed capturing what he thought was this species dur-

ing his survey of fishes of the Big South Fork of the Cumberland River

(Comiskey and Etnier 1972).

Kentucky Fishes

79

Percina caprodes (Rafinesque). Logperch. A,C,D,E,F,G,H,J,K,L.

Generally distributed from Clarks River eastward. Three records exist

from the extreme lower Ohio (INHS) and Mississippi River systems

(SIUC).

Percina copelandi (Jordan). Channel darter. C,F,H,J,K,L.

Sporadically distributed and uncommon in the upper Green, upper Cum-

berland (below the falls), upper Kentucky, middle Licking and upper Big

Sandy rivers. One preimpoundment record from Blood River, Calloway

County (UMMZ), is extant.

Percina evides (Jordan and Copeland). Gilt darter. C,F,J,K,L. Un-

common in the upper Green, upper Kentucky, middle Licking and upper

Barren rivers. There is a preimpoundment record from Blood River,

Calloway County (UMMZ).

Percina macrocephala (Cope). Longhead darter. F,H,J,K,L. Spo-

radic in the upper Barren, upper Green, Kentucky, and Licking rivers

and region L. Kirsch (1893) collected this species from Little South Fork

of the Cumberland River, Wayne County, but it has not since been re-

ported from the Cumberland River system in Kentucky. This species

once was common in the upper Barren River, as indicated by the many

specimens collected by KFW personnel using rotenone (Page 1978).

Percina maculata (Girard). Blackside darter. A,C,D,E,F,G,H,J,K,L.

Generally distributed throughout the state from Clarks River eastward.

Two records from Obion Creek, Flickman County (MSU), are known.

Percina ouachitae (Jordan and Gilbert). Ouachita darter. A,C,E.

This species, formerly known in Kentucky by the name P. uranidea, is

common in Obion Creek and Bayou du Chien. Single records from Blood

River, Calloway County (UL), and Jonathon Creek, Marshall County

(UMMZ), are preimpoundment collections. Woolman (1892) reported

this species from Rough River, Ohio County (USNM), and his specimens

were verified by Williams and Etnier (1977).

Percina oxyrhyncha (Flubbs and Raney). Sharpnose darter. J,K,L.

Specimens originally labeled P. phoxocephala were recently reidentified

as P. oxyrhyncha by Thompson (1978), and the species is now known to

be present in the upper Kentucky, upper Licking and upper Big Sandy

drainages.

Percina phoxocephala (Nelson). Slenderhead darter. C,D,E,F,G,J.

Present prior to impoundment in the lower Cumberland and Tennessee

rivers. Occasional throughout the Green and Salt River systems and the

lower Kentucky River drainage (Eagle Creek). Woolman (1892) recorded

it from Mayfield Creek.

Percina sciera (Swain). Dusky darter. A,B,C,D,E,F,G,H,J,L.

Sporadic throughout the entire state except the Licking River system,

from where I have been unable to locate specimens.

Percina shumardi (Girard). River darter. C,D,E,F,J,K. Sporadic and

uncommon in the main channel of the Mississippi and Ohio rivers and

80

Brooks M. Burr

the lower Tennessee, upper Green, upper Kentucky and middle Licking

rivers. Woolman’s (1892) records of this species from the lower Cum-

berland and lower Rough rivers have not been confirmed by recent

collecting efforts in these areas, but are considered valid.

Percina squamata (Gilbert and Swain). Olive darter. H. Known from

five localities in Rockcastle River, Rockcastle and Jackson counties

(EKU, UMMZ), and Big South Fork of the Cumberland River,

McCreary County (KFW; specimens discarded). Clay’s (1975) record of

this species from the Tennessee River, Marshall County, needs verifica-

tion, but I have been unable to locate the specimens on which the record

was based.

Percina species. Undescribed. Blackfin darter. F,J. This species was

long known under the name P. cymatotaenia, but is an undescribed

species common in the upper Barren, upper Green and upper Kentucky

River systems. The report of this species from Obion Creek (Smith and

Sisk 1969) was based on P. sciera.

Stizostedion canadense (Smith). Sauger. A,C,D,E,F,G,FI,J,K,L. Oc-

casional in the main channels of the Ohio and Mississippi rivers and the

lower reaches of their major tributaries.

Stizostedion vitreum (Mitchill). Walleye. C,D,E,FI,K,L. Sporadic

and uncommon in the main channels of the Ohio and Mississippi rivers

and the Tennessee, Cumberland, Licking and Big Sandy rivers.

Sciaenidae — drums

Aplodinotus grunniens Rafinesque. Freshwater drum. A,C,D,E,F,

G,FI,J,K,L. Generally distributed throughout the Ohio and Mississippi

rivers and their major tributaries; occasional in medium-size rivers

throughout the state.

PROBLEMATICAL LIST OF SPECIES

The five species in the following list of hypothetical additions to the

Kentucky fauna represent one family and three genera.

Cyprinidae — minnows and carps

Campostoma oligolepis Elubbs and Greene. Largescale stoneroller.

There is little question that this species (probably a new subspecies) oc-

curs in Kentucky in the lower Tennessee, lower Cumberland and upper

Green River drainages. Elowever, until a thorough study of variation is

completed I have chosen to refer to stonerollers in Kentucky as C.

anomalum.

Notropis coccogenis (Cope). Warpaint shiner. Recorded by

Woolman (1892) from the big Sandy River, but the specimens have not

been located and the record is probably an error (Gilbert 1964). A

specimen from Cumberland Gap, Bell County, collected by Walter

Faxon in 1875, is available (UMMZ). Except for Woolman’s record and

Kentucky Fishes

81

Faxon's specimen, the species has been considered endemic to the Ten-

nessee River drainage.

Notropis heterolepis Eigenmann and Eigenmann. Blacknose shiner.

Reported by Turner (1959) from Rough River, but no specific locality

was given and no specimens from the state are known to exist. Since N.

boops is present in the same area and is superficially similar, I assume the

record of N. heterolepis was actually based on N. boops. However, there is

a relict population of N. heterolepis in central Tennessee, so it possibly

occurs or once did occur in Kentucky. Until a verified record is available,

however, it seems best to delete it from the state faunal list.

Notropis hubbsi Bailey and Robison. Bluehead shiner. Known from

adjacent southern Illinois in Wolf Lake, Union County (Bailey and

Robison 1978, Smith 1979), and should be looked for in lowland lakes in

western Kentucky.

Rhinichthys cataractae (Valenciennes). Longnose dace. Known from

the Cumberland River in adjacent Tennessee where it is sporadically dis-

tributed. Possible in the upper Cumberland and Big Sandy River

drainages, Kentucky.

ACKNOWLEDGMENTS.— This list could not have been completed

without the efforts of many persons who over the years have

collected fishes in Kentucky. I am especially grateful to Lawrence M.

Page, Philip W. Smith, Marvin E. Braasch, Richard L. Mayden, Michael

E. Retzer, Steven D. Ogle, and Patti A. Burr for assistance in Field work.

I am grateful to the following curators and their institutions for

granting specimen loans, providing laboratory space, assisting with

locality records, and other courtesies: Tomio Iwamoto (CAS); Robert K.

Johnson (FMNH); Camm Swift, formerly at FSU; Philip W. Smith and

Lawrence M. Page (INHS); Benjie Kinman and Ronald R. Cicerello

(KFW); Donald Harker and Melvin L. Warren, Jr. (KNP); Marvin E.

Braasch, formerly at KU; Karsten Hartel (MCZ); Donald Johnson

(MSU); Neil H. Douglas (NLU); Royal D. Suttkus (TU); William D.

Pearson (UL); Reeve M. Bailey and Ellie Koon (UMMZ); A. P. Blair

and Hague Lindsey (UTULSAC); and Robert D. Hoyt (WKU).

The drainage maps were adapted by myself and Karen Schmidtt

from several base maps purchased from the Kentucky Department of

Commerce, Frankfort. The county map was compiled by the Kentucky

Department of Commerce, Frankfort (1964, Base Map Series B-5).

For critical review of the manuscript I am grateful to Drs. Philip W.

Smith and Lawrence M. Page, Illinois Natural History Survey.

82

Brooks M. Burr

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Cumberland. Ky. Dep. Fish Wildl. Resour. Fish. Bull. (50:1-69.

Bailey, Reeve M., and M.O. Allum. 1962. Fishes of South Dakota. Misc.

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, and F.B. Cross. 1954. River sturgeons of the American genus

Scaphirhynchus: characters, distribution, and synonymy. Pap. Mich.

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, and H.W. Robison. 1978. Notropis hubbsi, a new cyprinid fish

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from the United States and Canada. 3rd. ed. Am. Fish. Soc. Spec.

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Barbour, Roger W. 1951. Occurrence of the paddlefish in eastern Kentucky.

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1963. A record of the alligator gar in Kentucky. Trans. Ky.

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Bauer, Bruce H., and B.A. Branson. 1979. Distributional records for and

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Branson, Branley A. 1972. Fundulus notti in Kentucky. Trans. Ky. Acad. Sci.

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1977. Threatened fishes of Daniel Boone National Forest,

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Burr, Brooks M. 1978. Systematics of the percid fishes of the subgenus

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, and R.L. Mayden. 1979. Records of fishes in western Kentucky

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Call, R.E. 1896. Fishes and shells of the Falls of the Ohio. Memorial

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Clark, Minor E. 1940. A list of the fishes in northeastern Kentucky.

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Comiskey, Charles E., and D.A. Etnier. 1972. Fishes of the Big South Fork

of the Cumberland River. J. Tenn. Acad. Sci. 47(4): 140-145.

Evermann, Barton W. 1902. Description of a new species of shad, (Alosa

Kentucky Fishes

83

ohiensis ), with notes on other food fishes of the river. Rep. U.S. Fish.

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1918. The fishes of Kentucky and Tennessee: a distributional

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Garman, H. 1894. A preliminary list of the vertebrate animals of Kentucky.

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Gilbert, Carter R. 1961. Hybridization versus intergradation: an inquiry

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fauna, with two subspecies. Occas. Pap. Mus. Zool. Univ. Mich.

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, and W.R. Crowe. 1956. Preliminary analysis of the American

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the Cumberland, Whitley County, Ky., with descriptions of three

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U.S. Fish. Comm. 77:257-268.

Lachner, Ernest A., and R.E. Jenkins. 1967. Systematics, distribution, and

evolution of the chub genus Nocomis (Cyprinidae) in the southwestern

Ohio River basin, with the description of a new species. Copeia 1967

(3):557-580.

, and 1971. Systematics, distribution, and evolution

of the Nocomis biguttatus species group (Family Cyprinidae: Pisces)

with a description of a new species from the Ozark Upland. Smith-

son. Contrib. Zool. 97:1-28.

Miller, Lewis G. 1978. New distributional records for the rosyside dace in

Kentucky. Trans. Ky. Acad. Sci. 39(3-4): 142-144.

Minckley, W.L., and L.A. Krumholz. 1960. Natural hybridization between

the clupeid genera Dorosoma and Signalosa, with a report on the

distribution of S. petenensis. Zoologica 44(4): 171- 180.

Page, Lawrence M. 1978. Redescription, distribution, variation and life

history notes on Percina macrocephala (Percidae). Copeia 1978(4):

655-664.

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Brooks M. Burr

, and M.E. Braasch. 1976. Systematic studies of darters of the

subgenus Catonotus (Percidae), with the description of a new species

from the Duck River system. Occas. Pap. Mus. Nat. Hist. Univ.

Kans. 63: 1-18.

Pflieger, William L. 1975. The fishes of Missouri. Mo. Dep. Conserv.,

Jefferson City. 343 pp.

1978. Distribution and status of the grass carp ( Ctenopharyng -

odon idella) in Missouri streams. Trans. Am. Fish. Soc. 707(1): 1 13-1 18.

Rafinesque, Constantine S. 1820. Ichthyologia ohiensis. W.G. Hunt, Lex-

ington. 90 pp.

Sisk, Morgan E. 1969. The fishes of west Kentucky. I. Fishes of Clark’s

River. Trans. Ky. Acad. Sci. 20(3-4):54-59.

1973. Six additions to the known piscine fauna of Kentucky.

Trans. Ky. Acad. Sci. 24(3-4):49-50.

, and D.H. Webb. 1976. Distribution and habitat preference of

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Smith, Philip W. 1965. A preliminary annotated list of the lampreys and

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Turner, William R. 1959. Pre-impoundment surveys of six Kentucky streams.

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A ccepted 25 A pril 1 980

New Species Groups of Pseudanophthalmus from the Central

Basin of Tennessee (Coleoptera: Carabidae: Trechinae)

Thomas C. Barr, Jr.

School of Biological Sciences,

University of Kentucky, Lexington, Kentucky 40506

ABSTRACT. —Two new species groups of Pseudanophthalmus are

proposed. The simplex group includes P. simplex n. sp. and P. fowlerae

n. sp. The cumberlandus group includes P. cumberlandus Valentine, P.

productus n. sp., and P. inquisitor n. sp.; the following are full species

and also belong to the cumberlandus group: tiresias, catherinae, insularis,

occidentalis, acherontis, tullahoma, and bendermani. All species dis-

cussed occupy caves in the Central Basin of Tennessee.

Cave carabids of the very large genus Pseudanophthalmus Jeannel

(about 200 species) have traditionally been arranged in “species groups”

— neutral categories of supposedly monophyletic species assemblages

which are of lower rank than subgenera. The comparative homogeneity

of the genus does not readily permit division into clearly distinguishable

subgenera, but within certain limits there is substantial diversity at the

species group level. In preparing a new classification of Pseudan-

ophthalmus I was unable to assign two undescribed species from Jackson

and Clay counties, in the upper Cumberland River basin, Tennessee, to

an existing group. These species are described in the present paper and

placed in the new simplex group. Two other species with which they are

sympatric are close to P. cumberlandus Valentine, which I have made the

type species of a widely distributed new group, the cumberlandus group.

This group, occurring throughout the Central Basin of Tennessee and

with species in southwestern and southeastern Kentucky, the western

Tennessee River valley in Tennessee, and one county in north Alabama,

is the largest species group in the genus, including about 30 species.

simplex group (new group)

Size medium (4.0-5. 1 mm); robust, depressed; moderately pubescent

to nearly glabrous; elytral microsculpture a fine, transverse meshwork,

with hint of pruinosity in one species. Pronotum transverse, hind angles

right or nearly right. Labrum doubly emarginate. Anterior discal

puncture at level of 4th umbilicate; apical groove subparallel or slightly

oblique to suture, connected obliquely to 3rd stria in advance of anterior

apical puncture, or running to this puncture via prominent crosier;

humeri finely serrulate. Mentum tooth broad, emarginate. Mesosternum

Brimleyana No. 3: 85-96 July 1980.

85

86

Thomas C. Barr, Jr.

declivous. Last abdominal sternite of males with apical margin entire.

Aedeagus scarcely arcuate, basal bulb not conspicuously enlarged nor

deflexed, apex gradually attenuate and bluntly rounded at tip, only feebly

produced; transfer apparatus with rather heavily sclerotized right piece,

medially concave, apex knobbed or not, and broadly triangular, mem-

branous, spiny left piece with apex spatulate and twisted 90°, or right

piece as described and left piece absent; parameres moderately slender,

bearing 4 long setae at their apexes. Type species: P. simplex , new species.

Discussion.— This group consists of only two known species that occur

in caves of Jackson and Clay counties, Tennessee, at the northeast edge

of the Central Basin, in Ordovician limestones of the upper Cumberland

River drainage, on the south (left) side of the river. The somewhat

isolated troglobitic crayfish species Orconectes incomptus (Hobbs and

Barr 1972) also occurs in caves of this region.

The form of the copulatory sclerites and the serrulate humeri suggest

a close relationship with the pubescens group (see Barr 1979); very faint

pruinose microsculpture on the elytra of one species is additional

evidence favoring this view. Different features, however, require the es-

tablishment of a distinct group: depressed, robust form; more posterior

position of the anterior discal puncture (pleomorphic); reduced

pubescense and reduced pruinose microsculpture (probably apo-

morphic); and the general form of the aedeagus. The transfer apparatus

in simplex itself is similar in basic pattern to that of the pubescens group.

One can speculate that the simplex group may be a link between the

pubescens and menetriesi groups on the one hand and the numerous,

small to medium species of the cumberlandus group on the other. In the

cumberlandus group there is a single copulatory piece, presumably the left

one; its simple structure, together with the apparent loss of the right

piece, deprive the phylogenist of a key character in inferring relationships

of the group to other Pseudanophthalmus species groups. The left

copulatory piece in simplex is close enough to the cumberlandus transfer

apparatus that a relationship is at least feasible. Habitus and geographic

distribution of the simplex group, although weaker evidence supporting a

relationship to the cumberlandus group, are at least consonant with an

hypothesis that the two groups are phylogenetically close.

Pseudanophthalmus simplex, new species

Figs. 1, 2

Etymology.— Latin simplex, “simple.”

Diagnosis.— With the characters of the group as defined above; two

copulatory pieces present; elytral striae deeper, impunctate; elytral

microsculpture with a few pruinose patches near center of disc, which is

moderately pubescent; apical groove longer.

New Beetle Species Groups

87

Figs. 1-3. Pseudanophthalmus: simplex group. 1. P. simplex, n. sp. (4.6 mm). 2.

Aedeagus, P. simplex n. sp. (0.86 mm). 3. Aedeagus, P.fowlerae, n. sp. (0.68 mm).

Description. — Length 4.2-5. 1, mean 4.6 mm. Form robust and

depressed; elytral microsculpture finely transverse, forming meshes, with

trace of pruinose microsculpture near middle of disc, scarcely discernible

(50X). Head rounded, as wide as long; labrum distinctly trilobed.

Pronotum transverse, 0.8 as long as wide, disc feebly convex and very

sparsely pubescent; sides strongly arcuate in apical 0.8, subparallel in

basal 0.2; front angles rounded, apex about 0.85 maximum width, which

occurs at apical 0.20-0.22, base width nearly equal to maximum width,

hind angles large and more or less right or slightly acute, depending on

development of lateral emargination of base and secondary angles, which

are variably developed but always present in some form. Elytra more

than 1.5 times as long as wide, depressed, moderately pubescent; humeri

angular, somewhat obsoletely serrulate, prehumeral border not quite per-

pendicular to midline; inner 4 longitudinal striae moderately impressed,

intervals convex, outer striae obsolete; apical groove subparallel or

slightly oblique to suture, running to third stria in advance of anterior

apical puncture or running to anterior apical puncture via prominent

crosier. Aedeagus 0.78-0.89, mean 0.83 mm long; feebly arcuate, basal

bulb not appreciably expanded and not deflexed, apex gradually atten-

uate, scarcely produced, rounded at tip; right copulatory piece a more or

88

Thomas C. Barr, Jr.

less hyaline hemisheath, its apex a bluntly rounded knob, left piece a little

shorter, broadly triangular, densely but minutely spiny, with apex twisted

90° to right and thus appearing as narrow, nipplelike or broadly

spatulate, depending on orientation; parameres rather slender, with 4

apical setae.

Type series. — Holotype male (American Museum of Natural

History) and one female paratype, Carter Cave, 5.5 km ssw of Flynns

Lick, Jackson Co., Tennessee, 23 October 1960, T. C. Barr. Two ad-

ditional paratypes from same cave, 9 June 1955 and 20 July 1957, T. C.

Barr. Three paratypes from Hailes Cave, 0.5 km n of Flynns Licks, 6

May 1959, T. C. Barr, and 19 October 1948, J. M. Valentine and W. B.

Jones. (See Barr 1961 for detailed locations and descriptions of these and

other caves mentioned in this paper.)

Measurements (in mm).— Holotype male, total length 4.60, head

0.80 long X 0.80 wide, pronotum 0.90 long X 1.10 wide, elytra 2.66 long

X 1.72 wide, antenna 3.06, aedeagus 0.86 long.

Distribution. — The material seen of this species includes the type

series of 7 specimens and one specimen (not a paratype) from Cherry

Cave, northeast Jackson County, Tennessee. The three caves from which

the species is known are located at the northeast edge of the Central

Basin in the upper Cumberland River drainage; all of the caves are on the

south (left) side of Cumberland River.

Discussion. — In Carter Cave the species occurs sympatrically and

syntopically with a more abundant species, P. productus, of the

cumberlandus group, under rocks and on silt banks by the cave stream.

Females and undissected males can be distinguished by the more robust

and more depressed form and by the deeper elytral striation. The

aedeagus of simplex has a shorter and less produced apex than that of

productus, and the transfer apparatus consists of two prominent sclerites

instead of the small, single, very slender sclerite characteristic of the

cumberlandus group.

Pseudanophthalmus fowlerae, new species

Fig. 3

Etymology. — Patronymic honoring Mrs. Otia Fowler, owner of the

type locality cave.

Diagnosis.— Differs from simplex in shallow, irregularly punctate

elytral striae, glabrous and non-pruinose elytral disc, microsculpture with

New Beetle Species Groups

89

denser, transverse meshes; apical groove a little shorter; left copulatory

piece absent.

Description. — Length 4.0-4. 6, mean 4.3 mm. Form as in simplex but

paler, slightly more depressed, virtually glabrous, elytral disc highly

polished, transverse meshworks of microsculpture denser, and no trace of

pruinose microsculpture. Head slightly transverse, labrum doubly

emarginate. Pronotum as in simplex. Elytra with longitudinal striae more

shallowly impressed, feebly punctulate, inner 3 striae regular, fourth stria

irregular, outer striae obsolete. Aedeagus 0.66-0.70, mean 0.68 mm, of

same form as that of simplex but a little smaller and apex even less

produced; transfer apparatus a single sclerite, apparently homologous to

right piece of simplex : large, rather heavily sclerotized, margins parallel,

concave to left, apex bluntly rounded but scarcely knobbed.

Type series. — Holotype male (American Museum of Natural

History) and 4 paratypes, Sheals Cave, 0.8 km e of Celina, Clay Co., Ten-

nessee, 3 December 1960, T. C. Barr. Six additional paratypes, Sheals

Cave, 28 February 1959, 7 April 1960, and 11 August 1965, T. C. Barr

and J. R. Holsinger.

Measurements (in mm). — Holotype, total length 4.04, head 0.70 long

X 0.74 wide, pronotum 0.76 long X 0.90 wide, elytra 2.20 long X 1.48

wide, antenna 2.80, aedeagus 0.68.

Distribution.— Known only from the type locality cave, from which

1 1 specimens are available.

Discussion : Two beetles collected in August, 1965, are late tenerals,

suggesting midsummer eclosion. Most of the specimens were taken under

rocks in a depression in the floor of the entrance room, well within the

twilight zone, in an area kept moist by a constant drip from the ceiling, or

from the damp silt floor in a small domepit or along the cave stream not

far from the entrance. The entrance room is approximately 75 m long,

and the cave extends another 180 m along a narrow stream channel. The

species is sympatric with P. inquisitor, which is noticeably smaller and

belongs to the cumberlandus group. Both species are about equally abun-

dant, judging from the size of the type series (1 1 fowlerae, 10 inquisitor ),

which were obtained on four separate visits to the cave.

cumberlandus group (new group)

Size small to medium (3-5 mm); moderately pubescent, form

variable, usually rather slender and depressed, robust and convex in a few

90

Thomas C. Barr, Jr.

larger species; elytral microsculpture a fine, tranverse meshwork,

pruinose only in species from Rockcastle, Pulaski, and Wayne counties,

Kentucky. Labrum doubly emarginate, the median lobe varying from

weak to moderately prominent. Anterior discal at level of 4th umbilicate;

apical groove typically short and very rounded, joining 3rd stria at or

slightly anterior to anterior apical puncture; humeri weakly serrulate or

not serrulate. Mentum tooth broad, short, emarginate. Mesosternum

declivous. Last abdominal sternite of males with apical margin entire.

Aedeagus weakly and evenly arcuate, basal bulb not conspicuously

enlarged nor deflexed, aedeagal apex usually slender, more or less

produced, not very attenuate, bluntly rounded at tip; transfer apparatus

consisting of a single slender, small, elongate-triangular copulatory

sclerite; parameres rather short, with 4 setae at their apexes. Type species:

P. cumberlandus Valentine.

Discussion. — Pseudanophthalmus cumberlandus was placed in the

pubescens group by Jeannel (1949). Barr (1959) previously relegated 7

taxa described as polytypic P. tiresias to a section of the engelhardti

group, where they do not belong; they are small, mostly slender and

depressed species, but all have the typical cumberlandus group transfer

apparatus. Further study of the tiresias complex indicates that all 7 taxa

are full species.

The group is widely distributed in the Central Basin of Tennessee

but also extends into north Alabama (Limestone County), western Ken-

tucky (Christian, Caldwell, and Livingston counties), eastern Kentucky

(Rockcastle, Pulaski, and Wayne counties), the eastern Highland Rim of

Tennessee (Coffee County), and the western Highland Rim and western

valley of the Tennessee River in Tennessee (Montgomery, Dickson,

Hickman, and Lewis counties). In numbers of species (approximately 30)

the group exceeds the engelhardti group. The aedeagal form and simple

copulatory sclerite are highly diagnostic; the single sclerite is probably

homologous with the left piece of other groups. Some of the component

species, especially those at the periphery of the group’s collective range,

are sympatric with species of the pubescens, menetriesi, and simplex

groups. Some of the species in the southeastern and western parts of Mid-

dle Tennessee are geographically close to species of the engelhardti group

(P. loedingi humeralis Valentine and P. hesperus Barr, respectively), but

no sympatry has been demonstrated. In the case of the sympatric species

pair P. productus/ P. simplex the body size is approximately the same

(4. 2-5. 4 mm and 4.2-5. 1 mm respectively), and in the pair P. inquisitor/ P.

fowlerae the former species is much smaller (3. 1-3.9 vs. 4.0-4. 6 mm,

respectively). The species of the pubescens and menetriesi groups are in-

variably conspicuously larger than species of the cumberlandus group

with which they coexist.

New Beetle Species Groups

91

Pseudanophthalmus cumberlandus Valentine

Valentine 1937:96. Type locality, Piper Cave, Smith Co., TN.

The species is seasonally abundant in damp areas of Piper Cave, es-

pecially on previously flooded muddy floors and around rimstone pools

and bat guano, but it is usually rare or absent in winter. Unlike the ma-

jority of the species in the group, P. cumberlandus is unusually robust and

convex, with prominent humeri and nearly perpendicular prehumeral

borders, the elytral striae very shallow. I have also taken it in Skeleton

Cave, Smith County, and Ann White Cave, Macon County, Tennessee.

Pseudanophthalmus productus, new species

Fig. 5

Etymology.— Latin productus, “produced.”

Diagnosis. — Closely similar in the robust, convex body form and

nearly perpendicular prehumeral borders to P. cumberlandus Valentine,

differing in denser pubescence, wider pronotum base, deeper elytral

striae, and long, produced, aedeagal apex.

Description. — Length 4.2-5.4, mean 4.9 ± S.E. 0.3 mm (N =35).

Form robust and convex, pubescent, rufotestaceous. Head about 0.1

longer than wide; labrum doubly emarginate; antenna 0.7 body length.

Pronotum 0.87 as long as wide, transverse-subquadrate; base wider than

apex and 0.8 maximum width, which occurs in apical third; sides sinuate

in basal sixth, but very shallowly so, hind angles about right, sometimes

slightly acute or slightly obtuse, feebly emarginate behind, secondary

angles present but rather subdued. Elytra 1.6 times longer than wide,

prehumeral borders nearly perpendicular to midline, humeri prominent

and serrulate, inner 6 striae moderately impressed, intervals subconvex,

apical groove short and very rounded, joining third stria at level of an-

terior apical puncture. Aedeagus 0.62-0.77, mean 0.70 ± S.E. 0.04 mm

long (N = 30), moderately and evenly arcuate, apex elongate, slender,

and much produced; copulatory sclerite elongate.

Type series. — Holotype male (American Museum of Natural

History) and 27 paratypes, Neil Fisher Cave (=Rip Van Winkle Cave),

Smith Co., Tennessee, 25 March 1961, T. C. Barr; 33 paratypes, same

cave, 18 October 1948, J. M. Valentine and W. B. Jones.

Measurements (in mm).— Holotype, total length 4.80, head 0.92 long

92

Thomas C. Barr, Jr.

X 0.84 wide, pronotum 0.92 long X 1.06 wide, elytra 2.85 long X 1.79

wide, antenna 3.32 long.

Distribution.— One hundred seven specimens seen as follows. TEN-

NESSEE: Smith County.— Neil Fisher Cave (type locality), New Salem

Cave. Putnam County.— Bartlett, Bowen, Hargis, Jared Hollow, Petty,

and Sadler caves. Jackson County.— Burial, Harris, Mahaney, Medley,

and Carter caves (see Barr 1961 for cave locations).

Discussion. — This moderately large, robust, convex species is closely

similar to P. cumberlandus but occurs on the opposite (south, left) side of

Cumberland River. It is readily differentiated by characters given in the

diagnosis. In the northern part of its range, P. productus is sympatric and

syntopic with P. simplex, and at the southern margin of its range it is con-

junct with P. farrelli Barr ( robustus group), which occurs in nearby John

Fisher Cave (Barr 1962). Superficially both P. cumberlandus and P.

productus resemble species of the robustus group, but their distinctive

aedeagi readily differentiate them as members of different groups.

Although a few other species of the cumberlandus group are rather

robust, they are less convex and do not have the prominent humeri of P.

cumberlandus and P. productus. Neither species is obligatorily riparian,

although both are hygrophilous and somewhat seasonal in caves influ-

enced by cold air flowing into large entrances. The niche filled appears to

be much like that of P. robustus and its allies (other than P. valentinei

Jeannel, which is riparian), and this ecological similarity may explain the

conjunct, nonoverlapping ranges of P. farrelli and P. productus.

Pseudanophthalmus tiresias Barr, new status

Pseudanophthalmus tiresias tiresias Barr 1959:16. Type locality, Indian

Grave Point Cave, Dekalb Co., TN.

This is a species found in mesic cave microhabitats, occurring on wet

rotting wood and muddy floors rich in organic matter in the type locality

cave and in nearby Fox Cave. It is sympatric with P. farrelli, which is

conspicuously larger (see Barr 1962).

Pseudanophthalmus catherinae Barr, new status

Pseudanophthalmus tiresias catherinae Barr 1959:17. Type locality, Petty

Cave, Marshall Co., TN.

Known only from the type locality along the banks of the cave

stream, where it is moderately abundant. The species is rather robust but

more depressed than P. cumberlandus.

New Beetle Species Groups

93

P seudanophthalmus insularis Barr, new status

Pseudanophthalmus tiresias insularis Barr 1959:18. Type locality, Baker

Station Cave, Davidson Co., TN.

The type locality is a small stream cave in Silurian limestones of the

Highland Rim margin north of Nashville; known only from the type

locality, where it is rare. The species is small, slender, and depressed.

Pseudanophthalmus occidentalis Barr, new status

Pseudanophthalmus tiresias occidentalis Barr 1959:18. Type locality,

DePriest Branch Cave, Lewis Co., TN.

Known only from the type locality cave and nearby Cave Branch

Cave, Hickman County, Tennessee, this small, slender, depressed species

is rare in both caves. It is potentially sympatric with P. hesperus Barr

(engelhardti group), which occurs in the Blowing Caves, Perry County,

Tennessee, but the two species have not yet been shown to coexist.

Pseudanophthalmus acherontis Barr, new status

Pseudanophthalmus tiresias acherontis Barr 1959:20. Type locality, Echo

Cave, Rutherford Co., TN.

Known only from the type locality cave, apparently a branch of the

extensive Snail Shell Cave system (in the main trunk of which it has not

yet been collected), this small, slender, depressed species is found in

riparian microhabitats. All available specimens are unusually pale,

rufotestaceous.

Pseudanophthalmus tullahoma Barr, new status

Pseudanophthalmus tiresias tullahoma Barr 1959:20. Type locality,

Carroll Cave, Coffee Co., TN.

The species is known only from Carroll and Riley Creek caves, Cof-

fee County, Tennessee, both of which have been flooded by the Ten-

nessee Valley Authority’s Normandy Dam on Duck River. In Carroll

Cave this small, slender, depressed species occurred on rotting wood in a

wet area, and in Riley Creek Cave it occurred in a muddy, periodically

flooded area on rotting wood in which large numbers of Onychiurus

collembolans were found.

94

Thomas C. Barr, Jr.

Pseudanophthalmus bendermani Barr, new status

Pseudanophthalmus tiresias bendermani Barr 1959:21. Type locality, Ben-

derman Cave, Maury Co., TN.

At the present time this species is known only from the type locality,

an extensive stream cave containing much bat guano. The small, slender,

depressed beetles are rather abundant in riparian microhabitats.

Figs. 4-6. Pseudanophthalmus: cumberlandus group. 4. P. inquisitor, n. sp. (3.4

mm). 5. Aedeagus, P. productus, n. sp. (0.75 mm). 6. Aedeagus, P. inquisitor, n.

sp. (0.53 mm).

Pseudanophthalmus inquisitor, new species

Figs. 4, 6

Etymology. — Latin inquisitor, “searcher, inquirer.”

Diagnosis.— A slender and depressed species characterized by very

small size, non-pruinose elytral microscuipture, and the small, scarcely

arcuate aedeagus with deflexed apex.

New Beetle Species Groups

95

Description.— Length 3. 1-3.9, mean 3.5 0.3 mm. Head rounded;

labrum feebly trilobed; antenna 0.7 body length. Pronotum transverse,

0.87 as long as wide, margins arcuate in apical two-thirds, then oblique

and scarcely sinuate at all to approximately right (or slightly acute or

slightly obtuse) hind angles; widths at apex and base subequal and three-

fourths greatest width, which occurs in apical third; base emarginate

behind angles, secondary angles present; disc with rather long

pubescence. Elytra elongate-oval, strongly depressed, 1.6 times longer

than wide; microsculpture a fine, very tight, transverse meshwork;

humeri angular, prehumeral borders almost perpendicular to midline;

longitudinal striae 1-5 more or less complete and moderately impressed,

intervals subconvex; apical groove short and widely rounded or (as in

holotype) subparallel, joining third stria at anterior apical puncture.

Aedeagus 0.48-0.52 mm long, basal bulb large but not sharply set off

from median lobe, which is narrow and straight, its apex deflexed,

produced, and bluntly rounded at tip; transfer apparatus a single, slen-

der, elongate sclerite; parameres slender, with 3 apical setae.

Type series. — Holotype male (American Museum of Natural

History) and one paratype, Sheals Cave, Clay Co., Tennessee, 7 April

1960, T. C. Barr. Three additional paratypes, same cave, 3 December

1960, T. C. Barr, and 5 paratypes, same cave, 1 1 August 1965, T. C. Barr

and J. R. Holsinger. Known only from the type locality.

Measurements (in mm).— Holotype, total length 3.10, head 0.60 long

X 0.62 wide, pronotum 0.65 long X 0.77 wide, elytra 1.85 long X 1.15

wide, antenna 2.18 long.

Distribution.— Sheals Cave is 0.8 km e of Celina, Tennessee, near the

juncture of Obey and Cumberland rivers at the northeast margin of the

Central Basin.

Discussion. —The species is sympatric and syntopic with P. fowlerae

but is noticeably smaller. It is more readily collected in summer at the

back of the entrance room, but a few specimens occur throughout the

year along the stream in the back of the cave.

ACKNOWLEDGMENTS.— I thank Dr. John R. Holsinger for

assistance in the field, and Drs. J. Manson Valentine and the late Walter

B. Jones for contribution of specimens. The encouragement and coopera-

tion of Mrs. Otia Fowler, owner of Sheals Cave at the time of my visits, is

greatly appreciated.

96

Thomas C. Barr, Jr.

LITERATURE CITED

Barr, Thomas C., Jr. 1959. New cave beetles (Carabidae, Trechini) from Tennessee

and Kentucky. J. Tenn. Acad. Sci. 34: 5-30.

1961. Caves of Tennessee. Tenn. Div. Geol. Bull. 64, Nashville.

567 pp.

1962. The robustus group in the genus Pseudanophthalmus (Coleop-

tera: Carabidae:Trechini). Coleopt. Bull. 7(5:109-118.

1979. The taxonomy, distribution, and affinities of Neaphaenops,

with notes on associated species of Pseudanophthalmus (Coleoptera:

Carabidae). Am. Mus. Novit. 2682:1-20.

Hobbs, Horton, H., Jr., and T. C. Barr, Jr. 1972. Origins and affinities of the

troglobitic crayfishes of North America (Decapoda: Astacidae). II. Genus

Orconectes. Smithson. Contrib. Zool. 105:1-84.

Jeannel, Rene. 1949. Les coleopteres cavernicoles de la region des Appalaches.

Etude systematique, Notes Biospeol. 4 (Publ. Mus. Natl. Hist. Nat.,

Paris 12):37- 104.

Valentine, J. Manson. 1937. Anophthalmid beetles (fam. Carabidae) from

Tennessee caves. J. Elisha Mitchell Sci. Soc. 55:93-100.

Accepted 28 January 1980

Breeding Migrations, Population Size Structure, and

Reproduction of the Dwarf Salamander,

Eurycea quadridigitata, in South Carolina

Raymond D. Semlitsch

and

Michael A. McMillan

Savannah River Ecology Laboratory,

P.O. Drawer E., Aiken, South Carolina 29801

ABSTRACT. — Life history parameters of Eurycea quadridigitata from

two populations on the upper Coastal Plain of South Carolina were

examined. Breeding migrations into Carolina bays occurred during Sep-

tember and October in 1978 and July through October in 1979. The in-

itiation of breeding migrations was associated with a drop in air tem-

perature during late summer or early autumn. The sex ratio of E.

quadridigitata entering the breeding sites was 1:1. Egg deposition

probably occurs during November and early December. Sexual

dimorphism in snout-vent lenth (SVL) was not present, but there was

significant local variation in SVL of adults (x = 26 and 29 mm). The

number of ovarian eggs was positively correlated with SVL and varied

between populations (x = 21 and 33).

INTRODUCTION

Although the dwarf salamander, Eurycea quadridigitata, is a widely

distributed species in the southeastern United States (Dunn 1926; Mit-

tleman 1947), life history information in most parts of its range is based

on anecdotal observations (Bishop 1947; Dunn 1926; Neill 1949; Noble

1927). Sever (1975), in a more complete report, described seasonal varia-

tion of the nasolabial glands and included data on the male reproductive

cycle. Harrison (1973) and Semlitsch (in press) provided ecological infor-

mation on habitat, and reproductive and larval characteristics. Tax-

onomic accounts, distributional records, and other pertinent literature on

the species were reviewed by Mittlefnan (1967).

As previously shown, the drift fence and pitfall trap technique

provides an ideal sampling method for monitoring populations of

fossorial amphibians (Gibbons and Bennett 1974; Gill 1978; Shoop

1965). Since lack of an adequate sampling technique has hindered the

study of E. quadridigitata, this study was initiated to provide

quanititative information on life history parameters of dwarf salamander

populations from the upper Coastal Plain of South Carolina.

Brimleyana No. 3: 97-105 July 1980.

97

98

Raymond D. Semlitsch and Michael A. McMillan

MATERIALS AND METHODS

Two Carolina bays, Rainbow Bay and Ellenton Bay, represent

natural lentic habitats of the upper Coastal Plain (Schalles 1979) and

were selected as study sites on the U. S. Department of Energy’s Savan-

nah River Plant (SRP) near Aiken, South Carolina. Rainbow Bay is a

Carolina bay < 1 ha in area and is surrounded primarily by pine planta-

tions. Slash pine, Pinus elliotii, and loblolly pine, P. taeda, surround the

bay, with wax myrtle, Myrica cerifera, and blackberry, Rubus sp.,

providing the peripheral understory vegetation. Rainbow Bay is subjec-

ted to yearly drying, whereas Ellenton Bay is a more stable body of water,

drying only twice in the last 24 years. Ellenton Bay is an approximately

10 ha Carolina bay surrounded by an old field community undergoing

secondary succession (Gibbons 1970). Predominant plants peripheral to

the bay are bush clover, Lespedeza sp., dog fennel, Eupatorium sp., and

blackberry, Rubus sp.. Additional soil and vegetation characteristics are

described in Odum (1960). Rainbow Bay is approximately 12 km (7.5 mi)

from Ellenton Bay.

The drift fence and pitfall trap technique was employed at both bays.

The Rainbow Bay drift fence was checked daily from 21 September 1978

through 30 December 1979. Individuals caught in pitfall traps were coun-

ted, and sex and reproductive condition noted before an animal was

released on the opposite side of the fence. Samples for dissection were

collected from pitfall traps at Rainbow Bay from 24 September through

30 November 1978 and at Ellenton Bay from 30 September through 10

November 1978. Movements of E. quadridigitata at Ellenton Bay were

not monitored daily as they were at Rainbow Bay. Rainfall and

minimum and maximum air temperatures at Rainbow Bay were recorded

daily.

Salamanders kept for dissection were sacrificed in chloretone, pre-

served in 10% formaldehyde, and stored in 70% ethyl alcohol. Snout-vent

length (SVL) was measured on preserved specimens from tip of the snout

to posterior end of the cloaca. Individuals were dissected using a dis-

secting microscope to determine sex, reproductive condition, and ovarian

egg size. The diameters of six eggs, chosen at random, were measured

with a stage micrometer. Sexual maturity for males was determined by

the amount of pigmentation on testes and size of the testes and vasa

deferentia, and the presence of enlarged cirri (Gordon 1953; Sever 1975).

Females were considered mature if eggs ( > 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

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

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Crocker, Denton W. and D. W. Barr. 1968. Handbook of the Crayfishes of Ontario. Univ.

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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