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number 21 december 1 994

EDITORIAL STAFF

Richard A. Lancia, Editor

Suzanne A. Fischer, Assistant Editor

Eloise F. Potter, Production Manager

EDITORIAL BOARD

James W. Hardin

Professor of Botany

North Carolina State University

William M. Palmer

Director of Research and Collections

North Carolina State Museum

of Natural Sciences

Rowland M. Shelley

Curator of Invertebrates

North Carolina State Museum

of Natural Sciences

Robert G. Wolk

Director of Programs

North Carolina State Museum

of Natural Sciences

Brimleyana, the Zoological Journal of the North Carolina State

Museum of Natural Sciences, appears twice yearly in consecutively

numbered issues. Subject matter focuses on systematics, evolution,

zoogeography, ecology, behavior, and paleozoology in the southeastern

United States. Papers stress the results of original empirical field

studies, but synthesizing reviews and papers of significant historical

interest to southeastern zoology are also included. Brief communications

are accepted.

All manuscripts are peer reviewed by specialists in the Southeast

and elsewhere; final acceptability is determined by the Editor. Address

manuscripts and related correspondence to Editor, Brimelyana, North

Carolina State Museum of Natural Sciences, P.O. Box 29555, Raleigh,

NC 27626. Information for contributors appears in the inside back

cover.

Address correspondence pertaining to subscriptions, back issues,

and exchanges to Brimleyana Secretary, North Carolina State Museum

of Natural Sciences, P.O. Box 29555, Raleigh, NC 27626.

In citations please use the full name - Brimleyana.

North Carolina State Museum of Natural Sciences

Betsy Bennett, Director

North Carolina Department of Environment,

Health and Natural Resources

James B. Hunt, Jr., Governor

Jonathan B. Howes, Secretary

CODN BRIMD 7

ISSN 0193-4406

N.C. DOCUMENTS

CLEARI SE

The Mammals of the Ardis Igygal if&yy%

(Late Pleistocene), Harleyville, South Carolina

N.C. STATE LIBRARY

Curtis C. Bentley and James L. Kf$t£|GH

South Carolina State Museum, Natural History, 301 Gervais Street

P.O. Box 100107, Columbia, South Carolina 29202-3107

AND

Martin A. Knoll

University of the South, Department of Forestry and Geology,

Sewanee, Tennessee 37375-4003

ABSTRACT — The Ardis local fauna is contained within sediment-

filled solution cavities of late Pleistocene age, located in the

Giant Cement Quarry near Harleyville, Dorchester County, South

Carolina. This paper, the second on the fossil remains collected

from the Ardis site, documents 43 taxa of mammals recovered

from a group of interconnected solution cavities, including 24

taxa of mammals not previously reported as fossils from the

state. Fossil remains from the lowermost layers and the extreme

upper layer of the deposit were C14 dated at 18,940 ± 760

and 18,530 ± 725 y.b.p., respectively, and are considered

contemporaneous. Fossil remains were deposited near the height

of the Wisconsin glaciation and appear to reflect a mosaic edge

community, probably a patchwork of mixed hardwood and conifer

forest, interspersed with meadows, possibly associated with a marsh

or bog, located near a permanent stream or a river. The Ardis

local fauna is composed of a mammal community which has

no modern analogues ("disharmonious fauna"), and reflects a

more equitable climate, cooler summers and warmer winters,

than that presently occurring in the region.

INTRODUCTION AND GEOLOGY

As collections of fossils continue to be amassed from various

localities in South Carolina, the fossil record of the state likely will

be one of the richest in the eastern United States. Long known as a

source of marine mammal fossils of Eocene and Oligocene age, the

State has a large number of vertebrate faunas from various other time

periods. Although the Pleistocene epoch is one of the best represented

in South Carolina, only two assemblages (Allen 1926, Roth and Laerm

1980) have been reported in the literature to date.

Our purpose is to present data on a collection of late Rancholabrean

mammals, the Ardis local fauna (Appendix 1), found in the Giant

Brimleyana 21:1-35, December 1994

C. C. Bentley, J. L. Knight, and M. A. Knoll

Cement Plant pit near Harleyville, South Carolina. This is the second

(Bentley and Knight 1993) in a series of papers that reports the various

taxonomic assemblages collected from this site.

The Giant Cement Plant, located about 5 km north-northeast of

Harleyville, South Carolina (Fig. 1), is in Dorchester County, about

1 km from Interstate 26 at the Harleyville exit (33°14'N,80°26'W). It

is a large, open-pit mine, in operation for the past 45 years. The stratigraphic

assignments for the large exposures of middle Eocene Santee Limestone,

which is exploited for cement products, were determined by Sanders

(1974), who discussed the marine vertebrate and invertebrate material

collected from the pit. Based on reports from Cooke and MacNeil

(1952) and Malde (1959), Sanders (1974) refers to the layer which

Fig.l. Location of the Ardis local fauna.

Mammals of the Ardis Local Fauna

unconformably overlies the Santee Limestone as the "Cooper Marl"

of early to late Oligocene age. A more recent survey by Ward et al.

(1979) recognizes three members of the "Cooper Marl" (Cooper

Formation), the Ashley (Oligocene), the Harleyville (late Eocene), and

the Parkers Ferry (late Eocene). The Parkers Ferry and Ashley members

were absent at the Ardis fossil locality. Harris and Zullo (1991) refer

to the Harleyville member as the Harleyville Formation, and we follow

that practice in this paper. Since Sanders' (1974) study, fossil vertebrates

of presumed late Pleistocene origin have been collected by workman

and hobby collectors from several areas of the quarry, although no

stratigraphic assignment could be determined for these fossils. Most

were collected from spoil piles of unknown origin.

In April 1991, during a periodic visit to the quarry, the senior

author, accompanied by Mr. Vance McCollum, a hobby collector,

discovered a newly-mined area in the southeastern wall of the quarry

which contained a dense concentration of late Pleistocene terrestrial

vertebrate fossils. Giant Cement, in anticipation of further mining

operations, had used a dragline to remove overburden from the

underlying Santee Limestone. This was done in a single strip which

formed a trench at a right angle in the southernmost corner of the

pit. This tract of freshly mined earth had a maximum length of about

150 m, a width of 10 m, and a depth of 4 m. Fossils were initially

collected from a single row of spoil piles dumped adjacent to the area

from which they were mined. We determined that the fossil material

came from a series of localized, sediment-filled cavities formed in the

underlying Harleyville Formation and Santee Limestone. Groundwater

had preferentially dissolved away the upper portions of the Santee

Limestone, so that many of the solution cavities contacted the overlying,

clay-rich Harleyville Formation. This resulted in the occasional collapse

of the Harleyville Formation, opening several of the cavities to the

surface.

The diameter of the solution cavities varied from a few centimeters

to a maximum diameter of nearly 2.0 m, with many of the cavities

interconnected. The solution cavities reached a maximum depth below

the Harleyville Formation of about 4 m. The majority of the cavities

were infilled with clastic sediment from the surface. The sediment is

well stratified and ranges in size from clay to small pebbles. Multiple

fining-upward sequences within the cavity-filling deposits indicate episodic

sedimentation, perhaps corresponding to flood events within a nearby

fluvial system. The undisturbed clay layers within the cavity-filling

and the narrow interconnecting nature of the cavities indicate that

the entire deposition probably occurred over a relatively short time

C. C. Bentley, J. L. Knight, and M. A. Knoll

span. This tends to be confirmed by the limited range of carbon dates.

The Harleyville Formation was capped by a well sorted, cross-stratified

quartz sand having a maximum thickness of about 4 m. Fossils collected

from this layer were similar to those in the cavities, but fewer in number.

Locally, several cavities which had no connections to the surface

were exposed by mining operations in the Santee Limestone. Conse-

quently, these cavities were not infilled from above.

A systematic excavation was not possible because the area at

the time of discovery was actively being mined. The operators of the

mine graciously relocated their mining operation to another section of

the pit, allowing us the maximum possible study time and nearly unlimit-

ed access to the fossil deposit. Bert Ardis, an employee of the Giant

Cement Plant Company, played a crucial role in the discovery and

excavation of fossils from the quarry. In recognition of his efforts

the fauna was named in his honor.

Dating of the Deposit

Kruger Enterprises, Inc., Cambridge, Massachusetts, used C14

dated (C13 corrected) mammal and reptile bone apatite to date fossil

materials from the lower levels of the solution cavities and from the

well-sorted sand layer above the Harleyville Formation. Dating was

done on apatite due to the paucity of collagen. The fossil material

from the solution cavities (300 g of mixed mammal and reptile bone)

were dated at 18,940 ± 760 y.b.p. and the fossil material (300 g of

mammoth bone), from the overlying unnamed, quartz sand layer (about

1 km from the primary deposit), dated at 18,530 ± 725 y.b.p. The

overlap of the two dates suggests that the material in the solution

tunnels and the homogeneous sand layer above the Harleyville Forma-

tion are contemporaneous. These dates place the time of deposition at,

or near, the height of the Wisconsin glaciation (Bowen 1988).

Serious questions have recently been raised concerning the

reliability of C14 dating based on bone apatite and collagen (Stafford

et al. 1991). They state that C14 dates on noncollagenous bone may

yield dates potentially thousands of years too young. Efforts are under-

way to acquire an amino acid date, even though the apatite sample was

apparently very clean (Kruger Enterprises, Inc., personal communica-

tion). In light of recent work (Stafford et al. 1991), the C14 dates we

used will be considered a minimal age for this deposit. The maximum

age of the deposit is not expected to exceed 22,000 y.b.p. (Kruger

Enterprises, Inc., personal communication). Whatever the date, how-

ever, there is little doubt that the fauna represents a late Rancholabrean

age, and minor revisions in the absolute dates would have little effect

on the ecological or climatic interpretations suggested in our paper.

Mammals of the Ardis Local Fauna

Methods

All fossil material reported here was collected during periodic

trips into the quarry by the authors and a group of volunteers. We

collected fossil material from the surface of associated spoil piles as

well as by screen sifting the fossiliferous sand. Screens used were 20

gauge (1.0 mm) to quarter-inch (6.35 mm) mesh. Materials in situ

were collected by a combination of screen-washing and by applying a

gentle, controlled stream of water to the exposed sands. Fossils were

exposed as the sand was gently washed away.

Most specimens from the Ardis local fauna were deposited in

the South Carolina State Museum collections and are registered under

the accession base number of S.C. 93.105. We cite such specimens in

this paper only by the numbers following that base number. Fossils

that were accessioned separately in the South Carolina State Museum's

collections will be designated by S.C. followed by five to six digits.

Fossil specimens deposited in the National Museum of Natural History

have been accessioned under the number of 407498 and are indicated

by an institutional prefix of USNM in the text. Fossil material deposited

at the Florida Museum of Natural History will be referred to by an

institutional prefix of UF.

Taxonomy for extant and extinct taxa follows Wilson and Reeder

(1993) and Kurten and Anderson (1980), respectively. Common names

were taken from Webster et al. (1985), and Kurten and Anderson

(1980). Under Materials, capital "M" denotes upper dentition, lower

case "m" denotes lower dentition.

SYSTEMATICS

Class Mammalia

Order Didelphimorphia

Family Didelphidae

Didelphis virginiana - Virginia Opossum

Material: 1 left dentary with ml and m2 and an unerupted p3 (.1);

1 left dentary with m2 and m3 (.2); 1 left maxilla fragment with P3

and Ml (.3); 1 right M3 (.8); 1 right Ml (.9); 6 right partial dentaries

(3 USNM & 3 UF); 1 left dentary, proximal one third (USNM); 1

right proximal dentary fragment (USNM); 4 left humeri, distal ends

(.4-.7).

Remarks: The opossum is found throughout most of the United States

and northward into Canada. Late Pleistocene fossils are restricted

to the southern parts of the United States, and the species is believed

to have radiated into the northern part of the continent during historical

times (Kurten and Anderson 1980). The species is common today in

the vicinity of the Ardis locality.

C. C. Bentley, J. L. Knight, and M. A. Knoll

This is the first published report of fossil D. virginiana from

South Carolina.

Order Xenarthra

Family Megalonychidae

Megalonyx jeffersoni - Jefferson's Ground Sloth

Material: 1 upper right 3rd molariform (.194).

Remarks: This sloth has been found as far north as Canada and the

western coast of Alaska, and formerly ranged throughout the United

States except for the Rocky Mountains, Great Basin, and the arid or

desert areas of the Southwest. It is thought that the diet of this sloth

consisted of twigs and leaves and that it inhabited forested or

wooded areas (Kurten and Anderson 1980).

Previous reports of this species from South Carolina include

Hay (1923) and material from Edisto Island (Roth and Laerm 1980).

Family Dasypodidae

Dasypus bellus - Beautiful Armadillo

Material: 25 buckler osteoderms (.200- .210), (5 USNM), (9 UF);

11 movable osteoderms (.195- .199), (3 USNM), (3 UF); 1 cephalic

osteoderm (.211); 1 caudal vertebra (.212).

Remarks: This armadillo probably fed primarily on insects and that

diet might have restricted its range to moderate climes (Kurten and

• Anderson 1980). However, Voorhies (1987) questioned its reliability

as a warm weather indicator. Its occurrence with mostly boreal forms

in the Craigmile local fauna, Nebraska (Rhodes 1984), suggests that

it may have been able to withstand weather conditions considerably

cooler than D. novemcinctus. D. bellus fossil records are well established

in South America and the southern portions of the United States (Kurten

and Anderson 1980). This suggests that D. bellus is a fairly reliable

indicator of warm climates, and the material from Nebraska may represent

a fringe population. Although a common Pleistocene fossil, this is only

the second report of this species from South Carolina. It was first

reported by Roth and Laerm (1980) from Edisto Island.

Family Pampatheriidae

Holmesina septentrionalis - Northern Pampathere

Material: 1 partial buckler osteoderm (.227); 1 cephalic osteoderm (.228).

Remarks: Differentiation of the cephalic and buckler osteoderm was

based on the descriptions of Edmund (1985).

Mammals of the Ardis Local Fauna

Little is known about its diet, which probably consisted of coarse

vegetation (based on large, flat, high-crowned teeth of indeterminate

growth). Regarding its habitat, Holmesina is suspected to be intolerant

of cold climates (Kurten and Anderson 1980).

Holmesina was first reported from South Carolina by Roth and

Laerm (1980) at Edisto Island.

Order Insectivora

Family Soricidae

Blarina brevicauda - Northern Short-tailed Shrew

Material: 2 right maxillae with the Ml and M2 (.11, .12); 4 right

dentaries with m2 and m3 (USNM); cl, pi, p2, ml, m2 (.13); cl, ml

(.14); cl, pi, p2, ml (.15), respectively; 2 left maxillae with Ml, M2,

M3 (.16) and Ml (.17) respectively; 1 left dentary with ml-m3 (.18);

1 isolated cl (.19).

Remarks: Identification was made on the basis of characters listed

by Guilday (1962) and by direct comparisons to fossil and recent specimens.

Ardis specimens are indistinguishable from modern comparative materials

of B. brevicauda and are generally larger in size than samples of B.

carolinensis (Graham and Semken 1976). Apparently B. brevicauda

size reflects a positive Bergmann's response, suggesting that the fossils

from the Ardis site are a more northerly stock and not the smaller

southern species B. carolinensis (McNab 1971, Guilday et al. 1977,

Klippel and Parmalee 1982, Jones et al. 1984).

The northern short-tailed shrew occurs in a variety of terrestrial

habitats from forests, fields and meadows, to salt marshes. Today this

species occurs, in South Carolina, only in the extreme northwestern

portion of the State, nearly 300 km from the fossil locality. An isolated

population on the coast of North Carolina is about the same distance

from the site (Webster et al. 1985). B. carolinensis does occur in the

area of the Ardis site today.

This is the first fossil record of B. brevicauda reported from

South Carolina.

Sorex sp. cf. S. longirostris - Southeastern Shrew

Material: 1 left dentary with complete dentition (.10).

Remarks: Guilday (1962) and Guilday et al. (1969, 1977) separated

Sorex cinereus from S. longirostris based on a slightly larger mean

size of S. cinereus, because tooth morphology is nearly identical. The

C. C. Bentley, J. L. Knight, and M. A. Knoll

P4-M3 measurement of our specimen (3.64 mm) fell at the bottom

of the range for S. cinereus and below the mean given for that species

at both New Paris Cave No. 4, Pennsylvania, and Clark's Cave, Virginia

(Guilday 1964, Guilday et al. 1977). Because the Ardis local fauna

contains many extralimital species, both northern and southern, and

because of the morphological ambiguity between S. longirostris and

S. cinereus (Jones et al. 1991), the fossil specimen from the Ardis site

is tentatively assigned to S. longirostris and not S. cinereus, as only

the former occurs in the area of the Ardis site today. The Sorex

longirostris P4-M3 measurement is considerably larger than that of S.

hoyi (Guilday et al. 1977).

At present the southeastern shrew ranges from southeastern

Arkansas, east to central Florida, and north along the Atlantic coast

into Maryland. It is associated with moist, open fields and lowland

forests but can also be found in dry upland fields (Webster et al. 1985).

In the late Pleistocene this shrew has been reported only from the late

Rancholabrean Haile 11B and Arredondo sites in northern Florida (Kurten

and Anderson 1980, Webb and Wilkins 1984). The Ardis site is the

first identification of this shrew from the Wisconsin time period and

from South Carolina.

Family Talpidae

Condylura cristata - Star-nosed Mole

Material: 3 humeri (.20-. 22).

Remarks: Identification was made by comparisons to recent specimens.

Humeri of this species differ from those of S. aquaticus in having a

less robust humeral shaft and a smaller width at both proximal and

distal ends.

At present C. cristata occurs in the upland and the Coastal Plain

of South Carolina but is absent from the midland area. The star-nosed

mole is an excellent swimmer and inhabits areas that have moist soils

or are located near water (Webster et al. 1985). It may be found in

wooded areas, meadows or fields, swamps, and bogs. Today, it is found

in the area of the Ardis site.

This is the first fossil report of this mole from South Carolina.

Scalopus aquaticus - Eastern Mole

Material: 16 humeri (.23-. 26), (6 USNM), (6 UF).

Remarks: These humeri are separated from other South Carolina mole

species on the basis of their greater robustness, which may reflect

a more fossorial life style.

Mammals of the Ardis Local Fauna

The eastern mole ranges from the eastern to midwestern United

States, commonly inhabiting well-drained sand or loam soil types,

and less common in clay or gravel soils (Webster et al. 1985). Eastern

moles are found today in the vicinity of the fossil site.

This is the first fossil record of this species for South Carolina.

Order Carnivora

Family Canidae

Urocyon sp. cf. U. cinereoargenteus - Gray Fox

Material: 1 metapodial (.254).

Remarks: Felid metapodial elements are generally more robust and

have a more acute curvature to the shaft in comparison to modern

canid specimens in the Florida Museum of Natural History collec-

tions (Gary Morgan, Florida Museum of Natural History, personal

communication). The proximal articulation varies significantly from

all felids examined and most closely parallels those of canids. The

curvature of the elongated shaft, overall size, and simple articulations

of the proximal end suggest a close affinity to U. cinereoargenteus. A

small convexity dorsal and anterior to the proximal articulation was

the most significant variation between the fossil element and recent

specimens of the gray fox.

Roth and Laerm (1980) refer an Edisto Beach ulnar fragment to

this species but state that it may be recent.

Canis dims - Dire Wolf

Material: 1 brain case (SC 91.171.1); 1 left cl (.251); 1 metapodial

shaft (.252); 1 left jugal (.253).

Remarks: The dire wolf is one of the more common late Pleistocene

canids recovered from North American fossil sites (Kurten and

Anderson 1980). The material recovered from the Ardis site is believed

to represent "wash-ins," as many of the elements, particularly the

brain case, show signs of weathering and abrasions prior to fossilization.

This represents the second reported occurrence of the dire wolf from

this State, as C. dims was reported from Edisto Island (Roth and

Laerm 1980).

10 C. C. Bentley, J. L. Knight, and M. A. Knoll

Family Felidae

Subfamily Machairodontinae

cf. Smilodon fatalis. - Sabertooth

Material: 1 left occipital condyle (.266).

Remarks: The shape and size of the condyle has its closest affinities

with Smilodon and differs significantly from any other large mammal.

The identification is extremely tenuous, however, and after an exhaus-

tive search, represents the best possible solution to the identification

of this specimen.

Subfamily Felinae

Lynx rufus - Bobcat

Material: 1 right dentary lacking teeth, proximal end missing behind

m3 alveoli (.267).

Remarks: This specimen is extremely similar to mandibles of Lynx

rufus especially in general ramus shape, and alveolar count and placement.

The bobcat inhabits a range of habitats including deserts, swamps, and

upland forests (Kurten and Anderson 1980), and it occurs in the Ardis

local fauna area today (Webster et al. 1985).

This is the first fossil record of Lynx rufus from South Carolina.

Family Mustelidae

Subfamily Lutrinae

Lontra canadensis - River Otter

Material: 1 cranium missing the right zygomatic arch and retaining

only the left Pl-Ml (SC 91.116.1); 1 atlas vertebra (.242); 1 caudal

vertebra (.243).

Remarks: The river otter once occurred throughout the United States,

including Alaska, but has been extirpated from many areas of the Mid-

west and Appalachian Highlands. The otter occurs sparsely through-

out South Carolina, inhabiting a wide range of aquatic habitats. It

appears to be most abundant in coastal estuaries and the lower reaches

of rivers in the State (Webster et al. 1985).

This is the first fossil record of this taxon from South Carolina.

Subfamily Mephitinae

Spilogale putorius - Eastern Spotted Skunk

Material: 1 proximal end of the right dentary with the alveoli for a

partial m2 and for the m3 (.244).

Mammals of the Ardis Local Fauna 11

Remarks: The partial dentary was differentiated from Mustela vison

because the condyle tapers to a point lingually, whereas it is blunt

in Mustela vison. The ramus of Spilogale is generally smaller and less

robust, is laterally compressed, and lacks the distinct curvature found

in mink. It also lacks the flattened ventral proximal edge found in the

dentary of mink. It differed from other mustelids in the shape and size

of the M2 alveoli and overall jaw size and shape. It is significantly

smaller than skunks of the genera Conepatus and Mephitis and is most

similar to Spilogale in size and general jaw morphology.

The eastern spotted skunk is found typically in prairies, brushy

open forests, and mountain habitats. They no longer occur in the immediate

area of the fossil site and are found sparsely in the Piedmont of

South Carolina today (Webster et al. 1985).

This represents the first fossil record of this species from the

State.

Mephitis mephitis - Striped Skunk

Material: 1 partial right dentary containing the alveoli of the m3 (.245);

1 left Ml (.256).

Remarks: The partial dentary was assigned to this species based on

the length of the third molar and the presence of auxiliary roots. The

overall size of the dentary was intermediate between Spilogale and

Conepatus.

The striped skunk is found in the area of the site today and through-

out much of North America and well into Central America (Hall 1981).

It can be found in habitats varying from high mountain forests to brushy,

semi-open areas, but appears to be much less common in wetlands

(Webster et al. 1985).

This is the first report of the striped skunk in the fossil record

from South Carolina.

Conepatus sp. cf. C. robustus - Extinct Hog-nosed Skunk

Material: 1 humerus (.247); 1 femur (.248); 1 partial dentary with p3

and alveoli of ml (.249); 1 astragalus (.250).

Remarks: The fossil remains have been referred to this species based

on their extremely large size in comparison with the living hog-nosed

skunk Conepatus leuconotus (Martin 1978). The Ardis fossils exhibit a

30% size increase over C. leuconotus. No definable differences were

observed between the Ardis material and the type of C. robustus from

12

C. C. Bentley, J. L. Knight, and M. A. Knol

B.

3 cm

Fig. 2. Conepatus sp. cf. C. robustus material recovered from the Ardis

local fauna. A) Humerus B) Femur C) Astragalus D) Dentary with p3.

the late Rancholabrean Haile 14 in Florida (Martin 1978). The femur

and astragalus are the first of these elements to be reported for this

species and were assigned based on morphology and their large size

(Fig. 2). The extant species inhabit the southwestern United States,

southward into Central America, and as far east as eastern Texas

(Hall 1981).

This is the first report of this species from South Carolina and

the first fossil record of this extinct hog-nosed skunk from outside of

Florida.

Subfamily Mustelinae

Mustela vison - Mink

Material: 1 left dentary, complete dentition except for incisors (1

USNM); 1 left dentary, complete dentition except for incisors and

Mammals of the Ardis Local Fauna 13

canine (.229); 1 partial maxilla with P3 and Ml and partial zygomatic

arch (.231); 2 canines (.232, .233); 1 left dentary with p3 and ml

(.230); 1 left dentary with alveoli p2-m2 (.234); 1 left dentary (in the

private collection of Lee Hudson, Florence, S.C.); 2 right ulnae (.235,

.236); 1 thoracic vertebra (.237); 1 lumbar vertebra (.238); 1 right 3rd

metatarsal (.239); 1 left 2nd metatarsal (.240); 1 rib (.241).

Remarks: Mink remains are generally uncommon in Pleistocene deposits

(Kurten and Anderson 1980), but it was the most common carnivore

at the Ardis site. This abundance might be explained by its relatively

small size and ability to enter cavities that may have excluded larger

carnivores. Mink may have also been attracted to favored prey items,

in particular a large muskrat population which was using the cavities

for various reasons. Muskrats are commonly fed upon by mink, and

during certain times of the year, depending on availability, make up

the bulk of their diet (Proulx et al. 1987). Mink are good indicators of

nearby bodies of water and are found in a variety of habitats bordering

water including rivers and streams, swamps, drainage ditches, marshes,

and lakes (Webster et al. 1985). Mink are not abundant in South

Carolina today, although they probably occur in the area of the Ardis

site.

This is the first reported fossil record of this species from South

Carolina.

Family Procyonidae

Subfamily Procyoninae

Procyon lotor - Raccoon

Material: 1 left dentary with m3 present and ml alveoli (.255); 1 left

dentary with p4-m2 alveoli (.256); 1 left ml (.257); 1 right proximal

end of dentary with m2 alveoli (.258); 1 left humerus shaft with

partial distal end (.259); 1 left humerus distal end (.260); 1 right ulna

proximal end (.261); 1 right humerus partial distal end (.263); 1 radius

distal half (.262).

Remarks: The distribution of the raccoon is from Panama north through

Mexico and the United States into the central portions of Canada

(Hall 1981). Its habitat is ubiquitous, but is generally in or near

forested wetlands such as stream and river bottoms, marshes, swamps,

ponds, and lakes as well as upland and agricultural areas (Webster et

al. 1985). Fossil remains of P. lotor are generally common in the

United States during the late Pleistocene, especially in sinkhole and

cave deposits (Kurten and Anderson 1980). The first South Carolina

fossil of the raccoon was reported from Edisto Island (Roth and Laerm

14 C. C. Bentley, J. L. Knight, and M. A. Knoll

1980), based on mandibular fragments and postcranial remains. The

material from the Ardis local fauna represents the second report of

fossil P. lotor from South Carolina.

Family Ursidae

Tremarctos floridanus - Florida Cave Bear

Material: 1 right m3 (.264); 1 first right metatarsal (.265)

Remarks: The m3 of Tremarctos floridanus differs from Ursus americanus

in that it lacks the double root, and is substantially smaller than any

other ursid in the eastern United States. The Florida cave bear has

been recovered from Rancholabrean sites in Florida, Georgia, Tennessee,

Kentucky, New Mexico, Texas, and northern Mexico (Kurten and

Anderson 1980). Based on the fossil record it would appear that the

Florida cave bear had a more southeastern distribution during the late

Pleistocene. The material from the Ardis local fauna represents the

second report of this species (Roth and Laerm 1980), from South

Carolina. Both reports are from deposits along the Coastal Plain of

the State.

Order Proboscidea

Family Mammutidae

Mammut americanum - American Mastodon

Material: 1 partial milk tooth (.332); 2 partial molars (.333, .334).

Remarks: The American mastodon was first reported in the fossil

record for South Carolina from Edisto Island by Roth and Laerm

(1980).

Family Elephantidae

Mammuthus columbi - Columbian Mammoth

Material: 1 complete molar (.335); partial molar (.336); partial scapula

(in the collections of Lander University).

Remarks: All identifiable Mammuthus remains from the area of the

Ardis local fauna have been recovered from the quartz sand layer

above the Harleyville clay, and not in direct association with the

Ardis fauna proper. Identification is based on the lamellar frequency

(7) of the occlusal surface coupled with the late Pleistocene date of

the site (Kurten and Anderson 1980). Carbon dating of the bone from

a scapula associated with a complete and a partial molar of M. columbi

gave a date of 18,530 ± 725 y.p.b., within the range of the Ardis

local fauna and here considered contemporaneous with it.

Mammals of the Ardis Local Fauna 15

All other Mammuthus material reported previously from South

Carolina has been assigned to M. columbi as well (Hay 1923, Allen

1926, Roth and Laerm 1980).

Order Proboscidea

gen. et spec, indet.

Material: 1 distal fragment of tibia (.337); 1 vertebra (.340); 1 proximal

rib end (.338); 6 fragments of ivory (.339a-c), (3 USNM).

Remarks: Identification to a particular family is not possible because

of the fragmentary nature of these remains. Because the openings of

the solution cavities to the surface were small, and large mammals

are only represented by isolated fragments, these animals probably

died on the surface nearby and were washed in by periodic flooding.

Order Perissodactyla

Family Equidae

Equus cf E. complicatus - Complex-toothed Horse

Material: 2 partial incisors (2 USNM); 1 left Ml or M2 (1 USNM);

1 left DP3 or DP4 (.320); 1 right P3 or P4 (.319); 2 right M3s

(1 USNM), (1 UF); 1 left m3 (1 USNM); 1 canine (.322); associated-

1 axis, 3 caudal vertebrae, 1 incisor, and 3 rib fragments (.321a-h); 1

left distal radial end (.323); 1 right fourth metatarsal (.324); 1 left

second metacarpal (.331); 2 left cuneiforms (.325, .326); 1 right magnum

(.327); 1 right lunar (.328); 2 medial phalanx (.329, .330).

Remarks: The Equus cheek teeth from the Ardis local fauna have

been tentatively assigned to the species E. complicatus based on the

extremely complex nature of the occlusal surface. All other remains,

because of the ambiguity in postcranial elements between E. complicatus,

E. fraternus, and E. occidentalism are referred to only as Equus sp.

Fossil remains assigned to the genus Equus were reported from

South Carolina by Hay (1923), Allen (1926), and Roth and Laerm

(1980).

Family Tapiridae

Tapirus veroensis - Vero Tapir

Material: 1 partial right m3 (1 USNM); 1 partial m2 (UF); 2 partial

left m3 (.316, .317); 1 partial right molar (.318).

16 C. C. Bentley, J. L. Knight, and M. A. Knoll

Remarks: Tapir remains are among the more commonly collected

Pleistocene fossils in South Carolina. Several tooth fragments referred

to T. haysii were reported by Allen (1926) and represent the first

record of tapir from South Carolina. Roth and Laerm (1980) reported

numerous tooth fragments and some postcranial material from Edisto

Island, but because of the fragmentary nature of the fossils, were

unable to assign them to species. A virtually complete skull of T.

veroensis, reported on by Ray and Sanders (1984), was collected by a

diver in the Cooper River near Charleston, South Carolina. The mate-

rial from the Ardis site is referred to T. veroensis, based on the

generally smaller size of the molars in comparison to T. haysii. Ex-

tant species are semiaquatic browsers having a Neotropical distribu-

tion, with North American fossil localities suggesting a distribution

south of glaciated areas during the Pleistocene (Kurten and Anderson

1980).

Order Artiodactyla

Family Tayassuidae

Mylohyus nasutus - Long-nosed Peccary

Material: 1 right dp2 (.306); 1 ml or m2 (.307).

Remarks: This species, referred to as M. pennsylvanicus, was first

reported from South Carolina by Allen (1926). Fossils from Edisto

Island were referred to as M. cf. M. fossilise by Roth and Laerm

(1980). Both names are synonyms of Mylohyus nasutus, following

Kurten and Anderson (1980).

Family Camelidae

Palaeolama mirifica - Stout-legged Llama

Material: 1 partial lower molar (.268); 1 partial upper molar (.269);

1 left ectocuneiform (.270); 1 phalanx distal end (.271); 1 phalanx

proximal two-thirds (.272); 1 hoof core (.273); 1 partial calcaneum

(.274); 1 thoracic vertebra (.275); 1 metatarsal 3&4 proximal end

(.276); 1 astragalus (.277).

Remarks: Identification was based on the presence of a weak "llama

buttress" on the lower molar, and a weakly developed stylid (Webb

1974/?), and a low crowned cheek tooth. Distinctive postcranial remains

were compared directly to other camelid material.

It appears that the northern limits of this stout-legged llama

were generally restricted to low temperate latitudes, e.g., southern

California, the Gulf Coast of Texas, and Edisto Island, South Carolina

(Roth and Laerm 1980), although it has been recorded in Missouri as

Mammals of the Ardis Local Fauna 17

well. It is thought that the diet of this llama consisted of grasses as

well as the shoots and leaves from bushes and trees (Kurten and

Anderson 1980). The Ardis local fauna is the second report of this

species from South Carolina.

Family Cervidae

Subfamily Odocoileinae

Odocoileus virginianus - White-tailed Deer

Material: 1 left Ml (.278); 1 left ml (1 USNM); 1 right Ml (.282);

1 right Ml (1 USNM); 1 left P3 (.279); 1 right p3 (1 UF); 1 right PI

(.283); 1 right m3 (.284); 1 left m2 (1 UF); 2 dm's (.280, .281); 1 left

P2 (1 USNM); 1 right scapula proximal end (.285); 1 humerus distal

end (.305); 1 distal end of calcaneum (.286); 1 astragalus (.287);

3 cubonavicalur (.288, .289), (1 USNM); 1 right scaphoid (.290);

1 trapezoidomagnum (.304); 1 ectocuneiform (.315); 2 right proximal

ulnas (.291, .292); 1 sacral vertebra (.293); 2 distal right radii (.294,

.295) and 1 right radius of sub-adult (.296); 1 left proximal tibio-

fibula (.297); 1 ulna partial proximal end (.298); right articulated (2)

metatarsus and hoof core (.299a-c); 1 right metacarpal (.300); 1 right

rib proximal end (.301) and 1 unspecified rib proximal end (.302); 1

right partial antler including the burr and past first two tines both of

which are missing (1 USNM); 1 right antler (.303).

Remarks: The first fossil record of O. virginianus reported from

South Carolina was based on two basal antler fragments (Allen 1926).

A second record exists from Edisto Island (Roth and Laerm 1980),

based on antler fragments and other postcranial remains, but these

may be of recent origin. The white-tailed deer ranges from Canada

into northern South America, but is absent from hot arid areas of

North America. It occupies a wide variety of habitats today, including

coniferous and deciduous forests, high mountain areas, coastal marshlands,

grasslands, and suburban fringes. It is most commonly found in broken

habitats typical of agricultural areas (Webster et al. 1985).

This deer is very common around the Ardis site today. The

Ardis local fauna appears to have individuals from several different

age cohorts, based on tooth wear patterns.

Family Bovidae

Subfamily Bovinae

Bison antiquus

Material: 1 right m2 (UF); 1 left P4 (.308); 1 left M2 (.309); 1 left p3

(.310); 1 left p2 (.311); 1 right M3 (USNM); 1 left m3 (.312); 2

18 C. C. Bentley. J. L. Knight, and M. A. Knol

lumbar vertebrae (.313), (USNM); 1 metacarpus (.401); 1 lunar scaphoid

pisiform (.314); 1 neural process (USNM), 1 right hoof core (S.C.

92.22.1).

Remarks: Leidy (1860) reported the first bison material from South

Carolina, as B. latifrons. Hay (1923) mentions a fossil which he refers

to as Bison sp., and Allen (1926) identified a single molar as Bison

sp. cf. B. bison. Allen also mentions a horn core and suggests that it

belongs to a bovid. The Edisto Island fossil assemblage (Roth and

Laerm 1980) contains bison material and is identified only as Bison

sp. The molars and postcranial material from the Ardis site are assigned

to B. antiquus based on size. These specimens are larger than modern

B. bison and comparable in size to B. antiquus.

This is the first report of Bison antiquus from South Carolina.

Order Rodentia

Family Sciuridae

Glaucomys volans - Southern Flying Squirrel

Material: 1 left femur (.171).

Remarks: The specimen, a femur broken just below the distal end, is

assigned to G. volans based on the length of the shaft compared to

the width of the proximal end, and overall smaller size compared to

G. sabrinus.

This species is found commonly today in the eastern United

States. It ranges as far north as Canada and southwest into Mexico

and Guatemala (Kurten and Anderson 1980).

Typical habitats are deciduous and mixed hardwood forests. Glaucomys

fossils found in the southeastern United States are commonly collected

from cave deposits frequented by birds of prey during the time of

deposition (Guilday 1962; Guilday et al. 1969, 1977, 1978; Grady

and Garton 1982). The femur is believed either to have been "washed

in" or carried by a non-avian predator. There is no evidence of regurgitated

pellets from roosting raptors, and the entrances and the chambers of

the solution tunnels throughout the site were too small to facilitate

large roosting birds.

This represents the first fossil evidence of a southern flying

squirrel in South Carolina.

Mammals of the Ardis Local Fauna

L9

Q\

% %

D. E.

3 cm

_i

Fig. 3. Comparisons of recent squirrel humeri to the Ardis fossil specimen.

A) Spermophilus franklini (USNM-54153) B) Spermophilus lateralis (USNM-

250744) C) Tamiasciurus hudsonicus (USNM-505592) D) Spermophilus

tridecemlineatus (USNM-255383) E) Spermophilus tridecemlineatus Ardis

local fauna (.175) F) Tamias striatus (USNM-347965).

Sciurus carolinensis - Gray Squirrel

Material: 1 right humerus proximal half (.172); 1 right tibia proximal

half (.173); 1 left tibia distal one third (.174).

Remarks: The material is distinguished from Sciurus niger by its smaller

size and is indistinguishable from recent material of the gray squirrel.

This species ranges throughout the eastern half of the United States

north into Canada, where it inhabits deciduous and coniferous forests,

timbered streams, and bottomlands (Webster et al. 1985).

This is the first fossil record of the gray squirrel from South

Carolina.

Spermophilus tridecemlineatus - Thirteen-lined Ground Squirrel

Material: 1 left humerus (.175).

Remarks: In comparisons with recent humeri of mustelids, microtines,

cricetids, and a variety of ground squirrels, all others were eliminated

because of various morphological inconsistencies. Tamias striatus hu-

meri vary significantly from S. tridecemlineatus with differently shaped

proximal condyles and distal articulations. The humeral shaft is no-

ticeably more slender with respect to the total length in Tamias striatus

than in S. tridecemlineatus. Tamiasciurus hudsonicus humeri have a

20 C. C. Bentley, J. L. Knight, and M. A. Knoll

broader distal end and are considerably larger than S. tridecemlineatus.

The humerus of S. franklinii is significantly larger (2x) and varies

morphologically from S. tridecemlineatus. The fossil humerus is in-

distinguishable from that of Spermophilus tridecemlineatus and given

geographical and temporal considerations is assigned to this species

(Fig. 3).

The thirteen-lined ground squirrel has a modern range reaching

into central Canada, south into eastern Utah and central Texas, and

eastward through the Midwest into Ohio (Hall 1981). The fossil

specimen from the Ardis site is about 1,000 km southeast of its present

day distribution. S. tridecemlineatus inhabits shortgrass prairies and

does not occur in forested areas (Kurten and Anderson 1980). Clark's

Cave and Natural Chimneys, Virginia, and Baker Bluff Cave, eastern

Tennessee, are the closest fossil localities containing remains of S.

tridecemlineatus. Martin and Webb (1974) reported Spermophilus sp.

from Haile 14A in Florida, which is the most southeastern report of

this genus. This ground squirrel's presence in the East has been inter-

preted as indicating parkland or semi-prairie conditions (Guilday et

al. 1977, Kurten and Anderson 1980).

This is the first fossil record of Spermophilus tridecemlineatus

from South Carolina.

Family Castoridae

Castor canadensis - Beaver

Material: 1 left M3 (.176); 1 partial incisor (.177); 1 left M2 (.178);

1 radius (.180); 1 tibio-fibula distal end (.179); 1 metatarsal (.193);

1 phalanx (.181); 1 thoracic vertebra (.182); 1 caudal vertebra (.183).

Remarks: The beaver is found throughout most of North America

except for southern peninsular Florida, the arid regions in the Southwest,

and along Arctic shorelines (Kurten and Anderson 1980). The presence

of this semiaquatic species in the Ardis local fauna indicates the

presence of a nearby wooded, permanent body of water.

Presence in the South Carolina fossil record from Edisto Beach

was first established by Roth and Laerm (1980), although they state

that the single postcranial element may be modern.

Family Muridae

Subfamily Arvicolinae

Ondatra zibethicus - Muskrat

Material: 1 complete skull (.27); 2 partial skulls (1 USNM), (1 UF);

4 parietals (.28-. 30), (1 USNM); both maxillaries and palate missing

Mammals of the Ardis Local Fauna 21

only left Ml (.31); 9 right dentaries (.32-35), (3 USNM), (2 UF);

8 left dentaries (.36-. 38), (2 USNM), (3 UF); 1 left ml (.39); 1 left

m2 (.40); 1 right m2 (.41); 1 right m3 (.49); 1 left m3 (.50); 4 right

Ml (.47, .48), (2 USNM); 4 left Ml (.42-. 45); 1 right M3 (.46), 1 left

M3 (.39); 1 left and 1 right lower incisor (.51, .52); 1 upper left

incisor (.53); 12 left femora (.54-. 59), (3 USNM), (3 UF); 10 right

femora (.60-. 63), (3 USNM), (3 UF); 9 left humeri (.64- .66), (3

USNM), (3 UF); 3 right humeri (.67- .69); 12 left tibio-fibula (.70-

.75), (3 USNM), (3 UF); 13 right tibio-fibula (.76- .82), (3 USNM),

(3 UF), 8 left ulna (.83-. 86), (2 USNM), (2 UF); 9 right ulna (.87-

.89), (3 USNM), (3 UF); 2 radii (.90- .91); 1 partial ilium (.92); 11

left partial innominates (.93- .97), (3 USNM), (3 UF); 11 right partial

innominates (.98-. 104), (2 USNM), (2 UF); 2 left 1st metatarsals (.105,

.106); 2 left 2nd metatarsals (.121, .122); 3 left 3rd metatarsals (.115-

.117); 3 right 1st metatarsals (.118- .120); 7 right 2nd metatarsals

(.107- .109), (2 USNM), (2 UF); 5 right 3rd metatarsals (.110- .112),

(1 USNM), (1 UF); 2 right 4th metatarsals (.113, .114); 3 calcanea

(.123- .125); 1 atlas vertebra (.126); 2 axis vertebra (.127, .128); 1

3rd cervical vertebra (.129); 2 5th cervical vertebrae (.130, 131); 3

6th cervical vertebrae (.132- .133), (1 USNM); 15 caudal vertebrae

(.135- .143), (3 USNM), (3 UF); 3 articulated caudal vertebrae (.134a-

c); 1 lumber vertebra (.144); 1 thoracic vertebra (.145); 9 vertebrae

(.146- .148), (3 USNM), (3 UF); 1 vertebra (.150); 1 sacral vertebra

(1 USNM); 1 sacral vertebra and two associated caudal vertebra (1

UF); 1 partial hyoid process (.149); 2 proximal rib halves (.151, .152).

Remarks: The muskrat is by far the most common mammal from the

Ardis local fauna. The relatively high number of muskrat remains

from the site suggests that they may have been using the solution

cavities as temporary shelters. Given that the deposit at the Ardis

local fauna represents fluvial episodic events, these cavities may have

provided excellent temporary shelters if muskrats retreated from their

usual shelters onto higher ground during periodic flooding. This behavior

has been observed in muskrats using multiple shelters in areas with

seasonal fluctuations in water levels (Brooks 1985). In addition, pieces

of fossil turtle shell collected from the site had gnaw marks of a large

rodent, presumably muskrats, suggesting that feeding may have occurred

in these cavities. It is unlikely that the turtle shells were "wash-ins,"

as no signs of weathering, water wear, or abrasive breakage were

evident. There have been many reports of contemporary muskrats

feeding upon turtles (Errington 1941, Doutt et al. 1966, Parmalee

1989), but no reports of this have been recorded from the fossil record.

C. C. Bentley, J. L. Knight, and M. A. Knoll

The muskrat is found throughout much of North America, including

all of South Carolina. This is the most aquatic of the microtine rodents

and is usually found in close proximity to fresh or brackish waters.

This is the first fossil record of this species from South Carolina.

Neofiber alleni - Round-tailed Muskrat (Florida Water Rat)

Material: 1 right Ml (.153); 1 fragmentary molar (.154).

Remarks: N. alleni is endemic to Florida and southernmost Georgia,

and its northward limits may be maintained by aridity and cold tem-

peratures (Frazier 1977). It is not currently sympatric with O. zibethicus.

The enamel above the jaw line on the partial molar appears to

be chemically etched or corroded. This may be due to the ingestion of

this specimen by a carnivore, at which point only the dorsal portion

of the tooth would be exposed to stomach acids (Gary Morgan, Florida

Museum of Natural History, personal communications). Separation of

the molar from the dentary bone may have occurred during or after

fossilization. This species is a good indicator of nearby bodies of

permanent water; its diet consists mostly of aquatic vegetation. It

builds its nest in areas such as open savannahs, mangroves, and in

suitable stumps (Kurten and Anderson 1980).

This is the first fossil record of the round-tailed muskrat outside

of Florida and Georgia.

Synaptomys cooperi - Southern Bog Lemming

Material: 1 right Ml (.155).

Remarks: The molar from the Ardis site compares favorably to recent

specimens of this species. Separation of Synaptomys cooperi from

S. australis is based on size, as S. cooperi is generally 35% smaller

than S. australis (Simpson 1928, Olsen 1958). The occlusal length

(2.2 mm) of the Ardis Ml referred to S. cooperi is comparable to the

lowest range given by Guilday et al. (1977).

The southern bog lemming can be found in habitats that include

grasslands, moist meadows, woodlands, thickets, weedy fields, and

bogs (Webster et al. 1985).

Synaptomys cooperi does not occur in the State today; the nearest

populations are found in the Piedmont and mountains of Virginia and

North Carolina, respectively, extending north into Maryland, and westward

into Kansas and Nebraska (Hall 1981). S. cooperi also occurs in the

Great Dismal swamp and on the Coastal Plain of North Carolina

(Clark et al. 1993).

This is the first fossil record of the southern bog lemming from

South Carolina.

Mammals of the Ardis Local Fauna 23

Synaptomys australis - Florida Bog Lemming

Material: 1 right Ml (.156).

Remarks: This species, though morphologically similar to S. cooperi,

was distinguished by its significantly larger occlusal surface length of

3.4 mm than S. cooperi (Simpson 1928, Olsen 1958). Apparently this

is the first sympatric occurrence of S. cooperi and S. australis in a

fauna known to be contemporaneous. Both species were recovered

from Ladds quarry, Georgia, but may have come from strata of two

different ages in the deposit (Kurten and Anderson 1980).

There has been some debate over whether or not the extinct

S. australis is a full species, represents a cline for greater body size,

or a large subspecies of S. cooperi. Their sympatric occurrence in the

Ardis Local Fauna suggests that they were distinct.

This is the first fossil record of this species from South Carolina.

Microtus pennsylvanicus - Meadow Vole

Material: 2 right dentaries one complete (.158) and the other containing

only the ml (.159); 2 left dentaries, one missing its proximal third

and the m3 (1 USNM), and one fragment containing only the ml

(.160); 2 palatines, one with right Ml and M2 along with zygomatic

arch (.161) and one with right Ml and the left M2 (.162).

Remarks: The meadow vole today inhabits a wide variety of habitats

including upland grasslands, meadows, swamps, stream borders, salt

marshes, and forests (Webster et al. 1985).

M. pennsylvanicus is commonly collected from Pleistocene sites

in North America (Kurten and Anderson 1980), and now occurs in the

Piedmont and in isolated populations near Charleston, South Carolina,

and Cedar Key, Florida (Webster et al. 1985, Woods et al. 1982).

M. pennsylvanicus does not currently occur in the vicinity of the

Ardis fauna.

This is the first fossil record of this species from South Carolina.

Microtus pinetorum - Woodland Vole

Material: 1 right dentary with a complete dentition (.163); 1 right ml

(.164); 2 left dentaries, one missing proximal third and m3 (.165),

one missing proximal third and m2 and m3 (.166); 1 maxilla with left

Ml and the right M3 (.167).

Remarks: Identification was based on criteria established by Martin

and Webb (1974) and Martin (1991). The woodland vole typically

24 C. C. Bentley, J. L. Knight, and M. A. Knoll

inhabits woodland and old-field habitats and is well known for build-

ing extensive shallow tunnels (Webster et al. 1985). M. pinetorum

can be found throughout South Carolina and is common in the pine

forests of the South and in eastern deciduous forests (Kurten and

Anderson 1980).

This is the first fossil record of this species from South Carolina.

Subfamily Sigmodontinae

Oryzomys palustris. - Marsh Rice Rat

Material: 1 left dentary lacking dentition (.175).

Remarks: Identification was based on several features, including

double rooted molars, mental foramen at base of Ml, diastema short

and robust (compared to Peromyscus), anterior root of Ml ventral and

offset from other roots, foramen next to M3 much larger than Peromyscus,

and over-all size correlation to Oryzomys, and direct comparisons to

recent specimens.

The marsh rice rat inhabits marshlands, meadows, and wet grass-

lands on the Coastal Plain of South Carolina and has been recorded

along a mountain stream in South Carolina (Webster et al. 1985).

This specimen represents the first fossil record of this species

from the State.

Peromyscus sp.

Material: 1 right dentary containing only the incisor (.168).

Remarks: Identification was based on the alveolar length of the tooth

row, and similarities in morphology and size when compared to recent

Peromyscus. The tooth row is longer than that of Reithrodontomys.

Several species of Peromyscus inhabit the northwestern portion

of South Carolina. P. gossypinus, the largest member of this genus

inhabiting the State, is the only species of Peromyscus that can be

found today in the area of the Ardis site. The robustness of the fossil

specimen suggests this species, but, given the presence of numerous

extralimital species in the fauna, it is not possible to assign the fossil

to a particular species of Peromyscus with any certainty.

Neotoma floridana - Eastern Woodrat

Material: 1 left Ml (.169); 1 right innominate bone (.170).

Remarks: Woodrats are notorious collectors of natural history ob-

jects. Apparent rarity of this species in the Ardis local fauna notwith-

Mammals of the Ardis Local Fauna 25

standing, activities of this species may explain some of the more

unusual occurrences of other vertebrate species in the solution cavi-

ties. The three-rooted Ml compares favorably in morphology and site

(anterior-posterior length of Ml = 3.52 mm) to recent material. N.

floridana is the only species of woodrat found in the southeastern

United States.

This represents the first fossil record of this species from South

Carolina.

Family Hydrochaeridae

Material: 1 fragment of the posterior lamina of the M3 (.184).

Remarks: Identification to family was based on direct comparison

to fossil and recent specimens. However, the fragmentary nature of

the specimen precludes identification to genus or species.

The extremely thin enamel and the particular angle of the fragment

are diagnostic and cannot belong to any other taxon.

One living genus of capybara {Hydrochaeris) occurs in tropical

habitats in Central and South America and is semiaquatic, commonly

found along the edges of streams and the borders of marshes (Kurten

and Anderson 1980). The presence of this tropical family in the fossil

record was first established for South Carolina by Roth and Laerm

(1980).

Order Lagomorpha

Family Leporidae

Sylvilagus palustris - Marsh Rabbit

Material: 4 right dentaries (.185-. 187), (1 UF); 3 left dentaries (.188-

.189), (1 USNM); 1 isolated p3 (.190).

Remarks: Identification was based on the presence of multiple anterior

reentrants on the third premolar.

The marsh rabbit is a good swimmer and can be found in wet-

lands areas such as marshes, flood plains, and hummocks, and its

modern distribution ranges from southeastern Virginia along the

Atlantic Coastal Plain into Florida (Webster et al. 1985). S. palustris

probably occurs in the immediate vicinity of the Ardis site. It has

been recorded from many Pleistocene sites in Florida (Webb 1974,

Kurten and Anderson 1980), and Sylvilagus sp. was identified from

Edisto Island, South Carolina (Roth and Laerm 1980).

This represents the first fossil material identified as this species

from South Carolina.

26 C. C. Bentley, J. L. Knight, and M. A. Knoll

Sylvilagus floridanus - Eastern Cottontail

Material: 2 left (.191, .192) and 1 right dentary (1 USNM).

Remarks: Remains were identified by complete absence or presence

of a single anterior reentrant on the p3.

This species is distributed from southern Canada south to Argentina

(Kurten and Anderson 1980). The eastern cottontail typically inhabits

areas with a mixture of herbaceous and shrubby plants in a disturbed

environment at some stage of successional transition that occur in

and among a variety of habitats (Webster et al. 1985). The eastern

cottontail occurs in the area of the Ardis site today.

This is the first fossil record of this species reported from

South Carolina.

Sylvilagus sp.

Material: 2 right distal femora fragments (.213, .214); 3 left partial

femora (.215, .216), (USNM); 1 right humerus (.217); 1 left distal

half of a humerus (.218); 1 proximal end of a humerus (.226); 1 right

partial scapula (.219); 2 distal and 1 proximal tibio-fibia ends (3

USNM); 1 left partial radius (.220); 1 ulna proximal end (.221); 4 left

partial innominate bones (.222, .223), (2 USNM); 3 left calcanea

(.224), (2 UF); 3 right calcanea (.225), (2 USNM).

Remarks: All of the leporid postcranial elements from the Ardis local

fauna represent this genus, but we are unable to assign these elements

to a particular species in this genus.

DISCUSSION

Paleoecology and Paleoclimate

The Ardis local fauna is one of only a handful of Rancholabrean

sites reported from the Atlantic Coastal Plain north of Florida, and is

the only C14 dated fauna from South Carolina. It was deposited during

the full glacial phase of the Wisconsin. This interval is poorly represented

in the fossil record, with New Trout Cave, West Virginia (Grady and

Garton 1982) and Bakers Bluff, Tennessee (Guilday et al. 1978) being

the only other fossil sites in the southeastern United States known to

be temporally similar.

Two biases must be considered which may have skewed types

and frequency of remains recovered from the Ardis site: 1) collection

bias; a disproportional amount of the larger "easier to see" material

was collected. Comparatively little of the fossiliferous sediments was

screen-washed to retrieve the smaller material otherwise easily missed

Mammals of the Ardis Local Fauna 27

and 2) depositional and behavioral biases; the smaller taxa, particularly

those that used this site as a shelter or in hunting, should be represented

in disproportionately higher numbers when compared to the megafauna.

Behavioral and environmental factors influence the occurrence of the

smaller taxa which could easily enter the cavities from the surface.

Furthermore, some taxa may have been concentrated in the stomachs

of predators who were subsequently entombed in the cavities. The

above-mentioned factors, excluding predation, but including other taphonomic

factors, especially the size-restrictive nature of the cavity openings,

would govern the lower frequency of the larger megafauna occurrence

in the cavities. Therefore, the species composition most likely is not

proportional to its true occurrence in this particular Pleistocene community.

Relatively large faunal diversity may give a reasonably reliable

picture of the surrounding habitat (Guilday 1962). The taxa of the

Ardis local fauna represent a diversity of ecological niches including

semi-aquatic forms (Castor, Lontra, Ondatra, Tapirus, and Mustela

vison), arboreal forms (Glaucomys volans, Sciurus carolinensis), marsh

and meadow inhabitants (Synaptomys, Microtus, Oryzomys), grassland

or prairie forest transition inhabitant (Spermophilus tridecemlineatus),

and the large grazers and browsers (Palaeolama, Bison, Equus, Odocoileus,

Megalonyx, Mammut, and Mammuthus), suggesting that the Ardis fauna

sampled an ecological mosaic of community types. The depositional

features and fauna suggest a composite conifer and hardwood forest,

interspersed or bordered by a grassland/meadow, possibly giving way

in low-lying areas to a marsh or bog, with a permanent nearby stream

or river.

Hypothesized changes in vegetation during the late Pleistocene

(Dreimanis 1968), with climatic conditions unlike any experienced in

North America today (COHMAP Members 1988), are reflected in the

extralimital tropical to boreal species found in the Ardis fauna.

Of the 43 mammalian species collected from the site, 27 are

extant, 21 still occurring in the area today. Of the six extant extralimital

taxa, four have more northern affinities, one a midwestern affinity,

whereas only one has a range well south of the Ardis site. Of the 16

extinct taxa, five have affinities considerably south and west of the

Ardis locality (Kurten and Anderson 1980, Martin 1978).

The Ardis mammal fauna exhibits a mixture of southern, west-

ern, and northern forms, resulting from the radiation and convergence

onto the lower Atlantic Coastal Plain of taxa migrating along the

Gulf Coast corridor (Webb 1974) and taxa migrating from the north-

west Appalachian Mountains region. It is plausible that Spermophilus

tridecemineatus may have entered South Carolina from a northern

28 C. C. Bentley, J. L. Knight, and M. A. Knoll

route instead of along the Gulf Coast corridor, as attributed to the

fossil remains from the Haile 14A fauna (Webb 1974). Fossil locali-

ties reporting S. tridecemlineatus (Kurten and Anderson 1980) and

other fossils collected from the Ardis site, in particular a portion of

the turtle fauna (Bentley and Knight, submitted), strongly suggests

the existence of a northern corridor(s) onto the Atlantic Coastal Plain.

This would be a logical pathway for those glacially displaced species

of the Northeast found at the Ardis site.

Blarina brevicauda, Microtus pennsylvanicus, and Synaptomys

cooperi are interpreted as "boreal" or "cool climate" components of

the Ardis fauna, based on modern distribution and habitat orientation

(Hoffman and Jones 1970, Graham 1976, Webster et al. 1985). Neofiber

alleni is considered a sub-tropical or "warm climate" species based on

modern distribution and fossil records (Martin and Webb 1974, Frazier

1977, Kurten and Anderson 1980, Holman 1985).

Those extinct species collected from the Ardis fauna with extant

genera or families primarily tropical to sub-tropical in distribution

include: Tremarctos floridanus, Palaeolama mirifica, Conepatus robustus,

Tapirus veroensis, and the family Hydrochoeridae. Holmesina septentrionalis

has also been interpreted as indicating a mild climate (Kurten and Anderson

1980).

Generalization of ecological needs of an extinct species based on

the needs of extant relatives cannot always be considered reliable.

However, when several such groups are geologically recent, are in a

single locality, and represent a short depositional time interval, as at

the Ardis site, these assumptions become more credible.

The modern Coastal Plain fauna of South Carolina has six

species of microtines, three species of shrews, and two mole species

(Webster et al. 1985). Fossil remains of both mole species and two of

the three shrew species were collected from the Ardis fauna, along

with eight microtine species. Greater microtine densities in late Pleis-

tocene faunas have been correlated with reduced temperature and

moisture gradients (Graham 1976). This is also true for shrew species

(Graham 1976), but we did not observe it in the Ardis fauna. The

reduction in the number of shrew species at the Ardis site is probably

a result of collection bias and not a true reflection of the shrew

populations.

Martin (1968) suggested that the distribution of M. pennsylvanicus

is limited mostly by warmer temperatures and drier summers, but also

by the presence of Sigmodon hispidus, which was inexplicably absent

from the Ardis fauna. The mean July temperature for the southern

boundary of M. pennsylvanicus is 23.9 ± 1.1 C (Martin 1968), and

Mammals of the Ardis Local Fauna 29

the two species occur sympatrically in areas where the mean July

temperature is near 26.7 C (Martin 1968). Today, S. hispidus is com-

mon throughout South Carolina, but M. pennsylvanicus is present

only in the extreme western Piedmont and in a small relict population

near Charleston, South Carolina (Webster et al. 1985). Data from

Charleston, South Carolina, over the past 75 years yielded a July

daytime mean of 31.1 C and a minimum daily mean of 23.9 C (Pearce

and Smith 1984). Vegetational data for the late Wisconsin full glacia-

tion, taken from White Pond in South Carolina (Watts 1980), sug-

gested a 7 C to 20 C decrease in July temperatures compared to

present, and possibly a marginal reduction in precipitation. The pres-

ence of M. pennsylvanicus in the Ardis fauna and the absence of S.

hispidus may indicate slightly drier conditions and temperatures be-

low the mean summer temperature experienced today.

It has been postulated that mammals in the late Pleistocene reacted

to environmental changes based on their own tolerance limits and not

as a "community unit" (Graham 1976, 1979). This accounts for northern

species pushed south by glaciation and then integrating with the existing

biota. Furthermore, southern species also were integrated into the

resident community as cooler summers and warmer winters prevailed,

producing the "disharmonious fauna" collected from the Ardis fauna.

In general, the mammalian composition of the Ardis fauna, containing

distinctly southern, northern, and western extralimital forms, reflects

a climate more equitable than present. The less severe climatic gradients

would facilitate the sympatric occurrence of species now ecologically

incompatible. The Ardis local fauna coincides well with other late

Pleistocene fossil localities reporting disharmonious faunas with similar

temporal and topographical settings. This suggests that the late Pleistocene

in the southeastern United States was climatically more equitable and

ecologically more diverse prior to the dramatic shift towards a modern

assemblage, approximately 10,000 to 11,000 y.b.p. (Lundelius et al.

1983).

ACKNOWLEDGMENTS— First and foremost we wish to thank

personnel of the Giant Cement Plant for their generous cooperation

and patience. They provided, at our request, heavy equipment, opera-

tor time, and rerouting of mining operations, to give us maximum

possible time for excavations, and nearly unlimited access to the site.

A special acknowledgment goes to the plant's safety director, Bert

Ardis, for whom the fauna is named. Mr. Ardis went out of his way to

ensure our access to the quarry, to provide us food and refreshments,

a place to sleep and shower, and field assistance far beyond our

hopes.

30 C. C. Bentley, J. L. Knight, and M. A. Knoll

We also wish to recognize all those who toiled along with us in

the field: Vance McCollum, Linda Eberle, Craig and Alice Healy,

Derwin Hudson, Ray Ogilvie, Lee Hudson, Tom Reeves, Suzanne

Boehme, George Beighley, Jr., Martha Bentley, and Karin Knight.

We are grateful to David Webb and Gary Morgan of the Florida

Museum of Natural History, for allowing us access to the UF collec-

tion and critical reviews of this manuscript. We also wish to thank

Gary Morgan for his aid in the identification of ambiguous species.

Robert Martin kindly provided us with his unpublished manuscript,

helped us with initial identification of the microtines, and discussed

muskrat evolution. We also wish to express our appreciation to Mike

Trinkley and Debi Hacker of the Chicora Foundation, Columbia, South

Carolina, for the frequent use of their field equipment; Michael Runyon,

Lander University, for donating mammoth bone samples for dating

purposes; Claudia Angle, United States Fish and Wildlife Service;

and Bob Purdy and Clayton Ray of the National Museum of Natural

History for their courtesies extended. Wallace Dawson and Fontelle

Thompson of the University of South Carolina reviewed an earlier

draft of this paper. Finally, thanks go to Overton Ganong, Director of

the South Carolina State Museum, and Darby Erd for the superb illus-

trations.

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Received 30 November 1993

Accepted 27 January 1994

APPENDIX I

Taxa and Minimum Number of Individuals

Taxa Minimum Number of Individuals

Didelphis virginiana* 6

fDasypus bellus 1

^Holmesina septentrionalis(S) 1

"\Megalonyx jeffersoni 1

Sorex sp. cf. S. longirostris* 1

Blarina brevicauda*(N) 3

Scalopus aquaticus* 11

Condylura cristata* 2

Urocyon cf. U. cinereoargenteus 1

Mammals of the Ardis Local Faun;

35

t Can is dims

fcf.Smilodon fatalis

Lynx rufus*

Lontra canadensis*

Spilogale putorius*(N)

Meph itis m eph itis *

fConepatus robustus*(S)

Mustela vison*

Procyon lotor

fTremarctos floridanus(S)

fMammut americanum

t Ma mm u thus columbi

fEquus cf. E. complicatus

f Tapir us veroensis

iMylohyus nasutus

fPalaeolama mirifica(S)

Odocoileus virginianus

fBison antiquus

Glaucomys volans*

Sciurus carolinensis*

Spermophilus tridecemlineatus*(MW)

Castor canadensis

Ondatra zibethicus*

Neofiber alleni*(S)

Synaptomys cooperi*(N)

"\Synaptomys australis*(S)

Microtus pennsylvanicus* (N)

Microtus pinetorum*

Oryzomys palustris*

Peromyscus sp.*

Neotoma floridana*

Hydrochoeridae(S)

Sylvilagus palustris*

Sylvilagus floridanus*

Faunal list = 43 species

13

4

2

Total 89

* = first fossil record from South Carolina

(N) = extralimital northern (S) = extralimital southern

(MW) = extralimital mid-western

t = Extinct taxa

36

Comments on the Body Mass Trend

of Ondatra zibethicus (Rodentia: Muridae)

During the Latest Pleistocene

Curtis C. Bentley and James L. Knight

South Carolina State Museum, 301 Gervais Street

P.O. Box 100107, Columbia, South Carolina 29202-3107

ABSTRACT — Martin (1993) suggested, in his investigation of the

phyletic evolution in the rodent genus Ondatra, that an increase

in body mass through time has occurred in Ondatra zibethicus,

with the increase in size being concentrated in the last 600,000

years before present (y.b.p.). Ondatra zibethicus apparently obtained

its greatest body mass during the latest Pleistocene, followed by

a sharp decrease in body mass into Recent times, referred to as

a "dwarfing event." We examined fossil muskrats from late Pleistocene

sites in South Carolina, Florida, and additional Recent material

which do not support the proposed "dwarfing event" of O. zibethicus

at the close of the Pleistocene. As more fossil material becomes

available, future research could provide a clearer picture of the

body mass trend in Ondatra zibethicus.

METHODS

Two recently discovered late Pleistocene sites, Crowfield local

fauna and the Ardis local fauna (Bentley et al. 1994) yielded fossils

of Ondatra zibethicus. These remains, coupled with unpublished

material from the Aucilla River, Florida, and Recent specimens from

Iowa and Georgia provided new data elucidating body mass trend for

this taxon during the latest Pleistocene. It is not within the scope of

this paper to do a thorough review of the literature on muskrat body

mass nor extensive studies of museum collections. A project of this

magnitude would encompass an undertaking much larger than this

note. The data published here are intended to elicit further research

into the trends of muskrat body mass at the close of the Pleistocene.

Boyce (1978) noted a small degree of sexual dimorphism in

muskrats with males slightly larger than females. However, because

this difference is slight, and sex cannot be determined from fragmen-

tary fossil material, the effects of sexual dimorphism can not be assessed

in this study.

All measurements were done on the first lower molar (ml).

Measurements were taken three times with calipers and rounded to

the nearest 0.01 mm.

Brimlevana 21:37-43. December 1994 37

38

Curtis C. Bentley and James L. Knight

- E

E E

2 Z

E E

^ z

^ c.

— — r- r-- n

\H Q

en

x t^ x — r- r~- r~- r- r- n -

(N d n r, m M n tN r>) r, f

x c-~ in o^ x

C") rn ro co <o ro

r*~, co cs

^222

C/5 C/5 C/3 C/5 on

U U U U U

en x c/3 x x

c> c?ch?r, iy, y-ch-r, — i^sC in —

ri r, >c — — x (^ — x x t-- — xxxr^r^— ? ri

IX Q

x

in in in — in n <n in in in

C — t-- c> x oj c x n rn x x cn — — x c x

zzzzz ^ ^ ^ ^ ^ -

^^^^^^T^c^^PxZZ^zzz 3

-J -J — -J -J c~< c*i <■*", co co r^, x x r»i c/j en x ■

r-r-xxr^(^vCi^t^r^xr^scr^r^scr-i^r-r--r-sC

Muskrat Body Mass Trend 39

Fossil specimens from the Ardis local fauna, deposited in the

collections of the South Carolina State Museum (SCSM), are designated

by the base number S. C. 93.105. and cited in this paper by the two

digits following the base number. Specimens from the Crowfield

local fauna and Recent unaccessioned specimens at the South

Carolina State Museum are denoted by SCSM. Fossil material from

the Florida Museum of Natural History is indicated by the accession

numbers of 132680 - 131318.

We used a weighted, Wilcoxon rank analysis test to search for

statistical significance between our data and Martin's. The data were

analyzed over three time intervals: 1) 20.000 y.b.p. to Recent; 2)

20,000 y.b.p. to 10,000 y.b.p.; 3) 20,000 y.b.p. to 15,000 y.b.p. The

Wilcoxon rank analysis test was used to compare sample means, as

Martin's raw data were not available to us.

INTRODUCTION

Martin (1993) derived a regression formula based on the length

of the ml to estimate the body mass of arvicolines: M = 0.71 (I)359,

where M is body mass in kg and L is the length of ml in mm. He used

the formula to help determine the trend in body mass for the polytypic

genus Ondatra during the last 3.75 million years. Martin noted that

most of the change in body mass occurred during the last 600.000

years. Muskrats reached their greatest size (1.75 kg) during the latest

Wisconsin between 20.000 and 10,000 y.b.p. This larger form was

Ondatra zibethicus floridanus (Lawrence 1942). synonymized with

O. zibethicus by Martin (1993). Martin conjectured that approximately

10,000 years ago it appears that body mass dramatically decreased to

the levels he recorded for Recent samples. Martin referred to this

decline in body mass as a "dwarfing event.*' He was unclear as to the

cause but mentioned human culling and/or natural selection as possible

explanations. Applying Martin's regression formula to fossils from

several recently collected late Pleistocene sites in South Carolina and

Florida, as well as recent specimens from Iowa and Georgia (Table

1). gives more resolution to this short, but apparently dynamic time

interval.

RESULTS AND DISCUSSION

The Ardis local fauna (19.000 y.b.p.) (Bentley et al. 1994)

yielded 18 Ondatra mi's for which measurements could be taken

(Table 1). producing a mean body mass of 0.95 kg. An unpublished

fauna from South Carolina, the Crowfield local fauna (80.000 y.b.p.),

40 Curtis C. Bentley and James L. Knight

under study by Fred Grady of the National Museum of Natural

History, produced a sample of six muskrat mi's giving a mean of 0.92

kg. In Florida, an unpublished, late Pleistocene site (12,000-10,000

y.b.p.), from the Aucilla River (Priscilla site, Little River section),

under study by S. David Webb (Florida Museum of Natural History,

personal communication), yielded 15 mi's with a mean of 1.04 kg.

The age of this site is based on numerous radiocarbon dates (Dunbar

et al. 1989; S. D. Webb, personal communication).

The senior author obtained 22 muskrat carcasses from a fur

buyer in Roselle, Carroll County, Iowa, in December 1992. The sample

represents muskrats from populations within a 100-mile radius of Roselle

and were probably collected from many different sites. Using Martin's

regression formula, these samples produced a mean of 0.94 kg. Measure-

ments of two modern mi's from Georgia (no other data) had estimated

mean mass of 1.27 kg.

Comparing these data (Fig. 1) with those of Martin (unpublished

data) would suggest that the increase in body mass for the time span

covered by our samples was much more subtle than Martin's data

would indicate. After Martin's initial body mass increase at 600,000

y.b.p., the upward change in size until Recent is almost negligible. In

addition, the statistical analysis of the three time intervals yielded no

significant difference between our data and Martin's. Thus, our data

do not support Martin's ''dwarfing event" at the close of the Pleistocene.

We view the data on which the dwarfing hypothesis was based

as having several problems, the most significant of which is sample

size. The four samples between 10,000 and 20,000 y.b.p. which constitute

the much larger forms of O. zibethicus (compared to Recent specimens)

are made up of a total of seven mi's, from four sites (Table 2), an

extremely small sample size. Furthermore, all of Martin's 10,000-

20,000 y.b.p. specimens were recovered from cave faunas (Table 2),

a habitat in which modern muskrat populations do not naturally occur.

This would suggest that various selection pressures, probably predatory,

must be taking place in order for these remains to occur in cave

deposits. Thus, the samples from these caves may not be a true representation

of nearby local populations.

Assuming that Martin's regression for body mass holds true, it

seems highly probable that a "dwarfing event" has not occurred between

the latest Pleistocene and contemporary times based on this new material.

However, this has little consequence for the overall trend in muskrat

body mass over the past 3.75 million years. When this time period is

re-scaled to a common interval length, Martin (1993) states that the

dwarfing event ". . . appears to be only moderately pronounced."

Muskrat Body Mass Trend

41

1.8

1.6

1.2

0.8

0.6-

0.4

•500 -450 -400

-350 -300 -250 -200 -150

Time YBP (Thousands)

Fig. 1. Mean body mass trend of Ondatra zibethicus. Solid triangles:

sample means for non-cave faunas reported by Martin (1993). Plus

symbols: sample means for cave faunas reported by Martin (1993).

Open squares: sample means for non-cave faunas (from Table 1). Overlapping

samples are denoted by two location numbers. Numbers and letters

indicate location of the samples (from Table 2).

■500

-400

-350 -300 -250 -200 -150

Time YBP (Thousands)

Fig. 2. Comparison of raw versus mean data points for Ondatra zibethicus.

Solid triangles and plus symbols are Martins' (1993) sample means.

Open squares are Bentley and Knight's raw data (from Table 1).

42

Curtis C. Bentley and James L. Knight

Table 2. Location, sample size, and age of muskrat specimens.

Locality

Sample Size Age (y.b.p.) Source

Martin (unpublished)

We believe the "dwarfing event" is an artifact of small sample

sizes and selection bias and not a dramatic evolutionary response to

some environmental change. However, further data are needed to provide

a more definitive answer to the true body mass trend of Ondatra zibethicus

during the latest Pleistocene. As fossil collections are amassed from

the latest Pleistocene a clearer picture may develop, resolving trends

that cannot be discerned here.

ACKNOWLEDGMENTS— First and foremost we wish to thank

personnel of the Giant Cement Plant, Harleyville, South Carolina, for

their generous cooperation and patience while we worked in their

quarry.

We also wish to recognize all those who toiled along with us

in the field: Vance McCollum, Linda Eberle, Craig and Alice Healy,

Derwin Hudson, Ray Ogilvie, Lee Hudson, Tom Reeves, Suzanne

Boehme, Martha Bentley, and Karin Knight. We are grateful to S.

David Webb and Gary Morgan, Florida Museum of Natural History,

Gainesville, Florida, for allowing us access to the University of Florida

collection and for critical reviews of this manuscript. Particular thanks

Muskrat Body Mass Trend 43

go to Robert A. Martin who kindly provided us with his unpublished

manuscript, and whose critical review of this work greatly improved

its relevance. We also wish to express our appreciation to Mike Trinkley

and Debi Hacker of the Chicora Foundation, Columbia, South Carolina,

and to David Wethey, Sara Woodin, and Wallace Dawson for their

input on this manuscript. Finally, thanks go to Overton Ganong, Director

of the South Carolina State Museum, for other courtesies.

LITERATURE CITED

Bentley, C. C, J. L. Knight, M. A. Knoll. 1994. The mammals of

the Ardis local fauna (late Pleistocene), Harleyville, South Caro-

lina. Brimleyana 21:1-35.

Boyce, S. Mark. 1978. Climatic variability and body size variation

in the muskrat (Ondatra zibethicus) of North America. Oecologia

(Bed.) 36:1-19.

Dunbar, J., D. Webb, D. Crane. 1989. Culturally and naturally modified

bones from a paleoindian site in the Aucilla River, northern Florida.

Pages 473-497 in Bone modification. (R. Bonnichsen and M. Sorg,

editors). Center for the Study of the First Americans. Oregon

State University, Corvallis.

Lawrence, B. 1942. The muskrat in Florida. Proceedings of the New

England Zoological Club 19:17-20.

Martin, R. A. 1993. Patterns of variation and speciation in Quater-

nary Rodents. Pages 1-16 in Morphological change in Quarternary

mammals of North America. (R. A. Martin and A. D. Barnosky,

editors). Cambridge University Press, Cambridge, England.

Received 8 March 1994

Accepted 30 August 1994

44

Introductions of the Scorpions Centruroides vittatus

(Say) and C. hentzi (Banks) into North Carolina, with

Records of the Indigenous Scorpion,

Vaejovis carolinianus (Beauvois)

(Scorpionida: Buthidae, Vaejovidae)

Rowland M. Shelley

North Carolina State Museum of Natural Sciences, P. O. Box 29555,

Raleigh, North Carolina 27626-0555

ABSTRACT — The scorpions Centruroides vittatus (Say) and

C. hentzi (Banks), with subaculear tubercles on their telsons,

have been accidentally imported into piedmont and coastal North

Carolina and may become established in parts of these regions.

They are distinguished by the larger size of C. vittatus and by

the following differences in pigmentation: the presence of a darkly

pigmented, inverted triangular patch on the cephalothorax of C.

vittatus, as opposed to light mottled brownish coloration in C.

hentzi, and by the reticulated brown pigmentation on the dorsal

surfaces of the chelicerae of C. hentzi, in contrast to the unpigmented

condition in C. vittatus. The native scorpion, Vaejovis carolinianus

(Beauvois), which lacks the subaculear tubercle, occurs in southwestern

border counties adjoining South Carolina and Georgia and has

penetrated the western fringe of the State, occurring just inside

the Tennessee state line in the French Broad and Little Tennessee

river valleys. It is also recorded from Yancey, Haywood, Mecklenburg,

Iredell, Guilford, Wake, and Columbus counties, all probably representing

accidental human importations. A key, descriptive drawings, and

a map of occurrences are presented.

In April 1991, I was notified that employees in a north Raleigh

office building had encountered and trapped a scorpion in a hallway.

Vaejovis1 carolinianus (Beauvois) (family Vaejovidae), occurring

in southwestern border counties adjoining South Carolina and Georgia,

some 208 mi (333 km) from Raleigh (Shelley 1975a, b), is the only

scorpion native to North Carolina, so I was surprised to find

that they had a specimen of Centruroides vittatus (Say) (family

Buthidae), a species occurring in the southcentral and southwestern

United States and the adjacent states in northern Mexico (Stahnke and

Calos 1977). This individual had been unknowingly transported to

North Carolina in a shipment of mesquite lumber for an adjoining

1 Francke (1977) showed that Vaejovis and Vaejovidae, with an "a", are the correct

spellings for the genus and family, respectively, as opposed to the previous

orthography without this vowel.

Brimleyana 21:45-55, December 1994 45

46

Rowland M. Shelley

steak restaurant and had wandered into the office building. Occupants

reported seeing occasional scorpions for a year previously, but

thorough searches of the building, its grounds, and a nearby rocky

ditch in both daytime and at night, using a black light, produced no

more specimens. Shortly afterwards I learned that individuals of C.

vittatus had been encountered near downtown Raleigh and in a building

in the Research Triangle Park, in both cases near restaurants using

mesquite to broil steaks. The species has also been collected in Nash

and Dare counties, and a Florida scorpion, C. hentzi (Banks), has been

discovered in Carteret and Brunswick counties, on the North Carolina

coast, and in Durham County in the Piedmont. Thus, three scorpions

(dorsal views in Fig. 1) may now be encountered in North Carolina.

The origins of most accidental animal introductions cannot be

traced, but if reproducing populations of either C. vittatus or C. hentzi

become established in North Carolina through the introduction of a

gravid female or a mating pair, I believe they will have resulted from

.:*

i

J

Fig. 1. Dorsal views of, left to right, V. carolinianus, C. vittatus, and

C. hentzi.

Scorpions of North Carolina 47

human introductions that occurred primarily in the late 1980's and the

early 1990's. Both species can potentially survive and reproduce in

parts of North Carolina, thereby becoming components of the State's

fauna. Likewise, V. carolinianus can potentially survive and reproduce

north and east of Polk, and the adjacent fringe of Rutherford, counties.

If such populations are ever discovered, they too will date from the

late 1980s and early 1990s.

These scorpions are not dangerous to man, their stings being

roughly comparable to those of bees and wasps. Muma (1967) reported

that the sting of C. hentzi produced a localized burning sensation and

that the area was tender for a few hours; the venom of C. vittatus

produces a similar, though more painful, reaction. No information is

available on the venom of V. carolinianus, but no species of this genus

is known to be harmful (Muma 1967).

For the benefit of local naturalists, I announce the discovery of

these non-native arachnids in North Carolina, publish the available

records along with an identification key and pertinent illustrations, and

update the known localities of V. carolinianus. Acronyms of

sources of preserved study material are as follows:

AMNH - American Museum of Natural History, New York,

New York.

DEH - Division of Environmental Health, Public Health Pest

Management Section, North Carolina Department of

Environment, Health, and Natural Resources, Raleigh.

FMNH - Field Museum of Natural History, Chicago, Illinois.

FSCA - Florida State Collection of Arthropods, Gainesville.

MEM - Mississippi Entomological Museum, Mississippi State

University, Starkville.

MMNS - Mississippi Museum of Natural Science, Jackson.

NCDA - Division of Plant Industry, North Carolina Department

of Agriculture, Raleigh.

NCSM - North Carolina State Museum of Natural Sciences, Raleigh.

NCSU - Entomology Department, North Carolina State University,

Raleigh.

NMNH - National Museum of Natural History, Smithsonian

Institution, Washington, DC.

NSC - Natural Science Center, Greensboro, North Carolina.

RNH - Private collection of R. N. Henson, Boone, North Carolina.

SEM - Snow Entomological Museum, University of Kansas, Lawrence.

48

Rowland M. Shelley

Figs. 2-3. Comparison of telsons, setation omitted. 2,

V. carolinianus.

vittatus:

Key to the Scorpions in North Carolina

Telson with variable subaculear tubercle (Fig. 2); pigmentation

yellow to yellow brown with obvious, dorsal, longitudinal

stripes

Telson without subaculear tubercle (Fig. 3); pigmentation

generally dusky brown, without stripes; southeastern states

from Kentucky and southwestern North Carolina to south-

eastern Louisiana

Vaejovis carolinianus (Beauvois)

Cephalothorax light, mottled brown, without black, inter-

ocular triangle, chelicerae with reticulated brown pigmen-

tation (Fig. 4); small species, adults ca. 32-44 mm in length

including metasoma and telson; Florida

Centruroides hentzi (Banks)

Cephalothorax darkly pigmented, with well defined, black

triangle pointing caudad and extending just beyond ocular

tubercle, chelicerae unpigmented (Fig. 5); large species,

adults ca. 40-60 mm in length; southcentral to south-

western United States and northern Mexico

Centruroides vittatus (Say)

Scorpions of North Carolina

49

Figs. 4-5. Color patterns of the chelicerae and cephalothorax.

4, C. hentzi; 5, C. vittatus.

50

Rowland M. Shelley

Family Buthidae

Centruroides vittatus (Say)

Habitat — In its native range, C. vittatus occurs in a wide variety

of microhabitats in deserts, deciduous and pine forests, and grasslands.

It lives in cracks and crevices of rocky outcrops and canyons walls,

climbs into vegetation, occurs beneath yuccas in deserts and grasslands,

and commonly enters houses (W. D. Sissom, West Texas A&M University,

personal communication). The specimens from Nash and Dare counties,

the Research Triangle Park, and Bland Road, Raleigh, were discovered

inside buildings; those at the last two sites were walking across a room

and a hallway. The specimen from Wakefield Street, Raleigh, was found

by workers digging behind a building.

Fig. 6. Occurrences of scorpions in North Carolina. Dots, V. carolinianus;

stars, C. hentzi; asterisks, C. vittatus. The site record from near Cowans

Ford Dam, Mecklenburg County, is indicated by the eastern circle; that

from Transylvania County is obscured by a dot. The western circle, in

Jackson County, denotes Balsam Gap, the location from which the wood

containing V. carolinianus in Haywood County was obtained. Dashed

lines surround presumably indigenous records in western counties show-

ing the assumed expansions from adjoining states.

Scorpions of North Carolina 51

Distribution — The generalized range of the southcentral and

southwestern United States to northern Mexico can be stated more

specifically as from Louisiana to New Mexico east of the Rio Grande

and, latitudinally, from the Central Plains of the United States into

the adjoining northern states of Mexico (W. D. Sissom, personal

communication). North Carolina specimens were examined as follows

(Fig. 6):

Wake Co., Raleigh, office building along Bland Rd., 1 spmn.,

April 1991, J. Wigmore (NCSM) and along Wakefield St., 1 spmn.,

24 October 1986, M. A. Brittain (NCSU); and Research Triangle Park,

1 spmn., August 1991, collector unknown (NCSM). Nash Co., Rocky

Mount, 1 spmn., 26 July 1991, collector unknown (NCSM). Dare Co.,

Nags Head, in building behind self storage facility, 1 spmn., 10 May

1986, L. Griffin (NCSU).

Remarks — Because winters in North Carolina's piedmont are

considerably milder than those in its native range in the midwest, C.

vittatus potentially could become established in the central part of

the State. An introduced population now exists in Murfreesboro, Rutherford

County, Tennessee (W. D. Sissom and G. A. Polis, Vanderbilt

University, personal communication).

There is at least one widespread chain of restaurants that broil

steaks over mesquite, so introduced, reproducing populations of

C. vittatus may exist in major cities throughout the Southeast. Use of

mesquite chips for broiling, rather than whole logs or lumber, would

produce the same flavor and eliminate the possibility of importing

live scorpions, not to mention unknown numbers of insects that might

have an adverse economic impact on agriculture.

Centruroides hentzi (Banks)

Habitat — In Florida, C. hentzi is usually encountered under litter,

logs, and stones; it can also be found under bark of dead trees up to

20 ft (6 m) high and commonly invades houses (Muma 1967). The

Durham County specimen was discovered in a rolled towel in a dormitory;

those from Carteret and Brunswick counties were found in residences

in condominium buildings, the 1992 scorpion was encountered in a

bathtub. At Emerald Isle, four specimens were found in hallways and

closets on both floors in a beachfront Carteret County condominium

complex during a two-month period in summer 1993, and two more

were encountered in May 1994. Six condominum units were involved,

comprising the northern half of one building; scorpions were not

discovered in the other buildings of the complex. One scorpion was

found dead in a man's shoe, and another stung a young child, who was

52 Rowland M. Shelley

treated at Carteret General Hospital and released. A pest control operator

treated inside and outside the building with pesticide but could not

determine the source of the scorpions. I visited the site in October

1994 and found no specimens and little shelter near the building. The

yard of the complex was immaculate, and the building is bordered

by low bushes surrounded with pine straw. Some bushes had

grown into relatively dense hedges, and the only external shelter of

consequence was beneath these hedges. I spent two hours searching

around the building, elsewhere on the complex, and in nearby wooded

areas in Emerald Isle and the western tip of Bogue Banks without

finding any scorpions.

Distribution — According to Muma (1967), C. hentzi occurs

throughout Florida, occurring in Columbia County, on the border

with Georgia, and in Escambia, the westernmost county. It should

therefore be expected in the southern coastal islands of Georgia, where

Say (1821) collected scorpions. North Carolina specimens were

examined as follows (Fig. 6):

Durham Co., Durham, Duke Univ., 1 spmn., 8 September 1987,

C. Brock (NCSU). Carteret Co., Bogue Banks, Emerald Isle, 1 spmn.,

Sept. 1993, D. McCluskey (NCSM). Brunswick Co., Bald Head Island,

1 mi (1.6 km) E of Marina, 1 spmn., July 1992, collector unknown

(RNH) and unknown site on island, 1 spmn., February 1993, collector

unknown (NCSM).

Remarks — An individual of C. hentzi was encountered in Raleigh

on 10 March 1938 "in strawberries from Florida"; one of C. gracilis

(Latreille) was discovered in Raleigh in the fall 1940 "in box shipped

from Florida"; and an undetermined Neotropical scorpion was found

in Raleigh on 13 December 1937 "in bunch of bananas from Central &

South America" (all specimens in NCDA). Although not encountered

in North Carolina environments, these specimens confirm that commercial

activities, like importing foods and fruits from other states and foreign

countries, is a key mechanism through which allochthonus organisms

are accidentally introduced into distant areas. The importation of

Florida palm trees for planting along the North Carolina coast can only

aid the spreading of C. hentzi and may result in its becoming established

in warm areas like Bald Head Island, where winters are not much cooler

than those in northern Florida where the scorpion is common.

Family Vaejovidae

Vaejovis carolinianus (Beauvois)

Habitat — I collected V. carolinianus in Cherokee County from

beneath large rocks on a dirt road and leaves in a deciduous forest

Scorpions of North Carolina 53

(Shelley 1975a), but I typically encounter the scorpion in association

with decaying pine logs and stumps, particularly under loose bark.

Rossman (1979) encountered specimens in clay soil on a stream bank,

and beneath decaying logs, leaf litter, slabs of wood, and the bark of

a dead hardwood tree. Gibbons et al. (1990) stated that V. carolinianus

was restricted to moist woodland habitats, where it occurs beneath

leaves, logs, and other litter. Several specimens from Transylvania

and Polk counties were found in and around houses; that from

Guilford County was taken within a house; and the one from Iredell

County was discovered in a sink in the basement of a house, but it

could have been imported from north Georgia, where the collector

spent the previous week. The specimen from Yancey County was

discovered in a tent at a campground.

Distribution — The southeastern United States from the Ohio River

in central Kentucky through eastern Tennessee, southwestern North

Carolina, and the Fall Zone of South Carolina and Georgia, to eastern

Mississippi and westcentral Tennessee, with a disjunct population in the

Tunica Hills of southwestern Mississippi and southeastern

Louisiana (Rossman 1979, Gibbons et al. 1990, Shelley 1994). In North

Carolina, V. carolinianus is native to Polk, Transylvania, and Cherokee

counties, spreading into these areas and up the Toxaway and Hiwassee

river valleys from adjacent parts of northern Georgia and western South

Carolina It also penetrates the western periphery by extending up the

Little Tennessee and French Broad river valleys from eastern Tennessee.

The scorpion also has been encountered in seven other counties, five

in the interior of the State and two on the border with piedmont and

coastal South Carolina, which probably represent accidental human

importations and examples of intra-state introductions. The Haywood

County site, in the heart of the Blue Ridge Province and at 5,000 ft.

(1,500 m) elevation, the highest reported altitude for the scorpion,

surely reflects an importation, as V. carolinianus was found in a

woodpile that was brought from Balsam Gap, Jackson County. The

scorpion was probably transported from the latter site (open circle in

Fig. 6); Haywood County is also in the heart of the Blue Ridge and an

unlikely spot for a native population. North Carolina specimens were

examined as follows:

Yancey Co., Crabtree Meadows, along Blue Ridge Pkwy., 1 spmn.,

8 June 1960, L. Mason (SEM). Madison Co., 1 mi (1.6 km) SE Walnut,

along US hwys. 25/70, 1 spmn., 25 August 1981, B. Hill (NCSM).

Haywood Co., Mt. Pisgah Cpgd., 1 spmn., August 1993, B. Randolph

(RNH). Swain Co., 0.5 mi (0.8 km) N Tapoco, along Little Tenn. R.,

2 spmns., 13 August 1985, R. Gaul, J. Whitcomb, D. Anthony, R. Lee

54 Rowland M. Shelley

(NCSM). Cherokee Co., locality unknown, 2 spmns., J. Gallatin (NMNH)

and 1 spmn., 19 June 1988, collector unknown (DEH); 6 mi (9.6 km)

WNW Culberson, along co. rd. 1107, 0.2 mi (0.3 km) N jet. co. rd.

1108, 1 spmn., 27 June 1974, R. M. Shelley (NCSM); 5 mi (8 km) W

Murphy, along US hwy. 64, 1 spmn., 1 October 1987, F. Bailey (RNH);

7.2 mi (11.5 km) NW Murphy, along co. rd. 1326, 0.3 mi (0.5 km) W

jet. co. rd. 1406, 2 spmns., 27 June 1974, R. M. Shelley (NCSM); and

1.7 mi (2.7 km) N Murphy, 1 spmn., 22 June 1984, A. L. & A. B.

Braswell (NCSM). Transylvania Co., 4.5 mi (7.2 km) SW Rosman,

along co. rd. 1139, 1 spmn., 3 December 1980, A. Burdo (NCSM);

Brevard, 1 spmn., 5 September 1975, D. Sizemore (NCSM) and in

house, 1 spmn., 18 November 1985, M. Albertson (NCSU); and along

Bearcamp Cr., 0.5 mi (0.8 km) N SC border, 1 spmn., 25 June 1962, R.

C. Graves (NCSM). Polk Co., nr. Tryon, 1 spmn., 16 September 1934,

collector unknown (NMNH); Tryon, around houses under construction,

2 spmns., 10 April 1957, D. F. Ashton (NCSM) and in leaf litter, 1

spmn., 21 November 1949, L. Eisenach (FMNH); in house 1.5 mi (2.4

km) WNW Columbus, along co. rd. 1135, 4 spmns., 21 September

1984, O. R. Ammons (NCSU); and Columbus, 3 spmns. 22 April 1957,

G. D. Jones (NCSM, NMNH) and 1 spmn., 1973, P. Culberson (NCSM).

Mecklenburg Co., Charlotte, Farmingdale Dr., 1 spmn., 8 August 1970,

M. Overton (NCSM). Iredell Co., 5 mi (8 km) SSW Troutman, SR

1401 at Lake Norman, 1 spmn., 25 June 1990, K. Troutman (NCSU).

Guilford Co., Greensboro, 1 spmn., November 1992, collector unknown

(NSC). Wake Co., Raleigh, Fairway Ridge Dr., 1 spmn., 25 August

1990, F. Starnes (NCSM). Columbus Co., Whiteville, 1 spmn., 9 July

1976, J. Rogers (NCSM).

The following site records are also considered valid but not

substantiated by specimens:

Transylvania Co., Toxaway Gorge, under bark of logs in 1963

(J. R. Paul).

Mecklenburg Co., Cowans Ford Dam at Lake Norman, under

mat in a building ca. 1987 (K. L. Manuel).

There are also two specimens taken in the "western part of

state," exact location unknown, 5 October 1965, J. Gallion (AMNH).

Remarks — Because of the abundance of predominantly pine

forests like those in which it occurs in piedmont South Carolina and

Georgia, V. carolinianus could become established in central North

Carolina through accidental importations from its natural range in

this or other states.

ACKNOWLEDGMENTS— I thank the following curators, collection

managers, and university faculty for records from their collections or

loans from, and access to, specimens under their care: B. R. Engber

Scorpions of North Carolina 55

(DEH), D. Summers (FMNH), G. B. Edwards (FSCA), T. L. Schiefer

(MEM), R. L. Jones (MMNS), K. R. Ahlstrom (NCDA), R. L. Blinn

(NCSU), J. A. Coddington (NMNH), and R. W. Brooks (SEM). The sight

records are courtesy of K. L. Manuel, Duke Power Company, and

J. R. Paul. The following people assisted by bringing scorpions to my

attention: D. L. Stephan, D. McCluskey, J. Weems, and E. Kunickis.

D. McCluskey explained the situation with C. hentzi at the Carteret

County condominiums; D. S. Lee reviewed a preliminary draft of the

manuscript; and R. N. Henson provided records of C. hentzi and

V. carolinianus from his private collection (RNH) and the NSC. W. D.

Sissom, West Texas A&M University, Canyon, Texas, and NCSM

arachnid research associate, provided the AMNH record of V. carolinianus,

advised on scorpions, and commented on a preliminary draft of

the paper. R. G. Kuhler, NCSM scientific illustrator, prepared Figures

4-5; Figure 1 is courtesy of D. J. Lyons, NCSM exhibits section.

LITERATURE CITED

Francke, O. F. 1974. Two emendations to Stahnke's (1974) Vaejovidae

revision (Scorpionida: Vaejovidae). Journal of Arachnology 4:125-

135.

Gibbons, W., R. R. Haynes, and J. L. Thomas. 1990. Poisonous

Plants and Venomous Animals of Alabama and the Adjoining States.

University of Alabama Press, Tuscaloosa.

Muma, M. H. 1967. Scorpions, whip scorpions and wind scorpions of

Florida. Arthropods of Florida and Neighboring Land Areas 4:1-

28.

Rossman, D. A. 1979. Distribution of the southern unstriped scorpion,

Vaejovis carolinianus (Beauvois). Proceedings of the Louisiana Academy

of Sciences 42:10-12.

Say, T. 1821. An account of the Arachnides of the United States.

Journal of the Philadelphia Academy of Sciences 2:65-68.

Shelley, R. M. 1975a. North Carolina's scorpion. Wildlife in North

Carolina 39:8-9.

Shelley, R. M. 1915b. Occurrence of the scorpion, Vejovis carolinianus

(Beauvois), in North Carolina (Arachnida: Scorpionida: Vejovidae).

Journal of the Elisha Mitchell Scientific Society 91:29-30.

Shelley, R. M. 1994. Distribution of the scorpion, Vaejovis carolinianus

(Beauvois) — a reevaluation. Brimleyana 21:57-68.

Stahnke, H. L., and M. Calos. 1977. A key to the species of the

genus Centruroides Marx (Scorpionida: Buthidae). Entomological News

88:111-120.

Received 8 March 1994

Accepted 27 September 1994

56

Distribution of the Scorpion,

Vaejovis carolinianus (Beauvois) — a Reevaluation,

(Arachnida: Scorpionida: Vaejovidae)

Rowland M. Shelley

North Carolina State Museum of Natural Sciences, P. O. Box 29555,

Raleigh, North Carolina 27626-0555

ABSTRACT — Vaejovis carolinianus (Beauvois) is primarily an

upland scorpion ranging from the Ohio River in central Kentucky

to the inner Coastal Plain of Alabama and, east/west, from

the Fall Zone of South Carolina and Georgia to eastern Mississippi

and westcentral Tennessee. A disjunct population inhabits the

Tunica Hills, along the eastern side of the Mississippi River

in southwestern Mississippi and adjacent Louisiana. It is abundant

in the Cumberland Plateau of Kentucky, Tennessee, and Alabama,

but occurs only in the western tip of Virginia. The distribution

skirts the western and southern peripheries of the Blue Ridge

Province, including only certain border counties of North Carolina,

primarily those adjoining Georgia and western South Carolina.

The northward extension west of the Appalachians is much

greater than that to the east, and sporadic records from the

interior of North Carolina and southern border counties east of

the Appalachians apparently constitute accidental human importations.

Specific localities are detailed and plotted on a distribution

map.

The southern unstriped scorpion, Vaejovis carolinianus (Beauvois)

(Fig. 1), characterized by dusky brown pigmentation without stripes

and the absence of a subaculear tubercle on the telson, is the only

indigenous scorpion in the southeastern United States known to occur

north of Florida (Muma 1967, Shelley 1994). Rossman (1979) present-

ed a distribution map and summarized earlier records; Gibbons et al.

(1990) mapped the overall distribution and that in Alabama, and reported

that the scorpion occurs in scattered localities from central Kentucky,

eastern Tennessee, and western parts of Virginia and the Carolinas

through Georgia and Alabama, chiefly above the Coastal Plain. Gibbons

et al. mentioned that isolated records were available for eastern and

southwestern Mississippi, and the adjacent part of Louisiana, and that

the scorpion ranges south of the Fall Zone in Alabama as far as Dallas

County. However, the only specific locality reported in the latter areas

is Tunica Hills, along the eastern side of the Mississippi River in West

Feliciana Parish, Louisiana, and Wilkinson County, Mississippi, and

Brimleyana 21:57-68, December 1994 57

58 Rowland M. Shelley

Fig. 1. Vaejovis carolinianus, dorsal view.

few definite sites of occurrence have ever been recorded. The only

other published records known to me are the following by Brimley

(1938) and/or Shelley (1975a):

KENTUCKY: Laurel Co., 1 mi (1.6 km) W Baldrock. Bell Co.,

Pineville.

VIRGINIA: Lee Co., Cumberland Gap.

TENNESSEE: Sevier Co., Pigeon Forge and Laurel Creek, Great

Smoky Mountains National Park.

NORTH CAROLINA: Cherokee Co., near Culberson and Murphy.

Polk Co., Tryon and Columbus.

SOUTH CAROLINA: Pickens Co., Clemson. Laurens Co., Laurens.

Lexington Co., Lexington.

GEORGIA: Habersham Co., Tallulah Falls (incorrectly placed

in Rabun County). Fulton Co., Atlanta. Clarke Co., Athens.

For the past 15 years, I have examined unreported museum samples

of V. carolinianus and collected the scorpion in southeastern states;

from these records a clear picture of the species' range has emerged

Vaejovis carolinianus Distribution

59

(Fig. 2). The purpose of this contribution is to document the distribution

by providing specific localities and collection data, and to publish a

dot map that eliminates ambiguities in previous maps, in which counties

of occurrence are shaded. This procedure is misleading for peripheral

areas like Lee County, Virginia, where V. carolinianus occurs only in

Cumberland Gap, in the westernmost two miles of the County and

State. Concerted efforts to find V. carolinianus eastward in Lee County

have been unsuccessful (R. L. Hoffman, Virginia Museum of Natural

History, personal communication), so shading the entire county incorrectly

implies wider occurrence.

The overall distribution, as documented herein, extends southward

through the Cumberland Mountains from the Ohio River, the northern

boundary in central Kentucky, and spreads eastward to the Appalachians

Fig. 2. Distribution of V. carolinianus.

60 Rowland M. Shelley

in southeastern Tennessee. The border then swings southward around

the bulk of the Blue Ridge Province, penetrating border counties of

southwestern North Carolina by spreading up river valleys from adjacent

states. The easternmost natural occurrence in North Carolina is in Polk

County; other records in this State are believed to represent accidental

human importations (Shelley 1994). The distributional boundary then

angles southeastward through the Piedmont Plateau of South Carolina

to Columbia and follows the Fall Zone through Georgia into Alabama,

before spreading onto the Gulf Coastal Plain in western Alabama and

extending to the Mississippi River above Baton Rouge, Louisiana. Fewer

specimens are available from the western side of its distribution, but V.

carolinianus ranges northward through eastern Mississippi and western

Tennessee to the Ohio River in central Kentucky; the only specimens

taken farther west are those from Tunica Hills. Although these records

still are somewhat scattered, they are continuous enough to suggest

regular occurrence throughout the overall range except for the Tunica

Hills population, which is disjunct. Vaejovis carolinianus is therefore

an upland species occurring exclusively west of the Fall Zone in the

Carolinas and Georgia. Where it extends onto the Coastal Plain in

Alabama, it shows a marked preference for hills or ridges like Tunica

Hills and the Lime and Buhrstone Hills, and the Chunnenuggee Ridge,

in Clarke County, Alabama. The scorpion has not been taken in the

pine flatwoods that are abundant in the Coastal Plain of the Carolinas

and Georgia. Muma (1967) considered V. carolinianus to be a potential

inhabitant of the northern and panhandle counties of Florida, but the

known distribution shows that this possibility is remote, even for the

most proximate part of the State, i. e., the inner peripheries of Escambia

and Santa Rosa counties in the western panhandle.

The greater northward extension west of the Blue Ridge Province,

in contrast to that on the east, is particularly striking. East of the

Appalachians, the range angles southeastward from Polk County, North

Carolina, to Lexington County, South Carolina; to the west, however,

it extends some 230 mi (368 km) farther north, to the Ohio River in

central Kentucky. If the distribution east of the Appalachians were

equivalent to that on the west, it would reach to around Charlottesville,

Virginia! Vaejovis carolinianus is common in the Cumberland Mountains

of Kentucky and Tennessee, as it also is in the western Blue Ridge

Province in southeastern Tennessee. However, the only records from

the western periphery of North Carolina are from Madison and Swain

counties (Shelley 1994), which doubtlessly represent penetrations up

the French Broad and Little Tennessee river valleys, respectively. The

former also implies occurrence in adjacent Cocke County, Tennessee,

Vaejovis carolinianus Distribution 61

where the species has not been taken. The range swings northwestward

from Sevier, and presumably Cocke, counties into the Cumberland Plateau

and does not extend farther north in the Blue Ridge and Ridge and

Valley Provinces.

Vaejovis carolinianus typically occurs in association with decaying

pine logs, particularly beneath loose bark; it can be plentiful under

loose rocks on talus slopes and has been encountered in mesic deciduous

forests (Shelley 1975/?, 1994). In central Tennessee, specimens have

been taken at night with ultraviolet light from moss-covered vertical

surfaces of old highway roadcuts in limestone, suggesting that the scorpions

inhabit the cracks and crevices on these walls and forage either in the

cracks or on the surface itself (W. D.Sissom, West Texas A&M University,

personal communication). Rossman (1979) encountered specimens in

clay soil on a stream bank, and beneath decaying logs, leaf litter, slabs

of wood, and the bark of a dead hardwood tree. Gibbons et al. (1990)

stated that V. carolinianus was restricted to moist woodland habitats,

where it occurs beneath leaves, logs, and other litter. The scorpion

occasionally wanders into buildings and has been collected in homes in

Davidson County, Tennessee; Madison, Polk, and Transylvania counties,

North Carolina; Laurens, Spartanburg, Lexington, and Oconee counties,

South Carolina; and Tishomingo and Clarke counties, Mississippi. It

was also taken in a dormitory room at Mississippi State University,

Oktibbeha County, Mississippi.

The following locality records are supported by preserved

specimens and are presented in alphabetical order; those from North

Carolina are provided by Shelley (1994). Citations include the date of

collection, the name(s) of the collector(s), and the repository, indicated

by the following acronyms:

AMNH - American Museum of Natural History, New York,

New York.

ANSP - Academy of Natural Sciences, Philadelphia, Pennsylvania.

CAS - California Academy of Sciences, San Francisco.

CC - Biology Department, Columbus College, Columbus, Georgia.

CU - Entomology Department, Clemson University, Clemson

South Carolina.

FMNH - Field Museum of Natural History, Chicago, Illinois.

FSCA - Florida State Collection of Arthropods, Gainesville.

ILNHS - Illinois Natural History Survey, Champaign.

MCZ - Museum of Comparative Zoology, Harvard University,

Cambridge, Massachusetts.

MEM - Mississippi Entomological Museum, Mississippi State University,

Starkville.

62 Rowland M. Shelley

MMNS - Mississippi Museum of Natural Science, Jackson.

MPM - Milwaukee Public Museum, Milwaukee, Wisconsin.

NCDA - Pest Control Division, North Carolina Department of

Agriculture, Raleigh.

NCSM - North Carolina State Museum of Natural Sciences, Raleigh.

NCSU - Entomology Department, North Carolina State University,

Raleigh.

NMNH - National Museum of Natural History, Smithsonian

Institution, Washington, DC.

NSC - Natural Science Center, Greensboro, North Carolina.

RNH - Private collection of R. N. Henson, Boone, North Carolina.

SEM - Snow Entomological Museum, University of Kansas, Lawrence.

TAM - Entomology Department, Texas A&M University, College

Station, Texas.

UAAM - Univeristy of Arkansas Arthropod Museum, Fayetteville.

UGA - University of Georgia Museum of Natural History, Athens.

UL - Biology Department, University of Louisville, Louisville,

Kentucky.

UMMZ - University of Michigan Museum of Zoology, Ann Arbor.

VMNH - Virginia Museum of Natural History, Martinsville.

WDS - Private collection of W. D. Sissom, West Texas A&M

University, Canyon, Texas.

KENTUCKY: Bell Co., Pineville, 14 August 1952, collector unknown

(CAS); Pine Mountain St. Pk., 7 August 1949, H. S. Dybas (FMNH), 2

May 1975, C. C. Cornett (UL), and June 1975 and 1976, J. K. Ettman

(NCSM); Fern Lake nr. Middlesboro, 12 August 1959, T. Wolfe, W. L.

Burget (AMNH, VMNH); Cumberland Gap, date unknown, W. Faxon

(MCZ); and Balkan, 1927, C. F. Clayton (NMNH). Bullitt Co., Fort

Knox, date unknown, P. Burchfield (AMNH). Bullitt/Nelson Cos, State

Forest, 17 July 1947, H. Nadler (NMNH). Jefferson Co., Ft. Knox, 7

September 1993, G. Kettring (UL). Laurel Co., 1 mi (1.6 km) NW

Baldrock, 26 May 1952, L. Hubricht (NMNH). Lincoln Co., locality

unknown, 22 May 1966, B. Carter (UL). McCreary Co., Yahoo Falls

and Cumberland Falls, 24 May 1972, collector unknown (UL). Meade

Co., Quarry off KY hwy. 1638, 12 July 1967, C. Karnella (UL). Pulaski

Co., Mt. Victory, 27 May 1972, collector unknown (UL). Rockcastle

Co., 3 mi (4.8 km) N Mt. Vernon, Renfro Valley, 10 June 1954, R.

Barbour (NCSM). Taylor Co., Campbellsville, 7 May 1956, collector

unknown (FSCA). Wayne Co., locality and collector unknown, 21 September

1994 (UL). Whitley Co., Cumberland Falls St. Pk., nr. Cumberland

Falls, 27 June 1945 collector unknown (CAS); and 0.5 mi (0.8 km) E

Cumberland Falls St. Pk., 10 October 1992, R. W. VanDevender (RNH).

Vaejovis carolinianus Distribution 63

VIRGINIA: Lee Co., 0.5 mi (0.8 km) N Cumberland Gap, Willis

Hollow, 15 July 1958, C. Stoddard, W. L. Burget (AMNH, VMNH);

and Cumberland Gap, above Cudjo's Cave, 23 June 1950, J. H. Fowler

(CAS).

TENNESSEE: Anderson Co., Oak Ridge, August 1962, H. Phillips

(NMNH). Blount Co., Great Smoky Mts. Nat. Pk., The Sinks, 19 June

1960, I. McClurkin (FSCA); Chilowhee Mts., 21 July 1965, C. L. Wilder

(VMNH); and Townsend, 19 June 1960, I. McClurkin (FSCA). Cheatham

Co., Ashland City, summer 1994, R. Stevens (NCSM). Claiborne Co.,

Cumberland Gap, 11 June 1933, H. W. Chickering (MCZ). Cumberland

Co., 11 mi (17.6 km) NNW Crossville, US hwy. 127 at Lickfork Cr., 9

June 1979, R. M. Shelley, R. K. Tardell (NCSM). Davidson Co., Goodlettsville,

19 December 1949, Ross, Stannard (ILNHS); ca. 1.5 mi (2.4 km) SW

Marrowbone L., 28 April 1991, D. McGinnity (WDS); and Nashville,

23 May 1985, K. Collier (WDS). DeKalb Co., Edgar Evins, St. Pk., 20

May 1983, R. W. VanDevender (RNH); and 6 mi (9.6 km) NE Smithville

off TN hwy. 56 on S side Center Hill L., 20 July 1984, W. D. Sissom,

C. N. McReynolds (WDS). Hamilton Co., Chattanooga, October 1932,

L. Trenholm (NMNH). Hardin Co., Savannah, 1960, collector unknown

(AMNH); Counce, YMCA camp nr. Pickwick Dam, 5 August 1966,

A. Smoot (AMNH); and Pickwick Dam, 29 September 1963, M. Lou

(CAS). Madison Co., Jackson, 1958, A. F. Archer (AMNH). Marion

Co., US hwy. 64, 2.9 mi (4.6 km) E jet. TN hwy. 156, 26 March 1979,

R. Franz, R. E. Ashton, A. L. Braswell (NCSM); 6 mi (9.6 km) S

Jasper, along TN hwy. 156A, 0.2 mi (0.3 km) N AL line, 21 May

1983, R. M. Shelley, J. Staton (NCSM). Morgan Co., locality unknown,

28 August 1976, J. C. Mitchell, J. Byrd, J. Kline (VMNH); and Catoosa

WRA, between Lookout & Phil, 8 May 1976, R. L. Jones (MCZ).

Pickett Co., Pickett St. Pk., 1 May 1935, H. Augusby (NMNH). Polk

Co., exact site unknown, April 1985, R. W. VanDevender (RNH); and

Parksville Reserve, 7 August 1936, M. H. Hatch (AMNH). Putnam

Co., 2 mi (3.2 km) E Cookville, along 1-40, date and collector unknown

(FMNH). Rhea Co., 3 mi (4.8 km) N Spring City, 26 June 1962, F. N.

Young (FSCA); and Spring City, August 1919, E. R. Dunn (MCZ).

Roane Co., nr. Harriman, 11 July 1933, W. Ivie (AMNH); and Kingston,

12 and 14 July 1933, W. J. Gertsch, W Ivie (AMNH) and 15 April

1952, G. E. Smith (AMNH). Rutherford Co., ca. 7.4 mi (11.8 km) E

Murfreesboro, along US hwy. 70S at Cripple Cr., 18 April 1973, Czajka,

Ketchum (MPM). Sevier Co., Pigeon Forge, 29 August 1943, J. A. Bell

(CAS). Van Buren Co., Fall Creek Falls St. Pk., 11 August 1951, T.

Cohn (AMNH). Warren Co., nr. McMinnville, along TN hwy. 8, 15

August 1971, M. A. Morris (ILNHS). White Co., Rock Island, 18 April

64 Rowland M. Shelley

1917, collector unknown (NMNH). Wilson Co., Cedars of Lebanon

St. Pk., 16 April 1976, R. L. Jones, A. C. Ecternauer (MCZ).

NORTH CAROLINA: Cherokee, Columbus, Guilford, Haywood,

Iredell, Madison, Mecklenburg, Polk, Swain, Transylvania, Wake, and

Yancey counties, as per the listing in Shelley (1994) (NCSM, NCSU,

NMNH, NSC, RNH, SEM).

SOUTH CAROLINA: Abbeville Co., Sumter Natl. For., FS rd.

505 at Long Cane Scenic Area, 14 September 1980, R. M. Shelley, M.

Morgan (NCSM). Anderson Co., Pendleton, Aldwood, 18 May 1985, J.

H. Morse (CU); and along US hwy. 29, exact site unknown, July 1989,

A. Hill (RNH). Edgefield Co., 11.4 mi (18.2 km) W Edgefield, SC

hwy. 68, 0.2 mi (0.3 km) N jet. US hwy. 52, 8 August 1976, R. M.

Shelley (NCSM). Greenville Co., nr. NC border, exact location unknown,

29 August 1929, collector unknown (NMNH); and Greenville, date

and collector unknown (ANSP) and Camp Sevier, WWI Army Post

probably now in Greenville, date unknown, J. Leidy (ANSP). Laurens

Co., Kinards, 15 May 1975, G. F. Smith (NCSU); and Laurens, 1

November 1955, collector unknown (CAS). Lexington Co., nr. Chapin

on L. Murray, 19 February 1991, F. Authinreith (RNH); Leesville vie,

26 September - 7 October 1949, L. Brodie (FMNH); and Lexington,

date and collector unknown (CAS). McCormick Co., locality unknown,

1935, collector unknown (NMNH) and 15 June 1958, collector unknown

(MCZ); and De La Howe Forest, 10 July 1942, C. J. Goodnight (ILNHS).

Newberry Co., Newberry, July 1932, L. A. Savage (NMNH). Oconee

Co., Mountain Rest, 1973, R. F. Shriner (NMNH); Oconee St. Pk., 10

June 1991, E. G. Riley (TAM); South Cove Landing, date unknown, A.

Dozier (FSCA); nr. Clemson, 22 September 1985, J. R. Mayer (NCSU);

and 6.8 mi (10.9 km) S Westminster, along SC hwy. 67 at Choestoea

Cr., Hartwell Res., 10 June 1978, R. M. Shelley, W. B. Jones (NCSM).

Pickens Co., 1 mi (11.2 km) NE Pickens, 16 October 1976, collector

uknown (FSCA); and Clemson, 3 December 1926 and April 1928, C.

S. Brimley (NCDA), 25 September 1928, R. C. Fox (CU), February

1930, F. Smith (UAAM), 16 March 1940, E. C. Van Dyke (CAS), 22

April 1968, H. Harris (CU), and 8 September 1976, S. Prichard (CU).

Saluda Co., Saluda, 15 June 1958, N. B. Causey (MCZ); and 8 mi

(12.8 km) SW Leesville, along Brodie Rd., September 1992, B. Cockerel

(RNH). Spartanburg Co., Landrum, 4 August 1910, R. V. Chamberlin

(AMNH); and Glenn Springs, 17 October 1984, H. B. Matthews (NCSU).

GEORGIA: Baldwin Co., Milledgeville, 2 August 1894, collector

unknown (NMNH). Bartow Co., E of Cartersville, along GA hwy. 21,

1.8 mi (2.9 km) E jet. hwy. 1-75, 17 March 1984, G. T. Baker (MEM).

Butts Co., Indian Springs, date unknown, L. M. Underwood (NMNH).

Vaejovis carolinianus Distribution 65

Clarke Co., Athens, 1 October 1936, H. 0. Lund (AMNH), 11 October

1960-30 October 1972, N. Bein, W. Eissler, R. Muha, K. Douce (UGA),

and 27 November 1983, M. LaSalle (MEM); and Whitehall For., 9 July

1977-21 May 1984, C. L. Smith (UGA). Cobb Co., Austell, date unknown,

N. Banks (MCZ); and Kennesaw Mtn., 26 April 1963, P. W. Fattig

(NMNH). Columbia Co., Harlem, 25 July 1962, G. Jordan (FSCA).

DeKalb Co., Atlanta, September 1982, collector unknown (RNH); Stone

Mtn., 15 June 1958, N. B. Causey (MCZ) and 25 August 1966, Trobinck

(FSCA); and Dunwoody, Summerbrook Dr., September 1991, A. L.

Henson (RNH). Elbert Co., Bobby Brown St. Pk., 31 July 1977, R. M.

Shelley (NCSM); and 11 mi (17.6 km) ENE Elberton, along GA hwy.

368 at Pickens Cr., 20 July 1979, R. M. Shelley, R. K. Tardell (NCSM).

Floyd Co., Rome, Marshall For., 17 March and 25 April 1963, J. Parker

(NMNH). Fulton Co., Roswell, date unknown, Mrs. King (MCZ); Atlanta,

27 June 1943, H. Hoogstraal (FMNH), March 1945 and 21 September

1951, collector unknown (CAS), August 1951, G. W. Fraser (NMNH),

and 1957, H. D. Pratt (ILNHS); Chamblee, 15 September 1952, collector

unknown (ILNHS). Gordon Co., Horn Mtn., 12 June 1986, J. D. Lazell

(NMNH). Habersham Co., Tallulah, 1-4 July 1955, R. B. & R. L.

Hoffman (NMNH); Tallulah Falls, 1 April 1891 and April 1897, L. M.

Underwood (NMNH); N of Clarkesville, 27 April 1943, W. Ivie (AMNH);

and between Clarkesville and Toccoa, 28 April 1943, W. Ivie (AMNH).

Harris Co., exact sites unknown, June 1982-August 1992, J. A. Layne,

P. Jones, H. M. Johnson, B. A. Craighton, J. A. Riley, J. Chappell, and

R. D. Schiavone (CC). Jackson Co., locality not specified, 29 September

1960, Moody (UGA). Lumpkin Co., Frogtown Gap, 31 May 1934, F.

Harper (AMNH). McDuffie Co., between Thomson and Washington,

22 April 1943, W. Ivie (AMNH). Meriwether Co., Warm Springs, 4

May 1935, collector unknown (NMNH). Monroe Co., Forsyth, 1962,

A. F. Archer (AMNH). Murray Co., Fort Mtn. St. Pk., 14 July 1952, P.

M. Choate (FCSA). Muscogee Co., Columbus, 25 April 1971, W. M.

Petrasek (CC) and 5 April 1982, J. Losonsky (CC). Newton Co., Covington,

17 November 1984, J. Wheeler (NMNH). Putnam Co., 8 mi (12.8 km)

S Eatonton, Oconee Nat. For., 22 May 1981, J. M. Carpenter (MCZ).

Rabun Co., Rabun Jet., April and August 1887, collector unknown

(NMNH); and Clayton, 6-12 April 1940, E. C. Van Dyke (CAS) and 5-

9 September 1959, K. Ulman (AMNH). Rockdale Co., along Big Haynes

Cr., 16 September 1976, T. Schowalter (UGA). Stephens Co., Toccoa

Lake, 10 October 1928, collector unknown (NMNH); and Toccoa, June

1953, R. L. Hoffman (NMNH). Taliaferro Co., Crawfordville, 30 July

1971, M. & M. Cazier (WDS). White Co., Unicoi St. Pk., 8 July 1973

and 25 May 1975, H. O. Lund (UGA); and Sautee, 13 September 1959,

66 Rowland M. Shelley

H. O. Lund (UGA). County Unknown, Gowerville, date unknown, S.

M. King (MCZ).

ALABAMA: Bibb Co., nr. Wilton, 28 July 1962, R. E. Crabill,

A. B. Gurney (NMNH). Chambers Co., Fairfax, 1949-1950, A. F. Archer

(AMNH). Chilton Co., Maplesville, date and collector unknown (FSCA);

and Verbena, 22 July 1900, W. R. Maxon (NMNH). Clarke Co., Thomasville,

6 August 1933, C. E. Burt (NMNH); and Camp Maubila, a Boy Scout

camp. ca. 6 mi (9.6 km) S Grove Hill, 4 mi (6.4 km) E US hwy. 43, 23

June 1983, P. Cross (MEM). Clay Co., Cheaha Mtn., 3 September

1969, J. G. E. Rehn (ANSP). Cleburne Co., locality unknown, 8-9

September 1946, collector unknown (FSCA); and Talladega Natl. For.,

28 July 1962, R. C. & A. Graves (FSCA). Coosa Co., Hatchet Cr., 4

June 1940, A. F. Archer (AMNH). Cullman Co., Cullman, St. Boniface

Col., 17 June 1958, collector unknown (FSCA). DeKalb Co., Desoto

St. Pk., 10 May 1986, R. F. C. Naczi (UMMZ) and 18-19 May 1990,

D. Hildebrandt, T. L. Schiefer (MEM); and Collinsville, 10 July 1962,

H. B. Cunningham (ILNHS). Franklin Co., Ezell Cave, along AL hwy.

1060, 8 November 1969, F. Shires (NCSM). Jackson Co., 12 mi (19.2

km) N Scottsboro, National Bridge Cave, August 1970, R. C. Graham

(NCSM). Lauderdale Co., Wilson Dam, 13 July 1942, J. W. Belkin

(AMNH) and June 1953, R. V. Schick (AMNH). Lee Co., Auburn,

November 1895 and July 1896, L. M. Underwood (NMNH). Madison

Co., Huntsville, Monte Sano St. Pk., date unknown, Rosenburg (FSCA);

E of Huntsville, SW slope of Round Top Mtn., 19 July 1988, K. Woodstock

(NMNH); and Monte Sano St. Pk., 1946, J. Murphy (AMNH). Morgan

Co., locality and collector unknown, 27 July 1910 (AMNH). Russell

Co., exact site unknown, 29 July 1990, T. Mann (CC). Tallapoosa Co.,

Dadeville, 13 July 1914 (NMNH); and Alexander City, 31 October

1944, G. Nelson (MCZ). Tuscaloosa Co., Lake Lurleen St. Pk., 18

May 1988, C. M. & O. S. Flint (NMNH). Walker Co., Forks of Warrior

R., 20 October 1912, H. Smith (AMNH). Winston Co., Double Springs,

18 June 1958, N. B. Causey (MCZ).

MISSISSIPPI: Clarke Co., 0.8 mi (1.3 km) N Hurricane Cr., 3

December 1961, L. Hubricht (NMNH); and at AL line, 5.0 mi (8.0 km)

SE Lauderdale Co. line, 17 April 1987, R. L. Jones, J. Wiseman (MMNS).

Oktibbeha Co., Starkville, Mississippi St. Univ., 3 October 1983, collector

unknown (MEM). Tishomingo Co., Iuka, 31 October 1988, B. Hopper

(MEM); and Tishomingo St. Pk., 28 July 1983, P. R. Miller (MEM).

No specimens were available from the following additional, generalized

county records of Benton (1973), Rossman (1979), and Gibbons et al.

(1990). Dots are therefore placed centrally in these counties in Figure

2:

Vaejovis carolinianus Distribution 67

KENTUCKY: Jackson, Larue, Lee, and Leslie counties.

TENNESSEE: Claiborne, Grundy, and Monroe counties.

GEORGIA: Cherokee, Crawford, Dade, and Polk counties.

ALABAMA: Calhoun, Colbert, Dallas, Elmore, Fayette, Hale,

Lawrence, Marion, Randolph, and Shelby counties.

MISSISSIPPI: Lauderdale County.

The following additional localities were communicated by H. L.

Stahnke (in lift.) based on specimens in his personal collection, now

housed at the CAS. These samples were not represented in the material

loaned from that institution, but they are considered accurate and are

hence incorporated into Figure 2.

KENTUCKY: Meade Co., Muldraugh.

TENNESSEE: Blount Co., Great Smoky Mountains Nat. Pk., 1

mi (1.6 km) S The Sinks, and along Little Tennessee R. at Long Arm

Bridge. Sevier Co., Gatlinburg.

ALABAMA: Jefferson Co., Adger. Tuscaloosa Co., Peterson.

MISSISSIPPI: County Unknown, Pine Flat.

The following localities, communicated verbally by J. L. Knight

(South Carolina State Museum, Columbia), are also incorporated into

Figure 2.

SOUTH CAROLINA: Aiken Co., North Augusta, S of 1-20 along

N side of Savannah River. Lexington Co., West Columbia. McCormick

Co., L. Thurmond.

ACKNOWLEDGMENTS— I thank W. D. Sissom, West Texas A&M

University and NCSM arachnid research associate, for providing records

of V. carolinianus from his files, including those in the AMNH and

TAM holdings as well as his personal collection, and for reviewing a

preliminary draft of the manuscript. The following curators, collection

managers, and university faculty graciously provided access to, or loans

from, holdings under their care: D. Azuma (ANSP), C. E. Griswold

and V. F. Lee (CAS), G. E. Stanton (CC), J. H. Morse (CU), D. Summers

(FMNH), G. B. Edwards (FSCA), K. C. McGiffen and K. R. Methven

(ILNHS), H. W. Levi (MCZ), T. L. Schiefer (MEM), R. L. Jones (MMNS),

J. P. Jass (MPM), K. R. Ahlstrom (NCDA), R. L. Blinn (NCSU), J. A.

Coddington (NMNH), R. W. Brooks (SEM), J. B. Whitfield (UAAM),

C. L. Smith (UGA), C. V. Covell, Jr. (UL), M. F. O. Brien (UMMZ),

and R. L. Hoffman (VMNH). M. Ward, University of Alabama, provided

a copy of Benton's thesis (1973) and checked on Alabama localities; J.

Hall, Alabama Museum of Natural History, likewise provided valuable

advice on Alabama localities and physiographic features. R. N. Henson

kindly provided records from his personal collection (RNH) and the

68 Rowland M. Shelley

NSC, and J. Knight did likewise for specimens at the SCSM. D. S.

Lee also reviewed a preliminary draft of the manuscript. Figure 1 is

courtesy of D. J. Lyons, NCSM exhibits designer.

LITERATURE CITED

Benton, C. L. B., Jr. 1973. Studies on the biology and ecology of

the scorpion Vejovis carolinianus (Beauvois). Ph.D. Thesis, Uni-

versity of Alabama, Tuscaloosa.

Brimley, C. S. 1938. Insects of North Carolina. North Carolina

Department of Agriculture, Division of Entomology, Raleigh.

Gibbons, W., R. R. Haynes, and J. L. Thomas. 1990. Poisonous

plants and venomous animals of Alabama and the adjoining States.

University of Alabama Press, Tuscaloosa.

Muma, M. H. 1967. Scorpions, whip scorpions and wind scorpions

of Florida. Arthropods of Florida and Neighboring Land Areas

4:1-28.

Rossman, D. A. 1979. Distribution of the southern unstriped scor-

pion, Vaejovis carolinianus (Beauvois). Proceedings of the Louisi-

ana Academy of Sciences 42:10-12.

Shelley, R. M. 1975a. Occurrence of the scorpion, Vejovis carolinianus

(Beauvois) in North Carolina (Arachnida: Scorpionida: Vejovidae).

Journal of the Elisha Mitchell Scientific Society 91:29-30.

Shelley, R. M. 1975£>. North Carolina's scorpion. Wildlife in North

Carolina 39:8-9.

Shelley, R. M. 1994. Introductions of the scorpions Centruroides

vittatus (Say) and C. hentzi (Banks) into North Carolina, with

records of Vaejovis carolinianus (Beauvois) (Scorpionida: Buthidae,

Vaejovidae). Brimleyana 21:45-55.

Received 8 March 1994

Accepted 27 September 1994

Atlantic Ocean Occurrences of the Sea Lamprey,

Petromyzon marinus (Petromyzontiformes:

Petromyzontidae), Parasitizing Sandbar, Carcharhinus

plumbeus, and Dusky, C. obscurus (Carcharhiniformes:

Carcharhinidae), Sharks off North and South Carolina

Christopher Jensen and Frank J. Schwartz

Institute of Marine Sciences

University of North Carolina

Morehead City, North Carolina 28557

ABSTRACT — Sandbar and dusky sharks captured in 1993 in western

Atlantic Ocean waters off North and South Carolina were parasitized

by sea lampreys. All lampreys were females ranging from 165

to 343 mm total length. Removal of an attached lamprey revealed

round, reddish and/or bleeding areas on a shark's body. Blood

oozing from a lamprey's cloaca indicated that feeding was occurring

or had occurred.

The anadromous parasitic sea lamprey (Petromyzon marinus) is

widely distributed on both sides of the Atlantic Ocean. It occurs off

North America from Labrador southward to Florida, and along eastern

Europe from Varanger Fjord in Norway to the western Mediterranean

(Beamish 1980). Apparently it also formerly occurred in the Gulf of

Mexico (Vladykov and Kott 1980, Gilbert and Snelson 1992). Lampreys

are known from marine waters to depths of 4,099 m (Haedrich 1977).

Dempson and Porter (1993) note other western Atlantic captures of sea

lampreys in deep open ocean waters. Excellent reviews of sea lamprey

biology can be found in Hardesty and Potter (1971) and in the Proceedings

of the Sea Lamprey International Symposium (1980). We add the sea

lamprey as an external parasite of sharks and present meristic and morphometric

data for specimens captured off North and South Carolina.

Sea lampreys prey on a variety of fishes in freshwater and marine

habitats (Bigelow and Schroeder 1948). Sea lampreys have not been

reported from ocean habitats off North Carolina (personal observation)

or South Carolina (S. Van Sant, South Carolina Marine Resources Center,

personal communication), although lamprey captures are known from

inland North Carolina streams and Albemarle Sound (Smith 1907,

Menhinick 1991). Schwartz et al. (1982) reported a 140-mm total length

(TL), 3.9-g specimen (UNC 8501) entangled in a gill net on the west

side (Station 19 west) of the Cape Fear River, 4 km north of Southport,

North Carolina, from waters of 10. 2C and 10 ppt salinity on 19 February

1974. Whether it was attached to a fish caught in the net was unknown.

Brimleyana 21:69-72, December 1994 69

70 Christopher Jensen and Frank J. Schwartz

Previous sea lamprey-shark parasitism records

We know of two verified records of sea lamprey-shark parasitism.

One involves a female sea lamprey and a basking shark (Cetorhinus

maximus), specimen 965-2-3-1, of the Nova Scotia Museum (Bigelow

and Schroeder 1948). The lamprey was 290-mm TL when preserved in

formalin. The 7.6-m-long basking shark, caught 29 June 1965 in a gill

net off Hopson Island (near Prospect), Halifax County, Canada, was

alive when the lamprey was removed. Attachment was just above and

anterior to the base of the anal fin, although sea lampreys often attach

to pectoral fins and along the dorsal and body sides (Cochran 1985,

1986). The second was a record of two adult lampreys, 180- and 250-

mm TL (USNM 130791) taken from an unknown species of shark

captured 3 June 1885 off Cape Charles, Virginia, at Albatross Station

2422 at 37°08'30"N, 74°33'30"W (Jenkins and Burkhead 1993).

Recent sea lamprey-shark parasitism records

South Carolina — We captured a female sea lamprey (UNC 17398),

168-mm TL, 8.8 g, on 6 February 1993 while longlining 69 km off South

Carolina in 31.1 m of water. Set location began at 33°10.9'N, 78°17.45,W

and ended at 33°00'N, 78°24.08'W. It was still attached to a 1280-mm

fork length (FL), male sandbar shark (Carcharhinus plumbeus) along the

shark's right lateral flank midway between the rear tips of the pelvic and

dorsal fins on the gray portion of the skin. Removal of the lamprey

revealed a round reddish area on the side of the body, which indicates

that it had been attached for some time before the shark's capture. Blood

oozed freely from the female lamprey's cloacal opening.

North Carolina — We know of five recent occurrences (March 1993)

of female sea lampreys parasitizing sharks captured from two different

locations off North Carolina; the host in one case was a 3-m-FL dusky

shark {Carcharhinus obscurus), the others three 3-m-FL sandbar sharks

(C. plumbeus). A dusky shark and one sandbar shark, captured by

fishermen longlining 74-km east-southeast off Masonboro, North Carolina,

carried one feeding lamprey attached near the cloaca of each shark.

But the lampreys were not retained by the fishermen who captured the

sharks.

Three additional female sea lampreys (UNC 17403, Table 1)

165-, 178-, and 343-mm TL, weighing 6.4, 9.5, and 70.7 g, respectively,

were captured 23 March 1993 during nightime longlining sets 46.2 km

east of Cape Lookout in 31-36-m waters. All three specimens parasitized

3-m-TL female dusky sharks, one was attached to a pelvic fin, the

others to the white skin of the cloacal area. No masses were taken of

any shark at sea. Body proportions of the North Carolina preserved sea

Sea Lamprey Parasitism 71

Table 1. Meristic and morphometric data for sea lampreys captured parasitizing

dusky and sandbar sharks caught off South Carolina (UNC 17398) and

North Carolina (17403), 1993. Lengths are expressed as a percentage of

the total length.

'Host female sandbar shark.

2Hosts all female dusky sharks.

3Dark adult body coloration prevented accurate myomere count.

lampreys (Table 1) were larger than those reported for a 136-mm-TL

specimen from Florida (Vladykov and Kott 1980).

Conclusions

Sea lamprey-shark parasitism occurrences are rarely reported because

fishermen or scientists often think that a reddened bleeding area on the

body is simply a bruise rather than a wound caused by a lamprey.

Likewise, a lamprey might have fallen off once a shark was landed,

making the association of the injury with a lamprey difficult. Information

on sea lampreys from sharks caught at sea may shed more information

on their occurrence, seasonality, water depth frequented, host preferences,

and biology of sea lampreys than is presently known. Lamprey parasitism

may be more damaging to marine fishes than now suspected.

ACKNOWLEDGMENTS— We thank owner Mike Merritt, Captain

Jimmy Murry, Captain Richard West, Chuck Wagner, and Steve Francis

of the Reel Action II for taking part in the longlining operation and sea

lamprey association information. S. Van Sant and others of the South

Carolina Marine Resources Center, Charleston, South Carolina, commented

on the absence of South Carolina lamprey records. J. Gilhen of the

Nova Scotia Museum, Halifax, and W. B. Scott, Huntsman Marine

Science Center, St. Andrews, New Brunswich, Canada, supplied informa-

tion on the Canadian sea lamprey-basking shark record. R. Jenkins,

72 Christopher Jensen and Frank J. Schwartz

Roanoke College, Salem, Virginia, was helpful with the early Virginia

lamprey record. L. White typed the text.

LITERATURE CITED

Beamish, F. W. H. 1980. Biology of the North American anadromous

sea lamprey, Petromyzon marinus. Canadian Journal of Fisheries and

Aquatic Sciences 37:1929-1943.

Bigelow, H. B., and W. C. Schroeder. 1948. Fishes of the western

North Atlantic 2. Cyclostomes. Memoirs Sears Foundation Marine

Research 1:29-58.

Cochran, P. A. 1985. Size selective attack by parasitic lampreys:

consideration of alternate null hypothesis. Oecologia 67:137-141.

Cochran, P. A. 1986. Attachment sites of parasitic lampreys: Compari-

sons among specimens. Environmental Biology of Fish 17:71-79.

Dempson, J. B., and T. R. Porter. 1993. Occurrences of sea lamprey,

Petromyzon marinus, in a Newfoundland River, with additional records

from the Northwest Altantic. Canadian Journal of Fisheries and

Aquatic Sciences 50:1266-1269.

Gilbert, C. R., and F. F. Snelson. 1992. Petromyzon marinus Linnaeus,

sea lamprey. Pages 122-127 in Rare and endangered biota of

Florida (C. R. Gilbert, editor). Florida University Press, Gainesville.

Haedrick, R. C. 1977. Sea lamprey from the deep ocean. Copeia

1977:767-768.

Hardesty, M. W., and L. C. Potter (editors). 1971. The biology of

lampreys. Volume I. Academic Press, New York, New York.

Jenkins, R., and N. Burkhead. 1993. Fishes of Virginia. American

Fisheries Society, Bethesda, Maryland.

Menhinick, E. F. 1991. The freshwater fishes of North Carolina. North

Carolina Wildlife Resources Commission, Raleigh.

Proceedings of the 1979 Sea Lamprey International Symposium. 1980.

Canadian Journal Fisheries and Aquatic Sciences 37:1-2214.

Schwartz, F. J., W. T. Hogarth, and W. P. Weinstein. 1982. Marine

and freshwater fishes of Cape Fear estuary, North Carolina, and

their distribution in relation to environmental factors. Brimleyana

7:17-37.

Smith, H. M. 1907. The fishes of North Carolina. North Carolina

Geological and Economic Survey. Volume 2. Raleigh, North Carolina.

Vladykov, V. D., and W. I. Follett. 1968. Lampetra richardsoni, a

new nonparasitic lamprey (Petromyzontidae) from western North America.

Journal of the Fisheries Research Board of Canada 22:139-158.

Vladykov, V. D., and E. Kott. 1980. First record of the sea lamprey,

Petromyzon marinus L., in the Gulf of Mexico. Northeast Gulf

Science 4:49-50.

Received 18 August 1993

Accepted 1 November 1993

Clutch Parameters of Storeria dekayi Holbrook

(Serpentes: Colubridae) from Southcentral Florida

Walter E. Meshaka, Jr.

Archbold Biological Station

P.O. Box 2057

Lake Placid, Florida 33852

ABSTRACT — I examined clutch characteristics from a series of

Storeria dekayi collected in southcentral Florida from March to

July 1990. Clutch size averaged 8.5 and was not significantly

associated with female body size. The small clutch sizes of this

sample conformed to predictions of clutch size reduction in southern

populations. However, the data did not support predictions of

increased clutch number in southern populations. Possibly, high

relative clutch mass detected in this population and an unaltered

breeding season hinder production of more than one clutch annually.

Two latitudinal clines in snakes have been proposed that predict

differences in clutch size and number along a geographic gradient. The

first predicts a decrease in clutch size from north to south (Fitch 1985).

The second hypothesis predicts an increase in clutch number at lower

latitudes concomitant with a longer reproductive season (Fitch 1970).

The reproductive biology of Storeria dekayi in Florida is poorly

known; however, parturition dates of this species are available from

Florida (Iverson 1978), and they do not differ from parturition dates in

more northern populations (Fitch 1970). Iverson's (1978) data do not

support the prediction of a latitudinal cline in clutch number for this

species.

With few exceptions, Iverson (1978) found that most Florida snake

species he examined did not conform to the prediction of multiple

clutches in southern populations. In this article I present additional

reproductive data for female S. dekayi from south Florida which permit

testing Fitch's (1985) hypothesis of clutch size reduction in southern

populations and further evaluation of the likelihood of multiple clutch

production in this species at the southern limit of its geographic range

(Fitch 1970).

METHODS

Snakes were collected from 1830 to 2200 hours from a paved

road (C-621) near Lake Placid, Highlands County, Florida, during March-

July 1990. All snakes observed were collected, frozen within 3 hours

of capture, and dissected the next day.

Brimleyana 21:73-76. December 1994 73

74 Walter E. Meshaka, Jr.

Condition of follicles was staged according to Kofron (1979). I

estimated clutch size by counting enlarged follicles or conceptuses.

Relative clutch mass, the quotient of clutch mass divided by the sum

of the clutch mass and the female body mass (Seigel and Fitch 1984),

was measured in females with fully developed conceptuses. All specimens

are located in the Archbold Biological Station vertebrate collection.

RESULTS AND DISCUSSION

Eighteen snakes (3 males, 15 females) were collected during 22.5

hours of searching. Snout-vent lengths (SVL) of males collected in

March (n = 1) and June (n = 2) were 23.0, 25.0, and 27.5 mm, respectively.

Snout-vent lengths and clutch parameters of females are summarized in

Table 1. Estimated clutch size was not significantly correlated with

SVL (r = 0.31, P>0.05).

Table 1. Snout-vent lengths (SVL) and clutch parameters of Storeria

dekayi collected from one location in Highlands County, Florida, 1990.

Relative

Clutch

Date Female SVL Clutch Size Mass Neonate SVL (cm)

6.3 + 0.306

7.7 + 0.500

8.4 + 0.097

1 Denotes a specimen collected in 1992 from the same site. Data not

analyzed with 1990 sample.

Clutch Parameters of Storeria dekayi 75

Mean clutch size (8.5) was similar to that found in Everglades

National Park by Dalrymple et al. (1992), and the samples from both

regions had a female - biased sex ratio. Female S. dekayi from Iverson's

(1978) northern Florida sample had smaller SVL than females from my

study (t = 3.681; df = 10; P < 0.004), but the two samples did not

differ significantly in clutch size. Although both mean female SVL

(27.3 mm) and clutch size (14.0) from S. dekayi near Lake Erie (King

1993) were significantly larger than those of my study (t = 2.12 P =

0.05; and / = 7.53 P < 0.00, respectively; df = 38), the difference in

SVL was marginal. Mean clutch sizes, 14.9 in Louisiana (Kofron 1979)

and 14 in New York (Clausen 1936), are also substantially larger than

those for Florida, which supports Fitch's (1985) prediction of smaller

clutch sizes in southern latitudes.

A review of relative clutch mass in snakes indicates that there is

a reduction in relative clutch mass among viviparous forms that may

reduce the risk of mortality in gravid females (Seigel and Fitch 1984).

The cost of lowering relative clutch mass is a reduction of clutch size,

offspring size, or both. Resources could limit production of more than

one clutch (Bull and Shine 1979), but a large clutch could compensate

for a single brood (Seigel and Fitch 1984). Mean relative clutch mass

in .V. dekayi from southern Florida was high (0.367) and similar to that

(0.372) recorded for S. dekayi from Maryland (Jones 1976). A high

relative clutch mass in southern Florida .V. dekayi may compensate for

a single small clutch produced each season.

In northern Florida, females with fully developed conceptuses

were recorded from July to September (Iverson 1978). In southern

Florida, the earliest date was May (Iverson 1978, my study), and in

Everglades National Park captive females gave birth from June to September

(Dalrymple et al. 1992). Collectively, the breeding season of Florida

populations of S. dekayi (Iverson 1978, Dalrymple et al. 1992, my

study) falls within the range of other populations (Fitch 1970, Kofron

1979). Further, my results did not indicate a reduction of relative clutch

mass, which could facilitate multiple clutch production, in southern

Florida S. dekayi.

CONCLUSIONS

Clutch frequency of this species in southern Florida has not been

determined to date, and multiple clutch production, even if infrequent,

has not been excluded. An annual sample of specimens or mark-recapture

will best answer this question. Results of my study do not support

Fitch's (1970) prediction of multiple clutch production by S. dekayi in

the southern part of the range.

76 Walter E. Meshaka, Jr.

However, clutch sizes from south Florida S. dekayi were smaller

than northern populations as predicted by Fitch (1985) and unaffected by

female body size. Possibly, a high relative clutch mass and an

unaltered breeding season limit this population to one brood annually.

ACKNOWLEDGMENTS— I would like to thank John W. Fitzpatrick,

Station Director, for support for this project at Archbold Biological

Station. Betty S. Ferster assisted in collecting snakes. Henry R. Mushinsky,

James N. Layne, and Sam D. Marshall reviewed an earlier version of

this manuscript.

LITERATURE CITED

Bull, J. J., and R. Shine. 1979. Iteroparous animals that skip oppor-

tunities for reproduction. American Naturalist 114:296-303.

Clausen, H. J. 1936. Observations on the brown snake, Storeria dekayi

(Holbrook), with special reference to the habits and birth of young.

Copeia 1936:98-102.

Dalrymple, G. H., T. M. Steiner, R. J. Nodell, and F. S. Bernardino,

Jr. 1992. Seasonal activity of the snakes of Long Pine Key,

Everglades National Park. Copeia 1991:294-302.

Fitch, H. S. 1970. Reproductive cycles of lizards and snakes. Univer-

sity of Kansas Museum of Natural History Miscellaneous Publica-

tion. 42:1-247.

Fitch, H. S. 1985. Variation in clutch and litter size in new world

reptiles. University of Kansas Museum of Natural History Miscella-

neous Publication. 76:1-76.

Iverson, J. B. 1979. Reproductive notes on south Florida snakes. Florida

Scientist 41:201-207.

Jones, L. 1976. A large brood for a Maryland Storeria dekayi dekayi.

Bulletin of the Maryland Herpetology Society 12:102-103.

King, R. B. 1993. Determinants in offspring number and size in the

brown snake, Storeria dekayi. Journal of Herpetology 27(2): 175-

185.

Kofron, C. P. 1979. Female reproductive biology of the brown snake,

Storeria dekayi, in Louisiana. Copeia 1979:463-466.

Seigel, R. A., and H. S. Fitch. 1984. Ecological patterns of relative

clutch mass in snakes. Oecologia 61:293-301.

Received 15 June 1993

Accepted 15 September 1993

Influence of Environmental Conditions on Flight

Activity of Plecotus townsendii virginianus

(Chiroptera: Vespertilionidae)

Michael D. Adam1, Michael J. Lacki, and Laura G. SIioemaker

Department of Forestry, University of Kentucky,

Lexington, Kentucky 40546

ABSTRACT — Flight activity of the Virginia big-eared bat (Plecotus

townsendii virginianus) was measured in relation to eight environ-

mental variables during 1990 and 1991 in Lee County, Kentucky.

Activity, measured as the mean nightly detection frequency of

bats fitted with transmitters, was positively related to percent

relative humidity and negatively related to moon phase and wind

speed. Multiple regression analysis showed relative humidity to

have the strongest association with flight activity of all the environmental

variables tested. An explanation for this pattern was that bats

reduced their foraging activity on nights of low relative humidity

to avoid excess water loss because of extremes in vapor pressure

deficits during flight. Other explanations for the observed activity

patterns may exist, but they were not investigated in our study.

A wide range of abiotic environmental variables affect flight

activity of bats, including sunlight, moonlight, temperature, wind speed,

and precipitation. Sunlight inhibits flight activity and serves to

synchronize circadian periodicity (Erkert et al. 1980). Moonlight

reduces flight activity (Erkert 1974) and is known to induce shifts in

foraging patterns (Fenton et al. 1977). Flight activity of bats increases

with temperature, with shorter activity periods on cooler nights (Anthony

et al. 1981) and extended bouts of activity on warmer nights (O'Farrell

et al. 1967). Sufficiently strong winds suppress flight activity (O'Farrell

et al. 1967), but the influence of slower air speeds, if any, is unknown.

Responses of bats to precipitation is not consistent among, or

even within, species. Heavy rainfall reduced flight activity of

Pipistrellus pipistrellus (Stebbings 1968), but did not do so in another

study (Swift 1980). The timing of rainfall events is important

(Felton et al. 1977), with rain at dusk known to delay nightly emergence

in Nycticeius humeralis (Watkins 1972).

In contrast, effects of relative humidity on flight activity of

bats have been suggested (Watkins 1972, Lacki 1984) but quantitative

data are lacking. Studies have demonstrated the importance of water

balance to bats under both laboratory (Bassett 1980, Bassett and

1 Present address: Coastal Oregon Productivity Enhancement, Hatfield Marine

Science Center, 2030 South Marine Science Drive, Newport, Oregon 97365-5296.

Brimleyana 21:77-85, December 1994 77

78 M. D. Adam, M. J. Lacki, and L. G. Shoemaker

Wiebers 1980) and free-ranging conditions (Kurta et al. 1989, Kurta

et al. 1990). This would suggest that selection for adaptations to

minimize water loss should evolve in bats, particularly in association

with flight because of the high surface area to volume ratio of bat

wings.

Bats inhabiting arid environments show a direct relationship

between urine concentrating ability and evapotranspiration to pre-

cipitation ratio (Bassett 1986). In wetter regions, bats should encounter

fewer problems of water balance. Flexibility in foraging strategy, such

as reduced activity on less humid nights (i.e., increased vapor

pressure deficits), should help to maintain water balance.

Using radio telemetry we monitored flight activity of a temperate

zone insectivorous bat, Plecotus townsendii virginianus, and related

observed activity patterns with data for local environmental conditions.

We tested the hypothesis that flight activity of P. t. virginianus is

reduced as relative humidity declines, which would be expected if

water balance is a critical selective pressure for temperate zone bats.

METHODS

The study area was located in the Cumberland Plateau,

Lee County, Kentucky. Lee County is 80% forested and sparsely

populated by humans (Newton et al. 1974). Mixed mesophytic forest is

the dominant habitat (Braun 1950) with most stands being second

growth timber because of past logging practices. The climate of the

region is temperate, characterized by warm and humid summers and

moderately cold winters. Average maximum and minimum tempera-

tures are 34C in August and -18C in January (Newton et al. 1974).

Average monthly precipitation is 9 cm (Newton et al. 1974). July and

October are the wettest and driest months, respectively. Additional

site details are provided by Adam et al. (1994).

Sixty bats were fitted with transmitters during 1990 and 1991, 30

each summer. Because this subspecies is very sensitive to disturbance

(Bagley 1984), we studied males in 1990 and females in 1991 and

addressed any problems that occurred with males in 1990 before

handling females in 1991. Bats were mist-netted as they emerged from

bachelor and maternity roosts, along cliffs, and on an abandoned

logging road. Bats were weighed, sexed, aged, and then fixed with a

0.8-g transmitter (Type BD-2A, 172-173 MHz; Holohil Systems, Ltd.,

Ontario, Canada, and Wildlife Materials, Inc., Carbondale, Illinois)

on the dorsum between the scapulae. The surface was prepared by

trimming the fur with scissors and applying surgical cement

designed to hold the transmitter for about 10 days.

Flight Activity of Bats 79

Three telemetry periods, each spanning five nights, were

conducted in each year. In 1990 male bats were radiotracked from 2

to 6 June (n = 9 bats), 16 to 20 July (ti = 10), and 6 to 10 August

(n = 11). In 1991 females were tracked from 10 to 14 May (n = 9), 17

to 21 June (n = 10), and 5 to 9 August (n = 11). Bats were tracked

from both fixed and mobile stations. Fixed stations were positioned

on the top of cliffs enclosing a hollow with either a bachelor or

maternity roost. Distances between fixed stations averaged 857 m

in 1990 and 509 m in 1991. Mobile stations were along road routes

throughout the surrounding areas. Three TRXlOOOs receivers (Wildlife

Materials, Inc., Carbondale, Illinois) were used to locate bats, with an

additional TRX2000s receiver used in 1991. Receivers were coupled

to a 3- or 5-element yagi antenna. Signals were searched for at

20-minute intervals from sunset to sunrise. Bats may have been

detected in multiple intervals by more than one receiver.

Telemetry data were organized into nightly rates of bat activity by

converting signal responses into mean nightly detection frequencies

(NDF) calculated as

t n

NDF = 2(2 (dlo)ln)tl

;=1 i=\

where NDF = mean nightly detection frequency, t - number of time

intervals post-sunset, n - number of bats with transmitters, d - number

of receivers detecting a bat in an interval, and o - number of

receivers operating in an interval.

Patterns of activity were also derived for each sampling period by

converting signal responses to mean detection frequencies per time

interval (TIDF) calculated as

k n

TIDF = 2 ( 2 (dlo)ln)kx

7 = 1 i"=l

where TIDF = mean detection frequency per time interval, and k -

number of days sampled.

Nightly environmental conditions were obtained from the

Heidelberg, Kentucky, weather station located 11 km, from the bach-

elor roost. The foraging radius of bats from the bachelor and maternity

colonies was large (Adam et al. 1994), rendering sampling for environ-

mental conditions throughout the study site impractical. Data for

eight variables were analyzed: daily maximum temperatures (°C), daily

minimum temperature (°C), total precipitation the day of sampling (cm),

total precipitation on the day preceding sampling (cm), average daily

80 M. D. Adam, M. J. Lacki, and L. G. Shoemaker

relative humidity (%), average daily wind speed (km/hr), average daily

barometric pressure (millibars), and moon phase (% of full moon

illumination).

We initially tested NDF against all environmental variables

separately using simple linear regression, and only those variables meeting

the 0.10 probability level were retained. Backwards stepwise multiple

regression was then used for modeling NDF against environmental variables.

A probability of >0.10 was used for removal of a variable from the

model. Data for 1990 and 1991 were combined for analysis. Differences

between years (sexes) were checked using analysis of variance

(ANOVA), with the day of sampling as a block effect. Relative humidity

and moon phase were arcsine transformed to correct for nonnormality

of the data.

RESULTS

All bats that were fitted with transmitters were adults except

for five juvenile males in August 1990 and one juvenile female in

August 1991. Data for body mass and reproductive condition are pre-

sented elsewhere (Adam et al. 1994). Transmitters did not appear to

adversely affect the bats. Bats showed no difficulty flying upon

release and were located at considerable distances from known roosts.

During August 1991, two females were captured which had previously

been fitted with a transmitter. Masses for these females were not different

from the average mass of other females captured during that period.

Although transmitters on some bats emitted signals for up to 10 days,

we considered 5 days to be the normal life of transmitters in this study.

Data from all 60 bats were used, regardless of the life of the transmitters.

Analysis of variance demonstrated no day effect (F = 2.81; df =

4, 20; P > 0.10) and no interaction between day and year (F = 1.48; df

= 4, 20; P > 0.10). Activity rates were higher in 1991 than in 1990

(F = 22.9; df =1, 20; P < 0.0001) (Fig. 1). It is unclear whether this

difference was due to sex or varying conditions between summers, as

males and females were not tested in both summers.

Bats did not emerge to forage until 30 to 45 minutes post-sunset

(Fig. 1). Males in 1990 exhibited a pattern with highest activity during

the first few hours of the night (Fig. la). The activity of females in

1991 was more sustained throughout the night (Fig. lb). Females in

these periods were either pregnant (May) or lactating (June) (Adam et

al. 1994), suggesting the use of night roosts and/or shorter foraging

bouts which allowed them to return to the maternity roost to nurse

their young.

Flight Activity of Bats

81

Period I

Period II

Period

0.30

0.24

Q 0.18

h-

c

CD

0)

^ 0.12

0.06

0.00

10 15 20

Interval Post-sunset

25

30

0.30

10 15 20

Interval Post-sunset

Fig. 1. Mean frequency of detection by 20-minute time intervals

(I IDF) post-sunset for P. t. virginiamis comparing sampling periods; males

(a): Period 1 = June. Period II = July, Period III = August 1990;

females (b): Period I = May, Period II = June, Period III = August

1991). Lee Countv, Kentucky.

82 M. D. Adam, M. J. Lacki, and L. G. Shoemaker

Three environmental variables were related to NDF (Table 1);

NDF was positively associated with percent relative humidity, and negatively

related to both wind speed and moon phase. Stepwise regression demonstrated

a significant (F = 5.93, P = 0.0073, R2 = 0.30) relationship of NDF

with relative humidity and moon phase, eliminating wind speed from

Table 1. Regression analyses of mean nightly detection frequency (NDF)

with environmental variables for P. t. virginianus, Lee County, Ken-

tucky, summers 1990 and 1991.

Table 2. Values for nightly detection frequency (NDF) and environmental

variables for P. t. virginianus, Lee County, Kentucky, summers 1990 and

1991.

Range

Variable Mean CV High Low

NDF 0.08 88.5 0.30 0.01

Relative humidity (%)

Moon phase (%)

Wind speed (km/hr)

Barometric pressure (mb)

Maximum temperature (C)

Minimum temperature (C)

Precipitation (cm)

Prior precipitation1 (cm)

' Represents rainfall during the day prior to sampling.

Flight Activity of Bats 83

the final model. No pattern was observed between significant and

nonsignificant environmental variables using coefficients of variation

(Table 2), indicating no bias because of differences in the amount of

variability in these sets of variables.

DISCUSSION

We found a direct relationship between flight activity of P. t.

virginianus and ambient relative humidity, with bats exhibiting

reduced activity on nights with lower percent relative humidity. Using

mist-net captures as a measure of activity, Lacki (1984) observed a

similar pattern for Myotis lucifugus and suggested reduction in flight

activity as a behavioral mechanism for avoiding water loss on

nights when ambient conditions created large vapor pressure deficits.

Bats experience water loss in flight and can lose water because of

roost conditions (Carpenter 1969). Substantial water loss accompanies

digestion in M. lucifugus (Bassett and Wiebers 1980) with water

balance in female M. lucifugus (Kurta et al. 1989) and female Eptesicus

fuscus (Kurta et al. 1990) influenced by reproductive condition. Because

water loss by bats is dependent on ambient temperature and water

vapor pressure under laboratory conditions (Bassett 1980) and water

loss increases with higher levels of flight activity (Studier 1970), we

suggest that our data support the existence of a behavioral response

by bats for avoiding extremes in vapor pressure deficits during flight.

Factors such as prey activity and availability may also contribute

to the observed activity patterns, but were not investigated in our study.

Data comparing the abundance of insect prey with ambient relative

humidity are sparse; however, in one study no association was found

between relative humidity and activity of moths (Mizutani 1984).

Moths are the predominant item in the diet of P. t. virginianus (Sample

and Whitmore 1993).

The inverse relationships we observed between activity of P. t.

virginianus and both moon phase and wind speed are consistent with

other findings reported in the literature (O'Farrell et al. 1967, Erkert

1974, Fenton et al. 1977). Whether avoidance of moonlight by P. t.

virginianus was because of predators or availability of insect prey or

both is unclear. Several species of owls were common in the study

area, and abundance of insect prey has been shown to be negatively

related to moon phase (Anthony et al. 1981). Observations at a maternity

colony of a related subspecies, the Ozark big-eared bat (P. t. ingens),

found no patterns between flight activity and indicies of moon

brightness (Clark 1991).

84 M. D. Adam. M. J. Lacki, and L. G. Shoemaker

ACKNOWLEDGMENTS— We thank C. M. Cunningham, R. R.

Currie, B. J. Deetsch, J. R. MacGregor, D. Miller, D. C. Yancy, and R.

J. Yablonsky for assistance in the field. Funding was provided by the

United States Forest Service, the United States Fish and Wildlife

Service, the Kentucky Department of Fish and Wildlife Resources,

and The University of Kentucky, College of Agriculture. This

research was conducted with the approval of the Institutional Animal

Care and Use Committee of The University of Kentucky (Protocol

#90-0007A). This is a contribution of the Kentucky Agricultural

Experiment Station, paper number 93-8-129.

LITERATURE CITED

Adam, M. D., M. J. Lacki, and T. G. Barnes. 1994. Foraging areas

and habitat use of the Virginia big-eared bat in Daniel Boone

National Forest, Kentucky. The Journal of Wildlife Management

58:462-469.

Anthony, E. L. P., M. H. Stack, and T. H. Kunz. 1981. Night roosting

and the nocturnal time budget of the little brown bat, Myotis

lucifugus: Effects of reproductive status, prey density, and environ-

mental conditions. Oecologia 51:151-156.

Bagley, F. 1984. A recovery plan for the Ozark big-eared bat and the

Virginia big-eared bat. United States Fish and Wildlife Service,

Twin Cities, Minnesota.

Bassett, J. E. 1980. Control of post-prandial water loss in Myotis

lucifugus lucifugus. Comparative Biochemistry and Physiology 65A:497-

500.

Bassett, J. E. 1986. Habitat aridity and urine concentrating ability of

Nearctic, insectivorous bats. Comparative Biochemistry and Physiol-

ogy 83 A: 125-131.

Bassett, J. E., and J. E. Wiebers. 1980. Effect of food consumption

on water loss in Myotis lucifugus. Journal of Mammalogy 61:744-

747.

Braun, E. L. 1950. Deciduous forests of eastern North America. Hafner,

New York, New York.

Carpenter, R. E. 1969. Structure and function of the kidney and the

water balance of desert bats. Physiological Zoology 42:288-302.

Clark, B. S. 1991. Activity patterns, habitat use, and prey selection

by the Ozark big-eared bat (Plecotus townsendii ingens). Ph.D. Thesis,

Oklahoma State University, Stillwater.

Erkert, H. G. 1974. Der Einfluss des Mondlichtes auf die Aktivitatsperiodik

nachtaktiver Saugetiere. Oecologia 14:269-287.

Erkert, H. G., S. Kracht, and U. Haussler. 1980. Characteristics of

circadian activity systems in Neotropical bats. Pages 95-104 in

Proceedings of the Fifth International Bat Research Conference (D.

E. Wilson and A. L. Gardner, editors). Texas Tech Press, Lubbock.

Flight Activity of Bats 85

Fenton, M. B., N. G. H. Boyle, T. M. Harrison, and D. J. Oxley.

1977. Activity patterns, habitat use, and prey selection by some

African insectivorous bats. Biotropica 9:73-85.

Kurta, A., G. P. Bell, K. A. Nagy, and T. H. Kunz. 1989. Water

balance of free-ranging little brown bats (Myotis lucifugus) during

pregnancy and lactation. Canadian Journal of Zoology 67:2468-

2472.

Kurta, A., T. H. Kunz, and K. A. Nagy. 1990. Energetics and water

flux of free-ranging big brown bats (Eptesicus fuscus) during pregnancy

and lactation. Journal of Mammalogy 71:59-65.

Lacki, M. J. 1984. Temperature and humidity induced shifts in the

flight activity of little brown bats. Ohio Journal of Science 84:264-

266.

Mizutani, M. 1984. The influences of weather and moonlight on the

light trap catches of moths. Applied Entomological Zoology 19:133—

141.

Newton, J. H., C. W. Hail, T. P. Leathers, P. M. Love, J. G. Stapp, V.

Vaught, and P. E. Avers. 1974. Soil survey of Estill and Lee

counties, Kentucky. United States Department of Agriculture Soil

Conservation Service and Forest Service, Washington, D.C.

O'Farrell, M. J., W. G. Bradley, and G. W. Jones. 1967. Fall and

winter bat activity at a desert spring in southern Nevada. South-

western Naturalist 12:163-171.

Sample, B. E., and R. C. Whitmore. 1993. Food habits of the

endangered Virginia big-eared bat in West Virginia. Journal of

Mammalogy 74:428-435.

Stebbings, R. E. 1968. Measurements, composition and behavior of a

large colony of the bat Pipistrellus pipistrellus. Journal of Zoology

156:15-33.

Studier, E. H. 1970. Evaporative water loss in bats. Comparative

Biochemistry and Physiology 35:935-943.

Swift, S. M. 1980. Activity pattern of Pipistrelle bats (Pipistrellus

pipistrellus) in north-east Scotland. Journal of Zoology 190:285-

295.

Watkins, L. C. 1972. A technique for monitoring the nocturnal

activity of bats, with comments on the activity patterns of the

evening bat, Nycticeius humeralis. Transactions of the Kansas Academy

of Science 74:261-268.

Received 1 February 1994

Accepted 10 March 1994

86

The Pygmy Shrew, Sorex hoyi winnemana

(Insectivora: Soricidae), from the Coastal Plain

of North Carolina

Thomas M. Padgett

North Carolina Wildlife Resources Commission

Route 2, Box 583A

Elizabethtown, North Carolina 28337

AND

Robert K. Rose

Department of Biological Sciences

Old Dominion University

Norfolk, Virginia 23529-0266

ABSTRACT — Sorex hoyi winnemana from two counties in

extreme northeastern North Carolina represent the first docu-

mented specimens of pygmy shrews from the Coastal Plain

and the first collected in North Carolina in 50 years. The 15

pygmy shrews were collected from a variety of habitat types,

ranging from shrubby-grassy fields in ditched peaty wetlands

to managed pine plantations and upland hardwood forests.

The smallest of the North American long-tailed shrews (genus

Sorex) is Sorex hoyi, aptly named the pygmy shrew. At 2-3 g with

a total length of 70-86 mm, of which 25-33 mm is tail, the pygmy

shrew is the smallest mammal in North Carolina and among the

world's smallest mammals. The masked shrew (Sorex cinereus) and

the southeastern shrew (S. longirostris) are similar in appearance and

in proportions, but both are somewhat larger (Webster et al. 1985).

Besides having slightly longer bodies and tails, S. cinereus and S.

longirostris have five prominent unicuspids, whereas the pygmy shrew

has only three distinct unicuspids, the third and fifth being greatly

reduced.

The pygmy shrew is distributed throughout the northern tier of

states from Minnesota through Maine and extensively in Canada and

Alaska, with fingers of distribution extending southward in the Rocky

and Appalachian Mountains (Hall 1981), which suggests the animals

need boreal conditions. However, studies involving the extensive use

of pitfall traps have documented that S. hoyi is more widely distributed

and occupies a wider range of habitats in the southeastern part of its

geographic distribution than it does elsewhere. In Virginia, pygmy

shrews have been found in at least 20 counties, including seven counties

Brimleyana 21:87-90 December 1 QQ4 87

88 Pygmy Shrew

west of the Blue Ridge where they were unknown before 1980 (Pagels

1987).

In North Carolina, the pygmy shrew is considered to be rare and

of undetermined status (Webster 1987), in part because only three

specimens, all from the extreme western mountainous region of the

state, have been found. Of these, two specimens are from Bent Creek

(23 April 1928), Buncombe County, and the other (6 September 1941)

from Newfound Gap in nearby Swain County (Webster 1987).

During extensive studies of the southeastern shrew in the vicinity

of the Great Dismal Swamp National Wildlife Refuge (Refuge) of

southeastern Virginia and northeastern North Carolina from 1986-91,

we collected Sorex hoyi from Camden and Gates counties, in extreme

northeastern North Carolina. These represent the first documented

specimens of Sorex hoyi from North Carolina in nearly 50 years, as

well as the first of this species from the Coastal Plain (Lee et al.

1982).

METHODS

We used pitfall traps, consisting of #10 metal cans sunk flush

into the ground and half-filled with water, to collect small mammals.

Pitfall traps are very effective in trapping small cryptic mammals

such as shrews (Padgett 1991). These traps offer advantages in that

they do not need to be checked daily, are relatively maintenance free,

and capture small shrews (and some other species of small mammals)

that normally are difficult to collect in other traps. We placed pitfall

traps in transects along roads and trails at measured intervals or in

0.25-ha grids spaced 12.5 m apart in 5 x 5 arrays. Each transect or

grid was trapped for 3-4 weeks.

One Gates County site was located along Weyerhaeuser Ditch in

the Refuge 2 km north of Highway 158, and the other was a transect

extending from the escarpment on the western boundary of the Refuge

into the swamp. This transect was located 2 km northeast of the

intersection of Route 32 and State Route 1332. The Camden County

grids were located on a large tract 9 km east-northeast of South

Mills.

RESULTS AND DISCUSSION

We collected 15 Sorex hoyi, referable to S. h. winnemana, during

the course of our survey, nine from the southern section of the Refuge

in Gates County and six outside the Refuge on three grids in Camden

County. Within the Refuge, eight animals were collected from the

site adjacent to Weyerhaeuser Ditch. This site was located in an

Thomas M. Padgett and Robert K. Rose 89

upland (or mesic) area of the Dismal Swamp locally referred to as

"mesic islands"; these are remnant Pleistocene marine deposits oriented

in east-west directions. In contrast to the surrounding bottomland

forest of tupelo (Nyssa aquatica), red maple (Acer rubrum), and scattered

bald cypress (Taxodium distichum), these "island" habitats have such

species as American beech (Fagus grandifolia), swamp chestnut oak

(Quercus michauxii), blackgum (Nyssa sylvatica), and loblolly pine

(Pinus taeda). The remaining specimen from the Refuge was collected

along the Nansemond (or Suffolk) Escarpment, a Pleistocine feature

that delineates the western boundary of the Dismal Swamp, in a transi-

tional wetland forest composed of both upland and bottomland species.

The six pygmy shrews from Camden County were taken in a

variety of habitats, all with black peaty loam soils, sometimes with

some sand component. Five pygmy shrews came from recently (<2

years) clearcut sites regenerating mostly in mixed grasses and a few

shrubs, and one was collected from a 15-year-old loblolly pine plan-

tation with pine straw as virtually the only ground cover.

The specimens from Gates County ranged from 64 to 81 mm (x

= 75.44 mm), whereas those from Camden County ranged from 78 to

81 mm (x = 79.67 mm). In the Gates County specimens, the tails

averaged 36% of total length, compares to 33% in the Camden County

specimens. Although the sex of some individuals could be determined,

we were unable to establish useful sex ratios. Other shrews caught at

one or more sites yielding pygmy shrews include the Dismal Swamp

southeastern shrew, (Sorex longirostris fisheri), the short-tailed shrew

(Blarina brevicauda) and in Camden County, the least shrew (Cryptotis

parva).

As is typical of studies using pitfall traps in Virginia (John

Pagels, Virginia Commonwealth University and Kurt Buhlman,

Virginia Natural Heritage Program, personal communication), the

distribution of pygmy shrews appears to be patchy. For example, in

Camden County pygmy shrews were caught on three of 10 grids, four

on one site, and one on each of the others; seven other sites yielded

none. In Gates County in the Refuge, pygmy shrews were taken from

two of six sites in habitats that seemed comparable.

We verified that pygmy shrews occur in a range of habitats; we

caught shrews in fields in early succession, maturing loblolly pines,

and in mature deciduous forest. In the Refuge (Gates County), pygmy

shrews were collected on slightly higher and better drained sites than

the surrounding more typical Dismal Swamp forested swampland

dominated by tupelo, red maple, and bald cypress. The Camden County

site is located in the Pasquotank River drainage, and therefore is also

90 Pygmy Shrew

a part of the historic Dismal Swamp, but the land has been ditched,

creating slightly drier conditions than probably prevailed there before

development. John Pagels (personal communication), who has used

pitfall traps in extensive studies in many counties in Virginia, has

caught pygmy shrews in many different habitats but not in wetland

habitat. In Virginia, Pagels (1987) has collected Sorex hoyi with six

other soricid shrews, but not with Blarina. In conclusion, our studies

indicate that in North Carolina, as elsewhere, pygmy shrews have

patchy distributions in a range of habitats; they probably occur through-

out much more of the state than is presently known.

ACKNOWLEDGMENTS— We thank David Webster for useful

comments on an earlier draft and Mary K. Clark for receiving the

specimens into the collections of the North Carolina State Museum of

Natural Sciences in Raleigh and for verifying their identifications.

These studies were conducted under permits given by the North Caro-

lina Wildlife Resources Commission and the Virginia Department of

Game and Inland Fisheries.

LITERATURE CITED

Hall, E. R. 1981. The Mammals of North America. John Wiley &

Sons, New York, New York.

Lee, D. S., J. B. Funderburg, Jr., and M. K. Clark. 1982. A distributional

survey of the North Carolina mammals. Occasional Papers of the

North Carolina Biological Survey. 1987-10. North Carolina State

Museum of Natural History, Raleigh.

Padgett, T. M. 1991. The identification, distribution, and status of

the threatened Dismal Swamp Shrew {Sorex longirostris fisheri).

M.S. Thesis, Old Dominion University, Norfolk, Virginia.

Pagels, J. F. 1987. The pygmy shrew, rock shrew, and water shrew:

Virginia's rarest shrews (Mammalia: Soricidae). Virginia Journal

of Science 38(4):364-368.

Webster, Wm. D. 1987. Sorex hoyi winnemana. Pages 40-41 in

Endangered, threatened, and rare fauna of North Carolina, Part 1. A

re-evaluation of the mammals (M. K. Clark, editor). Occasional Papers

of the North Carolina Biological Survey. 1987-3. North Carolina

State Museum of Natural Sciences, Raleigh.

Webster, Wm. D., J. F. Parnell, and W. C Biggs, Jr. 1985. Mammals

of the Carolinas, Virginia, and Maryland. University of North Carolina

Press, Chapel Hill.

Received 1 June 1993

Accepted 30 July 1993

Additional Records of the Pygmy Shrew,

Sorex hoyi winnemana Preble (Insectivora: Soricidae),

in Western North Carolina

Joshua Laerm

Museum of Natural History and Institute of Ecology

University of Georgia, Athens, Georgia 30602

William M. Ford

Daniel B. Warnell School of Forest Resources

University of Georgia, Athens, Georgia 30602

Daniel C. Weinand

Museum of Natural History

University of Georgia, Athens, Georgia 30602

ABSTRACT — Additional records of the pygmy shrew, Sorex hoyi

winnemana Preble, are reported from 14 localities in 7 counties

of western North Carolina. Results of recent surveys in adjacent

regions of Tennessee and Georgia indicate that the species is

widely distributed in the extreme southern Appalachian Mountains,

including North Carolina, but is nowhere abundant.

The pygmy shrew, Sorex hoyi winnemana, has been regarded as

one of the rarest mammals in the southeastern United States. In 1980

(see Diersing 1980, Handley et al. 1980), there were only 17 records

known from southern Illinois east to Maryland and south throughout

the Appalachian highlands to the Carolinas and Georgia. However,

more recently, considerable information on the distribution, abundance,

and habitat associations of this species has become available from Indiana

(Caldwell et al. 1982, Cudmore and Whitaker 1984), Virginia (Handley

et al. 1980, Pagels 1987, Pagels et al. 1992, Mitchell et al. 1993),

Kentucky (Caldwell 1980, Caldwell and Bryan 1982), Tennessee (Kennedy

et al. 1979, Kennedy and Harvey 1980, Tims et al. 1989, Harvey et al.

1991, Harvey et al. 1992, Feldhamer et al. 1993), South Carolina (Mengak

et al. 1987), and Georgia (Wharton 1968). This information indicates

that this subspecies can be found over a wider range of habitats and

geographic area than previously known. Although nowhere abundant,

it may be common where it occurs.

The first North Carolina records of Sorex hoyi were of two individuals

collected by A. H. Howell and reported by Jackson (1928) from Bent

Creek Experimental Forest in Pisgah National Forest, Buncombe County.

Webster (1987) indicated that the Buncombe County specimens were

erroneously reported from Transylvania County by Smith et al. (1960),

Brimleyana 21:91-96, December 1994 91

92 J. Laerm, W. M. Ford, and D. C. Weinand

Diersing (1980), and Lee et al. (1982). However, a single specimen

was subsequently collected in Transylvania County, at Cedar Mountain,

and reported by Mengak et al. (1987). Additionally, Hoffmeister (1968)

reported a single specimen from Newfound Gap, Swain County. Thus,

until recently the species was represented by only four specimens

from North Carolina.

In August 1993 E. v. d. Berghe (Appalachian Environmental Research

Center, Frostburg, Maryland) submitted to J. L. collections of shrews

that were made in North Carolina as incidental captures in surveys for

carabid beetles. Included in these collections were two Sorex hoyi from

McDowell County and an additional specimen from Graham County.

Subsequently, we obtained a record of an additional specimen from M.

Steele (Wilkes University, Wilkes Barre, Pennsylvania), who recovered

S. hoyi from Mount Mitchell, Avery County, in pitfall studies for soricid

parasites.

In this volume, Padgett and Rose (1994) report on significant

new records of S. hoyi from the Dismal Swamp area in the extreme

northeastern portion of North Carolina. Because the species is listed as

special concern in North Carolina by the North Carolina Wildlife Resources

Commission (see also Webster 1987), additional information on its

distribution is needed. We report on new records of this species and

the results of preliminary surveys to document additional records in

western North Carolina.

METHODS

To document the occurrence of S. hoyi in regions from which it

had not previously been reported, we established pitfall traplines at

eleven sites in Clay, Cherokee, Jackson, and Macon counties in extreme

western North Carolina from December 1993 through January 1994.

Additionally, we established 15 pitfall trap lines at Coweeta Hydrological

Laboratory along an altitudinal gradient from 710 m to 1,525 m from

April through May 1994.

Traplines consisted of twenty, 32-ounce plastic cups (11-cm lip

diameter, 14-cm depth) placed flush with or below the surface of the

ground and adjacent to rotting logs, stumps, rocks, or other forest floor

debris. Pitfalls were placed approximately 10-m apart along a linear

transect and were set in a diversity of typical southern Appalachian

forest habitats and checked biweekly. Because the species is protected

in North Carolina, we were required to discontinue trapping after the

second record of S. hoyi was obtained at a site.

Pygmy Shrew Records 93

RESULTS AND DISCUSSION

Our survey of 13.200 trap nights yielded 10 records of Sorex

hoyi: one each from Cherokee. Clay, and Jackson counties and seven

from Macon County. Five of the Macon County records were obtained

at Coweeta Hydrological Laboratory. The western North Carolina sites

ranged in elevation from 700 m to 1,524 m in a variety of moderate to

mesic hardwood to mixed hardwood-pine sites. Sorex hoyi was taken

in a heath bald dominated by rhododendron {Rhododendron maximum):

cove hardwood communities dominated by yellow poplar (Liriodendron

tulipifera), northern red oak (Quercus rubra), white oak (Q. alba), and

buckeye (Aesculus octandra); moderately xeric sites dominated by white

oak, northern red oak, hickory (Carya spp.), chestnut oak (Q. prinus),

scarlet oak (Q. coccinea). and white pine (Pinus strobus); and streamside

communities dominated by eastern hemlock (Tsuga canadensis) and

rhododendron. Standard body measurements for the 13 new North Carolina

specimens available to us are as follows: total body length ( x = 68.7

mm, range = 65.0-73.5 mm), tail length (x = 26.2 mm, range = 24.0-

28.4), and hind foot length (x = 8.0, range = 7.0-8.5 mm).

The few historical collection records of this species from western

North Carolina probably do not necessarily reflect its rarity in the area

but rather inappropriate collecting methodology. In the past 10-12 years

significant information regarding this species has become available,

largely through pitfall trapping, which has been shown to be the most

(if not the only) effective method of collecting insectivores (Handley

and Kalko 1993). Trapping efforts by Harvey et al. (1991) in the southern

districts (Monroe and Polk counties) and Harvey et al. (1992) in the

northern districts (Unicoi, Johnson, and Carter counties) of the Cherokee

National Forest, Tennessee, have indicated S. hoyi to be widely distributed

but nowhere abundant. Harvey et al. (1991) reported 16 captures in

226,054 pitfall trap nights in a diversity of forest habitats in the southern

portions of the Cherokee National Forest ranging in elevation from

396 m to 1,122 m. Harvey et al. (1992) report 13 captures in 389,995

pitfall trap nights in a similar diversity of forest habitats in the northern

portions of the Cherokee National Forest ranging in elevation from

695 m to 1,524 m. Similarly, in 67,500 pitfall trap nights we have

recorded 72 S. hoyi from 42 localities throughout the entire Blue Ridge

Province of Georgia where the species is widely distributed in a variety

of forest habitat types, including clearcuts, early and mid-successional

forest stages, as well as mature stands in streamside, xeric, and mesic

communities at elevations ranging from 700 m to 1,372 m. In Georgia,

it is nowhere abundant, but is widely distributed.

94 J. Laerm, W. M. Ford, and D. C. Weinand

COLLECTION RECORDS

Records of Sorex hoyi from western North Carolina using acronyms

for the museum collections in which the specimens are housed follow

Yates et al. (1987).

Avery Co.: Mount Mitchell (1, M. Steele Collection). Buncombe

Co.: Bent Creek Experimental Station, Pisgah National Forest (2, USNM).

Cherokee Co.: Nancy Gap (2, UGAMNH). Clay Co.: 3.0 mi. E. Fires

Creek Recreation Area (1, UGAMNH). Graham Co.: 15 mi. NW Robbinsville,

Joyce Kilmer Memorial Forest (1, UGAMNH). Jackson Co.: 4.5 mi. S

Cashiers (1, UGAMNH). Macon Co. 3.0 mi. W. Highlands (1, UGAMNH),

0.5 mi. E. Winding Stair Gap on U.S. 64 (1, UGAMNH); Albert Mt.,

Coweeta Hydrological Laboratory (1 UGAMNH); Cold Spring Gap,

Coweeta Hydrological Laboratory (1 UGAMNH); Cold Spring Cove,

Coweeta Hydrological Laboratory (1 UGAMNH); Lick Branch, Coweeta

Hydrological Laboratory (2 UGAMNH). McDowell Co.: Balsam Gap,

along Blue Ridge Parkway at mile marker 357 (1, UGAMNH); Glassmine

Falls, along Blue Ridge Parkway at mile marker 362 (1, UGAMNH).

Swain Co.: Newfound Gap (1, UIMNH). Transylvania Co.: Cedar Mountain

(1, CUVC).

ACKNOWLEDGMENTS— Kathy Barker, Alex Menzel, Kendal

Cochran, Eric Fowler, Jane Ellis, Kate Schumacher, Joe Devivo, and

Mac Callahan worked in snow, rain, ice, and dirt. We thank Ron Escano

and Rod McClanahan of the Nantahala National Forest and Wayne

Swank of Coweeta Hydrological Laboratory for authorization to undertake

surveys on Forest Service lands. These surveys were conducted under

North Carolina Wildlife Resources Scientific Collecting Permits (93-

05, 94-05, 93-ES-65, 94-ES-65). Support for this project was provided

by The University of Georgia Museum of Natural History and NSF

grant BSR 9011661.

LITERATURE CITED

Caldwell, R. S. 1980. First records of Sorex dispar and Microsorex

thompsoni in Kentucky with distributional notes on associated species.

Transactions of the Kentucky Academy of Science 41:46-47.

Caldwell, R. S., and H. Bryan. 1982. Notes on the distribution and

habits of Sorex and Microsorex (Insectivora: Soricidae) in Ken-

tucky. Brimleyana 8:91-100.

Caldwell, R. S., C. K. Smith, and J. O. Whitaker, Jr. 1982. First

records of the smoky shrew, Sorex fumeus, and pygmy shrew, Microsorex

hoyi, from Indiana. Proceedings of the Indiana Academy of Science

91:606-608.

Pygmy Shrew Records 95

Cudmore, W. W., and J. O. Whitaker, Jr. 1984. The distribution of

the smoky shrew, Sorex fumeus, and the pygmy shrew, Microsorex

hoyi, in Indiana with notes on the distribution of other shrews.

Proceedings of the Indiana Academy of Science 93:469-474.

Diersing, V. E. 1980. Systematics and evolution of the pygmy shrews

(Subgenus Microsorex) of North America. Journal of Mammalogy

61:76-101.

Feldhamer, G. A., R. W. Klann, A. S. Gerard, and A. C. Drickell.

1993. Habitat partitioning, body size and timing of parturition in

pygmy shrews and associated soricids. Journal of Mammalogy 74:403-

411.

Handley, C. O., Jr., and E. K. V. Kalko. 1993. A short history of

pitfall trapping in America, with a review of methods currently

used for small mammals. Virginia Journal of Science 44:19-26.

Handley, C. O. Jr., J. F. Pagels, and R. H. De Rageot. 1980. Microsorex

hoyi winnemana Preble. Pages 545-547 in Endangered and threat-

ened plants and animals of Virginia (D. M. Linzey, editor). Center

for Environmental Studies, Virginia Polytechnic Institute and State

University, Blacksburg.

Harvey, M. J., C. S. Chaney, and M. D. McGimsey. 1991. Distribution,

status, and ecology of small mammals of the Cherokee National

Forest, Tennessee (Southern Districts). Report to the United States

Forest Service. Manuscript on file, Center for the Management, Utilization,

and Protection of Water Resources, Tennessee Technical University,

Cookville.

Harvey, M. J., M. D. McGimsey, and C. S. Chaney. 1992. Distribu-

tion, status, and ecology of small mammals of the Cherokee Na-

tional Forest, Tennessee (Northern Districts). Report to the United

States Forest Service. Manuscript on file, Center for the Manage-

ment, Utilization, and Protection of Water Resources, Tennessee

Technical University, Cookville.

Hoffmeister, D. F. 1968. Pygmy shrew, Microsorex hoyi winnemana,

in Great Smoky Mountains National Park. Journal of Mammalogy

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tailed shrews (genera Sorex and Microsorex). North American Fauna

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Tennessee rare vertebrates (D. C. Eager and R. M. Hatcher, editors).

Tennessee Wildlife Resources Agency and Tennessee Department of

Conservation, Nashville.

Kennedy, M. L., M. C. Wooten, and M. J. Harvey. 1979. Thompson's

pygmy shrew, Microsorex hoyi winnemmana, in Tennessee. Journal

of the Tennessee Academy of Science 54:14.

96 J. Laerm, W. M. Ford, and D. C. Weinand

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survey of North Carolina mammals. Occasional Papers of the North

Carolina Biological Survey. North Carolina State Museum of Natural

Sciences, Raleigh.

Mengah, M. T., D. C. Guynn, Jr., J. K. Edwards, D. L. Sanders, and

S. M. Miller. 1987. Abundance and distribution of shrews in

western South Carolina. Brimelyana 13:63-66.

Mitchell, J. C, S. Y. Erdle, and J. F. Pagels. 1993. Evaluation of

capture techniques for amphibian, reptile and small mammal com-

munities in saturated forested wetlands. Wetlands 13:130-136.

Padgett, T. M., and R. K. Rose. The pygmy shrew, Sorex hoyi winne-

mana (Insectivora: Soricidae), from the Coastal Plain of North Carolina.

Brimleyana 21:87-90.

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Virginia's rarest shrews (Mammalia: Soricidae). Virginia Journal of

Science 38:364-368.

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for the pygmy shrew (Sorex hoyi winnemana) in Tennessee. Journal

of the Tennessee Academy of Science 64:240.

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of the North Carolina Biological Survey, Raleigh.

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malogy 68(2), supplement: 1-76.

Received 24 February 1994

Accepted 3 May 1994

Food and Ectoparasites of the Southern Short-tailed

Shrew, Blarina carolinensis (Mammalia: Soricidae),

from South Carolina

John O. Whitaker, Jr.

Department of Life Sciences, Indiana State University,

Terra Haute, Indiana 47809

Gregory D. Hartman1

Museum of Southwestern Biology, Department of Biology,

University of New Mexico, Albuquerque, New Mexico 87131 and

Savannah River Ecology Laboratory,

Drawer E, Aiken, South Carolina 298022

AND

Randy Hein

Department of Life Sciences, Indiana State University,

Terra Haute, Indiana 47809

ABSTRACT — Food habitats and ectoparasites were examined in a

sample of 50 individuals of Blarina carolinensis collected in a

hardwood forest on the Coastal Plain of western South Carolina.

Both in terms of volume and frequency of occurrence, predominant

foods were slugs and snails (Mollusca), the hypogeous fungus

Endogone, earthworms (Annelida), and beetle (Coleoptera) adults

and larvae. Ectoparasites observed on B. carolinensis included

one species of flea (Doratopsylla blarina), one species of beetle

(Leptinus americanus), and 25 species of mites, the most frequent

being Orycteroxenus soricis, Asiochirus blarina, Echinonyssus blarinae,

Haemogamasus liponyssoides, and several species of Pygmephorus.

A good deal of information exists on the foods and ectoparasites

of the northern short-tailed shrew, Blarina brevicauda (Say 1823);

however, this is not the case for the southern short-tailed shrew,

B. carolinensis (Bachman 1837). We are not aware of any detailed

information on the foods eaten by B. carolinensis, and know of only

five species of ectoparasites that have been reported: the laelapids

Androlaelaps fahrenholzi and Haemogamasus harperi by Hayes and

Guyton (1958), Eulaelaps stabularis by Hayes and Guyton (1958) and

Jameson (1947), and Myonyssus jamesoni by Ewing and Baker (1947),

1 Present address: Department of Biological and Environmental Sciences, McNeese

State University, Lake Charles, Louisiana 70609.

2 Address for reprint requests.

Brimleyana 21:97-105, December 1994 97

98 J. O. Whitaker, Jr.. G. D. Hartman, and R. Hein

and the myobiid Blarinobia simplex by Ewing (1938). The purpose of

this paper is to present data on the food habits and ectoparasites of B.

carolinensis from South Carolina, and to compare these with data

that have been reported for B. brevicauda.

MATERIALS AND METHODS

Shrews were collected over a 21-day period in May 1986 from

the Savannah River Ecology Laboratory's Mill Creek small mammal

trapping grid. The Mill Creek grid is located in a mixed hardwood

cove forest located on the United States Department of Energy's Savannah

River Site, near Aiken, South Carolina, and is on the western-most

Coastal Plain of the State; specifics of grid dimensions and habitat

structure have been described elsewhere (Gentry et al. 1968, 1971).

One Museum Special and one Victor mouse trap were set at each

station of the grid; traps were baited with peanut butter and checked

daily. Captured shrews were placed in individual plastic bags and

frozen for later examination.

Stomach contents of each animal were removed and then identified

under a dissecting microscope. The volume of each item in each

stomach was estimated visually. Data were compiled as percent frequencies

(percentage of shrews with each item) and percent volumes (average

percentage of each food) of each item observed in the entire sample.

We collected ectoparasites by examining the fur with a dissecting

microscope. When ectoparasites were observed to occur in relatively

small numbers, all individuals were collected; when the numbers of

ectoparasites were large, the numbers individuals of each species were

estimated, and samples were taken of each. Data on food habits and

ectoparasites were compared to similar data for B. brevicauda from

Indiana (Mumford and Whitaker 1982). Shrews from Indiana were

collected from a variety of habitats.

RESULTS

Foods

Forty-five of the 50 individuals of B. carolinensis examined

contained food, totaling 23 items (Table 1). The five dominant foods

were slugs and snails (18.5% of total volume), the hypogenous fungus

Endogone and related genera (16.3%), earthworms (14.8%), unidentified

adult beetles (9.6%), and unidentified beetle larvae (5.8%). Total volumes

of Coleoptera, Lepidoptera, and Diptera were 17.8, 6.0, and 6.7%,

respectively. No single food item clearly was dominant in the sample

of B. carolinensis; slugs and snails, Endogone, and earthworms were

Food and Ectoparasites of Shrews 99

Table 1. Food items observed in the stomachs of 45 short-tailed shrews

(Blarina carolinensis) from South Carolina and 125 B. brevicauda from

Indiana (Mumford and Whitaker 1982).

100 J. O. Whitaker, Jr., G. D. Hartman, and R. Hein

represented about equally and collectively comprised about half of

the food in the sample. Whereas no food item was dominant in the

sample of B. carolinensis, earthworms clearly were dominant (37.5%

volume) in B. brevicauda, followed by slugs and snails (8.5%), lepidopterous

larvae (8.2%), Gryllidae (6.2%), and Chilopoda (4.5%) (Table 1).

Ectoparasites

Ectoparasites were observed on all B. carolinensis examined,

and individuals of 27 different species were collected (Table 2): one

species of flea (Doratopsylla blarina), one species of beetle (Leptinus

americanus), and 25 species of mites from eight families (Acaridae,

Anoetidae, Cyrtolaelapidae, Laelapidae, Listrophoridae, Myobiidae,

Pygmephoridae, and Trombiculidae). Both in terms of the percentage

of hosts infested and the mean numbers observed per host, the most

frequently observed ectoparasites on B. carolinensis were Echinonyssus

blarinae, Haemogamasus liponyssoides (Laelapidae), Asiochirus

blarina (Listrophoridae), Orycteroxenus soricis (Acaridae), and

Protomyobia blarinae (Myobiidae).

Doratopsylla blarina, the only flea observed on B. carolinensis,

was the second most abundant of the six flea species observed on

B. brevicauda. The acarid mite Orycteroxenus soricis occurred on

both species of shrew, but two acarid hypopi, Xenoryctes latiporus

(only on B. carolinensis) and Dermacarus hypudaei (only on B. brevicauda),

also were present. Asiochirus blarina was the only listrophorid

collected; it occurred on both species of shrew. Five species of

laelapid mites were collected from B. carolinensis, as compared to

eight species on B. brevicauda. The laelapid species Haemogamasus

liponyssoides was one of the two most abundant ectoparasites on

both species of shrew. Echinonyssus blarinae was more abundant on

B. carolinensis, whereas Androlaelaps fahrenholzi was the most

abundant laelapid on B. brevicauda. Seven species of Pygmephorus

were collected from B. carolinensis, and 12 species from B. brevicauda.

Of the 11 species of pygmephorids observed on B. carolinensis,

four were in the genus Bakerdania, a genus that was not observed on

B. brevicauda. Three species of Bakerdania probably are undescribed.

Thirteen species of ectoparasite (1 flea, 1 beetle, 11 mite species)

were common to both species of Blarina. Fourteen species of ectoparasites

were observed only on B. carolinensis: these 14 consisted of one

laelapid, one acarid, eight species of pygmephorids, two cyrtolaelapids,

an anoetid, and a chigger (Trombiculidae). Of the 32 species reported

for B. brevicauda, 19 were found only on that host. The 19 consisted

Food and Ectoparasites of Shrews 101

on five species of fleas, four species of laelapid mites, one species

of acarid mite, and a species of Pygmephorus.

DISCUSSION

Both B. carolinensis and B. brevicauda eat a wide variety of

foods: 23 categories of food items in the South Carolina material

(n = 45), compared with 36 categories in the much larger sample of

B. brevicauda (n = 125) from Indiana. The lower percent volume of

earthworms observed for B. carolinensis likely represents the low numbers

of earthworms that are supported by the sandy soils of the Savannah

River Site, rather than a difference in dietary preference between

B. carolinensis and B. brevicauda.

The hypogeous mycorrhizal fungus Endogone was one of the

more heavily eaten foods in the shrews from South Carolina at 16.3%

of the volume, but formed only 3.6% of the total volume in the foods

of Indiana shrews. Endogone (including related genera, see Castellano

et al. 1989) often is important as a food of small mammals (Whitaker

1962, Williams and Finney 1964). The small mammal-fungal relationship

is an important component of many communities because small

mammals act as dispersal agents for mycorrhizal fungi (Maser et al.

1978).

Of the five ectoparasitic species that previously had been

reported from B. carolinensis, all were found during our study (marked

with asterisks, Table 2) except for Haemogamasus harperi. The flea

most frequently observed on B. brevicauda, Ctenopthalmus pseudagyrtes,

was not collected from B. carolinensis; however, C. pseudagyrtes does

occur on the Savannah River Site, and has been observed on eastern

moles, Scalopus aquaticus (G. D. Hartman, unpublished data). The

most abundant mites on Blarina tend to be the tiny Asiochirus Marina

(Listrophoridae) and hypopi of Orycteroxenus soricis (Acaridae); both

of these ectroparasites likely were more abundant than the data indicate.

The number of ectoparasite species observed on B. carolinensis

was less than for B. brevicauda, in part because of the smaller number

of B. carolinensis examined. However, in spite of the different sample

sizes, there were notable differences between the ectoparasite assemblages.

Of 27 species observed on B. carolinensis, 14 were found only on this

host, and of the 32 species reported for B. brevicauda, 19 were found

only on that host. Anoetids, cyrtolaelapids, trombiculids, and the genus

Bakerdania (Pygmephoridae) were observed only on B. carolinensis.

Although the species of Pgymephorus found on the two host species

were not similar, this is not too surprising because pygmephorid

mites that occur on mammals are not host specific.

102

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104 J. O. Whitaker, Jr., G. D. Hartman, and R. Hein

CONCLUSION

We report the first study of the food habits of B. carolinensis,

and we increased, by more than five-fold, the number of species of

ectoparasites known to occur on this shrew. However, the specimens

of B. carolinensis that we examined were collected from a single locality

in less than one month. Further studies on B. carolinensis from different

localities, habitats, and times of the year are needed to account for any

temporal or spatial variation in food habits and the occurrence of ectoparasitic

species, and to further elucidate the differences or similarities between

B. carolinensis and B. brevicauda in their food habits and ectoparasite

assemblages.

ACKNOWLEDGMENTS— We wish to thank T. M. Padgett and an

anonymous reviewer for comments and suggestions that improved the

manuscript. Financial support for portions of this research were pro-

vided by contract DE-AC09-76SROO-819 between the United States

Department of Energy and the University of Georgia.

LITERATURE CITED

Castellano, M. A., J. M. Trappe, Z. Maser, and C. Maser. 1989. Key

to spores of the genera of hypogeous fungi of north temperate

forests. Mad River Press Incorporated, Eureka, California.

Ewing, H. E. 1938. North American mites of the subfamily Myobiinae,

new subfamily (Arachnida). Proceedings of the Entomological

Society of Washington 40:180-197.

Ewing, H. E., and E. W. Baker. 1947. Myonyssus jamesoni, a new

liponyssid mite (Acarina: Laelaptidae) from Blarina brevicauda (Say).

Journal of Parasitology 33:376-379.

Gentry, J. B., F. B. Golley, and M. H. Smith. 1968. An evaluation

of the proposed International Biological Program census method for

estimating small mammal populations. Acta Theriologica 13:313-327.

Gentry, J. B., F. B. Golley, and M. H. Smith. 1971. Yearly fluctua-

tions in small mammal populations in a southeastern United States

hardwood forest. Acta Theriologica 15:179-190.

Hayes, K. L., and F. E. Guyton. 1958. Parasitic mites (Acarina:

Mesostigmata) from Alabama mammals. Journal of Economic Ento-

mology 51:259-260.

Jameson, E. W., Jr. 1947. Natural history of the prairie vole (mam-

malian genus Microtus). University of Kansas Publications. Museum

of Natural History 1:125-151.

Maser, C, J. Trappe, and R. H. Nussbaum. 1978. Fungal-small

mammal interrelationships with emphasis on Oregon coniferous forests.

Ecology 59:799-809.

Food and Ectoparasites of Shrews 105

Mumford, R. E., and J. O. Whitaker, Jr. 1982. Mammals of Indiana.

Indiana University Press, Bloomington.

Whitaker, J. O., Jr. 1962. Endogone, Hymenogaster and Melanogaster

as small mammal foods. American Midland Naturalist 67:152-156.

Williams, O., and B. A. Finney. 1964. Endogone - food for mice.

Journal of Mammalogy 45:265-271.

Received 8 March 1994

Accepted 16 September 1994

106

Mensural Discrimination of Four Species of

Perotnyscus (Rodentia: Muridae) in the Southeastern

United States

Joshua Laerm

Museum of Natural History, University of Georgia, Athens,

Georgia 30602

AND

James L. Boone

Museum of Natural History, Institute of Ecology, and Savannah River

Ecology Laboratory, University of Georgia, Athens, Georgia 30602

ABSTRACT — We subjected 17 mensural characters from a total

of 460 cotton mice (Peromyscus gossypinus), white-footed mice

(P. leucopus), deer mice (P. maniculatus), and old-field mice

(P. polionotus) to discriminant analysis to maximally distinguish

among specimens of these species in the southeastern United

States. If external measurements are available, 13 characters are

necessary to correctly classify all specimens. If external measurements

are not available, 14 cranial characters discriminate at most 91%

of the specimens. In pairwise comparisons using external and

skull measurements, at least 98% of specimens can be separated

with one or two characters. In pairwise comparisons (except P.

leucopus-P. maniculatus) using only skull measurements, at least

95% of specimens can be correctly identified to species with

one or two characters. For P. leucopus and P. maniculatus, six

characters correctly separate 86% of the specimens, and two characters

separate 82%.

White-footed mice {Peromyscus, Golger) are among the most

widely distributed and ubiquitous North American mammals (Hall 1981),

are the most broadly studied native mammals (King 1968), and are

represented extensively in systematic collections. Despite their

commonness and familiarity to most biologists, it is still difficult to

distinguish among species when we use morphological characters

(Hooper 1968). Much literature has resulted from regional attempts to

provide for mensural discrimination among Peromyscus, especially

between and within Osgood's (1909) maniculatus and leucopus species-

groups. Papers have been published separating the white-footed mouse

(P. leucopus [Rafinesque]), from the deer mouse (P. maniculatus [Wagner])

in New England (Choate 1973), Kansas (Choate et al. 1979), Wisconsin

(Stromberg 1979), and Maryland (Feldhamer et al. 1983); separating

the white-footed mouse from the cotton mouse (P. gossypinus [Le Conte]),

in Alabama (Linzey et al. 1976) and eastern Texas (Engstrom et al.

Brimleyana 21:107-123, December 1994 1.07

108

Joshua Laerm and James L. Boone

1982); separating five Peromyscus species in New Mexico (Smart 1978);

and separating four Peromyscus species in Arkansas (McDaniel et al.

1983). These studies indicate that it is usually possible to distinguish

between morphologically similar species, but the characters necessary

to do so vary geographically. Thus, for example, the characters used to

distinguish between P. leucopus and P. maniculatus in New England

differ from those in Wisconsin or Kansas. This almost ad hoc approach

Fig. 1. Southeastern distribution of the four Peromyscus species showing

collection location of the specimens used to build the model (•) and

specimens used to test the model (*).

Mensural Discrimination of Peromyscus 109

to the problem has been necessary because several of the species,

particularly P. leucopus and P. maniculatus, have a high degree of

intraspecific variation in morphology.

In the southeastern United States the ranges of four species overlap

(Fig. 1). It is difficult to correctly identify these species using available

taxonomic keys (e.g., Golley 1962, 1966; Blair et al. 1968; Hall 1981)

based only on pelage features and/or cranial measurements. The four

species usually can be distinguished based on collection location,

habitat, and morphological data. Populations of Peromyscus maniculatus

in this region are referred to as P. m. nubiterrae and are typically

found in mesic forests at elevations higher than 900 m, and P. maniculatus

usually has a sharply bicolored tail that is longer than the head and

body. Peromyscus gossypinus is generally found in hardwood river

bottoms and coastal oak-palmetto (Quercus sp. and Serenoa repens)

forests and is the largest and heaviest of the four species. Peromyscus

polionotus is generally found in areas of sandy soil and has a very

short, distinctly bicolored tail. Peromyscus leucopus leucopus is generally

found at elevations below 900 m in relatively xeric woodlands. Its

tail is shorter than the head and body, and it is smaller and lighter in

mass than P. gossypinus. A plot of principal component scores generated

from the correlation structure of three standard external measurements

(body, tail, and hind foot lengths) illustrates the overlap in measurements

from specimens collected in the Southeast and graphically illustrates

the difficulty in separating these four species based on these features

(Fig. 2).

For museum personnel that acquire poorly curated public or private

collections, or who desire to reexamine their holdings, identification

of specimens from regions where ranges overlap may be difficult.

The objective of this study is to examine the effectiveness of statistical

procedures to distinguish these species in the southeastern United

States without the use of collection-location information and without

using statistically unsound ratios (Humphries et al. 1981). To do this,

we generate discriminant functions from both external and skull measurements

and from skull measurements alone.

METHODS

We used univariate and multivariate statistics to examine 460

Peromyscus museum specimens collected in the southeastern United

States for variation in 17 morphometric characters. We selected sample

sites based on the availability of large numbers of adult specimens

from throughout the region. Sample sites were selected to reduce

potential for incorrect a priori species identification by eliminating,

to some degree, consideration of localities where ranges overlap. These

criteria resulted in the distribution of sample sites in Figure 1.

110

Joshua Laerm and James L. Boone

P. maniculatus

Axis 1

P. polionotus

P. gossypinus

P. leucopus

Axis 2

Fig. 2. Distribution of principal component scores generated from external

measurements (body, tail, and hind foot lengths) illustrating overlap in

the measurements of these characters.

A priori identifications were based on specimen tag information.

We used only specimens we believed were correctly identified. We

wanted to create a robust generalized model, but we also wanted to

build the model based on, as much as possible, animals that we felt

were correctly identified. The selection procedure resulted in using

110 P. gossypinus, 108 P. leucopus, 112 P. maniculatus, and 110 P.

polionotus. The Appendix lists specimens examined. We used five

additional specimens of each species, generally selected from

locations not included in the model building process, to test the model.

One of us (JL) measured 14 cranial characters to the nearest 0.1

mm with dial calipers and recorded three external measurements from

specimen tags. We estimated age from pelage characters (no juvenile

gray), tooth wear (significant wear on all major cusps), and degree of

cranial suture fusion. We measured only adults (in age classes 4-6 of

Schmidly 1973) and excluded specimens with missing data from all

analyses. Mensural characters (Choate et al. 1973, DeBlase and Martin

Mensural Discrimination of Peromyscus 111

1981) included: head and body length (body), tail length (tail), hind

foot length (foot), greatest skull length (SL), basonasal length (BNL),

rostral breadth (RB), nasal length (NL), interorbital constriction (OC),

zygomatic breadth (ZB), bony palate length (PL), maxillary toothrow

length (MTL), total toothrow length (TTL), palatal width (PW), pterygoid

breadth (PB), bullar depth (BD), and anterior palatal (incisive) foramen

length (PFL). We measured rostral length (RL) from the anteriormost

point of the nasals to the anterior edge of the zygomatic arch. Body

length was calculated as the difference between total and tail lengths.

We excluded ear length due to predominance of missing data.

We performed statistical analyses with Systat 5.1a (Wilkenson

1989) and SPSS 4.01 (Norusis 1990). We tested normality and

homogeneity of variance by inspecting plotted residuals and by

Bartlett's test for homogeneity of group variances, respectively.

Differences among adult age classes and between sexes were tested

with analysis of variance, and type I error rates were corrected with

the Bonferroni adjustment (Rice 1989). We classified taxa using stepwise

discriminant analysis. Variables were included in the models based on

minimizing residual variance, prior probabilities were equal to sample

size, and varimax rotation was employed. Stepwise discriminant analysis

will find an optimal solution based on the data; however, depending on

where the analysis begins (i.e., which variables enter the model first),

it may find a local, rather than the global, optimum. To help avoid this

optimization problem, we removed variables that entered the model in

the first steps and repeated the analysis. In one case, that of discrimination

based on all external and skull measurements, we found that bullar

depth (BD) forced the model onto a local optimum. Therefore, we

eliminated this character from further consideration in that model. We

used stepwise discriminant analysis to produce two main predictive

functions from the smallest set of characters needed to separate all

four species — one for external and skull measurements and another for

skull measurements alone. In addition, we generated predictive functions

that used only one or two measurements to separate in pairwise comparisons

among species.

We performed all analyses on raw data without transformation

(because transformation did not result in homogeneous variances)

and without removing size (Burnaby 1966, Rohlf and Bookstein 1987)

because this produced the simplest tool for the identification of

questionable specimens in the future. Although there was significant

heterogeneity of variances among species for some characters, standard

transformations (e.g., logarithm, etc.) did not homogenize it, and raw

data were more effective in discrimination than log-transformed data.

112

Joshua Laerm and James L. Boone

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Mensural Discrimination of Peromyscus 113

RESULTS

In univariate tests, we found significant differences between the

sexes in P. gossypinus for body length (P<0.02), RL (P<0.02), and

NL (P<0.05); in P. leucopus for foot (P<0.()3) and PFL (P<0.04); in

P. maniciilatus for SL (P<0.05), PB (P<0.04), and PFL (P<0.01);

and in P. polionotus for body length (P<0.01). Although these differences

were individually significant, there was considerable overlap in character

ranges, and none was significant when we applied the Bonferroni correction

(table-wide significance began at P< 0.003). The differences between

the sexes of P. maniciilatus approached significance (P<0.07), but

none was significantly different (P<0.05) when subjected to two group

(i.e., male vs. female) discriminant analysis. We included gender in

the discriminant analysis of all characters, but its effect was not significant,

and it did not enter the final stepwise model. Table 1 contains means,

ranges, and standard errors for all characters.

Univariate analyses were marginally successful in identifying the

four species, but no single measurement unambiguously separated them.

Most characters separated the large P. gossypinus from the small P.

polionotus, but six of 17 characters showed overlapping distributions.

Tail length greater or less than 55 mm is the simplest method to separate

these two species. No single character could separate P. gossypinus

from P. leucopus or P. maniculatus, but anterior palatal foramen length

5.4 mm identified most (67%) P. gossypinus. Tail length 83 mm separated

81% of P. maniciilatus from the other three species, but four P. gossypinus

had tails longer than 83 mm. There was no overlap in the tail lengths

of P. polionotus and P. maniciilatus. No single character separated P.

leucopus from P. maniculatus.

Multivariate analyses using external and skull measurements were

successful in identifying the four species. Stepwise discriminant analysis

correctly classified all specimens using measurements of 13 characters

(in order of inclusion into model: Tail, SL, MTL, Foot, RL, OC, PFL,

Body, TTL, PB, BNL, PL, PW). The three axes accounted for 55.51,

37.16, and 7.34% of the variance (Fig. 3a). After a varimax rotation,

the variables most highly correlated with the first discriminant function

were TTL (0.87), SL (0.85), BNL (0.74), PL (0.69), RL (0.58), PFL

(0.58), ZB (0.57), MTL (0.53), and NL (0.52); those highly correlated

with the second function were BD (0.82), PB (0.40), and OC (0.37);

and those highly correlated with the third function were PFL (0.49),

RL (0.45), PB (0.23), and OC (-0.21).

Discriminant analysis using only skull measurements correctly

classified at most 90% of the specimens with 10 characters (in order

of inclusion into model: SL, BD, MTL, RL, PFL, OC, TTL, BNL, PB,

114

Joshua Laerm and James L. Boone

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PW). After a varimax rotation, the variables most highly correlated

with the first discriminant function were SL (0.74), TTL (0.74), BNL

(0.66), PFL (0.66), RL (0.65), PL (0.62), Foot (0.59), MTL (0.53),

ZB (0.53), and NL (0.50); the only variable highly correlated with the

second function was Tail (0.82, all others were less than ±0.17); at

-0.33, OC was most highly correlated with the third function. All the

misclassifications of the data were in separating P. leucopus and P.

maniculatus (Fig. 3b). This observation led us to implement a two-

step discrimination process as suggested by Thompson and Conley

(1983). First, we grouped P. leucopus and P. maniculatus and performed

discriminant analysis among P. gossypinus, P. polionotus, and P. leucopus-

P. maniculatus; then we separated P. leucopus and P. maniculatus.

However, this scheme did not improve the classification results.

In analysis of species pairs, at least 98% of specimens could be

separated using only one or two external and/or skull measurements

(Table 2). In pairwise comparisons using only skull measurements,

we could separate at least 95% of the specimens (except for P. leucopus-

P. maniculatus). For this species pair two characters separate 82% of

the specimens. The scores generated by the discriminant functions

(Table 2) approximately fall on either side of zero, such that scores

for one species are positive, and scores for the other species are

negative. However, these models do generate a few misclassifications;

therefore, specimens with scores near zero (e.g., ±0.5) should be subjected

to the full discriminant models.

DISCUSSION

Discrimination of these Peromyscus species is difficult when

collection location information or skins are missing, and we did not

achieve the ultimate goal of this project which was to correctly classify

any skull without external information. However, the great majority

of specimens can be correctly assigned to species, and the discriminant

function was useful in identifying likely misclassified and questionable

specimens in our museum collections. Additionally, the function allows

evaluation of specimens collected at the periphery of species' ranges.

The model using external and skull characters was reasonably

successful in classifying the test specimens, which suggests that we

captured enough of the variation within each species to make it useful

in classifying specimens from somewhat beyond the geographic distribution

of our samples. This is an improvement over the ad hoc approach

where each state or region requires a different discrimination model.

However, although the P. maniculatus test specimens classified correctly,

they tended to fall in the margins of the discriminant score distributions.

The model with only skull measurements was less successful in classifying

116

Joshua Laerm and James L. Boone

P. maniculatus

Axis 2

P. polibnotus R leuc°Pus

P. gossypinus

Axis 2

P. polionotus

maniculatus

Fig. 3. Distribution of discriminant scores generated from (a) external

and skull measurements and from (b) skull measurements alone plotted

on the first two canonical axes. Letters (the first letter of the specific

epithet for each species) designate the location of test specimens, letters

in parentheses mark misclassifications, and crosses mark group centroids.

Mensural Discrimination of Peromyscus 117

the test specimens, and results should be viewed with caution if that

model is used for specimens collected far outside the geographic distribution

of our samples.

Our results were similar to those of previous authors who found

that these species tend to differ significantly in most measurements,

but that there is generally some overlap in measurement that prevents

classification of some specimens based on single characters. For example,

Linzey et al. (1976) could separate most specimens using anterior

palatal foramen length and width or skull length. Choate (1973) could

separate most specimens with tail length. Engstrom et al. (1982) found

that P. gossypinus differed significantly from P. leucopus in every

character they measured, but that there was overlap in all characters.

McCarley (1954) found that hindfoot length separated most P. leucopus

from P. gossypinus.

Stromberg (1979) successfully used discriminant analysis on external

characters to separate P. maniculatus from P. leucopus. We found

that these characters could not be used in the extreme Southeast (Fig.

2). However, he found that ear length was especially useful, and we

were not able to include that character. We disagree with Stromberg's

(1979) statement that discrimination of external characters offers a

dependable alternative to cranial measurements in the identification

of P. maniculatus and P. leucopus. As in our study, McDaniel et al.

(1983) and Choate et al. (1979) were able to separate almost all of

their specimens using cranial measurements. Only Engstrom et al.

(1982) was able to separate all of their specimens using cranial measurements.

Choate (1973), Choate et al. (1978), and Engstrom et al. (1982)

found that variation among adult age classes was required in the

models for accurate classification. In contrast, we did not find that

age variation among adult age classes (4-6, Schmidly 1973) was significant.

We found statistical differences among age classes 4-6, but these

differences were small relative to the differences among species, and

thus age information was not important in our models.

Several authors have found ratios useful in identifying Peromyscus

species pairs. McDaniel et al. (1983) found that the ratio of interorbital

width to length of the nasal bone was useful in separating P. attwateri

from P. gossypinus. Feldhamer et al. (1983) found that the ratio of

tail length to body length in conjunction with body mass separated P.

leucopus from P. maniculatus (pregnant females excluded). McCarley

(1954) used the ratio of skull length to foot length to identify P.

gossypinus, P. leucopus, and their purported hybrids. Although ratios

may provide useful indices, we agree with Humphries et al. (1981)

and the references they provide that ratios should be avoided in morphometric

118 Joshua Laerm and James L. Boone

studies because of statistical and conceptual difficulties. Discriminant

analysis based on two characters has a similar result of separating

groups based on the magnitude of two measurements. It also has the

benefits of potentially better separation of groups by stretching the

axes (weighing measurements with discrimination function coefficients)

and an associated probability of group membership. Therefore, we

have presented results (Table 2) that use one or two measurements

rather than ratios to separate pairs of species with discriminant functions.

We agree with Choate (1973) that habitat and external features

(e.g., tail coloration, penciled tail, color, and degree of fur luxuriance)

can yield important information for classifying these species. For

example, we believe that the best ways to identify P. polionotus are

that it is found on sandy soils and by its short, strongly bicolored tail,

and the best ways to identify P. leucopus are that it is found in low

elevation exeric sites and that it has more reddish-orange on the sides

than P. gossypinus. Other qualitative characters may also be useful.

For example, Linzey et al. (1976) found that the skulls of P. leucopus

tend to be lighter and more fragile than those of P. gossypinus. However,

our goal was to identify these species with quantitative characters

rather than qualitative characters, and preferably with the skull alone,

as noted by Feldhamer et al. (1983), these qualitative characteristics

can be variable within species. Most of the classification problems

we encountered involved old skulls without associated skins.

Use of the discriminant function — Discriminant analysis combines

variables to generate a set of linear, independent axes upon which

specimens, after appropriate scoring, can be plotted and their classification

determined. The appropriate scoring method is to multiply each morphological

character variable (e.g., foot length, skull length) by its discriminant

function coefficient, sum the products, and add a constant (for each

axis separately). In general:

D, = Bw + BuX^ + £,2X2 + #13*3+ ... + BlnXn

Di = £20 + B2^X^ + £22*2 + Bz>X> + ... + BlnXn

where D\ is the specimen's discriminant score on the first axis, the

Bus are discriminant function coefficients estimated from the data

for the first axis (Bins are constants), and the Xt's are the values of

the original variables. This is done separately for each axis, and the

scores, D\, D2, ..., D«, form the coordinate of the specimen's location

in the ^-dimensional discriminant space. For example, to separate P.

gossypinus from P. leucopus using external and skull measurements,

the appropriate transformation is (only one axis is needed)

D = -34.125 + 0.593(hindfoot length) + 0.821(skull length).

Mensural Discrimination of Peromyscus 119

Table 3. Unstandardized canonical discriminant function coefficients,

external and skull characters, and skull characters only for four Peromyscus

species in the southeastern United States.

Table 4. Group centroids for external and skull characters and for

skull characters only based on unstandardized canonical discriminant

function coefficients of four Peromyscus species in the southeastern

United States.

120 Joshua Laerm and James L. Boone

Given an unknown specimen with hindfoot and skull lengths of 23.5

and 28.7 mm, respectively, and the coefficients of these measurements

from Table 3, this equation becomes:

D = -34.125 + 0.593(23.5) + 0.821(28.7)

D = 3.377

In this case, any positive value of D indicates P. gossypinus, and any

negative value of D indicates a P. leucopus (Table 2). Thus, this

specimen is a P. gossypinus.

If these two species required more than one axis, D\ and Di

would be calculated using discriminant coefficients from Table 3 for

external and cranial measurements or Table 4 for skull measurements

only. The bivariate coordinate (Di, Di) can be plotted on a 2-dimensional

graph (e.g., Fig. 3).

ACKNOWLEDGMENTS — We appreciate the assistance extended

by curatorial staffs of the following museums: Carnegie Museum of

Natural History (CMNH), Florida Museum of Natural History (UF),

Louisiana State University Museum of Zoology (LSUMZ), Harvard

University Museum of Comparative Zoology (MCZ), University of

Alabama Museum of Natural History (UAL), University of Georgia

Museum of Natural History (UGAMNH), and the United States National

Museum (USNM). Two anonymous reviewers offered helpful suggestions

on an early draft of the manuscript.

LITERATURE CITED

Blair, W. F., A. P. Blair, P. Brodkorb, F. R. Cagle, and G. A. Moore.

1968. Vertebrates of the United States. Second edition. McGraw

Hill, New York, New York.

Burnaby, T. P. 1966. Growth invariant discriminant functions and

generalized distances. Biometrics 22:96-110.

Choate, J. R. 1973. Identification and recent distribution of white-

footed mice (Peromyscus) in New England. Journal of Mammal-

ogy 54:41-49.

Choate, J. R., R. C. Dowler, and J. E. Krause. 1979. Mensural

discrimination between Peromyscus leucopus and P. maniculatus

(Rodentia) in Kansas. Southwestern Naturalist 24:249-258.

DeBlase, A. F., and R. E. Martin. 1981. A manual of mammalogy

with keys of the families of the world. Second edition. Wm.

Brown Company, Dubuque, Iowa.

Engstrom, M. D., D. J. Schmidly, and P. K. Fox. 1982. Nongeographic

variation and discrimination of species within the Peromyscus leucopus

species group (Mammalia: Cricetinae) in eastern Texas. Texas Journal

of Science 34:149-162.

Mensural Discrimination of Peromyscus 121

Feldhamer, G. A., J. E. Gates, and J. H. Howard. 1983. Field

identification of Peromyscus maniculatus and P. leucopus in Maryland:

Reliability of morphological characters. Acta Theriologica 28:417-

423.

Golley, F. B. 1962. Mammals of Georgia: A study of their distribu-

tion and functional role in the ecosystem. University of Georgia

Press, Athens.

Golley, F. B. 1966. South Carolina mammals. The Charleston Mu-

seum, Charleston, South Carolina.

Hall, E. R. 1981. The mammals of North America. Second edition.

John Wiley and Sons, New York, New York.

Hooper, E. T. 1968. Classification. Pages 27-74 in (J. A. King,

editor). Biology of Peromyscus (Rodentia). Special Publication of

the American Society of Mammalogists 2:1-593.

Humphries, J. M., F. L. Bookstein, B. Chernoff, G. R. Smith, R. L.

Elder, and S. G. Poss. 1981. Multivariate discrimination by

shape in relation to size. Systematic Zoology 30:291-308.

King, J. A. (editor) 1968. Biology of Peromyscus (Rodentia). Spe-

cial Publication of the American Society of Mammalogists 2:1-

593.

Linzey, A. V., D. W. Linzey, and S. E. Perkins, Jr. 1976. The

Peromyscus leucopus group in Alabama. Journal of the Alabama

Academy of Science 47:109-113.

McCarley, W. H. 1954. Natural hybridization in the Peromyscus

leucopus species group of mice. Evolution 8:314-323.

McDaniel, V. R., R. Tumlison, and P. McLarty. 1983. Mensural

discrimination of the skulls of Arkansas Peromyscus. Proceedings

of the Arkansas Academy of Science 37:50-53.

Norusis, N. J. 1990. SPSS base system user's guide. SPSS Incorpo-

rated, Chicago, Illinois.

Osgood, W. H. 1909. Revision of the mice of the American genus

Peromyscus. North American Fauna 82:1-285.

Rice, W. R. 1989. Analyzing tables of statistical tests. Evolution

43:223-225.

Rohlf, F. J., and F. L. Bookstein. 1987. A comment on shearing as

a method for "size correction." Systematic Zoology 36:356-367.

Schmidly, D. J. 1973. Geographic variation and taxonomy of Peromyscus

boylii from Mexico and the southern United States. Journal of

Mammalogy 54:111-130.

Smart, R. A. 1978. A comparison of ecological and morphological

overlap in a Peromyscus community. Ecology 59:216-220.

Stromberg, M. R. 1979. Field identification of Peromyscus leucopus

and P. maniculatus with discriminant analysis. Wisconsin Academy

of Science, Art and Letters 67:159-164.

122 Joshua Laerm and James L. Boone

Thompson, T. G., and W. Conley. 1983. Discrimination of coexist-

ing species of Peromyscus in south-central New Mexico. South-

western Naturalist 28:199-209.

Wilkenson, L. 1989. SYSTAT: The system for statistics. SYSTAT,

Incorporated, Evanston, Illinois.

Received 12 July 1993

Accepted 2 September 1993

APPENDIX

Specimens examined to build the model.

Museum acronyms are defined in the acknowledgments. Location

names are states and counties.

P. gossypinus — ALABAMA: Jackson; 9 USNM). Tuscaloosa; 7

(UAL). FLORIDA: Alachua; 13 (UF). GEORGIA: Burke; 12 (MCZ),

2 (USNM). Camden; 13 (UGAMNH). Charlton; 22 (UGAMNH). Ware;

8 (UGAMNH). NORTH CAROLINA: Gates; 8 (LSUMZ). SOUTH

CAROLINA: Charleston; 16 (CMNH).

P. leucopus— ALABAMA: Colbert; 1 (USNM). Jackson; 4 (USNM).

GEORGIA: Barrow; 1 (UGAMNH). Clarke; 40 (UGAMNH). Dekalb;

1 (UGAMNH). Elbert; 1 (UGAMNH). Oconee; 1 (UGAMNH). Rockdale;

1 (UGAMNH). Walton; 1 (UGAMNH). Wilkes: 1 (UGAMNH). NORTH

CAROLINA: Anson; 3 (USNM). Jackson; 8 (USNM). Macon: 2 (UGAMNH).

Wake; 7 (USNM). SOUTH CAROLINA: Abbeville; 1 (USNM). Greenville;

2 (USNM). Oconee; 7 (USNM). Pickens; 4 (USNM).

P. maniculatus— GEORGIA: Rabun; 9 (UGAMNH). Towns; 9

(UGAMNH). Union; 27 (UGAMNH). KENTUCKY: Bell; 7 (USNM).

Harlan; 12 (USNM). NORTH CAROLINA: Macon; 21 (UGAMNH).

TENNESSEE: Carter, 9 (USNM). Johnson; 2 (USNM). Sevier; 16

(USNM).

P. polionotus— ALABAMA: Autauga; 7 (USNM). Henry; 7 USNM).

Marshall; 1 (USNM). FLORIDA: Indian River, 5 (UGAMNH). Marion;

5 (UGAMNH). GEORGIA: Baker, 2 (UGAMNH). Barrow; 2 (UGAMNH).

Burke; 1 (UGAMNH). Clarke; 10 (UGAMNH). Decatur; 3 (UF). Dougherty;

2 (USNM). Gordon; 3 (USNM). Haralson; 3 (UGAMNH). Irwin; 2

(UF). Johnson; 2 (UGAMNH). Lowndes; 2 (UGAMNH). Marion; 1

(UF). Mcintosh; 3 (UGAMNH). Randolph; 10 (UGAMNH). Richmond;

2 (UGAMNH). Seminole; 2 (UF). Taylor; 1 (USNM). Tift; 13 (USNM).

SOUTH CAROLINA: Aiken; 12 (UGAMNH). Barnwell; 9 (UGAMNH).

Mensural Discrimination of Peromyscus 123

Specimens examined to test the model.

P. gossypinus— ALABAMA: Dekalb; 2 (UI). GEORGIA: Dougherty;

2 (UI). SOUTH CAROLINA: Aiken; 1 (UGAMNH).

P. leucopus— KENTUCKY: Bell; 1 (UGAMNH). NORTH CAROLINA:

Gates; 2 (UGAMNH). McDowell; 2 (UGAMNH).

P. maniculatus— GEORGIA: Fannin; 2 (UGAMNH). NORTH CAROLINA:

Watauga; 1 (USNM). TENNESSEE: Carter; 1 (UGAMNH). VIRGINIA

Giles; 1 (UGAMNH).

P. polionotus— ALABAMA: Marshall; 1 (USNM). FLORIDA:

Highlands; 2 (UGAMNH). Marion; 1 (UGAMNH). SOUTH CAROLINA:

Barnwell; 1 (UGAMNH).

124

Small Mammal Communities

in Streamside Management Zones

Dagmar P. Thurmond1 and Karl V. Miller2

Daniel B. Warnell School of Forest Resources

The University of Georgia, Athens, Georgia 30602

ABSTRACT — Populations of small mammals were sampled in six

streamside management zones (SMZs) of three widths: narrow

(15 m), medium (30 m), and wide (50 m), which extended through

a three-year-old pine plantation. We also sampled the pine plantation

and a nearby mature riparian forest. Two hundred and twenty-

eight small mammals from 12 species were captured in 8,640

trapnights. Overall, capture rates were not related to SMZ width.

During summer, capture rates were greater in the mature riparian

forest than in SMZs. Abundance of individual species varied among

the habitats sampled. SMZs supported populations of Oryzomys

palustris, Ochrotomys nuttalli, and Neotoma floridana, three species

not found in the pine plantation. Inclusion of SMZs in pine

plantation management can enhance habitat diversity and contribute

to local diversity of the small mammal community.

Approximately 8.5 million hectares in the southern United

States is maintained in pine plantations (United States Department of

Agriculture, Forest Service 1988), much of which is managed on short

rotations. Although young pine plantations provide seasonal habitat

needs for several mammalian species including white-tailed deer

(Odocoileus virginianus), eastern cottontails (Sylvilagus floridanus),

and oldfield mice (Peromyscus polionotus), other later-successional

species may be low in abundance or absent.

Streamside management zones are designed to protect water

quality from potential impacts of silvicultural operations. SMZs also

add habitat diversity to the surrounding pine plantations. Additionally,

SMZs create an area of edge, which increases the number of niches

available to wildlife.

Squirrel (Sciurus spp.) use of SMZs is greater than in adjacent

upland pine-hardwood areas in Mississippi (Warren and Hurst 1980)

and Alabama (Fischer and Holler 1991), and greater than in adjacent

pine plantations in Texas (McElfresh et al. 1980). Studies in eastern

Texas indicated that squirrels were more abundant in wide SMZs

(>55 m) than in narrow SMZs (<25 m). Conversely, small mammals

were more abundant in the narrow SMZs (Dickson and Huntley 1987,

1 Present address: Shoal Creek Ranger District, Talladega National Forest, Heflin.

Alabama 36264.

: Reprint requests.

Brimleyana 21:125-130. December 1994 125

126 Daemar P. Thurmond and Karl V. Miller

Dickson and Williamson 1988). Nevertheless, the relationships between

SMZ width and small mammal communities have not been investigated

adequately. We censused the small mammal communities in SMZs of

varying width, in adjacent pine plantations, and in mature riparian

areas.

MATERIALS AND METHODS

Study areas were located in the Upper Coastal Plain of Georgia

on the Ogeechee River drainage in Jefferson and Emanuel counties.

All SMZs were along first order streams in a 450-ha pine plantation

owned by Federal Paper Board Company. The stand was clearcut in

1985, the site prepared chemically, and planted in a 2-m X 3-m spacing

to loblolly pine (Pinus taeda) in 1987. All SMZs were selectively

harvested. Remaining overstory in the SMZs was dominated by

blackgum (Nyssa sylvatica), tulip poplar (Liriodendron tulipifera),

red maple (Acer rubrum), sweetgum (Liquidambar styraciflua), and

loblolly pine. Understory composition in the SMZs was dominated by

blackberry (Rubus spp.), greenbriar (Smilax spp.), poison ivy {Toxicodendron

radicans), gallberry (Ilex glabra), and fetterbush (Lyonia lucida).

Three width categories of SMZs were compared using two

replicates of each: narrow (15-18 m), medium (28-30 m), and wide

(49-53 m). Additional plots were established along two creeks in

mature, riparian forests on Old Town Plantation near Louisville, Georgia.

Streams in these forests averaged 2 m in width and were at least 100

m away from any forest edge. Dominant overstory included loblolly

pine, cypress (Taxodium distichum), hickory (Carya spp.), oaks (Quercus

spp.), sweetgum, and red maple.

Populations of small mammals were sampled by removal trapping

along a 200-m transect in the center of each SMZ and along the

stream in the mature riparian area. An additional transect was established

just inside the outer edges of the medium and wide SMZs. Parallel

200-m transects were sampled in the adjoining pine plantations, 50 m

from the SMZ edge. Transect paths were lightly cleared for access.

Ten trapping stations were placed at 20-m intervals along each

transect.

Small mammal populations were censused during four consecutive

nights in December 1990, June 1991, January 1992, and June 1992.

Sampling did not occur on rainy days. One Victor™ mouse trap, a

Victor™ rat trap, and a pitfall trap were placed at each trapping

station. Snap traps were baited with a mixture of peanut butter and

peanut oil. Pitfall traps (10-cm diameter, and filled to a depth of 7 cm

with water) were used to increase trapping success for shrews. Shrews

Streamside Management Zones 127

are difficult to catch with conventional snap or live traps (Szaro et al.

1988, Rose et al. 1989). Captured animals were donated to The University

of Georgia Museum of Natural History.

Captures were combined by season over the 2-year trapping

period and treated as replicates. Differences in mean capture rates

were tested by analysis of variance, and Duncan's Multiple Range

Test was used to separate means (a=0.05).

RESULTS AND DISCUSSION

We captured 228 small mammals from 12 species in 8,640

trapnights. Southern short-tailed shrews (Blarina carolinensis) accounted

for 24.5% of all captures. White-footed (Peromyscus leucopus) and

cotton mice (P. gossypinus) were grouped together as cotton mice,

because of the difficulty in positive identification. Morphological

criteria used to separate the species are of limited value when applied

to subadult mice (Dickson and Williamson 1988). Cotton mice accounted

for 20.2% of the animals caught, followed by cotton rats (Sigmodon

hispidus, 17.1%), old-field mice (Peromyscus polionotus, 13.6%), least

shrews (Cryptotis parva, 11.0%), rice rats (Oryzomys palustris, 7%),

and golden mice (Ochrotomys nuttalli, 3.1%). Other species captured

included the woodrat (Neotoma floridana), Eastern harvest mouse

(Reithrodontomys humulis), Southeastern shrew (Sorex longirostris),

pine vole (Microtus pinetorum), and Eastern mole (Scalopus aquaticus).

During winter sampling periods, small mammal capture rates

did not vary by treatment (P = 0.56). However, in summer total

capture rates were greater in the mature riparian forest than in the

other habitats sampled. Several species showed significant habitat

preferences (Table 1). During both winter and summer, cotton mice

were trapped more frequently in the mature riparian forest than in the

other habitats sampled. The preferred habitat for the cotton mouse is

bottomland hardwood forest subject to frequent flooding (Cothran

et al. 1991). Cotton mice were equally abundant in SMZs and pine

plantations.

In winter, oldfield mice were most common on the pine transects.

Several studies have reported the preference of oldfield mice for early

successional habitats (Golley et al. 1965, Brooks 1992). Oldfield mice,

harvest mice, and cotton rats prefer areas with stands of dense grass.

Cotton rats were caught most frequently in narrow SMZs in winter,

and no habitat preference was observed in summer. The rice rat was

not recorded in the pine plantations in either season. Southern short-

tailed shrews, which prefer moist habitats (Szaro et al. 1988), occurred

128

Dagmar P. Thurmond and Karl V. Miller

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Streamside Management Zones 129

in all habitats, although they tended to be caught most frequently

in the mature riparian forest.

In summer, mature riparian areas were dominated by the cotton

mouse, and, along with wide SMZs, were the only areas in which

golden mice and woodrats were found. These species often prefer

mature hardwood forest habitat, where they eat insects, twigs, green

leaves, berries, seeds, and nuts (Cothran et al. 1991). The wide SMZs

in our study provided some habitat for species associated with

mature stands, such as the golden mouse and the woodrat.

Our results suggest that the species composition of the small-

mammal community was affected by SMZ width. Only wide SMZs

(49-53 m) maintained populations of small mammal species that are

characteristic of mature riparian forests. Rice rats, golden mice, and

woodrats were captured in the SMZs, but not in the adjacent pine

plantations. Inclusion of SMZs in pine plantation management can

enhance habitat diversity and thereby contribute to local diversity of

the small mammal community.

ACKNOWLEDGMENTS— Funding for this study was provided by

the Georgia Forestry Commission, the National Council of the Paper

Industry for Air and Stream Improvement, Martha Black of Old Town

Plantation, and Mclntire-Stennis Project (GEO-0046-MS). Federal

Paper Board Company owned the SMZ treatment areas. B. R. Chapman,

A. S. Johnson, and R. J. Warren reviewed earlier drafts of this

manuscript and provided valuable suggestions.

LITERATURE CITED

Brooks, J. J. 1992. Chemical site preparation: effects on wildlife

habitat and small mammal populations in the Georgia Sandhills.

M.S. Thesis, The University of Georgia, Athens.

Cothran, E. G, M. H. Smith, J. O. Wolff, and J. B. Gentry. 1991.

Mammals of the Savannah River Site. Savannah River Site, National

Environmental Research Park, SRO-NERP-21.

Dickson, J. G, and J. C. Huntley. 1987. Riparian zones and wildlife

in southern forests: The problem and squirrel relationships. Pages

37-39 in Proceedings of managing southern forests for wildlife and

fish (J. G. Dickson and O. E. Maughan, editors). United States

Department of Agriculture, Forest Service, General Technical Re-

port SO-65.

130 Dagmar P. Thurmond and Karl V. Miller

Dickson, J. G., and J. H. Williamson. 1988. Small mammals in streamside

management zones in pine plantations. Pages 375-378 in Proceed-

ings of management of amphibians, reptiles, and small mammals in

North America (R. C. Szaro, K. E. Severson, and D. R. Paton,

technical coordinators). United States Department of Agriculture, Forest

Service, General Technical Report RM-166.

Fischer. R. A., and N. R. Holler. 1991. Habitat use and relative

abundance of gray squirrels in southern Alabama. The Journal of

Wildlife Management 55:52-59.

Golley, F. B., J. B. Gentry, L. D. Caldwell, and L. B. Davenport, Jr.

1965. Number and variety of small mammals on the AEC Savan-

nah River Plant. Journal of Mammalogy 46:1-18.

McElfresh, R. W., J. M. Inglis, and B. A. Brown. 1980. Gray squirrel

usage of hardwood ravines within pine plantations. Louisiana

State University Annual Forestry Symposium 19:79-89.

Rose, R. K., R. K. Everton, J. K. Stankavich, and J. W. Walke. 1989.

Small mammals in the Great Dismal Swamp of Virginia and North

Carolina. Brimleyana 16:87-101.

Szaro, R. C, L. H. Simons, and S. C. Belfit. 1988. Comparative

effectiveness of pitfalls and live-traps in measuring small mammal

community structure. Proceedings of management of amphibians, reptiles,

and small mammals in North Arizona. United States Department of

Agriculture, Forest Service, General Technical Report RM-166.

United States Department of Agriculture, Forest Service. 1988. The

South's fourth forest: alternatives for the future. Forestry Research

Report 24. Washington, D.C.

Warren, R. C, and G. A. Hurst. 1980. Squirrel densities in pine-

hardwood forests and streamside management zones. Proceedings of

the Annual Conference of the Southeastern Association of Fish and

Wildlife Agencies 34:492-498.

Received 30 May 1993

Accepted 31 August 1994

Home Range and Activity Patterns by Gray Foxes.

Urocyon cinereoargenteus (Carnivora: Canidae), in

East Tennessee

Cathryn H. Greenberg1

Ecology Program, Knoxville, Tennessee 37901-1071

AND

Michael R. Pelton

Department of Forestry, Wildlife and Fisheries,

Knoxville, Tennessee 37901-1071

ABSTRACT — We determined home-range size, spatial distribution,

and activity patterns of gray foxes (Urocyon cinereoargenteus)

(N = 10) between September 1986 and August 1987 in east

Tennessee. Average annual home-range size was 3.97 ± 1.51

(x ± SE) km:. There were no significant differences in home-

range size between sexes (females 3.67 ± 1.54; males 4.27 ±

1.59 km:) or age groups (adults 4.41 ± 1.46; subadults 3.20

± 1.62 km2). Home-range sizes were similar in three reproductive

seasons and in seasons of predominantly fruit (presumably abundant)

and predominantly flesh (presumably more scarce) diets. Home

ranges of adult male-female pairs and subadults coincided, suggesting

monogamy and exclusive area utilization by family groups. We

observed lower sunrise and/or daylight activity levels during breeding

and flesh diet seasons, and in months of low foliar cover.

The relationship of body size, metabolic needs, and dietary trophic

level to home-range size is well known. Relative to body size, flesh-

eaters have larger home ranges than plant-eaters, presumably due to

decreasing food base with ascending trophic level (McNab 1963).

However, most studies relating home-range size to trophic level are

based on interspecific comparisons.

Intraspecifically, seasonal shifts in home-range size appear to be

negatively correlated to food availability (Nicholson 1982). Sex, age,

and reproductive cycle (MacDonald 1980), population density (Trapp

and Hallberg 1975), inter- or intraspecific competition, and habitat

quality and dispersion (MacDonald 1980) are also thought to play a

role in affecting home-range size.

Most studies indicate that gray foxes are nocturnally active (Yearsley

and Samuel 1980, Nicholson 1982, Haroldson and Fritzell 1984). However,

1 Present address: United States Department of Agriculture Forest Service, South-

eastern Forest Experiment Station, P.O. Box 14524, Gainesville, Florida 32604.

Brimleyana 21:131-140, December 1994 131

132 Cathryn H. Greenberg and Michael R. Pelton

the amount of time required for foraging and temporal activity

patterns might be expected to change in relation to seasonal food

availability and/or type.

We obtained concurrent data on food habits, home range, and

activity patterns for gray foxes (Greenberg et al. 1988), which provided

an opportunity to compare seasonal shifts in dietary trophic level with

changes in home-range size. We predicted that home-range size would

become smaller as fruit became seasonally available (spring-fall) and

would expand in response to a presumably scarcer (predominantly

flesh) winter food supply. We also compared reproductive seasons,

sex, and age with home-range size.

METHODS

Study Area — The study area was located within the National

Environmental Research Park on the Department of Energy's Oak

Ridge Reservation, approximately 28 km west of Knoxville, Tennessee

(35°58' N, 84°56' W). Vegetation community types included pine and

pine-hardwood forests, loblolly pine (Pinus taeda) plantations, eastern

red cedar {Juniperus virginiana) barrens, oak-hickory forests, bottomland

hardwood forests, old fields, and developed areas. The Tennessee

Valley Authority's Melton Hill Reservoir and Watts Bar Lake border

the reservation on the west, south, and east; streams and springs

throughout the area provided water and wetland habitat.

The geology of the reservation is characteristic of the Southern

Appalachian Valley and Ridge Province. Parallel, southwest-northeast-

oriented ridges separated by valleys (elevation ranged from 226 to

413 m) lend additional diversity to the landscape.

Radio Telemetry — Foxes were captured in Number 1.5 Victor

soft-catch leg-hold traps with dirt-hole sets and drags. Attractants

included fox urine, fox gland lures, pork cracklings, fish oil, and

muskrat oil. Foxes were anesthetized with 5-10 mg/kg ketamine

hydrochloride (Ketaset) or not anesthetized (Nicholson 1982).

Animals were eartagged, fitted with radio collars equipped with a

mercury tip-switch activity sensor (Telonics, Mesa, Arizona) in the

150-151.84 frequency range, and released at the trap site. We classified

foxes as subadults (<l-year-old) or adults based on tooth wear

(Geir 1968).

There were 138 receiving stations established. Animals were

located with a four-element, hand-held, Yagi antenna and a portable

receiver. Locations and activities were recorded at 2-hour intervals

between the hours of 1600 and 0800 weekly. Occasional locations

were also recorded between 0800 and 1600. We used >2 compass

Gray Foxes 133

bearings with an intersecting angle >45° and <135°, and as close to

90° as possible (Heezen and Tester 1967) to plot locations. Activity

was recorded. Azimuths were converted to x:y coordinates by the

computer program Convxpoly (Boyle 1986), and the data were hand-

plotted on a 1:24,000 United States Geological Survey topographic

map with the Universal Transverse Mercator grid system.

We estimated home-range sizes by the minimum convex polygon

method (Mohr 1947). Atypical peripheral locations (known excursions)

were excluded based on subjective knowledge of typical home-range

use by the authors (Abies 1968).

Smith et al. (1981) found that three half-night radio-tracking periods

provided a larger estimate of coyote home ranges than 30 independent

daily locations, and that three or four nights provided good home-

range estimates for coyotes with small home ranges. We assumed that

their findings also applied to gray foxes. Hence, we considered >25

locations and at least three track-nights to be an adequate sample size

for home-range determinations.

Home ranges were calculated for three reproductive and two dietary

seasons. Three reproductive seasons included breeding (January-March),

pup-rearing (April-June), and nonbreeding (July-December) (Sullivan

1956, Nowak and Paradiso 1983). Dietary seasons included a dominantly

flesh diet (January-April) and dominantly insect or fruit diet (May-

December) (Greenberg and Pelton 1991). We compared annual and

seasonal home-range sizes between sexes and age groups. Due to small

sample sizes and high variance, we used descriptive statistics rather

than statistical tests in drawing our conclusions.

We calculated the percentage of "active" locations within four

time periods: two at sunrise (0.5 hours prior, 0.5 hours after sunset)

and two at night (0.5 hours after sunset, 0.5 hours prior to sunrise).

Data were pooled for all animals. We used Chi-square tests to detect

temporal differences in activity level, differences among repro-

ductive and dietary seasons, and differences between seasons of

low (November-April) and high (May-October) foliar cover.

RESULTS

We obtained 2,247 locations on 12 foxes captured between

September 1986 and August 1987 (Fig. 1 and 2). Five adult males, two

adult females, and five subadult females were captured. Only 10 animals

were included in home-range estimates. Because of variable tracking

periods among foxes, some animals could not be used in home-range

estimates of reproductive or dietary seasons (Table 1).

134 Cathryn H. Greenberg and Michael R. Pelton

^Zfy ADULT FEMALE

(74,80)

SUBADULT FEMALE

(62.63.64.66.72)

Fig. 1. Composite home ranges of two adult and five subadult female

gray foxes on the Oak Ridge Reservation in east Tennessee, Septem-

ber 1986-August 1987.

j^j] ADULT MALE (58, 69. 76. 82. 84)

Fig. 2. Composite home ranges of five adult male gray foxes on the

Oak Ridge Reservation in east Tennessee, September 1986-August 1987.

Gray Foxes 135

Table 1. Annual minimum cover polygon home-range estimates (km2)

for five male and seven female gray foxes radiotracked from Septem-

ber 1986 to August 1987, Oak Ridge Reservation in east Tennessee.

1 Omitted from reproductive season home-range analysis; insufficient data.

2 Omitted from dietary season home-range analysis; insufficient data.

3 Omitted from annual home-range analysis; insufficient data.

Mean annual home-range size (x ± SE) for 10 gray foxes was

3.97 ± 1.51 km2. Home-range size was similar between males (N = 5;

4.27 ± 1.59 km2) and females (N = 5; 3.67 ± 1.54 km2), and between

adult (N = 6; 4.41 ± 1.46 km2) and subadult (N = 4; 3.20 ± 1.62 km2)

foxes.

Home-range size was similar during fruit diet (N = 9; 2.92 ±

0.40) and flesh diet seasons (N = 9; 3.43 ± 0.48). Home-range sizes

were similar among reproductive seasons for all foxes (N = 8; 2.72 ±

0.17); (N = 7; 2.32 ± 0.43); and (N = 9; 2.83 ± 0.42) for breeding,

pup-rearing, and pre-breeding seasons, respectively. Within repro-

ductive seasons, male and female home-range sizes also were

similar (N = 4; 2.67 ± 0.11) versus (N = 4; 2.7 ± 0.30) for breeding;

(N = 3; 2.79 ± 0.59) versus (N = 4; 1.98 ± 0.55) for pup-rearing; and

(N = 4; 2.60 ± 0.30) versus (N = 5; 3.01 ± 0.67) for pre-

breeding seasons, respectively. We observed that whelping females

exhibited restricted movements during pup-rearing season.

Adjacent home ranges of four adult males were nearly exclusive

except for excursions by M 69 and M 76 into M 82's home range

during breeding season (Fig. 2). Subadults F 64, F 62, and F 78 home

ranges were contained within adult M 69's home range; F 64 and

136

Cathryn H. Greenberg and Michael R. Pelton

SUNRISE DAYLIGHT SUNSET NIGHT

Fig. 3. Diet-related (fruit versus flesh) activity levels (%) of gray foxes

radiotracked on the Oak Ridge Reservation in east Tennessee, September

1986-August 1987.

p 50

40

30

20

10

0

BREEDING

PUP-REARING

PREBREEDING

£J

SUNRISE DAYLIGHT SUNSET NIGHT

Fig. 4. Reproduction-related activity levels (%) of gray foxes radiotracked

on the Oak Ridge Reservation in east Tennessee, September 1986-

August 1987.

Gray Foxes

137

SUNRISE DAYLIGHT SUNSET NIGHT

Fig. 5. Cover-related activity levels (%) of gray foxes radiotracked on

the Oak Ridge Reservation, east Tennessee, September 1986-August 1987.

F 62 alternately used the same resting places. Adult male-female

pairs sharing home ranges include M 82 with F 80 (who died before

an adequate sample size was obtained for inclusion in home-range

analysis) and M 76 with F 74. Adult same-sex fox home ranges overlapped

little, whereas adult male-female pairs and adult-subadult home

ranges overlapped substantially. Subadult F 63 had an aberrantly

large "home range," which may have been explorations instead of a

home range at all.

Gray foxes were active on a greater proportion of locations

in evening and night hours than during sunrise and daylight hours.

Animals exhibited a lower sunrise activity level during flesh diet than

fruit diet season (x2 = 17.8, P < 0.0005) (Fig. 3). Lower sunrise

activity levels were observed during breeding season, and higher daylight

activity was observed during pup-rearing season than during other

reproductive seasons (x2 = 29.8, P < 0.0005) (Fig. 4). Lower sunrise

and daylight activity levels were observed during months of low foliar

cover than during months of high cover (x2 = 32.3, P < 0.0005)

(Fig. 5).

DISCUSSION

Gray fox home-range sizes were within the range of those

reported in other studies (Richards and Hine 1953, Fuller 1978, Yearsley

and Samuel 1980, Nicholson 1982, Hallberg and Trapp 1984,

Haroldson and Fritzell 1984, Wooding 1984). Nearly exclusive home

138 Cathryn H. Greenberg and Michael R. Pelton

ranges shared by adult male-female pairs and subadults suggested

that family units are spatially segregated. However, this conclusion

is tentative because uncollared foxes may have lived undetected

within the study area. Trapp and Hallberg (1975) suggest that a

family shares a home range exclusive of others, and they provide

some evidence for territoriality.

We were unable to detect any influence of seasonal dietary

composition or dietary trophic level on home-range size. Instead, we

suggest that diverse, interspersed habitat types within home ranges

might provide sufficient food supply in all seasons. Maintaining a

home range encompassing sufficient habitat area and types to provide

a year-round food supply might be a better strategy than shifting

home-range size in response to fluctuating patch productivity or

food availability (MacDonald 1980). Further study, including a larger

sample size, is warranted to determine the influence of seasonal diet

on gray fox home-range size.

Trends in home-range size indicate that males may range farther

than females during breeding season. High variance and small sample

size may obscure detection of seasonal differences in patterns of

home-range size or differences among age groups or sexes.

Predominantly nocturnal activity has been reported in other

studies (Nicholson 1982). Lower levels of sunrise activity during

flesh diet season, breeding season, and months of low foliar cover

could all be a response to sparse cover (November-April). Energy

conservation during a period of lower food availability may be a

factor. Higher daylight activity during pup-rearing season than during

other seasons might be due to increased energy requirements for

both parents and pups.

ACKNOWLEDGMENTS— This research was funded by the

Oak Ridge National Laboratory, Oak Ridge, Tennessee, operated by

the Martin-Marietta Energy Systems, Incorporated, for the U.S.

Department of Energy, under contract DE-AC05-840R2 1400. We wish

to thank P. D. Parr for her invaluable logistic support and assistance.

Thanks to H. A. Longmire, W. Sera J. Gittleman, A. Echternacht,

B. Dearden, T. Fendley, J. Evans, M. O'Neil, R. Sargeant, M. Sargeant,

and S. H. Crownover for advice and assistance.

Gray Foxes 139

LITERATURE CITED

Abies, E. D. 1968. Ecological studies on red foxes in southern

Wisconsin. Ph.D. Thesis. University of Wisconsin, Madison.

Boyle, K. A. 1986. A comparison of methods of home range analysis.

M.S. Thesis, Clemson University, Clemson, South Carolina.

Fuller, T. K. 1978. Variable home-range sizes of female gray foxes.

Journal of Mammalogy 59:446-449.

Greenberg, C. H., and M. R. Pelton. 1991. Food habits of gray

foxes (Urocyon cinereoargenteus) and red foxes {Vulpes vulpes)

in east Tennessee. Journal of the Tennessee Academy of Science

66(2):79-84.

Greenberg, C. H., M. R. Pelton, and P. D. Parr. 1988. Gray fox

ecology in the Oak Ridge National Environment Research Park:

food habitats, home range, and habitat use. Environmental Sciences

Division Publication No. 3101.

Geir, H. T. 1968. Coyotes in Kansas. Kansas State College Agricul-

tural Experiment Station Bulletin 393.

Hallberg, D. L., and G. R. Trapp. 1984. Gray fox temporal and

spatial activity in a riparian agricultural zone in California's

Central Valley. Pages 920-928 in California riparian systems:

ecology, conservation, and productive management. (R. E. Warner

and K. M. Hendrix, editors). University of California Press.

Berkeley.

Haroldson, K. J., and E. K. Fritzell. 1984. Home ranges, activity,

and habitat use by gray foxes in an oak-hickory forest. The Journal

of Wildlife Management 48:222-227.

Heezen, K. L., and J. R. Tester. 1967. Evaluation of radio-tracking

by triangulation with special reference to deer movements. The

Journal of Wildlife Management 31:124-141.

MacDonald, D. W. 1980. Resources dispersion and the social organiza-

tion of the red fox (Vulpes vulpes). Pages 918-949 in Worldwide

Furbearer Conference Proceedings, Volume II (J. A. Chapman and

D. Pursley, editors). Frostburg, Maryland.

McNab, B. K. 1963. Bioenergetics and the determination of home-

range size. American Naturalist 97:133-140.

Mohr. C. O. 1947. Table of equivalent populations of North American

small mammals. American Midland Naturalist 37:223-249.

Nicholson, W. S. 1982. An ecological study of the gray fox in east

central Alabama. M.S. Thesis, University of Georgia, Athens.

Nowak, R. M., and J. L. Paradiso. 1983. Walker's mammals of the

world. Volume 2. Fourth Edition. The Johns Hopkins University

Press, Baltimore, Maryland.

Richards, S. H., and R. L. Hine. 1953. Wisconsin fox populations.

Wisconsin Conservation Department Technical Wildlife Bulletin

Number 6.

140 Cathryn H. Greenberg and Michael R. Pelton

Smith, G. J., J. R. Gary, and O. J. Rongstad. 1981. Sampling strate-

gies for radio-tracking coyotes. Wildlife Society Bulletin 9:88-93.

Sullivan, E. G. 1956. Gray fox reproduction, denning range, and weights

in Alabama. Journal of Mammalogy 37:346-351.

Trapp, G. R., and D. L. Hallberg. 1975. Ecology of the gray fox.

Pages 164-178 in The wild canids (M. W. Fox, editor). Van

Nostrand Reinhold Company, New York, New York.

Wooding, J. B. 1984. Coyote food habits and the spatial relationship

of coyotes and foxes in Mississippi and Alabama. M.S. Thesis,

Mississippi State University, Starkville.

Yearsley, E. F., and D. E. Samuel. 1980. Use of reclaimed surface

mines by foxes in West Virginia. The Journal of Wildlife Manage-

ment 44:729-734.

Received 11 January 1994

Accepted 29 June 1994

External Morphometries of Blaek Bears, Ursus americanus

(Carnivora: Ursidae), in the Great Dismal Swamp

of Virginia and North Carolina

ERIC C. HELLGRhN

Campus Box 218, Caesar Kleberg Wildlife Research Institute

Texas A&M University-Kingsville, Kingsville, Texas 78363

And

Michael R. Vaughan

Virginia Cooperative Fish and Wildlife Research Unit, Virginia

Polytechnic Institute and State University,

Blacksburg, Virginia 24061

ABSTRACT — We report body measurements of black bears (Ursus

americanus) for Great Dismal Swamp, a seasonally-flooded, forested

wetland in the Atlantic Coastal Plain. For most body

measurements, males reached adult size by 5 years of age and

females by 3-4 years of age. Chest girth, and zygomatic cir-

cumference were the best (P < 0.001) predictors of body mass

for both sexes. External morphometries can be used to predict

nutritional condition. Growth models using mass or length data

can be developed and growth rates compared among popula-

tions. Such comparisons may shed light on black bear tax-

onomy or habitat quality.

Published data on black bear external morphometries, other

than body mass, are scanty, although common in unpublished theses.

Sauer (1975) reported a large data set of external measurements of

black bears from New York. Other publications containing data on

black bear morphometries include Poelker and Hartwell (1973:89-

104), Cherry and Pelton (1976), and Juniper (1978) from Washington,

Tennessee, and Quebec, respectively.

Body morphometries and growth are important characters in the

study of intraspecific regional variation. In a mammal with a wide

distribution such as the black bear, such data may provide insights

into relationships among environmental factors, particularly nutrition,

and genetic potential. For example, mean body masses of adult (>5-

year-old) male black bears range from 96 kg in western Montana

(Jonkel and Cowan 1971) to 183 kg in Pennsylvania (Alt 1980), a

range mirrored by differences in reproductive rates and attributed to

differences in food availability (Bunnell and Tait 1981). Kingsley et

al. (1988) found differences in growth curves and body size in three

Brimleyana 21:141-149, December 1994 141

142 Eric C. Hellgren and Michael R. Vaughan

disjunct populations of brown bears (Ursus arctos). They attributed

variation to differences in system productivity or bear density.

As part of a larger project studying black bear ecology and

physiology in Great Dismal Swamp, we collected body measurements

from live-captured black bears (Ursus americanus americanus) (Hall

1981). Our objectives were to provide baseline data on body measurements

by age for black bears in the Atlantic Coastal Plain and to produce

prediction equations for body mass based on morphometric measurements.

MATERIALS AND METHODS

We conducted research from April 1984 to August 1986 on a

555-km2 study area containing the 440-km: Great Dismal Swamp

National Wildlife Refuge, 57.5-km2 Dismal Swamp State Park, and

adjacent private land. Descriptions of the study area were reported

elsewhere (Hellgren and Vaughan 1988, 1989a). We captured 101

different bears 120 times with spring-activated cable snares during

April through December. Bears were immobilized with a 2:1 mixture

of ketamine hydrochloride and xylazine hydrochloride at a concentration

of 300 mg/mL administered intramuscularly at an initial dosage rate

of 6.6 mg/kg. Mass was measured to the nearest kg with a hanging

spring scale.

We took measurements on immobilized animals to the nearest

mm. Body length was measured from the tip of the snout to the distal

end of the last caudal vertebra while the animal was in lateral recumbency.

Head length was measured from the tip of the nose to the occiput.

Neck girth was measured in the middle of the neck. Chest girth was

measured immediately posterior to the scapulae. Circumferences of

wrist and elbow (at olecranon process) also were measured. Zygomatic

circumference was measured anterior to the ears. The above measure-

ments were taken with a cloth tape pulled snug. Tail length (from

base of tail to distal end of caudal vertebra), ear length (from inner

notch to tip of pinna), forepaw and hindpaw width (greatest distance

across pads), and forepaw and hindpaw length (longest distance along

length of pads) were measured with a steel tape. Canine measurements

were taken with dial calipers to the nearest 0.1 mm. Upper and lower

canine breadths were the distance between the tips of the right and

left maxillary and mandibular canines, respectively. Upper and lower

canine lengths were measured from the gum line to the tip of the

canine. Anterior-posterior lengths and lingual-labial widths of upper

and lower canines were measured at the gum line.

We used one-way analysis of variance to examine age differences

in physical characteristics within each sex. We did not analyze data

Black Bear Morphometries 143

for differences by sex because of obvious size dimorphism. Because of

small sample sizes and asymptotic growth, all animals >7 years

old were placed into one age category. Samples were pooled across

seasons, and all data were analyzed. When data collected after 15

September (n = 19) were deleted, mean body mass for males and

females decreased by a maximum of 2.1 and 6.0 kg for any year class.

Previous analyses showed an age-season (age catergorized as adults or

subadults) interaction (P = 0.06) in body mass for females and non-

significant seasonal variation (P = 0.11) for males (Hellgren and Vaughan

1989b), probably because of small samples in fall and, subsequently,

weak statistical power. We used Tukey's studentized range test to separate

means. Recapture data for individuals recaptured within the same year

were not included in any analyses. Recaptures in different years

(n = seven male, nine female) were treated as independent observations.

Simple linear regression was used to develop relationships among body

mass and body measurements.

RESULTS AND DISCUSSION

Ages ranged from 1 to 16 years for males (n = 71). All morphometric

variables measured varied (P < 0.001) by age except ear length (x±

SE) (119 ± 1 mm, n = 64) and tail length (73 ± 2 mm, n = 65) (Table

1). Based on means separation, we concluded that adult size was reached

for most body and canine measurements by 5 years of age. Body mass

continued to increase until 6 years of age, with a maximum mass of

198 kg for a 7-year-old individual captured in July.

It is interesting to note the lack of morphometric differences

(P > 0.05) between 3- and 4-year-old male bears. The stress of competing

for access to reproducing females may reduce body growth in these

young males, as nutrients are partitioned away from growth and into

demands for mate-searching and male-male aggression (Garshelis and

Hellgren 1994).

Females ranged in age from 1 to 9 years (n = 37). Body measure-

ments that did not vary by age (n = 34) were ear length (112 ± 1 mm),

tail length (74 ± 3 mm), forepaw width (83 ± 1 mm), forepaw length

(85 ± 1 mm), hindpaw length (79 ± 1 mm), and hindpaw length (147

± 1 mm). Female adult size was reached at an earlier age than male

adult size (Table 2). Adult size in body measurements was generally

reached by 3 or 4 years, whereas adult canine size was reached by 2

years of age. In New York, female bears attained adult size for all

measured characteristics by 2.5 years (Sauer 1975).

Morphometric data are limited for other southeastern wetland

bear populations. Adult (>3 years) males and females weighed an

144

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Eric C. Hellgren and Michael R. Vaughan

average of 120 and 88 kg, respectively, in Bladen County, North Carolina,

(Hamilton 1978) and 102 and 52 kg, respectively, in a bottomland

hardwood swamp in eastern Arkansas (Smith 1985). Harvested, adult

females from the periphery of Okefenokee Swamp, Georgia, ranged in

mass from 46 to 101 kg (Abler 1985). Smith (1985) reported that

males reached peak mass by 5 years of age. Although females reached

adult stature by age 2 or 3, they continued gaining mass and girth until

age 9 or 10. Bears in Great Dismal Swamp became heavier than bottomland

Arkansas bears (Smith 1985) by age 6 in males and age 5 in females.

Total length and chest girth measurements were also larger for Dismal

Swamp males by age 6.

Table 3. Significant (P < 0.0001) bivariate regression models with body

mass (kg) as the dependent variable for black bears in Great Dismal Swamp,

Virginia and North Carolina, 1984-1986.

Black Bear Morphometries 147

Regression analyses indicated that chest girth, neck girth, and

zygomatic circumference were the best predictors of body mass for

both sexes (Table 3). Chest girth has been used commonly to estimate

body mass in bears (Cherry and Pelton 1976, Glenn 1980, Nagy et al.

1984), although Swenson et al. (1987) cautioned that interpopulation

variation in measurement-mass relationships makes it impossible to

produce a single, species-specific equation. These authors also concluded

that gender variation warranted development of sex-specific prediction

equations.

Morphometric data can be used to predict nutritional condition

(Cattet 1990) and make intraspecific comparisons of body size. Differences

in body size and growth rates of black bears of different populations

resulting from variability in ecosystem productivity may lead to differences

in skull morphometry, a key tool in taxonomic analysis. If morphometric

variation between populations is best explained by phenotypic responses

to the environment, can morphometries be used to classify animals

into subspecies (Pelton 1990)? Such a question is germane to taxonomy

of black bears and other species.

Our paper reports on a single, southeastern Coastal Plain

population of black bears. We encourage other black bear researchers

to standardize the collection and reporting of data on external

morphometries to maximize their utility.

ACKNOWLEDGMENTS— Viz thank the United States Fish and

Wildlife Service, Virginia Department of Game and Inland Fisheries,

North Carolina Wildlife Resource Commission, North Carolina State

Parks Department, and the Department of Fisheries and Wildlife Sciences,

Virginia Polytechnic Institute and State University. Particularly helpful

were D. J. Schwab, R. D. McClanahan, and the entire staff of Great

Dismal Swamp National Wildlife Refuge. Technical field assistance was

provided by W. M. Lane, J. R. Polisar, and K. M. Meddleton. This

project was funded by the United States Fish and Wildlife Service.

LITERATURE CITED

Abler, W. A. 1985. Bear population dynamics on a study area in

southeastern Georgia. Georgia Department of Natural Resources, Atlanta.

Alt, G. L. 1980. Rate of growth and size of Pennsylvania black

bears. Pennsylvania Game News 51(12):7-17.

Bunnell, F. L., and D. E. N. Tait. 1981. Population dynamics of

bears — implications. Pages 75-98 in Dynamics of large mammal

populations. (T. D. Smith and C. Fowler, editors). John Wiley

and Sons, Incorporated, New York, New York.

148 Eric C. Hellgren and Michael R. Vaughan

Cattet, M. 1990. Predicting nutritional condition in black bears and

polar bears on the basis of morphological and physiological mea-

surements. Canadian Journal of Zoology 68:32-39.

Cherry, J. S., and M. R. Pelton. 1976. Relationships between body

measurements and weight of the black bear. Journal of the Ten-

nessee Academy of Sciences 51:32-34.

Garshelis, D. S., and E. C. Hellgren. 1994. Variation in reproductive

biology of male black bears. Journal of Mammalogy 75:175-188.

Glenn, L. P. 1980. Morphometric characteristics of brown bears on

the central Alaska Peninsula. International Conference on Bear

Research and Management 4:313-319.

Hall, E. R. 1981. The mammals of North America. Second edition.

John Wiley and Sons, Incorporated, New York, New York.

Hamilton, R. J. 1978. Ecology of the black bear in southeastern

North Carolina. M. S. Thesis, University of Georgia, Athens.

Hellgren, E. C, and M. R. Vaughan. 1988. Seasonal food habits of

black bears in Great Dismal Swamp, Virginia-North Carolina. Pro-

ceedings of the Annual Conference of the Southeastern Associa-

tion of Fish and Wildlife Agencies 42:295-305.

Hellgren, E. C, and M. R. Vaughan. 1989a. Demographic analysis

of a black bear population in the Great Dismal Swamp. The

Journal of Wildlife Management 53:969-977.

Hellgren, E. C, and M. R. Vaughan. 19896. Seasonal patterns in

physiology and nutrition of black bears in Great Dismal Swamp,

Virginia-North Carolina. Canadian Journal of Zoology 67:1837-1850.

Jonkel, C. J., and I. T. Cowan. 1971. The black bear in the spruce-

fir forest. Wildlife Monographs 27:1-57.

Juniper, I. 1978. Morphology, diet, and parasitism in Quebec black

bears. Canadian Field-Naturalist 92:186-189.

Kingsley, M. C. S., J. A. Nagy, and H. V. Reynolds. 1988. Growth in

length and weight of northern brown bears: Differences between

sexes and populations. Canadian Journal of Zoology 66:981-986.

Nagy, J. A., M. C. Kingsley, R. H. Russell, A. M. Pearson, and B. C.

Goski. 1984. Relationship of weight to chest girth in the grizzly

bear. The Journal of Wildlife Management 48:1439-1440.

Pelton, M. R. 1990. Black bears in the Southeast: To list or not to

list? Eastern Workshop on Black Bear Research and Management

10:155-161.

Poelker, R. J., and H. D. Hartwell. 1973. Black bear of Washing-

ton. Washington State Game Department Biological Bulletin Number

14. Olympia, Washington.

Sauer, P. R. 1975. Relationship of growth characteristics to sex and

age for black bears from the Adirondack region of New York.

New York Fish and Game Journal 22:81-113.

Black Bear Morphometries 149

Smith, T. R. 1985. Ecology of black bears in a bottomland hard-

wood forest in Arkansas. Ph.D. Thesis, University of Tennessee,

Knoxville.

Swenson, J. E., W. F. Kasworm, S. T. Stewart, C. A. Simmon, and

K. Aune. 1987. Interpopulation applicability of equations to

predict live weight in black bears. International Conference on

Bear Research and Management 7:359-362.

Received 28 June 1994

Accepted 14 September 1994

150

Nutrient Content of Squawroot, Conopholis americana,

and Its Importance to Southern Appalachian Black

Bears, Ursus americanus (Carnivora: Ursidae)

Steven G. Seibert1 and Michael R. Pelton

Department of Forestry, Wildlife and Fisheries,

The University of Tennessee, Knoxville, Tennessee 37901

ABSTRACT — Squawroot (Conopholis americana), a preferred late

spring and early summer food of black bears (Ursus americanus),

was collected from Pisgah National Forest, North Carolina, on a

weekly basis from 25 April to 4 July 1987. Proximate analysis

procedures were used to determine the nutrient content of the

plant. Samples were examined for nutrient differences between

the capsule and stems. Peak percentages were 13% crude protein

(capsule), 31% crude fiber (capsule), 3% fat [either extract] (capsule),

and 77% nitrogen-free extract [NFE] (whole plant). Gross energy

averaged 4.84 kcal/dry g. Levels of crude protein, crude fiber,

and either extract were similar to values reported for soft mast

species eaten by bears, and NFE was greater than herbaceous

material consumed in spring. Trends in protein and fat content

were higher in the capsules; protein decreased as crude fiber

increased. Nitrogen-free extract levels were relatively high throughout

the study and likely represent an important energy source for

bears feeding on squawroot.

Squawroot (Conopholis americana) is a perennial, parasitic plant

(Musselman 1982) common to the Piedmont and southern Appalachian

(Harvill et al. 1981). Little is known about the plant, but it appears to

grow only from the roots of oak trees (Musselman and Mann 1978),

probably by infecting young root tips (Musselman 1982).

Squawroot also is a common food eaten by black bears in spring

and early summer in the southern Appalachians (Beeman and Pelton

1980, Eagle and Pelton 1983, Garner 1986); the species is locally

abundant and may be nutritionally important to bears. Because of its

local abundance and time of maturity (often the first productive food

available), squawroot patches may influence movements of female

bears in the southern Appalachians. By locating readily-available,

high-energy foods, females may improve their energy benefit/cost

ratio, thereby increasing cub survival. The purpose of this study was

to determine the nutritional content of squawroot.

1 Present address: United States Fish and Wildlife Service, 6620 Southpoint Drive

South, Suite 310, Jacksonville, Florida 32216.

Brimleyana 21:151-156, December 1994 151

152

Steven G. Seibert and Michael R. Pelton

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Nutrient Content of Squawroot 153

STUDY AREA

The study was conducted on the Harmon Den Bear Sanctuary,

Pisgah National Forest, North Carolina. The area is part of the Blue

Ridge Physiographic Province (Fenneman 1938). The mountains are

sharply dissected and the terrain is steep. Elevations range from 439 to

1,411 m; slopes average over 30% (Finlayson 1957). The area is described

as a warm-temperate rain forest (Thornwaite 1948). Vegetation is diverse

and changes dramatically with aspect, elevation, soil, and drainage.

The general area is among the most botanically diverse temperate areas

in the world (Whittaker 1956).

The majority (89%) of Harmon Den is in hardwood cover types,

consisting of white oak (Quercus alba)-northem red oak (Q. rubra)-

hickory {Carya sp.) (45%), yellow poplar (Liriodendron tulipifera)-

white oak-northern red oak (26%), yellow poplar (10%), scarlet oak

(Q. coccinea) (5%), and chestnut oak (Q. prinus) (3%) (U.S. Forest

Service 1988).

METHODS

Whole plants of squawroot (20-30 plants) were collected weekly

from 25 April to 4 July 1987 at elevations ranging from 946 to 1,068

m. Quantities of 639 to 1,123 g were obtained each week; samples

were composited, therefore no statistical analysis could be performed.

Thus, the trends or differences noted in this paper may not be statistically

significant. Oven-dried samples were analyzed for crude protein, crude

fiber, nitrogen-free extract, fat (ether extract), and gross energy (Maynard

et al. 1979).

Capsules and stems were analyzed separately to detect nutritional

differences between them because, as squawroot matured, bears were

observed to selectively feed on the capsule portion of the plant. Capsules

were difficult to separate from the stem before 30 May.

RESULTS

Crude Protein

The trend for crude protein (x = 8.8%, SE = 0.51, range = 5.1-

12.3%) of the entire plant peaked during late May (Fig. 1). Capsules

(x - 7.8%, SE = 0.78, range = 5.5-13.4% contained more protein than

stems (x = 5.5%, SE = 0.58, range = 3.6-10.2) throughout the study.

The highest percentage of protein occurred in the capsules collected 30

May. However, much of this protein may have been in the seeds (Wainio

and Forbes 1941) and unavailable because bears do not crack or digest

the seed coat (Eagle and Pelton 1983).

154 Steven G. Seibert and Michael R. Pelton

Fat (Ether Extract)

Fat content in squawroot was low (Fig. 1) and similar to soft

mast species such as huckleberry (Gaylussacia sp.) and black gum

(Nyssa sylvatica) (Landers et al. 1979), which also are important to

bears. The greatest trend in percentage of fat occurred in the capsules

collected on 13 June. Fat trends were higher in capsules (x = 1.53%,

SE = 0.14, range = 1.0-3.0%) than stems (x = 0.75%, SE = 0.12,

range = 0.1-1.4%) for all except the final collection. Mean fat content

for the whole plant was 1.34% (SE = 0.69, range = 0.7-2.5%).

Crude Fiber

The trend in crude fiber for the stem (x = 19.23%, SE = 0.65,

range = 14.8-21.5%) was similar throughout the study (Fig. 1); this

might be expected because the stem is the only structural component

of the plant. Crude fiber increased in the capsules (x = 22.62%, SE =

2.24, range = 10.2-31.5%) throughout the study period and appeared

negatively correlated with protein. Crude fiber averaged 16.53% (SE

= 1.65, range = 6.4-29.0%) for the whole plant.

Nitrogen-Free Extract (NFE)

Nitrogen-free extract was highest in early spring and lowest in

late June in the whole plant (x = 69.1%, SE = 1.14, range = 61.1—

77.4%) and capsules (x = 63.7%, SE = 1.23, range = 58.4-70.7%)

(Fig. 1). NFE increased from late May to July in the stems (x =

70.7%, SE = 2.88, range = 65.1-74.9%).

Gross Energy

There was little variation in gross energy among weeks, or between

the different plant parts; this agrees with Robbins (1983) and Powell

and Seaman (1990). Gross energy averaged 4.8 kcal/dry g (SE = 0.03,

range = 4.7-5.1) (whole plant); 5.0 kcal/dry g (SE = 0.05, range =

4.9-5.2) (capsules); and 4.7 kcal/dry g (SE = 0.06, range = 4.3-5.0)

(stems).

The nutritional content of squawroot appeared to change over

time. Protein and nitrogen-free extract (NFE) concentrations were

greatest during the early weeks. Fat was greatest during middle of the

study, and crude fiber was lowest early and increased with time.

DISCUSSION

Spring diets of bears in the southern Appalachians contain large

amounts of herbaceous material (Beeman and Pelton 1980, Eagle and

Pelton 1983, Garner 1986). Herbaceous material is relatively high in

protein (Landers et al. 1979, Eagle and Pelton 1983). Nitrogen-free

Nutrient Content of Squawroot 155

extract, however, is lowest in spring foods (Landers et al. 1979).

Eagle and Pelton (1983) suggested that squawroot was probably an

important energy source for bears because the carbohydrates in squawroot

are readily absorbed.

Squawroot is likely a major source of carbohydrates (represented

by relatively high NFE concentrations) in the spring diet of bears in

the Harmon Den area. Nitrogen-free extract concentrations in squawroot

tended to remain relatively high throughout the study, and carbohydrates

available in squawroot appear to be easily absorbed (Eagle and

Pelton 1983); this may be important to a bear's overall spring and

early summer condition. The combination of high protein herbaceous

material and relatively rich carbohydrate squawroot may be important

for bears recovering from the denning period; particularly for lactating

females with cubs, because of their increased nutritional requirements

(Eagle and Pelton 1983, Rogers 1987).

The habitat types where squawroot occurs (i.e., mature oak stands)

should receive special concerns, as management of these stands for

peak acorn production also would maintain ample sources of squawroot

for bears throughout their range in the southern Appalachians.

LITERATURE CITED

Beeman, L. E., and M. R. Pelton. 1980. Seasonal foods and feeding

ecology of black bears in the Smoky Mountains. Proceedings of

the International Conference for Bear Research and Management

4:141-147.

Eagle, T. C, and M. R. Pelton. 1983. Seasonal nutrition of black

bears in the Great Smoky Mountains National Park. Proceedings

of the International Conference for Bear Research and Manage-

ment 5:94-101.

Fenneman, N. M. 1938. Physiography of the eastern United States.

McGraw-Hill Book Company, New York, New York.

Finlayson, C. P. 1957. The geology of the Max Patch Mountain

area, Lemon Gap Quadrangle, Tennessee-North Carolina. M.S. Thesis.

The University of Tennessee, Knoxville.

Garner, N. P. 1986. Seasonal movements, habitat selection, and food

habits of black bears (Ursus americanus) in Shenandoah National

Park. M.S. Thesis. Virginia Polytechnic Institute and State Uni-

versity, Blacksburg.

Harvill, A. M., Jr., T. R. Bradley, and C. E. Sterns. 1981. Atlas of

the Virginia flora. Part II Dicotyledons. Virginia Botanical Asso-

ciates, Farmville, Virginia.

Landers, J. L., R. J. Hamilton, A. S. Johnson, and R. L. Marchinton.

1979. Foods and habitat of black bears in southeastern North

Carolina. The Journal of Wildlife Management 43:143-153.

156 Steven G. Seibert and Michael R. Pelton

Maynard, L. A., J. K. Loosli, H. F. Hintz, and R. G. Warner. 1979.

Animal nutrition. Seventh edition. McGraw-Hill Book Company, New

York, New York.

Musselman, L. J. 1982. The Orobanchaceae of Virginia. Castanea 47:266-

275.

Musselman, L. J., and W. F. Mann, Jr. 1978. Root parasites of

southern forests. United States Department of Agriculture, Forest

Service General Technical Report SO-20.

Powell, R. A., and D. E. Seaman. 1990. Production of important

black bear foods in the southern Appalachians. Proceedings of the

International Conference for Bear Research and Management 8:183-

187.

Robbins, C. T. 1983. Wildlife nutrition and feeding. Academic Press,

Incorporated, New York, New York.

Rogers, L. L. 1987. Effects of food supply and kinship on social

behavior, movements, and population growth of black bears in northeastern

Minnesota. Wildlife Monograph 97:1-72.

Thornwaite, C. W. 1948. An approach toward a rational classification

of climate. Geogography Review 38:55-94.

Wainio, W. W., and E. B. Forbes. 1941. The chemical composition of

forest fruits and nuts from Pennsylvania. Journal of Agricultural

Research 62:627-635.

Whittaker, R. H. 1956. Vegetation of the Great Smoky Mountains.

Ecological Monograph 26:1-50.

Received 11 January 1994

Accepted 10 March 1994

157

NEW PUBLICATION

Bird Life of North Carolina 's Shining Rock Wilderness is the

most recent release in the Occasional Papers of the North Carolina

Biological Survey series published by the North Carolina State Museum

of Natural Sciences. Written by Marcus B. Simpson, Jr., M.D., of the

George Washington University Medical Center, Washington, D.C.,

the work summarizes the geomorphology, climate, vegetation, and

ornithological exploration of the wilderness area and presents an annotated

list of 130 species known to occur within its boundaries. Illustrations

include a map of Shining Rock Wilderness and adjacent Middle Prong

Wilderness, both managed by the United States Forest Service, an

agency of the U.S. Department of Agriculture. The 32-page soft-

cover book sells for $5 postpaid. Send orders to: Publications Secretary,

N.C. State Museum of Natural Sciences, P.O. Box 29555, Raleigh,

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Extension Fund.

DATE OF MAILING

Brimleyana 20 was mailed on 5 August 1994.

158

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Adams, J. J. 1977. Food habits of the masked shrew, Sorex cinereus

(Mammalia: Insectivora). Brimleyana 7:32-39.

Adams, J. J. 1988. Animals in North Carolina folklore. Second edition.

University of North Carolina Press, Chapel Hill.

Barnes, R. G. 1986. Range, food habits, and reproduction in Glaucomys

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and Tennessee. Ph.D. Thesis. North Carolina State University, Raleigh.

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24 pages, 1988, $3 postpaid

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David S. Lee and James F. Parnell, editors, 52 pages,

1990, $8 postpaid

Atlas of North American Freshwater Fishes

David S. Lee et al., 867 pages, 1980, $25 postpaid

Atlas of North American Freshwater Fishes, 1983 Supplement

D. S. Lee, S. P. Platania, and G. H. Burgess, 68 pages plus

looseleaf additions and corrections, 1983, $10 postpaid

A Distributional Survey of North Carolina Mammals

David S. Lee, John B. Funderburg, and Mary K. Clark,

70 pages, 1982, $5 postpaid

The Seaside Sparrow, Its Biology and Management

Thomas L. Quay et al., editors, 174 pages, 1983, $15 postpaid

Autumn Land-bird Migration on the Barrier Islands

of Northeastern North Carolina

Paul W. Sykes Jr., 50 pages, 1986, $5 postpaid

Potential Effects of Oil Spills on Seabirds and

Selected Other Oceanic Vertebrates Off the North Carolina Coast

David S. Lee and Mary C. Socci, 64 pages, 1989,

$8 postpaid

Bird Life of North Carolina's Shining Rock Wilderness

Marcus B. Simpson, Jr., 32 pages, 1994, $5 postpaid

Send orders to: N.C. State Museum of Natural Sciences

Attention: Beverly Craven

P.O. Box 29555, Raleigh, NC 27626-0555

Please make checks payable in U.S. currency to Museum Extension Fund.

BRIMLEYANA NO. 21, DECEMBER 1994

CONTENTS

The Mammals of the Ardis Local Fauna (Late Pleistocene), Harleyville,

South Carolina. Curtis C. Bentley, James L. Knight, and Martin A. Knoll 1

Comments on the Body Mass Trend of Ondatra zibethicus (Rodentia: Muridae)

During the Latest Pleistocene. Curtis C. Bentley and James L. Knight 37

Introductions of the Scorpions Centruriodes vittatus (Say) and C. hentzi (Banks) into

North Carolina, with Records of the Indigenous Scorpion, Vaejovis carolinianus

(Beauvois) (Scorpionida: Buthidae, Vaejovidae). Rowland M. Shelley 45

Distribution of the Scorpion, Vaejovis carolinianus (Beauvois) — a Reevaluation,

(Arachnida: Scorpionida: Vaejovidae). Rowland M. Shelley 57.

Atlantic Ocean Occurrences of the Sea Lamprey, Petromyzon marinus

(Petromyzontiformes: Petromyzontidae), Parasitizing Sandbar,

Carcharhinus plumbeus, and Dusky, C. obscurus (Carcharhiniformes:

Carcharhinidae), Sharks off North and South Carolina.

Christopher Jensen and Frank J. Schwartz 69

Clutch Parameters of Storeria dekayi Holbrook (Serpentes: Colubridae) from

Southcentral Florida. Walter E. Meshaka,Jr 73

Influence of Environmental Conditions on Flight Activity of Plecotus

townsendii virginianus (Chiroptera: Vespertilionidae).

Michael D. Adam, Michael J. Lacki, and Laura G. Shoemaker 77

The Pygmy Shrew, Sorex hoyi winnemana (Insectivora: Soricidae), from the

Coastal Plain of North Carolina. Thomas M. Padgett and Robert K. Rose 87

Additional Records of the Pygmy Shrew, Sorex hoyi winnemana Preble

(Insectivora: Soricidae), in Western North Carolina.

Joshua Laerm, William M. Ford, and Daniel C. Weinand 91

Food and Ectoparasites of the Southern Short-tailed Shrew, Blarina carolinensis

(Mammalia: Soricidae), from South Carolina.

John O. Whitaker, Jr., Gregory D. Hartman, and Randy Hein 97

Mensural Discrimination of Four Species of Peromyscus (Rodentia: Muridae) in the

Southeastern United States. Joshua Laerm and James L. Boone 107

Small Mammal Communities in Streamside Management Zones.

Dagmar P. Thurmond and Karl V. Miller 125

Home Range and Activity Patterns by Gray Foxes, Urocyon cinereoargenteus

(Carnivora: Canidae), in East Tennessee.

Cathryn H. Greenberg and Michael R. Pelton 131

External Morphometries of Black Bears, Ursus americanus (Carnivora: Ursidae),

in the Great Dismal Swamp of Virginia and North Carolina.

Eric C. Hellgren and Michael R. Vaughan 141

Nutrient Content of Squawroot, Conopholis americana, and Its Importance to Southern

Appalachian Black Bears, Ursus americanus (Carnivora: Ursidae).

Steven G. Seibert and Michael R. Pelton 151

Miscellany 157