december 1 980

EDITORIAL STAFF

John E. Cooper, Editor

Alexa C. Williams, Managing Editor

John B. Funderburg, Editor-in-Chief

Alvin L. Braswell, Curator of

Lower Vertebrates, N.C.

State Museum

John C. Clamp, Associate Curator

(Invertebrates), N.C.

State M useum

Martha R. Cooper, Associate

Curator ( Crustaceans ), N.C.

State Museum

James W. Hardin, Department

of Botany, N.C. State

University

Board

David S. Lee, Chief Curator

of Birds and Mammals, N.C.

State Museum

William M. Palmer, Chief Curator

of Lower Vertebrates, N.C.

State Museum

Thomas L. Quay, Department

of ^oology, N.C. State

University

Rowland M. Shelley, Chief

Curator of Invertebrates, N.C.

State Museum

Bnmleyana, the Journal of the North Carolina State Museum of Natural His-

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Subject matter will focus on taxonomy and systematics, ecology, zoo-

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In citations please use the full name — Bnmleyana.

North Carolina State Museum of Natural History

North Carolina Department of Agriculture

James A. Graham, Commissioner

CODN BRIMD 7

ISSN 0193-4406

The Milliped Fauna of the Kings Mountain Region

of North Carolina (Arthropoda: Diplopoda)

Marianne E. Filka

Department of Zoology,

North Carolina State University,

Raleigh, North Carolina 27607

AND

Rowland M. Shelley

North Carolina State Museum of Natural History,

P.O. Box 27647 , Raleigh, North Carolina 2761 D

ABSTRACT. — The millipeds of the Kings Mountain region of Cleve-

land and Gaston counties, one of five inselberg areas in the Piedmont

Plateau of North Carolina, were sampled to determine seasonal varia-

tion in faunal composition. Comparative collections also were made at

Spencer Mountain, an inselberg located northeast of Gastonia. Of 24

species taken, only Narceus americanus (Beauvois) and Auturus

erythropygos (Brandt) were present as adults and juveniles in all three

sampling seasons. The most diverse assemblage was encountered in Oc-

tober. Five species were more common in April and October than in

July, four were more common in July than in either of the cooler

months, and five others were collected in but a single season (three in

July, two in October). A more diverse fauna was encountered in the con-

tiguous Kings-Crowders ridge than at the isolated Spencer Mountain

outcrop, from which three xystodesmids were conspicuously absent. A

notable difference between millipeds of the two localities involved color

pattern of the intergrades of Sigmoria latior (Brolemann). Ptyoiulus was

the sole genus represented by more than one species, and the overall

species/genus ratio is indicative of a lowland rather than a montane

fauna.

The Kings Mountain region shares eight species with the eastern

Piedmont and five with the Appalachian Mountains. Seven widespread

species occur in all three regions, but three species are unique to the

Kings Mountain region. This area is the northeastern range limit of the

genus Pachydesmus; and the easternmost populations of four montane

diplopods, the westernmost population of Ptyoiulus ectenes (Bollman),

and the southernmost known locality of Cleidogona medialis Shelley, oc-

cur there. The conservation status of three species of concern to North

Carolina is discussed, and the Kings Mountain region is considered a

“cluster area” due to its unique diplopod fauna. The ranges of Boraria

stricta (Brolemann) and Deltotaria lea Hoffman are extended into South

Carolina. A key to genera and species is provided along with pertinent

diagnostic illustrations.

'Adjunct Assistant Professor of Zoology, North Carolina State University

1

Brimleyana No. 4: 1-42. December 1980.

2

Marianne E. Filka and Rowland M. Shelley

INTRODUCTION

The importance of the Appalachian Mountains to the arthropod

class Diplopoda has been evident since 1969, when Hoffman identified

the mountains as a global center of milliped evolution. This opinion was

based on the diversity and abundance of known indigenous taxa. Four

other areas also were cited as important global centers of evolution and

dispersal, and since all are mountainous to some extent, Hoffman sur-

mised that vertical relief probably allows for a greater variety of

ecological niches than occur in lowland or flat areas.

The Appalachian Highlands, one of eight physiographic divisions of

the United States, is comprised of seven physiographic provinces (Hunt

1967). The most important in terms of known diplopod faunas are the

Ridge and Valley and Blue Ridge Provinces, especially the southern sec-

tion of the latter (the region south of the Roanoke River). The

Xystodesmidae, the dominant Nearctic polydesmoid family, attains its

greatest known diversity in the part of the southern Blue Ridge Province

south of the Kanawha River System (Hoffman 1969). The bulk of the

southern Blue Ridge Province is in western North Carolina, where it is

demarcated from the Piedmont Plateau by a prominent escarpment, the

Blue Ridge Front. Thus, for all practical purposes one of the five regions

of greatest milliped diversity in the world lies in the western part of this

state.

Although most of the mountains of North Carolina are west of the

Blue Ridge Front, a number of prominent hills and ridges also occur to

the east in the Piedmont Plateau. Some of these are quite properly called

mountains and extend to altitudes of well over 300 meters. These isolated

mountains protruding from a surrounding flat plain are known as in-

selbergs and are erosional remnants of previously more extensive moun-

tain masses (Kesel 1974). Five main groups of inselbergs occur in Pied-

mont North Carolina (Fig. 1): the Sauratown Mountains of Stokes

County (including Pilot Mountain, Surry County); the Brushy Moun-

tains of Wilkes, Caldwell, Alexander and Iredell counties; the South

Mountains of Burke, Rutherford, McDowell and Cleveland counties; the

Kings Mountain region of Cleveland and Gaston counties; and the

Uwharrie Mountains of Davidson, Randolph, Montgomery, and Stanly

counties. The faunas of these inselberg regions are of particular

biogeographic interest and raise a number of questions. Do they, for ex-

ample, reflect previous direct connection with the Blue Ridge chain? If

so, their later isolation may have separated previously continuous

diplopod populations and led to speciation by geographical isolation.

Accordingly, knowledge of the inselberg diplopod faunas may provide

insights into processes affecting milliped evolution, and an investigation

of one such area was conducted in this study.

The Kings Mountain region straadles the South Carolina border

about 13 km southwest of Gastonia. Preliminary studies there had dis-

closed a milliped fauna with southern elements found nowhere else in

Kings Mountain Milliped Fauna

3

BLUE RIOGE FRONT ^*l-L LINE

North Carolina. The other inselberg regions have no faunas of such

singular importance to the state. Information on unique areas in North

Carolina is timely in regard to recent concerns about environmental

management and planning, as reflected by the North Carolina Environ-

mental Policy Act of 1971; the State Land Policy and Coastal Area

Management Acts, both enacted in 1974; and the Symposium on En-

dangered and Threatened Biota of North Carolina (see Cooper et al.

1977). Moreover, the North Carolina Natural Heritage Program, ad-

ministered by the Department of Natural Resources and Community

Development through a contract with The Nature Conservancy, is

presently conducting an inventory of the state’s most significant natural

areas. In order to realize the goals of these programs and to effectively

manage the resources of the state, knowledge of its indigenous flora and

fauna must be substantiated. Another objective of this project, therefore,

was to furnish such knowledge for the Diplopoda of the Kings Mountain

area, and categories of concern are suggested in some of the species ac-

counts.

This report includes a key to genera and species, and gonopod il-

lustrations to assist in determinations. Accounts are presented for each

milliped species collected, along with synonymies for the two species

previously reported from the region or vicinity. Numeric ratios of or-

ders/families/genera/species (o/f/g/s) and species/genera (s/g) are used

to show diversity and seasonal variation within the Kings Mountain

fauna and to compare it with the faunas of the eastern Piedmont and the

Great Smoky Mountains (Tables 10-12). Comments on seasonal oc-

currence of adults and juveniles are provided in the species accounts and

summarized in Table 10. Noteworthy behavior and gonopodal variation

are discussed for each species where appropriate. Localities are listed for

species collected from fewer than six sites and for three diplopods con-

sidered of Special Concern in North Carolina, as defined in Cooper et al.

(1977). Due to present nomenclatorial confusion and in deference to

current work by other specialists, as explained in appropriate accounts.

4

Marianne E. Filka and Rowland M. Shelley

specific names cannot be assigned for two millipeds and provisional

names are used for two others. The major concern of this study was the

fauna of natural habitats, and synanthropic diplopods were thus incom-

pletely sampled. Additional species that might be discovered in future in-

vestigations are discussed in the final section.

/

The Kings Mountain Region

The Kings Mountain range extends northeastward as a linear ridge

some 26.5 km from the southern tip of Cherokee and York counties,

South Carolina, to the southeastern corner of Cleveland and south-

western part of Gaston counties. North Carolina (Fig. 2). It is bounded

on the east and west by Crowders and Kings creeks, respectively, and sur-

rounded by Piedmont Plateau. Isolated outlying peaks, inselbergs of the

Kings Mountain ridge, continue northeastward approximately 64 km to

Anderson’s Mountain in Catawba County. The bulk of the region is

located about 136 km east of the Blue Ridge Front in Cleveland and Gas-

ton counties, where it covers an area of approximately 3108 hectares. It

consists of four main groups of lowlying peaks separated by gaps. Mean

elevation is 361 m above sea level with maxima of 570 m (the Pinnacle)

and 474 m (Crowders Mountain). Spencer Mountain, a 378 m high in-

selberg of the Kings Mountain ridge, is located about 14.5 km northeast

of Crowders Mountain on the opposite side of Gastonia.

The Kings Mountain geologic belt, composing the range, is a narrow

zone of metamorphosed sedimentary rocks (schist, marble, and

quartzite) of Paleozoic age (Stuckey 1965). The porous nature of this

rock produces a bountiful supply of ground water, and natural springs

and seeps are characteristic of the area (Keith 1931). Soil composition

Kings Mountain Milliped Fauna

5

varies from thick black peaty humus in forested areas, to exposed red

clays and yellow silts on eroded downslopes, to glittering micaquartzite

sand along stream banks and on summits. Xeric scrub forests similiar to

those found in the Blue Ridge characterize these summits, and hardwood

forests, remnants of the previous oak-hickory and beech-maple climaxes,

distinguish relatively undisturbed regions on surrounding lower slopes.

In clear-cut or burned areas, dense stands of Virginia pine, Pinus

virginiana, and shortleaf pine, P. echinata, dominate to the exclusion of

other species. Various pine-hardwood mixtures occur in disturbed areas

throughout the Kings Mountain region (Burney 1974).

MATERIALS AND METHODS

Millipeds were sampled in July and October 1976, and April 1977, to

investigate seasonal variation in faunal composition. Collecting was done

primarily in the contiguous ridge area around Kings and Crowders out-

crops, but four sites around Spencer Mountain also were sampled for

comparison. The South Carolina state line was selected as the southern

boundary, and collecting limits were set in other directions using

topographical and county road maps. Collecting sites, chosen to provide

a maximum diversity of habitats, included pine, mixed pine-hardwood,

and deciduous forests; banks of streams and ponds; seepage areas; bor-

ders of flat meadows; gradual slopes and steep hill terrains; bases, slopes,

and summits of outcrops; and trash dumps. Climatic conditions varied

from hot and dry in July to cool and damp in October and April.

Twenty-five sites were examined during the first trip (July). Five of

these yielded few millipeds because of scant leaf litter, so only the twenty

remaining sites were routinely sampled on all trips. Several new prospec-

tive sites were visited during each ensuing trip. Specimens were collected

from beneath leaf litter, bark of decaying logs, and large rocks, and

preserved in 70% isopropanol. Pine, hardwood, and mixed pine-hard-

wood litter samples were collected for extraction with Berlese funnels.

Notes on color, behavior, and habitat were recorded at each site.

Measurements of length and width in mm were taken with verniei

calipers. Drawings of most gonopods and all other structures were

prepared with the aid of a grid reticle with 0.5 mm squares, but a camera

lucida was used for the smallest gonopods, which were mounted tem-

porarily in glycerine jelly and examined with a compound microscope.

All other structures were examined using a stereomicroscope, with the

specimens immersed in 70% isopropanol and stabilized by cotton. More

than 1000 specimens were examined, some of which were collected in

preliminary studies. All specimens are deposited in the invertebrate

collection of the North Carolina State Museum of Natural History

(NCSM), the invertebrate catalogue numbers of which are indicated in

parentheses with appropriate citations. A single pertinent specimen was

found in the collection of the American Museum of Natural History

(AMNH).

In the species accounts and legends, CR means country road.

6

Marianne E. Filka and Rowland M. Shelley

KEY TO GENERA AND SPECIES OF

KINGS MOUNTAIN DIPLOPODS

1. Body soft, exoskeleton noncalcified; clusters of modified setae

adorning head and tergites; terminal setal tufts present posterior-

ly; males without gonopods; adults less than 3 mm long (Penicil-

ata, Polyxenida, Polyxenidae Polyxenus fasciculatus Say

Exoskeleton hard, calcified; setae normal, scattered, males with

gonopods on 7th or 7th and 8th segments; adults varying in size

but always longer than 3 mm (Helminthomorpha) 2

2. Head reduced; males with 8 pairs of legs preceding gonopods . . 3

Head normal; males with 7 pairs of legs preceding gonopods ... 4

3. Three pairs of ocelli present; paranota absent; segments arched, con-

vex dorsally (Polyzoniida, Polyzoniidae)

Polyzonium strictum Shelley

Ocelli absent; paranota present, bilobed on segment five (Fig. 4);

segments narrow, flattened (Platydesmida, Andrognathidae)

A ndrognathus corticarius CopQ

4. Ocelli present; paranota absent; adults with more than 20 seg-

ments 5

Ocelli absent; paranota present; adults with 19 or 20 segments

(Polydesmida) 16

5. Segments with dorsal longitudinal crests 6

Without this character 9

6. Collum enlarged, hoodlike, covering epicranial region of head . 7

Collum reduced, head completely exposed (Callipodida, Caspiope-

talidae) 8

7. Epiproct trilobed (Fig. 11); adults with 30 segments; adult length

not exceeding 26 mm (Chordeumida, Striariidae)

S triaria sp.

Epiproct entire; adults with more than 30 segments; adults 40-50 mm

long (Spirostreptida, Cambalidae) .... Cambala annulata (Say)

8. Coxal process of gonopod thin, translucent, ensheathing stalk of

telopodite; flagellum absent (Figs. 25-26)

Delophon gear gianum ChdimbQxWn

Coxal process of gonopod thick, opaque, bent laterad apically, not

ensheathing stalk of telopodite; flagellum present (Fig. 20)

Abacion magnum {Loomis)

9. Adults with 28-30 segments (Chordeumida) 10

Adults with more than 30 segments 11

10. Ocelli arranged curvilinearly, 6 per row; adults not exceeding 5 mm

long (Trichopetalidae) Trichopetalum dux {CbdimhQxWn)

Ocelli in triangular patch, 26 per patch; adults longer than 5 mm

(Cleidogonidae) Cleidogona medialis Shelley

11\ Coxae of legs 3-7 of males with lobed extensions (Fig. 16); ocelli

in ovoid patch; gonopods concealed within body; adults large,

robust, 80-100 mm long (Spirobolida, Spirobolidae)

Narceus americanus (Beauvois)

Kings Mountain Milliped Fauna

7

Coxae of pregonopodal legs of males without lobed extensions;

ocelli variable in arrangement; gonopods completely visible or

concealed within body; adults slender, never exceeding 5 mm

long(Julida) 12

1 2. Ocelli arranged linearly (Blaniulidae) Nopoiulus minutus (Brandt)

Ocelli in triangular patch 13

13. Gonopods completely concealed within body; first pair of legs of

male reduced, hooklike, dorsum with 2 yellow longitudinal

stripes containing narrow median black line (Julidae)

Brachyiulus lusitanus Verhoeff

Gonopods visible externally; first pair of legs of male enlarged;

body uniformly gray in color, without stripes (Parajulidae) . 14

14. Epiproct decurved Teniulus sp.

Epiproct extending into straight spine 15

15. Peltocoxites of anterior gonopods with flared, serrate calyx (Fig. 6)

Ptyoiulus impressus (Say)

Calyx of peltocoxites cupped, smooth (Fig. 5)

Ptyoiulus ectenes (Bollman)

16. Midbody metatergites with transverse groove; rim of paraprocts

without setae (Paradoxosomatidae) Oxidus gracilis (Koch)

Midbody metatergites without transverse groove; rim of para-

procts with one pair of setae 17

1 7. Prefemora of legs with ventrodistal spines; gonopod usually bearing

prefemoral process; adults large, robust, color bright yellow-

black or yellow-brown (Xystodesmidae) 18

Prefemora of legs without ventrodistal spines; gonopod without

preformal process; adults slender, coloration otherwise .... 22

18. Gonopods with coxal apophysis (Figs. 55-57) 19

Without this character 20

19. Cranial setae present on frons and epicranium in both sexes; gono-

pods with one prefemoral and two tibiotarsal processes (Fig. 57);

podosterna present

Pachydesmus crassicutis incursus Chamberlin

Cranial setae absent from frons and epicranium in both sexes;

gonopod with or without small prefemoral process; telopodite

broadly curved, falcate in shape (Figs. 55-56); sterna unmodified

Deltotaria lea Hoffman

20. Telopodite of gonopod with irregularly notched expansion along

proximomedial edge; prefemoral process large, extending beyond

tip of telopodite (Fig. 54); membrane of cyphopod enlarged and

folded, protruding through medial portion of aperture.

Croatania catawba Shelley

Without these characters 21

21. Prefemoral process short, blunt, never two-thirds length of telopo-

dite; telopodite curved, with medial flange at midlength (Fig. 60);

adults 40-45 mm long Sigmoria latior (Brolemann)

8 Marianne E. Filka and Rowland M. Shelley

Prefemoral process of gonopod acicular, approximately two-thirds

length of telepodite; telopodite nearly straight, bent slightly

mediodorsad distally, without flange (Fig. 47); adults 28-32 mm

long Boraria St ricta (fix o\Qmdir\x\)

22. Epiproct broad, truncate; adults gray with orange paranota and

middorsal spots (Platyrhacidae)

A uturus erythropygos (Brandt)

Epiproct subtriangular; adults with essentially uniform coloration

(Polydesmidae) 23

23. Adults with 19 segments; metatergites with four rows of small seti-

ferous tubercles Scytonotus granulatus {Sdiy)

Adults with 20 segments; dorsum without setae and distinct rows of

tubercles Pseudopoly desmus branneri (Bollman)

CLASSIFICATION OF KINGS MOUNTAIN DIPLOPODS

CLASS DIPLOPODA

SUBCLASS PENICILLATA

ORDER POLYXENIDA

Family Polyxenidae

Polyxenus fasciculatus Say

SUBCLASS HELMINTHOMORPHA

ORDER POLYZONIIDA

Family Polyzoniidae

Polyzonium strictum Shelley

ORDER PLATYDESMIDA

Family Andrognathidae

Andrognathus corticarius Cope

ORDER JULIDA

Family Blaniulidae

Nopoiulus minutus (Brandt)

Family Julidae

Brachyiulus lusitanus Verhoeff

Family Parajulidae

Ptyoiulus ectenes (Bollman)

Ptyoiulus impressus (Say)

Teniulus sp.

ORDER CHORDEUMIDA

Family Cleidogonidae

Cleidogona medialis Shelley

Family Trichopetalidae

Trichopetalum dux (Chamberlin)

Family Striariidae

S triaria sp.

Kings Mountain Milliped Fauna

9

ORDER SPIROBOLIDA

Family Spirobolidae

Narceus americanus (Beauvois)

ORDER CALLIPODIDA

Family Caspiopetalidae

Abacion magnum (Loomis)

Delophon georgianum Chamberlin

ORDER SPIROSTREPTIDA

Family Cambalidae

Cambala annulata (Say)

ORDER POLYDESMIDA

Family Paradoxosomatidae

Oxidus gracilis (Koch)

Family Polydesmidae

Pseudopolydesmus branneri (Bollman)

Scytonotus granulatus (Say)

Family Platyrhacidae

Auturus erythropygos (Brandt)

Family Xystodesmidae

Boraria stricta (Brolemann)

Croatania catawba Shelley

Deltotaria lea Hoffman

Pachydesmus crassicutis incursus Chamberlin

Sigmoria latior (Brolemann)

SPECIES ACCOUNTS

Polyxenidae

Polyxenus fasciculatus Say, 1821

Polyxenus fasciculatus, a small, pale milliped, was recovered from

pine and mixed pine-hardwood litter using Berlese funnels but was absent

from hardwood litter. More adults and juveniles were taken in July than

in October or April. Previous North Carolina records are from Duke

Forest (Brimley 1938; Wray 1967) and the eastern Piedmont in general

(Shelley 1978). The species is known to range from Long Island through

the southeastern and midwestern United States to Texas (Chamberlin

and Hoffman 1958). The presence of P. fasciculatus in the Appalachian

Mountains is questionable, since Chamberlin and Hoffman (1958) re-

ported it absent or very scarce there.

10

Marianne E. Filka and Rowland M. Shelley

Figs. 3-8. 3, Polyzonium strictum, anterior gonopods, cephalic view. 4,

Andrognathus corticarius, head and first six segments, dorsal view. 5, Ptyoiulus ec-

tenes, anterior gonopods, caudal view, calyx (c) and peltocoxites (p). 6, Ptyoiulus

impressus, anterior gonopods, caudal view, calyx (c) and peltocoxites (p). 7-8

Teniulus sp. 7, anterior gonopods, caudal view. 8, left posterior gonopod, lateral

view. Scale line = 0.1 mm.

Kings Mountain Milliped Fauna

11

Polyzoniidae

Polyzonium strictum Shelley, 1976

Fig. 3

Yellow adults of P. strictum were taken from beneath the bark of

decaying logs in July and October. A large number of juveniles was ex-

tracted from mixed pine-hardwood Berlese samples in October. In North

Carolina P. strictum ranges from the mountains to the inner Coastal

Plain, and it also occurs in the mountains of Virginia (Shelley 1976a).

Andrognathidae

Andrognathus corticarius Cope, 1968

Fig. 4

This slender, cream-colored diplopod typically occurs beneath the

bark of decaying pine logs (Shelley 1978), a habitat that was examined ex-

tensively in the study area. Only two specimens were encountered, how-

ever, both in July from a single log at a Spencer Mountain site. Cham-

berlin and Hoffman (1958) reported ^4. corticarius from western Virginia,

southeastern Kentucky, Tennessee, Georgia, and northern Florida.

Gardner (1975) examined material from Graham and Madison counties

in the Appalachian Mountains of North Carolina, and Shelley (1978)

noted that the species was more prevalent in the southern subregion of

the eastern Piedmont than north of the Deep-Cape Fear Rivers.

Locality. Gaston Co. — 7.2 km NE Gastonia, along CR 2200, 2.2 km

SW jet. NC Hwy. 7, base of Spencer Mountain, 2 9,1 July 1976, M.

Filka and W.W. Thomson (NCSM A 1032).

Blaniulidae

Nopoiulus minutus (Brandt, 1841)

This narrow brown milliped occurs in habitats similar to those of A.

corticarius. Four immature specimens were encountered, in July and Oc-

tober, but no adults were found. The dearth of specimens in the Kings

Mountain region contrasts markedly with the abundance of the species

farther east in the fall zone region of North Carolina, where it also occurs

in summer and autumn (Shelley 1978). Nopoiulus minutus is widespread

east of the Great Plains, ranging from Illinois, Indiana, and Ohio south

to Georgia (Enghoff and Shelley 1979).

Localities. Cleveland Co. — 9.1 km SE Kings Mountain (town), along

CR 2286, 1.6 km S jet. CR 2283, 8 July 1976, 1 juv., M. Filka and W.W.

Thomson (NCSM A 1973); and 1.9 km SW Kings Mountain (town).

12

Marianne E. Filka and Rowland M. Shelley

along 1-85, jet. NC Hwy. 161, 3 juvs., 18 October 1976, M. Filka and G.

Wicker (NCSM A2197).

Julidae

Brachyiulus lusitanus Verhoeff, 1898

Brachyiulus lusitanus is easily identified by its characteristic dorsal

color pattern — two pale longitudinal stripes surrounding a narrow black

mid-dorsal line. Introduced from Europe, B. lusitanus has been reported

from developed areas of North America as far south as the “Triangle”

(Raleigh-Durham-Chapel Hill) region of North Carolina, where it was

erroneously reported as B. pusillus (Leach) by Shelley (1978). However,

three females were encountered during this study, all in April, under

debris at a public campsite. The Kings Mountain region thus becomes

the southernmost known locality for B. lusitanus in the New World.

Locality. Cleveland Co. — 1.9 km SE Kings Mountain (town), jet.

1-85 and NC Hwy. 161, 3 9 , 10 April 1977. M. Filka.

Parajulidae

Ptyoiulus ectenes (Bollman, 1888)

Fig. 5

Juvenile and female Ptyoiulus are unidentifiable to species. Those

found associated with males of a single species (all only in October) were

adjudged to be that species and are so shown in Table 3. Those collected

without associated males (all only in July) are tabulated by genus in

Table 3. No juveniles or females were found with males of both species at

a single collecting site. These identification problems may have influ-

enced the apparent seasonal distribution patterns of both species of

Ptyoiulus in the region, although examination of Table 3 reveals similar

July and October patterns for the two. Adults of P. ectenes were most

numerous in October but also occurred in April; immatures were found

only in July and October. These data suggest that P. ectenes reproduces

during fall and spring, and juveniles mature the following summer and

fall. All specimens were collected from deciduous litter. Gonopods of the

31 males were examined but no variation was apparent.

This species was reported from the fall zone region as Ptyoiulus sp.

by Shelley (1978) who declined to assign a specific name in deference to

studies being conducted by the late Dr. Nell B. Causey. The oldest

available specific name is in doubt, but that of Bollman is used tentatively

here since it is one of the earliest names and the first applied to specimens

from North Carolina. However, there is some question as to whether

ectenes is referable to Ptyoiulus, since Bollman (1887) remarked that the

Kings Mountain Milliped Fauna

13

species differed in its “slender body and peculiar form of the male geni-

talia,” which he neither illustrated nor described verbally in his descrip-

tion. Unfortunately, the male from the type series is lost, but collections

made by Shelley in and around the type locality — Chapel Hill, Orange

County, North Carolina — have produced male parajulids whose

gonopods are virtually identical to that illustrated in Figure 5. This

suggests that ectenes may be the species under consideration here, but a

final judgment can only result from a comprehensive revision of

Ptyoiulus in which female cyphopods are studied. Wray (1967) may have

been correct in transferring ectenes to Aniulus, and this combination may

be a senior subjective synonym of A. orientalis Causey, the only other

parajulid known from the “Triangle” region of the state.

Ptyoiulus impressus (Say, 1821)

Fig. 6

Adults of P. impressus were abundant in October and absent in April

and July; juveniles were taken only in July and October. Thus, P. im-

pressus appears to have a slightly different life history from that of P. ec-

tenes, with summer growth and maturation preceding fall reproduction.

Both species are uniformly gray and both were found in deciduous forest

litter. Adult P. ectenes are slightly smaller and less robust than adult P.

impressus, although this difference can be misleading and should not be

the sole criterion for identification. The most reliable character is the

configuration of the calyx of the peltocoxites of the anterior gonopod

(Figs. 5-6, c, p,), which is flared and serrate distally in P. impressus and

cupped and smooth in P. ectenes. As with its congener, the gonopods of

P. impressus were essentially uniform.

Ptyoiulus impressus ranges from the northeastern United States west

to Indiana and south to western North Carolina and Kentucky (Cham-

berlin and Hoffman 1958). Shelley (1978) deleted this species from the

eastern Piedmont fauna, stating that it was known definitely only from

the mountains and western Piedmont. The Kings Mountain region is the

easternmost authentic locality in North Carolina.

Teniulus sp.

Figs. 7-8

This uniformly gray species is similar in appearance to both species

of Ptyoiulus, but is distinguished by the decurved epiproct. Adults were

collected in October from moist deciduous leaf litter in association with

both species of Ptyoiulus. No juveniles were found.

The genus currently contains only two species, T. parvior and T.

setosior, both described by Chamberlin (1951) from Gatlinburg, Sevier

County, Tennessee. Gatlinburg is about 200 km west-northwest of the

14

Marianne E. Filka and Rowland M. Shelley

Kings Mountain region, and such wide geographic separation suggests

that the forms in the two areas are not conspecific. If true, the Kings

Mountain species is undescribed. However, drawings by Chamberlin

(1951) accompanying the descriptions of the Tennessee species are un-

clear and were prepared from different views, which prevents close com-

parisons. Examination of the type specimens of both species by Shelley

revealed that males and/or gonopods were absent. These two species may

be synonymous, but adult males are needed before their identities can be

determined and an accurate statement can be made on the status of the

Kings Mountain form.

Locality. Gaston Co. — 9.9 km S Bessemer City, along CR 1112, 0.3

km E jet. CR 1125, 6, 5 9, 17 October 1976, M. Filka and G. Wicker

(NCSM A2193).

Cleidogonidae

Cleidogona medialis Shelley, 1976

Figs. 9-10

The Kings Mountain region is the second known locality for this

light brown chordeumid, whose range is now extended some 117 km

from Blowing Rock, Watauga County, North Carolina (Shelley 1976b).

The single male and female taken during this study and all those from the

type locality were collected in October, suggesting autumnal maturation.

Juveniles would therefore be expected in the summer; although none

were encountered in July, they may be present in late August or early

September.

The medial processes of the gonopods of the Kings Mountain speci-

men were more jagged than those of the holotype (Shelley 1976b, Fig.

10), which conforms to known variation in the species. A single oversized

telopodite variant was reported in one male paratype, but those of the

Kings Mountain male were as illustrated for the holotype. No other

gonopodal variations were observed. Two additional records are cited

below from material that has recently become available.

Localities. Gaston Co. — 9.9 km SE Bessemer City, along CR 1126,

0.8 km SW jet. CR 1 1 13, 9, 16 October 1976, M. Filka and G. Wicker

(NCSM A2770); and 9.9 km SW Bessemer City, along CR 1104, 1.3 km

W jet. CR 1115, r7, 17 October 1976, M. Filka and G. Wicker (NCSM

A2771). Davidson Co. — Boone’s Cave State Park, c?, 6 November 1976,

R.M. Shelley (NCSM A 1434). Watauga Co. — 16 km NE Deep Gap, o',

9,17 October 1965, J. & W. Ivie (AMNH).

Kings Mountain Milliped Fauna

15

Figs. 9-19. 9-11, Chordeumida. 9-10, Cleidogona medialis. 9, anterior gonopods,

cephalic view, sternum broken in dissection. 10, left anterior gonopod, lateral

view. 11, Striaria sp., epiproct, dorsal view. 12-19 Narceus americanus. 12, an-

terior gonopods, cephalic view. 13, left posterior gonopod, cephalic view. 14-15,

distal portions of left posterior gonopods, cephalic views, showing variation in

prefemoral endite (pe). 16, coxae and lobes of legs 3-7 of male, ventral view. 17-

19, coxae and prefemora of left third legs of females, cephalic views, showing

variation in coxal lobes. Scale line = 0.1 mm.

16

Marianne E. Filka and Rowland M. Shelley

Trichopetalidae

Trichopetalum dux (Chamberlin, 1940)

The specific identification of this milliped is tentative and based

solely on previous North Carolina records. Only one female was collec-

ted, discovered in berlesate from a deciduous litter sample taken in Gas-

ton County in April. This is the only species of Trichopetalum known

from North Carolina, where it was previously reported from Duke

Forest (Chamberlin 1940a; Wray 1967) and Chatham County (Shelley

1978). Positive identification of the Kings Mountain species awaits the

collection of males.

Locality. Gaston Co. — 8 km NE Gastonia, base of Spencer Moun-

tain, along CR 2200, 1.7 km SW jet. CR 2003, 9 , 10 April 1977, M. Filka

(NCSM A2185).

Striariidae

S triaria sp.

Fig. 11

Striaria is readily distinguished from the other chordeumids by its

enlarged collum, crested segments, and trilobed epiproct. One adult

female and two juveniles were taken in Gaston County in October and

April, respectively. The adult female and one juvenile were brown with a

pale white collum, while the other juvenile was uniformly brown. Three

species of Striaria are known from North Carolina: two with a white

collum — S. zygoleuca Hoffman, from Highlands, Macon County (Hoff-

man 1950; Wray 1967), and an undescribed form from High Falls, Moore

County (Shelley 1978) — and one with a brown collum, S. causeyae

Chamberlin, from eight counties in the eastern Piedmont (Shelley 1978).

The presence of differently colored collums on the Gaston County

material suggests the presence of at least two species, but again the ab-

sence of males precludes final determinations. For the purposes of this

report, therefore, only one species is considered.

Localities. Gaston Co. — 4.0 km S Bessemer City, along CR 1 125, 0.2

km S jet. U.S. Hwy. 74^29, 9 , 17 October 1976, M. Filka andG. Wicker

(NCSM A2215); juv., 9 April 1977, M. Filka (NCSM A2219); and 6.4 km

SE Bessemer City, along CR 1103, jet. CR 1112, juv., 9 April 1977, M.

Filka (NCSM A2202).

Spirobolidae

Narceus americanus (Beauvois, 1805)

Figs. 12-19, Tables 1-2

Body coloration of N. americanus is dark brown with the head, legs,

and body segments bordered in red. As indicated in Table 3, this species

is common in the Kings Mountain region, and adults and juveniles were

Table 1. Comparison of the number of segments, clypeal and labral setae, ocelli, and formula value for the Kings Mountain spirobolid (KM),

Narceus annularis (Nan), and N. americanus (Nam). Data for Narceus from Keeton (1960). d = difference from the Kings Mountain

spirobolid.

Kings Mountain Milliped Fauna

17

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18

Marianne E. Filka and Rowland M. Shelley

collected from a diverse array of habitats on each trip. Continuous

breeding and maturation throughout the year is suggested by these data.

Two species of Narceus occur in North Carolina; N. americanus ,

known to range throughout the southeastern United States, and N.

annularis (Rafinesque), known from the northeastern and midwestern

United States (Chamberlin and Hoffman 1958). Both are reported from

the mountains (Keeton 1960) and eastern Piedmont (Shelley 1978) of

North Carolina.

In his monograph on the Spirobolidae, Keeton (1960) distinguished

between N. annularis and N. americanus by a formula computing values

based on somatic features (see footnote Table 1 for explanation) and by

comparison of gonopodal characters. He found that differences were dif-

ficult to define due to overlap of characters. Consequently, identifica-

tions are difficult and distinctions between the species are vague, facts

corroborated by the Kings Mountain material.

Thirty-four adults were collected but only 22 of these, 16 males and 6

females, were in sufficiently good condition for detailed study. As shown

in Tables 1-2, the number of segments, clypeal setae, and labral setae of

the Kings Mountain spirobolid are closer to the values for N. americanus:

but the mean number of ocelli is closer to that for N. annularis. Mean

length and width, the length/width ratio, and the formula value, also are

nearer the figures for N. americanus.

The distal prefemoral endite of the posterior gonopod, normally

rounded in N. annularis and acute in N. americanus, is rounded in 72% of

the specimens (Figs. 13-14, pe) and acute in the rest (Fig. 15, pe).

Likewise, the cephalic groove on the coxal lobes of the third pair of legs

of males, a characteristic of N. americanus, is absent from all Kings

Mountain males (Fig. 16). Furthermore, the third coxal lobe of adult

females, enlarged in N. americanus but only slightly produced ventrad in

N. annularis, conforms to the latter condition in all but one specimen

(Figs. 17-19). The Kings Mountain spirobolid therefore could be iden-

tified as either species of Narceus depending upon the character used, and

the question becomes one of the relative importance of the characters.

Although gonopods are the most important taxonomic character in

the Diplopoda, many genera show few specific gonopodal differences

whereas there is wide variation in body forms. This appears to be the

situation in Narceus, and the few gonopodal similarities between the

Kings Mountain spirobolid and N. annularis do not seem to outweigh the

close agreement of nearly all the somatic features with those of N.

americanus. Thus, the Kings Mountain spirobolid is identified as N.

americanus. The great variability of the somatic features of Narceus, as

demonstrated by Keeton (1960) and our tables, indicates a need for

reassessment of the status of the two nominal eastern species. Such a

study might show them to be conspecific.

Kings Mountain Milliped Fauna

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20

Marianne E. Filka and Rowland M. Shelley

Caspiopetalidae

Abacion magnum (Loomis, 1943)

Figs. 20-24

Abacion magnum is a crested diplopod, brown with a light middorsal

stripe. It was one of the few species found in drier parts of deciduous,

pine, and mixed leaf litter. Juveniles of the two callipodids, A. magnum

and D. georgianum, could not be identified to species, but all were found

with adult males of a single species and, as with Ptyoiulus, were identified

as such. Adults of A. magnum were taken during all three months, while

juveniles were collected only in July and October. This implies that

reproduction and maturation occur throughout the year. The coxal

processes of the gonopods of the eastern Piedmont specimens varied in

degree of apical serration and configuration of the midlength angulation

(Shelley 1978). These structures were found to vary similarly in the Kings

Mountain specimens (Figs. 21-24). No other gonopodal variations were

detected. This species has been collected in Macon and Transylvania

counties in the North Carolina mountains (Hoffman 1950) and in eight

counties of the eastern Piedmont (Shelley 1978).

Delophon georgianum Chamberlin, 1943

Figs. 25-26

This callipodid is similar in coloration to A. magnum but is smaller

and differs in the structure of the gonopod. Abacion magnum has a

flagellum and a serrate coxal process lateral to the telopodite (Fig. 20, fl.

cp). Delophon georgianum lacks the flagellum, and its coxal process

ensheaths the stalk of the telopodite (Figs. 25-26, cp) (Shelley 1979a). Lit-

tle gonopodal variation was found in this study. Like A. magnum, adults

of D. georgianum were taken on all three trips, but only one juvenile was

encountered, in April. Delophon georgianum was typically found in

moister habitats than Abacion.

This species has been previously reported from the mountains of

North Carolina as D. carolinum Hoffman (Hoffman 1950; Chamberlin

and Hoffman 1958; Wray 1967). However, Shelley (1979a) concluded

that this binomial was a synonym of D. georgianum. The Kings Moun-

tain population is disjunct from that occurring in the Appalachians, and

no specimens have ever been taken in the intervening lowlands. Shelley

speculated that the Kings Mountain population might be a Pleistocene

relict that has survived due to a slightly cooler microclimate afforded by

the peaks and coves of the area. Hardin and Cooper (1967) concluded

that this was the explanation for the occurrence of several disjunct pop-

ulations of montane plants, most notably Tsuga canadensis L. and Pinus

strobus L., in the Piedmont.

Kings Mountain Milliped Fauna

21

Figs. 20-27. 20-26, Callipodida. 20-24, Abacion magnum. 20, left gonopod, caudal

view, coxal process (cp), flagellum (fl). 21-24, coxal processes, lateral views, of

four specimens from the Kings Mountain Region. 25-26, Delophon georgianum.

25, left gonopod, lateral view, coxal process (sheath) (cp). 26, left gonopod,

caudal view, coxal process indicated. 27, Cambala annulata, left posterior

gonopod, lateral view. Scale line = 0.1 mm.

22

Marianne E. Filka and Rowland M. Shelley

Cambalidae

Cambala annulata (Say, 1821)

Fig. 27

Shelley (1978) reported that C. annulata seemed to prefer cooler tem-

peratures, and this was apparent in the Kings Mountain region where the

dark purple adults were abundant in both April and October. Only one

adult, a female, was collected in July. Juveniles were taken in July and

October but not in April. Hoffman (1958) indicated that individuals of

this species were usually found grouped together, but in the Kings Moun-

tain region this was true only of females; adult males were always found

alone. All stages were collected from moist humus. Hoffman found no

gonopodal variation in material from high elevations, and Shelley

(1979b) noted homogeneity in the gonopods of C. annulata throughout

its range. This was evident in the Kings Mountain material, as the struc-

tures were virtually uniform. Cambala annulata has been reported from

the northeastern and central subregions of eastern Piedmont North

Carolina (Shelley 1978), and its distribution in the Appalachian Moun-

tains was illustrated by Hoffman (1958).

Paradoxosomatidae

Oxidus gracilis (Koch, 1847)

Causey (1943) reported nearly year around oviposition by O. gracilis

under favorable conditions in a Durham County greenhouse, and Shelley

(1978) collected fifth instar juveniles (adults are the seventh instar) in Oc-

tober from William B. Umstead State Park in the eastern Piedmont. The

preponderance of juveniles in October and April and of adults in July

and October in the Kings Mountain region suggests that maturation oc-

curs in the fall and spring and breeding in the late summer and early fall.

White juveniles often populated several square meters of deciduous leaf

litter, and shiny black adults also were common. Oxidus gracilis is nearly

worldwide in distribution, and was introduced into the United States

from the East Indies via imported soil in greenhouses (Causey 1943).

Polydesmidae

Pseudopolydesmus branneri (Bollman, 1887)

Figs. 28-40

Pseudopolydesmus branneri is the sole representative of its genus in

the Kings Mountain region; neither P. collinus Hoffman nor P. serratus

Kings Mountain Milliped Fauna

23

(Say), both of which occur in the eastern Piedmont (Shelley 1978), were

encountered during this study. This species has been previously reported

from Rutherford, Wilkes and Alexander counties in the western Pied-

mont, as well as the mountains and eastern Piedmont (Shelley 1978), so

its occurrence in the Kings Mountain region was expected. Juveniles and

adults were collected in April and October, but none were found in July.

Adults were dull reddish brown with light brown paranoia, similar to

Richmond County specimens (Shelley 1978). Material from the two areas

was also similar in length.

Past discussion of gonopodal variation in the Polydesmidae has been

hampered by the absence of a standardized nomenclature for the spines,

branches, and other projections of the posterior faces of the telopodites.

Hoffman (1974) devised a labeling system based upon orientation of

these processes, with letters “m” and “e” designating mesial and ectal

position, respectively, and numbers indicating position relative to the

base of the telopodite, the most proximal designated by 1. Thus, in P.

branneri the four mesial processes are labeled ml, m2, m3, and m4; the

four ectal processes are el, e2 + 3 (indicating that they share a common

pedicel), and e4 (Figs. 28-29). Examination of the left gonopods of 51

specimens collected from the Kings Mountain region revealed consid-

erable gonopodal variation.

The only evident variation in the mesial processes involved suppres-

sion and division of m4. Nearly three-fourths of the individuals examined

had a normal m4 lobe with a projecting setaceous shoulder (Figs. 28, 30,

34); the remaining individuals had a reduced or vestigial m4, with a

shoulder lacking setae (Figs. 31-33). In 75% of the specimens, m4 con-

sisted of a large lobe contiguous with a smaller shoulder (Figs. 28, 30-33).

This lobe was divided into two separate processes in the remaining

specimens (Fig. 34). The other mesial processes, ml-m3, were virtually

uniform, although one individual lacked m2 (Fig. 35).

The most variable ectal process, el, displayed four configurations;

spiniform (Figs. 28, 36) in 31% of the males; reduced (Fig. 37) in 29%;

vestigial (Fig. 38) in 18%; and absent (Fig. 39) in 18%. Two individuals

(4% of the males) carried a vestigial secondary spine distal to a reduced el

spine (Fig. 40). The other ectal processes, e2+3 and e4, were uniform.

Terminal macrosetae of the telopodite varied in abundance and dis-

tribution. Thirty-one percent of the males had numerous macrosetae oc-

curing from just distal to m4 to the telopodite tip (Figs. 28-30, 33-34), and

fifty-one percent carried fewer macrosetae distributed from e4 to the tip

(Fig. 31). Most remaining individuals (14%) possessed very few mac-

rosetae, occurring only apically on the telopodite (Fig. 35). Two in-

dividuals (4%) lacked terminal macrosetae (Fig. 32).

Hoffman (1974) reported that m2, m3, e2, and e3 were the most

variable processes in P. branneri, but these were found to be the most

stable in the Kings Mountain population, where most of the variation in-

volved m4, el, and the terminal macrosetae. Many combinations of these

24

Marianne E. Filka and Rowland M. Shelley

Figs. 28-41. Polydesmidae. 28-40, Pseudopoly desmus branneri. 28, left gonopod,

medial view. 29, the same, lateral view, medial processes (ml-m4), ectal processes

(el-e4), endomerite (end), terminal macrosetae (tm). 30-34, distal ends of

telopodites, medial views, showing variation of m4 and terminal macrosetae. 35,

distal half of telopodite, medial view, showing absence of process m2. 36-40, en-

domerite regions of telopodites, medial views, showing variation of process el.

41, Scytonotus granulatus, left gonopod, medial view. Scale line = 0.1 mm.

Kings Mountain Milliped Fauna

25

gonopodal variants occurred, and there was no correlation between

them. The length of the patch of terminal macrosetae and the configura-

tion of m4 and el varied independently, and seem to be controlled by dif-

ferent genes.

These findings greatly expand current knowledge of variation for the

species and illustrate the degree of variability that may occur within a

local population. To Hoffman’s (1974) characterization of P. branneri

may now be added the occasional appearance of a new process, the

secondary el spine, the division of a single process into separate compo-

nents, m4 lobe and shoulder, and the occasional loss of all terminal

macrosetae.

Hoffman (1974) described P. collinus as differing from P. branneri in

the absence of m3 and either the absence or vestigial condition of el. In

the Kings Mountain specimens, el and/or m3 were present on all

specimens, although el varied considerably in size. Consequently, only

one species, P. branneri, is represented by this material.

Scytonotus granulatus (Say, 1821)

Fig. 41

This species was rare in the Kings Mountain region. Isolated brown

adults were found in April and October, and white juveniles were taken

in July. Both were found in moist humus. The widespread occurrence of

the species in western North Carolina and several other states was noted

by Hoffman (1962), and Shelley (1978) reported additional localities in

eastern Piedmont North Carolina.

Platyrhacidae

Auturus erythropygos (Brandt, 1841)

Figs. 42-46

Adults of A. erythropygos exhibit striking body coloration, with each

blue-gray metatergite bearing a bright orange middorsal spot and orange

paranota. Juveniles, though lighter, have a similar pattern. All stages

were collected from under bark of decaying deciduous logs, in humus un-

der logs, or in associated bark litter. Both adults and juveniles were most

abundant in October, but the species was quite common in April and

July.

Flattened, round molting chambers, built under the bark of logs in-

habited by A. erythropygos, were observed on each collecting trip (Figs.

45-46). They are constructed of cemented wood particles and provide

protection from desiccation and predation during intermolts. The dimen-

sions were proportional to the inhabitant’s size, the largest being 20-22

mm diameter. An adult or juvenile accompanied by cast exuvium was

26

Marianne E. Filka and Rowland M. Shelley

seen in each chamber. The exoskeleton of newly molted individuals was

whitish and incompletely sclerotized.

This species was unknown from North Carolina until reported from

Northampton County by Shelley (1978), who listed it as georgianus

Chamberlin, now considered a junior synonym.

Figs. 42-54. 42-46, Autunis erythropygos. 42, left gonopod, medial view. 43, the

same, lateral view. 44, distal end of telopodite, cephalic view. 45-46, molting

chamber. 45, side view in situ on log, bark lifted. 46, top view. 47-53, Boraria

stricta. 47, left gonopod, medial view. 48-51, distal halves of prefemoral processes

of left gonopods, medial views. 52-53, molting chamber. 52, side view in situ,

attached to plant roots. 53, top view. 54, Croatania catawba, telopodite of left

gonopod, medial view. Scale line = 0.1 mm.

Kings Mountain Milliped Fauna

27

Xystodesmidae

Boraria stricta (Brolemann, 1896)

Figs. 47-53

The color of B. stricta, black with yellow paranota, is typical of most

xystodesmid species in the Kings Mountain region. Adults were most

abundant in April, and juveniles were common in October and April.

Large colonies were discovered in wet mud-clay soils lining the banks of

streams throughout the region. Individuals often were captured in tun-

nels beneath shallow layers of detritus. Round molting chambers, simi-

lar to those described for this species by Hoffman (1965), were observed

in the vertical shafts of several tunnels in April (Figs. 52-53). Each cham-

ber was formed of clay attached to exposed plant roots, and inhabited by

a newly molted milliped with its cast exuvium. As can be seen by compar-

ing illustrations (Figs. 45-46, 52-53), the molting chamber of B. stricta is

spherical with an apical “chimney” and is attached at its base, whereas

that of A. erythropygos is round, flattened in a vertical plane, and at-

tached at both ends. These distinctions reflect the different biotopes in-

habited by the species.

Hoffman (1965) reported that the known range of B. stricta

coincided closely with the southern section of the Blue Ridge physi-

ographic province and predicted only slight extensions at the northern

and southern extremities. Discovery of the species in the Kings Mountain

region represents an extension of slightly less than 64 km east into the

Piedmont Plateau. The species was not reported by Shelley (1978) from

the fall zone region, and extensive investigations in the Uwharrie Moun-

tains also have failed to produce it. Hence, the Kings Mountain popula-

tion is the easternmost known and is probably peripheral. Specimens also

have been collected from a number of other Piedmont localities in the

past eight years by Shelley, and since the Piedmont is geologically and

climatically distinct from the southern Appalachians, material from the

entire range was examined to determine if recognition of geographic

races was warranted. Hoffman (1965) noted the homogeneity of B. stricta

gonopods, with only slight differences detected. The lobes of the distal

subhastate end of the telopodite varied in size relative to each other, and

the degree of bending at midlength of the telopodite and apically on the

prefemoral process varied. These differences were scattered and inconsis-

tent, not conforming to any geographic pattern. The prefemoral proc-

cesses of seven percent of males in the Kings Mountain population

(including material from York County, South Carolina), however, were

apically bifurcate (Figs. 47-48, 51), a condition never before reported for

either the genus or species. These bifurcate males were intermixed with

normal individuals, although there were differences in the apical pre-

femoral bend of the latter (Figs. 49,50).

In summary, no significant geographical variation was observed in

Marianne E. Filka and Rowland M. Shelley

28

B. stricta, and the homogeneity noted previously by Hoffman (1965) also

applies to Piedmont populations. The bifurcate prefemoral process is

new, however, and its occurrence solely in the Kings Mountain popula-

tion may represent a peripheral population effect. This occurs in too

small a sample of the Kings Mountain population, however, to justify

taxonomic recognition. The known range of B. stricta is expanded con-

siderably to include Gaston and Cleveland counties. North Carolina, and

York (Kings Mountain State Park) and Spartanburg (Croft State Park)

counties. South Carolina.

Croatania catawba Shelley, 1977

Fig. 54

Croatania catawba Shelley, 1977:306, Figs. 1-2, 7, 11-12, 16.

Croatania catawba was one of the few millipeds encountered pri-

marily in July; only three adults were found in April and October. The

preference of species of Croatania for hot, dry conditions was discussed

by Shelley (1977), who also presented a description of the habitat at the

type locality in Cleveland County. Individuals collected during the

present study, however, were taken from cool, moist seepage areas under

deciduous leaf piles and from under large, decaying deciduous logs. As

reported by Shelley (1977), adults were typically black with lemon yellow

paranota and a variable yellow stripe along the anterior edge of the

collum. One female displayed an orange tinted collum stripe similar to

that reported by Shelley for two Union County, South Carolina

specimens. No significant gonopodal variation was discerned.

Shelley (1977) suggested that the distribution of C. catawba in North

Carolina might be associated with the Kings Mountain range, which ex-

tends northeastward through a series of hills and ridges to Anderson

Mountain in Catawba County. Except for one Lincoln County specimen

taken in 1952, however, the milliped has not been collected in North

Carolina outside the contiguous ridge portion of the range in Cleveland

and Gaston counties. Croatania catawba is thus essentially restricted to

this small area in North Carolina, and therefore is considered to be a

species of special concern in the state, as defined by Cooper et al. (1977).

Localities. Cleveland Co. — 9.3 km S Kings Mountain (town), along

CR 2245, 0.2 km N jet. CR 2288, 9 cJ, 5 9, 16 September 1975, R.M.

Shelley and J.C. Clamp (NCSM A450) TYPE LOCALITY; 1.9 km SW

Kings Mountain (town), along 1-85, jet. NC Hwy. 161,9 , 10 April 1977,

M. Filka (NCSM A1040), d^, 9, 7 July 1976, M. Filka and W.\Y. Thom-

son (NCSM A1048), and 5 o’, 9 , 6 juvs., 10 July 1976, M. Filka and

W.W. Thomson (NCSM A 1049). Gaston Co. — 7.7 km SW Gastonia,

along CR 1 131, 0.2 km NW jet. CR 1133, 9 , 9 July 1976, M. Filka and

W.W. Thomson (NCSM A 1340); and 7.2 km S Bessemer City, along CR

1 125, jet. CR 1 106, 9, 16 October 1976, M. Filka and G. Wicker (NCSM

A1418).

Kings Mountain Milliped Fauna

29

Deltotaria lea Hoffman, 1961

Figs. 55-56

Deltotaria lea Hoffman, 1971:33, Figs. Ic, 3a.

Only ten specimens of D. lea, all adults, were collected in this study,

all during the cooler weather of April and October. No juveniles were en-

countered. The original description by Hoffman (1961) was based on a

single preserved male, and specimens from the Kings Mountain region

supplement knowledge of the species in live coloration and gonopodal

variation. Adults are black with yellow paranota and broad yellow

stripes along the caudal edges of the metaterga and the anterior edge of

the collum. All specimens display a metallic sheen quite unlike the glossy

surfaces of species like C. catawba. Gonopodal variations included

presence or absence of a minute prefemoral process, dimension of the dis-

tal end of the telopodite, and diameter of the telopodite arc (Figs. 55-56).

In North Carolina, D. lea is known to be concentrated in the Kings

Mountain region. Like C. catawba, it also was collected in Lincoln

County in the 1950’s but has not been found there recently. Thus, in

North Carolina D. lea also may be restricted to the Kings Mountain

region and is considered of special concern in the state. The species has

been collected from three South Carolina counties by Shelley, as reported

below.

Localities. NORTH CAROLINA. Cleveland Co. — 1.9 km SE Kings

Mountain (town), along 1-85, jet. NC Hwy. 161, d', 9,18 October 1976,

M. Filka and G. Wicker (NCSM A2284); 6.5 km SW Kings Mountain

(town), along CR 2245, 0.3 km N jet. CR 2283, 6', 9, 30 April 1976,

R.M. Shelley (NCSM A720); 9.3 km S Kings Mountain (town), along

CR 2245, 0.2 km N jet. CR 2288, cd, 30 April 1976, R.M. Shelley (NCSM

A723); 6.2 km S Kings Mountain (town), along NC Hwy. 161, 0.3 km S

jet. CR 2354, 2 d, 30 April 1976, R.M. Shelley (NCSM A725); and 6.2

km SE Kings Mountain (town), along NC Hwy. 161, 1.6 km S jet. CR

2289, d', 8 April 1977, M. Filka (NCSM A2204). Gaston Co. — 2.4 km

NE Crowders Mountain, along CR 1122, 0.2 km N jet. CR 1131, d, 18

April 1976, M.R. and J.E. Cooper (NCSM A733); and 13.6 km SW Gas-

tonia, along CR 1104, 0.5 km S jet. CR 1115, d , 29 April 1976, R.M.

Shelley (NCSM A713).

SOUTH CAROLINA. York Co.— Kings Mountain State Park, d,

30 April 1977, R.M. Shelley (NCSM A 1479). Cherokee Co. — 7.4 km SE

Blacksburg, along SC Hwy. 5, 0.5 km S jet. SC Hwy. 68, d, 10 May

1977, R.M. Shelley (NCSM A 1480). Chester Co. — 17.9 km NE Chester,

along SC Hwy. 32, 1.1 km N jet. SC Hwy. 46, d, 9 , 1 May 1977, R.M.

Shelley (NCSM A 1498); and 19.8 km W Lowrys, along SC Hwy. 9 at

Broad River, d, 1 May 1977, R.M. Shelley (NCSM A1501).

Pachydesmus crassicutis incursus Chamberlin, 1939

Figs. 57-59

This is the largest polydesmoid milliped in North Carolina and is

30

Marianne E. Filka and Rowland M. Shelley

Figs. 55-64. 55-56, Deltotaria lea, left gonopods, ventrolateral views, coxal

apophysis (ca). 57-59, Pachydesmus crassicutis incursus. 57, telopodite of left

gonopod, lateral view, coxal apophysis (ca), primary tibiotarsus (ptt), second

tibiotarsus (stt). 58-59, distal ends of secondary tibiotarsi, lateral views. 60-64,

Sigmoria latior. 60, telopodite of left gonopod, medial view. 61-62, distal ends of

telopodites, medial views. 63-64, prefemoral processes, medial views. Scale line =

0. 1 mm.

known definitely only from the Kings Mountain region (Shelley and

Filka 1979). It is approximately 7 cm long, and dusky brown with yellow

paranoia. As with C. catawba, most specimens were found in July, con-

centrated in wet spots such as seepage areas. Shelley and Filka presented

illustrations of gonopodal variation and showed changes in body dimen-

sions that occur with latitude. Individuals of both sexes are larger and

Kings Mountain Milliped Fauna

31

more brightly colored in the Kings Mountain region than farther south in

South Carolina, probably a reflection of more favorable environmental

conditions in the former area.

Gonopod comparisons revealed variation in primary and secondary

tibiotarsi (Figs. 57-59, ptt, stt). As reported by Shelley and Filka, the sub-

terminal process of the secondary tibiotarsus was pointed, blunt, or ab-

sent. Since, in North Carolina, P.c. incursus is apparently restricted to the

Kings Mountain region, it is considered to be endangered within the

state, as defined in Cooper et al. (1977).

Localities. Cleveland Co. — 6.6 km SW Kings Mountain (town) along

CR 2245 at Dixon Branch Creek, 0.8 km NW jet. CR 2283, 2 c?, 2 9, 1

juv., 16 August 1975, R.M. Shelley and J.C. Clamp (NCSM A537); 9.3

km S Kings Mountain (town), along NC Hwy. 245, 0.2 km N jet. CR

2288, 3 9, 16 August 1975, R.M. Shelley and J.C. Clamp (NCSM A541);

9.1 km SW Kings Mountain (town), along CR 2283, 1.3 km NE jet. NC

Hwy. 216, <? , 8 July 1976, M. Filka and W.W. Thomson (NCSM

A 1060); and 4.8 km S Kings Mountain (town), along CR 2289, 1.0 km W

NC Hwy. 161, 2 9, 18 October 1976, M. Filka and G. Wicker (NCSM

A2239). Gaston Co. —8.5 km SW Gastonia, along CR 1122, 1.4 km wjet.

CR 1131, along Crowders Creek, 2 ^5', 2 9, 16 August 1975, R.M. Shelley

and J.C. Clamp (NCSM A547); 6.4 km SW Gastonia, along CR 1 126, 0.8

km S jet. CR 1113, 9 , 16 August 1975, R.M. Shelley and J.C. Clamp

(NCSM A549); 7.7 km SW Gastonia, along CR 1 131, 0.2 km NW jet. CR

1 133, 9 , 8 July 1976, M. Filka and W.W. Thomson (NCSM A1091); and

1.9 km W Gastonia, along CR 1 106, 2.4 km E jet. CR 1236, d, 16 Oc-

tober 1976, M. Filka and G. Wicker (NCSM A2255).

Sigmoria latior (Brblemann)

Figs. 60-64

This was the most common xystodesmid in the region of study.

Adults and juveniles were discovered beneath decidous leaf litter and on

open substrate in July, but only four adults were taken in both April and

October. In North Carolina the species ranges from the northwestern

mountains to the eastern Piedmont, and intergrades of the three sub-

species were reported from McDowell County eastward to Scotland and

Hoke counties (Shelley 1976c), an area which includes the Kings Moun-

tain region.

Shelley (1976c) noted that all specimens available from south of the

Catawba and Deep-Cape Fear rivers, including intergrades of 5. 1. latior

(Brolemann) X S. /. hoffmani Shelley, had stripes along the caudal edges

of the metaterga. The nominate subspecies, occurring north of these

rivers, lacked stripes. During our study, however, specimens of both

color patterns were discovered. At Spencer Mountain they exhibited the

striped pattern, whereas around Kings-Crowders ridges the metaterga

32

Marianne E. Filka and Rowland M. Shelley

were black and without stripes. In both areas the stripe and/or paranotal

color was yellow and did not vary through shades of orange-red, as

reported by Shelley for the nominate subspecies and intergrades.

Since this is the first report of unstriped specimens in the zone of in-

tergradation, the gonopods of 18 striped and 13 unstriped males were ex-

amined for possible differences. Depth of the flange and broadness of the

distal curvature of the telopodite varied, but the flange always extended

below the tip of the telopodite (Fig. 60). The subterminal tooth varied in

prominence and was double in two individuals (Figs. 60-62), and the

prefemoral process ranged from simple to bifurcate with variation in the

relative lengths of the components, although the vertical branch was

always larger (Figs. 63-64). All are typical intergrade variations and do

not correlate with either color pattern. Thus, the solid black metatergal

color is interpreted to represent the nominate subspecies trait, just as

some intergrade gonopods more closely resemble those of one subspecies

than the other two.

DISCUSSION

Seasonal Variation of the Fauna

Although the Kings Mountain region was not sampled quanti-

tatively and only limited conclusions can be drawn concerning numbers

of species present in each season, the area was studied with sufficient in-

tensity to reflect general trends in seasonal differences (Table 3). The

overall abundance of millipeds increased from April to October, with

only two species, Narceus americanus and Auturus erythropygos, present

as both adults and juveniles in all three months. Adults of other species

varied seasonally, with juveniles present simultaneously or in other

months. The more common species that particularly exemplify these

seasonal variations are discussed by month below.

Diplopods were least abundant in April. Adults of Pseudopoly-

desmus branneri and Boraria stricta, and juveniles of Oxidus gracilis,

dominated the fauna, while adults and juveniles of N. americanus, and

adults of Cambala annulata and A. erythropygos were moderately abun-

dant. Adults of Ptyoiulus ectenes, Delophon georgianum, and Deltotaria

lea were less common, and juveniles of these species were absent or nearly

so. Only a few specimens of the remaining species were found. Adults of

D. georgianum, P. branneri, B. stricta, and D. lea, and juveniles of Stri-

aria sp., O. gracilis, and B. stricta were more numerous in April than in

any other month. Two species, Brachyiulus lusitanus and Trichopetalum

dux, were collected only in April.

A different group of diplopods dominated the fauna in July. Narceus

americanus, Abacion magnum, O. gracilis, A. erythropygos, Pachydesmus

crassicutis incursus, and Sigmoria latior were the prevalent adult forms,

while Polyxenus fasciculatus, Ptyoiulus sp., and N. americanus were com-

mon in immature stages. Intermediate numbers of A. erythropygos and

Kings Mountain Milliped Fauna

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34

Marianne E. Filka and Rowland M. Shelley

S. latior juveniles also were present, and the remaining species were

represented only by scattered individuals. Five species, P. fasciculatus, A.

magnum, Croatania catawba, P.c. incursus, and S. /an’or were more abun-

dant as both adults and juveniles during July than at any other time. Im-

matures of Nopoiulus minutus and Scytonotus granulatus also were most

numerous in July, although juveniles of the former were taken in October

as well. Species more common as adults in July than in April included

Polyzonium strictum, N. americanus, and O. gracilis. Species less common

as adults during July than in October or April included C. annulata and

B. stricta. Three species — P. ectenes, P. branneri, and D. lea — were ab-

sent from the July collections. Andrognathus corticarius was collected

only in July.

The greatest abundance and diversity of species occurred in October.

Those most abundant as adults were Ptyoiulus ectenes, P. impressus, C.

annulata, P. branneri, and A. erythropygos. Those most common in im-

mature stages were P. ectenes, O. gracilis, P. strictum, N. americanus, and

A. erythropygos. More juveniles of the last three species were encountered

in October than in either of the other months. The first two species were

moderately abundant as either adults or juveniles. Both Teniulus sp. and

Cleidogona medialis were found only in October and as adults. Millipeds

less common in the adult stage in October than in July included N.

americanus, A. magnum, O. gracilis, C. catawba, P.c. incursus, and S.

latior.

The species/genus ratio (S/G) for all three months (1.04) was essen-

tially unity (April and July S/G = 1.00, October S/G = 1.05), with the

slightly higher fraction of October reflecting the presence of both species

of Ptyoiulus. Seasonal changes in the faunal composition ratios (or-

ders/families/genera/species, O/F/G/S) from April to October were

more significant than changes in the S/G ratios. One more order, the

same number of families, and one less genus and species occurred in July

(8/12/16/16) than in April (7/12/17/17). The same number of orders,

one more family, four more genera, and five more species occurred in

October (8/13/20/21) than in July. Thus, the spring and summer faunas

were less diverse than the October fauna. These fluctuations reflect varia-

tions in times of maturation and breeding of the different species.

The overall O/F/G/S ratio for the three months combined

(9/16/23/24) showed one more order, three more families, three more

genera, and three more species than occurred in any single month. This

reflects the appearance and disappearance of species during the year,

which is also indicated by the following seasonal trends. Five species —

P. ectenes, C. annulata, P. branneri, B. stricta and D. lea — were more

common in April than in July and again increased in abundance during

October. A different five species — P. strictum, P. ectenes, C. catawba,

P.c. incursus and S. latior — were more common in July than in either of

the cooler months. Three species — P. strictum, P. ectenes, and P. im-

pressus — were most abundant in October, and two — N. americanus and

Kings Mountain Milliped Fauna

35

A. erythropygos — were common in all three months as both juveniles

and adults. Seven diplopods were encountered rarely (less than ten

specimens) and were collected during only one month {A. corticarius, B.

lusitanus, Teniulus sp., C. medialis, and T. dux), or two months {N.

minutus, and Striaria sp.) These data indicate that milliped faunas should

be sampled on a seasonal basis, a practice not generally followed to date,

and that collections in spring and fall may produce species not available

in summer.

Comparison of Faunas and Significance to North Carolina

Spencer Mountain is separated from the contiguous Kings-Crow-

ders ridge by approximately 15 km of urbanized Piedmont, and as shown

in Table 4 fewer milliped species occur at the inselberg. At both Spencer

Mountain (S/G = 1.00) and Kings-Crowder ridge (S/G = 1.05) every

genus is represented by one species with the sole exception of Ptyoiulus,

for which both species are present at Kings-Crowders ridge. At Spencer

Mountain, however, three less families, seven less genera, and eight less

species (8/11/14/14) were encountered than at Kings-Crowders ridge

(8/14/21/22). The two areas had 12 species in common — P.fasciculatus,

P. ectenes, N. americanus, A. magnum, D. georgianum, C. annulata, O.

gracilis, P. branneri, S. granulatus, A. erythropygos, B. stricta, and S.

latior. Two species collected only at Spencer Mountain, A. corticarius

and T. dux, were found in such low numbers (Table 3) that their absence

from the Kings-Crowders ridge could well be a collecting artifact.

The same is true of the apparent absence of six species from Spencer

Mountain — P. strictum, B. lusitanus, N. minutus, Teniulus sp., C.

medialis, and Striaria sp. Of the remaining five species absent from Spen-

cer Mountain, P. impressus, B. stricta, and P.c. incursus have western or

southern ranges that may well end at Kings-Crowders ridge. Two

xystodesmids, C. catawba and D. lea, could occur at Spencer Mountain,

since both were collected from Lincoln County in the 1950s. Their

presence seems doubtful, however, since the extensive searches for

diplopods at Spencer Mountain would surely have revealed these large,

brightly colored, and obvious millipeds. Thus, the absence of these five

species from Spencer Mountain may be real.

In addition to faunal distinctions between the two areas, color pat-

tern variation was noted in S. latior. As discussed in the species account,

specimens from Spencer Mountain displayed yellow paranota and stripes

along the caudal edges of the metaterga, whereas those from Kings-

Crowders ridge had yellow paranota but lacked the metatergal stripes.

No anatomical differences were detected, and both color patterns are

representative of intergrades. This is the first report of S. latior inter-

grades without stripes, a trait characteristic of the nominate subspecies.

The diplopod fauna of the Kings Mountain region is also compared

with the faunas of the eastern Piedmont and Appalachian Mountains

(numerical data for the Great Smoky Mountains) in Table 4, and is

36

Marianne E. Filka and Rowland M. Shelley

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<N

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^from Shelley (1978) for the eastern Piecimont

^from Hoffman (1969) for the Great Smoky Mountains, ordinal figure not provided.

Kings Mountain Milliped Fauna

37

shown to have a lower S/G ratio and fewer taxa below the level of order

than either. Comparison with the entire eastern Piedmont is somewhat

misleading, however, since the land area investigated by Shelley (1978)

was much larger and contained a greater variety of biotopes than the

Kings Mountain region. A more meaningful comparison is with the three

smaller areas that he sampled in detail — Medoc Mountain and William

B. Umstead state parks, and the hardwood locality near Ellerbe — each

more comparable in size to the Kings Mountain region. The ratios for

these three sites are as follows: Medoc Mountain State Park (5/6/7/7,

S/G = 1.00); William B. Umstead State Park (8/12/14/15, S/G = 1.07);

and Ellerbe (7/10/12/12, S/G = 1.00). The Kings Mountain fauna is

higher in each taxonomic category than any of these sites, but their S/G

ratios still reflect the occurrence of essentially one species per genus. Only

Ptyoiulus, with P. ectenes and P. impressus in the Kings Mountain region,

and Narceus, with N. americanus and N. annularis in Umstead State Park,

are represented by more than one species at a site. The greater numbers

of taxa in the Kings Mountain region may reflect its mountainous

character, but the region is still unable to support significantly more than

one species per genus. Compared to the Appalachian Mountains in

general and the Great Smoky Mountains in particular, the Kings Moun-

tain region has fewer taxa in every category (the number of orders for the

Great Smoky Mountains was not reported by Hoffman 1969) and a

much lower S/G ratio. Many Appalachian genera are represented by

more than one species, a reflection of the greater variety of niches af-

forded by the rugged, heterogeneous terrain.

Despite the numerical differences, however, there are similarities

between the Kings Mountain region and the other areas. Eight species of

widespread distribution are common to all three: P. strictum, A. cor-

ticarius, N. americanus, A. magnum, C. annulata, O. gracilis, P. branneri,

S. granulatus, and one or possibly two species of Striaria (taxonomic

problems exist within this genus). Some of the 24 species found in the

Kings Mountain region also occur in one of the others but not both.

Seven typically Piedmont inhabitants currently unknown from the moun-

tains are shared with the eastern Piedmont — P. fasciculatus, N. minutus,

B. lusitanus, P. ectenes, T. dux, A. erythropygos, and S. latior. Most were

expected in the Kings Mountain region at the outset of the study. Five

species are likewise shared with the Appalachians — P. impressus,

Teniulus sp., C. medialis, D. georgianum, and B. stricta. Their discovery in

the Kings Mountain region was a complete surprise and a significant

range extension for each. Fifteen species reported from the eastern Pied-

mont by Shelley (1978) were not found in the Kings Mountain region,

although three, Cylindroiulus truncorum (Silvestri), Ophyiulus pilosus

(Newport), and Apheloria tigana Chamberlin, are considered potential

inhabitants. The first two are synanthropic millipeds that could have

been overlooked in our study since we did not sample urban environ-

ments. Apheloria tigana is so common in the eastern Piedmont and in the

38

Marianne E. Filka and Rowland M. Shelley

more proximal Uwharrie Mountains that it must be considered a

possibility for the Kings Mountain region. Five millipeds known from

both the Appalachian Mountains and eastern Piedmont must also be

considered potential occupants of the Kings Mountain region due to its

location between these two areas. These five species are Polyzonium

rosalbum (Cope), known from Madison and Moore counties (Shelley

1976a, 1978); Cleidogona caesioannulata (Wood), reported from Macon,

Jackson, Transylvania, and Swain counties (Shear 1972) and Granville,

Orange, Durham, and Johnston counties (Shelley 1978); Branneria

carimta (Bollman) cited from Transylvania and Macon counties (Shear

1972), and Wake County (Shelley 1978); Pseudopolydesmus serratus,

collected in 14 eastern piedmont counties (Shelley 1978), and reported

generally from the mountains (Chamberlin and Hoffman 1958); and

Pleuroloma Jlavipes Rafinesque, recorded as Zinaria brunnea from

Watauga and Moore counties (Wray 1967) and as Pleuroloma sp. from

Orange and Wake counties (Shelley 1978).

In addition to species shared with the eastern Piedmont and/or Ap-

palachian Mountains, a fourth group of three xystodesmids is unique to

the Kings Mountain region: C. catawba, D. lea, and P.c. incursus. The

last is known in North Carolina only from the contiguous Kings-

Crowders ridge, but the others also have been recorded from Lincoln

County (Shelley 1978; Hoffman 1961), in the area that is the north-

eastward extension of the range to Anderson Mountain, Catawba

County. Croatania catawba and P.c. incursus are more common in South

Carolina and are basically southern forms which extend into North

Carolina along the Kings Mountain range. Together these two millipeds

lend a southern aspect to the Kings Mountain fauna, which is not found

in any other part of North Carolina. Deltotaria lea appears to be endemic

to a narrow section of the Carolinas, ranging from Lincoln County,

North Carolina, to Chester County, South Carolina.

Thus, the Kings Mountain milliped fauna is characterized by its own

species and the transitional ones it shares with the eastern Piedmont

Plateau and Appalachian Mountains, together and separately. Only five

of these species, however, are shared with the Appalachians alone. This,

plus the low diversity and the lowland nature of the fauna militate against

a prior direct topographic connection between the Blue Ridge Front and

the Kings Mountain region. Aside from a general Cretaceous peneplain

there is no geological evidence for such a connection, just as there is no

faunal evidence from the diplopods. Unlike the Appalachians, the Kings

Mountain region does not seem to have ever been a center of milliped

evolution and dispersal. The five Appalachian species in the area may be

relicts of a continuous Pleistocene or pre-Pleistocene distribution, as

suggested by Shelley (1979a) for D. georgianum. The most significant as-

pect of the Kings Mountain region is its position at the known range

periphery of several diplopods. It is the northern distribution limit of P c.

incursus and the northeastern of the genus Pachydesmus (Shelley and

Kings Mountain Milliped Fauna

39

Filka 1979), and two montane millipeds, D. georgianum and B. stricta,

reach their eastern terminus in the area. It is also the easternmost limit

for P. impressus and the genus Teniulus, the southeasternmost known site

for C. medialis, and the western limit for P. ectenes. The Kings Mountain

region is therefore a unique area in North Carolina, in the southern ele-

ments of its milliped fauna, in being a transitional area between

predominantly eastern and western faunas, and in forming a part of the

range periphery for four genera.

Teulings and Cooper (1977) used the term “cluster areas” to denote

places in North Carolina where species of concern are grouped. Four

rivers systems and four land areas in the Piedmont Plateau Province were

so identified. In a preliminary report, Filka and Shelley (1977) indicated

that, on the basis of its diplopod fauna alone, the Kings Mountain region

also would qualify as a cluster area. Three species considered of concern

in North Carolina occur in the region — P.c. incursus (endangered), and

C. catawba and D. lea (special concern). Moreover, the range peripheries

of P. ectenes, P. impressus, Teniulus sp., C. medialis, D. georgianum, B.

stricta and P.c. incursus lie there. The area also contains a unique

gonopod variant of B. stricta, and is distinguished by southern elements

of its fauna (C. catawba and P.c. incursus). As far as millipeds are con-

cerned the Kings Mountain region is of singular importance to North

Carolina, and investigations of other animal groups may provide further

evidence of its uniqueness. One state park, Crowders Mountain, exists in

the area, and every effort should be made to expand it to include the

deciduous bottomlands where most milliped species occur, including

those now considered of concern in the state. No millipeds were found

during this study in the dry, predominantly pine habitats of the existing

park.

One objective of this study, that of gaining insight into evolutionary

processes affecting millipeds in the southern Appalachians, went unmet.

With only five species in common and a lowland-type faunal diversity,

the Kings Mountain region adds little to current knowledge of milliped

biogeography that might be applied to such an objective. Moreover, none

of the five shared species belong to the xystodesmid tribe Aphelorini,

which is the single most diverse and abundant element of the Appala-

chian fauna. Aside from the ubiquitous Sigmoria latior, which ranges

from the mountains of West Virginia to the Coastal Plain of southern

South Carolina (Shelley 1976c), the great southeastern aphelorine fauna

is absent from the Kings Mountain region.

The study was, however, the first attempt to document seasonal

occurrence of milliped species in a discrete part of the southeast, an en-

deavor that should receive more attention. Seasonal sampling of juveniles

and adults can yield valuable information on life histories, for example,

and basic biological knowledge of this type has never been determined

for most North American diplopods. Although direct rearing of larvae

and adults, and breeding experiments, would provide the best such infor-

mation, inferences can nevertheless be gained from seasonal collections.

40

Marianne E. Filka and Rowland M. Shelley

ACKNOWLEDGMENTS.— take pleasure in thanking the per-

sons who helped collect specimens from the area of study, particularly

John C. Clamp, William W. Thomson, and Gerri W. Wicker. The as-

sistance of C.F. Lytle in providing equipment and laboratory space for

Filka is also gratefully appreciated. The specimens of C. medialis from

Deep Gap, Watauga County, North Carolina, were kindly loaned by

Norman I. Platnick, American Museum of Natural History; Shelley’s

collecting in Boone’s Cave and Crowders Mountain state parks was

done with permission of the North Carolina Department of Natural

Resources and Community Development, Division of State Parks.

Specimens of B. stricta and D. lea collected by Shelley in Kings Mountain

and Croft state parks. South Carolina, were secured through courtesy of

the South Carolina Department of Parks, Recreation, and Tourism. This

research was partly funded by the North Carolina State Museum of

Natural History and by the Department of Zoology, North Carolina

State University, and constituted part of a Master of Science thesis sub-

mitted to the latter institution by Filka.

LITERATURE CITED

Bollman, Charles H. 1887. Descriptions of fourteen new species of North Ameri-

can myriapods. Proc. U.S. Natl. Mus. 70:617-627.

Brimley, C.S. 1938. Insects of North Carolina. N.C. Dep. Agric. Div. Entomol.,

Raleigh. 560 pp.

Brolemann, Henri W. 1895. Liste de myriapodes des Etats-Unis, et principale-

ment de la Caroline du Nord, faisant partie des collections de M. Eugene

Simon. Ann. Soc. Entomol. France 65:43-70.

Burney, D.A. 1974. A preliminary interpretive prospectus of the Crowder’s

Mountain-King’s Pinnacle area of Gaston County, North Carolina. Unpubl.

rept. Div. Parks Rec., State Parks Sec. Raleigh. 14 pp.

Causey, Nell B. 1940. Ecological and systematic studies on North Carolina

myriapods. Ph.D. dissert., Duke Univ., Durham. 181 pp.

1943. Studies on the life history and ecology of the hothouse milli-

pede, Orthomorpha gracilis (C.L. Koch, 1847). Am. Midi. Nat. 5:670-682.

Chamberlin, Ralph V. 1940a. On some chilopods and diplopods from North

Carolina. Can. Entomol. 72:56-59.

1940b. Four new polydesmoid millipeds from North Carolina

(Myriapoda). Entomol. News 57:282-284.

1951. On eight new southern millipeds. Great Basin Nat. 77:19-26.

, and R.L. Hoffman. 1958. Checklist of the millipeds of North

America. U.S. Natl. Mus. Bull. 212. 236 pp.

Cooper, John E., S.S. Robinson and J.B. Funderburg (eds.). 1977. Endangered

and Threatened Plants and Animals of North Carolina. N.C. State Mus.

Nat. Hist., Raleigh, xvi + 444 pp.

Enghoff, Henrik, and R.M. Shelley. 1979. A revision of the millipede genus

Nopoiulus (Diplopoda:Julida:Blaniulidae). Entomol. Scand. 70:65-72.

Filka, M., and R.M. Shelley. 1977. The Kings-Crowders Mountain Region: A

milliped “cluster” area in North Carolina (Diplopoda). ASB Bull. 24{2):29.

Abstract.

Gardner, Michael R. 1975. Revision of the milliped family Andrognathidae in

the Nearctic Region. Mem. Pac. Coast Entomol. Soc. 5:1-61.

Kings Mountain Milliped Fauna

41

Hardin, James W,, and A.W. Cooper. 1967. Mountain disjuncts in the eastern

Piedmont of North Carolina. J. Elisha Mitchell Sci. Soc. 55:139-150.

Hoffman, Richard L. 1950. Records and descriptions of diplopods from the

southern Appalachians. J. Elisha Mitchell Sci. Soc. 66:11-31

1958. Appalachian Cambalidae: Taxonomy and distribution (Dip-

lopoda: Spirostreptida). J. Wash. Acad. Sci. 45:90-94.

1961. Revision of the milliped genus Deltotaria (Polydesmida:

Xystodesmidae). Proc. U.S. Natl. Mus. 775:15-35.

1962. The milliped genus Scytonotus in eastern North America,

with the description of two new species. Am. Midi. Nat. 67:241-249.

1965. Revision of the milliped genera Boraria and Gyalostethus

(Polydesmida:Xystodesmidae). Proc. U.S. Natl. Mus. 777:305-347.

1969. The origin and affinities of the southern Appalachian diplo-

pod fauna, pp. 221-246 in P.C. Holt (ed). The distributional history of the

biota of the southern Appalachians, part I: Invertebrates. Res. Div.

Monogr. 1, Va. Polytech. Inst. Blacksburg. 295 pp.

1974. A new polydesmid milliped from the southern Appalachians

with remarks on the status of Dixidesmus and a proposed terminology for

polydesmid gonopods. Proc. Biol. Soc. Wash. 57:345-350.

Hunt, Charles B. 1967. Physiography of the United States. W.H. Freeman & Co.,

San Francisco. 480 pp.

Keeton, William T. 1960. A taxonomic study of the milliped family Spiro-

bolidae (Diplopoda:Spirobolida). Mem. Am. Entomol. Soc. 11. 146 pp.

Keith, A. 1931. Geologic Atlas of the United States, Gaffney-Kings Mountain

Folio, South Carolina-North Carolina. U.S. Geol. Survey #222. 13 pp. + 4

maps.

Kesel, Richard H. 1974. Inselbergs on the Piedmont of Virginia, North Carolina,

and South Carolina: Types and characteristics. Southeast. Geol. 76:1-30.

Shear, William A. 1972. Studies in the milliped order Chordeumida (Diplopoda):

A revision of the family Cleidogonidae and a reclassification of the order

Chordeumida in the new world. Bull. Mus. Comp. Zool. 774:151-352.

Shelley, Rowland M. 1976a. Two new diplopods of the genus Polyzonium from

North Carolina, with records of established species (Polyzoniida:Poly-

zoniidae). Proc. Biol. Soc. Wash. 55:373-382.

1976b. A new diplopod of the genus Cleidogona from North

Carolina (Chordeumida:Cleidogonidae). Fla. Entomol. 59:325-327.

1976c. Millipeds of the Sigmoria latior complex (Polydesmida:

Xystodesmidae). Proc. Biol. Soc. Wash. 89:\l-3^.

1977. The milliped genus Croatania (Polydesmida:Xystodesmi-

dae). Proc. Biol. Soc. Wash. 96:302-325.

1978. Millipeds of the eastern Piedmont region of North Carolina,

U.S. A. (Diplopoda). J. Nat. Hist. 72:37-79.

1979a. A revision of the milliped genus Delophon, with the proposal

of two new tribes in the subfamily Abacioninae (Callipodida:Caspio-

petalidae). Proc. Biol. Soc. Wash. 92:533-550.

1979b. A synopsis of the milliped genus Cambala, with a description

of C. minor Bollman (Spirostreptida:Cambalidae). Proc. Biol. Soc. Wash.

92:551-571.

, and M. Filka. 1979. Occurrence of the milliped Pachydesmus crassi-

cutis incursus Chamberlin in the Kings Mountain region of North Carolina

42

Marianne E. Filka and Rowland M. Shelley

and the Coastal Plain of South Carolina (PolydesmidaiXystodesmidae).

Brimleyana 1:147-153.

Stuckey, Jasper L. 1965. North Carolina: Its Geology and Mineral Resources.

N.C. Dep. Cons. Devel., Raleigh. 550 pp.

Teulings, Robert P., and J.E. Cooper. 1977. Cluster Areas, pp. 409-431 in

J.E. Cooper, S.S. Robinson, and J.B. Funderberg (eds.). Endangered and

Threatened Plants and Animals of North Carolina. N.C. State Mus. Nat.

Hist., Raleigh, xvi -I- 444 pp.

Wray, David L. 1967. Insects of North Carolina, Third Supplement. N.C. Dep.

Agric. Div. Entomol., Raleigh. 181 pp.

Accepted 15 September 1980

Electrophoretic Analysis of Three Species of Necturus

(Amphibia: Proteidae), and the Taxonomic Status

of Necturus lewisi (Brimley)

Ray E. Ashton, Jr. and Alvin L. Braswell

North Carolina State Museum of Natural History,

P.O. Box 27647, Raleigh, North Carolina 27611

AND

Sheldon I. Guttman

Department of Zoology,

Miami University, Oxford, Ohio 45056

ABSTRACT.— E\Qcirop\iorQi\c analyses of 10 Necturus maculosus from

Minnesota, 10 from Massachusetts, and 1 from the Mills River, Hen-

derson County, North Carolina, were compared with those of 20

Necturus lewisi and 8 unspotted Necturus punctatus from the Neuse

River drainage and 8 spotted N. punctatus from Naked Creek, Robeson

County, North Carolina. Evaluation of 17 loci showed that the three

samples of N. maculosus were indistinguishable (Nei’s D = 0.000) while

N. lewisi were unequivocally different from N. punctatus at four loci and

from N. maculosus at six loci. The two N. punctatus populations were in-

distinguishable from each other but were distinguishable from N.

maculosus at 6 loci. These data indicate that N. maculosus, N. lewisi and

N. punctatus are distinct, long isolated species.

INTRODUCTION

Necturus lewisi is one of several endemic species of vertebrates and

invertebrates found in the Tar and Neuse River drainages of North

Carolina. This waterdog was originally described by Brimley (1924) as a

subspecies of Necturus maculosus because of the “spotted larvae”. Viosca

(1937) briefly described the previously unknown striped larvae of N.

lewisi and used its medium size and overall spotting as the apparent

criteria for elevating it to full species status. Ashton and Braswell (1979)

compared N. lewisi hatchlings and striped larvae with larvae of N.

maculosus and N. punctatus, and found that the striped larvae of N. lewisi

were quite distinctive. No electrophoretic studies in the genus have been

reported. Our study compared electrophoretic data for all three of these

Necturus species, in an attempt to evaluate the taxonomic status of N.

lewisi.

METHODS AND MATERIALS

Ten N. maculosus from Minnesota were obtained from Nasco, Fort

Atkinson, Wisconsin and ten from Massachusetts were purchased from

Connecticut Valley Biological Supply Company, Southampton,

Massachusetts. One additional N. maculosus was collected in the Mills

River, Henderson County, North Carolina. Twenty N. lewisi and eight

unspotted N. punctatus were captured in the Neuse River drainage. Eight

Brimleyana No. 4: 43-46. December 1980.

43

44 Ray E. Ashton, Jr., Alvin L. Braswell, Sheldon I. Guttman

spotted N. punctatus were collected from Naked Creek, PeeDee River

drainage, in the Sandhills region of Robeson County, North Carolina.

Animals were killed in the laboratory and an organ homogenate pre-

pared from the heart, liver, rinsed stomach and upper part of intestine,

and kidney and tongue of each. The specimen remains are housed in the

North Carolina State Museum collection. The tissues of individual

animals were then homogenized in an equal volume of 2% 2-

phenoxyethanol and centrifuged at 25,000 g at 4° C for 45 minutes. The

supernatant of soluble proteins was then decanted and stored at -70° C

until used a maximum of 48-hours following preparation.

The 17 loci coding for proteins consistently resolved are as follows:

malate dehydrogenase (NAD-dependent) (Mdh-1); indophenol oxidase

(Ipo-1); ^-glycerophosphate dehydrogenase (a-Gpdh-l); isocitrate

dehydrogenase (NADP-dependent) (Idh-1); phosphoglucomutases, three

loci (Pgm-1, Pgm-2, Pgm-3); glutamate oxalate transaminases, two loci

(Got-1, Got-2); glutamate dehydrogenase (Gdh-1); phosphoglu-

coisomerase (Pgi-1); malic enzyme, two loci (Me-1, Me-2); 6

phosphogluconate dehydrogenase (6-Pgdh-l); sorbitol dehydrogenase

(Sdh-1), glyceraldehyde-3-phosphate dehydrogenase (G-3-pdh-l); and

lactate dehydrogenase (Ldh-2).

Techniques of horizontal starch gel electrophoresis and protein

staining were similar to those described by Selander et al. (1971), with the

following modifications: Idh, Pgm, Mdh, Gdh, and Me were examined

with their continuous tris-citrate buffer (pH 8.00); 6-Pgdh, Got, Sdh and

G-3-pdh were demonstrated with the tris-borate-EDTA buffer of Ayala

et al. (1973). Staining methods for Gdh, G-3-pdh and Sdh were as

described by Brewer (1970). All gels were 12.5% starch (Electrostarch Lot

mi).

Genetic inferences from electrophoretic results are based on the pat-

terns being consistent with known molecular configurations for the pro-

teins analysed, i.e. two-banded patterns are observed for the

heterozygotes for a protein that is a monomer and three-banded patterns

are observed for a dimeric heterozygote. The genes coding for each en-

zyme are represented by italicized abbreviations.

If several forms of the same enzyme are present and each is con-

trolled by a separate gene locus, the hyphenated numeral serves to dif-

ferentiate the loci. The enzyme with the greatest anodal migration is

designated one, the next two, and so on. When allelic variation occurs,

the allele with the greatest anodal migration is called a, the next b, and so

on.

RESULTS

The two N. maculosus samples were essentially identical genetically

(genetic distance, D = 0.000; Nei 1972). One heterozygote was found at

each of the two loci (Pgm-2, Idh-1) in the Massachusetts sample, the only

heterozygotes found.

Necturus Electrophoresis

45

The only variants found in the N. lewisi sample were at the Got-1

locus. Six individuals were heterozygous for the a and b alleles, one was

homozygous for b.

Two unspotted N. punctatus were each heterozygous at single loci

(Pgi-1, Pgm-2). The only difference between the spotted and unspotted

samples was in the frequency of the Pgi-1 alleles (Table 1).

Table 1. Fixed genetic differences in three Necturus species.

Locus

Allele

The three species were unequivocally different at four {N. lewisi vs.

N. punctatus) or six {N. maculosus vs. N. lewisi or N. punctatus) of the

seventeen loci investigated; alleles were not shared at these loci (Table 1).

Nei’s standard genetic distance estimates between each pair of species

(Table 2) are indicative of a long history of isolation of the gene pools.

Electrophoretic examination of one N. maculosus from the Mills River,

Henderson County, North Carolina confirms the genetic distinction

found between the larger samples of allopatric N. maculosus and N.

lewisi.

Table 2. Standard genetic distance (D) between species of Necturus examined.

12 3 4

46 Ray E. Ashton, Jr., Alvin L. Braswell, Sheldon I. Guttman

DISCUSSION

Electrophoretic analysis of ten individuals from each of two popula-

tions of N. maculosus and one individual from a third population showed

that they were indistinguishable using this technique. Necturus lewisi and

N. punctatus, however, were highly distinguishable from each other and

from N. maculosus, indicating that each species has been genetically

isolated for some time. Two populations of N. punctatus, the spotted

form inhabiting the Sandhills region of North Carolina and the uni-

formly gray-black form inhabiting the Neuse River, were in-

distinguishable from each other.

In conclusion, the specific status of N. lewisi is confirmed by elec-

trophoretic data as well as by the distinct larvae described by Ashton and

Braswell (1979). Further, N. punctatus appears to have been reproduc-

tively isolated from sympatric N. lewisi and from allopatric N. maculosus

for a considerable period of time, and spotted N. punctatus from the

PeeDee River drainage (North and South Carolina) appear on the basis

of electrophoresis to be genetically similar to the unspotted populations

of the Neuse River system.

ACKNOWLEDGMENTS. — The authors wish to express their ap-

preciation to field technicians Angelo Capparella, Paul Freed, and Jerry

Reynolds, and laboratory assistants Gary Trakshel, Kim Haikyong, and

Ernie Flowers. We also thank John E. Cooper for his critical review of

the manuscript. This project was in part supported by funds from a U.S.

Fish and Wildlife Service (Office of Endangered Species) cooperative

agreement with the North Carolina Wildlife Resources Commission.

LITERATURE CITED

Ashton, Ray E,, Jr, and A.L. Braswell. 1979. Nest and larvae of the Neuse River

Waterdog, Necturus lewisi (Brimley) (Amphibia: Proteidae). Brimleyana

7:15-22.

Ayala, Francisco J., D. Hedgecock, G. Zumwalt and J. Valentine. 1973. Genetic

variation in Tridacna maxima, an ecological analog of some unsuccessful

evolutionary lineages. Evolution 27:177-191.

Brewer, George. 1970. Introduction to isozyme techniques. Academic Press,

New York. 186 pp.

Brimley, Clement S. 1924. The Water Dogs {Necturus) of North Carolina.

J. Elisha Mitchell Sci. Soc. 40(3-4): 166- 168.

Nei, Masatoshi. 1972. Genetic distance between populations. Am Nat

706:283-292.

Selander, Robert K., M.H. Smith, S.Y. Yang, W.E. Johnson and J.B. Gentry.

1971. Biochemical polymorphism and systematics in the genus Peromyscus.

I. Variation in the old-field mouse {Peromyscus polionotus). Stud. Genet. VI.

Univ. Texas Publ. 7103:49-90.

Viosca, Percy, Jr. 1937. A tentative revision of the genus Necturus, with descrip-

tions of three new species from the southern Gulf drainage area Copeia

1937(2):120-138.

Accepted 14 October 1980

Vertebrates of the Okefenokee Swamp

Joshua Laerm, B.J. Freeman, Laurie J. Vitt

Museum of Natural History and

Department of Zoology

Joseph M. Meyers

Institute of Ecology

AND

Lloyd Logan

Museum of Natural History and

Department of Zoology

University of Georgia, Athens, Georgia 30602

ABSTRACT. —Four hundred nineteen vertebrate species and sub-

species are known from the Okefenokee Swamp region of Georgia and

adjacent Florida. These include 36 fishes, 37 amphibians, 66 reptiles,

232 birds, and 48 mammals. The vertebrates occurring in the

Okefenokee represent a typical southeastern Atlantic Coastal Plain

fauna. There are no endemic species. Eleven species, recognized as

threatened or endangered under state and/or federal guidelines, occur

in the swamp.

INTRODUCTION

The Okefenokee Swamp region of southeastern Georgia and adja-

cent Florida contains an extremely diverse vertebrate fauna. However,

with the exception of biological surveys conducted by Cornell University

in the early decades of this century, that fauna has received little atten-

tion. At present there exists no comprehensive information on the ver-

tebrates of the swamp. Most of the available literature is semipopular,

anecdotal, or, at best, outdated.

Accurate faunal information is essential to understanding the Oke-

fenokee Swamp ecosystem. The long term value and credibility of the

systems ecology studies presently being undertaken in the swamp will, in

large part, be determined by the extent to which base level natural history

information can be incorporated into definitive analyses and models.

Base level faunal surveys provide information on species diversity and

patterns of habitat use that are crucial for biogeographic and systematic

research. Furthermore, comprehensive faunal studies serve also as dated

testaments to species composition and distribution within specific

habitats, which are crucial for enviromental impact assessments

associated with management practices.

For these reasons, we have undertaken vertebrate faunal surveys

within the Okefenokee Swamp and surrounding uplands. We report here

the results of these surveys. We present, too, a review of pertinent

historical foundations of our present knowledge of the swamp’s ver-

tebrate fauna, a comparison of the fauna with that of adjacent south-

eastern regions, and a preliminary analysis of habitat distributions of ver-

tebrates known to occur within the swamp.

Brimleyana No. 4: 47-73. December 1980.

47

48

Joshua Laerm, et al.

GENERAL HABITAT CHARACTERISTICS

The Okefenokee Swamp is one of the largest freshwater wetlands in

the United States. Situated in Charleton, Clinch, Echols and Ware coun-

ties, Georgia, and Baker and Columbia counties, Florida, the Oke-

fenokee watershed includes both swamp (189,000 ha) and surrounding

uplands (181,000 ha). It lies within the humid subtropical climatic zone

(Trewartha 1968) and is characterized by warm moist springs, hot wet

summers, warm dry falls, and cool moist winters. Weather is

predominantly influenced by tropical maritime air masses from the Gulf

of Mexico and the tropical Atlantic Ocean in spring, summer and fall,

but by continental air masses in winter. Annual precipitation averages

100-150 cm (Hunt 1972).

The Okefenokee consists of a variety of vegetational habitat types.

Plant specimens are on file at the University of Georgia Herbarium.

a. Two prairie habitat types are identified, comprising approx-

imately 21% of the swamp. (1) Aquatic macrophyte prairies are

dominated by emergent, floating-leaved, and submerged hydrophytes

such as white water lily, Nymphaea odorata; yellow water lily, Nuphar

luteum; neverwet, Orontium aquaticum; floating heart, Nymphoides

aquaticum; yellow eyed grass, Xyris smalliana; pickerel weed, Pontederia

cordata; redroot, Lachnanthes caroliniana; and bladderwort, Utricularia

spp. (2) Grass-sedge prairies are characterized by various species of

sedges, Carex; panic grasses, Panicum; and beak rush, Rhynchospora, as

well as broomsedge, Andropogon virginicus; giant chain fern, Woodwardia

virginica; and Sphagnum moss.

b. Shrub swamps cover approximately 34% of the swamp and are

predominated by hurrah bush, Lyonia lucida; fetter bush, Leucothoe

racemosa; titi, Cyrilla racemijiora; sweet spire, Itea virginica; pepper

bush, Clethra alnifolia; and dahoon. Ilex cassine.

c. Blackgum forests cover less than 6% of the swamp. Blackgum,

Nyssa sylvatica var. biflora, with a small amount of dahoon and pond

cypress, Taxodium ascendens, dominates the canopy, with red maple,

Acer rubrum, and dahoon the predominant understory plants.

d. Bay forests also cover less than 6% of the swamp. Loblolly bay,

Gordonia lasianthus; red bay, Persea borbonia; and sweet bay. Magnolia

virginiana, are the predominant canopy species although occasional pond

cypress, blackgum, and slash pine, Pinus elliottii, are seen.

e. Mixed cypress forests are characterized by pond cypress domi-

nated canopy and subcanopy, but loblolly bay, dahoon, and blackgum

are frequently scattered in the subcanopy. This and the following habitat

make up approximately 23% of the swamp.

f. Pure cypress forests are limited in extent but consist almost en-

tirely of a cypress canopy with a sparse subcanopy or understory.

g. There are approximately 70 islands in the swamp and they ac-

count for roughly 12% of the area. Vegetation is dominated by loblolly

pine, Pinus taeda; slash pine; longleaf pine, Pinus palustris; water oak.

Okefenokee Swamp Vertebrates

49

Quercus niger; and live oak, Quercus virginiana.

The uplands surrounding the swamp are intensively managed pine

forests. Historically, the area was dominated by longleaf and slash pine

with an understory dominated by saw palmetto, Serenoa repens; small

gallberry. Ilex glabra; and various forbs and grasses. Fire was the major

factor maintaining successional stages (Monk 1968). Today the uplands

are dominated by slash pine plantations with a similiar understory

managed by prescribed periodic burns. Remnants of hardwood and

mixed hardwood-pine forests are very limited but occur in scattered loca-

tions on some islands and at the periphery of the swamp. Management

for pine, including prescribed burns, is responsible for the virtual absence

of hardwoods in the uplands.

FISHES

Historical Foundations

Scientific collections of fishes in the Okefenokee Swamp span 68

years. The earliest significant collections were undertaken in 1912 by

A.H. Wright and Francis Harper, both from Cornell University. The ac-

count of Palmer and Wright (1920), based primarily on these collections,

represents the only published information on fishes of the swamp. Subse-

quent collections, resulting from various museum expeditions and the ac-

tivities of Okefenokee National Wildlife Refuge (ONWR) personnel,

were made by R.A. Chesser in 1922, R.T. Berryhill in 1924, T.

Reichelderfer in 1935, M.S. Verner, Jr. in 1936, B. Cadbury in 1937, C.B.

Obrecht and M. Godfrey in 1941, H.A. Carter in 1941-1942, Southern

Piedmont and Coastal Plain Survey in 1941, T. Rodenberry in 1941, and

R.J. Fleetwood in 1947. Collecting activities ceased in the 1950s and

began again in the 1960s (E. Cypert in 1960, 1963; T. Cavender in 1965;

and M.W. Bohlke in 1966), and have continued to the present (B.J.

Freeman, 1978-1980). Additional studies in the southeastern lower

Coastal Plain (Gassaway 1976; Holder and German 1977) contributed

much to existing knowledge of swamp ichthyofauna. Voucher specimens

of significant collections are deposited in the National Museum of

Natural History, Philadelphia Academy of Natural Sciences, Cornell

University, University of Georgia Museum of Natural History, and Uni-

versity of Michigan Museum of Zoology.

Comparison With Regional Fauna

The ichthyofauna of Okefenokee Swamp consists of 36 species

representing 13 families (Table 1). The most remarkable character of the

fauna is the absence of minnows (Cyprinidae). The remaining fish fauna

is not substantially different from adjacent southeastern drainages.

Average faunal resemblance values (Ramsey 1965) were computed for

Okefenokee Swamp and major drainages in the area. Values can range

50

Joshua Laerm, et al.

from 0 to 1, with 0 indicating no species in common and 1 indicating all

species in common. The river systems compared were the Suwannee

River (from Fargo, Georgia to its junction with the Alapaha River); the

Alapaha River (a Suwannee River tributary); the Withlacoochee River (a

Suwannee River tributary); the St. Mary’s River, and the Satilla River.

The values ranged from a minimum of .84 for the Withlacoochee to a

maximum of .90 for the St. Mary’s. The Alapaha, Suwannee, and Satilla

were intermediate, with resemblance values of .86, .86, and .88, respec-

tively. These differences are due entirely to absence from the swamp of

minnows, which otherwise are widely distributed in adjacent drainages.

Their absence appears to be due to substantially lower pH values in the

swamp.

Swamp pH ranges from 3.1 to 4.2; pH values for surrounding

streams (where minnows occur) range from 4.8 to 6.9. In a study of

Carolina bay lakes in North Carolina, Frey (1951) noted that in two lakes

with a pH of 4.3 there were no minnows, while lakes with higher pH

values (up to 5.9) had some minnows present. These were Notropis

chrysoleucas, N. chalybaeus, and N. petersoni — three of the species that

occur near the Okefenokee Swamp. Comparing minnow distributions

with pH shows that appearance of minnows in the St. Mary’s River coin-

cides with a pH of 4.8 or higher. The pH values for surrounding streams

are even higher. Although detailed pH data for these streams are not

available (especially for the Suwannee River section) the general pattern

suggests that increasing acidity might limit, or at least influence, minnow

distributions. This possibility deserves more critical attention.

Habitat Distribution of the Fishes

The 36 species of fish occurring in the swamp are distributed in a

heterogeneous series of aquatic habitats that can be broadly classified as

lake, aquatic prairie, and stream.

Lakes are open bodies of water of .25 ha or larger with depths of .5

m or more. The bottom is generally unconsolidated peat, which may have

a depth of .3 m to greater than 1 m; some lakes, however, have hard sand

bottoms. The margins are heavily vegetated with rooted and floating

aquatic plants as well as submergent vegetation. The topography around

the lakes grades into aquatic prairie (when the water levels are not low)

composed of a variety of rooted aquatic macrophytes, floating vegeta-

tion, sedges, and small shrubs. Water depth may range from several cm

to over 1 m. Current in these two areas varies from none in the lakes to

noticeable in the prairies. Streams generally have noticeable to moderate

current, consolidated banks, and sandy bottoms. Some aquatic vegeta-

tion and backwater areas are at the water margins.

The streams are located primarily in the northwest part of the

swamp and on some of the islands. The prongs of the Suwannee River

and the Suwannee Canal also provide stream habitat. Elements of the

prairie habitat, i.e. heavily vegetated areas, can be found bordering lakes

Okefenokee Swamp Vertebrates

51

and streams as well as in the large, open expanses of the swamp.

Aquatic habitats are not discrete units in the swamp but are graded

and sometimes mixed. The distribution of fishes reflects this. The habitat

associations of the swamp ichthyofauna indicates the fishes are rather

uniformly distributed (Table 1). Comparison of water current preferences

among the species does, however, indicate some degree of habitat

segregation. Noturus leptacanthus and Percina nigrofasciata will

generally be found in water with noticeable to moderate current. Umbra

pygmaea, Fundulus chrysotus, Fundulus cingulatus, Fundulus lineolatus,

Leptolucania ommata, Heterandria formosa, Elassoma evergladei and

Elassoma okefenokee generally are found in areas with no current but

with abundant aquatic vegetation. The remaining fishes occur in areas

with water currents ranging from none to noticeable. This wide range of

current tolerances helps explain the overlap observed in fish distributions

across obvious physically different habitats.

Table 1. List of fishes of the Okefenokee Swamp. Based on museum records and

data from Dahlberg and Scott (1970), Gasaway (1976), Holder and

German (1977), and personal observations (B.J. Freeman). Scientific

and common names based on Bailey et al. (1970). L = lake, P= prairie,

S = stream.

SPECIES

ORDER SEMIONOTIFORMES

Family Lepisosteidae

Lepisosteus platyrhincus, Florida gar

ORDER AMIIFORMES

Family Amiidae

Amia calva, Bowfin

ORDER ANGUILLIFORMES

Family Anguillidae

Anguilla rostrata, American eel

ORDER OSTEOGLOSSIFORMES

Family Esocidae

Esox americanus, Redfin pickerel

Esox niger, Chain pickerel

Family Umbridae

Umbra pygmaea, Mudminnow

ORDER CYPRINIFORMES

Family Catostomidae

Erimyzon sucetta, Lake chubsucker

Minytrema melanops. Spotted sucker

HABITAT

PREFERENCE

L P S

L S

52

Joshua Laerm, et al.

HABITAT

SPECIES PREFERENCE

ORDER SILURIFORMES

Family Ictaluridae

/c/^z/wn/5 Yellow bullhead EPS

Ictalurus nebulosus, Brown bullhead L P

Ictalurus punctatus, Channd catfish L S

N o tu ms gyrinus, Tadpole madtom EPS

Noturus leptacanthus, Speckled madtom S

ORDER PERCOPSIFORMES

Family Aphredoderidae

Aphredodems sayanus,V\xatc perch EPS

ORDER ATHERINIFORMES

Family Cyprinodontidae

Fundulus chrysotus, Golden topminnow EPS

Fundulus cingulatus, landed topminnov^ EPS

Fundulus line olatus, fined topminno^N EPS

Leptolucania ommata,T)/gmy kWhfish EPS

Family Poeciliidae

Gambusia affinis, Mosquitofish EPS

Heterandria formosa, Eeast killifish EPS

Family Atherinidae

Labidesthes sicculus, Brook silverside EPS

ORDER PERCIFORMES

Family Elassomidae

Elassoma evergladei, Everglades pygmy sunfish EPS

Elassoma okefenokee, Oke^enokee pygmy sixnfish EPS

Family Centrarchidae

A cantharcus pomotis, Mndsixnfish EPS

Centrarchus macroptems, Flier EPS

Enneacanthus chaetodon, ^lackhanded sixnfish E P

Enneacanthus gloriosus,^hxespottedsi\nfish EPS

Enneacanthus obesus, Banded sunfish EPS

Lepomis gulosus, Warmouth EPS

Lepomis macrochims, Bluegill EPS

Lepomis marginatus, DoWav sunfish EPS

Lepomis punctatus, Spotted sunfish EPS

Microptems salmoides, Largemouth bass EPS

Pomoxis nig romaculatus, Black cvapp'ie E S

Family Percidae

Etheostoma fusiforme, Swamp darter EPS

Percina nigrofasciata, Blackbanded darter S

AMPHIBIANS AND REPTIEES

Historical Foundations

Serious investigations of the herpetofauna of Okefenokee Swamp

began in 1912 with the first in a series of surveys conducted by Cornell

University. Prior to this only anecdotal accounts of the reptiles and

Okefenokee Swamp Vertebrates

53

amphibians are known (Fountain 1901 [cited in Wright and Funkhouser

1915]; Reese 1907). At least three herpetologists participated in the Cor-

nell collections: A.H. Wright, W.D. Funkhouser, and S.C. Bishop. Dur-

ing the same period (and possibly with the same expedition) F. Harper

began recording observations on some of the reptiles and amphibians. In

two summary publications (Wright and Bishop 1915; Wright and

Funkhouser 1915), 9 chelonians, 6 saurians (actually 7, as Wright and

Funkhouser had 2 species of Ophisaurus), 21 serpents and 1 crocodilian

were recorded. This represents less than half the currently known fauna.

Harper (1934) discussed aspects of the ecology and behavior of several

Okefenokee reptiles and amphibians based on his visits, and numerous

short papers on aspects of the biology of Okefenokee species, mostly

authored by A.H. Wright, appeared in various scientific journals. Many

of the observations on anurans in Wright (1932) and Wright and Wright

(1949) were based on Okefenokee studies. Since these early visits to the

swamp (up to about 1946), there has only recently been a renewed in-

terest in its herpetofauna. Several southeastern herpetologists made small

collections in the area, including W.T. Neill and F.L. Rose, but the

collections made by C.H. Wharton and his students at Georgia State

University are by far the most extensive. Additional surveys have been

conducted by L. Vitt and J. Laerm. Significant collections of Okefenokee

material can be found at Cornell University, Florida State Museum,

National Museum of Natural History, University of Georgia Museum of

Natural History, and University of Michigan Museum of Zoology.

Comparison With Regional Fauna

The Okefenokee Swamp contains a diverse herpetofauna of 103

species and subspecies including 2 crocodilians, 15 chelonians, 38 ser-

pents, 11 saurians, 16 urodeles, and 21 anurans (Table 2). The present

herpetofauna can be considered a typical southeastern Atlantic Coastal

Plain fauna (see Conant 1975). There are no species endemic to the

swamp. In general species diversity within the swamp and surrounding

uplands is greater than in similar sized areas in the adjacent southeastern

Atlantic Coastal Plain, primarily because of the high habitat diversity

associated with the swamp. However, the high species diversity can also

be attributed to the fact that at least 20 species of reptiles and amphibians

reach the limit of their natural range in the region of the swamp (see Co-

nant 1975). Thus, the faunal diversity is somewhat greater in the

Okefenokee region in comparison to other Atlantic Coastal Plain

localities to the immediate north or south.

Habitat Distribution of the Amphibians and Reptiles

Unlike the other vertebrates, most amphibians and reptiles in the

Okefenokee are not usually associated with a particular vegetational

habitat (blackgum swamps, for example) but rather seem to be associated

with structural habitats (water courses, sandy bottoms, etc.) Thus, it

54

Joshua Laerm, et al.

serves little purpose to group species by vegetation habitats recognized by

biologists. Ecological distribution of the herpetofauna can, however, be

summarized in terms of general habits of the animals. Many species, for

example, are entirely aquatic and use most if not all aquatic habitats in

the swamp. Other categories also are useful in respect to ecological dis-

tribution of the reptiles and amphibians. For descriptive purposes, the

herpetofauna is partitioned into six “ecological” groups (Table 2): 1) En-

tirely aquatic species are those that spend nearly all of their lives in water;

2) Semi-aquatic species are those that spend a major part of their lives in

water, but may often be found on land (does not include species entering

water only for breeding); 3) Fossorial species are those that spend most of

their lives underground (they may become surface active for breeding or

limited foraging); 4) Terrestrial species are those most often encountered

on the surface and that spend most of their active time there; 5)

Terrestrial-arboreal species may spend nearly as much time in arboreal

habitats as on the surface; 6) Arboreal species are those that spend nearly

all of their lives in vegetation (some of these may enter water to breed, or

lay eggs on the ground).

Of the Okefenokee Swamp herpetofauna, 25 (24.3%) species are en-

tirely aquatic, 21 (20.4%) are semiaquatic, 10 (9.7%) are fossorial, 29

(28.2%) are terrestrial, 9 (8.7%) are terrestrial-arboreal, and 9 (8.7%) are

arboreal. Most turtles are either aquatic or semiaquatic, most lizards

tend to be terrestrial, terrestrial-arboreal or arboreal, most snakes are

terrestrial (but there are large numbers of species in other groups), most

salamanders are aquatic, semiaquatic or fossorial, and frogs (including

toads) tend to be semiaquatic or arboreal (Table 2).

Table 2. List of amphibians and reptiles of the Okefenokee Swamp. Based on

museum records and data from Wright and Funkhouser (1915), Wright

and Bishop (1915), Wright (1932), Harper (1934), Wright and Wright

(1949), and personal observations (L. Vitt, J. Laerm). Most scientific

and all common names based on Collins et al. (1978). Aq = aquatic,

Ar = arboreal, F = fossorial, Sa = semi-aquatic, T = terrestrial,

T-Ar = terrestrial-arboreal.

SPECIES

CLASS AMPHIBIA

ORDER ANURA

HABITAT

Family Bufonidae

Bufo quercicus, Oak Toad

Bufo terrestris, Southern Toad

T

Family Hylidae

A cris gryllus dorsalis, Florida Cricket Frog

Hyla chrysoscelis. Gray Treefrog

Sa

Ar

Okefenokee Swamp Vertebrates

55

SPECIES

Hyla cinerea cinerea, Green Treefrog

Hyla crucifer bartramiana, Southern Spring Peeper

Hyla femoralis, Pine Woods Treefrog

Hyla gratiosa, Barking Treefrog

Hyla squirella, Squirrel Treefrog

Limnaoedus ocularis, Little Grass Frog

Pseudacris nigrita nigrita, Southern Chorus Frog

Pseudacris ornata. Ornate Chorus Frog

Family Microhylidae

Gastrophryne carolinensis, Eastern Narrow-mouthed Toad

Family Pelobatidae

Scaphiopus holbrooki holbrooki, Eastern Spadefoot Toad

Family Ranidae

Rana areolata aesopus, Florida Gopher Frog

Rana catesbeiana, Bullfrog

Rana clamitans clamitans, Bronze Frog

Rana grylio, Pig Frog

Rana heckscheri, River Frog

Rana utricularia, Southern Leopard Frog

Rana virgatipes, Carpenter Frog

ORDER CAUDATA

Family Ambystomatidae

Ambystoma cingulatum, Flatwoods Salamander

Ambystoma opacum, Marbled Salamander

Ambystoma talpoideum, Mole Salamander

Ambystoma tigrinum, Tiger Salamander

Family Amphiumidae

Amphiuma means, Two-toed Amphiuma

Family Plethodontidae

Desmognathus fuscus auriculatus , Southern Dusky Salamander

Eurycea bislineata cirrigera. Southern Two-lined Salamander

Eurycea quadridigitata. Dwarf Salamander

Plethodon glutinosus glutinosus. Slimy Salamander

Pseudotriton montanus floridanus. Gulf Coast Mud Salamander

Stereochilus marginatus^, Many-lined Salamander

Family Salamandridae

Notophthalmus perstriatus. Striped Newt

Notoph thalamus viridescens louisianensis. Central Newt

ORDER TRACHYSTOMATA

Family Sirenidae

Pseudobranchus striatus sppA, Dwarf Siren

Siren intermedia intermedia. Eastern Lesser Siren

Siren lacertina. Greater Siren

HABITAT

Ar

Sa

F,Sa

F

Aq

Sa

F'

Aq

Sa

T

Sa

Aq

Sa

Aq

56

Joshua Laerm, et al.

SPECIES HABITAT

CLASS REPTILIA

ORDER CROCODILIA

Family Alligatoridae

Alligator mississippiensis, American Alligator Sa

Caiman sclerops^. Spectacled Caiman Sa

ORDER SQUAMATA

Family Anguidae

Ophisaurus attenuatus longicaudus, Eastern Slender Glass Lizard T

Ophisaurus compressus, Island Glass Lizard T

Ophisaurus ventralis, Eastern Glass Lizard T

Family Iguanidae

Anolis carolinensis, Green Anole Ar

Sceloporus undulatus undulatus, Southern Fence Lizard T-Ar

Family Scincidae

Eumeces egregius similis. Northern Mole Skink F

Eumeces fasciatus, Five-lined Skink T-Ar

Eumeces inexpectatus, Southern Five-lined Skink T-Ar

Eumeces laticeps, Broad-headed Skink Ar

Scincella laterale, Ground Skink T

Family Teiidae

Cnemidophorus sexlineatus sexlineatus, Six-lined Racerunner T

Family Colubridae

Cemophora coccinea copei, Northern Scarlet Snake F

Coluber constrictor priapus. Southern Black Racer T-Ar

Diadophis punctatus punctatus. Southern Ring-necked Snake T

Drymarchon corais coupe ri, Indigo Snake T

Elaphe guttata guttata, Corn Snake T-Ar

Elaphe obsoleta quadrivittata. Yellow Rat Snake T-Ar

Elaphe obsolete spiloides, Gray Rat Snake T-Ar

Earancia abacura abacura. Eastern Mud Snake Aq

Earancia erytrogramma, Rainbow Snake Aq

Heterodon platy rhinos, Eastern Hognose Snake T

Heterodon simus. Southern Hognose Snake T

Lampropeltis calligaster rhombomaculata. Mole Snake F

Lampropeltis getulus getulus. Eastern Kingsnake T

Lampropeltis getulus getulus x floridana, intergrade kingsnake T

Lampropeltis triangulum elapsoides. Scarlet Kingsnake T-Ar

Masticophis flagellum flagellum. Eastern Coach whip T-Ar

Nerodia cyclopion floridana, Florida Green Water Snake Aq

Nerodia erythrogaster erythrogaster. Red-bellied Water Snake Sa

Nerodia fasciata fasciata. Banded Water Snake Sa

Nerodia fasciata pictiventris, Florida Water Snake Sa

Nerodia taxispilota. Brown Water Snake Sa

Opheodrys aestivus. Rough Green Snake Ar

Pituophis melanoleucus mugitus, Florida Pine Snake T

Regina alleni. Striped Swamp Snake Aq

Regina rigida rigida. Eastern Glossy Water Snake Aq

Okefenokee Swamp Vertebrates

57

Rhadinaea flavilata, Pine Woods Snake T

Seminatrix pygaea pygaea, North Florida Black Swamp Snake Aq

Storeria dekayi victa, Florida Brown Snake T

Storeria occipitomaculata obscura, Florida Red-Bellied Snake T

Thamnophis sauritus sackeni, Eastern Ribbon Snake T

Thamnophis sirtalis sirtalis, Eastern Garter Snake T

Virginia striatula. Rough Earth Snake T

Virginia valeriae valeriae, Eastern Smooth Earth Snake T

Family Elapidae

Micrurus fulvius fulvius, Eastern Coral Snake F

Family Viperidae

Agkistrodon piscivorus conanti, Florida Cottonmouth Sa

Crotalus adamant eus, Eastern Diamondback Rattlesnake T

Crotalus horridus atricaudatus, Canebrake Rattlesnake T

Sistrurus miliarius barbouri. Dusky Pigmy Rattlesnake T

ORDER TESTUDINATA

Family Chelydridae

Chelydra serpentina serpentina, Common Snapping T urtle Aq

Macroclemys temmincki, Alligator Snapping Turtle Aq

Family Emydidae

Chrysemys nelsoni, Florida Redbelly Turtle Aq

Deirochelys reticularia reticularia, Eastern Chicken Turtle Sa

Pseudemys {= Chrysemys) floridana floridana, Florida Cooler Aq

Pseudemys { = Chrysemys) scripta elegans, Red-eared Slider Aq

Pseudemys { = Chrysemys) scripta scripta, Yellowbelly Slider Aq

Terrapene Carolina bauri, Florida Box Turtle T

Terrapene Carolina Carolina, Eastern Box Turtle T

Family Kinosternidae

Kinosternon bauri palmarum. Striped Mud Turtle Sa

Kinosternon subrubrum subrubrum. Eastern Mud Turtle Aq

Stemotherus minor minor. Loggerhead Musk Turtle Aq

Sternotherus odoratus,Si\nk\)oi Aq

Family Testudinidae

Gopherus polyphemus. Gopher Tortoise T

Family Trionychidae

Trionyx ferox, Florida Softshell Aq

'Larvae aquatic, adults aquatic and/or terrestrial during breeding only.

^May occur around the periphery of the swamp.

’Three subspecies are recognized in the region (Conant 1975). It is possible that each occurs

on different sides of the swamp.

^An introduced species. Known from a single specimen. It is not known if a population

exists, although populations are apparently established in Florida.

BIRDS

Historical Foundations

The earliest observations on birds of the Okefenokee Swamp were

made by W. Bartram (1958). There is, however, some doubt that he ac-

tually visited the swamp (Harper 1920). In the 1880s C.F. Batchelder and

58

Joshua Laerm, et al.

M. Thompson (see Wright and Harper 1913) recorded observations on

birds, but the most significant avian surveys in the swamp were not to

begin until 1912 and the Cornell expeditions. Wright and Harper (1913)

listed 94 bird species based on the first expedition. Subsequent avian sur-

veys were conducted by Cornell personnel until 1937, but only incidental

accounts were published (Wright 1926). Frederick V. Hebard, an

amateur ornithologist, accumulated records of birds of the swamp for

almost 50 years. Although he published an annotated winter bird

checklist (Hebard 1941), the completion of a more comprehensive list

was interrupted by his death. Extensive observations of birds compiled

by H.A. Carter, E.R. Green, and F.V. Hebard were to be published. This

was never realized because of disagreement and communication prob-

lems among the authors.

A number of ONWR biologists (especially H.A. Carter, R.J.

Fleetwood, E. Cypert, and L. Walker) have made significant contribu-

tions to our knowledge of Okefenokee birds. Carter compiled the most

extensive and detailed reports from 1940 to 1942. Later, Fleetwood

collected bird notes and conducted a series of quantitative breeding cen-

suses (Fleetwood 1947a, 1947b, 1948). Cypert organized the bird records

and compiled the current checklists (Anon. 1971, 1974). Recent surveys

have been conducted by J. Meyers.

Comparison With Regional Fauna

The Okefenokee Swamp is an important wintering and breeding

area for approximately 232 bird species (Table 3). The coastal region of

Georgia, including the swamp, and the Southeast in general have fewer

breeding bird species but higher winter bird densities and species richness

than the northeastern United States. Approximately 120 bird species are

known to breed in the Coastal Plain of Georgia (Burleigh 1958), although

fewer breeding birds actually are found in the Okefenokee Swamp. Salt

marsh or coastal breeding birds generally are not observed in the swamp

but occasionally are recorded during spring and fall migrations and

following strong eastern winds (Table 3). Several Coastal Plain breeding

species are either absent or unconfirmed from the swamp (Table 4). Ab-

sence of some of these species is due to lack of appropriate breeding

habitat, but lack of observations for others may be due to their rarity.

The wintering birds of the swamp are similar to those of the sur-

rounding Coastal Plain with only a few exceptions. The most notable one

is the large population of Sandhill Cranes, Grus canadensis pratensis,

occurring on the refuge (Sanderson 1977).

Habitat Distribution of the Birds

Birds, perhaps more than other vertebrates, are habitat specific. The

diversity of Okefenokee Swamp habitats, in comparison to the sur-

rounding uplands, at least partly accounts for the large diversity of resi-

dent, breeding, and wintering species. While the specified habitat types

Okefenokee Swamp Vertebrates

59

(see above) in the swamp have, for the most part, differing avifaunas,

quantitative investigation of several of these is not yet complete and

others (pure cypress and shrub swamp) have just begun. Despite this it is

possible, based on present knowledge, to provide a fairly accurate general

picture of the habitat distribution of birds in the swamp by grouping the

previously identified habitat types into more general categories: swamp

forest, upland forest, and prairie (Table 3). These general categories in-

clude all specific habitat types known to occur in the swamp and sur-

rounding uplands.

Swamp forests include bay, cypress, and blackgum forests in various

stages of succession as well as a considerable portion of the shrub

swamps. Upland forests are dominated by slash and longleaf pine with

associated understory. Remnants of hardwood and mixed pine-

hardwood forests exist in the upland forests, but prescribed burning has

substantially reduced them. Upland forests are found on several swamp

islands, but generally occur on the periphery of the swamp. Prairies con-

tain grass-sedge meadows and aquatic macrophytes as described, with

treehouses (small floating islands with an overstory) and shrub batteries

scattered more or less throughout.

Table 3. List of birds of the Okefenokee Swamp and their status (A = accidental,

B = breeding, H = hypothetical, R = permanent resident, S = summer

resident, T = transient, W = winter resident). Based on Wright and

Harper (1913), Hebard (1941), checklists (Anon. 1971, 1974), ONWR

records, and personal observations (J. Meyers). All common names are

those standardized and listed with scientific names by the American

Ornithologists’ Union check-list committee (A.O.U. 1957, 1973, 1976).

P = prairie, Sf = swamp forest, Uf = upland forest.

60

Joshua Laerm, et al.

Okefenokee Swamp Vertebrates

61

62

Joshua Laerm, et al.

Okefenokee Swamp Vertebrates

63

64

Joshua Laerm, et al.

SPECIES STATUS

Catharus guttata, W

Catharus ustulata, Swainson’s Thrush T

Catharus minima, Gray-cheeked Thrush T

Catharus fuscescens, Veery T

Sialia sialis. Eastern Bluebird B,R

Family Sylviidae

Polioptila caerulea. Blue-gray Gnatcatcher B,S

Regulus satrapa. Golden-crowned Kinglet W

Regulus calendula. Ruby-crowned Kinglet W

Family Montacellidae

Anthus spinoletta,y^?^.QX p'xpQi W

Family Bombycillidae

Bombycilla cedrorum. Cedar Waxwing W

Family Laniidae

Lanius ludovicianus , Loggerhead shrike B,R

Family Sturnidae

Stumus vulgaris. Starling B,R

Family Vireonidae

Vireo griseus. White-eyed Vireo B,S

Vireoflavifrons, Yellow-throated Vireo B,S

Vireo solitarius. Solitary Vireo W

Vireo olivaceus. Red-eyed Vireo B,S

Family Parulidae

Mniotilta varia. Black-and-white Warbler W

Protonotaria citrea, Prothonotary Warbler S,B

Limnothlypis swainsonii, Swainson’s Warbler S,B

Helmintheros vermivorus. Worm-eating Warbler T

Vermivora chrysoptera, Golden-winged Warbler T

Vermivora pinus. Blue-winged Warbler T

Vermivora bachmanii, Bachman’s Warbler H

Vermivora celata. Orange-crowned Warbler W

Parula americana. Northern Parula B,S,W

Dendroica petechia. Yellow Warbler T

Dendroica magnolia. Magnolia Warbler T

Dendroica tigrina. Cape May Warbler T

Dendroica caerulescens. Black-throated Blue Warbler T

Dendroica coronata, Yellow-rumped Warbler W

Dendroica virens. Black-throated Green Warbler T

Dendroica cerulea. Cerulean Warbler T

Dendroica fusca, Blackburnian Warbler T

Dendroica dominica. Yellow-throated Warbler B,S,W

Dendroica pensylvanica. Chestnut-sided Warbler T

Dendroica striata, Blackpoll Warbler T

Dendroica pinus, V\x\QVxIdiXb\Qx B,R

Dendroica discolor. Prairie Warbler S

Dendroica palmarum. Palm Warbler W

Seiurus aurocapillus,OyQX\h\xd T

Seiurus noveboracensis. Northern Waterthrush T

HABITAT

Uf

Sf,Uf

p

Uf

Sf,Uf

Uf

Sf,Uf

Uf

Sf

Uf

Sf

Uf

Sf,Uf

Uf

P,Sf,Uf

Uf

P,Uf

Uf

Sf

Okefenokee Swamp Vertebrates

65

66

Joshua Laerm, et al.

SPECIES STATUS HABITAT

A/^/o5/7/za Song Sparrow W P,Uf

'Last reported in 1948; current status unknown.

Table 4. Georgia Coastal Plain breeding birds that are absent or unknown as

breeding birds in the Okefenokee Swamp.

MAMMALS

Historical Foundations

The earliest account of the mammals of the Okefenokee Swamp

(Jones 1876), though largely anecdotal, resulted from a joint expedition

Okefenokee Swamp Vertebrates

67

sponsored by the State Geological Survey and X\\q Atlanta Constitution in

1875. Somewhat more extensive observations resulted from a brief survey

by B.T. Gault in 1903 (reported in Harper 1927). The mammals of the

swamp, however, are known primarily as a result of the work of F. Har-

per. Harper’s (1927) classic Mammals of the O kef inokee Swamp, based on

surveys conducted by him, other Cornell personnel, and local hunters,

represents the definitive work on the mammals of the region. Subsequent

observations and records by Refuge personnel, particularly R.J.

Fleetwood, have provided valuable historical data including the first

records of Myotis austroriparius and Tadarida brasiliensis. More recently

small mammal surveys have been conducted in the swamp and sur-

rounding uplands by L. Logan and J. Laerm. Significant collections of

mammals from the swamp have been deposited in the Philadelphia

Academy of Natural Sciences, Cornell University, Florida State

Museum, National Museum of Natural History, and University of

Georgia Museum of Natural History.

Comparison With Regional Fauna

The 48 species and subspecies of mammals known to occur in the

Okefenokee Swamp and surrounding uplands represent a typical

southeastern fauna (Table 5). It is interesting to note that 12 (25%) of the

mammals occurring in or around the swamp are at or very near the limits

of their ranges. The Okefenokee Swamp region of Georgia and adjacent

Florida represents the southern limit of the range of Condylura cristata

and Castor canadensis, and the northern limit of the range of Neofiber

alleni. The southern limit of the ranges of the subspecies Cryptotis parva

parva, Eptesicus fuscus fuscus, and Sciurus niger niger occurs in the region

of the swamp and the northern limits of Geomys pinetis floridanus, Ursus

americanus floridanus, and Procyon lotor elucus occur also at or near the

swamp. In general the subspecific affinities of these mammals is poorly

understood and to a large extent suspect. Proper subspecific affiliation

must therefore await systematic study.

The composition of the mammalian fauna of the swamp has

changed little since Harper’s (1927) surveys. There are, however, ap-

parent changes in population level of a number of species.

Nycticeius humeralis, once the most common bat in the swamp (Har-

per 1927) is one of the most uncommon today. Its decrease is probably

due to the decrease in man-made structures, common nursery colony

sites (Watkins 1972), since the establishment of ONWR. Similarly, pop-

ulations of Plecotus rafinesquii, Rattus rattus, and Mus musculus have ap-

parently decreased due to the reduction of such structures.

The most striking change in the mammalian fauna of the swamp has

been the extirpation of Canis rufus niger and probably Felis concolor

coryi. These two carnivores were becoming rare during Harper’s surveys.

There have been a few unconfirmed recent reports of Felis in the area.

Odocoileus virginianus was driven nearly to local extinction by hunters

68

Joshua Laerm, et al.

during the first two decades of this century. Harper (1927) reported see-

ing only a single live individual during his extensive field work from 1912

and 1922. Since that time populations have recovered dramatically due in

large part to introductions and the establishment of the ONWR. Today

deer are the most common large mammals on the refuge.

Dasypus novemcinctus, introduced into Florida after Harper’s sur-

veys (Humphrey 1974) has extended its range significantly in recent

years. Although it moved into Georgia in the 1950s (Fitch et al. 1952) it

was not known from the swamp until 1968. Today it extends well up the

Coastal Plain.

Habitat Distribution of the Mammals

The habitat distribution of mammals occurring in the swamp and

surrounding uplands, insofar as it is presently known, is shown in Table

5. With few exceptions it is difficult to define a single habitat or assem-

blage of habitats to which a particular mammal is restricted; the majority

may be found in most of them. Nine of the mammals occurring within

the swamp and surrounding uplands have been reported from every

habitat defined within the swamp. These include Pipistrellus subflavus,

Lasiurus seminolus, Sylvilagus palustris, Sciurus carolinensis, Peromyscus

gossypinus, Ursus americanus, Procyon lotor, Lynx rufus, and Odocoileus

virginianus. Only two species, Geomys pinetis and Neofiber alleni, are

restricted to a single habitat. The former occurs only in well-drained

sandy uplands, while the latter is restricted to boggy prairies (Harper

1927; Birkenholz 1972).

Table 5. List of mammals of the Okefenokee Swamp. Based on museum records.

Harper (1927), and personal observations (L. Logan, J. Laerm). Scien-

tific and common names based on Jones et al. (1973). U = uplands,

I = islands, P = prairies, SS = shrub swamp, BG = blackgum forest,

PB = pure bay forest, PC = pure cypress, MC = mixed cypress.

SPECIES

ORDER MARSUPIALIA

Family Didelphidae

Didelphis virginiana pigra, Virginia Opossum

ORDER INSECTIVORA

Family Soricidae

Blarina carolinensis. Southern Short-tailed

Shrew

Cryptotis parva parva, Least Shrew

Family Talpidae

Scalopus aquaticus australis, Eastern Mole

Condylura cristata cristata. Star-nosed Mole

ORDER CHIROPTERA

Family Vespertilionidae

HABITAT

U,I,BG,PC

U,I

PC,MC

Okefenokee Swamp Vertebrates

69

SPECIES

Myotis austroriparius austroriparius, South-

eastern Myotis

Pipistrellus subflavus subflavus, Eastern

Pipistrelle

Eptesicus fuscus fuscus, Big Brown Bat

Lasiurus borealis borealis, Red Bat

Lasiurus seminolus, Seminole Bat

Lasiurus cinereus cinereus, Hoary Bat

Lasiurus intermediusfloridanus, Northern

Yellow Bat

Nycticeius humeralis humeralis, Evening Bat

Plecotus rafmesquii, Rafinesque’s Big-eared

Bat

Family Molossidae

Tadarida brasiliensis cynocephala, Brazilian

Free-tailed Bat

ORDER EDENTATA

Family Dasypodidae

Dasypus novemcinctus mexicanus\ Nine-

banded Armadillo

ORDER LAGOMORPHA

Family Leporidae

Sylvilagus palustris palustris, Marsh Rabbit

Sylvilagus floridanus mallurus, Eastern

Cottontail

ORDER RODENTIA

Family Sciuridae

Sciurus carolinensis carolinensis, Gray

Squirrel

Sciurus niger niger, Fox Squirrel

Glaucomys volans querceti, Southern

Flying Squirrel

Family Geomyidae

Geomys pinetis pinetis, Southeastern Pocket

Gopher

Geomys pinetis floridanus, Southeastern

Pocket Gopher

Family Castoridae

Castor canadensis carolinensis, Beaver

Family Cricetidae

Oryzomys palustris palustris. Marsh Rice Rat

Reithrodontomys humilus humilus. Eastern

Harvest Mouse

Peromyscus polionotus polionotus, Oldfield

Mouse

Peromyscus gossypinus gossypinus. Cotton

Mouse

HABITAT

I

U,I,P,SS,BG,PB,PC,MC

U,I,H

U,I,BG,MC

U,I,P,SS,BG,PB,PC,MC

I

U,BG

U,I,BG,PC

U,I

I

U,SS,MC

U,I,P,SS,BG,PB,PC,MC

U,I,SS

U,I,P,SS,BG,PB,PC,MC

U,I,PC,MC

U,I,BG,MC

U

U,I

u,i,p,ss,bg,pb,pc,mc

u

U,I

U,I,P,SS,BG,PB,PC,MC

70

Joshua Laerm, et al.

SPECIES

HABITAT

Ochrotomys nuttalli aureolus, Golden Mouse

Sigmodon hispidus hispidus, Hispid Cotton

Rat

Neotoma jloridana floridana, Eastern

Woodrat

Micro tus pinetorum parvulus, Woodland Vole

Neofiber alleni exoristus, Round-tailed

Muskrat

Family Muridae

Rattus rattus rattus\ Black Rat

Rattus rattus alexandrinus\ Black Rat

M us mus cuius musculus\ House Mouse

ORDER CARNIVORA

Family Canidae

Canis rufus niger, Red Wolf

Urocyon cinereoargenteus Jloridanus, Gray

Fox

Family Ursidae

Ursus americanus Jloridanus, Florida Black

Bear

Family Procyonidae

Procyon lotor elucus, Raccoon

Family Mustelidae

Mustela frenata olivacea, Long-tailed Weasel

Mustela visonmink, Mink

Mephitis mephitis elongata, Striped Skunk

Lontra canadensis vaga, River Otter

Family Felidae

Felis concolor coryi, Florida Panther

Lynx rufus Jloridanus, Bobcat

ORDER ARTIODACTYLA

Family Suidae

Sus scrofa\ Wild Pig

Family Cervidae

Odocoileus virginianus virginianus, White-

tailed Deer

U,I,BG,PC,MC

U,I

U,I,P,SS,BG,MC

U

P

U,I

U,I,MC

U,I,SS,MC

U,I,P,SS,BG,PB,PC,MC

U,I,PC,MC

U

U,I

U,P,PC,MC

U,I,P

U,I,P,SS,BG,PB,PC,MC

U,I,SS,BG,MC

U,I,P,SS,BG,PB,PC,MC

'Introduced.

DISCUSSION

At this time insufficient data are available to accurately assess the

patterns of habitat preference for all species or to quantify patterns of

species diversity in the various habitat types present in Okefenokee

Swamp. However, sufficient data are available to provide an accurate

faunal list and to allow some habitat correlation.

Okefenokee Swamp Vertebrates

71

A total of 419 vertebrate species or subspecies occurs in Okefenokee

Swamp and surrounding uplands. These include 36 fish, 37 amphibians,

66 reptiles, 232 birds, and 48 mammals. The vertebrates represent a fairly

typical southeastern Atlantic Coastal Plain fauna. There are no species

endemic to the swamp. In general vertebrate diversity in the swamp is

greater than in any area of similar size in the adjacent Southeast. This is

due primarily to the swamp’s habitat diversity, but another factor is the

prevalence of heavily managed pine forests throughout much of the adja-

cent southeastern region.

The role of ONWR in the preservation of the swamp as an

ecosystem is crucial, particularly insofar as threatened or endangered

wildlife is concerned. Eleven species or subspecies are considered either

threatened or endangered under the guidelines of federal and/or state

agencies (Table 6). The status of populations of these species in

Okefenokee Swamp has not yet been determined.

Table 6. List of Threatened or Endangered species in Okefenokee Swamp. Legal

status as of 15 February 1980, defined by United States Fish and Wild-

life Service (USFWS), Florida Game and Freshwater Fish Commission

(GFWFC), and Georgia Department of Natural Resources (DNR).

E = Endangered; T = Threatened.

SPECIES

Alligator mississippiensis, American Alligator

Drymarchon corais couperi, Eastern Indigo Snake

Mycteria americana, Woodstork

Haliaeetus leucocephalus, Bald Eagle

Falco peregrinus, Peregrine Falcon

Grus canadensis pratensis, Florida Sandhill Crane

Campephilus principalis, Ivory-billed Woodpecker

Picoides borealis, Red-cockaded Woodpecker

Vermivora bachmanii, Bachman’s Warbler

IJrsus americanus floridanus, Florida Black Bear

Felis concolor coryi, Florida Panther

STATUS

USFWS GFWFC DNR

T T E

T T T

E

E T E

E EE

T

E EE

E T E

E EE

T

E EE

ACKNOWLEDGMENTS.— acknowledge our indebtedness to

the many persons involved in collections and observations of the ver-

tebrates of the Okefenokee Swamp throughout this century. While in-

dividual efforts might have been small, their total contribution has been

great. We are especially grateful to those persons mentioned in the

72

Joshua Laerm, et al.

historical foundations sections above. The field assistance of C.G.

Tompson, G.S. Wise, J.H. Rappole, and J. Howland is much ap-

preciated. The understanding of our respective wives and ladies during

our seemingly endless trips to the swamp should not go unrecognized.

Finally, the assistance and support given us by ONWR personnel, es-

pecially J. Eadie and T. Wilkins, made our efforts possible. This work

was supported largely by NSF Grants DEB-78-08842 and DEB-79-

22633, and partly by grant-in-aid funds under section 6 of the En-

dangered Species Act of 1973 (P.L. 93-205). This is paper No. 1 1, Univer-

sity of Georgia Okefenokee Ecosystem Investigations and a contribution

of the University of Georgia Museum of Natural History.

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Okefenokee Swamp Vertebrates

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specific variation. J. Elisha Mitchell Sci. Soc. 67:(l):l-44.

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497 pp.

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reconnaissance of the Okefinokee Swamp in Georgia: The reptiles. Proc.

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pp. 107-139. in A biological reconnaissance of the Okefinokee Swamp in

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Accepted 25 August 1980

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New Records, Distribution and Diagnostic

Characters of Virginia Ictalurid Catfishes

With An Adnexed Adipose Fin

Noel M. Burkhead and Robert E. Jenkins

Department of Biology,

Roanoke College, Salem, Virginia 24153

AND

Eugene G. Maurakis

Environmental Affairs Department,

Potomac Electrical Power Company,

1900 Pennsylvania Avenue NW, Washington, DC 20068

ABSTRACT.— KQCQWi introductions of Ictalurus melas and Pylodictis

olivaris and the discovery (possible introduction) of /. brunneus has

raised the number of ictalurids with an adnexed adipose fin to eight

species in the Roanoke River drainage. Introduction of /. furcatus to

other drainages raised the Virginia total to nine. Although most of these

species are widely distributed in North America, none is native in all

Virginia drainages. Most species have been variously introduced, and /.

brunneus and I. platycephalus exhibit an atypical distributional in-

terrelationship. Key characters for separating these two flathead

bullheads from typical bullheads (/. melas, 1. natalis, 1. nebulosus) are

described, and some diagnostic characters different from those generally

used are emphasized for distinguishing I. brunneus from /. platycephalus,

and I. melas from /. nebulosus. These and characters of other ictalurids

with an adnexed adipose fin are discussed; a key is provided for the

species of Virginia drainages.

INTRODUCTION

Exceptional or significant new records, occasional recent misiden-

tifications, vexatious old records, and an atypical distributional in-

terrelationship between two Virginia ictalurids led to this report.

Although most species have been long known, the ictalurids with an ad-

nexed (free) adipose fin still present at least local problems in identifica-

tion, and consequently may subvert zoogeographic studies in North

America. Problems have extended elsewhere. For example, Banarescu

(1968) found that the bullhead widely introduced in Europe was actually

Ictalurus melas (Rafinesque) instead of I. nebulosus (Lesueur). Important

external characters are few, and most are variable and widely shared

among similar appearing, often sympatric species; no consistently present

external character state of juvenile and adult Ictalurus appears unique to

a single species.

The Roanoke River drainage is now known to harbor eight species

of ictalurids with an adnexed adipose fin, a larger complement than oc-

curs in the Mississippi River basin. A total of nine species is now known

from Virginia: /. brunneus (Jordan), snail bullhead; I. catus (Linneaus),

white catfish; I. furcatus (Lesueur), blue catfish; /. melas, black bullhead;

75

Brimleyana No. 4: 75-93. December 1980.

76

Noel M. Burkhead, et al.

/. natalis (Lesueur), yellow bullhead; /. nebulosus, brown bullhead; /.

platycephalus (Girard), flat bullhead; I. punctatus (Rafinesque), channel

catfish; Pylodictis olivaris (Rafinesque), flathead catfish. Of this assem-

blage, /. melas and Pylodictis are considered to be introduced, and /.

brunneus probably so, to the Roanoke drainage; the same is true for the

status of I. furcatus and /. brunneus in Virginia.

This report discusses the distribution of each species in Virginia and

extralimitally where pertinent. Diagnostic characters with the greatest

utility and ease in identifying these species within the study area are dis-

cussed and employed in a key. Osteological and other differences among

Ictalurus species are found in Paloumpis (1964), Yerger and Relyea

(1968), Smith and Lundberg (1972), and Lundberg (1975).

Concepts of genera, subgenera, and species groups follow Lundberg

(1975). However, regarding our discussion of species identification, for

practical purposes we artificially group /. catus with the “forktail cat-

fishes,” /. furcatus and /. punctatus of the subgenus Ictalurus. Ictalurus

catus actually is placed, in the subgenus Amiurus, in the catus group with

the species we collectively refer to as “flathead bullheads,” i.e., I. brun-

neus, /. platycephalus and /. serracanthus Yerger and Relyea. The other

three species, our “typical bullheads,” /. melas, /. natalis and /. nebulosus,

are referred by Lundberg to the natalis group of the subgenus Amiurus.

METHODS AND MATERIALS

Methods of counting and mensuration follow those outlined by

Hubbs and Lagler (1958) and Yerger and Relyea (1968), with one dif-

ference from the latter study. Removal of the gill arch for gill raker

counts was necessary only in the smallest specimens; otherwise a slit at

the dorsal and ventral junctions of the operculum, and adduction of the

latter, were sufficient to expose gill rakers. All rakers on the right arch in-

cluding rudiments on lower limb, were counted; fused rakers were coun-

ted as one. To count anal rays it was necessary to expose them by slitting

the anal fin base and peeling the skin back. All anterior rudimentary rays

were counted; the last two rays with a basal conjuncture were counted as

one.

Measurements were made using Helios dial calipers for all propor-

tionally expressed characters (as % SL) and for standard length (SL) of

smaller specimens; they were recorded to the nearest 0.1 mm. The SLs of

large specimens were obtained with a beam compass and a steel rule, and

recorded to the nearest 0.5 mm. Counting was aided by the use of a

variable magnification stereo dissecting microscope. The counts from

one I. melas, 33.6 mm SL, were omitted from tabulation due to extreme

low counts (rakers incompletely developed).

Complete locality data on specimens examined are on file at

Roanoke College. Flathead bullhead localities are depicted in Figure 1

and are listed in sequence from downstream to upstream. Typical

bullhead localities are presented by basin or drainage and therein

Virginia Ictalurid Catfishes

77

alphabetically by state and tributary. All specimens were from Virginia

drainages, except for /. melas, which was supplemented by material from

other states. Localities for both sections are followed by institutional ab-

breviation and catalog number. Roanoke College and Virginia Com-

monwealth University are followed by collector’s initials and field num-

ber.

Institution and agency abbreviations used are:

ACE, U.S. Army Corps of Engineers

CU, Cornell University

DPC, Duke Power Company

FWS, U.S. Fish arid Wildlife Service

EC, Lynchburg College

RC, Roanoke College

SCS, Soil Conservation Service

UMMZ, University of Michigan Museum of Zoology

UNC, University of North Carolina at Charlotte

USNM, National Museum of Natural History, Smithsonian

UT, University of Tennessee at Knoxville

VCGIF, Virginia Commission of Game and Inland Fisheries

VCU, Virginia Commonwealth University

VFU, Virginia Cooperative Fisheries Unit

VIMS, Virginia Institute of Marine Science

VPI, Virginia Polytechnic Institute and State University

Collections from the following sources are housed at Roanoke College:

ACE, FWS, LC, SCS, VCGIF; some of the collections originally at VCU

are also at Roanoke College. Numbers that follow these series refer to

collection reference numbers used for a data bank concerning the

freshwater fishes of Virginia.

Ictalums brunneus

Dan River system. - VA: Dan R. RC VCGIF 230; RC FWS 8; NC:

Country Line Creek RC ACE 15; Rattlesnake Creek RC ACE 18; VA:

RC VCGIF 229; NC: Pumpkin Creek RC ACE 16; VA: Dan R. RC

EGM Va-23; Fall Creek RC ACE 2; Dan R. RC FWS 6; Dan R. RC

FWS 1; NC: Dan R. DPC 50101-09 and -11; DPC 50101-10, 50107-06;

DPC 50101-18; DPC 50101-12 and -19; UNC 76-95.

Ictalums platycephalus

Chowan River system. — VA: Great Creek RC SCS 8; N. Meherrin R.

RC HJP 44, VCU HJP 100.

Lower Roanoke River system (below Dan River mouth). — VA: Flat

Creek VPI 1029; Miles Creek VPI 1037; NC: Grassy Creek RC REJ 865;

VA: Beaver Pond Creek RC REJ 863.

Lower Dan River system (below Smith River mouth). — VA: Banister.

R. VCU HJP 84; NC: Cascade Creek ACE 89; Dan R. DPC 50107-07.

Smith River and tributaries (Dan River system). — VA: Leatherwood

78

Noel M. Burkhead, et al.

Creek RC ACE 80; Beaver Creek CU 13921; Town Creek VPI 984;

Green Brook RC ACE 134; Smith R. RC HJP 60.

Upper Dan River system (above Smith River mouth). — NC: Buffalo

Creek RC ACE 108; Jacob Creek RC ACE 115; Belews Lake DPC

50107-12 and -18.

Upper Roanoke River system (above Dan River mouth). — VA: Dif-

ficult Creek RC REJ 856; Twittys Creek VFU 109; Wards Fork USNM

101324; Turnip Creek RC REJ 873; Falling R. RC HJP 59; Little Falling

R. RC HJP 75; Seneca Creek VCU HJP 82; trib. Little Otter R. LC 30;

Leesville Reservoir RC REJ 333; Pigg R. RC REJ 402; Blackwater R.

VPI 989; Maggodee Creek VPI 990, 974 and CU 43587 (split collection);

Ellie Creek VPI 975; Blackwater R. VPI 690; 2187; South Fork

Blackwater R. VPI 2188; North Fork Blackwater R. VPI 1755.

Ictalurus natalis

York River drainage. — VA: Pond Creek RC JRR 124; Smoots Pond

RC JRS 23; Ta River RC JRR 134.

James River drainage. — VA: Barrows Creek RC TZ 156; Maury R.

RC NMB 73; Tuckahoe Creek RC JRR 131.

Roanoke River drainage. — VA: Beaver Pond Creek RC REJ 863;

Great Creek RC SCS 8; Lake Jordan RC JRS 22; Little Buffalo Creek

RC REJ 869; Roanoke R. RC REJ 781.

Tennessee River drainage. — VA: Clinch R. RC REJ 503; Clinch R.

RC REJ 611; North Fork Holston R. RC NMB 153; RC NMB 157.

Ictalurus nebulosus

York River drainage. — VA: Bunch Creek RC SCS 24; Smoots Pond

RC JRS 23.

James River drainage. — VA: Herring Creek RC TZ 157; Jordans

Branch Creek RC VCU-B-JB-1.

Roanoke River drainage. — NC: Anderson Swamp Creek RC REJ

867; Belews Lake DPC 50106-15; Flat Creek RC REJ 866; VA: Back

Creek RC WJM; Ballows Creek RC ACE 34; Banister R. RC HJP 80;

Beaver Pond Creek RC REJ 863; Dan R. RC FWS 6; Falling R. RC REJ

815; Grassy Creek RC REJ 860; Green Branch RC ACE 134; Lawsons

Creek RC ACE 26; Mason Creek RC REJ 524; Old Woman Creek RC

REJ 401; Pigg R. RC REJ 402; RC DLJ 26; RC DLJ 6; Lake Drummond

VPI 1218; Lake Jordan RC JRS 22.

New River drainage. — VA: Meadow Creek RC JRR 207.

Ictalurus melas

Roanoke River drainage. — NC: Belews Lake DPC 50105-07; VA:

Grassy Creek RC REJ 860; Little Buffalo Creek RC REJ 869.

Peedee River drainage. — NC: trib. Yadkin R. UMMZ 138401.

Tennessee River drainage. — TN: Big Sandy R. UT 48. 1 14; Dry Creek

UT 48.202; Duck R. UT 48.294; Sims Spring Branch UT 48.284; VA:

Virginia Ictalurid Catfishes

79

Copper Creek RC REJ 348.

Cumberland River drainage. — TN: East Fork Stones R. UT 48.7.

Green River drainage. — TN: Hurricane Creek RC REJ 560.

Coosa River drainage. — TN: Coahulla Creek UT 48.57; Mill Creek

UT 48.56; UT 48.285.

Mississippi River basin. — TN: backwater Mississippi R. UT 48.26.

Hatchie River drainage. — TN: ditch UT 48.109.

Forked Deer River drainage. — TN: Nixon Creek UT 48.249; slough

UT 48.250.

Red River drainage. — LA: ditch VPI 2758; Shepherd Bayou VPI

2342.

Sabine River drainage. — LA: Sabine R. VPI 2652.

DISTRIBUTION

The ictalurids with an adnexed adipose fin generally occur in

moderate to large streams and main river channels of all physiographic

provinces in Virginia except the Blue Ridge, from which they are essen-

tially absent except for upper New River. Most species readily adapt to

reservoir and farm pond habitats, and a few, notably /. catus, I.furcatus

and /. punctatus, tolerate estuarine conditions. One species, /. brunneus,

commonly occurs in moderate currents (Yerger and Relyea 1968; Bryant

et al. 1979; D. Cloutman, pers. comm.) as well as sluggish currents and

backwaters with soft bottoms (M. Corcoran, pers. comm.), which are

typically inhabited by the remaining species. When collected during

daylight most of these species are associated with cover such as undercut

banks, logs and boulders. The following discussion includes considera-

tion of native or introduced status in the drainages.

Ictalurus brunneus. — The snail bullhead is known in the Roanoke

drainage only from the Dan River system above Kerr Reservoir, North

Carolina and Virginia (Fig. 1). It was first collected from the lower Dan

in 1976 just above this reservoir, and was subsequently taken in low num-

bers from the main channel and a few tributaries. Prior to the Dan

records, Yerger and Relyea (1968) reported its northern limits as the up-

per Cape Fear and Peedee River drainages. North Carolina, both adja-

cent on the south to the Dan.

The distributional relationship in the Roanoke drainage of /. brun-

neus and /. platycephalus, the closest relative of /. brunneus (Lundberg

1975), appears unique. Yerger and Relyea (1968) found that, although

the species are broadly sympatric and occasionally syntopic in several

drainages, /. brunneus tends to be more frequently found in, and perhaps

differentially favors, higher gradient areas in the upper parts of those

drainages. In the Mobile drainage, where only /. brunneus occurs, this

species was found only in the upper section, in Georgia, over hard bot-

tom in riffles and moderate currents (Bryant et al. 1979). Although both

species occupy upper and lower reaches of many streams, this distribu-

tion pattern was not regarded as atypical since higher and lower gradient

80

Noel M. Burkhead, et al.

t

—

NC 'X'l

O Ictalurus platycephalus

^Ictalurus brunneus

/I

vi'

«0 hM

30 Mi

Fig. 1. Distribution of Ictalurus brunneus and I. platycephalus in the Roanoke

River drainage, North Carolina and Virginia.

regimes occur in many parts of these streams (Yerger and Relyea 1968).

Extensive surveys of the Neuse drainage for Ictalurus, and less extensive

surveys of other Carolinean Atlantic Slope drainages, revealed that I.

brunneus is more abundant, sometimes greatly so, than /. platycephalus

(M. Corcoran, pers. comm.). Corcoran also determined that, at least in

the Neuse, both species are generally absent from the Coastal Plain.

Thus, previous concepts of a preference by 7. brunneus for upper stream

sections may partly relate to its numerical abundance over /.

platycephalus. However, in the Roanoke drainage, only /. platycephalus

appears to currently occur in the main trunk Roanoke system and the

Smith River tributary of the Dan River. In both these systems it extends

upstream well through moderate gradients into Blue Ridge foothills.

The apparent absence of /. brunneus from most of the Roanoke

drainage, including the Chowan system of the lower Roanoke, and the

wide geographic and ecological range of /. platycephalus therein, suggest

that /. brunneus was recently introduced to the Dan. Prior absence of /.

brunneus would have allowed /. platycephalus to become widely es-

tablished. The apparent current exclusion of /. brunneus from montane

sections of the Dan system thus may relate to former establishment of /.

platycephalus.

Belews Lake, an upper Dan system impoundment (Fig. 1), was

reportedly stocked with /. melas by a “concerned citizen” to improve

fishing (W. Smith, pers. comm.). These introduced I. melas may have

Virginia Ictalurid Catfishes

81

been transferred from the Yadkin system of the Peedee, where both I.

brunneus and 1. melas occur. Although I. brunneus is not known from the

lake (D. Cloutman, pers comm.) it occurs in the immediate area, and the

/. melas stocking may have included the superficially similar /. brunneus.

Belews Creek was impounded in 1970 and the lake reached full pool in

1973 (Harrell et al. 1973). If /. brunneus dispersed from the Belews Lake

area, its mobility would have been similar to that of introduced Pylodictis

olivaris now spreading in the Cape Fear drainage (M. Corcoran, pers.

comm.). However, the Belews Lake area may not have been the point of

origin; possibly more than one stocking occurred.

Ictalurus catus. — White catfish are native to the major Atlantic slope

drainages of Virginia, occurring widely in Piedmont and Coastal Plain

parts of large streams and reservoirs. Jordan (1889) reported it from

Maury (North) River and elsewhere in the upper James drainage in the

Ridge and Valley. It also has been taken in South Fork Shenandoah

River (Potomac drainage) in the Ridge and Valley. Clay (1975) noted

that I. catus introduced to Kentucky were from the James River.

Ictalurus furcatus. — The presence in Virginia of the blue catfish, a

primarily Mississippi basin and Gulf slope species, has been widely

reported, but the species has only recently been verifed as introduced. It

was stocked in lower Rappahannock (1975 and 1977) and James (1977)

rivers by the Virginia Commission of Game and Inland Fisheries (L.

Hart, pers. comm.). Juvenile specimens from these stockings have since

been collected by Virginia Institute of Marine Science personnel (J.

Gourley, pers. comm.; VIMS specimens examined by us). It is not known

whether the species is reproducing. Ictalurus furcatus is unknown from

the Potomac, York, New, Roanoke and Tennessee (in Virginia) River

drainages. Records from the Potomac and New River drainages are dis-

cussed in detail.

Ictalurus furcatus may have been introduced into the Potomac River

near Washington, D.C., between 1898 and 1905. The old U.S. Fish Com-

mission rearing and holding ponds in that area were an early active center

of fish dispersal. It was not recorded by Smith and Bean (1898), but was

reported as introduced (probably with /. punctatus) based on 1905

records by Bean and Weed (1911), and by McAtee and Weed (1915)

based on two specimens collected in 1912. We located an adult /.

punctatus (USNM 70281) previously misidentified as /. furcatus,

apparently one of the specimens on which McAtee and Weed (1915)

based their record. Radcliffe and Welsh (1916) reported I. furcatus from

the Chesapeake and Ohio canal, along the Potomac River, Maryland.

The single specimen was reportedly sent to Washington, but it was not

found by us at the USNM. It is unknown whether /. furcatus was in-

troduced and failed to establish, or if all records are actually of /.

punctatus. Elser (1950) and Manville (1968) based their records of I. fur-

catus on these early reports. A second body of literature (Wiley 1970;

82

Noel M. Burkhead, et al.

Jenkins et al. 1972; Lee et al. 1976; Stauffer et al. 1978) reported I. fur-

catus from the Potomac based on records of Schwartz (1961). Frank J.

Schwartz (pers. comm.) later felt that these specimens were “odd /.

punctatus"\ no Potomac /. furcatus were found in collections of

Chesapeake Biological Laboratory, Lfniversity of North Carolina In-

stitute of Marine Sciences, and Virginia Institute of Marine Science,

which house Schwartz’s collection (F. Schwartz, J. Stauffer, J. Gourley,

pers. comm.). Ictalurus furcatus has not been collected in recent extensive

surveys of the Potomac River from Maryland— West Virginia (Energy

Impact Associates), along Virginia above Great Falls (E. Enamait, pers.

comm.), or from Washington, D.C., downstream (J. Gourley, pers.

comm.). If ever introduced into the Potomac River near Washington,

D.C., it probably shares extirpated status with Percopsis omiscomaycus

(Walbaum) and Percina caprodes (Rafmesque).

In the New drainage, /. furcatus was reported introduced into the

West Virginia section (Schwartz in Jenkins et al. 1972), but no specimens

were seen. Cope’s (1868) record of /. “caerulescens” in the Virginia sec-

tion was based on /. punctatus (Fowler 1945:81). Addair’s (1944) records

from West Virginia of /. “anguilla’ probably are of only /. punctatus. His

New River, West Virginia, specimens at the UMMZ are /. punctatus.

Hocutt et al. (1978) listed I. furcatus as a hypothetical inclusion to the

Greenbrier River fauna based on Addair (1944). Ross (1959) repeated

reports by game wardens of “blue catfish” from the New River in three

Virginia counties. Also, a single record was reported (specimen discar-

ded) by personnel of the VCGIF from Claytor Lake, New River im-

poundment. The above two reports of I. furcatus are considered to be of

I. punctatus, based on the absence of I. furcatus from extensive New River

surveys by Hocutt et al. (1973), Stauffer et al. (1975, 1976) and others,

and because nonspotted channel catfish have often been misidentified as

blue catfish.

Ictalurus melas. — The black bullhead probably is native to Virginia in

only the Tennessee and Big Sandy drainages. Until recent collections in

the Roanoke drainage, it was thought to be absent from Atlantic slope

drainages. Hence, we considered records of collections and literature

compilations (Abbott et al. 1977) for I. melas to be I. nebulosus, a species

with which it is sometimes confused. However, recent records from

Belews Lake (see I. brunneus), from Dan River above Belews Lake, North

Carolina (UNC 76-93), from two Virginia tributaries of Kerr Reservoir,

and two specimens (UMMZ 138480) taken in 1940 from the North

Carolina section of the upper Peedee drainage, prompted us to reconsider

records from the Atlantic slope. Collections of I. nebulosus from the

Roanoke drainage (including Kerr Reservoir preimpoundment collec-

tions and the Chowan system) in Virginia were examined, and no I. melas

were discovered. Also, none were reported from extensive surveys of the

North Carolina parts of the Roanoke and Chowan systems (Smith 1963;

Carnes 1965). The absence of I. melas from earlier collections strongly

Virginia Ictalurid Catfishes

83

suggests that its presence in the Roanoke is the result of single or multiple

introductions. Because of difficulties of identification, until specimens

are examined we still consider /. melas to be absent elsewhere on the

Atlantic slope in Virginia. Other species recently introduced into the

Roanoke drainage in the North Carolina part of the Dan at Belews Lake

are Notropis lutrensis (Baird and Girard) and Pimephales promelas

(Rafinesque) (DPC 30407-04 and 31201-02, respectively).

The occurrence of /. melas in the New River drainage is also

problematic. It appears to have been introduced but now possibly extir-

pated. The only extant specimens known are four juveniles from Fries, a

town on New River, taken in 1939 by B. Smith (USNM 109467). The

only other record of /. melas is from Reed Creek at Wytheville based on

unretained specimens (Wollitz 1968). Wollitz (pers. comm.) thought the

Reed Creek specimens resulted from introduction. Ictalurus melas has

not been taken in recent extensive New River and tributary surveys in

Virginia, or from New River tributaries in West Virginia (Hocutt et al.

1978, 1979). Hocutt et al. (1978) reported /. melas as stocked in Sherwood

Lake, Greenbrier River system. West Virginia. Like other bullheads, it

may be widely introduced in farm ponds.

Ictalurus natalis. — The yellow bullhead is native to Virginia, occur-

ring in all drainages except the New. The only record for the latter, from

the Gauley River system of the lower New, West Virginia, may represent

an introduction (Hocutt et al. 1979).

Ictalurus nebulosus. — The brown bullhead is native to the Atlantic

slope of Virginia; it occurs in all Atlantic slope drainages as well as being

the only ictalurid known from the diminutive freshwater ichthyofauna of

the southern part of the Delmarva Peninsula. Ictalurus nebulosus is

probably introduced to the New drainage. In Virginia it is known from

only two collections, both from tributaries entering New River below

Claytor Lake: a juvenile (VPI 2039) was rotenoned in 1971 from East

River just above its mouth and immediately upstream from the Virginia

— West Virginia state line; and two juveniles were collected in 1972 from

Meadow Creek, Montgomery County. Hocutt et al. (1979) reported

another specimen taken in 1976 from a lower New River tributary

system. West Virginia. It may have been stocked in farm ponds in much

of the state, but is unknown from the Tennessee River drainage in

Virginia.

Ictalurus platycephalus. — The flat bullhead occurs only in the

Roanoke drainage, including the Meherrin River branch of the Chowan

system, in Virginia; this is the northern limit of its distribution (Yerger

and Relyea 1968; Fig. 1). The species generally occurs in small to

moderate-size streams draining the Piedmont, where it inhabits sluggish

waters and is known from reservoirs. In the Roanoke drainage it extends

into smaller streams than it is “typically” associated with elsewhere on

the Atlantic Slope. The possible historical absence of /. brunneus in the

84

Noel M. Burkhead, et al.

Roanoke drainage may have allowed it to invade smaller stream habitats

thought to be typically occupied by only I. brunneus when the two species

are sympatric. The first life history study of I. platycephalus was conduc-

ted by Olmstead and Cloutman (1979).

Ictalurus punctatus. — The channel catfish is native in the Tennessee

and Big Sandy drainages in Virginia and, based on Cope’s 1867 record

{sQQ 1. furcatus), probably native in the New River above Kanawha Falls.

It has been introduced in all Atlantic slope drainages in the state (Jenkins

et al. 1972).

Pylodictis olivaris. — The flathead catfish is native to the Tennessee,

Big Sandy, and New drainages in Virginia. It has recently been in-

troduced into the James and Roanoke drainages. Introduction into lower

James River near Surry accidentally occurred in 1965 when a temporary

holding pond at Hog Island Game Refuge washed out during a storm

and released about 50 P. olivaris. A 20 to 30 pound P. olivaris was seen by

Dean Estes (Virginia Electric Power Co. biologist) in 1977; it was taken

on a trotline near Surry (J. Gourley, pers. comm.). Hart (1978) reported

the introduction into Smith Mountain Reservoir (the most upstream

reservoir on the Roanoke River, Fig. 1) of one specimen in 1976 and five

in 1977. Specimens from 10 inches long to 10 pounds weight, taken from

the Roanoke River near Brookneal below Smith Mountain Lake, were

observed in 1978-79 by L. Hart (pers. comm.).

DIAGNOSTIC CHARACTERS

The following account and critique of distinguishing characters in-

cludes summaries of our data as well as characters abstracted from the

literature. Discussion of diagnostic features is supplemented by fre-

quency distributions of counts for Virginia Ictalurus (Amiurus) in Tables

1-3, comparison of eye sizes (Fig. 2), and fins (shape and pigmentation),

as well as premaxillary teeth configurations (Fig. 3). We emphasize that

the following discussion pertains to Virginia Ictalurus (except where sup-

plemented; see Methods and Materials), and is limited to characters with

known or reputed utility in identifying species. Mention of somatic and

fin pigmentation is generally avoided, as many aspects of coloration are

variable in all species of Ictalurus. Although the Virginia Ictalurus fauna

is artificially enriched in species, the species are easily distinguished. To

reduce redundancy, diagnostic features are discussed by the following

groups: the flathead bullheads, the typical bullheads, and the Ictalurus

with forked tails. Characters of the monotypic genus Pylodictis are listed

only in the key at the end of this section.

Flathead bullheads. — This group is represented in Virginia by I. brun-

neus and I. platycephalus. The species were clearly distinguished first by

Yerger and Relyea (1968), who recognized the flathead bullheads as a

group but did not provide a key character for its separation from the

Virginia Ictalurid Catfishes 85

Fig. 2. Relationship of eye length (as % SL) to SL in Virginia Ictalurus (Amiurus)

with emarginate caudal fins.

typical bullheads group. The flathead bullheads are best distinguished

from other bullheads by the presence of a large dark basal blotch, its up-

per edge straight or convexly rounded, in the dorsal fin (Fig. 3D). The

blotch was also recognized as a key character and figured for /.

platycephalus by Eddy (1969), and depicted for /. brunneus by Smith-

Vaniz (1968). Eye size is secondarily useful in separating the groups, the

size being moderate in flatheads and small in typical bullheads. Although

size varies allometrically relative to SL, more pronouncedly in small

juveniles (Fig. 2), the differences between the groups are generally ob-

vious, with little overlap when comparing specimens of similar lengths.

The third member of the flathead bullhead group, /. serracanthus, a

primarily Floridean species, also has the dorsal blotch and moderate eye

size character states (figure and description in Yerger and Relyea 1968).

Head shape of flathead and typical bullheads is variable, from essentially

flat to slightly convex dorsally in flatheads, versus usually more elevated

or markedly convex in typical bullheads. Overlap in head shape and eye

size is such that sole reliance on either character for group separation will

result in some misidentifications.

Ictalurus brunneus and /. platycephalus are best distinguished from

each other by barbel pigmentation and premaxillary teeth configuration,

and secondarily by meristics. Most juveniles and adults of I. brunneus

examined had profusely dark pigmented mental (chin) barbels, whereas

most specimens of /. platycephalus had unpigmented or slightly pigmen-

ted, pale mental barbels. The absence of profusely developed mental bar-

bel pigment in /. brunneus usually occurs in specimens smaller than 100

mm SL. In I. platycephalus the presence of slightly pigmented mental bar-

bels occurs mostly in adults, particularly in the lateral pair of barbels; the

86

Noel M. Burkhead, et al.

medial mental barbels rarely possess melanophores, and then only

basally. Additionally, the maxillary barbels of 1. platycephalus usually ap-

pear bicolored (leading edge pale, posterior edge dark), whereas in /.

brunneus these barbels are uniformly dark.

Fig. 3. Diagnostic features of some Virginia ictalurids: A, ventral aspect showing

premaxillary tooth patch of Pylodictis; B, premaxillary tooth patch of I. brun-

neus; C, premaxillary tooth patch of /. platycephalus; D, flathead bullhead dorsal

fin with dark basal blotch; E, dorsal fin of typical bullhead; F, fin profiles of /.

furcatus; G. fin profiles of /. punctatus; FI, fin profiles of I. catus; I, profile of an

emarginate caudal fin.

The premaxillary tooth patch of /. brunneus differs from /.

platycephalus in being wider, fairly uniform in width, and usually possess-

ing lateral edentations in the patch (Fig. 3B). The cardiform teeth of /.

brunneus are more numerous along an anterior-posterior axis (teeth not

forming rows) than in /. platycephalus, and occur in two distinct sizes.

Large cardiform teeth are positioned along the anterior margin of the

tooth patch, and are often additionally arranged in a medial, triangular

configuration. The tooth patch of /. platycephalus usually lacks lateral

edentations and is occasionally slightly constricted medially (Fig. 3C).

Virginia Ictalurid Catfishes

87

The cardiform teeth of /. platycephalus are small and fairly uniform in

size. The premaxillary tooth patch and cardiform teeth size differences

are not evident in small specimens. These characters were first recognized

by Lundberg (1970).

The greatest meristic differences between /. brunneus and I.

platycephalus are in anal rays and a character index (Tables 2 and 3). Fre-

quency ranges of all counts differed slightly from data of Yerger and

Relyea (1968), indicating possible geographic variation. A slightly higher

range of character index values exists in Roanoke drainage /. brunneus

when compared to data of Yerger and Relyea (1968) from some more

southerly drainages, and results in greater meristic overlap between the

two species in the Roanoke.

The difference in mouth position between the species conformed to

Yerger and Relyea’s (1968) description; however, we do not advocate

general use of the character, as the difference seems to be only an average

one and is not as obvious as barbel pigmentation.

Typical bullheads. — Three species of typical bullheads (or the natalis

group of Lundberg 1975) inhabit Virginia waters: /. natalis, /. nebulosus

and /. melas. These are best separated from the flathead bullhead group

by the absence of a discrete dark blotch at the base of the dorsal fin (Fig.

3E) and by small eye size (Fig. 2).

Ictalurus natalis is easily distinguished from the others by its un-

pigmented mental barbels. The dark blood pigments in vessels of these

barbels should not be confused with the presence of melanophores.

Preserved blood in mental barbels appears as a dark line. The remaining

species, /. nebulosus and /. melas, have often been reported to be

separable by the character of the serrae on the posterior edge of the pec-

toral spine: moderate serrae in /. nebulosus, weak serrae in /. melas

(Trautman 1957; Blair et al. 1957; Hubbs and Lagler 1958; Pflieger 1975;

and others). The posterior pectoral spine serrae in /. melas are variable,

being absent to moderately developed. Although most often weakly

developed in adult /. melas, the pectoral serrae are unreliable for con-

sistently distinguishing /. melas from /. nebulosus. Ictalurus melas is best

distinguished from I. nebulosus by higher (rarely overlapping) gill raker

counts (Table 1). The single I. melas possessing 15 gill rakers on the right

arch had 17 on the left arch.

Fin pigmentation differences have also been reported. Of these

characters, only the depigmented “bar” at the caudal base of I. melas is

consistently present, and then only in larger juveniles and adults.

However, it is often evident only when directly compared to specimens of

I. nebulosus.

Forked-tail Ictalurus. — Of the three species in this group, I. catus is

readily separated by a moderately forked tail (Fig. 3H) and low anal ray

counts, usually 22-24, (22-25, x = 23.1). Variation exists in the anal ray

count ranges reported for I. catus: 19-22 (Jordan and Evermann 1896);

Table 1. Frequency distribution of gill raker counts (total) for subgenus Amiurus of Virginia.

Noel M. Burkhead, et al

88

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Virginia Ictalurid Catfishes

89

18-21 (Blair et al. 1957); 18-24 (Trautman 1957); 19-23 (Eddy 1969); 18-22

in key, 19-23 in text (Clay 1975). The larger ranges and lower extremes of

these counts may have resulted from failure to count all anterior

rudimentary rays in at least some of the material examined by these

authors. Smith-Vaniz (1968) reported a count range similar to ours (21-

26, X = 23). The gap in the “bony ridge” between the head and dorsal fin

(a disjuncture between the supraoccipital and the anterior process of the

first pterygiophore), reported in keys by Hubbs and Lagler (1958) and

Clay (1975), was consistently present in /. catus. However, a disjuncture

also occurs in juveniles of /. punctatus and /. furcatus.

Until recently, anal ray counts were reported to have little overlap

between 1. furcatus and /. punctatus (Trautman 1957; Blair et al. 1957;

Pflieger 1975; and others). However, Clay (1975) reported anal ray count

ranges to be 27-34 for /. furcatus and 23-29 for 7. punctatus and

emphasized the need to consider anal fin shapes (Fig. 3F, G). An obvious

character when present are the spots on young to adult /. punctatus, but

adults often lack them. W. Ralph Taylor (pers. comm.) informed us of a

difference in the gas bladders of L furcatus and I. punctatus; that of I. fur-

catus has an elongate posterior extension and that of /. punctatus does

not. The gas bladder difference is illustrated by Pflieger (1975).

KEY TO VIRGINIA ICTALURIDS WITH AN ADNEXED ADIPOSE FIN

1. Premaxillary tooth patch with posterolateral

extensions (Fig. 3A); upper lobe caudal fin

partially depigmented (varies in adults) Pylodictis olivaris.

Premaxillary tooth patch without posterolateral

extensions; upper lobe caudal fin not partly

depigmented 2.

2. Caudal fin deeply forked (Fig. 3F, G) 3.

Caudal fin moderately forked to emarginate

(Fig.3H,I) 4.

3. Anal fin margin usually rounded (Fig. 3G); anal rays

23-29; young to small adults often with spots Ictalurus punctatus.

Anal fin margin straight (Fig. 3F); anal rays 27-34;

never spotted I. furcatus.

4. Caudal fin moderately forked (Fig. 3H); anal rays

usually 22-24 (22-25) /. catus.

Caudal fin emarginate (Fig. 31) 5.

5. Dorsal fin with dark basal blotch (Fig. 3D); eye size

moderate (flathead bullheads) 6.

Dorsal fin without dark basal blotch (Fig. 3E); eye

size small (typical bullheads) 7.

6. Mental barbels usually without pigment (pigment may

be present in large specimens on lateral barbels, rarely

on medial); leading edge of maxillary barbels pale

90

Noel M. Burkhead, et al.

(appearing bicolored); premaxillary tooth patch of

large juveniles and adults as in Figure 3C; gill rakers

usually 10-14 (10-17); anal rays usually 22-24 (21-26) I. platycephalus.

Mental barbels usually profusely pigmented (occasionally

pigment only developed basally in small specimens);

maxillary barbels uniformly dark; premaxillary tooth

patch in large juveniles and adults as in Figure 3B; gill

rakers usually 12-16 (11-18); anal rays usually

18-20 (18-22) I. brunneus.

7. Mental barbels usually pale; anal rays usually 24-27

(24-28); gill rakers usually 12-15 (12-18) I. natalis

Mental barbels usually profusely pigmented 8.

8. Gill rakers usually 17-20 (15-24); a rectangular

depigmented area often present at base of caudal

fin in adults I. melas.

Gill rakers usually 13-15 (13-16); caudel base with

uniform pigmentation I. nebulosus.

ACKNOWLEDGMENTS. — The following people kindly provided

specimens for this study; Edward B. Brothers, Cornell University;

Donald G. Cloutman, formerly Duke Power Company; David A. Etnier,

University of Tennessee, Knoxville; E. David Frankensteen, U. S. Army

Corps of Engineers; John Gourley, Virginia Institute of Marine Science;

Larry Hart, Virginia Commission of Game and Inland Fisheries; Elbe

Koons, University of Michigan Museum of Zoology; William J.

Matthews, formerly Roanoke College; O. Eugene Maughan and Morris

Mauney, formerly Virginia Cooperative Fisheries Unit; Edward F.

Menhinick, University of North Carolina at Charlotte; James P. Oland,

U. S. Fish and Wildlife Service; W. Ralph Taylor and George Van Dyke,

U. S, National Museum of Natural History; Bruce Turner, Virginia

Polytechnic Institute and State University; and Shirley K. Whitt,

Lynchburg College,

Locality data, comments on critical records, or stocking information

were provided by the following: Michael Corcoran, Duke University;

Donald G. Cloutman, DPC; Edward C. Enamait, Maryland Fish Ad-

ministration; John Gourley, VIMS; Larry Hart, VCGIF; Frank J.

Schwartz, University of North Carolina Institute of Marine Sciences;

William B. Smith, North Carolina Wildlife Resources Commission; Jay

R. Stauffer, University of Maryland; W. Ralph Taylor, USNM; David

K. Whitehurst, VCGIF; and Ralph W. Yerger, Florida State University.

W. Ralph Taylor also made us aware of the gill raker difference between

I. melas and /. nebulosus. We are grateful to Michael Corcoran, Duke

University, for a review of a draft of this paper, Susan Karnella (USNM)

provided us with the catalog number of an Ictalurus punctatus.

LITERATURE CITED

Abbott, Tom M., K. L. Dickson and W. A. Potter. 1977. Notropis cerasinus

Virginia Ictalurid Catfishes 91

(Cope) record from the Appomattox River drainage. Va. J. Sci. 28(4):

167-168.

Addair, John. 1974. The fishes of the Kanawha River system in West Virginia

and some factors which influence their distribution. Ph.D. dissert.,

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Banarescu, P. 1968. Pozita sistematica a somnuli pitic American

adimatizat in apele Romaniei [The systematic position of the North

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Bean, Barton A., and A. C. Weed. 1911. Recent additions to the fish fauna

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Vertebrates of the United States. McGraw-Hill Co., New York. 819 pp.

Bryant, Richard T., B. H. Bauer, M. G. Ryon and W. C. Starnes. 1979. Distri-

butional notes on fishes from northern Georgia with comments on the

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Carnes, William C. 1965. Survey and classification of the Roanoke River

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Clay, William M. 1975. The Fishes of Kentucky. Ky. Dep. Fish Wildl. Resour.,

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Eddy, Samuel. 1969. How to Know the Freshwater Fishes. 2nd Ed. Wm. C.

Brown Co., Dubuque. 268 pp.

Elser, Harold J. 1950. The common fishes of Maryland. How to tell them

apart. Chesapeake Biol. Lab. No. 88. 45 pp.

Fowler, Henry W. 1945. A study of the fishes of the southern Piedmont and

Coastal Plain. Acad. Nat. Sci. Phila. Monogr. 7. 408 pp.

Harrel, R. Duane, R. L. Fuller and T. J. Edwards. 1978. An investigation of

the fish community of Belews Lake, North Carolina. Duke Power Co.

Rpt. 78-07. 64 pp.

Hart, Larry G. 1978. Project completion report for Virginia Dingell-Johnson

reservoir research study. Va. Com. Game Inland Fish., Richmond.

120 pp.

Hocutt, Charles H., P. S. Hambrick and M. T. Masnik. 1973. Rotenone

methods in a large river system. Arch. Hydrobiol. 72(2):245-252.

, R. F. Denoncourt and J. R. Stauffer. 1978. Fishes of the Greenbrier

River, West Virginia, with a drainage history of the central Appala-

chians. J. Biogeogr. 5:59-80.

, , and 1979. Fishes of the Gauley River, West

Virginia. Brimleyana 7:47-80.

Hubbs, Carl L., and Karl F. Lagler. 1958. Fishes of the Great Lakes Region,

2nd Ed. Cranbrook Inst. Sci. Bull. 26. 213 pp.

Jenkins, Robert E., E. A. Lachner and F. J. Schwartz. 1972. Fishes of the

central Appalachian drainages: their distribution and dispersal, pp.

43-117 in P. C. Holt (ed.). The distributional history of the biota of the

southern Appalachians, Part III: Vertebrates. Res. Div. Monogr. 4, Va.

Polytech. Inst. State Univ., Blacksburg. 306 pp.

92

Noel M. Burkhead, et al.

Jordan, David S. 1889. Report of investigations made during the Summer

and Autumn of 1888, in the Allegheny region of Virginia, North Carolina,

and Tennessee, and in western Indiana, with an account of the fishes

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, and B. A. Evermann. 1896. Fishes of North and Middle America,

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McAtee, W. L., and A. C. Weed. 1915. First list of the fishes of the vicinity

of Plummers Island, Maryland. Proc. Biol. Soc. Wash. 25:1-14.

Olmstead, Larry L., and D. G. Cloutman. 1979. Life history of the flat bull-

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Montgomery County, Maryland, region. Proc. Biol. Soc. Wash. 24:

39-46.

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the New (upper Kanawha) River system in Virginia. Va. Polytech. Inst.

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Amer. Mus. Nat. Hist. 148:40-154.

Smith, Hugh M., and B. A. Bean. 1898. Fishes known to inhabit the waters of

the District of Columbia and vicinity. Bull. U. S. Fish Com. 18:179-187.

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tributaries. North Carolina. N. C. Wildl. Resour. Com., Raleigh. 43 pp.

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tudinal distribution of the fishes of the East River, West Virginia-

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, K. L. Dickson, J. Cairns and D. S. Cherry. 1976. The potential and

realized influences of temperature on the distribution of fishes in the

New River, Glen Lyn, Virginia. Wildl. Monogr. 50. 40 pp.

, C. H. Hocutt and D. S. Lee. 1978. The zoogeography of freshwater

Virginia Ictalurid Catfishes

93

fishes of the Potomac River basin, pp. 44-54 in K. C. Flynn and W. T.

Mason (eds.). The Freshwater Potomac: Aquatic communities and

environmental stresses. Interstate Com. Potomac River Basin.

Trautman, Milton B. 1957. The Fishes of Ohio. Ohio State Univ. Press,

Columbus. 683 pp.

Wiley, Martin L. 1970. Fishes of the lower Potomac River. Atl. Nat. 25(4):

151-159.

Wollitz, Robert E. 1968. Smallmouth bass stream investigations. Job. No. 1

New River study. Va. Com. Game Inland Fish., Richmond. 59 pp.

Yerger, Ralph W., and K. Relyea. 1968. The flatheaded bullheads (Pisces:

Ictaluridae) of the southeastern United States, and a new species of

Ictalurus from the Gulf Coast. Copeia 1968(2):361-384.

Accepted 13 October 1980

9

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Geographic Variation in the Snake Storeria occipitomaculata

(Storer) (Serpentes: Colubridae) in Southeastern

United States

Douglas A. Rossman and Robert L. Erwin'

Museum of Zoology, Louisiana State University,

Baton Rouge, Louisiana 70893

ABSTRACT. -The populations of Storeria occipitomaculata occurring

in the Gulf Coastal Plain from eastern Texas to the Carolinas differ

from the nominate subspecies in nuchal pattern, ventral coloration,

relative tail length, and subcaudal number. To accommodate these pop-

ulations nomenclaturally the concept of S.o. obscura Trapido is expand-

ed and redefined.

The Florida red-bellied snake, Storeria occipitomaculata obscura,

was described by Trapido (1944), who distinguished it from the nominate

race on the basis of the former having a black head, the light supralabial

spot touching the edge of the lip, a light nuchal collar, fewer ventrals, and

more subcaudals. He envisioned the range of S.o. obscura as encompass-

ing peninsular Florida and coastal plain Georgia, with intergradation oc-

curring to the north and west of this area. Subsequent authors (Wright

and Wright 1957; Cliburn 1959; Mount 1975) reported the occurrence of

some of the distinguishing features of S.o. obscura in Mississippi and

Alabama populations, and we discovered them in Louisiana snakes.

These observations, along with the more than fivefold increase in

specimens available from the Southeast since the time of Trapido’s study,

prompted our investigation of variation in the species throughout this

region of the country.

METHODS

We examined 523 specimens of Storeria occipitomaculata from

southeastern United States (Fig. 1), and for each specimen we recorded

sex, ventral and subcaudal numbers, tail length as percent of total length,

nuchal pattern, and supralabial light spot condition. To facilitate

analysis, data for specimens from geographically proximate and

physiographically similar localities were pooled. We also examined living

snakes from Wisconsin, Pennsylvania, Georgia, Florida, Alabama, Mis-

sissippi, Louisiana, Arkansas, and Oklahoma, and noted various aspects

of their color pattern, particularly the ventral coloration.

'Present address: Department of Molecular Virology, Abbott Laboratories, North Chicago

IL 60064.

Brimleyana No. 4: 95-102. December 1980.

95

96

Douglas A. Rossman and Robert L. Erwin

Fig. 1. Distribution of Storeria occipitomaculata in southeastern United States.

Solid circles represent localities from which specimens were examined in this

study. Dotted lines delimit sample boundaries.

Storeria Variation

97

ANALYSIS OF CHARACTERS

Head Pattern. —Specimens of Storeria occipitomaculata frequently

darken if left in formalin for even a few days, so we were not able to

determine natural head coloration in many of the individuals we ex-

amined. Nevertheless, it is apparent that snakes having a very dark head

that contrasts with a lighter dorsum are not confined to the originally

described range of S.o. obscura, but occur in coastal plain populations

from at least South Carolina through Louisiana. Many of these same

populations do, however, also contain animals whose head does not con-

trast markedly with the dorsum, hence we question the taxonomic

usefulness of this character.

The presence or absence of a dark bar separating the light

supralabial spot from the edge of the lip is virtually impossible to deter-

mine in formalin-darkened specimens, and we lack these data for the ma-

jority of specimens examined.

Nuchal Pattern. —Although Storeria occipitomaculata obscura was

characterized as having a complete light nuchal collar as opposed to the

three light nuchal spots of S.o. occipitomaculata, 25% of the specimens we

examined (9 of 36) from the described range of S.o. obscura have separate

spots rather than a complete collar. Moreover, a complete collar occurs

fairly frequently in most coastal plain populations (for instance in 10 of

13 animals from the Florida Parishes of Louisiana and 15 of 27 from

west-central Louisiana.)

A more consistent character for distinguishing the coastal plain pop-

ulations from their more northern counterparts is whether or not the

light nuchal marks (spots or collar) are in contact with the light colora-

tion of the venter (see Fig. 2). Such contact on both sides of the neck oc-

curs in nearly all specimens examined from Florida, western Georgia,

Alabama, Mississippi, Louisiana, eastern Texas, and southern Arkansas.

In northern Arkansas, Tennessee, and extreme northern Alabama the

frequency of animals having the light nuchal marks separated from the

venter on at least one side of the neck ranges from 84 to 100%. Farther

east, in northern Georgia and North Carolina, the frequency ranges from

60 to 75%. In the geographically intermediate areas (southeastern

Oklahoma, western Tennessee, eastern Georgia, and South Carolina) the

light nuchal marks are separated from the venter in 27 to 50% of the

animals.

Ventral Coloration. —For reasons previously cited we have relatively

sparse information about ventral color in living S. occipitomaculata.

Nonetheless, from our personal observation and from color notes and

color transparencies provided by other workers, we can say that the

coastal plain snakes rarely have the bloodred or crimson venter typical of

the nominate race. Instead, ventral color may range from lemon yellow

through pale orange to butterscotch tan.

98

Douglas A. Rossman and Robert L. Erwin

Fig. 2. Lateral view of left side of head and neck in Storeria occipitomaculata oc-

cipitomaculata (above) and S.o. obscura (below) showing the differences in nuchal

pattern.

Relative Tail Length. —As can be seen from Fig. 3, the tail is propor-

tionally longer in the coastal plain populations than in those farther in-

land. The longest tail occurs in animals from the Florida Panhandle,

southern Alabama, and southern Mississippi.

Ventrals. —The number of ventrals exhibits no consistent differences

between coastal plain and inland populations (see Fig. 4). Ventral num-

ber is markedly higher in Louisiana, southern Arkansas, and eastern

Texas, a trend that may reflect the closer geographical proximity of those

populations to the Mexican S.o. hidalgoensis Taylor, which has even

more ventrals and exhibits an increasing north-south dine therein

(Trapido 1944; pers. obs.).

Suhcaudals. -As might be anticipated, geographic variation in sub-

caudal number (see Fig. 5) parallels that exhibited by relative tail length.

In this instance, however, the coastal plain populations from Georgia,

South Carolina and North Carolina agree more closely with inland pop-

ulations than they do with the coastal plain snakes from farther south

and west.

Storeria Variation

99

100

Douglas A. Rossman and Robert L. Erwin

Storeria Variation

101

CONCLUSIONS

The general concordance of geographic variation in nuchal pattern,

ventral coloration, relative tail length, and subcaudal number suggests

Fig. 5. Variation in subcaudal number of Storeria occipitomaculata in

southeastern United States. Upper figures in each pair represent sample mean

and sample size for males, lower figures represent similar data for females.

102

Douglas A. Rossman and Robert L. Erwin

that there is sufficient justification for giving taxonomic recognition to

the Gulf Coastal Plain populations of Storeria occipitomaculata. This ex-

panded concept of S.o. obscura requires, however, that the taxon be

redefined. As reconstituted, S.o. obscura can be distinguished from the

nominate race by having: the light nuchal marks usually contacting the

venter (versus usually separated from the venter); the venter yellow,

orange, or tan (versus venter some shade of red); the sample means for

relative tail length exceeding 25% in males, 22% in females (versus sample

means less than 25% in males, 22% in females); the sample means for sub-

caudal number exceeding 53 in males, 45 in females (versus sample means

less than 49 in males, 42 in females). Thus defined, S.o. obscura ranges

from eastern Texas through southern Arkansas and Louisiana to

Florida; it appears to intergrade with S.o. occipitomaculata in

southeastern Oklahoma, western Tennessee, northern Alabama, Georgia,

and the Carolinas (exclusive of the mountains, where the nominate race

occurs). Data on ventral coloration in animals from the Carolinas may

help to more clearly delimit the zone of intergradation.

ACKNOWLEDGMENTS. or the loan of material in their care

we are grateful to the curators of the following collections: American

Museum of Natural History; Auburn University Museum; Carnegie

Museum of Natural History; Charleston Museum; Chicago Academy of

Sciences; Field Museum of Natural History; University of Florida/

Florida State Museum; Florida State University; University of Kansas

Museum of Natural History; E.A. Liner, private collection; Louisiana

State University at Shreveport; Louisiana Tech University; McNeese

State University; University of Michigan Museum of Zoology; Mis-

sissippi Museum of Natural Science; National Museum of Natural

History; North Carolina State Museum of Natural History; Northeast

Louisiana University; Northwestern State University of Louisiana; Uni-

versity of Oklahoma; Stephen F. Austin State University; University of

Southern Mississippi; Tall Timbers Research Station; Texas Cooperative

Wildlife Collection; and Tulane University. For providing living

specimens, color transparencies, or color notes we thank W. Auffenberg,

R.M. Blaney, E.A. Liner, C.J. McCoy, D.B. Means, C.W. Myers, and

K.L. Williams.

LITERATURE CITED

Cliburn, J. William. 1959. The distribution of some snakes in Mississippi.

Am. Midi. Nat. 62:218-221 .

Mount, Robert H. 1975. The Reptiles and Amphibians of Alabama. Agric. Exp.

Sta., Auburn Univ., Auburn, Alabama. 347 pp.

Trapido, Harold. 1944. The snakes of the genus Storeria. Am. Midi. Nat. 3/;l-84.

Wright, Albert H., and Anna A. Wright. 1957. Handbook of Snakes of the

United States and Canada. Vol. 2. Comstock Publ. Assoc., Ithaca, N.Y.

pp. 565-1105.

Accepted 16 May 1980

Effects of Microhabitat Size and

Competitor Size on Two Cave Isopods

David C. Culver

AND

Timothy J. Ehlinger

Department of Biological Sciences,

Northwestern University, Evanston, Illinois 60201

ABSTRACT. isopods are common in cave streams in northern

West Wxfmxdi—Caecidotea cannulus (Steeves) and Caecidotea holsingeri

(Steeves). Laboratory experiments using an artificial stream demon-

strated that (1) for any given size of isopod, washout rate depended

on the size of gravels in the stream bed, i.e., larger isopods had low

washout rates in larger gravels and vice versa, and (2) competition (as

reflected in washout rate) decreased as size differences among competi-

tors increased. Field evidence provided support for the first result. In

particular, there was a concordance between isopod size and gravel size

distributions. There was no evidence for character displacement.

Appalachian cave stream communities are relatively simple systems,

dominated by one to three isopod and amphipod species (Culver 1976).

The limited amount of utilizable habitat is of great importance in struc-

turing these communities. Utilizable habitat is limited to the underside of

stream gravels, which provide concentrations of detritus, the major food

source, and hiding places from the brunt of the current (Culver 1971,

Estes 1978). Dislodgment from a rock often follows the encounter of two

individuals and results in significant mortality (Culver 1973). Since

washout is density-dependent, competition results, and is important in

determining annual population size fluctuations (Culver 1971, Estes

1978), microhabitat separation (Culver 1973), and resistance of com-

munities to invasion by other species (Culver 1976).

In this study we examined the effect of size on competition in two

species of isopods found in northern West Virginia caves. We first tested

the hypothesis that washout rates of different size isopods and gravel size

are correlated. This should result in a “match” between isopod sizes and

gravel sizes in cave streams. Second, we tested the hypothesis that in-

terspecific competition is reduced by size divergence of the two species.

This should result in a “mismatch” between isopod size and gravel size.

Using an artificial stream in the laboratory, we showed that washout rate

depends on both gravel and isopod size, and that size differences among

isopods reduces competition. Then, using data on the size distribution of

isopods and stream gravels in various caves, we assessed the importance

of these two factors.

Brimleyana No. 4; 103-113. December 1980.

103

104

David C. Culver and Timothy J. Ehlinger

METHODS AND MATERIALS

The isopods Caecidotea holsingeri (Sleeves) and Caecidotea cannulus

comprise the great majority of individuals and biomass of the

macroscopic fauna in most cave streams of the Monongahela River

drainage in West Virginia. Collections were made in every known loca-

tion for the species in this drainage. This covers the entire range of C.

cannulus. Caecidotea holsingeri is also known from caves in the New,

Greenbrier, Elk, and James River drainages (Holsinger et al. 1976),

where it often occurs with a rich amphipod fauna (Culver 1970). For

comparative purposes, collections of C. holsingeri were taken from Lin-

wood Cave in the Elk River drainage. This is the closest known locality

to caves in the Monongahela River drainage. Caves visited are listed in

Table 1.

Table 1. List of caves where collections were taken. All collections were made by

the authors except for Mill Run, Nelson, and Cave Hollow caves where

the collections were made by Dr. J.R. Holsinger. All caves are in the

Monongahela River drainage except Linwood Cave, which is in the Elk

River drainage. Cave locations and descriptions are in Davies (1965) and

Medville and Medville (1972).

' Male second pleopods, the critical taxonomic character, were not removed, but the bi-

modal size distribution is strong presumptive evidence that both species were present.

In four caves (Bowden, Glady, Harman and Linwood) stream

gravels were collected from riffles where isopods were found. Gravels

were sorted by diameter into 0.3 cm intervals up to 2.5 cm. The larger

gravels were divided into those less than and greater than 5 cm in

diameter.

Cave Isopod Ecology

105

For laboratory stream studies the isopods were collected alive and

measured using an ocular micrometer on a dissecting microscope. Collec-

tions were sorted into two size classes: small (less than 4.5 mm long) and

large (greater than 8 mm long). All individuals, including intermediate

sizes, were eventually preserved, measured, and identified. All small in-

dividuals were C. holsingeri and all large individuals were C. cannulus.

Two experiments were done in an artificial stream in the laboratory

(see Culver 1 97 1 for design of the stream). To measure the effect of gravel

size on washout rate, two 10 cm by 10 cm areas of rocks were used in the

stream. One consisted of gravels between 0.2 cm and 1 cm in diameter;

the other consisted of gravels between 2.2 cm and 3.8 cm in diameter. The

washout rate of each size class of isopods was measured for each size

class of gravels. Five individuals were used in each run, and each run was

repeated at least four times. Animals that washed out were collected at

the end of the artificial stream bed. Individuals washing out in the first 30

minutes were placed back in the stream, and then the number washed out

after 12 hours was recorded.

The second experiment measured interspecific effects on washout

rate, as reflected by size differences of the two species. To measure in-

trasize washout rates, 10 isopods of a single size class were placed in a 15

cm X 15 cm section of gravels of various sizes, patterned after a cave

stream. Each of these runs lasted 24 hours. Intersize class competition

was measured by following the procedure outlined above, but using five

small isopods and five large isopods. Thus, the total number of isopods

present at the beginning of a run was always 10. Each run was repeated

five times.

All isopods collected were measured and identified. Identification

posed some problems. Species can be separated only on the basis of the

second male pleopod. Most males can be identified since the second

pleopod sclerotizes at an early age. We found the following two charac-

ters to be reliable; the comparative length of the endopodial groove, and

the angle between the endopodite tip and the cannula. Since males are

usually scarcer than females, and none of the populations are large (we

collected less than 50, usually less than 25, in any one cave), it was

necessary to try to include females. There is little if any size dimorphism,

so we called all females C. holsingeri that were smaller than the largest C.

holsingeri male, and all females C. cannulus that were larger than the

smallest C. cannulus male. Ambiguous cases were randomly assigned to

the two species. In practice, such random assignments were only

necessary for Glady Cave. We also present size histograms to obviate

identification problems.

RESULTS

Due to the relatively small number of isopods available, it was often

necessary to use the same individual for more than one run in the arti-

106

David C. Culver and Timothy J. Ehlinger

ficial stream. Since individuals that washed out sometimes died, this in-

troduces a potential bias if there are differences in washout propensity

among individuals of the same size class. This would result in higher

washout rates in earlier runs of the same experiment. There was no

evidence that this happened.

There is a very clear tendency for small C. holsingeri to have higher

washout rates in large gravels than in small gravels, and for large C. can-

nulus to have higher washout rates in small gravels than in large gravels

(Table 2). The washout frequency of C. holsingeri is significantly higher

(P < 0.01) in large gravels (x = 0.75) than in small gravels (x = 0.15).

Similarly, the washout frequency of C. cannulus is significantly higher (P

<0.05) in small gravels (x = 0.55) than in large gravels (x = 0.24). These

data suggest that there should be a “match” between gravel and isopod

size distributions in natural streams.

Table 2. Washout rates of small C. holsingeri ( <C.4.5 mm) and large C. cannulus

( ^ 8 mm) in small gravels (.2 cm diam. 1 cm) and large gravels

(2.2 cm <: diam ^3.8 cm). Statistical analysis used washout frequency

transformed to sin"' VT, where x is the washout frequency.

X

sim' V\ = y

t P

5.46 0.01

2.07 -=^0.05

There was also evidence that size differences among isopods reduced

the washout rate in an artificial stream with a variety of gravel sizes

(Table 3). The washout frequency of large C. cannulus is significantly

lower when small C. holsingeri are present (x = 0.08) than when the same

total density of isopods, all of them large, are present (x = 0.44). The

washout frequency of small isopods was slightly lower when large

isopods were present (0.36 compared to 0.42), but the difference was not

significant (Table 3).

The difference in washout frequency shown in Table 3 could be due

either to size differences per se or to species differences per se. Washout

experiments with intermediate-size isopods (Culver and Ehlinger, in

preparation) indicate that size per se is more important. In these experi-

ments, five small C. holsingeri (or five large C. cannulus) were put in the

Cave Isopod Ecology

107

artificial stream with five intermediate-size C. holsingeri or C. cannulus. If

species differences are important, then total number of isopods washing

out when all individuals are the same species should be different than

when two species are involved. In fact, total washout of conspecifics (x =

0.28, S.E. = 0.03, N = 6) was nearly identical to total washout of in-

terspecifics (x = 0.32, S.E. = 0.03, N = 4).

Table 3. Fraction of isopods washing out in artificial stream with various sizes

of gravel. Statistical analysis used washout frequency (x) transformed to

sin'' /x. Each run was started with 10 individuals (10 of one size, or 5

each of two sizes).

x sin'' /x = y

Very limited information is available concerning Caecidotea in three

of the caves listed in Table 1 . Nelson and Bazzle caves have large popula-

tions of the amphipod Gammarus minus. Only two C. holsingeri were

found in Bazzle Cave after extensive searching. A situation is similar in

Nelson Cave. No isopods could be found in this cave in February 1980.

Mill Run Cave has a large isopod population, but is closed by the

owner. Specimens in National Museum of Natural Flistory collections

were measured, and had a bimodal size distribution. Male second

pleopods were not removed for examination, but the bimodal size dis-

tribution is strong presumptive evidence that both species were present.

Length measurements for C. cannulus and C. holsingeri are sum-

marized in Table 4. In all cases of syntopy, C. cannulus is at least 1.9 times

as large as C. holsingeri, all the differences being statistically significant.

However, consideration of allotopic populations complicates the issue.

The allotopic population of C. cannulus in Cave Hollow Cave is slightly

smaller (x = 7.05 mm) than syntopic populations of the same species (x =

7.75 mm, S.E. = 0.25), but the difference is not statistically significant

(t =-0.83). The allotopic population of C. holsingeri in Harman Cave is

smaller (x = 2.8 mm) than syntopic populations (x = 3.1 mm, S.E. =

0.1), but the difference is not statistically significant (t = 1.42). On the

other hand, the allotopic population of C. holsingeri in Linwood Cave

(which is in a different drainage basin) is larger (x = 5.7 mm) than syn-

topic populations, and the difference is statistically significant (t = 1 1.34,

108

David C. Culver and Timothy J. Ehlinger

P <C 0.01). In summary, there is no clear evidence for divergence in size

where the species are syntopic.

Table 4. Length measurements of Caecidotea holsingeri and Caecidotea cannulus,

in mm.

C. holsingeri C. cannulus

Cave X S.E. N x S.E. N

suggest the two distributions should be correlated, although the actual

gravel sizes that an isopod of a given size uses are not known. However,

there are some striking similarities in the qualitative aspects of the gravel

and isopod size distributions for each cave, which are summarized in

Table 5. In Bowden Cave, both distributions are strongly bimodal. In

Glady Cave, the gravel size distribution is weakly bimodal, while isopod

size distribution is apparently unimodal with a broad size range. By con-

trast, in Harman Cave both gravel and isopod size distributions are

strongly unimodal and skewed to the left. In Linwood Cave, gravel sizes

are uniformly distributed for small gravels, and skewed to the right

overall. Isopod sizes are strongly unimodal, with a narrow size range.

There are two discrepancies between isopod and gravel size distributions.

First, in Glady Cave, gravel size distributions are weakly bimodal and

isopod sizes are not, although they do have the expected broad size range.

Second, in Linwood Cave there is a broad size range of gravels and a

narrow size range of isopods, and there is little concordance of the two

distributions. With these two exceptions, there was a good fit between the

two distributions.

DISCUSSION

There are two interrelated hypotheses about body size of Caecidotea

cannulus and C. holsingeri. The first is that small isopods suffer less

washout (and less mortality) in small gravels and large isopods suffer less

Cave Isopod Ecology

109

washout (and less mortality) in large gravels. Thus, minimization of mor-

tality (whether through natural selection or developmental adjustment of

body size) should result in a concordance between body and gravel size

distributions. Supporting this, previous work (Culver 1971, 1973) in-

dicated that washout is due to encounters between two individuals, and

thus is density-dependent. The second hypothesis is that interspecific

competition is size-dependent. If the first hypothesis is true, then the ex-

tension to interspecific competition should result in character displace-

ment, and at least a partial discordance between body and gravel size

distributions.

Both laboratory and field data strongly support the first hypothesis.

The artificial stream experiments (Table 2) clearly show that in the ab-

sence of other factors gravel size and isopod size should be correlated.

Furthermore, field data support this hypothesis (Fig. 1 , Table 5). In Bow-

den Cave, both distributions are bimodal. In Glady Cave, large gravels

are less frequent (Table 3), and C. cannulus is smaller than in Bowden

Cave. In Harman Cave, small gravels predominate, and C. holsingeri is

small. The low frequency of large gravels in Harman Cave may be the

reason for the absence of C. cannulus. Only in Linwood Cave is there a

discordance difficult to explain by the first hypothesis (Fig. 1, Table 5).

Larger isopods should be present. However, Linwood Cave is separated

from the range of C. cannulus by a major drainage divide. Thus, the most

reasonable explanation for the absence of large C. cannulus is historical.

The evidence for the second hypothesis is mostly negative. Artificial

stream experiments suggest that size differences reduce interspecific com-

petition (Table 3), at least for the larger species. On the other hand, field

evidence provides no support for the character displacement hypothesis.

The allotopic population of C. cannulus in Cave Hollow Cave is slightly

but not significantly smaller than syntopic populations. Unfortunately

for the testing of the hypothesis, C. cannulus in Cave Hollow Cave does

not occur in a gravel-bottom stream area. Only four individuals were

collected in three trips to the cave, and all of these were collected on

bedrock in the stream (Holsinger, pers. comm.). Gravel-bottom riffles

had large populations of the amphipod Gammarus minus. The size of C.

cannulus in water flowing over limestone bedrock may be subject to op-

timization, but we have no idea what that optimal size is. Thus, even if in-

dividuals in Cave Hollow Cave were significantly smaller, its significance

would be hard to interpret. Caecidotea holsingeri sizes also provide no

support for character displacement. One allotopic population (Linwood

Cave) is larger than all syntopic populations, and one (Harman Cave) is

smaller than most syntopic populations. In addition, gravel size distribu-

tions provide no support for character displacement. If character dis-

placement were occurring, then discordances between gravel and isopod

distributions should result in greater separation of isopod sizes. But, in

the case of Glady Cave, sizes of the two isopod species are unimodal

while gravel sizes are bimodal (Fig. 1, Table 5).

110

David C. Culver and Timothy J. Ehlinger

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Cave Isopod Ecology

111

It is worth pointing out that C. cannulus is, overall, larger than C.

holsingeri. The basic question is what determines body size, both in the

ultimate and proximate sense. It is unlikely that size differences are due

to differences in age structure, since age (or at least size) structure is

skewed toward older (or larger) individuals (Culver 1971). However, it is

ISOPODS

GRAVELS

10-

s-

HARMAN

Fig. 1. Gravel and isopod sizes for Harman, Linwood, Bowden, and Glady caves.

See Table 5 for description of the distribution.

impossible, with available data, to determine whether size differences are

genetically or environmentally determined. Dickson’s (1977) finding that

drip pool populations of the amphipod Crangonyx antennatus had con-

sistently larger size individuals than stream populations, but showed no

112

David C. Culver and Timothy J. Ehlinger

genetic differences (Dickson et al. 1979), suggests environmental deter-

mination of size. On the other hand, the extensive genetic differentiation

of the amphipod Gammams minus in narrow anticlinal valleys similar to

those in our study area (Gooch and Golladay, in press) suggests that dif-

ferences in body size on a small geographic scale may be genetically

determined.

Even if C. cannulus is genetically larger than C. holsingeri, it is not

clear that this is the result of past competition. The two species may have

descended from a common surface ancestor, and the larger size of C. can-

nulus may be the result of past competition. However, the two species

may have descended from separate ancestors that differed in body size.

There is no way to determine which of the two scenarios is the correct

one.

Finally, elimination of some of the data would result in the conclu-

sion that character displacement is important. In particular, if data for

Harman Cave and Glady Cave are eliminated, character displacement

would be indicated. Individuals in allotopic populations of C. holsingeri

are large (Linwood Cave) and individuals of C. cannulus in such popula-

tions are small (Cave Hollow Cave). In fact, we originally studied these

two species because they appeared to show character displacement (see

Holsinger et al. 1976). This supports the critique of character displace-

ment by Strong et al. (1979) and shows that sampling should not be

ended merely because the “correct” results have been obtained.

ACKNOWLEDGMENTS. John R. Holsinger and Mr.

William K. Jones assisted in the field. Dr. Thomas Bowman loaned

specimens from Mill Run Cave and provided measurements of the type

series of C. cannulus from Cave Hollow Cave. Mr. Julian Lewis helped

with identifications. Dr. John R. Holsinger provided information on his

collections in the area. TJE was supported by funds from the Orlando

Park Fund of Northwestern University.

LITERATURE CITED

Culver, David C. 1970. Analysis of simple cave communities. 1. Caves as islands.

Evolution 2^:463-474.

1971. Analysis of simple cave communities. III. Control of abun-

dance. Am. Midi. Nat. (55:173-187.

1973. Competition in spatially heterogeneous systems: an analysis

of simple cave communities. Ecology 54:102-110.

1976. The evolution of aquatic cave communities. Am. Nat.

y/t}:945-957.

and T.J. Ehlinger. In preparation. Determinants of body size of

two subterranean ho^od?,~Caecidotea cannulus and Caecidotea holsingeri

(Isopoda: Asellidae).

Davies, William M. 1965. Caverns of West Virginia, with supplement. W. Va.

Geol. Survey, Morgantown. 401 pp.

Cave Isopod Ecology

113

Dickson, Gary. 1977. Variation among populations of the troglobitic amphipod

crustacean Crangonyx antennatus Packard living in different habitats.

I. Morphology. Int. J. Speleol. 9:43-58.

, J.C. Patton, J.R. Holsinger and J.C. Advise. 1979. Genetic variation

in cave-dwelling organisms: the troglobitic amphipod crustacean Crangonyx

antennatus. Brimleyana 2:1 19-130.

Estes, James A. 1978. The comparative ecology of two populations of the trog-

lobitic isopod crustacean Lirceus usdagalun (Asellidae). M.S. Thesis,

Old Dominion University, Norfolk. 85 pp.

Gooch, James L., and S.W. Golladay. In press. Genetic population structure in

an amphipod species. Int. J. Speleol.

Holsinger, John R., R.A. Baroody and D.C. Culver. 1976. The invertebrate cave

fauna of West Virginia. W. Va. Speleol. Survey Bull. 7. 82 pp.

Medville, Douglas, and H. Medville. 1971. Caves of Randolph County, West

Virginia. W. Va. Speleol. Survey Bull. 1. 218 pp.

Strong, Donald S., L.A. Szyska and D.S. Simberloff. 1979. Tests of community

wide character displacement against Null hypotheses. Evolution 35:897-913.

Accepted 15 September 1980

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Life History of the Mottled Sculpin, Cottus bairdi, in

Northeastern Tennessee (Osteichthyes: Cottidae)

Jerry W. Nagel

Department of Biological Sciences, East Tennessee State University,

Johnson City, Tennessee 37614

ABSTRACT. ^Cottus bairdi in northeastern Tennessee spawns for the

first time at the end of its second year. Mature males are larger than

mature females. Spawning occurs in early April and is completed within

a one week period. Comparison with northern and western populations

indicates lower fecundity, larger eggs, and larger hatchlings for C. bairdi

in northeastern Tennessee. Population estimates revealed that immature

fish were concentrated in disturbed, exposed habitats, whereas mature

fish were evenly distributed between disturbed and undisturbed

habitats.

INTRODUCTION

Several studies reported on the life history and ecology of the mot-

tled sculpin, Cottus bairdi, in the northern and western parts of its range

(Hann 1927; Koster 1936; Bailey 1952; Zarbock 1952; McCleave 1964;

Ludwig and Norden 1969; Patten 1971). This study presents information

on the life history of C. bairdi in the southeastern part of its range.

MATERIALS AND METHODS

Specimens of C. bairdi (N = 795) were collected from a 2.5 km sec-

tion of Straight Creek, a tributary to the Nolichucky River, 7.5 km south

of Johnson City, Washington County, Tennessee. Stream width in this

section ranges from 0.5 to 2.0 m and averages 1.0 m. The stream has a

gravel and rubble substrate and is 95% riffle (depth = 5-15 cm) with 5%

pool habitat (depth =0.5- 1.0 m). Streamside canopy is either 100% in un-

disturbed areas or 0% in areas disturbed by road construction 15 years

prior to this study. Stream gradient in this section is 39 m km-' . Water

temperatures were recorded with a mercury thermometer at the time of

collection; highest recorded temperature was 19°C on 11 July 1977 and

on 5 August 1977.

Collections were made by electrofishing at monthly intervals from

March 1977 to April 1978. All specimens were preserved in 10% formalin

(buffered with CaC03) within one hour of capture.

A supplementary collection of C. bairdi (N = 102) was made from

North Indian Creek in Unicoi County, Tennessee on 25 February 1978 to

provide comparative information from a larger stream. North Indian

Creek at this collection site is 5 to 10 m wide, 0.2 to 0.5 m deep in riffle

areas, and 2 to 3 m deep in pools.

Total length (TL) of all specimens was measured to the nearest 1.0

mm and gonads from all specimens greater than 45 mm TL were weighed

Brimleyana No. 4: 115-121. December 1980.

115

116

Jerry W. Nagel

5

10

_ FEMALES MARCH 1977 _

FEMALES MARCH 1978

MALES

30 40 50 60 70 80

TOTAL LEHGTH, MM

Fig. 1. Frequency distributions of total lengths of all C. bairdi collected from

Straight Creek, March 1977 and 1978. Each bar represents a 2-mm interval. Solid

bars = immatures: open bars = breeding adults.

to the nearest 0.1 mg. Measurements of follicle diameter, egg diameter,

and TL of hatchlings were made with an ocular micrometer in a dis-

secting microscope.

Mark and recapture population estimates were made in April and

May 1 979 in a 195 m section of Straight Creek. This included a 100 m sec-

tion (#1 ) with 100% canopy cover and a 95 m section (^2) with 0% canopy

cover. All specimens less than 50 mm TL were marked with a left pelvic

clip while specimens over 50 mm TL were tagged with a fingerling tag

(Floy Tag and Mfg,, Inc., Seattle, Washington 98105) fastened with vinyl

thread under the anterior part of the dorsal fin. Some tag loss was noted,

but the duration of the study was short enough that tagging wounds were

Cottus Life History

117

MONTH

Fig. 2. Mean monthly gonad weights for all C. bairdi over 45 mm TL collected

from Straight Creek, March 1977 to April 1978.

easily detected and fish with lost tags could be identified for population

estimates of this size class.

RESULTS

Population —Length/frequency distributions of all

sculpins collected from Straight Creek in March 1977 and March 1978

are presented in Figure 1 and suggest that both males and females

reproduce for the first time at the end of their second year. Mature males

were distinctly larger than mature females in this population. Com-

parison of the rather small samples of immatures in the March 1977 and

118

Jerry W. Nagel

Fig. 3. Number of enlarged follicles related to total length in gravid females from

Straight Creek.

March 1978 length/frequency distributions suggests between-year varia-

tion in growth rates for the Straight Creek population during this study.

Reproduction. ^y[oni\\\y gonad weights for mature specimens are

presented in Figure 2 and indicate that spawning occurred in early April.

In 1978, weekly samples were taken during March and April to more

precisely indicate the time of spawning. On 4 April all seven mature

females were still gravid, but on 1 1 April all but 1 of 12 mature females

were spent. Afternoon water temperatures on these dates were 14°C and

12°C, respectively.

Cottus Life History

119

Fecundity estimates were based on gravid females collected just

prior to spawning in March 1977 and March/April 1978. Figure 3 pre-

sents the relationship between TL and number of mature follicles for

1977 and 1978. Correlation coefficients were 0.69 (N = 18, P 0.01) and

0.82 (N = 47, P <C 0.001), respectively. Regression lines were calculated

by Bartlett’s 3-group method for Model II regression (Sokal and Rohlf

1969); 95% confidence limits for the slopes were 0.85 to 3.43 (1977) and

2.08 to 3.28 (1978). Although the average total lengths for gravid females

in 1977 (x = 63.9 mm, N = 18, 95% confidence limits = ±2.14 mm) and

1978 (x = 63.2 mm, N = 47, 95% confidence limits = ± 2.00 mm) were

similar, the average number of mature follicles in 1978 (x = 67.7, N = 47,

95% confidence limits = ± 5.98) was significantly higher than in 1977 {X

= 56.8, N = 18, 95% confidence limits = ± 6.34). Gravid females col-

lected from North Indian Creek in February 1978 were similar in TL (x =

63.2 mm, N = 53, 95% confidence limits = ±1.69 mm) to the Straight

Creek samples and had an average follicle count of 55.5 (N = 53, 95%

confidence limits = ± 3.62).

Average follicle diameter in gravid females collected just prior to

spawning in April 1978 was estimated by measuring five follicles each

from nine different females. Average diameter was 3.32 mm (N = 45,

95% confidence limits = ± 0.08 mm, range = 2.8-3. 8 mm). A clutch of

eggs collected on 3 May 1978 had an average egg diameter of 3.73 mm (N

= 10, 95% confidence limits = ± 0.05 mm, range = 3. 6-3. 8 mm). This

clutch was incubated at 15°C and hatched on 8 May 1978. Sac fry at

hatching had an average TL of 9.80 mm (N = 5, 95% confidence limits =

± 0.22 mm, range =9.6-10.1 mm).

Population density and recapture population estimates were

calculated from N = (M±l) (C±1)/(R±1), where N = estimated pop-

ulation, M = number of marked fish, R = number of marked fish recap-

tured, and C = total sample of marked and unmarked fish in recapture

sample (Ricker 1975:78, equation 3.7). Calculation of 95% confidence

limits was based on the 95% confidence limits of R (Ricker 1975:78 and

Append. II). Population estimates and supporting data are presented in

Table 1 .

Table 1. Mark and recapture population estimates of C. bairdi in Straight Creek,

April 1979.

120

Jerry W. Nagel

At the time of the April/May 1979 population estimates specimens

less than 50 mm TL were mainly immature individuals ending their first

year, whereas specimens more than 50 mm TL were mature individuals

that were two years or older (Fig. 1), Mature specimens were equally

abundant in the two sections, whereas the immature size class

showed strikingly higher density in the exposed and disturbed habitat of

section §2.

DISCUSSION

In this study one year old fish were all immature and most two year

old fish were mature individuals in breeding condition (Fig. 1). This is

similar to maturity patterns found by investigations in Michigan (Hann

1927); New York (Koster 1936); Montana (Bailey 1952); Wisconsin

(Ludwig and Norden 1969); and Washington (Patten 1971). Koster

(1936), however, noted that the lake-dwelling form, C.b. kumlieni,

appeared to mature at the end of its first year at an average TL of approx-

imately 61 mm (conversion from standard length based on Bailey 1952).

The sexual dimorphism in size found for breeding individuals in this

study was noted in other studies (Flann 1927; Koster 1936, for C.b. bairdi

but not for C.b. kumlieni: Simon and Brown 1943; Bailey 1952; Zarbock

1952; Ludwig and Norden 1969).

In this study spawning occurred at water temperature of 12° - 14°C

during a one week period in early April. Previous investigations of

spawning by C. bairdi reported water temperatures ranging from

5° - 18°C and dates from late February to late May (reviewed in Ludwig

and Norden 1962). Hann (1927) and Koster (1936) reported brief spawn-

ing periods similar to the findings in this study, but Simon and Brown

(1943), Bailey (1952), and Ludwig and Norden (1962) reported spawning

periods of a month or more.

Several authors presented information on the average number of

enlarged follicles in gravid females for western and northern populations

of C. bairdi (257, Hann 1927; 120 for C.b. bairdi and 135 for C.b.

kumlieni, Koster 1936; 629, Simon and Brown 1943; 203, Bailey 1952;

328, Ludwig and Norden 1969; 95, Patten 1971). Although no particular

geographic pattern in this variation is evident in these populations, these

estimates are consistently higher than fecundity estimates reported in this

study (three separate estimates = 56, 57, and 68) even though the size

ranges of gravid females were quite similar in all studies. Correlated with

this, average follicle diameter in northeastern Tennessee (3.32 mm) was

greater than diameters reported by other authors (2.0-2. 5 mm, Hann

1927; 2.2 mm, Simon and Brown 1943; 1.88 mm, Ludwig and Norden

1969). This correlation continues when comparing average egg size (in

nest) and average hatchling size. In this study eggs averaged 3.73 mm and

sac fry 9.8 mm. Corresponding estimates from other studies are; sac fry

- 6.4 mm (Hann 1927); eggs ~ 2.6 and 2.7 mm, sac fry ~ 6.9 and 7.9

mm (Koster 1936); 5-day fry 6.9 mm (Simon and Brown 1943); sac fry

Coitus Life History

121

— 8.1 mm (Bailey 1952); sac fry — 5.9 mm (Ludwig and Norden 1969).

This striking emphasis on “quality” (larger and fewer eggs) by C. bairdi

in northeastern Tennessee deserves further investigation in other

southern populations to determine whether or not it is a consistent

geographic trend.

LITERATURE CITED

Bailey, Jack E. 1952. Life history and ecology of the sculpin Coitus bairdi

punctulatus in southwest Montana. Copeia 1952(4);243-255.

Hann, Harry W. 1927. The history of the germ cells of Coitus bairdi Girard. J.

Morphol. Physiol. 45:427-497.

Koster, William J. 1936. The life-history and ecology of the sculpins (Cottidae)

of central New York. Ph.D. Dissert., Cornell Univ., Ithaca. 87 pp.

Ludwig, Gerald M., and C.R. Norden. 1969. Age, growth and reproduction of

northern mottled sculpin {Coitus bairdi bairdi) in Mt Vernon Creek,

Wisconsin. Milw. Public Mus. Occas. Pap. Nat. Hist. No. 2, 67 pp.

McCleave, James D. 1964. Movement and population of the mottled sculpin

{Coitus bairdi Girard) in a small Montana stream. Copeia 1964(3):506-513.

Patten, Benjamin G. 1971. Spawning and fecundity of seven species of northwest

American Coitus. Am. Midi. Nat. 55:493-506.

Ricker, William E. 1975. Computation and interpretation of biological statistics

of fish populations. Bull. Fish. Res. Board Can. 191, 382 pp.

Simon, James R., and R.C. Brown. 1943. Observations on the spawning of

the sculpin Coitus semiscaber. Copeia 1943(l):41-42.

Sokal, Robert R., and F.J. Rohlf. 1969. Biometry. W.H. Freeman and Co.,

San Francisco. 776 pp.

Zarbock, William M. 1952. Life history of the Utah sculpin. Coitus bairdi semi-

scaber (Cope) in Logan River, Utah. Trans. Am. Fish. Soc. 57:249-259.

Accepted 25 August 1980

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Notes on the Distribution and Ecology of the Black

Mountain Dusky Salamander Desmognathus welteri

Barbour (Amphibia: Plethodontidae) in Tennessee

William H. Redmond

Office of Natural Resources,

Tennessee Valley Authority,

Norris, Tennessee 37828

ABSTRACT.— Desmognathus welteri was found at 16 localities in the

Cumberland Mountains and the northern half of the Cumberland

Plateau. The observed intermittent seasonal nature of most southern

Cumberland Plateau small streams, combined with the strong aquatic

tendencies of the species, may be responsible for the apparent absence of

D. welteri in this region. Cursory observations indicate that D. welteri

and D. monticola may be competitors, while D. welteri and D. fuscus

probably partition the streamside habitats according to gradient and

substrate particle size. Alteration of streams by coal strip mine opera-

tions and extensive use of the species for fish bait have resulted in the

decline of many local populations. Considering these factors, D. welteri

should continue to be considered a species “in need of management” in

Tennessee.

INTRODUCTION

Desmognathus welteri was originally described from Big Black

Mountain, Harlan County, Kentucky, as a subspecies of D. fuscus

(Barbour 1950). Subsequently, Barbour (1971) believed that sufficient

evidence was available to treat it as a distinct species. Recent studies of

Caldwell (1977, 1980), Caldwell and Trauth (1979), and Juterbock (1975,

1978) provided substantial morphological evidence that supports this

proposal. Barbour (1971) noted the range of D. welteri as the eastern

third of Kentucky with disjunct populations in east central Alabama and

northern West Virginia. Caldwell (1977) stated that D. welteri probably

does not occur in Alabama and that most reports of the species from the

state were based on misidentified D. monticola. Juterbock (1975)

provided the first Tennessee record of D. welteri, from Cumberland Gap

National Historic Park, Claiborne County. Caldwell (1977) and Caldwell

and Trauth (19'^9) reported the species from Pickett, Fentress, Cum-

berland, and Scott counties. Redmond and Jones (1978) noted the Ten-

nessee distribution to include the northern half of the Cumberland

Plateau Physiographic Province. Caldwell and Trauth (1979) believed

that the distribution of D. welteri reached its southern limit in the Crab

Orchard Mountains near the northern end of Walden Ridge, Cum-

berland County, Tennessee.

In 1975, the Tennessee Wildlife Resources Agency included D.

welteri in a list of species designated as “wildlife in need of management.”

This designation includes species that are potential candidates for

threatened status, but whose status needs further evaluation in the state

123

Brimleyana No. 4: 123-131. December 1980.

124

William H. Redmond

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Desmognathus Distribution and Ecology

125

Table 1. Localities from which D. welteri were examined, with a list of associated

desmognathine species and numbers collected at each locality.

Locality

Campbell County

Bear Branch just off Hwy. 63, 17.4 km NW of

Caryville, elev. 500 m; 5 specimens, UTKVZC

01135-39, 20 August 1977.

Small trib. to New River, 2.1 km NNW of Shea

on gravel road, elev. 463-518 m; 5 specimens,

UTKVZC 01 144-48, 20 August 1977.

Small trib. to Davids Creek, 0.6 km W of

peak Cross Mountain on gravel road, elev. 805 m;

9 specimens, UTKVZC 01 156-64, 20 August 1977.

Small trib. to Clear Fork, 4.3 km E of jet.

Hwys. 25W and 1-75 on 25W, elev. 305-335 m; 9

specimens, UTKVZC 01337-45, 18 September 1977.

Small trib. to Clear Fork, 6.2 km E ofjet.

Hwys. 25W and 1-75 on 25W, elev. 366-396 m; 1

specimen, UTKVZC 01 199, 18 September 1977.

Small trib. to Hickory Creek, 1 .8 km W of jet.

Hwys. 25W and 90 on 25W, elev. 366-396 m; 6

specimens, UTKVZC 01219-24, 18 September 1977.

Small trib. to Big Creek, approx. 5.6 km NE

ofjet. Hwys. 25W and 63 on 25W, elev. 427-442 m;

6 specimens, UTKVZC 01232-37, 18 September 1977.

Cumberland County

Genesis Creek at Genesis Road, 11.0 km S of

Morgan Co. line; 1 specimen, AUM 24602.

Small trib. to Renfro Creek, 4.8 km SE ofjet.

Hwys. 70 and 1-40 on 70, elev. 488 m; 1 specimen,

UTKVZC 01245, 1 October 1977.

Small trib. to Renfro Creek, 5.5 km SE of jet.

Hwys. 70 and 1-40 on 70, elev. 482-488 m; 4 spec-

imens, UTKVZC 01267-70, 1 October 1977.

Small trib. to Jewett Branch, 4.6 km NW of

Cumberland-Bledsoe Co. line on Jewett Road, elev.

634-671 m; 12 specimens, UTKVZC 01298-309,

1 October 1977.

Associated species (N)

D. fuscus (7, and 1

egg clutch)

D. monticola (3)

D. fuscus (6)

D. fuscus ( 1 )

D. monticola (12)

D. fuscus (1)

D. fuscus ( 14)

D. monticola (2)

D. fuscus ( 1 )

D. monticola (3)

unknown*

D. ochrophaeus (5)

D. fuscus (7)

D. ochrophaeus (7)

D. fuscus (6)

D. ochrophaeus (6)

Fentress County

Deer Gap, Buffalo Cove, approx. 5.6 km S of unknown*

Jamestown on Hwy. 127; 25 specimens, AUM 24648-72,

2 September 1975.

Northupp Falls; 45 specimens, AUM 24603-47, unknown*

1 September 1975.

Small trib. to Campbell Hollow Branch, 3.0 none

km NW of Jamestown on Hwy. 52, elev. 470 m;

2 specimens, UTKVZC 03277-78, 30 April 1978;

126

William H. Redmond

Locality

5 specimens, UTKVZC 02032-36, 9 November 1977.

Small trib. to Stuart Creek, 2.4 km (airline)

W of Sharp Place, elev. 463-488 m; 2 specimens,

UTKVZC 02049-50, 9 November 1977.

M organ County

North Prong of Flat Fork, approx. 6.6 km

(airline) WNW of Fork Mountain; 2 specimens,

UTKVZC 02704-05, 1976.

Catoosa Wildlife Management Area, Pennykin

Branch, elev. 494 m; 4 specimens, UTKVZC

01169-72, 28 August 1977.

Pickett County

Small trib. to Rock Creek, 7.7 km (airline)

NE of Sharp Place, elev. 408 m; 4 specimens,

UTKVZC 02057-60, 9 November 1977.

Scott County

Small trib. to Bill’s Branch nr. USGS

weather station, elev. 425-460 m; 8 specimens,

UTKVZC 02267-74, 2 October 1976.

* Localities not collected by the author.

Associated species (N)

D.fuscus (4)

none

D. fuscus (24)*

D. monticola (1)

(Tennessee Wildlife Resources Agency 1978). The purpose of this study

was to further delineate the range of D. welteri and to provide comments

concerning its ecology and factors threatening the species in Tennessee.

METHODS

Field investigations were conducted from July 1976 to May 1979.

Forty-one collection sites were visited and two people spent approx-

imately, one hour at each site (Fig. 1). Because most published reports in-

dicate that the range of D. welteri is predominantly within the Cum-

berland Plateau and Mountain regions, most field efforts were concen-

trated in these regions of Tennessee. However, field searches were con-

ducted in the adjacent Highland Rim and Ridge and Valley.

Eighty-five specimens of D. welteri from sixteen localities were

collected and deposited in The University of Tennessee Vertebrate

Zoology Collection, Knoxville (UTKVZC). At each locality, general

habitat characteristics were noted, and associated desmognathine species

were collected and deposited in UTKVZC (Table 1). General habitat

characteristics and desmognathine species present were also noted for

collection sites where D. welteri was not found (Table 2). Seventy-one

specimens of D. welteri from three localities were examined from the

Auburn University Museum (AUM) (Table 1). Morphological charac-

teristics used to distinguish D. welteri from other sympatric

desmognathine species were taken from Caldwell (1977, 1980), Caldwell

and Trauth (1979), and Conant (1975).

Desmognathus Distribution and Ecology

127

Table 2. Localities where D. welteri was not found, with a list of desmognathine

species and numbers collected at each locality.

128

William H. Redmond

Desmognathine

Locality species (N)

10.6 km (airline) NW of Surgoinsville, elev.

469-488 m, 22 March 1978.

Moore County

Small southwestern trib. to Shipman Creek, 0.3 D.fuscus (5)

km NW of Ledford’s Mill, elev. 305 m, 4 October 1979.

Overton County

Small headwater stream of Big Laurel Creek

0.8 km S of jet Hwys. 85 and 164 on 164, elev.

530-549 m, 12 May 1979.

Pickett County

Small seepage below sandstone bluff at Natural

Bridge, 4.9 km (airline) NE of Sharp Place, elev.

494 m, 9 November 1977.

Putnam County

Unnamed Creek and spring run, 1.2 km S of jet.

Hwys. 70N and 84 on 84, elev. 518 m, 12 May 1979.

Eastern trib. to Mill Creek along gravel road,

1 .6 km S of Mill Creek Baptist Church, elev. 381 m,

12 May 1979.

Sequatchie County

Reynolds Creek, 9.2 km (airline) NNW of Dunlap,

20 July 1976.

Van Buren County

Small stream, 0.5 km SW of Spencer on Hwy. 30,

elev. 533 m, 19 November 1977.

Small stream, 3.3 km E of jet. Hwys. 30 and

1 1 1 on 30, elev. 488 m, 19 November 1977.

Small stream, 5.3 km E of jet. Hwys. 30 and

1 1 1 on 30, elev. 396 m, 2 September 1978. _

White County

Near jet. of Clifty creek and Millsea Branch,

0.9 km NE of Mobra, elev. 463 m, 19 November 1977.

Base of waterfall in upper headwaters of Wildcat

Branch, 2.6 km SW of Bon Air, elev. 549 m,

1 9 November 1977.

Virgin Falls and Cave, southern slope of Little

Chestnut Mountain, elev. 335 m, 2 September 1978.

D. fuscus ( 1 2)

D. fuscus (6)

D. fuscus (9)

D. fuscus (14)

D.fuscus (14)

none

D. fuscus (3)

D.fuscus (1 with

egg clutch)

none

D. fuscus (2)

RESULTS AND DISCUSSION

As determined in this study, the range of D. welteri in Tennessee is

Desmognathus Distribution and Ecology

129

shown in Figure 1. The species appears limited to the Cumberland

Mountains and northern half of the Cumberland Plateau. The

southernmost locality is on the south slope of Brady Mountain at the

northern end of the Sequatchie Valley, Cumberland County. This locality

is approximately 16 km south of Crab Orchard Mountain, which

Caldwell and Trauth (1979) speculated to be the southern distributional

limit of the species. In Tennessee, D. welteri was commonly found along

small- to medium-size streams flowing through mesophytic forests. In-

habited streams were typically permanent, and D. welteri usually oc-

curred in areas with steep to moderate gradients and with bedrock to

coarse gravel substrates. Specimens were taken from 305 to 805 m eleva-

tion.

The factors responsible for the apparent absence of D. welteri from

the southern half of the Cumberland Plateau are obscure. Seemingly

suitable habitats occur in several gorges and along stream courses on the

eastern and western escarpments of the region. Caplenor (1979) described

these gorge forests and contrasted them with the more xeric forests of the

Plateau’s tablelands. A general observation made during this study was

that most seemingly suitable streams on the southern Plateau were often

dry during extended droughts, especially in late summer and fall.

Desmognathus welteri has been characterized as a semiaquatic species

which seldom ventures far from water (Juterbock 1975; Caldwell 1977).

Juterbock (1975) noted that, in a given stream, D. welteri was found in

lower numbers in late summer when streamflow was decreased than dur-

ing periods of higher flow. The observed seasonal intermittent nature of

southern Plateau streams, combined with the strong aquatic tendencies

of the species, may be responsible for the apparent absence of D. welteri

in this region.

Listed in order of abundance, the desmognathine species found

closely associated with D. welteri were D. fuscus, D. monticola, and D.

ochrophaeus (Table 1). Where they were found together, D. welteri and D.

fuscus were seldom in the same habitats. Desmognathus fuscus was

typically taken along those sections of stream where the gradient was

gentle to moderate and where the substrate was silt, sand, or small gravel;

it was often found several meters from the stream. Desmognathus welteri

was most frequently found in areas with steep to moderate gradient

where the stream substrate was predominantly large rock, gravel, or

bedrock; it was seldom found more than a meter from the stream. In

Alabama, Folkerts (1968) described a similar phenomenon involving D.

fuscus and D. monticola, where D. monticola was the typical inhabitant of

the rocky, swift areas of a stream.

During this study, D. welteri and D. monticola were taken from

remarkably similar areas along the streams surveyed. Desmognathus

welteri was usually the more abundant in habitats where both species

were found. Based on these cursory observations, it appears that D. mon-

ticola and D. welteri may be competitors, with D. welteri being the domi-

130

William H. Redmond

nant form. However, detailed studies are needed to adequately describe

the ecological interactions between these two species. The observed par-

titioning of streamside habitats in Tennessee by D. welteri and D.fuscus

according to gradient and substrate particle size, and the possible com-

petitive relationship between D. welteri and D. monticola, are consistent

with the findings of Juterbock (1975), Caldwell (1977), and Caldwell and

Trauth (1979).

Alteration of streams by coal strip mine operations, and extensive

use of the species by bait fishermen, have resulted in the decline of many

local populations. Desmognathus welteri was never taken from stream

habitats where strip mining operations had removed bank vegetation, or

from streams with high silt, sand, and heavy metal concentrations.

However, because the species can inhabit relatively small streams, it was

often found in isolated, unaltered coves adjacent to orphan mine lands.

Within its range and in surrounding regions, D. welteri was one of the

most common “spring lizards” found in bait shops. Its large size makes it

a sought after fish bait. Up to 300 individuals were observed in one bait

shop holding box in Norris, Anderson County, Tennessee. This

collecting pressure is greatest during spring, summer, and early fall, and

probably results in the removal of many large, reproductively active

females.

Widespread use of D. welteri as a live bait may have resulted in

numerous introductions and alterations of the natural range of the

species in Tennessee. Martof (1953) discussed the distributional and

genetic ramifications of the commercial use of salamanders for fish bait.

Many D. welteri populations studied were found along small rivulets

which drained into nearby, often-fished streams. Although the data are

inconclusive, this distributional pattern may indicate past introductions

by fishermen.

Considering the lack of knowledge concerning factors limiting its

distribution, the rapid habitat degradation occurring in the Cumberland

Mountains and Plateau, and the widespread use of the species as fish

bait, I believe that D. welteri should continue to be considered as “in need

of management” in Tennessee.

ACKNOWLEDGMENTS.~¥ox his constant assistance in the field

and many helpful suggestions, I am grateful to Robert L. Jones, Ecology

Program, University of Tennessee, Knoxville. R.S. Caldwell, Kentucky

Nature Preserves Commission, Frankfort, Kentucky, verified the iden-

tification of many specimens taken during the study. Thanks are also due

A.C. Echternacht, Ecology Program, University of Tennessee, Knoxville,

and two anonymous reviewers, for providing many helpful comments.

LITERATURE CITED

Barbour, Roger W. 1950. A new subspecies of the salamander Desmognathus

fuscus. Copeia 1950(4):277-278.

Desmognathus Distribution and Ecology

131

1971. Amphibians and reptiles of Kentucky. Univ. Kentucky

Press, Lexington. 334 pp.

Caldwell, Ronald S. 1977. Cranial osteology of the salamander genus Desmogna-

thus Baird (Amphibia: Plethodontidae). Ph.D. Thesis, Auburn Univ.,

Auburn. 161 pp.

1980. Lens morphology as an identification tool in the salamander

subfamily Desmognathinae. J. Tenn. Acad. Sci. 55(1):15-17.

, and S.E. Trauth. 1979. Use of toe pad and tooth morphology

in differentiating three species of Desmognathus (Amphibia, Urodela, Pletho-

dontidae). J. Herpetol. yi(4):491-497.

Caplenor, Donald. 1979. Woody plants of the gorges of the Cumberland

Plateau and adjacent Highland Rim. J. Tenn. Acad. Sci. 54(4): 139-145.

Conant, Roger. 1975. A field guide to reptiles and amphibians of eastern and

central North America. Houghton Mifflin, Boston. 429 pp.

Folkerts, George W. 1968. The genus Desmognathus Baird (Amphibia: Pletho-

dontidae) in Alabama. Ph.D. Thesis, Auburn Univ., Auburn. 129 pp.

Juterbock, J. Eric. 1975. The status of Desmognathus welteri Barbour (Caudata:

Plethodontidae) and a comparison with two sympatric cogeners. M.S. Thesis,

Ohio State Univ., Columbus. 169 pp.

1978. Sexual dimorphism and maturity characteristics of three

species of Desmognathus (Amphibia, Urodela, Plethodontidae). J. Herpetol.

/2(2):217-230.

Martof, Bernard S. 1953. The “spring-lizard” industry: a factor in salamander

distribution and genetics. Ecology i4(2):436-437.

Miller, Robert A. 1974. The geologic history of Tennessee. Tenn. Div. Geol.

Bull. 74. Nashville. 63 pp.

Redmond, William H., and R.L. Jones. 1978. Preliminary comments on the

distribution and ecology of Desmognathus welteri (Black Mountain Dusky

Salamander) in Tennessee. J. Tenn. Acad. Sci. 5i(2):76 (Abstract).

Tennesseee Wildlife Resources Agency. 1975. Endangered and threatened species

of Tennessee. Proclamation Numbers 75-15 and 75-16. Nashville. 6 pp.

1978. Endangered species operational plan. Action year 1978-1979

through 1982-83. Nashville. 39 pp.

Accepted 4 September 1980

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Morphological and Habitat Variability in

Gammarus minus Say (Amphipoda: Gammaridae)

James L. Gooch

Department of Biology, Juniata College,

Huntingdon, Pennsylvania 16652

AND

Jeffrey S. Wiseman

Department of Biology, Muhlenberg College,

Allentown, Pennsylvania 18104

ABSTRACT. —The surface dwelling ecotype of Gammarus minus Say in

the central Appalachian Mountains varies in morphology partly in con-

formance to a habitat scale from fully open to borderline hypogean.

Characters investigated were eye facet density and relative lengths of eye

and three appendages to body length. There also is considerable inter-

demic variability not related to habitat scale. This is inferred to be due

to isolation of some demes, which have undergone differentiation, and

to gene flow among others, which has prevented full ecotypic dif-

ferentiation.

INTRODUCTION

Geographical variation in morphology within species of gammarid

amphipods is known to occur in Gammarus minus (Cole 1970, Minckley

and Cole 1963, Holsinger and Culver 1970); G. bousfieldi (Minckley and

Cole 1963); G. oceanicus (Croker and Gable 1977); G. pulex (Pinkster

1971, 1972); and Crangonyx antennatus (Dickson 1977). Intraspecific

variation in freshwater species has been linked to diet, light, current

velocity, substratum, competing and predatory species, and genetic drift.

This study examines interdemic morphological variation in epigean

populations of Gammarus minus Say. This species ranges from the mid-

Atlantic piedmont westward through the middle Appalachian Moun-

tains, Interior Low Plateaus, and portions of the Mississippi Valley and

Ozarks uplift, reaching peak abundances in caves, springs, and springfed

streams underlain by carbonate rocks (Holsinger 1976). Nine popula-

tions were examined, seven from Huntingdon and Centre counties in cen-

tral Pennsylvania, and one outlier population each in Virginia and West

Virginia. Our investigation was directed to the question: is morphological

variation systematically related to habitat variation? Habitats vary in

substratum, velocity, macrofauna, and many other attributes. This study

focuses on variation in habitats grading from open surface, i.e. fully

epigean, to cave associated or borderline hypogean. The study by

Holsinger and Culver (1970) indicated that the morphology of Gammarus

minus is particularly sensitive to this habitat spectum. All future

references to Holsinger and Culver will be to this 1970 paper. These

authors reported differences in eye shape and size and ratio of appendage

to body length in cave and surface dwelling populations in the mid-

Appalachians. Shoemaker (1940) earlier recognized a distinct cave form

Brimleyana No. 4; 133-147. December 1980. 133

134

James L. Gooch and Jeffrey S. Wiseman

with reduced eyes and elongate antennae, and Hubricht (1943) further

described an intermediate type between the surface and deep cave forms.

Holsinger and Culver distinguished three intergradational forms or

ecotypes, each associated with a specific habitat. Form I amphipods have

considerably reduced eyes, bluish body color, and elongate appendages,

and are confined to a few large cave systems in isolated karst areas of

Virginia and West Virginia. Form II individuals have slightly reduced

eyes, bluish bodies, and slightly lengthened appendages, and occur widely

in mid-Appalachian caves. Form III amphipods are brownish and ro-

bust, with large eyes and relatively shorter appendages, and are found in

surface habitats throughout the species’ range. The three forms occur in

proximity in the karst areas of southeastern West Virginia. Many popula-

tions there are sharply genetically distinct, genetic breaks frequently coin-

ciding with divides between karstic sub-basins (Gooch and Hetrick 1979).

The present study is limited to Form III populations and inter-

gradational Form II, which comprise the great majority of G. minus

populations. We used, among others, the labile eye and appendage

characters of the Holsinger and Culver study. The nine populations oc-

cupied habitats which we rank ordered from most open or epigean to

least open (or marginal hypogean). Measurements were taken on each

population to determine the degree to which ecotype typifying

morphology conformed to the habitat scale. Strong conformance as

shown by similar rank order would indicate a predominant influence of

environmental factors related to habitat openness. Weak conformance

would indicate that other factors such as local adaptation to biotic or

physical conditions or the interplay between genetic drift and gene flow

strongly influence morphology.

METHODS AND MATERIALS

Samples of Gammams minus were taken using a Surber sampler and,

on highly irregular bottoms, a dip net. From each sample 50 sexually

mature individuals, 25 of each sex, were chosen at random. The following

measurements were made on one side, indiscriminantly right or left, on

each individual:

(1) Packing density and regularity of the eye. The facets bordering

the eye may be tightly and regularly packed or loose and irregular,

producing, respectively, a border that is smooth or one that is ragged and

embayed. Holsinger and Culver treated this character in some detail and

depicted eye shape in typical members of the three habitat forms (their

Fig. 3). The latitude of regularity was much less in our populations. Eyes

were scored on an estimated ordinal scale of high, intermediate, and low

density and regularity.

(2) Lobe orientation. The eyes of typical epigean G. minus are short

reniform with the lower lobe usually broader than the upper (Cole 1970).

In some individuals the lobes are equal or the upper lobe is broader. Eyes

Gammarus Variability

135

were scored as broad lobe up, equal, or down. This character is not ob-

viously related to habitat but is simply an easily scored trait that may

show interdemic variation.

(3) Lobe angle. The reniform eye has a shallow, slightly variable

angle opening anteriorly. This angle was not measured but was scored

relatively on an ordinal scale from 1 to 3 indicating, respectively, more

acute, intermediate, and more obtuse. Like lobe orientation this is not an

ecotype differentiating character.

(4) Eye length. The length of the eye is successively less in Form II

and Form I amphipods. It was measured (mm) parallel to the long dor-

soventral axis of the eye.

(5) Length of antenna 1. Elongate first antennae are particularly

striking in cave ecotypes. The first antenna was measured from pedun-

cular base to flagellar tip.

(6) Length of pereopod 7. The seventh pereopod is relatively longer

in hypogean habitats. It was measured extended from coxal base to tip of

dactyl.

(7) Length of uropod 3. This is a highly variable biramous structure

in G. minus (Holsinger 1976) and is usually longer in cave forms. The long

ramus (exopod) was measured.

Measurements were taken on population samples from sites listed

below in inferred rank order, from most nearly hypogean to most open

eipigean.

(i) Emma Spring, Huntingdon Co., PA: strongly shaded spring dis-

charging from a subterranean conduit in a limestone rockface.

(ii) Smoke Hole Spring, Giles Co., VA: partially shaded spring pool

formed from outflow of Smoke Hole Cave.

(iii) Greenland Gap, Grant Co., WV: partly shaded runoff 4 m

downstream from a spring.

(iv) Cunninghams, Huntingdon Co., PA: unshaded runoff 7 m

downstream from spring.

(v) Marklesburg, Huntingdon Co., PA: shaded, heavily vegetated

first order stream collected 12 m downstream from small spring.

(vi) Church Camp, Centre Co., PA: shaded runoff 15 m downstream

from series of large ground seeps.

(vii-viii) Petersburg I and II, Huntingdon Co., PA: large, partly

shaded springfed stream; site I is a large, open, impounded pool, site II is

runoff about 20 m downstream from the pool.

(ix) James Creek, Huntingdon Co., PA: unshaded site in a second-

order stream about 3 km downstream from spring sources.

This rank order is subjectively based on proximity to subterranean

water source, likelihood that discharge is from a sizeable cave or conduit,

and amount of cover or shade. Emma Spring is very secluded. Smoke

Hole Spring receives the immediate discharge of Smoke Hole Cave,

which probably contains Form II amphipods as does nearby Tawneys

Cave, a site of Holsinger and Culver. These localities would be expected

to harbor populations bordering on Form IT The James Creek site is

136

James L. Gooch and Jeffrey S. Wiseman

completely open and relatively remote from hypogean environments, so

the Form III morphology should be well developed there. These are the

maximally contrasting sites and they define the extremes of the Form III

habitat spectrum. Other localities differ less strikingly and are ranked in

only an approximate way.

RESULTS

The ordinal scale scores of eye packing density, broad lobe orienta-

tion, and lobe angle of population samples are given by sex in Table 1.

Localities are in rank order with the most epigean at bottom. Packing

density is relatively uniform, with most samples distributed in about a

40:60 ratio between high- and intermediate-density scores. Only a few

scattered individuals, constituting less than 1% of the total, have the low-

density irregular borders indicative of the typical Form II ecotype. There

is a shift from intermediate- to high-density eyes in increasingly open

populations. James Creek has the highest proportion of regular-eyed in-

dividuals, although it does not differ significantly from Emma Spring,

using the R x C Chi-square test, with pooling of low and intermediate

scores. However, the second to fourth ranked sites have low high-density

scores and a 2 x 2 contingency table of the four highest ranking popula-

tions with the five lowest, pooled by site and sex, yields a significant dif-

ference (X- = 6.53, 1 df, p C.05). Much of the difference is contributed

by Smoke Hole Spring, whose entire sample consists of intermediate den-

sity eyes. When sexes pooled over populations are compared by con-

tingency table there is a significantly greater proportion of high-density

eyes in females (2C = 9.83, 1 df, p <^.01). We have no hypothesis to ac-

count for this difference, but since females are smaller it may be related to

size rather than sex.

Table 1 shows little interdemic variation in broad lobe orientation or

lobe angle. All pairwise contingency table tests were performed on the

sample distributions of both characters, with broad lobe up pooled with

equal lobe scores. Of 36 tests none indicated significant differences be-

tween localities for lobe orientation. The overall distribution, to which all

population samples conform fairly closely, is in the ratio 7:33:60, broad

lobe up, equal, and down, respectively. This confirms Cole’s (1970) ob-

servation that the lower lobe is usually broader in G. minus. Although

lobe orientation is quite variable within populations, interdemic varia-

tion is too low for this character to be useful in geographic studies. Lobe

angle is only slightly more variable, with 4 of 36 pairwise tests yielding

significant differences between sites. All involve the Petersburg I sample,

which has a high proportion (0.49) of individuals with more obtuse

angles. The overall ratio is 16:51:33, more acute, intermediate, and more

obtuse angle, respectively. This character also appears to have little value

in studies of geographical variation.

The characters eye, antenna 1, pereopod 7, and uropod 3 lengths will

Gammarus Variability

137

Table 1. Scores of eye facet packing density (high, intermediate, low), lobe

orientation (broad lobe up, equal, down), and lobe angle (more acute,

intermediate, more obtuse). Localities are in rank order, with the most

epigean at bottom.

Packing density Broad lobe Lobe angle

be treated together. These characters, except for eye length, also were in-

vestigated by Holsinger and Culver. They are correlated with body length

by the relative growth equation T = a + c, in which Y is any one of the

above characters as the dependent variable; X is the independent

variable, body length; a is the slope of the regression line; b is the coef-

ficient of allometry; and c is the intercept of the regression line on the or-

dinate. If b is other than 1 there is nonlinear relative growth (allometry)

of parts. If ^ = 1 there is no allometry and the equation simplifies to T =

a X c, the linear regression equation. Allometry in populations of dif-

fering size distributions would make meaningful comparisons more dif-

138

James L. Gooch and Jeffrey S. Wiseman

Table 2. Linear regression of eye and appendage lengths against body length.

Values are for 25 males (upper row) and 25 females (lower row) per

locality, averaged over 9 localities. Further explanation in text.

Dependent variable

slopes Y-interceptc

ficult. Holsinger and Culver obtained values of b close to unity and con-

cluded that allometry was not important in the adult growth of antenna

1, pereopod 7, or uropod 3 relative to body length. We also tested for

allometry by solving for b in the equation Y = a b log X (Frazzetta

1975), obtaining the slope b from plots of sample means of eye and ap-

pendages against mean body lengths of males and females separately.

Values of b ranged between 0.78 and 1.15 for appendage growth, none of

which was significantly different from unity. For eye length, b for males

was 1.08 ± 0.21 and for females 0.67 ± 0.28. The latter figure suggests

negative allometry, but the large standard error and the anomalously low

coefficient of determination for females (0.12) make this estimate of b

meaningless. With qualification for eye length in females, we conclude

with Holsinger and Culver that over the range of measurements used in

adult G. minus allometry is a minor factor in determining length ratios.

Interdemic regression data on sample mean eye and appendage

lengths against mean body lengths are presented in Table 2. There are no

significant differences in slope between sexes. Excepting female eye

length, the coefficients of determination are between 0.61 and 0.80, in-

dicating moderate scatter of locality means and probably reflecting non-

uniform slopes and intercepts of growth equations of different popula-

tions. The slopes for appendage growth are generally slightly higher than

those obtained by Holsinger and Culver for Form III amphipods, but not

significantly so.

The ratios of eye and appendage length to body length in rank or-

dered habitats are given in Table 3. Mean length of males is 9.28 ±0.16

mm, about 1 mm less than Holsinger and Culver determined for III

habitats. Mean female length is 6.73 ± 0. 14 or 73% of male length. There

is considerable interdemic variation in female/male length ratio, from

Gammarus Variability

139

IX

9.28 ±.16 6.73 ±.14 0.050 0.054 0.576 0.546 0.483 0.451 0.201 0.169

140 James L. Gooch and Jeffrey S. Wiseman

Table 4. Pairwise comparisons of eye/body length ratios vs. antenna 1/body

length ratios (top) and uropod 3/body length ratios vs. pereopod 7/

body length ratios (bottom) of males of rank ordered localities. Single

asterisks indicate significant differences at the 0.05 level, double asterisks

at the 0.01 level, using the Mann-Whitney U Test.

Eye/body

0.64 at Marklesburg to 0.90 at Cunninghams. Although Holsinger and

Culver found generally larger amphipods in cave populations, there is no

evident trend toward larger size in either sex in our less epigean sites.

The last four columns of Table 3 display length ratios and feature

two points of interest: nonsystematic interdemic variation in ratios, and

systematic variation associated with rank in the habitat scale. Non-

systematic variation was assessed by testing the ratio arrays of all charac-

Per-7/body Ant-l/body

Gammarus Variability

141

Table 5. Pairwise comparisons in females. Captions as in table 4,

Eye/body

ters in pairwise locality comparisons using the Mann-Whitney V test.

The results for males are shown in Table 4 and for females in Table 5. In-

terdemic variation in length ratios is extensive. Of 288 pairwise tests, 144

per sex, 166 or 58% show significant (p <C.05) or very significant (/?<

.01) differences between localities. Variation is greater in males (65% of

tests) than females (50%), which may be related to size rather than sex.

Among characters with sexes pooled, interdemic differences are greatest

in eye length ratio (51 of 72 comparisons, 71%) and are at roughly the

same level for appendage ratios (antenna 1, 56%; pereopod 7, 57%;

uropod 3, 47%). Some of this variation is due to comparison of samples

from strongly contrasting habitats at opposite ends of the ranking. Some

142

James L. Gooch and Jeffrey S. Wiseman

is nonsystematic and unrelated to the epigean-hypogean scale as shown

by the fact that many similarly ranked sites differ significantly in ratios.

Tables 4 and 5 show this qualitatively: ratio variation due purely to

habitat difference would produce a pattern of significant differences

(asterisks) in comparisons of widely rank separated sites and non-

significant differences (dashes) in comparisons of similar rank. However,

asterisks and dashes are so interspersed as to indicate numerous signifi-

cant differences between like ranked localities.

Systematic variation lies in the association of ratios with habitat

rank. If the eye and appendage size differences between ecotypes are also

distinguishable within the single Form III ecotype, eye length ratio

should increase and appendage length ratios should decrease with in-

creasing (more epigean) rank. The predicted trends are roughly con-

firmed in Table 3. Some ratios, however, do not conform to expectation

for their rank. This is especially so for eye length in both sexes and

pereopod 7 length in females, which reveal no discernible trends.

Holsinger and Culver obtained appendage ratios in males of the

three ecotypes. Their mean values, followed by ours in parentheses, are:

antenna 1, I .720, II .644, III .570, (.576); pereopod 7, I .496, II .463, III

.442 (.483); uropod 3, I .21 1, II .190, III .184 (.169). Our figures approx-

imate the published ones for Form III amphipods except for pereopod 7,

which has almost the relative length of Form I. Agreement is good con-

sidering the wide range of ratios found in both investigations and the

small sample sizes used in the earlier study.

In 24 of 27 comparisons male appendage ratios are higher than

female ratios. The sex differences are significant at the 0.05 level by the

Mann-Whitney U test for all 3 appendages. Ruling out allometry, these

data indicate generally greater appendage elongation in males than

females. The effect does not extend to eye length, which does not differ

significantly between sexes. Curiously, the three instances of higher ap-

pendage ratio in females belong to the James Creek population. Unless

parasitism or an unusual environmental factor was responsible (neither

were evident), this suggests that relative appendage elongation between

sexes is genetically labile and occasionally prone to variation in semi-

isolated populations.

The findings on all habitat associated characters are summarized in

Table 6, which presents each locality rank ordered by its score or ratio,

progressing to epigean typical values to the right. For each site there is

considerable scatter in rank by both sex and character. Nevertheless the

pattern of low rank for borderline type II habitats and high rank for fully

open type III habitats does emerge. Emma Spring and Smoke Hole

Spring, for example, rank near the hypogean pole for most characters

and James Creek and Petersburg I and II usually rank near the epigean.

At the foot of Table 6 overall rank order is given and each locality name

is provided below in parentheses by its mean rank over all characters.

This provides us with two scales of rank order— the habitat scale

Gammarus Variability

143

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144

James L. Gooch and Jeffrey S. Wiseman

previously given and this character scale. A one-to-one conformance of

ranks would indicate that interdemic variation in these characters is com-

pletely determined by openness of habitats and their proximity to

hypogean environments. The scales conform well for six localities, which

conform from zero to two rank positions. Three large discrepancies do

occur and require explanation: (1) Cunninghams has the highest

hypogean character score, very narrowly over Smoke Hole Spring, but

ranks as only the fourth most hypogean habitat; (2) Greenland Gap

ranks eighth in character scale, third in habitat; (3) James Creek ranks

sixth and ninth, respectively. It should be underscored that neither scale

is an absolute standard to which the other can be compared. A difference

in rank may mean faulty ranking of habitat, i.e. the habitat is less or more

epigean to the amphipods than to the observer, or that factors other than

habitat scale affect the characters and their rank. Our discussion can take

factors of the second type into account more easily than the first. For ex-

ample, James Creek is almost certainly the most fully epigean site, yet its

character rank does not reflect this (note, however, that its mean rank is

only 0.75 positions higher than that of the ninth ranked locality). Its

overall rank is higher due simply to the high appendage ratios of its

females, as was mentioned earlier and is apparent from Tables 3 and 6.

Presumably slender-limbed females is a genetic characteristic indigenous

to the James Creek population and is unrelated to the habitat scale.

The rank discrepancies of Greenland Gap and Cunninghams are at

first puzzling. Greenland Gap is about 175 km southwest of seven of the

localities and could obey different ecogeographic rules. However, Smoke

Hole Spring and most of the localities of Holsinger and Culver are even

more distant and they match the Pennsylvania populations quite closely

in character trends. One potential influence on ecotype that has not been

considered is migration and attendant gene flow. Populations subjected

to high migration rates from contrasting habitats would not undergo as

marked ecotypic differentiation as isolated demes. This is especially true

if the phenotypic expression is largely under genetic control. The

Greenland Gap site is only 2 m upstream from North Fork Patterson

Creek, which is an open, typical Ill-habitat harboring a population of G.

minus. It is probable that gene exchange between the populations has

prevented the Greenland Gap population from developing the characters

associated with its habitat type. Cunninghams, on the other hand, feeds

immediately into Standing Stone Creek, a stream that lacks G. minus. We

infer that Cunninghams is thus a highly isolated deme that has developed

strongly habitat specific characters. No other locality appears to be as ex-

posed to gene flow or as isolated as these two, although this factor has

probably influenced character scores everywhere to some extent.

DISCUSSION AND CONCLUSIONS

A summary analysis of our data falls into three categories. First, a

Gammarus Variability

145

morphological profile of the common epigean ecotype was done, ex-

tending the pioneering work of Holsinger and Culver to additional pop-

ulations, both sexes, and larger sample sizes. Second, the level of inter-

demic variability was determined and variability was related to the

epigean-hypogean habitat spectrum and to other factors. Last, the

overall pattern of interdemic variation was evaluated, which also may be

applicable to other freshwater species.

All characters displayed high intrademic variability, and eye density

and length ratios of eye and appendages also varied significantly among

demes. Lobe orientation and lobe angle scores were distributed quite

uniformly over populations and thus have less value in geographical

studies in the Appalachians. Growth of appendages was approximately

linear on body length in the adult size range, without significant dif-

ferences in slope between sexes or large differences in intercepts. Growth

equations were different enough among populations, however, to give

coefficients of determination usually less than 0.70 on locality mean

lengths. Female amphipods averaged 73% the length of males, but

proportionate size of sexes varied widely. We have no evidence as to

genetic or directly environmental causes of size differences. Females also

had statistically significant smaller appendage length ratios than males.

Interdemic character variation was both systematic, that is habitat

scale related, and nonsystematic. The former is evidenced by the finding

that Form III demes in shaded, secluded springs near cave or conduit dis-

charge usually had more irregular eyes and more elongate appendages

than more open populations. At Emma Spring and Cunninghams, at

least, there are no known Form II populations. This suggests that

morphology is an in situ adaptation and not the result of genetic mixing

with cave ecotypes. These intergradational populations would be ex-

pected to possess slightly reduced eyes as well. Although we have in-

cluded eye length ratio in the composite character scale there is no clear

trend toward eye reduction (Table 3). We conclude that ecotypical

characters are intergradational among epigean demes and that the

characters rank roughly in the same order as habitats scaled from fully

epigean to semihypogean.

The conformation of character and habitat rankings is very inexact.

Aside from sampling errors and imprecision in assigning habitat rank,

there is extensive nonsystematic and statistically significant interdemic

variability. Some of this variability may actually be systematic adapta-

tion to habitat variables not apparent to the observer. Minckley and

Cole (1963) found variation, mostly of setation patterns, in G. minus in

two Kentucky streams to be associated with lotic and lentic microen-

vironments, aquatic vegetation, and the presence of the probable com-

petitor Gammarus bousfieldi. Dickson (1977) noted that type and amount

of food influenced pigmentation and body and antenna length in the

troglobitic species Crangonyx antennatus.

A baseline datum in the present study is the existence of extensive

146

James L. Gooch and Jeffrey S. Wiseman

genetic differentiation among demes. This is true of the karst region

where the ecotypes were first described (Gooch and Hetrick 1979) and in

the central Pennsylvania localities studied here (Gooch and Golladay, in

press). The genetic investigations were done on three polymorphic

allozyme loci. Genetic patterns have not been found to correspond to

habitats or ecotypes and will not be discussed here. They do clearly in-

dicate that the most geographically isolated populations often carry

atypical alleles in high frequencies, either due to local adaptation or

genetic drift, and that demes linked by likely avenues of migration have

not undergone as much genetic differentiation.

James Creek, with its aberrant ratio of female/male appendage

lengths, is probably an example of a deme that has evolved mor-

phological characters in partial isolation from other populations.

Greenland Gap, on the other hand, apparently has been prevented from

acquiring habitat specific characters due to strong gene flow from other

habitats. The mid-Appalachian area is an environmental mosaic of

epigean, intermediate, and hypogean habitats. The result in G. minus has

been the evolution of markedly differentiated ecotypes. Streams and

divides, however, provide clear avenues and barriers to migration,

leading to highly anisotropic gene flow. Superimposed on ecotypic varia-

tion and sometimes discordant with it is the local differentiation of

isolated demes. Ecotypic distinctions are further modified by gene flow

among demes that are open to migration.

ACKNOWLEDGMENTS.— Jhh research was carried out during

the tenure of NSF-URP Grant No. SPI-7827183, which provided a sti-

pend to the junior author. Funding under NSF Grant No. DEB75-03302

AOl to the senior author supported the initial research. We are indebted

to Drs. Gerald A. Cole, John R. Holsinger, and William R. Rhodes for

their constructive comments on earlier drafts of the manuscript.

LITERATURE CITED

Cole, Gerald A. 1970. Gammarus minus: Geographic variation and description

of new subspecies G.m. pinnicollis (Crustacea, Amphipoda). Trans. Am.

Microsc. Soc. (59:514-523.

Croker, R.A., and M.F. Gable. 1977. Geographic variation in western Atlantic

populations of Gammarus oceanicus Segerstrale (Amphipoda). Crustaceana

32:55-76.

Dickson, Gary W. 1977. Variation among populations of the troglobitic amphi-

pod crustacean Crangonyx antennatus Packard living in different habitats.

I. Morphology. Int. J. Speleol. 9:43-58.

Frazzetta, T.H. 1975. Complex adaptations in evolving populations. Sinauer

Associates, Inc., Sunderland, Mass. 267 pp.

Gooch, James L., and S. Golladay. In Press. Genetic population structure in

an amphipod species. Int. J. Speleol.

, and S.W. Hetrick. 1979. The relation of genetic structure to environ-

mental structure: Gammarus minus in a karst area. Evolution 33:192-206.

Gammarus Variability

147

Holsinger, John R. 1976. The freshwater amphipod crustaceans (Gammaridae)

of North America. Biota of Freshwater Ecosystems. Ident. Man. 5, U.S.

Environ. Protect. Agency, Washington. 89 pp.

, and D. Culver. 1970. Morphological variation in Gammarus minus

(Amphipoda, Gammaridae) with emphasis on subterranean forms. Postilla

746:1-24.

Hubricht, Leslie. 1943. Studies on the Nearctic freshwater amphipoda, III. Notes

on the freshwater Amphipoda of the eastern United States, with descrip-

tions of ten new species. Am. Midi. Nat. 29:683-712.

Minckley, W.L., and G.A. Cole. 1963. Ecological and morphological studies on

gammarid amphipods {Gammarus sp.) in spring-fed streams of northern

Kentucky. Occasj Pap. Adams Cent. Ecol. Stud. West. Mich. Univ. 70:1-35.

Pinkster, S. 1971. Members of the Gammarus pulex group (Crustacea-

Amphipoda) from North Africa and Spain, with description of a new

species from Morocco. Bull. Zool. Mus. Univ. Amst. 2:45-61.

1972. On members of the Gammarus pulex group (Crustacea -

Amphipoda) from western Europe. Bijdr. Dierk. 42:164-191.

Shoemaker, Clarence R. 1940. Notes on the amphipod Gammarus minus Say and

description of a new variety, Gammarus minus var. tenuipes. J. Wash. Acad.

Sci. 50:388-394.

Accepted 3 May 1980

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Ozarka, A New Subgenus of Etheostoma

(Pisces: Percidae)

James D. Williams

U. S. Fish and Wildlife Service,

Office of Endangered Species, Washington, D. C. 20240

AND

Henry W. Robison

Department of Biological Sciences,

Southern Arkansas University, Magnolia, Arkansas 71753

ABSTRACT. — A new subgenus of Etheostoma is diagnosed and briefly

described. It consists of five species, Etheostoma punctulatum, E. cragini,

E. pallididorsum, E. boschungi, and E. trisella, which have similar

breeding colors, tubercle patterns and spawning habitats. The species

are distributed from the Arkansas River drainage in Colorado to the up-

per Coosa River drainage in north Georgia and southeast Tennessee.

Distribution, dispersal and relationships of the species are discussed,

and a key to the species is presented.

INTRODUCTION

The nominal darter genera were reduced to four {Ammocrypta Jor-

dan, Etheostoma Rafinesque, Hadropterus Agassiz, and Percina

Haldeman) by Bailey (1951) and further reduced to three {Ammocrypta,

Etheostoma, and Percina) by Bailey (in Bailey et al. 1954). Bailey and

Gosline (1955), in a review of the vertebral counts of American percids,

placed the 70 species of Etheostoma in 12 subgenera {Boleosoma, loa,

Etheostoma, Ulocentra, Allohistium, Nothonotus, Oligocephalus,

Austroperca, Psychromaster, Catonotus, Hololepis, and Microperca).

Based primarily on the presence and distribution of breeding tubercles,

Collette and Yerger (1962) and Collette (1965) recognized an additional

subgenus, Villora. The most recent works by Collette and Banarescu

(1977) and Page (1977) recognized the additional subgenera Doration,

Litocara, and Vaillantia, for a total of sixteen.

The subgenus Oligocephalus is the most wide-ranging, large, struc-

turally diverse (Bailey and Richards 1973), complex, and speciose (Ram-

sey and Suttkus 1965) subgenus of Etheostoma. Bailey and Gosline (1955)

assigned 19 species and Collette (1965) 21 species to this subgenus. Ram-

sey and Suttkus ( 1 965) discussed the E. asprigene species group within the

subgenus Oligocephalus.

The purpose of this paper is to remove E. punctulatum, E. cragini, E.

pallididorsum, and E. boschungi from the subgenus Oligocephalus, and E.

trisella from the subgenus Psychromaster, and to erect a subgenus for

these closely related species. The close relationship among E. punc-

tulatum, E. cragini, and E. pallididorsum was first recognized by Blair

(1964). Wall and Williams (1974) described E. boschungi and presented

additional data clarifying the relationships among E. boschungi, E.

punctulatum, E. cragini, and E. pallididorsum.

Brimleyana No. 4: 149-156. December 1980.

149

150

James D. Williams and Henry W. Robison

Ozarka, new subgenus

Type Species. — Etheostoma punctulatum {\g2iss\z)= Poecilichthys

punctulatus Agassiz 1854 (original description, type locality Osage River,

Missouri).

Species Included in Ozarka.— £. punctulatum, E. cragini, E. pallididor-

sum, E. boschungi, and E. trisella. Etheostoma trisella was originally

placed in the subgenus Psychromaster by Bailey and Richards (1963)

while the other four species have been referred to the subgenus

Oligocephalus (Bailey and Gosline 1955; Collette 1965; Wall and

Williams 1974). The subgenus name Ozarka is taken from the Ozark

Mountains Physiographic Provice, which we believe to have been the

center of dispersal for the subgenus.

Diagnosis. — A subgenus of the genus Etheostoma of moderate to small

size, adult males and females ranging from 40-70 mm standard length

(SL). Cheeks and opercles usually naked or with embedded scales, except

E. trisella which has scales on these structures. Lateral line incomplete,

except complete in E. trisella. Lateral line scale rows 40-80, anterior por-

tion not arched upward. Transverse scale rows 12-24, caudal peduncle

scales 20-29. Branchiostegal membranes narrowly joined to overlapping;

branchiostegal rays usually 6-6 and unsealed. Frenum present, broad and

well developed. Preoperculomandibular canal pores usually 10, infraor-

bital canal complete with 7 or 8 pores, supratemporal canal complete or

interrupted. Preopercle entire. Caudal peduncle deep. Vertebrae 32-39,

usually 34-37. Caudal vertebrae usually 18, 19, or 20. First interneural

spine between the neural spines of third and fourth or fourth and fifth

vertebrae.

Dorsal spines VI-XII usually IX-XI; dorsal rays 10-15, usually 11-

14; pectoral rays 9-14, usually 11-13; anal spines 2 except usually 1 in E.

trisella; anal rays 6-10, usually 7-9. Males with breeding tubercles on

scales of belly, around base of anal fin posteriorly to caudal peduncle,

and on anal spines and rays and ventral surface of pelvic rays; tubercles

absent in females. Sexual dichromatism pronounced in breeding adults,

males brightly colored, females not. Breeding males with bold blue-black

subocular bar, width one-third to two-thirds diameter of orbit. First dor-

sal fin of breeding male with black marginal band (usually narrow an-

teriorly, increasing in width posteriorly), submarginal orange to red-

orange band, and blue-green to blue-black basal band. Submarginal and

basal bands approximately equal in width. Second dorsal of breeding

males without bright colors. Venter from pectoral and pelvic fin inser-

tions to caudal peduncle suffused with varying concentrations of orange

to red-orange pigment. Branchiostegal and gular region orange except in

E. trisella. Genital papilla tubular in females, not long and tubular in

males. Anus not surrounded by fleshy villi.

New Etheostoma Subgenus

151

Additional Characters. — Body slender to moderately stout; snout

moderately to slightly decurved; mouth terminal, slightly oblique;

premaxillary frenum present, generally broad and well developed. Head

moderately large; eye breaking dorsal contour of head in lateral view;

caudal fin slightly rounded; branchiostegal membranes separate to

narrowly joined, rays 6-6; preopercle entire; pectoral fin length usually

shorter than head length. Lateral line complete or incomplete, arching

gently anteriad; lateral line scales 40-80; vertebrae 32-39, usually 34-37.

Transverse scale rows 12-24; caudal peduncle scales 20-29. Dorsal fin

spines VI-XII, usually IX-XI; dorsal fin soft rays 10-15, usually 11-14;

anal spines II, except in E. trisella which usually has I anal spine; anal fin

soft rays 6-10, usually 7-9; branched caudal fin rays 12-17; pectoral fin

rays 9-14. Supratemporal canal complete or interrupted; lateral canal

complete with 5 pores; single coronal pore; postorbital, interorbital,

posterior nasal and anterior nasal pores present; preoperculomandibular

canal complete with 10 pores; infraorbital canal complete with 7 or 8

pores. Nape scaly with exposed or embedded ctenoid scales. Cheeks and

opercles naked or scaly; prepectoral region generally naked or with a few

scales; anterior portion of belly naked to fully scaly; breast naked or with

a few embedded scales. Nuptial tubercles present in males only; tubercles

on scales around base of anal fin, anal spines and rays and ventral sur-

faces of pelvic rays. Breeding tubercles absent in females. Genital papilla

sexually dimorphic.

Body generally olivaceous to grayish brown. Well developed hu-

meral spot present in all species except E. trisella where it may be in-

distinct. Bold, dark subocular bar present. Lateral blotches variable; dis-

continuous or fused into irregular lateral band in E. cragini. Dorsal sad-

dles 3-9, conspicuous, and dark. Sexual dichromatism pronounced in all

species. Well developed orange to red-orange in breeding males located

on the venter from pelvic fins posteriad to caudal fin. Spinous dorsal fin

with black marginal band (may be absent in some individuals), usually

narrow anteriorly, increasing in width posteriorly. Submarginal orange

to red-orange band below the dark marginal band and followed by blue-

green to blue-black basal band of equal width. All species lack bright

breeding colors in soft dorsal fin. Soft dorsal fin with six to eight in-

distinct horizontal bands formed by dark spots of pigment on rays, and

darker pigment on fin membranes giving a barred appearance to fins.

Caudal fin with spots confined to rays, arranged in four to six irregular

vertical rows. Spots on pectoral and pelvic fins aligned in irregular ver-

tical rows. Pelvic fins typically dusky to gray-black; pigment usually

restricted to rays or proximal membranes of fin. Anal fin with scattered

spots on rays and membranes. Gular and branchiostegal regions orange.

Females with generally olivaceous or brownish-gray bodies. Bright

orange coloration not observed in females. Spinous dorsal fins, while oc-

casionally tinged with orange or yellow pigment, never brightly banded

as in males. Soft dorsal fin without bright colors; some irregular spotting

of rays occurs.

152

James D. Williams and Henry W. Robison

Distribution. — The geographic range of the five species of the subgenus

Ozarka is centered in the Ozark Mountains Physiographic Region. The

species are found from eastern Colorado {E. cragini) to southeastern Ten-

nessee and northwestern Georgia {E. trisella). They are allopatric except

for E. punctulatum and E. cragini, which are sympatric in southwestern

Missouri and northeastern Oklahoma. Etheostoma punctulatum, E.

cragini, and E. pallididorsum are found west of the Mississippi Embay-

ment, and E. boschungi and E. trisella east of the Embayment. None of

the species is known from the Coastal Plain Province.

The stippled darter, Etheostoma punctulatum, is known from the

Arkansas River drainage in northwestern Arkansas (Buchanan 1973),

northeastern Oklahoma (Miller and Robison 1973), extreme south-

eastern Kansas (Cross 1967), and southern Missouri (Pflieger 1975). It

also occurs in the Missouri River drainage in central Missouri and the

Castor River system, tributary to the Mississippi River, in southeastern

Missouri (Pflieger 1975) as well as in the White River drainage of

southern Missouri and northern Arkansas (Buchanan 1973; Pflieger

1975). Distribution of the Arkansas darter, E. cragini, is in the Arkansas

River drainage from eastern Colorado (Ellis 1914; Ellis and Jaffa 1918),

southern Kansas (Cross 1967), northeastern Oklahoma (Miller and

Robison 1973), and extreme southwestern Missouri (Pflieger 1975). In

the original description, E. pallididorsum was reported from the Caddo

River of the Ouachita River drainage in western Arkansas (Distler and

Metcalf 1962). Robison (1974a) reported an additional population from

the upper Ouachita River drainage.

East of the Mississippi Embayment, Etheostoma boschungi is known

from widely separate tributaries of the Tennessee River in western Ten-

nessee and northern Alabama (Wall and Williams 1974). Etheostoma

trisella is known from isolated localities in the Coosa River drainage in

northeastern Alabama (Bailey and Richards 1963; Ramsey 1976),

northwest Georgia (Howell and Caldwell 1967), and southeastern Ten-

nessee (Etnier 1970). Based on available distributional data, E. trisella

has the most limited distribution of the species in the subgenus. The only

extant population known is in the Conasauga River near the Tennessee-

Georgia border.

Habitat. — Members of the subgenus Ozarka typically inhabit gentle

riffles and slackwater areas of small to medium-size shallow, upland

tributary streams. Etheostoma punctulatum is generally restricted to

small, clear, moderate to high gradient permanent streams or spring

branches with substrates of gravel or rubble (Blair 1959; Miller and

Robison 1973; Pflieger 1971, 1975). It is frequently found in vegetation or

in detritus in quiet side pools and backwaters away from the main current

(Branson 1967). Moore and Paden (1950) reported that this form was

taken principally in heavily vegetated springs with slight gradient and in

small leaf-filled indentations along the shore.

New Etheostoma Subgenus

153

Etheostoma cragini prefers quiet pools of the smallest spring

branches. It also occurs in spring-fed creeks where it is most often found

along the shallow margins of pools and riffles in thick growths of water-

cress, Nasturtium officinale (Ellis and Jaffa 1918; Blair 1959; Branson

1967; Cross 1967; Pflieger 1971, 1975; Miller and Robison 1973). Moore

and Cross (1950) collected E. cragini in small clear streams of moderate

current over mud, gravel, and sand substrates in quiet pools in which

aquatic vegetation flourished (Ranunculus, Potamogeton, Myriophyllum,

Callitriche, and Radicula).

Etheostoma pallididorsum inhabits small, spring-fed brooks, 0.6 m in

average width and 5 cm in depth over mud, gravel and/or rubble bottoms

(Distler and Metcalf 1962). The species typically prefers shallow (15-30

cm) backwater pool areas with leaf-litter and small gravel-rubble bot-

toms when found in larger streams (Robison 1974a, b; Hambrick and

Robison 1979).

Etheostoma boschungi inhabits clear, medium-current, second and

third order streams ranging in width from 3 to 6 m, and ranging in depth

from less than 15 cm to 1 .7 m (Wall and Williams 1974). Boschung (1976)

collected this species over gravel infiltrated by silt, and over silt and mud,

but never over clean gravel. Individuals seemed to prefer accumulations

of detritus in areas of relatively low water velocity.

Etheostoma trisella was hypothesized by Bailey and Richards (1963)

to live in springs, although the holotype was collected from a small,

sluggish pasture stream with a bottom of silt mixed with sand and gravel

and heavily overgrown with emergent Diathera. Etnier (1970) reported

that E. trisella appears to inhabit riffles and almost stagnant quiet

backwaters of small, low-gradient streams.

Breeding habits and habitats of all species except E. punctulatum

have been examined to some degree (Ellis and Jaffa 1918; Distler 1972;

Boschung 1976; M. Ryon, pers. comm.; and HWR and JDW, pers. ob-

serv.). Although no actual observations of spawning of E. punctulatum

have been made, field data from several workers attest to the presence of

nuptial males during the spring (W. Pflieger, pers. comm.; L. Knapp,

pers. comm.; and HWR, pers. observ.).

The unique spawning habitat of the species of Ozarka affirms their

close phylogenetic relationship. All typically live in or enter small tribu-

tary streams during later winter and spawn during early spring (March-

April). Etheostoma pallidisorsum, E. boschungi, and E. trisella enter and

spawn in tiny spring-fed rivulets or seepage water in open fields that

drain into nearby streams. More specific life history information linking

these three species is available to us and will be published later by HWR

(E. pallididorsum), H. T. Boschung (£. boschungi), and M. Ryon (E.

trisella). A detailed analysis of the systematics of the five species of

Ozarka will be forthcoming from the authors and B. R. Wall.

154

James D. Williams and Henry W. Robison

Key to Species of the Subgenus Ozarka

1. Anal spines one; lateral line complete with 44-52 scales; three distinct

dorsal blotches Etheostoma trisella.

(Coosa River system in Alabama and Georgia, Conasauga River

system in Tennessee.)

Anal spines two; lateral line incomplete with 5-59 pored scales;

dorsal blotches variable 2

2. Lateral line with more than 30 pored scales 3

Lateral line with 5-25 pored scales 4

3. Lateral line scales 58-80; soft dorsal fin usually 14 or 15 rays

Etheostoma punctulatum.

(Arkansas River drainage in Arkansas, Oklahoma, Kansas and

Missouri, White River drainage in Arkansas and Missouri, and

Missouri River drainage and Castor River system of Missouri.)

Lateral line scales 43-58 (usually 34-38 pored); soft dorsal fin

usually 1 1 or 12 rays Etheostoma boschungi.

(Tennessee River drainage in northern Alabama and southcentral

Tennessee.)

4. Prominent pale mid-dorsal stripe present; venter behind pelvics

naked; prepectoral areas naked Etheostoma pallididorsum.

(Upper part of Caddo River system and Hallmans Creek in upper

Ouachita River system, Arkansas.)

No prominent pale mid-dorsal stripe; venter behind pelvics fully

scaly; prepectoral areas with few scales Etheostoma cragini.

(Arkansas River drainage in Colorado, Kansas, Oklahoma, and

Missouri.)

ACKNOWLEDGMENTS.— VsIq acknowledge with thanks the

assistance of the following individuals. For the loan of material we are

grateful to H. T. Boschung, University of Alabama; N. H. Douglas,

Northeast Louisiana University; E. A. Lachner, National Museum of

Natural History; D. A. Etnier and M. Ryon, University of Tennessee;

and J. T. Collins, University of Kansas. Thanks are due H. T.

Boschung, G. Clemmer, M. Ryon, D. A. Etnier, L. W. Knapp, and

W. Pflieger for use of color slides and notes and miscellaneous life

history information concerning members of Ozarka. Special thanks

are due Dr. George A. Moore, Oklahoma State University, for

suggesting the name Ozarka for the new subgenus.

LITERATURE CITED

Agassiz, Louis. 1854. Notice of a collection of fishes from the southern bend

of the Tennessee River, in the State of Alabama. Am. J. Sci. Arts, Ser. 2,

77:297-308, 353-369.

Bailey, Reeve M. 1951. A checklist of the fishes of Iowa, with keys for identifi-

cation. pp. 185-237 in R. Harlan and E. B. Speaker. Iowa Fish and Fishing.

Iowa Conserv. Comm., Des Moines.

New Etheostoma Subgenus

155

, and W. A. Gosline. 1955. Variation and systematics significance of

vertebral counts in the American fishes of the family Percidae. Univ.

Mich. Mus. Zool. Misc. Publ. 9i:l-44.

, and W. J. Richards, 1963. Status of Poecilichthys hopkinsi Fowler

and Etheostoma trisella, new species, percid fishes from Alabama, Georgia,

and South Carolina. Occas. Pap. Mus. Zool. Univ. Mich. 630:\-2\.

, H. E. Winn and C. L. Smith. 1954. Fishes from the Escambia River,

Alabama and Florida, with ecologic and taxonomic notes. Proc. Acad.

Nat. Sci. Phila. 706:109-164.

Blair, Albert P. 1959. Distribution of the darters (Percidae, Etheostomatinae)

of northeastern Oklahoma. Southwest. Nat. 4(1):1-13.

1964. Habitat and other notes on Etheostoma pallididorsum

(Etheostomatini: Percidae). Southwest. Nat. 9(2): 105-107.

Boschung, Herbert T. 1976. Amplification of final environmental impact state-

ment for Cypress Creek watershed. USDA Soil Conserv. Serv., Auburn,

Alabama. 49 pp.

Branson, Branley A. 1967. Fishes of the Neosho River system in Oklahoma.

Am. Midi. Nat. 75(1): 126-154).

Buchanan, Thomas M. 1973. Key to the fishes of Arkansas. Arkansas Game

Fish Comm., Little Rock. 68 pp., 198 maps.

Collette, Bruce B. 1965. Systematic significance of breeding tubercles in fishes

of the family Percidae. Proc. U.S. Natl. Mus. 777:567-614.

, and P. Banarescu. 1977. Systematics and zoogeography of the fishes

of the family Percidae. J. Fish. Res. Board Can. 34(10): 1450-1463.

, and R. W. Yerger. 1962. The American percid fishes of the subgenus

Villora. Tulane Stud. Zool. 9(4):21 3-230.

Cross, Frank B. 1967. Handbook of fishes of Kansas. Univ. Kans. Mus. Nat.

Hist. Misc. Publ. 45. 357 pp.

Distler, D. A. 1972. Observations on the reproductive habits of captive Etheo-

stoma cragini Gilbert. Southwest. Nat. 76(3-4):439-441 .

, and A. L. Metcalf. 1962. Etheostoma pallididorsum, a new percid fish

from the Caddo River system of Arkansas. Copeia 1962(3):556-561 .

Ellis, M. M. 1914. Fishes of Colorado. Univ. Colo. Stud. Ser. Biol. 77(1):1-136.

, and B. B. Jaffa. 1918. Notes on Cragin’s darter, Catonotus cragini

(Gilbert). Copeia 1918(59):73-75.

Etnier, David A. 1970. Additional specimens of Etheostoma trisella (Percidae)

from Tennessee. Copeia 1970(2):356-358.

Hambrick, P. S., and H. W. Robison. 1979. Life history aspects of the paleback

darter, Etheostoma pallididorsum (Pisces: Percidae) in the Caddo River

system, Arkansas. Southwest. Nat. 24(3):475-484.

Howell, W. M., and R. D. Caldwell. 1967. Discovery of a second specimen of the

darter, Etheostoma trisella. Copeia 1967(l):235-236.

Miller, Rudolph J., and H. W. Robison. 1973. The fishes of Oklahoma. Okla.

State Univ. Mus. Nat. Hist. Ser. 11. 246 pp.

Moore, George A., and F. B. Cross. 1950. Additional Oklahoma fishes with

validation of Poecilichthys parvipinnis (Gilbert and Swain). Copeia 1950

(2): 139-148.

, and J. M. Paden. 1950. The fishes of the Illinois River in Oklahoma

and Arkansas. Am. Midi. Nat. 44(l):76-95.

Page, Larry M. 1977. The lateralis system of darters (Etheostomatini). Copeia

1977(3):472-475.

156

James D. Williams and Henry W. Robison

Pflieger, William L. 1971. A distributional study of Missouri fishes. Univ. Kans.

Publ. Mus. Nat. Hist. 20(3):225-570.

1975. The fishes of Missouri. Mo. Dep. Conserv., Jefferson City.

viii -I- 343 pp.

Ramsey, John S. 1976. Freshwater fishes, pp. 53-65 in H. Boschung (ed.). En-

dangered and Threatened Plants and Animals of Alabama. Bull. Alabama

Mus. Nat. Hist. No. 2. 93 pp.

, and R. D. Suttkus. 1965. Etheostoma ditrema, a new darter of the

subgenus Oligocephalus (Percidae) from springs of the Alabama River Basin

in Alabama and Georgia. Tulane Stud. Zool. 72(3):65-77.

Robison, Henry W. 1974a. An additional population of Etheostoma pallididorsum

Distler and Metcalf in Arkansas. Am. Midi. Nat. 9y(2):478-479.

1974b. Threatened fishes of Arkansas. Proc. Ark. Acad. Sci. 25:59-64.

Wall, Benjamin R., and J. D. Williams. 1974. Etheostoma boschungi, a new percid

fish from the Tennessee River drainage in northern Alabama and western

Tennessee. Tulane Stud. Zool. 75(4): 172-182).

Accepted 24 October 1980

Stomach Contents of Some

Snakes from Eastern and Central North Carolina

Richard F. Collins^

Department of Zoology, North Carolina State University,

Raleigh, North Carolina 27607

ABSTRACT. — Stomach contents of eight species of snakes from the

Coastal Plain and Piedmont Plateau regions of North Carolina were

identified. The majority of the snakes were of the genus Nerodia: N.

sipedon sipedon contained primarily amphibians; N. taxispilota con-

tained fishes; and single specimens of N. erythrogaster erythrogaster

and N.fasciata fasciata contained frogs. Agkistrodon piscivorus pisci-

vorus was omnivorous, and A. contortrix contortrix contained a frog

and a small mammal.

Food habits of snakes from various localities in the United States have

been noted by a number of authors (see Brown 1979). That author also

provided food records for a number of snakes in North and South

Carolina. In addition to those studies cited by Brown, several other

authors furnished pertinent information on snake food habits. Mushinsky

and Hebrard (1977) and Oliver (1970) gave records for Nerodia spp. and

Elaphe obsoleta, respectively. Stomach contents of 17 Agkistrodon pisci-

vorus leucostoma were noted by Collins and Carpenter (1970) and of 4

A. p. piscivorus by Goodman (1958). The food items of 93 individuals of

the latter subspecies were provided by Wharton (1969). He studied

snakes from Sea Horse Key, Florida, and also listed the findings of a

number of other authors. Kofron (1978) studied several species of

Nerodia as well as ^4./?. leucostoma from a variety of habitats. Although

the subspecies were different from those in my study, their food habits

appeared to be similar in terms of species designations. However, the

small number of Nerodia spp. examined in both studies precludes any

definite conclusions.

Arthropods are not common foods of most colubrids or crotalids.

However, as Brown noted, it is probably erroneous to assume that some

species do not take arthropods under certain conditions. Of the species 1

examined, only a single specimen of Elaphe o. obsoleta contained

arthropod material, a larva of Phengodes sp. (glowworm). This snake was

a mature female that did not contain any other food remains. Brown did

not report arthropods from 39 specimens of this snake. However,

arthropod remains (primarily lepidopteran larvae) were noted by him in 9

Agkistrodon contortrix. I found a small mammal and a Rana catesbei-

iCurrent address: College of Osteopathic Medicine and Surgery, 3200 Grand Avenue,

Des Moines, Iowa 50312

Brimleyana No, 4: 157-159. December 1980.

157

158

Richard F. Collins

Table I. Stomach contents of snakes from eastern and central North Carolina

ana in the two individuals in my study. Amphibians were not found in

any of the 35 A . contortrix examined by Brown.

The Nerodia species examined in my study comprise the majority of

individuals examined. There are no major differences between the classes

of food items found by Brown and by me. However, some additional

species were seen. Amphibians were the major food of N. s. sipedon. In

addition to the items listed by Brown, one individual contained an adult

Notophthalnius viridescens. This snake was captured in a farm pond. A

N . f. fasciata contained a Rana palusfris, an item not found by Brown in

12 snakes of this species. This snake was captured in a swamp near the

Northeast Cape Fear River in Duplin County. Natrix taxispilota was not

studied by Brown. As noted above, this species is primarily riverine in

eastern North Carolina. Unlike the other species of Nerodia studied,

which apparently feed mainly on amphibians, its primary food source

probably is fish. This was also noted by Laughlin (1959) for this species

from a lacustrine habitat in Oklahoma.

A^kistrodon p. piscivorus exhibited a more varied diet than any of the

other snakes examined. These data, along with those of Hamilton and

Snake Stomach Contents

159

Pollack (1955), Goodman (1958), Laughlin (1959), Wharton (1969),

Collins and Carpenter (1970), Kofron (1978), and Brown, show that this

species will feed on certain members of every class of vertebrates. In

addition, Collins and Carpenter (1970) reported insects in the stomach

contents of two snakes.

Only three individuals contained mixed categories of food items: a

Nerociia s. sipeclon contained four toads and a fish; an Agkistroc/on p.

piscivorus contained a small mammal and a bass; and an A . c. confortrix

contained a Rana catesheiana and a small mammal. None of the other

snakes contained more than a single class of food item. The total number

of snakes containing any food items was therefore 24, including the three

individuals mentioned above.

Whether certain snake species can be characterized as “opportunists”

in terms of food items consumed is conjectural. Of the species examined,

A^kistrocion piscivorus most readily can be characterized this way, but

some species of Nerociia also may be opportunists. Brown mentioned

observing N . f. fasciata eating road-killed anurans; I have seen N. ery-

throgaster transversa eating chunks of fresh fish. Snakes generally are

considered to be predators, but these kinds of observations indicate that

under certain circumstances some species can adopt other feeding habits.

LITERATURE CITED

Brown, E.E. 1979. Some snake food records from the Carolinas. Brimleyana

1:113-124.

Collins, Jospeh T., J.E. Huheey, J.L. Knight and H.M. Smith. 1978. Standard

common and current scientific names for North American amphibians .and

reptiles. Soc. Study Amphib. Reptiles Herpetol. Circ. No. 7. 36 pp.

Collins, Richard F., and C.C. Carpenter. 1970. Organ position-ventral scute

relationship in the water moccasin (A^ki.strodon piscivorus leucosionia),

with notes on food and distribution. Proc. Okla. Acad. Sci. 49:15-18.

Goodman, John D. 1958. Material ingested by the cottonmouth, AykistroJon

piscivorus, at Reelfoot Lake, Tennessee. Copeia 1958 (2): 149.

Hamilton, W. J.,. Jr., and J.A. Pollack. 1955. The food of some crotalid snakes

from Fort Benning, Georgia. Nat. Hist. Misc. (Chic.) No. 140:1-4.

Kofron, Christopher P. 1978. Food and habitats of aquatic snakes (Reptilia,

Serpentes) in a Louisiana swamp. J. Herpeto! 72:543-554.

Laughlin, Harold E. 1959. Stomach contents of some aquatic snakes from

Lake McAlester, Pittsburgh County, Oklahoma. Tex. J. Sci. 77:83-85

Mushinsky, Henry R., and J.J. Hebrard. 1977. Food partitioning by five

species of water snakes in Louisiana. Herpetologica JJ: 162-166.

Oliver, George V., Jr. 1970. Black ratsnake predation upon nesting barn

and cliff swallows. Bull. Okla. Ornithol. Soc. i: 17-20.

Wharton, C.H. 1969. The cottonmouth moccasin on Sea Horse Key, Florida.

Bull. Fla. State Mus. Biol. Sci. 74:227-272.

Accepted 18 June 1980

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A

Mandibular Dental Anomaly in

White-tailed Deer

George A. Feedhamer and Joseph A. Chapman

Appalachian Environmental Laboratory,

Gunter Hall, Frosthurg State College Campus,

Frost burg, Maryland 21532

A BSTRACT. — Agenesis of the second mandibular premolar in four

White-tailed deer from Dorchester County, Maryland, was considered

to be of genetic origin. The body condition of anomalous animals

apparently was not affected.

The dentition of most mammalian species has been well studied

because of its importance in systematics and evolution, and in estimation

of individual age. As a result, associated dental anomalies have been

described for a variety of species encompassing many mammalian orders

(Choate 1968; Colyer 1936; Lavelle and Moore 1972; Pavlinov 1975;

Sheppe 1963). This paper describes a dental anomaly found in 4 of 24

yearling and adult White-tailed deer, Odocoileus virginianus, obtained

during the 1976 through 1978 hunting seasons in southern Dorchester

County, Maryland. The skull and dentary bone were collected from each

animal. Skulls were solicited and obtained from hunters at two deer

check stations. No samples were examined prior to solicitation and all

hunters entering the stations were approached. Thus, the sample was

considered to be random.

Bilateral agenesis of the second premolar (p2) (Fig. 1) was found in two

female and one male yearlings, all in which the permanent dentition had

erupted. A unilateral P2 agenesis occurred in an adult female. In all four

occurrences, the anomaly was considered to be genetic in origin because:

1) there was no evidence of previous traumatic injury; 2) no alveoli were

present at the P2 position; and 3) X-rays of the dentary revealed no

vestigial or impacted teeth in the underlying bone tissue at the P2 positions.

The mean total length of the mandibular tooth row for animals with the

anomaly was significantly less than that of animals with a normal

complement of mandibular cheek teeth(t = 4.80; p< 0.001). Flowever, all the

anomalous animals had a normal complement of maxillary cheek teeth,

exhibited normal occlusion and showed no unusual wear.

Benson ( 1 957) attributed a missing second premolar in two White-tailed

deer to traumatic injury. No sample size was given and only a single

mandible was available from each animal. A unilateral absence of P2 in a

Mule deer, Odocoileus hemionus, similarly was attributed to injury (Short

and Short 1 964). Apparent genetic agenesis of the second premolar involv-

Brimleyana No. 4; 161-163. December 1980.

161

162

Richard F. Collins

Fig. 1. Representative bilateral agenesis of second premolar (P2) in a White-tailed

deer (AEL #1 103) from Dorchester County, Maryland.

ing one or both mandibles was noted in only 5 of 422 ( 1 .2%) White-tailed

deer from New York (Free et al. 1972), and 8 of 401 (2.0%) White-tails

from northern Minnesota (Mech et al. 1970). This anomaly also has been

described in Roe deer, Capreolus capreolus, by Meyer ( 1977). It is interest-

ing that this anomaly apparently did not occur among 33,337 White-tailed

deer examined over a 3-year period in Michigan (Ryel 1963). Although

from a limited sample, the occurrence of this characteristic in 16.7 percent

of the White-tailed deer examined from Dorchester County suggests the

trait may be well established in this population.

Missing second premolars probably were not detrimental to the overall

condition or survival of the individual deer (Pekelharing 1968). Body

weights and standard measurements from two anomalous animals from

the 1978 sample were comparable to those animals with normal dentition.

Weights and measurements of the remaining three anomalous animals

were not available. This lends support to the suggestion of previous

investigators (Manville 1963, Smith et al. 1977) that dental anomalies,

although of intrinsic interest, probably are of little significance to the total

dynamics of the population.

Deer Dental Anomaly

163

ACKNOWLEDG M ENTS. Dr. R. W. Bosely, DDS, kindly X-rayed

the deer jaws. These findings were incidental to a continuing study funded

through Federal Aid to Wildlife Restoration Grant W-49-R, Maryland.

This is Contribution No. 1090, Appalachian Environmental Laboratory,

Center for Environmental and Estuarine Studies, University of Maryland.

All specimens are housed in the AEL museum.

LITERATURE CITED

Benson, D. A. 1957. Abnormal dentition in White-tailed deer. J. Mammal.

3(^:140.

Choate, Jerry R. 1968. Dental abnormalities in the short-tailed shrew, Blarina

hrevicauda. J. Mammal. 49:251-258.

Colyer, F. 1936. Variations and diseases of the teeth of animals. John Bale, Sons

and Danielsson, Ltd. London. 750 pp.

Free, Stuart L., A. S. Bergstrom and J. E. Tanck. 1972. Mandibular and dental

anomalies of White-tailed deer. N. Y. Fish Game J. /9:32-46.

Lavelle, C. L. B., and W. J. Moore. 1972. The incidence of agenesis and

polygenesis in the Primate dentition. Am. J. Phys. Anthropol. i<^:67 1-680.

Manville, Richard H. 1963. Dental anomalies in North American lynx. Z

Saeugetier. 28: 1 66- 1 69.

Mech, L. David, L. D. Frenzel, Jr., P. D. Karns and D. W. Kuehn. 1970.

Mandibular dental anomalies in White-tailed deer from Minnesota. J.

Mammal. 57:804-806.

Meyer, Von P. 1977. Innate hypodontia in Roe deer (Capreolus capreolus L.). Z.

Jagdwiss. 23:98-100.

Pavlinov, 1. Y. 1975. Tooth anomalies in some Canidae. Acta Theriol. 20:507-5 1 9.

Pekelharing, C. J. 1968. Molar duplication in red deer and wapiti. J. Mammal.

49:524-526.

Ryel, Lawrence A. 1963. The occurrence of certain anomalies in Michigan white-

tailed deer. J. Mammal. 44:79-98.

Sheppe, Walter. 1963. Supernumerary teeth in the deer mouse, Peronivscus

Z. Saeugetier. 29:33-36.

Short, Henry L., and C. P. Short. 1964. Abnormal dentition in a Colorado

mule deer. J. Mammal. 45:3 1 5.

Smith, James D., H. H. Genoways and J. K. Jones, Jr. 1977. Cranial and

dental anomalies in three species of Platyrrhinc monkeys from Nicaragua.

Folia Primatol. 2(^: 1-42.

Accepted 5 June 1980

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DATE OF PUBLICATION

Brimleyana No, 3 was mailed on 30 July 1980.

ERRATUM

The following error appeared in Brimleyana No. 3:

Page 95, line 1 — insert “ihS.E.” between 3.5 and 0.3 mm.

JOURNAL RECEIVES STC INTERNATIONAL AWARD

The Society for Technical Communications bestowed an Award for Merit on

Brimleyana in its Eleventh Annual International Technical Communications

Competition for 1980. The journal was entered in the Complete Periodicals

category of the competition, through STC’s Carolina Chapter. STC is the world’s

largest professional society devoted to technical communications, and has more

than 65 chapters in the United States and abroad.

164

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CONTENTS

The Milliped Fauna of the Kings Mountain Region of North Carolina

(Arthropoda: Diplopoda). Marianne E. Filka and Rowland M.

Shelley 1

Electrophoretic Analysis of Three Species of Necturus (Amphibia:

Proteidae), and the Taxonomic Status of Necturus lewisi (Brimley).

Ray E. Ashton, Jr., Alvin L. Braswell and Sheldon /. Guttman 43

Vertebrates of theOkefenokee Swamp. Joshua Laerm, B.J. Freeman,

Laurie J. Vitt, Joseph M. Meyers diX\d Lloyd Logan 47

New Records, Distribution and Diagnostic Characters of Virginia

Ictalurid Catfishes with an Adnexed Adipose Fin. Noel M.

Burkhead, Robert E. Jenkins and Eugene G. Maurakis 75

Geographic Variation in the Snake Storeria occipitomaculata (Storer)

(Serpentes: Colubridae) in Southeastern United States. Douglas A.

Rossman and Robert L. Erwin 95

Effects of Microhabitat Size and Competitor Size on Two Cave Isopods.

David C. Culver and Timothy J. Ehlinger 103

Life History of the Mottled Sculpin, Cottus bairdi, in Northeastern

Tennessee (Osteichthyes: Cottidae). Jerry W. Nagel 115

Notes on the Distribution and Ecology of the Black Mountain Dusky

Salamander Desmognathus welteri Barbour (Amphibia: Plethodon-

i\ddiQ)\n'TQnv\QssQQ. William H. Redmond 123

Morphological and Habitat Variability in Gammarus minus Say (Amphi-

poda: Gammaridae). James L. Gooch and Jeffrey S. Wiseman .... 133

Ozarka, a New Subgenus of Etheostoma (Pisces: Percidae). James D.

Williams and Henry W. Robison 149

Stomach Contents of Some Snakes from Eastern and Central North

CaxoWna. Richard F. Collins 157

Mandibular Dental Anomaly in White-tailed Deer. George A. Feldhamer

and Joseph A. Chapman 161

Errata and Miscellany

164