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number 8 december 1982

EDITORIAL STAFF

John E. Cooper, Editor

Alexa C. Williams, Managing Editor

John B. Funderburg, Editor-in-Chief

Board

Alvin L. Braswell, Curator of David S. Lee, Chief Curator

Lower Vertebrates, N.C of Birds and Mammals, N.C

State Museum State Museum

John C. Clamp, Associate Curator William M. Palmer, Chief Curator

(Invertebrates), N.C of Lower Vertebrates, N.C

State Museum State M

useum

Martha R. Cooper, Associate Rowland M. Shelley, Chief

Curator (Crustaceans), N.C. Curator of Invertebrates, N.C.

State Museum State Museum

James W. Hardin, Department

of Botany, N.C. State

University

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

tory, will appear at irregular intervals in consecutively numbered issues. Con-

tents will emphasize zoology of the southeastern United States, especially North

Carolina and adjacent areas. Geographic coverage will be limited to Alabama,

Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North

Carolina, South Carolina, Tennessee, Virginia, and West Virginia.

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

geography, evolution, and behavior. Subdiscipline areas will include general in-

vertebrate zoology, ichthyology, herpetology, ornithology, mammalogy, and

paleontology. Papers will stress the results of original empirical field studies, but

synthesizing reviews and papers of significant historical interest to southeastern

zoology will be included.

Suitability of manuscripts will be determined by the Editorial Board, and ap-

propriate specialists will review each paper adjudged suitable. Final ac-

ceptability will be decided by the Editor. Address manuscripts and all cor-

respondence (except that relating to subscriptions and exchange) to Editor,

Bnmleyana, N. C. State Museum of Natural History, P. O. Box 27647, Raleigh,

NC 27611.

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

Terrestrial Isopods (Crustacea: Isopoda:

Oniscoidea) from North Carolina

George A. Schultz

75 Smith Street, Hampton, New Jersey 08827

ABSTRACT.— The 18 terrestrial isopod crustaceans from North

Carolina are reviewed with synonymies, illustrations, ecological notes

and a key to species. Maps of distribution also are included. The spe-

cies can be divided into two major groups — species exclusively from

the seashore and species from more upland locations. There is only a

small overlap since some species from more upland locations also

occasionally are abundant locally at the edge of brackish and marine

waters. Species from more upland locations are further divided accord-

ing to the moisture of their habitat. Species of Ligidium, like related

Ligia exotica of the seashore, are found in permanently moist habi-

tats. Species such as Hyloniscus riparius also live in very wet habitats.

Others, such as species of Miktoniscus, are found in the moist, rotted

logs and leaf litter of forested river bottomlands. Species such as Por-

cellio scaber and P. laevis are common around human habitation in

relatively dry places. Species such as Armadillidium vulgare and A.

nasatum are more tolerant of moisture and dryness and are found in

the greatest number of widely distributed habitats in the state.

INTRODUCTION

Terrestrial isopod crustaceans (Oniscoidea) have not previously

been studied in detail in North Carolina or any other part of the south-

eastern United States. The first actual record of an oniscoid in North

Carolina was of Ligia sp. (most probably L. exotica Roux) from Ft.

Macon (Harger 1878). Brimley (1938) later included six species in an

account of the insects and some other arthropods of the state. Schultz

(1962) recorded Miktoniscus halophilus Blake from the Piedmont and

Coastal Plain (Schultz 1976) and Philoscia vittata (Say) from the coast

(Schultz 1963, 1974). Ligidium blueridgensis Schultz (1964) from the

western part of the state was then described, along with a record of

Miktoniscus medcofi (Van Name) from a cave. Schultz (1975, 1977)

recorded species from coastal Georgia, some of which range into North

Carolina, and Kelley (1978) recorded terrestrial isopods from the coastal

zone of South Carolina, some of which also range into North Carolina.

This paper is based on specimens collected by the author and on

many others kindly made available by Dr. Rowland M. Shelley, North

Carolina State Museum of Natural History, Raleigh. Some other small

collections are also included. Specimens of 17 of the 18 species (except

M. medcofi) are in the collection of that museum. Distributions within

the state are recorded on Maps 1-12. A short discussion and an illustra-

tion of each species is included so the records of the 18 species from

Brimleyana No. 8:1-26. December 1982 1

2 George A. Schultz

North Carolina are brought together in an orderly manner for the first

time. A key to the species is provided, and their classification is given in

Table 1.

MATERIALS AND METHODS

Isopods were collected by searching leaf litter, and looking under

bark and in and under decaying logs. They were sought under rocks and

trash along roadsides, in fields and in woods. Leaf litter and moss along

shaded streams and in swamps were also examined. Maritime drift lines

along margins of bays and the ocean were examined, including the

undersides of all flotsam. Isopods were preserved in 70% ethyl alcohol.

Whole animals were examined with a binocular microscope, and their

appendages (in glycerine mounts) with a compound microscope.

Table 1 . Checklist of terrestrial isopod crustaceans (Oniscoidea) recorded from

North Carolina.

Ligiidae

Ligia exotica Roux 1828

Ligidium elrodii (Packard 1873)

Ligidium blueridgensis Schultz 1964

Scyphacidae

Armadilloniscus ellipticus (Harger 1878)

Philosciidae

Philoscia vittata Say 1818

Trichoniscidae

Trichoniscus pusillus Brandt 1833

Miktoniscus halophilus Blake 1931

Miktoniscus medcofi (Van Name 1940)

Hyloniscus riparius (Koch 1838)

Haplophthalmus danicus Budde-Lund 1885

Oniscidae

Oniscus asellus Linnaeus 1758

Cylisticidae

Cylisticus convexus (De Geer 1778)

Porcellionidae

Porcellio scaber Latreille 1 804

Porcellio laevis Latreille 1 804

Porcellio virgatus (Budde-Lund 1885)

Porcellionides pruinosus (Brandt 1833)

Armadillidiidae

Armadillidium nasatum Budde-Lund 1885

Armadillidiwn vulgare (Latreille 1804)

North Carolina Terrestrial Isopods 3

DEFINITIONS

The following terms are used in the key and discussions. They will

be helpful to the non-specialist.

FLAGELLUM — thin apical part of antenna 2 divided into two or

more ARTICLES.

OCELLUS — unit of eye (compound eye has two or more ocelli); the

eye can have one to many ocelli in isopods.

UROPODS — posterior appendages ("tails" in some species).

PLEOPODS — appendages of ABDOMEN (pleon), or posterior five

segments plus pleotelson of body. Each pleopod has an EXOPOD

(outer branch) and an ENDOPOD (inner branch). Both branches of

pleopods 1 and 2 (the first and second small appendages behind the

seventh pair of walking legs) are sexually modified in male oniscoids.

The endopod of each of the pleopods usually is elongate and serves as a

secondary sexual organ. Males can be distinguished from females by

the presence of these modified pleopods.

PLEOTELSON — sixth abdominal segment, fused with telson (last

body segment) and with uropods as appendages.

THORAX (peraeon) — main seven segments of body behind head, in

front of abdomen.

EPIMERE — lateral extensions of thoracic and abdominal segments in

oniscoids.

PROCESS— a projection.

TUBERCULATE— with small bumps.

Van Name (1936) included a general account of the morphology of

terrestrial isopods in his introduction.

KEY TO SPECIES OF TERRESTRIAL ISOPODS

OF NORTH CAROLINA

All 18 species of oniscoids so far recorded within the borders of

North Carolina are included here. Certain species not yet recorded also

might be present. Specimens of such species could be erroneously identi-

fied by using only "key" characters, so comparison of specimens with

illustrations (especially the pleopods) is essential. Specimens not fitting

the key perhaps can be identified by use of the comprehensive works of

Van Name (1936, 1940, 1942). Color is a useful character for identifying

freshly caught isopods, but color fades on preservation and is only of

limited value as an aid for identifying preserved specimens. Color

phases also are common in populations of some species. Base color (all

species are multicolored) is included in parentheses in the key.

Many people think that tiny or small isopods are the young of

larger species. This might be true, especially in spring and early summer,

but some small specimens are distinct species and live in association

with larger specimens of other species. Specimens of all sizes then must

4 George A. Schultz

be carefully examined with a binocular microscope. The length of a

large specimen is recorded with the discussion of each species, and a line

representing 1 mm is present at the lower left of each illustration. With

the key one should be able to identify all sexually mature and larger

specimens of each species in North Carolina.

More than eight, or many flagellar articles on antenna 2;

uropods long and thin 2

Flagellar article number four, three, two or indistinct; uro-

pods inconspicuous or relatively short 4

Many flagellar articles (gray) Ligia exotica

Eight to 1 5 flagellar articles (brown) 3

With lateral process on apex of endopod of male pleopod 2

(Fig. 19) Ligidium blueridgensis

With apex of endopod of male pleopod 2 rounded (Fig. 18)

or squarish (Fig. 20) Ligidium elrodii

Eye of many ocelli 9

Eye of one ocellus or three ocelli 5

Eye of one ocellus 6

Eye of three ocelli (reddish) Trichoniscus pusillus

With body pigment; body surface dull with tiny bumps or

body surface shiny 7

Without body pigment; body surface strongly tuberculate

Haplophthalmus danicus

Body surface dull with many tiny bumps; four inconspicuous

flagellar articles (salmon red) 8

Body surface shiny; five or more inconspicuous flagellar

articles (burgundy red) Hyloniscus riparius

Exopod of male pleopod 1 elongate, ending in point; tip of

endopod long and round (Fig. 15)

Miktoniscus halophilus

8b. Exopod of male pleopod 1 short; apex produced into small

process; tip of endopod long and flattened (Fig. 14)

Miktoniscus medcofi

9a (4). Two flagellar articles on antenna 2 12

9b. Four or three flagellar articles on antenna 2 10

10a (9). Three flagellar articles; moderately large specimen 11

10b. Four flagellar articles; small elliptical specimen

Armadilloniscus ellipticus

1 la (10). Body broad, elliptical; epimeres long Oniscus asellus

1 lb. Body narrow, posterior (abdomen) abruptly narrower than

anterior part; epimeres short Philoscia vittata

12a (9). Uropods extend past posterior body margin; some roll into

ball 14

12b. Uropods contained within general margin of body; roll into

ball 13

13a (12) With broad projection or "shelf on anterior margin of head

(Fig. 30) (gray) Armadillidium nasatum

North Carolina Terrestrial Isopods 5

13b. With no broad projection on anterior margin of head (Fig.

30) (black or yellow-brown) Armadillidium vulgare

14a (12). Does not roll into ball 15

14b. Rolls into ball (black) Cylisticus convexus

15a (14). Head relatively smooth 16

15b. Head strongly tuberculate (Fig. 22) (brown)

Porcellio scaber

16a (15). Posterior margin of thoracic segment I straight; pleotelson

triangular (Fig. 24) (gray) Porcellio laevis

16b. Posterior margin of thoracic segment I recurved laterally;

pleotelson produced into point 17

17a (16). Proximal article of flagellum longer than distal article;

exopod of pleopod 1 of male with small spines (reddish-

brown — frosted) Porcellionides pruinosus

17b. Proximal article of flagellum about as long as distal article;

exopod of pleopod 1 of male with large spines (Fig. 27)

(brown) Porcellio virgatus

SPECIES DISCUSSIONS

Ligia exotica Roux

Figs. 1,2

Ligyda exotica (Roux). Richardson 1905:676, figs. 716-718.

Ligia exotica Roux. Van Name 1936:48, fig. 8. Stephenson and Ste-

phenson, 1952:29.

Ligia sp. Harger 1878:310.

Harger's 1878 record of Ligia sp. from Ft. Macon most probably is

the first record of the species from North Carolina. Richardson (1905)

recorded it from nearby Beaufort and it is here recorded from around

Beaufort and at Cape Hatteras. The species is abundant on rocky jetties,

piers, and other hard substrates, both natural and man-made, from

Florida to Virginia and perhaps north to New Jersey. If relatively cool

daytime retreats are present the nocturnally active species can be very

abundant at any site. The species is also present along the margin of salt

water bays and estuaries, which are so prevalent on the North Carolina

coast, but details of its habitats along these margins have never been

studied. Gravid females were collected in August and September, but

they probably are present from early summer until fall. The species is

distinguished from others with which it might be confused by the con-

figuration of the apex of the endopod of male pleopod 2 (Fig. 2). Spec-

imens, especially in autumn, range up to 32 mm long. Published records

of the species in North Carolina are plotted on Map 1, but it is undoubt-

edly present in many suitable habitats along most of the coastline of the

state.

George A. Schultz

North Carolina Terrestrial Isopods

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George A. Schultz

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North Carolina Terrestrial Isopods

10

George A. Schultz

North Carolina Terrestrial Isopods

11

12

George A. Schultz

Figs. 1-7. 1, Ligia exotica. 2, apex endopod pleopod 2 of male L. exotica. 3,

Armadilloniscus ellipticus. 4, Philoscia vittata. 5, pleopods 1 of male P. vittata.

6, Porcellionides pruinosus. 7, exopod pleopod 1 of male P. pruinosus. Line at

lower left of all illustrations represents 1 mm.

North Carolina Terrestrial Isopods

13

Figs. 8-16. 8, Oniscus asellus. 9a, exopod pleopod 1 of male O. asellus. 9b,

exopod pleopod 1 of female O. asellus. 10, lateral view head of Trichoniscus

pusillus. 11, Hyloniscus riparius. 12, exopod and endopod pleopod 1 of male H.

riparius. 13, Miktoniscus sp. 14, exopod and endopod pleopod 1 of male M.

medcofi. 15, exopod and endopod pleopod 1 of male M. halophilus. 16,

Haplophthalmus danicus.

14 George A. Schultz

Ligidium elrodii (Packard)

Figs. 17, 18, 20

Ligidium longicaudatum Stoller. Van Name 1936:70, fig. 24. Brimley

1938:503. Schultz 1961:194.

Ligidium elrodii (Packard). Schultz 1970:39, figs. 4-11.

The species was discussed by Schultz (1970) who pointed out that

L. longicaudatum Stoller was a junior synonym of L. elrodii. In eastern

United States it ranges from near Chicago to New York state, and from

Arkansas to North Carolina. It was first recorded in North Carolina by

Brimley (1938) from Pungo Lake, Bunnlevel, and the Duke Forest. Dis-

tribution is shown on Map 6. In the Durham region the species lives

around the edges of springs and along the margins of shaded permanent

streams. In the Beaufort region it lives in wet, swampy old-fields near

woodlands. It is abundant in both places. The species is shiny dark

brown and can attain a length of 10 mm. It can run quickly when dis-

turbed and was found to be active when snow was removed from near a

spring. In summer, as the water sources dry and moisture diminishes,

the species retreats to damper places. Only species of Ligia have more

flagellar articles and larger eyes than do most members of Ligidium.

Since members of the genus are apparently endemic to North America,

more work should be done studying the distribution and ecology of the

species.

A different morphology is present on the endopod of male pleopod

2 of some specimens of L. elrodii (Figs. 18, 20) and they are from loca-

tions shown on Map 6. What the taxonomic status of this variant is has

yet to be determined. Distribution of the two species of Ligidium (Map

6) overlap, although specimens of L. elrodii from the western parts of its

range in the United States have not been examined morphologically.

Comparison of the tips of the endopods of pleopods 2 can be used to

distinguish L. elrodii from L. blueridgensis.

Ligidium blueridgensis Schultz

Fig. 19

Ligidium blueridgensis Schultz 1964:90, figs. 1-19. Schultz 1970:38

This species was originally described from northern Georgia just'

south of the North Carolina border (Schultz 1964). It is here recorded

from several places in western North Carolina (Map 6). The species is

easily distinguished from others in the genus by the long process project-

ing laterally from the apex on male pleopod 2 (Fig. 19). A male was

collected from leaf mold at 2500 ft. (762.5 m) on Ramsey Prong, near

Greenbrier, Robertson County, Tennessee, which is about 25 mi. (40

km) north of Nashville. This record considerably extended its range and

further established it as a species. The specimens are in the collection of

North Carolina Terrestrial Isopods

15

Figs. 17-25. 17, Ligidium sp. 18, exopod and endopod pleopod 2 of male L.

elrodii. 19, exopod and endopod pleopod 2 of male L. blueridgensis . 20,

exopod and endopod pleopod 2 of male L. elrodii (variant). 21, Porcellio

scaber. 22, front view of head of P. scaber. 23, exopod of pleopod 1 of male P.

scaber. 24, Porcellio laevis. 25, exopod of pleopod 1 of male P. laevis.

16 George A. Schultz

the American Museum of Natural History (AMNH 8737). It apparently

lives in and near very wet habitats and can get to be 10 mm long.

Armadilloniscus ellipticus (Harger)

Fig. 3

Armadilloniscus ellipticus (Harger). Van Name 1936:102, fig. 45. Schultz

1963:26.

The species is abundant in the maritime drift line in and near Beau-

fort, and is especially abundant on and under old planks, stones and

other detritus beneath dead plant matter. Philoscia vittata occurs fre-

quently with A. ellipticus in the dead vegetation. The species has been

found all along the east coast of the United States, from Woods Hole,

Massachusetts, to south Florida. It is small and does not run when dis-

turbed, but instead clings to the bottom of hard substrates where it

remains long after the habitat has been disrupted. Because of its small

size (to 3.5 mm long), elliptical shape, and the four flagellar articles on

antenna 2 (Fig. 3), it cannot be confused with other species of the shore.

Philoscia vittata (Say)

Figs. 4, 5

Philoscia vittata (Say). Van Name 1936:115, fig. 2. Schultz 1974:147,

figs. 3b, c, f.

Philoscia robusta Schultz 1963:26, figs. 1-22.

Philoscia muscorum (Scapoli). Schultz 1965a: 107.

Confusion about the name and identification of this species was

reviewed by Schultz (1974) who clarified some misidentifications on

which his own error was based. Although the species probably does not

belong in Philoscia Latreille (Old World species only?), it should be

kept in the genus until all halophilic members of the Philosciidae from

the New World are reviewed. The species lives under dead seaweed and

detritus of the maritime drift, at the upper part of the beach and edges

of marshes along inland bays. It lives in the grasses of high tide washes

and swamps wet with tidal water.

The species is always taken near and often with Armadilloniscus

ellipticus, but is much more active and less secretive than that species.

Some ecology of the species in the Beaufort region was recorded by

Schultz (1963). I took the species at Hammocks Beach State Park,

North Carolina (Map 3), and it has been recorded on the coast of Geor-

gia (Schultz 1975), including St. Catherines Island (Schultz 1977). Male

pleopods 1 are large, with the apexes of the distal portion of the endo-

pods folded laterally (Fig. 5), setting the species apart from others in the

genus (see Schultz 1974 for comparisons). It ranges to 5 mm long.

North Carolina Terrestrial Isopods 17

Trichoniscus pusillus Brandt

Fig. 10

Trichoniscus demivirgo Blake. Van Name 1936:76, fig. 29.

The species T. demivirgo Blake has been shown to be the imported

T pusillus and not a distinct endemic species in North America as had

been thought. It is fairly common in northeastern United States, espe-

cially in association with Hyloniscus riparius in the leaf litter of wooded

river bottoms. It was found only in mountainous Watauga County. The

species attains a length of 5.5 mm, slightly smaller than Hyloniscus ripar-

ius, and is a paler red than that deep wine-red species (see Schultz

1965b). It has three ocelli in a characteristic triangular pattern (Fig. 10),

not one ocellus as in H. riparius. There are also fewer flagellar articles

on antenna 2 when compared to H. riparius. Most populations of the

species are composed of only females since the species reproduces par-

thenogenetically. It is probably more abundant in North Carolina than

the single record indicates (Map 7).

Hyloniscus riparius (Koch)

Figs. 11, 12

Hyloniscus riparius (Koch). Schultz 1965:131, figs. 1-20.

Schultz (1965b) redescribed and summarized the ecology of the

species in eastern United States. He included illustrations and described

the population structure of specimens from New Jersey. The species dis-

tribution in North Carolina is shown on Map 5. It is common in leaf

litter of wooded streams and river bottoms where it might share habitat

with Trichoniscus pusillus as it does in New Jersey. River bottoms in

North Carolina contain Miktoniscus halophilus (perhaps also M. med-

cofi) and the species of Ligidium. The presence of the other isopods

might influence establishment and spread of H. riparius in North Carol-

ina. The species is distinguished from T. pusillus by its shiny deep wine-

red color and single ocellus. It grows to a length of about 5.5 mm. Identi-

fication can be confirmed by examining the apex of the endopod of

male pleopod 1 (Fig. 12).

Miktoniscus medcofi (Van Name)

Fig. 14

Miktoniscus linearis (Patience). Schultz 1962:47; 1964:94, figs. 20-32;

1976:36.

Miktoniscus alabamensis Muchmore. Cooper and Cooper, 1977:211.

Specimens from North Carolina include an adult male and two

adult females taken from Linville Caverns (Map 7), McDowell County

(Schultz 1964). The species is probably more abundant in the western

18 George A. Schultz

part of the state than records indicate. Schultz (1976) summarized

knowledge of all members of the genus in North America. The species

differs from M. halophilus from eastern North Carolina in the structure

of male pleopod 1 (Fig. 14). It is about 5 mm long.

Miktoniscus halophilus Blake

Fig. 15

Miktoniscus halophilus Blake. Van Name 1936:88, fig. 36. Schultz

1976:29, figs. 1-39, 46.

Miktoniscus grayi Schultz 1962:47, figs. 1-19.

Miktoniscus sp. Schultz 1961:194.

This species lives in rotten logs and dense leaf litter in the Durham

region and in similar microhabitats in the salt marshes and other

marshes near the coast. It ranges from Massachusetts through New Jer-

sey to North Carolina (Schultz 1976), south to Georgia (Schultz 1975).

In forested land the species inhabits leaf litter and pulpy rot under loose

bark of well decayed deciduous logs, near spring and small streams

which are permanent and not subject to seasonal flooding or drying. In

coastal marshes it perhaps is inundated at extremely high tides. It lives

in leaf litter at the bases of reed grasses in salt marshes, frequently with

Porcellio scaber. The species is apparently common in eastern North

Carolina (Map 4) and is distinguished from M. medcofl by the differ-

ence in the configuration of male pleopod 1 (Fig. 15). Its length reaches

5.2 mm.

Haplophthalmus danicus (Budde-Lund)

Fig. 16

Haplophthalmus danicus (Budde-Lund). Van Name 1936:90, fig. 37.

Schultz 1965b: 134.

Because the species is tiny, unpigmented, and readily feigns death,

it can very easily be overlooked when making collections. It is the smal-

lest species in North Carolina, being only to 3.4 mm long. Its body is

cream colored (pigmentless) and highly tuberculate, and there is only

one ocellus. In the Durham region it lives in continually damp, but not

wet, places, such as well decayed, pulpy logs, or shaded leaf litter where

moisture is retained throughout the year. It was found in abundance

near Durham even in winter, in the pulpy remains of a damp, shaded

sycamore log that was partially covered with leaf litter. Gravid females

were collected in summer, but breeding season dates were not estab-

lished. At another site several hundred specimens were taken in a space

measuring about a square foot in area and two inches deep, in and

around part of a broken, decaying board. Since there were many pieces

of the board at the site, the isopods must have numbered in the thou-

sands. The species is easily identified (Fig. 16).

North Carolina Terrestrial Isopods 19

Oniscus asellus Linnaeus

Figs. 8, 9a, 9b

Oniscus asellus Linnaeus. Van Name 1936:182, fig. 97.

So far the only record in North Carolina is from the west (Map 7).

In more northern locations, such as Long Island, New York, it is com-

mon near the coast where it lives in leaf litter and under loose tree bark.

It clings to such hard substrates as the logs and the sides of rocks and

bricks and remains motionless and inconspicuous. It is a large, flat,

elliptical isopod with a slate-gray color and yellow spots. It is distin-

guished from other common large species by three, not two, flagellar

articles on antenna 2. It grows to 12 mm long.

Cylisticus convexus (De Geer)

Figs. 28, 29

Cylisticus convexus (De Geer). Van Name 1936:259, figs. 147A, 148.

Brimley 1938:502. Schultz 1961:194; 1965b:134.

Cylisticus convexus, like both Armadillidium nasatum and A. vul-

gare, rolls into a ball, but the ball is less spherical than that formed by

the Armadillidium species. Also, its uropods project well beyoud the

general margin of the body (Fig. 28). The species apparently ranges

throughout North Carolina, but records are scarce on the Coastal Plain

except in the Beaufort region where isopods were more than casually

sought. It lives in moderately moist habitats that must remain moist

throughout dry periods, especially late summer. In particular habitats

with more or less permanent moist retreats the species can occur in

extremely large numbers, especially in the spring. It often inhabits

decayed logs after carpenter ants have left. The species is shiny gray-

black and has long uropods, which separates it from species of Armadil-

lidium. It grows to 16 mm long. In the Durham region breeding takes

place from late May until late August.

Porcellio scaber Latreille

Figs. 21-23

Porcellio scaber Latreille. Van Name 1936:226, figs. 2, 3, 127A, 128.

Brimley 1938:502. Schultz 1961:194.

This is one of the most abundant and widespread species in North

Carolina, being found from the coast to the mountains. It frequently is

present in the maritime drift line and in leaf litter of reed grass swamps

on the coast. It also is occasionally present on and near rocky jetties

where it might sometimes be covered by marine water, and is common

in leaf litter and under logs in upland forests. It is abundant around

human habitations and in remote locations as well. Several color

phases, from chocolate brown to variegated yellow-brown frequently

are seen in populations throughout the United States. It is most easily

20

George A. Schultz

Figs. 26-31. 26, Porcellio virgatus. 27, exopod of pleopod 1 of male P. virgatus.

28, Cylisticus convexus. 29, exopod pleopod 1 of male C. convexus. 30, dorsal

view of head of Armadillidium nasatum. 31, Armadillidium vulgare.

distinguished from other species of Porcellio by the presence of tuber-

cles (scabers) on the head (Fig. 22). Such head tubercles are absent in

other species from the New World. They are present on the segments of

the thorax as well in P. scaber. From a few scattered records the species

breeds from late May to November in the Durham region. Specimens to

16 mm long are common.

Porcellio laevis Latreille

Figs. 24, 25

Porcellio laevis Latreille. Van Name 1936:229, fig. 129. Brimley 1938:502.

Schultz 1961:194.

In North Carolina this common, large species is collected almost

exclusively around human habitations. It is rare in woods and other

remote locations. In both the Durham region and at Beaufort it was

taken almost exclusively near building foundations and in refuse heaps.

North Carolina is probably the southern limit of its range, since it was

North Carolina Terrestrial Isopods 21

not found in comparable habitats in Georgia (Schultz 1975, 1977). The

adult is a large, elliptical, slate gray animal easily distinguished from

other species of Porcellio. Smaller specimens must be examined closely,

however, since they can be confused with small specimens of Porcellio-

nides pruinosus, which frequents the same sites. The species breeds from

early June to late July in the Durham region, and the brood time

extends to August at Beaufort. It ranges to 20 mm long in the Durham

region.

Porcellio virgatus (Budde-Lund)

Figs. 26, 27

Porcellionides virgatus (Budde-Lund). Van Name 1936:241, fig. 125.

North Carolina is apparently the northern limit of the range of this

species. It has been taken on and near marine shores, from northern

Mexico through Florida to North Carolina, and is well established in

live oak logs and leaf litter of the coastal forests in the state. It is com-

mon in the same habitats in Georgia (Schultz 1975, 1977). Since P. vir-

gatus is easily confused with other species of Porcellio, the exopod of

male pleopod 1 (Fig. 27) must be examined to correctly identify it. It

grows to 1 1 mm long.

Porcellionides pruinosus (Brandt)

Fig. 6

Porcellionides pruinosus (Brandt). Van Name 1936:238, figs. 127 A, 133,

134A. Brimley 1938:502.

Porcellionides pruinosus is a cosmopolitan species recorded from

all continents of the world except Antarctica. It is widely distributed in

the United States although it is never abundant at any one location.

Many specimens were taken with Porcellio scaber in the Durham region

and some with other species in the same region. A few specimens were

collected in the Beaufort region in habitats near, but not on, the shore

(Map 7). It also was present in coastal Georgia (Schultz 1975). Live

individuals are reddish and "frosted," but the "frost" is lost in preserva-

tive, where specimens become red-brown. The species is distinctive

because the distal article of antenna 2 in adults is much longer than the

proximal article. The configuration of the exopod of male pleopod 1

(Fig. 7) should be compared with that of other species for accurate iden-

tification. It grows to 12 mm long.

Armadillidium nasatum Budde-Lund

Fig. 30

Armadillidium nasatum Budde-Lund. Schultz 1961:193; 1965b:134.

Records from North America were reviewed by Schultz (1961). It is

recorded from the coast to the mountains in North Carolina (Map 11).

22 George A. Schultz

Interestingly, it was not recorded in coastal Georgia (Schultz 1975,

1977) where it was diligently sought. It lives in a wide range of habitats

from very dry (but not as dry as some of those of A. vulgare) to very

moist (but not as moist as some of those of Ligidium), and has been

collected in dense, moist leaf litter with Miktoniscus halophilus. It can

be found under loose tree bark and in dense leaf litter, especially near

houses and outbuildings. During prolonged rainy weather the species

frequently is seen on the sides of buildings where it has moved to avoid

the very wet conditions in the soil at the foundation. It is distinguished

from A. vulgare, also a "pill bug," by the "shelf on the front of the

head between the eyes (Fig. 30). Large individuals get to be 12 mm long.

The species is gray and white, and mature males are much darker than

females. They never are shiny black like males of A vulgare.

Armadillidium vulgare (Latreille)

Fig. 31

Armadillidium vulgare (Latreille). Van Name 1936:276, figs. 157-159.

Brimley 1938:503. Schultz 1961:194; 1962:47; 1964:194; 1965b:139.

This species is common in eastern United States and is very com-

mon in North Carolina (Map 12). It inhabits the driest habitats of all

species in the state and is found in a very wide range of dry-moist habi-

tats. It is very abundant around the foundations of buildings, in dense

leaf litter, and under boards and rocks, but rarely occurs far from

human habitations. The species has been recorded on "floating sea-

weed" on the coast of South Carolina (Kelley 1978), but that reference

is probably to a species of Tylos, a genus of isopods from almost com-

pletely marine habitats which also rolls into a ball and in many ways

superficially resembles large females of A. vulgare (Van Name 1936:408).

It can be distinguished from another "pill bug,'M. nasatum, by the lack

of a "shelf on the frontal margin of the head (Cf. Figs. 30, 31). Adult

males are ebony black with small yellow blotches. Females and juvenile

males are variegated yellow-brown. In the Durham region the species

breeds from the end of May until late July. It grows to 14 mm long.

DISCUSSION

The oniscoids of North Carolina can be divided into several groups

according to habitat preference. The first group contains three species

that live exclusively at or very near the sea and never are found far from

a marine shore. They are Ligia exotica, Armadilloniscus ellipticus and

Philoscia vittata. Ligia exotica spends much time in sea water and

frequently retreats there when disturbed. The habitats of Armadillonis-

cus ellipticus and P. vittata are on salt or brackish water shores and

occasionally habitats and inhabitants might be covered with salt water.

The two species are present up to the point of furthest penetration of

North Carolina Terrestrial Isopods 23

marine waters into the land. They are quite common in such places as

the maritime drift line, the high or storm tide line of dead vegetation

and flotsam.

Other species living at the North Carolina seashore are not exclu-

sive inhabitants there. A species such as Porcellio virgatus rarely lives at

the shoreline proper, but in leaf litter of nearby upland deciduous vege-

tation like the live oak forest of the Coastal Plain from North Carolina

to Mexico. It probably is less tolerant of brackish conditions than either

Miktoniscus halophilus or Porcellio scaber, which frequently are encoun-

tered on the shore itself and in reed grass litter of brackish swamps.

Occasionally they are encountered in the maritime drift line. However,

both species also live in upland habitats such as rotting logs and leaf

litter in the deciduous forests of river bottoms. Porcellio laevis has been

found under refuse on the sandy upper beaches of North Carolina, but

lives mainly around the bases of buildings and in refuse heaps in upland

locations.

Most species of terrestrial isopods in North Carolina must be

included with the upland species. In general they are distributed accord-

ing to the moisture content of their habitats, which range from wet to

dry, and the moisture content of the habitat is determined by rainfall

and evaporation. They all live in decayed organic vegetation such as leaf

litter or decomposed parts of grasses, shrubs and deciduous trees. Iso-

pods are only rarely encountered in evergreen logs or evergreen leaf

litter. The numbers and types of moist retreats available when condi-

tions become seasonally warm and dry is critical to the local distribu-

tion of terrestrial isopods. Seasonal temperature extremes probably

have the greatest influence on regional distribution of particular species

in temperate locations such as North Carolina. The types and amounts

of organic matter also probably help determine if a particular species

occupies a particular habitat. No real measure of moisture or organic

content can be easily made, but certain subjective and qualitative

determinations are obvious from simple observation. A swamp is wet; a

shaded retreat in a refuse heap probably changes very little during the

day and only slightly during the season: a pulpy, rotted, moist log in a

shaded place provides a fairly constant environment the year around;

and organic matter such as leaf litter, dry to the touch and in the sun at

the side of a building, has a very low moisture content. The upland

species are briefly discussed here in order of habitat moisture content,

from wet to dry.

The species of Ligidium need the most moisture and are present

only in habitats that are very wet with fresh water throughout the year.

They live at the edge of swamps and along the margins of permanent,

shaded streams. The detailed habitat preferences of each species of the

genus remains to be discovered, but in general L. bluer idgensis occurs in

24 George A. Schultz

the cool mountains and L. elrodii in the swamps of the Piedmont Pla-

teau and Coastal Plain. Altitude and temperature also probably affect

regional distribution of the species of the genus.

The five species of Trichoniscidae are small to tiny in size. They

require sheltered habitats where moisture is more or less uniform

throughout the year. Such habitats are found in dense leaf litter and in

well decayed logs of wooded areas. Miktoniscus halophilus is wide-

spread in eastern North Carolina, being abundant in leaf litter and

decayed logs of river bottoms in upland locations. As the leaf litter dries

in summer the species becomes more restricted to the moist parts of its

habitat. Miktoniscus medcofi generally occurs in habitats similar to

those of M. halophilus (see Schultz 1976), but in North Carolina it has

been collected only in a cave in the western part of the state. Thus, its

preferred habitat cannot now be discussed. Like many oniscoids it can

easily live in caves where temperature and moisture are relatively con-

stant. So far no species modified for cave life has been collected in

North Carolina.

Hyloniscus riparius in North Carolina occurs in habitats similar to

those of Miktoniscus halophilus, but generalizations about its distribu-

tion cannot be made because it was recorded only once, near Durham.

Trichoniscus pusillus has also been recorded only once in the state, but

lives with H. riparius in New Jersey (Schultz 1965b) and might do so in

North Carolina. Haplophthalmus danicus is apparently present in many

moist locations, but is especially abundant in pulpy, shaded logs. A syc-

amore log in an advanced stage of decay was one such habitat in the

Durham region. Organic content of habitat probably plays an impor-

tant role in the distribution of H. danicus.

The other species are the more commonly encountered "woodlice"

or "pill bugs," which also live in a variety of habitats. Cylisticus con-

vexus was found in places that seemed to provide moist retreats most of

the year, although the species occasionally was seen wandering in rela-

tively dry places. It was especially abundant where human refuse — old

furniture, boards, and other non-food trash — was present. It occurred

frequently in decayed logs that had been invaded and later abandoned

by carpenter ants. Porcellio scaber was present in leaf litter at the bases

of trees in parks and near building at many locations. It frequently

shared habitat with Porcellionides pruinosus. They never were as

abundant as the other large species, but their presence was always pre-

dictable. Porcellio laevis was occasionally encountered in the same habi-

tats, but was most abundant where the habitat was quite moist or where

human refuse was present. Oniscus asellus was recorded only once, so

its habitat preference in North Carolina is not well known.

Armadillidium nasatum occurred in the widest range of wet-dry

habitats, although it was not present in habitats as dry as those of A.

North Carolina Terrestrial Isopods 25

vulgare. At one site A. nasatum was present in abundance under the

muddy vegetation of a rock near which Ligidium elrodii was common.

Several thousand A. nasatum occasionally were encountered under

refuse and dense vegetation, places where apparently they had aggre-

gated to spend the winter. However, the species was widespread in any

leaf litter and was frequently collected with all of the larger species.

Armadillidium vulgare is the most tolerant of dry conditions of any

species in the state. Leaf litter at the base of a house during summer was

one particularly dry habitat where about 100 specimens of several sizes

of the species were collected. It was not encountered in the very moist

habitats where A. nasatum was occasionally found. Both species of

Armadillidium frequently were together in habitats of intermediate

moisture, and the large aggregations of A. nasatum often contained

small numbers of A. vulgare.

At night or on moist, cloudy days when isopods are generally most

active, any one of the larger, more common species could be found far

from the typical habitat. Although only a few were ever seen at any

time, the movements during moist times probably account for the

spread of such species throughout North Carolina and the southeastern

states.

ACKNOWLEDGMENTS.— The author would like to thank Dr.

Rowland M. Shelley, North Carolina State Museum of Natural His-

tory, Raleigh, for providing many of the specimens on which much of

this work was based. He would also like to thank Dr. John E. Cooper

of the State Museum for his help with the manuscript. Thanks also is

due to Harold and Norma Feinberg for the contribution of several

important collections from western North Carolina.

LITERATURE CITED

Brimley, C. S. 1938. The Insects of North Carolina. N.C. Dep. Agric. Div.

Entomol., Raleigh. 560 pp.

Cooper, John E., and M. R. Cooper. 1977. Miktoniscus alabamensis Muchmore.

Small Alabama Sowbug. pp. 21 1-212 in J. E. Cooper, S. S. Robinson and

J. B. Funderburg (eds.). Endangered and Threatened Plants and Animals

of North Carolina. N.C. State Mus. Nat. Hist., Raleigh, xvi + 444 pp.

Harger, Oscar. 1878. Descriptions of new genera and species of Isopoda, from

New England and adjacent regions. Am. J. Sci. Arts (3)75:373-379.

Kelley, B. J. Jr. 1978. Order Isopoda. pp. 167-170 in Zigmark, R. J. (ed.). An

annotated checklist of the biota of the coastal zone of South Carolina.

Univ. South Carolina Press, Columbia, xii + 364 pp.

Richardson, Harriet. 1905. Monograph on the isopods of North America. Bull.

U.S. Natl. Mus. 54:\\\-121.

Schultz, George A. 1961. Distribution and establishment of a land isopod in

North America. Syst. Zool. 10(4): 193-196.

26 George A. Schultz

1962. Miktoniscus grayi, a new species of terrestrial isopod from

North Carolina. J. Elisha Mitchell Sci. Soc. 7S(1):47-51.

1963. Philoscia robusta, a new species of terrestrial isopod crusta-

cean from southeastern United States. J. Elisha Mitchell Sci. Soc.

79( l):26-29.

1964. Two additional data on terrestrial isopod crustaceans: Lig-

idium blueridgensis, sp. no v., from Georgia and a North Carolina cave

location for Miktoniscus linearis (Patience, 1908). J. Elisha Mitchell Sci.

Soc. <S0(2):9O-94.

1965a. The reduction of Philoscia vittata Say, 1818, to a synonym

of Philoscia muscorum (Scopoli, 1763). Crustaceana 8(1): 107-108.

1965b. The distribution and general biology of Hyloniscus riparius

(Koch) (Isopoda, Oniscoidea) in North America. Crustaceana 5(2): 131-140.

1970. Descriptions of new subspecies of Ligidium elrodii (Packard)

comb. nov. with notes on other isopod crustaceans from caves in North

America (Oniscoidea). Am. Midi. Nat. 84(\):2>6-45.

1974. The status of the terrestrial isopod crustaceans Philoscia mus-

corum, P. vittata, P. robusta and P. miamiensis in the New World (Onis-

coidea, Philosciidae). Crustaceana 27(2): 147-153.

1975. Terrestrial isopod crustaceans (Oniscoidea) from coastal sites

in Georgia. Bull. Ga. Acad. Sci. 34(4): 185-194.

1976. Miktoniscus halophilus Blake, M. medcofi (Van Name) and

M. morganensis n. comb., reconsidered with notes on New World species

of the genus (Crustacea, Isopoda, Trichoniscidae). Am. Midi. Nat.

95(1):28-41.

1977. Terrestrial isopod crustaceans (Oniscoidea) from St. Cather-

ines Island, Georgia. Ga. J. Sci. 35:151-158.

Stephenson, T. A., and A. Stephenson. 1952. Life between the tide-marks in

North America. II. North Florida and the Carolinas. J. Ecol. 40:1-49.

Van Name, Willard G. 1936. The American land and fresh-water isopod Crus-

tacea. Bull. Am. Mus. Nat. Hist. 77:1-535.

1940. Supplement to American isopod Crustacea. Bull. Am. Mus.

Nat. Hist. 77:109-142.

1942. A second supplement to the American land and fresh-water

isopod Crustacea. Bull. Am. Mus. Nat. Hist. 50:299-329.

Accepted 10 December 1982

Aquatic Macroinvertebrates

of the

Upper French Broad River Basin

David L. Penrose, David R. Lenat and K. W. Eagleson

North Carolina Department of Natural Resources

and Community Development,

P. O. Box 27687, Raleigh, North Carolina 27611

ABSTRACT. — The aquatic macroinvertebrates of the Upper French

Broad River were sampled over a two-year period beginning in May

1977. The information gathered in this study, along with additional

data for other select tributary streams, were combined to generate a

list of 267 invertebrate taxa. The faunas of the French Broad River

and tributaries are compared to those of other rivers and streams of

the southern Appalachians to define "normal" faunal characteristics of

lotic systems in this geographic area. Information is presented on taxa

richness and abundance for the six major taxonomic groups. Temporal

and spatial changes in the benthic macroinvertebrate communities

were associated with changes in flow rates, watershed sizes, average

temperatures, canopies, substrate characteristics, and gradients.

INTRODUCTION

A chronicle of the French Broad River by Dykeman (1955) referred

to the river as "the classic example of an Appalachian River". Dykeman

presented the French Broad as a study of paradoxes: allure and despair,

problems and promises. These paradoxes are especially evident today as

we consider the inevitable growth and development of the upper French

Broad River basin in North Carolina, balanced against the assimilative

capacity of the river. Can the river continue to absorb perturbations and

remain a valuable natural resource?

Water quality records for the French Broad River in Transylvania

County are of little help in answering this question. A water quality

monitoring site is located in Rosman, North Carolina, and is main-

tained by the Division of Environmental Management. The water qual-

ity is described as very good, with few stream contraventions recorded

(N.C. Department of Natural Resources and Community Development

1976). This site is also monitored by the United States Geological Sur-

vey, and continuous stream flow records have been collected since 1955.

The average stream discharge at Rosman, located just below the conflu-

ence of the North and West Forks, is 242 cfs (6.85 m3/sec) and the

average rainfall there is 122.9 cm/ year (United States Geological Survey

1979). Average flow of the river during the period of the study herein

reported can be seen in Figure 1. The watershed above Rosman is

Brimleyana No. 8:27-50. December 1982 27

28

David L. Penrose, David R. Lenat, K. W. Eagleson

FLOOD

M J S N

1977

M M J S

1978

M M

1979

MONTH/YEAR

Fig. 1. Average flow of the French Broad River at Rosman, North Carolina,

May 1977-July 1979.

approximately 104 km2, generally ranges from 610 to 1524 m elevation,

and consists primarily of forest and farm land. A cursory inspection of

topographic maps reveals that much of the watershed lies in the Pisgah

National Forest. The East Fork of the French Broad joins the mainstem

below Rosman.

This paper describes the composition of the aquatic macroinverte-

brate fauna of the Upper French Broad River (Transylvania County)

and selected tributaries. This fauna is then compared to that of several

other Appalachian streams and rivers to define "normal" faunal charac-

teristics of lotic systems in the geographic area. Water quality is depend-

ent upon watershed characteristics (Lotspeich 1980), and the nature of

the aquatic community is shown to be similarly dependent upon these

characteristics. Therefore, faunal characteristics can be used as baseline

information, along with water quality records, to describe the current

environmental quality of the watershed and indicate how this may

change with growth and development.

French Broad River Macroinvertebrates

29

MATERIALS AND METHODS

Monitoring Stations

Biological monitoring stations were established along an 1 1 km sec-

tion of the French Broad River in Transylvania County (Fig. 2). Two

stations were selected above Rosman to serve as examples of "Upper

River-Montane" (UR) stations, areas of nearly complete canopy cover

and higher gradient. Below Rosman the river begins to meander through

a fairly large floodplain. Two stations were selected in this area to serve

as examples of "Upper River-Lowland" (LR) stations, areas with little

canopy cover and lower gradient. The two upper river stations are

French Broad 1 and 2. French Broad 1 (FB-1) was located on the West

Fork above the US 64 bridge, while French Broad 2 (FB-2) was located

on the North Fork at the bridge over state rural road (SRR) 1322. The

two lower river stations are French Broad 3 and 4; FB-3 was located at

SRR 1129 and FB-4 at SRR 1331.

■*c

S*.

f&

kc*

&\

178V

§**

pon(i

EAST FORK

FRENCH BROAD RIVER

1Km.

1mile

H

Fig. 2. Biological monitoring stations: French Broad River, Morgan Mi]

Creek, and Cherryfield Creek.

30 David L. Penrose, David R. Lenat, K. W. Eagleson

In addition to the four river stations, five tributary stations were

located on Cherryfield and Morgan Mill creeks, which are second order

streams. Two stations were established on Cherryfield Creek: a control

0.1 km above US 64 along SRR 1332, and a downstream station 1.0 km

below US 64 along SRR 1331. Three stations were located on Mor-

gan Mill Creek: a control 0.6 km above US 64 along SRR 1331, another

immediately below US 64, and a third at the SRR 1331 bridge. Cherry-

field and Morgan Mill creeks converge before flowing into the French

Broad near river mile 212. The Cherryfield Creek watershed is 7.04 km2

while the Morgan Mill Creek watershed is 2.93 km2.

Collection Methods

Samples were collected using the "kick net" method. The net was

positioned upright on the streambed while an upstream area of approx-

imately 1 m2 was physically disrupted. This technique is less quantitative

than others but may be used across a wide range of habitats. White and

Fox (1980) found that the kick technique was superior to other methods

by providing the largest number of species per unit time. The kick net

was constructed of window screening with 1 mm openings; smaller mesh

is available, but has a tendency to clog with organic debris and silt,

causing some flow to be diverted around the net. We had success col-

lecting even very small Chironomidae using this method. The samples

were then rinsed into a wash bucket (brass wire cloth mesh No. 30),

placed in quart freezer containers, and preserved in ethanol. Two sam-

ples were collected from each stream station and three from each river

station.

Little time could be devoted to qualitative collections (pools, bank

sweeps, and others), as this study was being conducted to determine the

probable effects of road construction on stream and river fauna of this

geographic area. Therefore, samples were collected from habitats that

normally would produce the most diverse fauna. Ten collections were

made from the river stations over a twenty-five month period (June

1977-July 1979). Nine collections were made from the tributary stations

over a twenty-two month period (February 1978-December 1979). Col-

lection dates and locations were chosen to coincide with construction

activities.

A list of taxonomic references used in our identifications is available

from the authors.

RESULTS AND DISCUSSION

Taxa Richness: Rivers

Total taxa richness from the four French Broad River stations was

221. Five other surveys conducted on Appalachian mountain river sys-

tems were selected for comparison. They were conducted above most

French Broad River Macroinvertebrates

31

potential pollution sources and were therefore assumed to be relatively

undisturbed enough to serve as controls. None produced as many total

taxa as our study, and the lower taxa richness (Table 1) is attributable

to less frequent sampling and/ or less precise taxonomy. Collection

methodology was designed, in each survey, to produce the most diverse

fauna.

Table 2 presents average taxa richness (per collection) for six refer-

ence sites on Appalachian mountain rivers. The level of taxonomic reso-

lution accounts for at least some variability between surveys. For exam-

ple, Starnes (1976) identified chironomids only to family in her survey

of the Little Pigeon River, while in our survey we identified 74 taxa

Table 1. Total taxa richness (S) comparisons for six Appalachian river studies.

S # Collections # Station Method Reference

4 Kick net This paper

2 Kick net Lenat et al.

(1979)

2 Kick net Lenat et al.

(1979)

4 Assorted Harned (1977)

1 Kick net Seagle & Hen-

dricks (1978)

1 Surber Starnes (1976)

1 Stations FB-1 and FB-2, upper river-montane only.

Table 2. Average taxa richness comparisons by taxonomic group for four

Appalachian river studies. Abbreviations as in Table I.

Station

Order FB-1 FB-2 MR-1 MR-2 NT-1 LPR

32 David L. Penrose, David R. Lenat, K. W. Eagleson

within the family Chironomidae. Because several of the surveys listed in

Table 1 were conducted by our group, reduction in taxa richness

between these surveys cannot be accounted for by differences in collec-

tion and taxonomic techniques. For example, the low average taxa

richness in the Trichoptera for the North Toe River cannot be accounted

for taxonomically.

Abundance (percent of total abundance): Rivers

Table 3 summarizes average community composition by taxonomic

order (as percent of total abundance) for several Appalachian mountain

control stations. Differences within watersheds are small for both the

French Broad and Mills rivers. However, large differences exist between

watersheds, and much of the variability can be attributed to watershed

characteristics.

Average annual water temperature is higher for the French Broad

(12.8° C) than for the Mills River (11.3° C) (N.C. Department of Natu-

ral Resources and Community Development 1977). The cooler water

temperature of the Mills River might favor the development of the Plec-

optera community observed there (Baumann 1979). Also, several of the

dominant Plecoptera genera in the Mills River are chironomid preda-

tors, which may account for the low percentage of Diptera in this area.

Average annual water temperature of the North Toe River also is high

(14.0° C) and might limit Plecoptera.

The highest percent abundance for Trichoptera was noted for the

North Toe River, and is related to watershed size. The North Toe

watershed (193 km2) is substantially larger than either the French Broad

(97 km2) or Mills River (43 km2) watersheds. Larger drainage areas

result in increased flow (Edington 1968) and increased amount of fine

particulate organic matter (Wallace 1975). These variables influence the

distribution of hydropsychid Trichoptera (Gordon and Wallace 1975)

because of the net building and filter feeding habits of this group. The

North Toe River at the sampling site is dominated by hydropsychid

Table 3. Average percent community composition comparisons by taxonomic

order for four Appalachian mountain river studies. Abbreviations as in

Table 1.

Order FB-1 FB-2 MR-1 MR-2 NT-1 LPR

French Broad River Macroinvertebrates

33

caddisflies, as are many other riverine systems with large watersheds

(Ross 1944; Rhame and Stewart 1976). Anderson (1976) said that

hydropsychids are "particularly" characteristic of large rivers.

Taxa Richness: Streams

The total number of taxa for the two French Broad tributaries (5

stations) was 199 (Table 4). Surveys conducted by the Division of

Environmental Management in the seven other Appalachian streams

shown in Table 4 did not record as many taxa, but this may simply

reflect the far fewer samples collected. It has long been known that taxa

richness is a logarithmic function of collection area (Gleason 1922). A

larger number of samples increases collection efficiency, hence produces

greater taxa richness.

Average taxa richness per collection for the taxonomic orders is

presented in Table 5. This table also includes information from the

seven other mountain stream surveys listed in Table 4.

Abundance (percent of total abundance): Streams

Table 6 gives abundance (as a percent of total abundance) for tax-

onomic orders. As before, we include comparisons from other presuma-

bly unstressed Appalachian streams. We have also been able to include

data from Tebo and Hassler (1961), Talak (1977), and Dysart et al.

34

David L. Penrose, David R. Lenat, K. W. Eagleson

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French Broad River Macroinvertebrates 35

(1973), because tabulation of percent abundance of major taxonomic

groups requires identification only to the level of order.

Many differences in abundance can be related to watershed size

and stream gradient. These factors were grouped and included in Table

6 when available. High percentages of Plecoptera (> 20%) were observed

only in streams with very small watersheds (SW-1 and BR-1). This phe-

nomenon was not evident at BC-1 (tributary to Beech Creek), a site that

was not sampled in the fall when most Plecoptera would probably be

present. In these very small first order streams, low flow rates allow

large accumulations of leaves. Such accumulations ("leaf packs") are the

preferred habitat of "shredder" Plecoptera genera, especially Allocap-

nia, Pel toper la and Taeniopteryx.

High gradient streams with watersheds of from 2 to 4 km2 (PC-1,

BL-1, and Cox-1) were strongly dominated by Ephemeroptera, espe-

cially Epeorus {Iron) and Rithrogenia. These streams are characterized

by many small waterfalls. Water velocity is sufficient to quickly remove

leaf accumulations and the substrate often has a very "clean" appear-

ance. We suspect that these watershed characteristics were also true for

those studied by Talak (1977).

Larger streams (CF-1, TM-1, DR-1; and Tebo and Hassler 1961)

were dominated by a combination of Ephemeroptera and Trichoptera.

These streams are characterized by somewhat less canopy cover, greater

amounts of fine particulate organic matter, and higher flow. The com-

position of the benthic fauna for this group of streams is similar to that

described for mountain rivers in Table 3.

Seasonal Trends

Abundance: River Stations. — Abundance values showed large

between-year and between-station variability (Fig. 3). However, several

distinct trends may be observed.

Organism abundance was usually greatest at station 4 and can be

correlated with substrate characteristics. Station 4 is located in an area

of heavy Podostemon (riverweed) growth. According to Anderson and

Sedell (1979), macrophytes may increase the density of benthic

macroinvertebrates although not serving as food. Macrophytes structur-

ally modify the river bed by increasing the available benthic area and

reducing flow. A reduction in flow in turn results in both less scour and

greater retention of detritus. This hypothesis is supported by the con-

vergence of abundance values (Fig. 3) at all stations during winter and

early spring, a period of Podostemon dieback.

All stations appeared to have a "refuge" from scouring. Following

the 1979 spring flood (Fig. 1), the number of benthic invertebrates did

not appear much lower than those observed in the spring of 1978, de-

spite a lower 1978 flow. Extremely high flows appeared to have minimal

36

David L. Penrose, David R. Lenat, K. W. Eagleson

MONTH/YEAR

Fig. 3. Average number of organisms per sample, French Broad River, June

1977-July 1979.

effect on benthic fauna in the French Broad River.

A sharp increase in the number of benthic organisms was asso-

ciated with low discharge rates, especially in the fall of 1978 (Fig. 1).

However, a decline in the number of organisms at station 2 was

observed in October 1978. This decline may have been caused by activi-

ties in the watershed of the North Fork. We learned that a fish kill

occurred just above station 2 in late July 1978. According to the Ashe-

ville Citizen (1978), 1,200 fish (mostly brown trout) were killed in John-

nies Creek, Tucker Creek, and parts of the North Fork of the French

Broad. The suspected cause of the kill was a pesticide in the Johnnies

Creek watershed. Passage of a pesticide (as a "slug dose") through sta-

tion 2 could have caused the fluctuations in density observed there. Plec-

optera, which are highly sensitive to a variety of pesticides (Jensen and

Gaufin 1964; Courtemanch and Gibbs 1977; Eidt 1975), were not col-

lected from this area in September 1978.

Taxa Richness: River Stations. — Only 1978 and 1979 data are pres-

ented in Figure 4, because pre- 1978 collections were identified mostly to

genus. Several trends may be observed. Taxa richness at station 1 was

remarkably constant, particularly for the last six sample periods. This

pattern suggests that total taxa richness in this area is independent of

French Broad River Macroinvertebrates

37

Fig

MONTH/YEAR

4. Total taxa richness, French Broad River, February 1978-June 1979.

both season and discharge rate. However, total taxa richness at other

stations showed considerable seasonal fluctuation. This fluctuation is

not random; two distinct patterns may be observed. First, taxa richness

values at these other stations were roughly parallel, suggesting that the

same factor (above station 2) controlled the benthos at all stations.

Second, reductions in taxa richness occurred primarily during the spring

and/ or summer period. Taxa richness values at all stations tended to

converge in winter and fall.

Abundance: Stream Stations. — Although five stream stations were

sampled, data are presented only for the "control" stations of Morgan

Mill (MM-1) and Cherryfield (CF-1) creeks (Fig. 5). There appear to be

two peaks (spring and fall) per year. This is an expected pattern for

temperate streams (Hynes 1972). Organism abundance was greater at

station MM-1 than that observed at station CF-1, but seasonal trends

were parallel for these two areas.

Taxa Richness: Stream Stations. — Taxa richness values suggest lit-

tle seasonal variation, although values tended to be slightly higher in

winter and early spring (Fig. 6). A marked divergence between stations

MM-1 and CF-1 was observed only during the period October 1978

through January 1979. This is the same period when the fish kill

occurred above station 2 on the French Broad.

38

David L. Penrose, David R. Lenat, K. W. Eagleson

MONTH/YEAR

Fig. 5. Average number of organisms per sample, Cherryfield (CF-l) and

Morgan Mill (MM-l) creeks, February 1978-December 1979.

Species (or taxa) Level Information

Table 7 lists all benthic macroinvertebrates collected in the Upper

French Broad River Basin, and includes 267 taxa, derived from identifi-

cation of 81,000 organisms. Data on frequency and seasonal distribu-

tion are included. Frequency data are based on the number of times

each taxon was collected rather than on any quantitative measure of

abundance.

Table 7 also lists the "months of maximum abundance." Since this

is a highly qualitative procedure, the analysis was very cautious. When

no clear seasonal pattern was apparent this column was left blank. It

was also left blank for all rare taxa.

Life History Data. — Many taxa were found to have two or three

distinct seasonal peaks in abundance. In many cases, these peaks may

correspond to separate generations. Multiple peaks in abundance were

recorded for Ephemeroptera (12 taxa), Trichoptera (4 taxa) and Diptera

(15 taxa). Most of these taxa are grazers or filter-feeders. Since this

group of organisms included many classified as "abundant", a large

proportion of the benthos would be expected to have two or three gen-

erations per year.

French Broad River Macroinvertebrates

39

80

70

60

50

40

30

20

10

MM-1

^CF-1

III!

Ill III ( I I I I

I t I \_L

J M M J S N

1978

J M M J S N

1979

MONTH/YEAR

Fig. 6. Total taxa richness per collection, Cherryfield (CF-1) and Morgan Mill

(MM-1) creeks, February 1978-December 1979.

The growth rate of many benthic species varies geographically,

principally due to differences in the temperature regime (Vannote and

Sweeney 1980). Water temperature will affect either the number of gen-

erations per year or the reproductive success of polyvoltine species.

Neves (1979) studied the benthic macroinvertebrates of a Massachusetts

stream and found that 14% of the standing crop was based on species

with two to four generations per year. These polyvoltine species com-

prised 32% of the total production. Neves' study was conducted in a

stream with a mean annual temperature of 8.3° C. In the French Broad

River, higher mean temperatures (about 13° C) might be expected to

increase the number of polyvoltine species. We found that taxa with two

to three peaks per year comprised 44% of the standing crop at stream

stations and 55% of the standing crop at river stations (unpublished

data). This implies that polyvoltine species make up a large proportion

of the French Broad River fauna. Production studies in the southeast-

ern United States must be careful to evaluate this factor.

For species with similar morphology and feeding type, differences

in seasonal distribution may help to avoid competition. Many such spe-

cies pairs can be seen in Table 7. A few examples include: Epeorus

40 David L. Penrose, David R. Lenat, K. W. Eagleson

Table 7. List of organisms recorded from the French Broad River and tributar-

ies, showing relative abundance in each major area and months of

maximum abundance. S = streams, UR = upper river-montane, UL =

upper river lowland. R = Rare; present in 1-5% of samples. C = Com-

mon; present in 6-25% of samples. A = Abundant; present in at least

26% of samples. + = identification to species only during last collection.

Frequency Month(s) of max-

imum abundance

Taxon S UR LR

Originally identified to genus level, species later identified from selected sam-

ples using Morihara and McCafferty (1979).

Epeorus sp. 2 was distinguished by difference in coloration and the presence of

spatulate claws.

Early instars of other species may have been lumped with S. modestum.

Some taxonomists now consider previous Ephemerella subgenera to have

generic status.

French Broad River Macroinvertebrates

41

5 Probably includes E. dorothea.

These genera misidentified in early samples and confused with Hastaperla.

42 David L. Penrose, David R. Lenat, K. W. Eagleson

ODONATA

Cordulegaster sp.

Lanthus parvulus

Stylurus scudderi

Boyeria vinosa

Aeschna sp.

Calopteryx sp.

COLEOPTERA

Gyrinus sp.

Dineutes sp.

Bidessus sp.

Hydroporus spp.

Ectopria nervosa

Psephenus herricki

Macronychus glabratus

Optioservus sp.

Oulimnius latiusculus

Promoresia elegans

P. tardella

Stenelmis sp.

Anchytarsus bicolor

TRICHOPTERA

Brachycentrus sp.

Micrasema wataga

Agape t us sp.

Glossosoma nigrior

Matrioptila jeanae

Helicopsyche borealis

Arctopsyche irrorata

Cheumatopsyche spp.

Diplectrona modesta

Hydropsyche betteni

H. demora

H. mississippiensis

Symphitopsyche bronta

S. macleodi

S. morosa

S. sparna

Parapsyche cordis

Ochrotrichia sp.

Lepidostoma sp.

Oecetis spp.

Set odes sp.

Goer a sp.

Hydatophylax argus

Neophylax spp.

Pseudostenophylax

uniformis

Pychopsyche guttifer

P. lepida

Doliphilodes sp.

Wormaldia sp.

French Broad River Macroinvertebrates 43

7 Paper in preparation by John Morse, Clemson University, and co-workers

indicates this genus includes Cernotina and Plectrocnemia.

8 Primarily N. fasciatus, but N. serricornis also present.

44

David L. Penrose, David R. Lenat, K. W. Eagleson

9 In-house keys for the Cricotopus j Or thocladius group (C/O), and reports from

the Biological Monitoring Unit, Division of Environmental Management, use

a number for various species and/ or species groups. This number is included

for comparison with other DEM reports.

French Broad River Macroinvertebrates 45

D-

Euorthocladius and other subgenera of Orthocladius are poorly known; this

group may contain several (probably undescribed) subgenera.

Follows William Bode's unpublished key to species groups.

46 David L. Penrose, David R. Lenat, K. W. Eagleson

Nais behningi R - -

N. elinguis R R R

N. simplex R R -

Pristina idrensis R - R

P. longiseta (?) R

Specaria josinae R - -

CRUSTACEA

Lirceus sp. R

Crangonyx sp. R

Cambarus spp. C R -

MOLLUSCA

Goniobasis sp. A C

Gyraulus sp. R

Ferrissia sp. C A C

Sphaerium sp. R R

Pisidium sp. - - R

"OTHER"

Nematoda - R -

Hydracarina R R -

Dugesia tigrina R - -

Cura foremanii R - R

Prostoma graecens - R C

(Iron) and Stenonema ithaca; Promoresia elegans and P. tardella; Mic-

rasema wataga and Brachycentrus; and Prosimulium mixtum and Simu-

lium spp..

Spatial Patterns. — Table 8 shows the number of taxa (by group)

with a preference for each area of the river basin. Data are not tabu-

lated for unusual taxa. Within this group, 86 taxa (64%) showed some

spatial preference.

Longitudinal zonation along lotic systems has been demonstrated

many times in rivers throughout the world, and Hynes (1972) listed over

100 examples. Our intent here is simply to illustrate the expected pat-

tern in a small southeastern river.

Comments on Particular Taxa. — The French Broad River data set

contained many unusual collection records, especially for Ephemero-

ptera and Chironomidae. An extremely high diversity was found within

the genus Ephemerella (24 species). These species often had a slight

degree of spatial and temporal overlap; single river stations in the spring

of 1979 had up to 20 species of Ephemerella.

The "new genus near Centroptilum" is a highly unusual mayfly,

characterized by long gills on the fore coxae. The morphology of this

species will be described by Carlson and Lenat (in prep.). Brachycercus

nitidis is infrequently collected in routine stream collections (Berner

1977). We found this species only in samples taken near the river bank.

Baetisca berneri was recently described from West Virginia by Tarter

French Broad River Macroinvertebrates 47

Table 8. Number of taxa by group, showing spatial preferences for second

order streams (S), Upper River-Montane (UR), Upper River-Lowland

(LR), streams and Upper River-Montane (S-UR) and Upper River-

Montane and Upper River-Lowland (UR-LR).

# with spatial preference

Total Taxa S UR LR S-UR UR-LR

and Kirchner (1978). Several specimens were collected at our stream

stations.

Many stream/ river surveys neglect the Chironomidae, citing taxo-

nomic difficulties. However, recent work in this important group has

made identifications much easier. Mozley (1980) provided a generic key

to North Carolina Orthocladiinae that should further reduce the diffi-

culty of identifications within this subfamily. We identified 74 taxa

within the Chironomidae, which probably included more than 85 spe-

cies. This number of taxa agrees well with many other intensive surveys

of lotic chironomids (see data cited by Lindegaard-Peterson 1972).

However, many more species might be expected with more precise tax-

onomy (especially if pupae and adults are used). Coffman (1973) identi-

fied 143 chironomids from a single Pennsylvania stream, and Lehmann

(1971) recorded 245 species from the Fulda River. The latter study

included data from many stations along a 220-km river segment.

The chironomid fauna of the upper French Broad River is domi-

nated by the Orthocladiinae and Diamesinae, a trend expected for cool

lotic systems (Oliver 1971). The Tanytarsini were also important, espe-

cially Micropsectra (stream stations) and Rheotanytarsus (river sta-

tions). Several unusual Diamesinae were collected, especially Pagastia

sp. These specimens were verified by D. T. Oliver (pers. comm.), who

stated they were "similar to an undescribed species near P. orthogonia"

Most recent chironomid keys omit this genus, but it is illustrated in

Mason (1973) under the name Pseudodiamesa pertinax.

48 David L. Penrose, David R. Lenat, K. W. Eagleson

The Cricotopus/ Orthocladius group contained 15 taxa, many of

which are species groups. The " Euorthocladius" group (which may con-

tain several subgenera) is particularly in need of taxonomic revision.

ACKNOWLEDGMENTS.- We would like to acknowledge the

capable assistance of several individuals, particularly Kay Dechant,

Ross Green, Barbara Burchard, and Jeanne Whittlesey, who were

responsible for sample collection and processing. Taxonomic assistance

was received from Paul Carlson (Ephemeroptera), Jerry Louten (Odo-

nata), John Morse (Trichoptera), and many chironomid specialists: Sam

Mozley, Pat Hudson, Jan Doughman, Len Femmington, Dave Smith,

M. W. Boesel, and Robert Bode. Three anonymous reviewers provided

helpful comments on the manuscript.

Biological monitoring activities in North Carolina are supported by

a grant from the U.S. Environmental Protection Agency.

LITERATURE CITED

Anderson, Norman H. 1976. The distribution and biology of the Oregon Tri-

choptera. Oreg. Aric. Exp. Stn. Tech. Bull. No 134. 152 pp.

, and J.R. Sedell. 1979. Detritus processing by macroinvertebrates in

stream ecosystems. Annu. Rev. Entomol. 24:351-377.

Asheville Citizen. 1978. Large fish kill reported. July 26.

Baumann, Richard W. 1979. Nearctic stonefly genera as indicators of eco-

logical parameters (Plecoptera: Insecta). Great Basin Nat. 59:241-244.

Berner, Lewis. 1977. Distributional patterns of southeastern mayflies (Ephem-

eroptera). Bull. Fla. State Mus. Biol. Sci. 22:1-55.

Courtemanch, David, and K.E. Gibbs. 1977. The effects of one and two

year applications of sevin-4-oil on non-target stream invertebrates. Unpubl.

report, Dep. Entomol., Univ. Maine, Orono. 17 pp.

Coffman, Wiliam P. 1973. Energy flow in a woodland stream ecosystem: II.

The taxonomic composition and phenology of the chironomidae as deter-

mined by the collection of pupal exuviae. Arch. Hydrobiol. 77:281-322.

Dykeman, Wilma. 1955. The French Broad. Univ. Tenn. Press, Knoxville.

371 pp.

Dysart, Benjamin C, A.R. Abernathy, H.J. Gover, R.D. Hatcher and B.R.

Ingram. 1973. Bad Creek Environmental Study. Report to Duke Power

Company. 217 pp.

Edington, J.M. 1968. Habitat preferences in net-spinning caddis larvae with

special reference to the influence of water velocity. J. Anim. Ecol. 57:675-692.

Eidt, D.C. 1975. The effects of fenitrothion from large-scale forest spraying on

benthos in New Brunswick headwater streams. Can. Entomol. 707:743-760.

Gleason, Henry A. 1922. On the relation between species and area. Ecology

5:158-162.

Gordon, A. Elizabeth, and J.B. Wallace. 1975. Distribution of the family

Hydropsychidae (Trichoptera) in the Savannah River Basin of North Caro-

lina, South Carolina and Georgia. Hydrobiologia 45:405-423.

French Broad River Macroinvertebrates 49

Harned, W. Douglas. 1977. A qualitative survey offish and macroinvertebrates

of the French Broad River and selected tributaries. Div. Water Resour.

TVA, Tech. Note B35. 37 pp.

Hynes, H.B.N. 1972. The Ecology of Running Waters. Univ. Toronto Press,

Toronto. 555 pp.

Jensen, L.D., and A.R. Gaufin. 1964. Effects of ten organic insecticides on two

species of stonefly naiads. Trans. Am. Fish. Soc. 95:356-367.

Lehmann, Jens. 1971. Die Chironomiden der Fulda (Systematische, okologische

und faunistische untersuchungen). Arch. Hydrobiol. Suppl. 57:466-555.

Lenat, David R., D.L. Penrose and K.W. Eagleson. 1979. Biological evaluation

of non-point source pollutants in North Carolina streams and rivers. N.C.

Dep. Nat. Resour. Comm. Dev., Raleigh. 168 pp.

Lindegaard-Peterson, Claus. 1982. An ecological investigation of the chironom-

idae (Diptera) from a Danish lowland stream (Linding A). Arch. Hydro-

biol. (59:465-507.

Lotspeich, Fredrick B. 1980. Watersheds as the basin ecosystem: This concep-

tual framework provides a basis for a natural classification system. Water

Resour. Bull. 7(5:581-586.

Mason, William T. 1973. An introduction to the identification of Chironomid

larvae. U.S. Environ. Prot. Agency, Cincinnati. 90 pp.

Morihara, D.K., and W.P. McCafferty. 1979. The Baetis larvae of North

America (Ephemeroptera: Baetidae). Trans. Am. Entomol. Soc. (Phila.) 105:

139-221.

Mozley, Samuel C. 1980. Biological indicators of water quality in North Caro-

lina: I. Guide to the generic identification of orthocladiinae chironomidae

(Diptera). Final Report to N.C. Div. Environ. Manage., Raleigh. 90 pp.

Neves, Richard J. 1979. Secondary production of epilithic fauna in a woodland

stream. Am. Midi. Nat. 102:209-224.

N.C. Department of Natural Resources and Community Development. 1976.

Water Quality Management Plan. French Broad River Basin.

1977. North Carolina Water Quality Inventory.

Oliver, D.R. 1971. Life history of the Chironomidae. Annu. Rev. Entomol.

7(5:211-230.

Penrose, David L., D.R. Lenat and K.W. Eagleson. 1980. Biological evaluation

of water quality in North Carolina streams and rivers. N.C. Dep. Nat.

Resour. Comm. Dev., Raleigh. 181 pp.

Rhame, Roy E., and K.W. Stewart. 1976. Life cycles and food habits of three

Hydropsychidae (Trichoptera) species in the Brazos River, Texas. Trans.

Am. Entomol. Soc. 702:65-99.

Ross, Herbert H. 1944. The caddisflies, or Trichoptera, of Illinois. 111. Nat. Hist.

Surv. Bull. 25:1-326.

Seagle, Henry H. Jr., and A.C. Hendricks. 1978. Benthic macroinvertebrates of

the Shenandoah Basin, pp. 36-43 in K.C. Flynn and W.T. Mason (eds.).

The Freshwater Potomac: Aquatic Communities and Environmental

Stresses. Interstate Comm. Potomac River Basin. 194 pp.

Starnes, Lynn B. 1976. Comparison offish and benthic faunal compositions in

polluted and pristine forks of a mountain river. Tenn. Wildl. Resour.

Agency Tech. Rep. No. 77-55. 102 pp.

50 David L. Penrose, David R. Lenat, K. W. Eagleson

Talak, Anthony, Jr, 1977. The recovery of stream benthic insect communities

following coal strip mining in the Cumberland Mountains of Tennessee.

Unpubl. M.S. Thesis. Univ. Tenn., Knoxville. 81 pp.

Tarter, Donald C, and R.F. Kirchner. 1978. A new species of Baetisca from

West Virginia. Entomol. News 59:209-213.

Tebo, L.B., Jr., and W.W. Hassler. 1961. Seasonal abundance of aquatic insects

in Western North Carolina trout streams. J. Elisha Mitchell Sci. Soc.

77:249-259.

United State Geological Survey. 1979. Water resources data for North Carolina,

water year 1979. 541 pp.

Vannote, Robert L., and B.W. Sweeney. 1980. Geographic analysis of thermal

equilibria: a conceptual model for evaluating the effects of natural and

modified thermal regimes on aquatic insect communities. Am. Nat.

7/5:667-695.

Wallace, J. Bruce. 1975. Food partioning in net-spinning Trichoptera larvae:

Hydropsyche venularis, Cheumatopsyche etrona and Macronema zebra turn

(Hydropsychidae). Ann. Entomol. Soc. Am. 65:463-471.

White, Tina R., and R.C. Fox. 1980. Recolonization of stream insects following

channelization. Clemson Univ. Water Resour. Res. Inst. Tech. Rep. No.

87(1). 120 pp.

Accepted I August 1982

Northern Limits of the Southeastern Shrew, Sorex

longirostris Bachman (Insectivora: Soricidae), on the

Atlantic Coast of the United States

John F. Pagels and Carol S. Jones

Department of Biology,

Virginia Commonwealth University, Richmond, Virginia 23284

AND

Charles O. Handley, Jr.

Division of Mammals,

U. S. National Museum of Natural History,

Smithsonian Institution, Washington, D. C. 20560

ABSTRACT. — Records of captures of 61 specimens of the southeast-

ern shrew, Sorex longirostris longirostris Bachman, were obtained for

37 localities in the District of Columbia, Maryland, and Virginia.

Twenty-four localities are in the Piedmont, eleven in the Coastal Plain,

and two in the Ridge and Valley Province. Shrews were taken at eleva-

tions averaging about 91 m (300 ft.), and ranging from near sealevel to

366 m (1200 ft.). Capture records indicate a wide choice of habitats,

from old-fields to upland hardwood forests, but nearly all can be

construed as disturbed. Japanese honeysuckle or other plants that pro-

vide dense ground cover are important features of most habitats of the

southeastern shrew.

INTRODUCTION

Knowledge of distribution, natural history, and systematic relation-

ships of shrews of the Middle Atlantic states has increased rapidly in

recent years. For example, the rock shrew, Sorex dispar Batchelder,

only recently has been found in Maryland (Mansueti and Flyger 1952).

In Virginia, the rock shrew and water shrew, Sorex palustris Richard-

son, were added to the State's list even more recently (Handley 1956,

and Pagels and Tate 1976, respectively). The Dismal Swamp short-

tailed shrew, which was in the past accorded the rank of a species, now

is considered to be a subspecies, Blarina brevicauda telmalestes Mer-

riam (Handley 1979). The southern short-tailed shrew, formerly thought

to be a smaller subspecies of B. brevicauda, now is regarded as a spe-

cies, Blarina carolinensis Bachman (Genoways and Choate 1972; Tate et

al. 1980). The pygmy shrew, Sorex hoyi Baird (formerly Microsorex

hoyi, but see Diersing 1980), long known from only two sites in Virginia

(Handley and Patton 1947), since 1950 has been collected at a montane

locality and at several Piedmont and Coastal Plain locations in the

State (Handley et al. 1980).

Brimleyana No. 8:51-59. December 1982 51

52 John F. Pagels, Carol S. Jones, Charles O. Handley, Jr.

Based on few specimens from six scattered localities in Piedmont

and mountain counties, Handley and Patton (1947) suggested that the

southeastern shrew, Sorex longirostris longirostris Bachman, had a

nearly statewide distribution in Virginia. Later, Stout (1967) reported a

capture in Hanover County, and noted that this record, along with the

Fairfax County and Brunswick County records of Handley and Patton

(1947) established the easternmost distributional limit of S. I longiros-

tris very near the Fall Line in Virginia. Gardner (1950) and Paradiso

(1969) reported five specimens from four localities in Maryland and one

from the District of Columbia, all taken in the Coastal Plain. Recent

collections and studies stressing the biology of shrews, for example

those of Tate et al. (1980) in Virginia, obtained many additional speci-

mens of S. longirostris and provided much ecological data.

This paper summarizes ecological and distributional data on 5. /.

longirostris in the District of Columbia, Maryland, and Virginia, and

makes comparisons with accounts from other sections of the eastern

United States (French 1980a, b). Sorex I. fisheri Merriam, which occurs

in the Dismal Swamp in southeastern Virginia and northeastern North

Carolina, is not considered here. Handley (1979, 1980) discussed this

larger subspecies.

MATERIALS AND METHODS

We found records of 61 specimens of S. I. longirostris that have

been taken in our area: one from the District of Columbia, five from

Maryland, and fifty-five from Virginia (see Fig. 1 and the list of speci-

mens examined). The specimens are in the Carnegie Museum of Natural

History (CM), North Carolina State Museum of Natural History

(NCSM), U.S. National Museum of Natural History (USNM), and Vir-

ginia Commonwealth University Mammal Collection (VCU).

Specimens Examined

DISTRICT OF COLUMBIA.— Washington, 1 (USNM).

MARYLAND.— Anne Arundel Co.: Shady Side, 1 (USNM). Calvert Co.:

Camp Roosevelt, nr. Chesapeake Beach, 2 (USNM); Chesapeake Beach, 1

(USNM). Prince Georges Co.: West Branch of Patuxent River, 4 mi W Hall, 1

(USNM).

VIRGINIA.— Amelia Co.: Virginia Secondary Highway (SH) 609, 6 mi NNE

Amelia, 2 (VCU); US Highway (US) 360, 3 mi NW SH 698 (2 mi N, 4 mi E

Amelia), 1 (VCU); US 360, ca. 3 mi SE SH 604 (ca. 6 mi ENE Amelia), 1

(VCU); Amelia Court House, 4 (USNM). Appomattox Co.: Appomattox-

Buckingham State Forest, 3 mi N, 9.8 mi W Appomattox, 1 (VCU). Arlington

Co.: Little Pimmit Run, 2 mi SW Chain Bridge, 1 (USNM). Brunswick Co.:

Seward Forest, nr. Triplett, 8-10 mi N Virginia-North Carolina boundary, 2

Northern Limits of Southeastern Shrew

53

Fig. 1. Distribution of the southeastern shrew, Sorex longirostris longirostris, in

the District of Columbia (DC), Maryland (MD), and Virginia (VA). Asterisk in

WV indicates the only reported locality of S. longirostris in West Virginia

(French 1976). Ridge and Valley Province and Piedmont Province are shaded.

Solid circle = specimen examined; open circle = specimen reported.

54 John F. Pagels, Carol S. Jones, Charles O. Handley, Jr.

(USNM). Buckingham Co.: Virginia Primary Highway (PH) 15 at north bank

of Willis River (2 mi NE Sheppards), 1 (VCU). Charles City Co.: PH 5, 0.25 mi

E PH 156 (2 mi SE Shirley), 1 (VCU). Chesterfield Co.: US 360, 2 mi W Swift

Creek (4 mi E Skinquarter), 1 (VCU). Chesterfield- Powhatan cos. boundary:

Keswick Farm, 4 mi N Midlothian, 1 (USNM). Culpeper Co.: Lignum, 10 mi

SE Culpeper, 1 (USNM). Cumberland Co.: US 60, 5-6 mi W Cumberland Court

House (nr. Buckingham Co. line), 2 (VCU). Essex Co.: 3.5 mi SW Center Cross,

2 (USNM). Fairfax Co.: Falls Church, 2 (USNM); Springfield, 1 (USNM); 2 mi

NW Vienna, elev. 125 m (410 ft.), 1 (USNM); nr. Burke, 5.5 mi SSE Fairfax, 1

(USNM). Fauquier Co.: US 17, 4 mi SE Marshall, 1 (VCU). Hanover Co.: 1.5

mi S Montpelier, 1 (USNM). Henrico Co.: 1-2 mi E Varina, 3 (VCU). Mecklen-

burg Co.: 2 mi S, 1 mi E Clarksville, 3 (VCU); 3.5 mi N, 4 mi W Clarksville, 1

(VCU). New Kent Co.: 0.5 mi NE Bottoms Bridge, 1 (VCU). Page Co.: Shenan-

doah National Park, headquarters area, 4.7 mi ENE Luray, elev. 366 m (1200

ft.), 3 (USNM). Powhatan Co.: US 60 at east bank of Sallee Creek (4 mi WNW

Powhatan), 1 (VCU). Prince Edward Co.: no exact locality, 1 (USNM). Prince

George Co.: 3.4 mi W Disputanta, 1 (VCU). Prince William Co.: 4 mi SE

Manassas, 2 (USNM). Surry Co.: Pipsico Scout Reservation, nr. Scotland, 4 mi

NE Surry, 1 (USNM). Warren Co.: Browntown Rd. (SH 649), 3.5 mi SSW

Front Royal, elev. 305 m (1000 ft.), 1 (CM).

Additional Virginia specimens, not seen: Arlington Co.: Little Pimmit Run,

2 mi SW Chain Bridge, 6 (Bray 1939). Brunswick Co.: Seward Forest, nr. Tri-

plett, 2 (Lewis 1943). Hanover Co.: nr. Ashland, 1 (NCSM).

RESULTS AND DISCUSSION

Distribution

The 61 specimens were taken at 37 localities: 11 in the Coastal

Plain, 24 in the Piedmont, and 2 in the Ridge and Valley Province.

Elevation of captures ranged from near sealevel to 366 m (1200 ft.) and

averaged about 91 m (300 ft.).

Not surprisingly, the localities cluster in two groups, around Rich-

mond, Virginia, and Washington, D. C, the home bases of active col-

lectors (Fig. 1). The localities are distributed about equidistant from the

Fall Line in both clusters. Thus, in the north, where Coastal Plain and

Piedmont are narrow, S. I. longirostris has been found throughout,

from coast to mountains, probably approximating the true extent of the

range of the species there. On the other hand, in southern Virginia,

where both Coastal Plain and Piedmont are wide, S. I. longirostris has

been found only in the upper Coastal Plain and lower Piedmont. We

suppose that this distribution is an artifact of collecting.

In Maryland, S. longirostris probably occurs only in the Coastal

Plain in the southern sector of the Western Shore. It is absent from the

Eastern Shore, the northern sector of the Western Shore, and the entire

Piedmont and mountainous portions of the State.

Northern Limits of Southeastern Shrew 55

It seems likely that in Virginia S. I. longirostris actually occurs

throughout the Piedmont and Coastal Plain, except on the Eastern

Shore (and in the extreme Southeast, where it is replaced by S. I.

fisheri). Distribution of this shrew in the western, mountainous portion

of Virginia remains to be determined. It definitely occurs west of the

Blue Ridge near Luray, Page County, and near Front Royal, Warren

County, from whence we have examined specimens. It also has been

reported at Mountain Lake, Giles County (Odum 1944); Blacksburg,

Montgomery County (Handley and Patton 1947); and Big Levels,

Augusta County (Bruce 1937). We examined the Giles County speci-

mens collected by Odum and agree with French (1980a) that they are

young masked shrews, Sorex cinereus Kerr. The Montgomery and

Augusta County specimens apparently have been lost. Handley et al.

(1980) supposed that those specimens also were misidentified. It is pos-

sible that S. I. longirostris makes incursions into the lower sectors of the

Blue Ridge and Ridge and Valley provinces in the drainages of the

James, Roanoke, and Tennessee rivers, as it does through the Potomac

watershed into the lower Shenandoah Valley, but this has yet to be

verified.

Sorex longirostris seems to be limited in its westward distribution

in Virginia and in its northward distribution in Maryland by the pres-

ence of S. cinereus. This species is widespread and locally abundant in

western Virginia and in Maryland except on the Western Shore south of

the latitude of Annapolis. These similar species appear to exhibit con-

tiguous allopatry in Virginia and Maryland. There are few if any known

exceptions to their allopatry in this region. The reported occurrence of

S. c. fontinalis Hollister in northern Virginia (Arlington County, Bray

1939), an area inhabited by S. longirostris, is not an exception. The

specimen (USNM 267569) identified formerly as S. c. fontinalis actually

is S. longirostris (Handley 1982).

A possible exception is a specimen of S. c. fontinalis (USNM

219050, identification verified by us), collected in 1918 by Titus Ulke

and labeled "Hollywood, Maryland". Hollywood post office is in St.

Marys County, in extreme southern Maryland, well within the range of

S. longirostris. There is also a "Hollywood" community about two miles

northeast of the center of College Park, Prince Georges County. This

"Hollywood" is within the known range of S. c. fontinalis. Gardner

(1950) and Paradiso (1969) attributed USNM 219050 to Hollywood,

Prince Georges County. There are no original labels with Ulke's speci-

men and the accession files shed no light on the whereabouts of Ulke's

"Hollywood". However, local history is helpful in pinning down the

locality. The Hollywood in Prince Georges County was founded around

1900 by Edward Daniels as "Hollywood-on-the-Hill". About 1902,

when the Rhode Island Avenue trolley line from the District of Colum-

56 John F. Pagels, Carol S. Jones, Charles O. Handley, Jr.

bia terminated there, the locality, by then consisting of about a dozen

Victorian style houses, had become known simply as "Hollywood". By

1918 (when Titus Ulke collected the shrew) the trolley line had been

extended to Laurel, but Hollywood was still a stop. Evidently there was

confusion at that time over the status of that "Hollywood". Fred De

Marr, who grew up in the Hollywood area of Prince Georges County,

remembers that Christmas cards addressed to "Hollywood" eventually

reached his family by way of the post office 50 miles away in Holly-

wood, St. Marys County. Titus Ulke was a resident of the District of

Columbia. It would have been easy for him to reach Hollywood in

Prince Georges County by trolley and not surprising for him to label a

specimen collected there "Hollywood, Maryland." It would have been

more difficult for him to reach Hollywood in St. Marys County. Since

the Prince Georges' Hollywood is within the known range of S. cinereus

and the St. Marys' Hollywood is not, it is reasonable to suppose that

Titus Ulke's specimen came from Prince Georges County.

The specimens of S. longirostris from Page and Warren counties,

Virginia, appear on a map to be within the range of S. cinereus, but

actually the two species probably are ecologically as well as altitudinally

segregated. These examples suggest that factors other than or in addi-

tion to physiography and ecology operate to delineate the contiguous

margins of the ranges of S. cinereus and S. longirostris. Additional

study in southern Maryland and western Virginia, in and near areas of

contiguity (or of potential sympatry) of the ranges of these species,

should provide information on factors limiting their distribution.

Ecology

We have ecological data for 41 of the specimens of S. I. longirostris

that have been taken in Virginia. Capture locations were almost evenly

distributed between open country (22) and forest (19). Those taken in

the open (19) were mostly in old abandoned fields, usually in dry sites in

dense sedge and/ or grass, with tangles of Japanese honeysuckle and

Rubus, and scattered saplings of pine, red cedar, black locust, or other

small trees and shrubs. A few specimens have come from older succes-

sional stages where saplings and shrubs predominated. Lewis (1943)

reported old-field captures of S. I. longirostris in damp sites: in pros-

trate honeysuckle in a small, wet swale; in a heavy growth of sedge at

the edge of a small sphagnum bog in a field where there were scattered

small pines; and in a dense growth of tall sedge and grass at the edge of

a former mill pond. We also have specimens from a weedy ditch bank in

a corn field, from grass and Rubus in a narrow ecotone between a cultiv-

ated field and a swamp, and from a swimming pool in a suburban yard.

Forests yielding specimens of S. I. longirostris most often have been

young forests (16 of 19 records). Bray (1939) caught seven Sorex, pre-

Northern Limits of Southeastern Shrew 57

sumably all S. I. longirostris, and Lewis (1943) caught one, in second

growth bottomland deciduous forest with deep leaf litter, numerous

decaying logs, and a dense overgrowth of honeysuckle. We have several

specimens from honeysuckle tangles in windfall openings in young pine

plantations, and Lewis (1943) had one from dense sedge and grass

beside a small pond in a pine plantation. Four captures were in second

growth mixed pine-hardwood forest with leaf litter but scant herbaceous

cover.

Only three captures were in older forest: one in Vaccinium and

herbs in deciduous forest on a steep, rocky bluff (Lewis 1943), another

in dry upland mixed woodland with deep leaf litter and a cover of Vac-

cinium, and a third in mixed bottomland forest with deep litter and a

lush growth of herbs and grass.

In summary, almost all of our habitat records for S. I. longirostris

are for disturbed situations: cultivated fields, abandoned fields, thickets

of saplings and shrubs, and young forest. Only 3 of 40 records pertain

to older forest. Another common thread through most of the records is

a dense ground cover of grass, sedge, herbs, or honeysuckle. There is no

evident preference for upland or lowland, dryness or moisture, type of

old-field, or type of young forest.

French (1980a,b) reviewed published ecological information on S. /.

longirostris throughout its range and noted that it has been found in

habitats varying from planted fields to dry upland hardwood forest to

moist floodplain forest. He observed (1980b:2) that, "In all habitats 5. /.

longirostris has been most often associated with a heavy ground cover

of grasses, sedges, rushes, blackberry, Japanese honeysuckle and/ or

thick mats of decaying leaves." The majority of S. I. longirostris

reported by Tuttle (1964) in Tennessee and by Rose (1980) in southern

Indiana were taken in association with honeysuckle.

Since S. I. longirostris has been found in so many kinds of habitat,

it may be that the microhabitat represented by honeysuckle or equally

dense ground cover is more important to the shrew than is the major

habitat type. Preference for disturbed habitats may also reflect the need

for dense ground cover. Such a microhabitat presumably would supply

shelter from both nocturnal and diurnal predators, primarily raptorial

birds, and also would provide a microclimate suitable for both the

shrew and its largely invertebrate food supply.

Very likely Sorex /. longirostris has benefited from the vast destruc-

tion of forests and habitat alteration that has occurred in the past 300

years in the southeastern United States. As has been suggested for sev-

eral other species (Pagels 1980), it would not be surprising to find that

this shrew is more abundant and widespread today than it was before

the European settlement of this region.

58 John F. Pagels, Carol S. Jones, Charles O. Handley, Jr.

ACKNOWLEDGMENTS.— We are grateful to many individuals

who provided specimen or ecological data and/ or allowed us to exam-

ine specimens under their care. They included J. A. Cranford, C. E.

Close, V. E. Diersing, T. W. French, J. B. Funderburg, A. L. Gardner,

H. H. Genoways, M. S. Hensley, H. G. M. Jopson, G. L. Kirkland, Jr.,

J. Laerm, D. S. Lee, J. F. Merritt, R. K. Rose, F. Sibley, M. D. Tuttle,

and S. L. Williams. Tom French, in addition to providing discussion

and information, critically reviewed the manuscript. An anonymous

reviewer provided other helpful comments. C. M. Tate assisted in the

collection of specimens. D. W. Handley determined the location of an

equivocal specimen locality with the aid of J. Crivak and F. S. De

Marr. Pippa Vanderstar constructed the distribution map.

LITERATURE CITED

Bray, Robert S. 1939. Sorex fontinalis in Virginia. J. Mammal. 20:102.

Bruce, James A. 1937. Sorex longirostris longirostris in Augusta County, Vir-

ginia. J. Mammal. 75:513-514.

Diersing, Victor E. 1980. Systematics and evolution of the pygmy shrews (sub-

genus Microsorex) of North America. J. Mammal. (57:76-101.

French, Thomas W. 1976. The first record of the southeastern shrew, Sorex

longirostris, in West Virginia. J. W. Va. Acad. Sci. ¥5:120-122.

1980a. Natural history of the southeastern shrew, Sorex longirostris

Bachman. Am. Midi. Nat. 704:13-31.

1980b. Sorex longirostris. Mamm. Species 143:1-3.

Gardner, Marshall C. 1950. A list of Maryland mammals. (Part I. Marsupials

and Insectivores). Proc. Biol. Soc. Wash. 65:65-68.

Genoways, Hugh H., and J. R. Choate. 1972. A multivariate analysis of system-

atic relationships among populations of the short-tailed shrews (genus

Blarina) in Nebraska. Syst. Zool. 27:106-116.

Handley, Charles O., Jr. 1956. The shrew Sorex dispar in Virginia. J. Mammal.

57:435.

. 1979. Mammals of the Dismal Swamp: A historical account. Pp.

297-357 in Kirk, P. W., Jr. (ed.). The Great Dismal Swamp. Univ. Press

Virginia, Charlottesville. 427 pp.

. 1980. Sorex longirostris fisheri Merriam. Pp. 535-536 in Linzey,

D.W. (ed.). Endangered and Threatened Plants and Animals of Virginia.

Center for Environmental Studies, VPI & SU, Blacksburg. 665 pp.

. 1982. Deletion of Sorex cinereus fontinalis from taxa known to

occur in Virginia. J. Mammal. 65:319.

, and C. P. Patton. 1947. Wild Mammals of Virginia. Comm. Game

Inland Fisheries, Richmond, vi + 220 pp.

, J. F. Pagels and R. H. de Rageot. 1980. Microsorex hoyi winne-

mana Preble. Pp. 545-547 in Linzey, D. W. (ed.). Endangered and Threat-

ened Plants and Animals of Virginia. Center for Environmental Studies,

VPI & SU, Blacksburg. 665 pp.

Northern Limits of Southeastern Shrew 59

Lewis, John B. 1943. Notes on the Bachman shrew. J. Mammal. 24:97-98.

Mansueti, Romeo, and V. F. Flyger. 1952. Long-tailed shrew (Sorex dispar) in

Maryland. J. Mammal. 53:250.

Odum, Eugene P. 1944. Sorex longirostris at Mountain Lake, Virginia. J.

Mammal. 25:196.

Pagels, John F. 1980. [Virginia mammals:] The changing scene. Pp. 603-609 in

Linzey, D. W. (ed.). Endangered and Threatened Plants and Animals of

Virginia. Center for Environmental Studies, VPI & SU, Blacksburg. 665

pp.

, and C. M. Tate. 1976. Shrews (Insectivora: Soricidae) of the Paddy

Knob-Little Back Creek area of western Virginia. Va. J. Sci. 27:202-203.

Paradiso, John L. 1969. Mammals of Maryland. N. Am. Fauna 66. 193 pp.

Rose, Robert K. 1980. The southeastern shrew, Sorex longirostris, in southern

Indiana. J. Mammal. (57:162-164.

Stout, Jack. 1967. An additional record of Sorex longirostris from eastern

Virginia. Chesapeake Sci. 8:264.

Tate, Cathy M., J. F. Pagels and C. O. Handley, Jr. 1980. Distribution and

systematic relationship of two kinds of short-tailed shrews (Soricidae:

Blarina) in south-central Virginia. Proc. Biol. Soc. Wash. 95:50-60.

Tuttle, Merlin D. 1964. Additional record of Sorex longirostris in Tennessee. J.

Mammal. 45:146-147.

Accepted 1 November 1982

Use of Lepomis macrochirus Rafinesque nests by Spawning

Notemigonus crysoleucas (Mitchill) (Pisces: Centrarchidae

and Cyprinidae)

David J. DeMont

Department of Zoology,

North Carolina State University, Raleigh, North Carolina 27650

ABSTRACT. — Golden shiners, Notemigonus crysoleucas, were observed

spawning in bluegill, Lepomis macrochirus, nests in a North Carolina

pond. Fifty-three percent of the larvae that hatched from eggs taken from

one of the nests were golden shiners. This spawning behavior may affect

bluegill reproductive success.

Several cyprinid fishes are known to spawn in the nests of other

cyprinid species (Raney 1940; Lachner 1952), but cyprinid spawning in

the guarded nests of centrarchids has been infrequently observed. Latta

(1957) reported that common shiners, Notropis cornutus (Mitchill),

spawned in smallmouth bass, Micropterus dolomieui Lacepede, nests in

Lake Michigan. Similar observations have been made with regard to

redfin shiner, Notropis umbratilis (Girard), in green sunfish, Lepomis

cyanellus Rafinesque, nests (Hunter and Wisby 1961) and golden shiner,

Notemigonus crysoleucas (Mitchill), in largemouth bass, Micropterus

salmoides Lacepede, nests (Kramer and Smith 1960). Pflieger (1975)

mentioned collecting golden shiners over green sunfish nests "where

they evidently spawn." Recently, I observed golden shiners spawning in

bluegill, Lepomis macrochirus Rafinesque, nests in North Carolina.

Observations were made from the dock of the North Carolina State

University Research Laboratory at Yates Pond, 10 km south of Raleigh,

North Carolina, on 14 May 1980. Water temperature at the time was

24° C. The bluegill nest affording the best view of golden shiner activity

was located 3 m from shore in approximately 30 cm of water. It occu-

pied a clear spot behind (inshore of) beds of smartweed, Polygonum

hydropiperoides and P. punctatum, which lined the shore to a depth of

1.5m.

Small groups of adult golden shiners (12-18 cm total length) left a

large school that was milling about just beyond the smartweed beds and

entered the shallow water where several male bluegills were guarding

nests. In groups of two to five, the shiners entered a nest which was

temporarily unguarded, perhaps due to the presence of the observer.

Each group remained in the nest for several seconds and upon leaving

was quickly replaced by another small group. The male bluegill that had

Brimleyana No. 8:61-63. December 1982 61

62 David J. DeMont

been guarding the nest remained absent during golden shiner spawning

but returned shortly after their activity had ceased several minutes later.

No antagonistic behavior of male bluegills toward golden shiners was

observed, although threats and attacks directed at juvenile bluegills and

other nesting males were frequent.

On 17 May 1980, a sample of the substrate from the nest in which

shiners had spawned was carefully transferred to a petri dish filled with

pond water for examination in the laboratory. This material (fibrous

roots of shoreline vegetation) contained 71 adhesive eggs. Twenty-four

eggs were dead and cove/ed with fungus while the remaining forty-seven

contained developing embryos. Forty-five larvae hatched from the vi-

able eggs during the next two days. Twenty-four (53%) were golden shin-

ers and the remainder were Lepomis sp. (presumably bluegills since a

male bluegill had been guarding the nest). Dead eggs were not

identifiable.

The environment provided by bluegill nests might be expected to

reduce egg and early larval mortality of golden shiners. The clean sub-

strate would help insure adhesion of the eggs and reduce the chance of

their being lost in pond sediments. Ideal conditions for golden shiner

spawning obtain when rising water levels in spring inundate shoreline

vegetation. The stems and leaves of this vegetation provide clean sub-

strate for the adhesive eggs (Guidice et al. 1981). In Yates Pond the

surface level falls in spring. The only available spawning substrate is

provided by aquatic macrophytes such as smartweed and spatterdock,

Nuphar luteum, but these are usually heavily encrusted with periphyton

and microinvertebrates. Therefore, the clean substrate in the centrarchid

nests would be expected to be attractive to spawning golden shiners.

Parental care by the male bluegill would also provide advantages to

the golden shiner eggs and prolarvae by protecting them from predators

and insuring a nearly constant supply of aerated water. The obser-

vations of Kramer and Smith (1960) in Lake George, Minnesota, sup-

port the contention that these expected benefits are real. They found

that golden shiner reproduction was most successful in successful bass

nests. In bass nests that had been prepared but unused, golden shiner

eggs disappeared shortly after spawning. Also, golden shiner young-of-

the-year were more abundant in summer bag-seine catches in Lake

George in years in which the percentage of bass nests used by golden

shiner adults was greatest.

It would be interesting to know what effects this spawning behavior

has on the reproductive success of centrarchids. In Yates Pond, larval

and early juvenile bluegills have been frequently observed schooling

with golden shiners of similar size. This activity could afford the bluegill

some protection from predators. On the other hand, increased egg den-

Spawning Notemigonus Using Lepomis Nests 63

sity in nests might lower dissolved oxygen concentration while elevating

levels of carbon dioxide and ammonia. Since developing golden shiners

are not normally dependent upon constant parental ventilation, slight

degradation of water quality in the nest would be expected to be less

harmful to them than to the developing bluegills. These factors, in addi-

tion to competition for food at the time of yolk sac absorption, would

be detrimental to centrarchid reproductive success. If these kinds of

negative interactions are real, their existence might help explain the fre-

quently observed decline in fishery value of bass/bluegill ponds follow-

ing the introduction of golden shiners.

ACKNOWLEDGMENTS.— I wish to thank Drs. Melvin T. Huish

and John M. Miller, and an anonymous reviewer, for helpful comments

on the manuscript. Paper number 8081 of the Journal Series of the

North Carolina Agricultural Research Service, Raleigh, N.C., 27650.

LITERATURE CITED

Guidice, John J., D. L. Gray and J. M. Martin. 1981. Manual for bait fish

culture in the South. Univ. Arkansas Coop. Extension Serv. Publ. EC550.

Hunter, John R., and W. J. Wisby. 1961. Utilization of the nests of green

sunfish {Lepomis cyanellus) by the redfin shiner {Notropis umbratilis

cyanocephalus). Copeia 1961(1): 1 13-1 15.

Kramer, Robert H., and L. L. Smith, Jr. 1960. Utilization of nests of large-

mouth bass, Micropterus salmoides, by golden shiners, Notemigonus

crysoleucas. Copeia 1960(l):73-74.

Lachner, Ernest A. 1952. Studies on the biology of the cyprinid fishes of the

chub genus Nocomis of the Northeastern United States. Am. Midi. Nat. 48:

433-466.

Latta, William C. 1957. The ecology of the smallmouth bass, Micropterus d.

dolomieui Lacepede, at Waugoshance Point, Lake Michigan. Unpubl.

Ph.D. Dissert., Univ. Mich., Ann Arbor. 127 pp.

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

City. 343 pp.

Raney, Edward C. 1940. The breeding behavior of the common shiner, Notropis

cornutus (Mitchill). Zoologica 25.1-14.

Accepted 19 November 1982

Systematics of the Troglobitic Caecidotea

(Crustacea: Isopoda: Asellidae)

of the Southern Interior Low Plateaus

Julian J. Lewis

Department of Biology and Water Resources Laboratory,

University of Louisville, Louisville, Kentucky 40292

ABSTRACT. — Caecidotea meisterae is synonymized with Caecidotea

whitei, which is reduced to a subspecies of Caecidotea bicrenata. This

species is now divided into two subspecies: Caecidotea bicrenata

bicrenata and Caecidotea bicrenata whitei. Caecidotea b. bicrenata

occurs in caves from northern Alabama to central Tennessee. From

northern Tennessee to southern Illinois it is replaced by C. b. whitei.

Fleming (1972a), following Bresson (1955) and Steeves (1963;

1964), considered Caecidotea alabamensis Stafford (1911 to be a wide-

spread troglobite inhabiting caves of practically the entire Interior Low

Plateaus, from Alabama to Indiana and Illinois. Lewis and Bowman

(1981) pointed out the morphological and zooeographical dissimilarities

of C. alabamensis to species occurring in caves of the Interior Low Pla-

teaus, restricting the known distribution of C. alabamensis to the type-

locality, Auburn, Alabama. Caecidotea j or dani (Eberly) was resurrected

for a distinct species in central Indiana, as was Caecidotea bicrenata

(Steeves) for the troglobitic species in northern Alabama. Three new

species closely related to C. jordani and C. bicrenata were described by

Lewis and Bowman (1981) to encompass "alabamensis" collections from

southern Illinois, Kentucky, and northern Tennessee. These were Caeci-

dotea beattyi, Caecidotea meisterae, and Caecidotea whitei.

Additional information now necessitates modification of oart of the

scenario of Lewis and Bowman (1981) for the "alabamensis" species.

While that paper was in press, collecting in Mammoth Cave National

Park produced both Caecidotea whitei and C meisterae, in addition to

Caecidotea stygia. The presence of both C. stygia and C. whitei in the

ecologically complex Mammoth Cave System was explained by Lewis

and Lewis (1980). However, the presence of three species stretched

credence, pointing to the possibility that C. meisterae and C. whitei are

conspecific.

Study of numerous specimens from the base level cave rivers of the

Mammoth Cave System revealed intergradations between specimens

with the weakly developed gnathopods of C. whitei and those with the

more fully differentiated gnathopods of C. meisterae. As the morph-

ology of the gnathopods was the primary character used to distinguish

these two species, it became apparent that they were identical.

A gray area left unconsidered by Lewis and Bowman (1981) was

central Tennessee, where numerous collections previously called

"alabamensis" were assigned neither to C. bicrenata nor any of the new

Brimleyana No. 8:65-74. December 1982 65

66 Julian J. Lewis

species. Examination of many collections showed that C. bicrenata's

range extends from northern Alabama to north central Tennessee. In

northern Tennessee and southern Kentucky C. whitei occurs, continuing

north to Hart County, Kentucky, and west into southern Illinois. The

only substantial morphological difference between C. bicrenata and C.

whitei is the shape and placement of the lateral process of the male

second pleopod endopod tip. Although this difference in morphology is

consistent, other distinguishing characteristics are too variable to be

reliable. Furthermore, significant dispersal barriers that might provide

reproductive isolation do not appear to exist along the boundary

between the ranges of C. whitei and C. bicrenata.

Caecidotea meisterae is herein synonymized with Caecidotea

whitei, and C. whitei reduced to a subspecies of Caecidotea bicrenata.

The rest of the taxonomy proposed by Lewis and Bowman (1981) for

other "alabamensis" species, i.e., Caecidotea antricola, C. beattyi and C.

jordani, remains unchanged.

Caecidotea bicrenata bicrenata, new status.

Asellus alabamensis. — Bresson, 1955:51-59, 65, 70. — Chappuis,

1957:39, 41-42, (in part, C. antricola Missouri specimens). —

Steeves, 1964:503-504; 1966:394-396, 401-402; 1969:521. — Barr,

1967:190-191. — Cooper and Cooper, 1968:22. — Fleming, 1972a:

230-231, 245-247 (Alabama records); 1972b:498; 1973:287-291, 294,

300, 302-303.

Asellus bicrenatus Steeves, 1963:471, 474-476, 478, 480.

Conasellus alabamensis. — Henry and Magniez, 1970:356.

Material examined — ALABAMA: Colbert Co., Cobbs Bear Pit

Cave, 25 Oct 1969, F. Shires and R. Cobb, 3<3<3, 1$. Jackson Co.,

Flatworm Cave, 7 May 1969, R.C. Graham, 2$$, 3$$; New Fern Cave,

W. Torode, July 1969, 5$$. Marshall Co., Beech Spring Cave, July

1969, W. Wilson, R.C. Graham, 4#<5, 3$$; Off Limits Pit Cave, Feb

1971, R.C. Graham, 1Q, 29$. Morgan Co., B & J Cave, 23 July 1970,

W. Torode, 1$$, 6tf<$. TENNESSEE: Bedford Co., Reese Cave, 8.6

mi. S Shelbyville, 22 Dec' 1956, L. Hubricht, 1<J, 7?$. Cannon Co.,

cave 3.5 mi. SSW Bradyville, 21 Aug 1967, S. Peck, A. Fiske, 1<J, 1$;

Fisher Cave, 1 July 1973, S. Peck, 1<J, 29$; Ten Penney Cave, 2 mi.

NW Woodbury, 9 Sep 1967, S. Peck, A. Fiske, 2$$, 3$$. Davidson

Co., Brents Cave, 18 Nov 1956, T.C. Barr, 3<5<3, 23$$. DeKalb Co.,

Ted Cave, 5 mi. E Smithville, 29 Aug 1939, L. Hubricht, \2$$, 13$$.

Franklin Co., Caroline Cove Cave, 5.5 mi. SE Belvidere, 11 July 1967,

S. Peck, A. Fiske, 1$$, 20$$; Lost Cove Cave, 5 mi. N Sherwood, 27

Aug 1968, S. Peck, 3<$<$, 10$$; Pitcher Ridge Cave, 6 mi. N Hytop, 19

Aug 1967, S. Peck, A. Fiske, 2$$, 3$$; Putnam Spring Cave, 9 mi. S

Belvidere, 19 July 1967, S. Peck, A. Fiske, 7Q$, 7$$; seep, 3.8 mi. N

Sherwood, 9 May 1954, L. Hubricht, 18<3<5, 5$$; seep, 6.5 mi. S Sewa-

nee, 23 May 1961, L. Hubricht, 1$$, 6$$, Marion Co., Crystal Cave,

Monteagle, 17 Mar 1931, collector unknown, \$. Rutherford Co., Echo

Systematics of Troglobitic Caecidotea 67

Cave, 1.2 mi. N Rockvale, 22 Oct 1956, T.C. Barr, 1<$, 799; Rainbow

Cave, 2.3 mi. SW Walter Hill, 1 June 1941, L. Hubricht, 2$$, 599.

White Co., Haskell Cave, 2 mi. E Doyle, 24 Dec 1956, L. Hubricht, \$\

Indian Cave, 2.5 mi. SE Quebec, 23 Dec 1956, L. Hubricht, 4<$#, 19;

Moore Cave, 28 Oct 1969, J. Holsinger, R. Baroody, %$$, 1199. Wil-

son Co., Hayes Cave, 1 mi. SE Statesville, 8 Aug 1967, S. Peck, A.

Fiske, 2$$, 2099; Jackson Cave, Cedars of Lebanon State Park, 22

Sep 1967, S. Peck, A. Fiske, 2$$y 699.

Diagnosis of male. — Antenna 1 esthete series uninterrupted,

esthete formula from 3-0-0 to 5-0-0. Pereopod 1, palm of propodus with

proximal spines, bicuspate, usually low, mesial and distal processes.

Pleopod 1, protopod with 3 retinacula, exopod broadly rounded dis-

tally, slightly concave laterally, setae along distal and lateral margins,

most elongate laterally. Pleopod 2 endopod tip consisting of 2 pro-

cesses, cannula beak-shaped, extending perpendicular to axis of endo-

pod, lateral process subterminal parallel to cannula, originating within

margin of endopod, recurved. Pleopod exopod 4 with single sigmoid

suture.

Caecidotea b. bicrenata may be distinguished from C. b. whitei by

the position of the lateral process of the male second pleopod endopod

tip. In the former subspecies, the lateral process is placed within the

margin of the endopod and is recurved, often strongly, while in the

latter subspecies the lateral process is found on the margin of the

endopod and is straight. In mature specimens, the bicuspate processes

of the propod are usually lower in C. b. bicrenata than in C. b. whitei.

Range. — Caves, from northern Alabama to northcentral Ten-

nessee, approximately to the Cumberland River but not reaching the

southern extension of the Pennyroyal plateau (Fig. 1).

Caecidotea bicrenata whitei, new combination

Asellus alabamensis. — Fleming, 1972a: 247-248 (in part)

Asellus antricolus. — Fleming, 1972a: 245 (Twin Level Cave)

Caecidotea sp. no. 1. — Peck and Lewis, 1978: 44.

Caecidotea sp. no. 2. — Peck and Lewis, 1978: 44.

Caecidotea sp. — Lewis and Lewis, 1980: 23-27. — Lewis, 1981a: 21;

1981b: 234-236.

Caecidotea meisterae Lewis and Bowman, 1981: 28-32.

Caecidotea whitei Lewis and Bowman, 1981: 51-59.

Material examined. — TENNESSEE: Davidson Co., Crocker

Springs Cave, 12 Nov 1956, C.K. Barr, 3<3<J, 499; Nashville, 3 Mar

1901, E.B. Williamson, 6$$, 699. Sumner Co., Escue Cave, 2 mi. NE

Portland, 18 Apr 1958, L. Hubricht, 10<5<3, 1799. KENTUCKY: Edmon-

son Co. , Mammoth Cave National Park, Cedar Sink Cave, 5 mi. SW

Mammoth Cave, 31 Aug 1939, L. Hubricht, 7>$$. Cave over Styx River

Spring, 26 June 1981, J. Lewis, T. Lewis, 4<5<$, 19 . Mammoth Cave:

Flint Dome in Jessup Avenue, 6 Sep 1981, J. Lewis, J. Eckstein, 45$,

999; Styx River, near Natural Bridge, 28 June 1980, J. Lewis, T. Lewis,

68

Julian J. Lewis

Fig. 1. Distribution of the troglobitic Caecidotea of the southern Interior Low

Plateaus: Caecidotea bicrenata bicrenata (triangles); Caecidotea bicrenata whitei

(solid circles); Caecidotea bicrenata unidentified subspecies, lacking lateral pro-

cess on second pleopod endopod tip (stars); Caecidotea stygia (squares). The

stippled region, which comprises most of the range of C. b. whitei, is the Penny-

royal Plateau.

Systematics of Troglobitic Caecidotea 69

15; Hawkins River, 2 Aug 1980, J. Oberlies, 5$$, 999; Roaring River,

Shrimp Pools area, 19 Aug 1980, J. Lewis, T. Lewis, 1$$, 1499; pools

in Carlo's Way, 17 Oct 1981, J. Lewis, T. Lewis, M. Hale, 4$$, 399;

same location, 28 Dec 1981, J. Lewis, T. Lewis, J. Eckstein, 1$, 19-

Parker Cave, Parker River, 20 Aug 1980, J. Lewis, T. Lewis, 8<$<$,

1599. Barren Co., Mill Hole, 20 Aug 1980, J. Lewis, T. Lewis, 1$$,

1499. Simpson Co., Old Smokey Cave, 20 mi. SW Bowling Green, 1

July 1981, J.R. Holsinger, 2\$$, 1999.

Diagnosis. — Caecidotea b. whitei can be distinguished from the

nominate subspecies by the straight lateral process of the male second

pleopod endopod tip of C. b. whitei, which is placed directly on the

margin of the endopod. In mature males where both the medial and

distal processes are well developed, the medial process usually appears

as a large triangular process with a shoulder distally, rather than a well

developed bicuspate process. This characteristic, however, is also shared

with some populations of C. b. bicrenata.

Range. — Caecidotea b. whitei occurs from northcentral Tennessee

north to Hart County, Kentucky, where the faulted, sandstone Hart

County Ridge is apparently a barrier to its dispersal. To the west the

species occurs across the Pennyroyal Plateau, from Mammoth Cave

into western Kentucky, and the extension of the plateau in southern

Illinois. Caecidotea b. whitei appears to exclude C stygia in the Penny-

royal west of Mammoth Cave. In the Kentucky and Illinois counties

adjacent to the Ohio River, C stygia again occurs, and at least in Har-

din County, Illinois, C. b. whitei is absent. In southwestern Illinois, C.

b. whitei again replaces C. stygia, although C stygia is reported from

western Illinois and eastern Missouri (Fleming 1972a, b; Lewis and

Bowman 1981; Peck and Lewis 1978). In the Mammoth Cave System of

central Kentucky, both species occur syntopically, with C. stygia in

small streams in the upper levels of the cave and C. whitei in the base

level cave rivers (Lewis and Lewis 1980; Lewis 1981a)

Discussion. — In support of the synonymy of Caecidotea meisterae

with C. whitei, illustrations of the gnathopods of both forms from a

habitat in Mammoth Cave are given in Figure 2. Figure 3 illustrates the

tip elements of male second pleopod endopods. Figure 4 illustrates male

pleopod 1, and Figure 5 shows the palmar margin of the propodus of

male first pereopods. Although C. meisterae is the more differentiated

of the two forms, C. whitei is chosen as the senior synonym for two

reasons. First, in the numerous collections examined both here and in

Lewis and Bowman (1981), the morphology of C. whitei is by far the

more prevalent and typical of the species. Second, the type-locality of C.

whitei, Cricket Cave, is a well known but remote locality that is cur-

rently unthreatened by man. In contrast, the type-locality of C. meiste-

rae in Johnson County, Illinois, lies adjacent to an active limestone

quarry. Although still some distance from the cave, this quarry has

already consumed one cave (Bretz and Harris 1961), and local residents

believe that the quarry operations may eventually consume other sec-

tions of White Hill.

70

Julian J. Lewis

Fig. 2. Palmar margin of propodus of male first pereopods of Caecidotea bicren-

ata whitei from Roaring River Shrimp Pools, Mammoth Cave, KY: (a) 8 mm

individual, C. meisterae form; (b) 6 mm individual, C. whitei form; (c) 4 mm

individual, immature.

Caecidotea whitei is redefined as a subspecies of Caecidotea bicre-

nata due to the slight morphological differences that distinguish them,

and the lack of dispersal barriers that might provide reproductive isola-

tion along the contact between C. whitei and C. bicrenata. Apparently

the ranges of C. b. bicrenata and C. b. whitei contracted at some time in

the past, then range expansion followed. If this occurred, some secon-

dary contact phenomenon might be expected, either hybridization or

character displacement. Along the contact between the two subspecies,

occasional populations occur in which the lateral process of the male

second pleopod endopod tip (Fig. 3) is either vestigial or absent. Con-

sidering the close morphological similarity of the two subspecies, it

seems more likely that this phenomenon is caused by breakdown of any

isolating mechanisms developed, rather than reinforcement. However,

without experimentally crossbreeding individuals taken from popula-

tions of each subspecies it is impossible to say with certainty that

hybridization is occurring. Furthermore, the presence of specimens lack-

ing the lateral process in one stream in Mammoth Cave (Mystic River)

complicates the situation. Mammoth Cave is relatively distant from

Systematics of Troglobitic Caecidotea

71

Fig. 3. Tip elements of male second pleopod endopods: (a) Caecidotea bicren-

ata whitei, Carter Cave, Jackson Co., TN; (b) Caecidotea bicrenata whitei, Mill

Hole, Barren Co., KY; (c) Caecidotea bicrenata bicrenata, B&J Cave, Morgan

Co., AL; (d) Caecidotea bicrenata, Mystic River, Mammoth Cave, Edmonson

Co., KY; (e) Caecidotea bicrenata, Dunbar Cave, Montgomery Co., TN; (0

Caecidotea bicrenata, Columbia Caverns, Dickson Co., TN.

72

Julian J. Lewis

Fig. 4. Pleopod 1 of male Caecidotea: (a) Caecidotea bicrenata, Mystic River,

Mammoth Cave, Edmonson Co., KY; (b) Caecidotea bicrenata whitei, Mill

Hole, Barren Co., KY; (c) Caecidotea bicrenata bicrenata, Beech Spring Cave,

Marshall Co., AL.

other populations exhibiting this modification of the second pleopod,

such that if hybridization is occurring it is across a rather wide geogra-

phic range. In isopods from other rivers flowing through the Mammoth

Cave System that have been sampled, the lateral process is present. In

the absence of the lateral process, critical in identifying specimens to the

subspecific level, these populations are here identified only as Caecido-

tea bicrenata.

Caecidotea bicrenata (Steeves)

Material examined. — TENNESSEE: Dickson Co., stream in

Columbia Caverns, 2 mi. SW VanLeer, 22 June 1957, L. Hubricht,

4$$, 3$$. Macon Co., Ann White Cave, 6 mi. W Lafayette, 19 Apr

1958, L. Hubricht, 9$$, 7§9. Montgomery Co., Dunbar Cave, 1.5 mi.

S St. Bethlehem, 15 June 1957, L. Hubricht, 13<3<3, 1$. KENTUCKY:

Edmonson Co. , Mammoth Cave National Park, Mammoth Cave, Mys-

tic River near Mystic River tributary, 27 Dec 1981, J. Lewis, T. Lewis,

J. Eckstein, T. Leitheuser, 5<5<?.

Systematics of Troglobitic Caecidotea

78

Fig. 5. Palmar margin of propodus of male first pereopods: (a) Caecidotea

bicrenata whitei, Mill Hole, Barren Co., KY; (b)Caecidotea bicrenata bicrenata,

Beech Spring Cave, Marshall Co., AL; (c) Caecidotea bicrenata bicrenata, B&J

Cave, Morgan Co., AL.

ACKNOWLEDGMENTS. — I would like to thank Drs. Thomas

E. Bowman, John R. Holsinger, and Gerald A. Cole for reading the

manuscript. Much of the material examined was provided by Dr. Bow-

man from the National Museum of Natural History, Smithsonian Insti-

tution, and by Dr. John E. Cooper, North Carolina State Museum of

Natural History. The Cave Research Foundation provided use of its

field station in Mammoth Cave National Park, and many members of

CRF aided in the collection of asellids in Mammoth Cave, especially

Mrs. Teresa M. Lewis and Mr. James Eckstein. I also thank Mr. Robert

Deskins, Mr. James Wiggins and the rest of the staff at Mammoth Cave

National Park for their cooperation in expediting my research at

Mammoth Cave. Funds were provided by a grant from the Research

Advisory Committee and the 1980 Stone Graduate Research Award of

the National Speleological Society, and a grant from the Graduate

School of the University of Louisville. This paper is Contribution #207

(New Series) of the Department of Biology, University of Louisville.

74 Julian J. Lewis

LITERATURE CITED

Barr, Thomas C, Jr. 1967. Ecological studies in the Mammoth Cave System of

Kentucky, I: The biota. Int. J. Sp61e*ol. 5:147-204.

Bresson, J. 1955. Aselles de sources et de grottes D'Eurasie et D'Amerique du

Nord. Arch. Zool. Exp. Gen. 92(2):45-77.

Bretz, J Harlan, and S. Harris. 1961. Caves of Illinois. 111. State Geol. Surv.

Rep. Invest. 215. 87 pp.

Chappuis, P.A. 1957. Un Asellide nouveau de l'Amerique du Nord. Notes

Biospeol. 12:37-43.

Cooper, John E., and M.R. Cooper. 1968. Cave associated herpetozoa II:

Salamanders of the genus Gyrinophilus in Alabama Caves. NSS Bull.

30(2): 19-24

Fleming, Laurence E. 1972a. The evolution of the eastern North American

isopods of the genus Asellus (Crustacea: Asellidae). Int. J. Speleol. 4:

221-256.

1972b. Four new species of troglobitic Asellids (Crustacea:Isopoda)

from the United States. Proc. Biol. Soc. Wash. S¥(57):489-500.

1973. The evolution of the eastern North American isopods of the

genus Asellus (Crustacea: Asellidae). Int. J. Sp61e*ol. 5:283-310.

Henry, Jean-Paul, and G. Magniez. 1970. Contribution a la systematique des

Asellides (Crustacea Isopoda). Ann. Speleol. 25(2):335-367.

Lewis, Julian J. 1981a. Distribution of aquatic isopods in Mammoth Cave.

Cave Research Foundation 1979 Annual Report:21.

1981b. The subterranean Caecidotea of the Interior Low Plateaus.

Proc. Eighth Int. Congress Speleol. 1:234-236.

, and T.E. Bowman. 1981. The subterranean Asellids (Caecidotea) of

Illinois (Crustacea:Isopoda:Asellidae). Smithson. Contrib. Zool. 335 :1 -66.

, and Teresa M. Lewis. 1980. The distribution and ecology of two

species of subterranean Caecidotea in Mammoth Cave National Park. Cave

Research Foundation 1980 Annual Report:23-27.

Peck, Stewart B., and Julian J. Lewis. Zoogeography and evolution of the

subterranean invertebrate faunas of Illinois and southeastern Missouri.

NSS Bull. 40(2):39-63.

Stafford, B.E. 1911. A new subterranean freshwater isopod. Pomona J.

Entomol. 5:572-575.

Steeves, Harrison R., III. 1963. The troglobitic Asellids of the United States:

The Stygius Group. Am. Midi. Nat. 59(2):470-481.

1964. Asellus bicrenatus, a synonym of A. alabamensis. Am. Midi.

Nat. 7/(2):503-504.

1966. Evolutionary aspects of the troglobitic Asellids of the United

States: The Hobbsi, Stygius and Cannulus Groups. Am. Midi. Nat.,

75(2):392-403.

. 1969. The origin and affinities of the troglobitic Asellids of the

southern Appalachians, pp. 51-65 in Perry 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.

Accepted 12 July 1982

Aboriginal and Modern Freshwater Mussel

Assemblages (Pelecypoda: Unionidae)

from the Chickamauga Reservoir, Tennessee

Paul W. Parmalee and Walter E. Klippel

Department of Anthropology,

University of Tennessee,

Knoxville, Tennessee 37996

AND

Arthur E. Bogan

Department of Malacology,

Academy of Natural Sciences, Philadelphia, Pennsylvania 19103

ABSTRACT.— From December 1976 through March 1979, ca. 40,500

mussel valves were collected from 28 aboriginal shell middens in the

Chickamauga Reservoir (TRM 495-528), Rhea and Meigs counties,

Tennessee. Approximately 46 species were identified in 27,875 speci-

mens from 14 of these Middle Woodland through Mississippian com-

ponent sites. Valves of Pleurobema spp., Elliptio spp., Actinonaias

ligamentina, and Dromus dromas comprised 75% of the total. About

28 species represented in the shell middens are now either extinct, or

are extirpated from the reservoir. Five species have invaded and

become established in the reservoir, and four others, rare in prehistoric

times, have greatly increased their range and abundance since

impoundment.

INTRODUCTION

Accumulations, often huge, of freshwater mussel valves and snail

shells along the banks of the Tennessee and other major rivers in south-

eastern United States attest to the degree aboriginal man used this easily

accessible food resource. Although Parmalee and Klippel (1974) showed

that this subsistence resource is relatively low in food energy compared

with most other meat animals, and that it was exploited as a supplement

rather than a staple, mollusks were nevertheless taken in great quantities

and did provide an abundant food supplement. The prehistoric naiad

(freshwater mussel) fauna of the Tennessee and Cumberland River sys-

tems was one of the richest in the world and consisted of at least 90

species. Although the Indian harvested vast quantities of mussels over a

period of thousands of years, there is no evidence to suggest that this

activity brought about the extinction of even one species or was detri-

mental to naiad populations in general. However, the effect of dam con-

struction, impoundment, siltation, and overharvesting by commercial

shellers over the last 75 years is a different matter (Isom 1969).

Relatively few detailed ecological and taxonomic studies involving

Brimleyana No. 8:75-90. December 1982 75

76 Paul W. Parmalee, Walter E. Klippel, Arthur E. Bogan

shell assemblages from aboriginal middens that are so prevalent along

the Tennessee River have been undertaken. One notable exception was

the analysis of over 100,000 mussel valves studied by Morrison (1942)

from Archaic and Woodland shell middens that were located in what is

now the Pickwick Landing Dam reservoir in northwestern Alabama.

The more recent study by Warren (1975) involving analysis of 60,350

naiad specimens recovered from the Widows Creek site in northeastern

Alabama provided an effective account of the species assemblage, the

limnological and other habitat conditions reflected by this assemblage,

and the extent of use of the naiads by the Woodland inhabitants of the

site. Although extensive archaeological field work was carried out dur-

ing 1937-1939 at Hiwassee Island (Lewis and Kneberg 1946), now within

the Chickamauga Reservoir, and extensive shell middens were encoun-

tered, no study of this material was made and samples of unmodified

shell were not retained.

Our interest in sampling a series of aboriginal shell middens that

still remain along the shores of the middle and upper Chickamauga

Reservoir was based on several factors: (1) no previous studies of pre-

historic shell middens located in this section of the Tennessee River had

been undertaken to assess the naiad species formerly present and their

relative abundance at these sites; (2) since impoundment, much of this

nonrenewable archaeological resource has been inundated and that

which remains is subject to continual destruction through wave action,

erosion, digging activities of artifact collectors, and general weathering;

(3) these shell middens provided an opportunity to evaluate similarities

and dissimilarities among cultural groups who once lived along this sec-

tion of the river as to how extensively they exploited this food resource;

and (4) recent surveys of naiad populations in the Chickamauga Reser-

voir, compared with shell samples collected from aboriginal middens,

could provide significant data relative to changes in species assemblages

during the last 2000 years.

MATERIALS AND METHODS

A total of 28 shell middens located along an approximately 50 km

stretch of the middle and upper Chickamauga Reservoir (TRM 495-528)

was sampled over a 2x/i year period from December 1976 to March

1979. Some middens were extensive (Fig. 1), and shell that had eroded

from the banks covered the beach (observable only during winter low

water levels) for a distance of 300 m or more at certain sites. Most of

the middens reflect an occupation by Late Woodland (ca. AD 600-1000)

cultural groups; however, at least one single component Middle Wood-

land (ca. AD 1-600) site and two Mississippian component sites (ca. AD

1000-1600) were encountered. Lithic and ceramic samples were collected

Chickamauga Reservoir Mussels

77

Fig. 1. Top: Aboriginal shell midden eroding from the bank of the Tennessee

River (Chickamauga Reservoir). Bottom: Shell scattered by wave action along

the beach below the same midden.

78 Paul W. Parmalee, Walter E. Klippel, Arthur E. Bogan

from the surface of the eroding shell banks and beaches, but a special

effort was made to recover all such cultural material found within the

portions of undisturbed shell lenses that were removed for this study.

Since many of the deposits were comprised of residue from several cul-

tural groups that spanned a considerable period of time, only those

sherds and other artifacts found in direct association with the selected

samples were used to define the temporal periods involved.

Considerable variation in midden size was apparent, with some

containing only a few hundred valves and others literally hundreds of

thousands. Most also contained shells of freshwater gastropods. Because

of the typically compact nature of the shell lenses, we could remove only

a small part of the shell comprising the midden and yet obtain a large

but relatively unbiased sample of the species assemblage. All samples

were returned to the Department of Anthropology, University of

Tennessee, Knoxville, where the shell was washed and determinations of

species were made by comparisons with fresh specimens in the Zooar-

chaeology Section mollusk collection. Approximately 40,500 freshwater

mussel valves, representing about 50 species, were identified from the

samples collected during this study. For this report we evaluated collec-

tions from 15 of the sites (Fig. 2) for which the temporal assessment was

most exact; samples from these 14 sites consisted of 27,875 valves. All

specimens recovered during this study are housed in the Zooarchaeol-

ogy Section, Department of Anthropology, University of Tennessee,

Knoxville.

RESULTS AND DISCUSSION

Present Status of Naiad Fauna in Middle and Upper Chickamauga

Reservoir

During the past two decades several investigators attempted to

determine the status of freshwater mussel populations inhabiting the

middle and upper Chickamauga Reservoir from ca. TRM 495 to TRM

529 (Scruggs 1960; Isom 1969; Pardue 1981). The study by Scruggs

(1960) was undertaken primarily to determine the abundance, effects of

shelling operations, and potential for recovery of commercially valuable

species, especially Pleurobema cor datum (Raf. 1820), on specific beds in

the Tennessee River (Wheeler and Chickamauga reservoirs). Although

also concerned with similar aspects of naiad ecology and economics, the

comprehensive study by Isom (1969) provided a more complete assess-

ment of the total mussel fauna assemblage and evaluated the factors

that brought about changes in extant species and population abundance

since impoundment.

On the basis of these studies approximately 20 species (synonymiz-

ing Anodonta corpulenta Cooper 1834 with Anodonta grandis Say

Chickamauga Reservoir Mussels

79

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ex

E

03

c

"5b

c

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u

D

DO

80 Paul W. Parmalee, Walter E. Klippel, Arthur E. Bogan

1829) were found still inhabiting the middle and upper Chickamauga

Reservoir (Pardue 1981). In addition, we collected specimens of three

other species (Anodonta suborbiculata Say 1831, Anodonta imbecillis

Say 1829, Leptodea laevissima Lea 1830) in the shallow mud-bottom

bays at the mouth of the Hiwassee River. These results reflect the extir-

pation of approximately half of the naiad fauna known to have inhab-

ited this part of the Tennessee River in prehistoric times (as evidenced

from species represented in aboriginal shell middens) and in the early

20th century prior to impoundment (Ortmann 1918). Isom (1969)

expressed the view that sedimentation, one of the consequences of

impoundment resulting from a slowed current that prevents the sub-

strate from being swept clean, and overharvesting by commercial shellers,

are the two major factors in limiting mussel population growth and

expansion in Tennessee River reservoirs.

Although populations of several species that are typical inhabitants

of shallow mud-bottom bays and shorelines of the reservoir (e.g. Ano-

donta spp., Leptodea sp., Potamilus spp.) may become locally large,

communities in the main river channel are composed primarily of Ellip-

tio crassidens (Lamarck 1819), Pleurobema cordatum, Quadrula pustu-

losa (Lea 1831), and Ellipsaria lineolata (Raf. 1820). Several species still

present in the reservoir are rare and their populations appear to consist

of old nonbreeding adults: these are Plethobasus cooperianus (Lea

1834), Cyprogenia irrorata (Lea 1829), Obovaria olivaria (Raf. 1820),

and Dromus dromas (Lea 1834). Until additional extensive sampling is

carried out, the status of several other species, including Lampsilis

orbiculata (Hildreth 1828), Plethobasus cyphyus (Raf. 1820), and Acti-

nonaias ligamentina (Lamarck 1819), remains uncertain.

Prehistoric Mussel Assemblages Based on Midden Samples

Table 1 shows the mussel species represented by 27,875 valves col-

lected from one Middle Woodland, eight Late Woodland, two Missis-

sippian, one Middle/ Late Woodland, and two Late Woodland/ Missis-

sippian sites along the middle and upper Chickamauga Reservoir, Meigs

(MG) and Rhea (RH) counties, Tennessee. Although Archaic (8000-

1000 BC) and Early Woodland (1000 BC-AD1) shell middens occur in

considerable numbers and are often of great magnitude in the middle

(Alabama) and lower (West Tennessee) reaches of the Tennessee, none

have been located in the section of the river between Knoxville and

Chattanooga. Shell middens left by peoples of the Early Woodland cul-

ture appear to be rare in the upper Tennessee River; we were unable to

locate any Early Woodland sites, and found only one single component

Middle Woodland site (40MG51) during our survey. For whatever rea-

sons), therefore, it appears that aboriginal man did not begin to harvest

freshwater mussels as a food resource in any quantity in the upper Ten-

Chickamauga Reservoir Mussels 81

nessee River until ca. AD 100.

In evaluating similarities and differences among mussel species

assemblages collected by people of various cultural groups, factors such

as individual preference or selection (e.g. large vs. small specimens),

location of the habitation site in relation to available mussel beds, sea-

son of the year collected, and probable differences in species comprising

the beds along a section of river, must be taken into account. On the

basis of samples collected during this study it is apparent that certain

species, such as A. ligamentina and D. dromas, or an assemblage of

closely related forms (e.g. Pleurobema spp.), were common and gener-

ally distributed throughout this section of the river during the past 2000

years. Considering the total sample from all cultural units, valves of

Pleurobema spp., A. ligamentina, and D. dromas comprised 75% of all

valves collected.

Valves of D. dromas varied from 24% in the Middle Woodland site

sample to 45% in the multicomponent Middle Woodland/ Mississippian

sites, averaging 35% of the total sample. Of the 60,350 valves studied by

Warren (1975) from the Widows Creek site shell midden (Tennessee

River, northeastern Alabama: Early Woodland to Late Middle Wood-

land), D. dromas comprised 23% of all shells. Of the seven shell mounds

sampled in the Pickwick Landing Basin (Tennessee River, northwestern

Alabama: Archaic, Woodland), D. dromas was "One of the most

abundant species in these shell deposits . . . and made up a major part

of the total mussel fauna gathered for food" (Morrison 1942:359).

Although a form of D. dromas is still fairly common in unimpounded

reaches of the upper Powell and Clinch rivers in East Tennessee, the

large river form appears to be surviving at only one restricted locality in

the Tennessee River (Chickamauga Reservoir) (Pardue 1981).

Ortmann (1918:556) stated that E. crassidens is "In the Tennessee

below Knoxville, and down to Chattanooga, it is extremely abundant";

it is reported to now be "by far the most abundant species in the upper

Tennessee River" (Pardue 1981:44). Valves of this large, big-river mussel

occurred in all middens sampled and accounted for about 6% of the

total shell sample. Specimens of the related species, Elliptio dilatatus

(Raf. 1820), also occurred at all sites but had been collected in greater

numbers (ca. 12% of the total) than E. crassidens, possibly because it

was more numerous in shallower riffles and shoals and consequently

more accessible to the Indian. Today, populations of E. dilatatus have

been greatly reduced in the Tennessee River, probably as a result of

impoundment and its concomitant adverse factors. Actinonaias liga-

mentina is another species that occurred consistently in all middens;

valves of this species varied from ca. 4.5% of the total from Late Wood-

land/Mississippian sites to nearly 9.5% at the Middle Woodland site

(average for all sites, 7.5%). At the Widows Creek site (Warren 1975) its

82

Paul W. Parmalee, Walter E. Klippel, Arthur E. Bogan

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84 Paul W. Parmalee, Walter E. Klippel, Arthur E. Bogan

valves comprised 8% of the total. Although Scruggs (1960) reported A.

ligamentina as still present in Chickamauga and Kentucky reservoirs,

Pardue (1981) failed to recover it during mussel surveys of Watts Bar,

Chickamauga, and Nickajack reservoirs. With the possible exception of

a few relic individuals, this species appears to have been extirpated from

all stretches of the middle and upper Tennessee River.

At least four and possibly five species (or forms, depending upon

the taxonomic approach) of Pleurobema were represented in the mid-

dens, and combined their valves comprised nearly 16% of the total. Of

the Pleurobema complex, valves of P. cor datum and P. plenum (Lea

1840), both of which today are usually found inhabiting large rivers at

depths of 3 to 6 m, were the most numerous. Pleurobema pyramidatum

(Lea 1834), another medium-to-large river deep water species, ranked

third in abundance within this complex. Of interest is the fact that com-

bined, all species of Pleurobema comprised approximately 24% of the

shells recovered at the two Mississippian sites, 16% of those at the Late

Woodland sites, and 23% of those at the Middle Woodland site. At

Widows Creek, valves of Pleurobema amounted to about 13% of the

total (Warren 1975). Continued harvesting of these species by the Indian

over a long period of time attests to their former abundance and proba-

ble habitation of the more shallow shoal areas of the river. Pleurobema

cordatum appears to be the only species of this complex still inhabiting

the upper Tennessee River.

Ortmann (1918, 1925) considered Pleurobema clava (Lamarck

1819) a species of the Ohio River drainage, replaced in the upper Ten-

nessee River system by the form or species Pleurobema oviforme (Con-

rad 1834). The relationship between these two is still not clear, but they

do not appear to intergrade; Warren (1975) recorded both from the

Widows Creek site. Although specimens from the Chickamauga Reser-

voir middens should theoretically tend more toward P. oviforme, and

admittedly some were questionable, shells of this complex more closely

approached in appearance those of P. clava than those of P. oviforme

from the Duck River and tributary streams of the Tennessee River in

the northeastern part of the state. In any case, P. clava appears to have

been a part of the upper Tennessee River mussel assemblage since at

least Middle Woodland times, but it evidently was uncommon (valves

amounted to < 1% of the total). Lexingtonia dolabelloides (Lea 1840), a

"Cumberlandian" species once locally common in the middle and upper

Tennessee River and its tributaries, has been extirpated throughout

most of its range. Although valves of this species amounted to < 1% of

the total, they occurred in all middens examined.

Shells of four species of Quadrula occurred in the middens, but

combined they totaled only 388, less than 2% of all specimens collected.

Valves of Q. metanevra (Raf. 1820) were the most numerous, followed

Chickamauga Reservoir Mussels 85

by those of Q. pustulosa; both species still occur as viable populations

in the Chickamauga Reservoir, apparently having adapted to

impoundment conditions. Both Q. cylindrica (1817) and Q. intermedia

(Conrad 1836) are apparently now extirpated in the upper Tennessee

River; judging by the relatively few specimens obtained in the middens,

both were uncommon in prehistoric times. Today, limited populations

of Q. intermedia, a "Cumberlandian" species that typically inhabits

small-to-medium size rivers, occur in the upper Powell and Clinch rivers

of northeast Tennesseee and in the Duck River of Middle Tennessee. In

prehistoric times, however, the species occurred throughout the upper

and middle Tennessee River; Morrison (1942) reported it present but

not common in Pickwick Basin shell mounds, northwestern Alabama.

Amblema plicata (Say 1817), a species now relatively common in the

Chickamauga Reservoir (Pardue 1981), appears to have been uncom-

mon to rare in prehistoric times (valves comprised < 0.5% of the total).

The combined number of valves (477) of the three species of Pleth-

obasus constituted less than 2% of the total. However, valves occurred

in all cultural components, but the low numbers apparently reflect their

limited abundance in the upper Tennessee during the last 2000 years. At

the Widows Creek site they comprised less than 3% of the total (Warren

1975), and Morrison (1942) reported only a few specimens of P.

cyphyus and P. cicatricosus (Say 1829) and none of P. cooperianus

from the Pickwick Basin middens. Ortmann (1925:338) commented that

P. cyphyus occurred sparingly in the main river in the lower Tennessee

drainage, while P. cooperianus "goes into the upper Tennessee up to the

Knoxville region, but it is rare there." Although P. cicatricosus was

recorded from below Wilson Dam by Stansbery (1964), it is now near-

ing extinction and is represented by a few old relic individuals. Based on

a few valves recovered from shelter's cull piles (taken in the early 1970s),

P. cyphyus appears to continue to survive in the Chickamauga Reser-

voir. One valve of P. cooperianus was also found in a cull pile.

Although Scruggs (1960) recorded it from the Washington Ferry area

(TRM 518), Pardue (1981) failed to find it anywhere in the middle or

upper reaches of the Chickamauga Reservoir.

One of the most interesting naiad assemblages represented in the

Chickamauga Reservoir middens consists of 12 species belonging to the

genus Plagiola (-Dysnomia-Epioblasma) (Johnson 1978). Six of these

(arcaeformis Lea \S34,flexuosa Raf. 1820, haysiana Lea 1834, propin-

qua Lea 1857, stewardsoni Lea 1852, turgidula Lea 1858) are now

extinct. Five others (capsaeformis Lea \834, florentina Lea 1834, torul-

osa Raf. 1820, interrupta Lea 1831, obliquata Raf. 1820) are either

extremely rare, or their former ranges have been reduced to a few local-

ized habitats, or both. All of those species inhabit (or did inhabit) small-

to-medium size rivers, or the shallow riffle and shoal areas of large riv-

86 Paul W. Parmalee, Walter E. Klippel, Arthur E. Bogan

ers such as the Tennessee having a sandy-gravel substrate with rapid

currents (Stansbery 1971). Although a few individuals of some of these

species (e.g. interrupta, obliquata) may continue to survive after im-

poundment (Parmalee et al. 1980), they do not or cannot propagate,

and the species within the affected sections of the river eventually die

out. Only P. triquetra still occurs in viable populations in upper East

Tennessee and locally in the Midwest and Great Lakes states.

All shells of the 12 species of Plagiola represented in the middens

amounted to approximately 8% of the total sample; valves of arcaeform-

is, propinqua, and torulosa comprised 6.5% of this total. Two distinct

forms of P. torulosa have been recognized (Ortmann 1918): P. t. torul-

osa, with the more inflated shell with a radial row of prominent knobs

across the middle, occurred from about Knoxville downstream; the

form P. t. gubernaculum, with a more compressed shell and poorly devel-

oped or wanting knobs, occurred in streams above Knoxville. Although

valves of the form torulosa predominate in the Chickamauga Reservoir

middens, a few approach the typical upstream form gubernaculum.

In a recent taxonomic revision of this complex, Johnson (1978)

synonymized the species P. lewisi with P. flexuosa, although not all

malacologists (e.g. Stansbery 1971) agree with this approach. Valves of

these two species, or the species P. flexuosa, occurred only sparingly in

the middens, which may reflect the former rarity of P. flexuosa in the

Tennessee. Except for arcaeformis, propinqua, and torulosa, the same

may be said of the other species of Plagiola. Of interest is the fact that

only five valves of P. triquetra, the one species still locally common and

widespread in streams of eastern Tennessee and the Great Lakes area,

were recovered in the middens. Warren (1975) listed only four valves of

this species in a sample of 60,350 valves from the Widows Creek site;

Morrison (1942) reported it absent from the Pickwick Basin mounds.

This rather ubiquitous inhabitant of numerous smaller streams and

tributaries of the Tennessee apparently did not become well established

in the shoals and riffles of the main river. This also seems to have been

the case with capsaeformis, turgidula, interrupta, and florentina, all spe-

cies of small-to-medium size rivers. It is of further interest that all shells

of Plagiola spp. comprised 1% of the valves from the Mississippian

component sites, but averaged about 6% of the totals from the Middle

and Late Woodland sites.

Fusconaia subrotunda (Lea 1831) occurred in all middens and var-

ied from about 3.5% of all valves in the Late Woodland and Mississip-

pian sites to 7.5% in the multicomponent Middle/ Late Woodland sites.

Pardue (1981) found very few specimens of this mussel in the Chicka-

mauga and Watts Bar reservoirs and we doubt that it will ever return to

a semblance of its former abundance. Cyclonaias tuber culata (Raf.

1820), however, another ubiquitous although slightly less common shell

Chickamauga Reservoir Mussels 87

than F. subrotunda in the middens, is presently a common component

of the Chickamauga Reservoir naiad fauna. With reference to Obovaria

retusa (Lamarck 1819), Ortmann (1925:348) commented that "In the

upper Tennessee it goes to Knoxville region, but is very rare." With the

exception of one Late Woodland site, valves of this distinct species were

recovered from all sites, but totaled < 2% for all samples. As a species it

appears on the verge of extinction; a few old relic specimens are occa-

sionally taken below Pickwick Dam and in the middle Cumberland

River, Tennessee (Parmalee and Klippel 1982). It is now extirpated in

the middle and upper Tennessee River.

Valves of a few species typical of small-to-medium size rivers, such

as Fusconaia barnesiana (Lea 1838), Lampsilis fasciola (Raf. 1820),

Lemiox rimosus (Raf. 1831), Obovaria subrotunda (Raf. 1820), and

Ptychobranchus subtentum (Say 1825), occurred sparingly in some of

the middens. Combined shells of these species comprised < 1% of all

samples. Only eight valves of Ligumia recta (Lamarck 1819) were reco-

vered (six in 40MG50, a Mississippian site). Although fairly common

and widespread throughout the Tennessee River and its major tributar-

ies today, this species was apparently a rare shell in the middle and

upper Tennessee during aboriginal times. Cyprogenia irrorata (Lea

1829) is another species once widely distributed throughout the Ohio,

Cumberland, and upper Tennessee River drainages. It occurred in most

of the Chickamauga Reservoir middens, but sparingly. Warren (1975)

identified 127 valves of this species (only 0.39% of the total) form the

Widows Creek site. Cyprogenia irrorata, like most species adapted to

riffle and shoal habitats in the Tennessee River, has been extirpated or

reduced to relic populations of old individuals in river sections affected

by impoundment.

Recent Adaptations to Impoundment

Impoundment generally produces a reduced rate of current flow,

increased water depth with often daily or seasonal extremes in water

level and temperature fluctuations, and siltation. The adverse effects

these conditions can exert in extirpating a species or reducing it to a few

nonpropagating individuals have been discussed. The fate of at least a

dozen species within the genus Plagiola provides ample documentation.

On the other hand, changes in the aquatic habitat brought about by

impoundment and the establishment of a "river-lake" has proved favor-

able to several species. Bates (1962) discussed this phenomenon for the

lower Tennessee River (Kentucky Reservoir). Three species of Anodonta

— A. grandis (Say 1829), A. imbecillus (Say 1829), and A. suborbicu-

lata (Say 1831) — have now become well established locally in the shal-

low mud-bottom bays and/ or shorelines throughout most of the reser-

voirs. No valves of Anodonta were recovered in the Chickamauga

88 Paul W. Parmalee, Walter E. Klippel, Arthur E. Bogan

Reservoir middens or at Widows Creek (Warren 1975), and Morrison

(1942:359) listed A. grandis along with a few other species as being

"present in small numbers only, if represented by more than one speci-

men each." Leptodea fragilis (Raf. 1820) often reaches its greatest

abundance in habitat similar to that preferred by Anodonta. It now

occurs locally throughout the Tennessee River although, as evidenced

by recovery of only one valve of 60,350 collected at Widows Creek

(Warren 1975) and none from the Pickwick Basin and Chickamauga

Reservoir middens, it was extremely rare in prehistoric times. A single

valve of Potamilus alatus (Say 1817) occurred in the Middle Woodland

site midden and two shells in the multicomponent Middle/ Late Wood-

land site. It was not present at the Widows Creek site (Warren 1975),

and Morrison (1942) identified only "a few individuals" from the Pick-

wick Basin mounds. Potamilus alatus is another species that has adapted

to and flourishes in Tennessee River reservoirs.

Viable populations of four other species now inhabiting the middle

and upper Tennessee River provide an interesting contrast with their

apparent prehistoric abundance and distribution. They are Ellipsaria

lineolata (Raf. 1820), Tritogonia verrucosa (Raf. 1820), Obliquaria

reflexa (Raf. 1820), and Megalonaias gigantea (Barnes 1823). All four

appear to have their center of origin in the Interior Basin or Mississippi

River drainages. Ellipsaria lineolata is presently relatively common in

the Chickamauga and Nickajack reservoirs (Pardue 1981). Ortmann

(1925) commented that it was rare in the upper Tennessee, and the pauc-

ity of shells from Pickwick Basin ("found only as scattered individuals

in two of the mounds"; Morrison 1942:361) and Widows Creek (one

valve), and no specimens from the Chickamauga Reservoir middens,

attest to its rarity in prehistoric times. Tritogonia verrucosa is another

species that has become established in the Chickamauga Reservoir,

apparently in modern times, as no shells of this mussel were found in

the midden samples we examined and it was not encountered by Warren

(1975) at the Widows Creek site or by Morrison (1942) in Pickwick

Basin mounds.

Pardue (1981) reported taking M. gigantea at one station in the

Nickajack Reservoir at Chattanooga and from three localities in the

Chickamauga Reservoir. Like T. verrucosa, M. gigantea has become

established in the middle and upper reservoirs of the Tennessee River in

recent times. No specimens were encountered in the Pickwick Basin

mounds, at Widows Creek, or in the Chickamauga Reservoir middens.

Megalonaias gigantea may well continue to extend its range and in-

crease in abundance, since it seems to adapt well to a river-lake habitat.

For example, M. gigantea is the most abundant species in the im-

pounded stretches of the middle Cumberland River, Tennessee (Parma-

lee et al. 1980). The last of these four river species that has thrived and

Chickamauga Reservoir Mussels 89

increased in abundance as a result of impoundment of the Tennessee

River is O. reflexa. Although Morrison (1942) did not encounter O.

reflexa in the Pickwick Basin mounds, it was present during prehistoric

times in the upper Tennessee River as evidenced by three valves from

the Widows Creek site (Warren 1975) and two shells from the Chicka-

mauga Reservoir middens (one valve from each of two multicomponent

sites, 40RH18 and 40RH39, not included in this study). It is a typical

river species that can adapt to a lake environment (Klippel and Parma-

lee 1979).

Data recovered from archaeological sites along the banks of the

Chickamauga Reservoir have provided information that greatly enhan-

ces our understanding of the diachronic variability in the mussel fauna

of the Upper Tennessee River. Pardue (1981), by considering presence

/absence of species recovered from the upper Tennessee between 1918

and 1980, has clearly delineated the magnitude of change that has taken

place in less than a century. Our data, recovered from archaeological

contexts, suggests that comparatively little change took place over as

much as two millennia preceding EuroAmerican settlement along this

section of the river. Although the invasion and establishment of "new"

species in the impounded stretches of the Tennessee River is noteworthy

and may provide a potentially valuable commerical resource, these

latecomers fail to equal the loss of so many of the river's endemic spe-

cies that were once a part of one of the richest naiad faunas in the

world.

ACKNOWLEDGMENTS. — We would like to express our thanks

to Betty W. Creech for typing the manuscript and to Terry Faulkner for

preparing Figure 2. Appreciation is extended to Tennessee Valley

Authority officials, Division of Property and Services, for granting

permission to sample aboriginal shell middens on property easements

held by that agency.

LITERATURE CITED

Bates, John M. 1962. The impact of impoundment on the mussel fauna of

Kentucky Reservoir, Tennessee River. Am. Midi. Nat. tf#(l):232-236.

Isom, Billy G. 1969. The mussel resource of the Tennessee River. Malacologia

7(2-3):397-425.

Johnson, Richard I. 1978. Systematics and zoogeography of Plagiola (-Dysno-

mia-Epioblasma), an almost extinct genus of freshwater mussels (Bivalvia:

Unionidae) from middle North America. Bull. Mus. Comp. Zool. Harv.

Univ. /45(6):239-320.

Klippel, Walter E., and P.W. Parmalee. 1979. The naiad fauna of Lake

Springfield, Illinois: an assessment after two decades. Nautilus 94

(4): 189-197.

Lewis, Thomas M.N., and M. Kneberg. 1946. Hiwassee Island. Univ. Tenn.

Press, Knoxville. 188 pp.

90 Paul W. Parmalee, Walter E. Klippel, Arthur E. Bogan

Morrison, Joseph P.E. 1942. Preliminary report on mollusks found in the shell

mounds of the Pickwick Landing Basin in the Tennessee River valley, pp.

377-392 in William S. Webb and David L. DeJarnette. An archaeological

survey of Pickwick Basin in the adjacent portions of the states of Alabama,

Mississippi, and Tennessee. Bur. Am. Ethnology Bull. 129. 536 pp.

Ortmann, Arnold E. 1918. The nayades (freshwater mussels) of the upper

Tennessee drainage. With notes on synonymy and distribution. Proc. Am.

Philos. Soc. LVII:521-626.

. 1925. The naiad-fauna of the Tennessee River system below

Walden Gorge. Am. Midi. Nat. 9(7):32 1-372.

Pardue, W. Jeffrey. 1981. A survey of the mussels (Unionidae) of the upper

Tennessee River — 1978. Sterkiana 77:41-51.

Parmalee, Paul W., and W.E. Klippel. 1974. Freshwater mussels as a prehistoric

food resource. Am. Antiquity 39(3):42 1-434.

, and 1982. A relic population of Obovaria retusa in the

middle Cumberland River, Tennessee. Nautilus 96(1 ):30-32.

, and A.E. Bogan. 1980. Notes on the prehistoric and pres-

ent status of the naiad fauna of the middle Cumberland River, Smith

County, Tennessee. Nautilus 94(3): 93-105.

Scruggs, George D., Jr. 1960. Status of fresh-water mussel stocks in the

Tennessee River. USFWS Spec. Sci. Rep. Fish. No. 370. 41 pp.

Stansbery, David H. 1964. The mussel (muscle) shoals of the Tennessee River

revisited. Am. Malacol. Union Inc. Annu. Rep. 1964:25-28.

. 1971. Rare and endangered freshwater mollusks in eastern United

States, pp. 5-18 in S.E. Jorgensen and R.W. Sharp (eds.) Rare and

Endangered Mollusks (Naiads) of the U.S. USFWS Region 3, Twin Cities,

MN. 79 pp.

Warren, Robert E. 1975. Prehistoric unionacean (freshwater mussel) utilization

at the Widows Creek site (IJA305), northeast Alabama. Upubl. M.A.

thesis, Univ. Nebraska, Lincoln. 245 pp.

Accepted 8 October 1982

Notes on Distribution and Habitats of Sorex and

Microsorex (Insectivora: Soricidae)

in Kentucky

Ronald S. Caldwell

Department of Biology,

Texas College, Tyler, Texas 75702

AND

Hal Bryan

Division of Environmental Analysis,

Kentucky Department of Transportation,

Frankfort, Kentucky 40622

ABSTRACT. — New distributional information on long-tailed shrews

in Kentucky is presented. Sorex fumeus, S. cinereus, S. longirostris,

and Microsorex hoyi are shown to be more widespread in the state

than previously thought. Two apparently isolated populations of M.

hoyi occur. Habitat information and a key to Kentucky Soricidae are

included.

INTRODUCTION

Of the eight species of long-tailed shrews (genera Sorex and Micro-

sorex) shown by Junge and Hoffman (1981) to occur in the eastern

United States, only three — Sorex longirostris Bachman, Sorex cinereus

Kerr, and Sorex fumeus Miller — were reported from Kentucky by Bar-

bour and Davis (1974). Caldwell (1980) additionally documented the

presence in the state of Sorex dispar Batchelder and Microsorex hoyi

(Baird), bringing the number of long-tailed species to five. Additional dis-

tributional information were reported for Sorex longirostris and Sorex

fumeus by Bryan (1979), and for Sorex cinereus by French (1978). A

sixth species, the Water Shrew, Sorex palustris Richardson, may yet be

discovered in the higher elevations of eastern Kentucky.

This paper updates the known ranges in the state for each species

of long-tailed shrew (Figs. 1 A-E). Figure IF shows counties where pit-

falls have been placed. We also present a key to the species of Soricidae

in Kentucky (including short-tailed species), and habitat and ecological

information. Exact locality data are available from the authors upon

request. Catalogued specimens are deposited in the collections of the

Kentucky Nature Preserves Commission, the Division of Environmental

Analysis, the University of Kentucky vertebrate museum, and the per-

sonal collections of the authors.

Brimleyana No. 8:91-100. December 1982 91

92

Ronald S. Caldwell and Hal Bryan

Fig. 1. County records of long-tailed shrews known to occur in Kentucky, and

pitfall locations. Open circles are records from Barbour and Davis (1974), closed

circles are additional recent records.

Notes on Kentucky Shrews 93

METHODS AND MATERIALS

Both standard snap-back mouse traps and pitfall traps were used to

capture shrews, usually with 5 to 20 pitfall traps and 30 to 300 snap-

traps in suitable habitat. Snap-traps were removed after one to three

nights, but pitfalls often remained in place at each locality for 3 to 10

months.

Many researchers have commented on the effectiveness of pitfall

traps for capturing Sorex (Briese and Smith 1974; Brown 1967; Hudson

and Solf 1959; Rose 1980; Wolfe and Esher 1981; and others). In this

study, several sizes of pitfall cans were often used. Number 10 cans (153

mm opening) were preferred, but smaller sizes were also used. The top

of each can was positioned just below the surface of the ground, and

leaves were placed around the lip to present a more natural environ-

ment. As was suggested by MacLeod and Lethiecq (1963) water was

added to some cans to prevent shrews from escaping. Brown (1967) gave

suggestions for setting pitfalls in talus.

RESULTS AND DISCUSSION

Annotated List

Sorex cinereus Kerr, Masked Shrew — North of Kentucky, the

masked shrew is common in many habitats (Buckner 1966; Getz 1961),

but Kentucky specimens have all been taken in woodlands. Two subspe-

cies occur in the state (Fig. 1A). Sorex c. cinereus occurs in the eastern

mountains. Sorex c. lesueurii Duvernoy was first found in Henderson

County, in a floodplain forest along the Ohio River (French 1978).

More recently it was taken in similar habitat along the Ohio River in

Union County (Floyd Scott, unpubl. data), and along the Green River

in Henderson County about 1 1 km from its confluence with the Ohio

River. Specimens were collected in floodplain forests beside fallen logs

and other large debris. Eastern Kentucky specimens {S. c. cinereus) were

taken along rockfaces close to small order streams and seepages in

mesic woodlands at higher elevations (above 500 msl).

Sorex longirostris Bachman, Southeastern Shrew — French (1980)

stated that the favored habitats of Sorex longirostris in Alabama are

grassy freshwater marshes with rotting logs, and along river floodplains

dominated by hardwood forest. However, he noted that collections in

other states were made in many habitats, such as upland old-fields, dry

sandy areas, and pine plantations. In Kentucky this species often inhab-

its wet weedfields. However, one specimen was collected within 1 m of

the water's edge on a Kentucky River rockbar with no vegetative cover.

We have also taken it in riparian forests that annually flood along the

Kentucky River in Franklin County, and along East Fork Lynn Camp

94 Ronald S. Caldwell and Hal Bryan

Creek in Knox County (Fig. IB). The Knox County specimen is the first

record of this shrew in the Eastern Coal Field (elevation 326 msl). We

also have a specimen from the edge of the Inner Bluegrass on the

Franklin-Scott County line. We have found it most frequently in bot-

tomland hardwoods of the Mississippi Embayment, in the same cans

with the small short-tailed shrew, Blarina carolinensis.

Sorex fumeus Miller, Smoky Shrew — This is the most common

long-tailed shrew in the Eastern Coal Field (Fig. 1C). The series of

smoky shrews taken by Bailey (1933) at Mammoth Cave National Park,

Edmonson County, was once thought to represent a disjunct population

(Blair et al. 1968). However, our recent records support the hypothesis

of Barbour and Davis (1974) that these animals are "probably distrib-

uted across southern Kentucky where suitable habitat is available". We

collected smoky shrews across the Mississippian Plateau west to Todd

County. In addition, they have been collected along the wooded corri-

dor of the Kentucky River north to Franklin County, and probably

occur in other counties of central and north-central Kentucky. The

western limit of the species' range is as yet undetermined.

The species is usually found in relatively mature mesic forests with

deep organic litter (Hamilton 1940). Along the Kentucky River in

northern Franklin County it is the most abundant shrew in the forest

above the five-year floodplain. Individuals seldom move into the annu-

ally inundated riparian forest of cottonwood, silver maple, and syca-

more, where Blarina brevicauda is the most common shrew. In rich

mesic forest we have taken as many as 15 smoky shrews in 2 cans left in

place for 5 days. We recently took a smoky shrew in a wooded ravine in

Hardin County over upper Mississippian limestone in what was thought

to be the Kentucky "Barrens" (Dicken 1935).

Sorex dispar Batchelder, Long-Tailed Shrew — Sorex dispar was

taken at only two localities, both woods at high elevations on Pine

Mountain (Fig. ID). Here specimens were captured beside logs in

water-filled pitfall traps, and along wet rockfaces in snap-traps. Wet,

moss-covered rockfaces, and cool, shaded talus slopes at high eleva-

tions, are the preferred habitats. Sorex dispar should be sought at addi-

tional sites on Pine, Cumberland, and Big Black mountains in Pike,

Letcher, Harlan, and Bell counties.

Microsorex hoyi (Baird), Pygmy Shrew — Pygmy shrews are the

smallest mammals endemic to North America. Diersing (1980) revised

the pygmy shrews and reduced Microsorex to subgeneric status. The

taxonomic status of this shrew in Kentucky is currently under investiga-

tion by Caldwell and Smith. Long (1972a,b; 1974) considered the pygmy

shrew to be a boreal animal restricted in the southern parts of its range

Notes on Kentucky Shrews 95

to the higher elevations of the Appalachian Mountains, with extensions

along riparian corridors draining the high slopes. There is, however, a

recent record from Wabash County in southern Illinois (see Diersing

1980). North of Kentucky the pygmy shrew has been reported from a

variety of habitats, including bluegrass pasture, mature woodland,

marsh, and brushland (Long 1972b), often where two habitats are in

close proximity (Brown 1967; Spencer and Pettus 1966).

Barbour and Davis (1974) noted that there was a specimen from

the state with no locality data in the University of Kentucky collection.

Caldwell (1980) reported the first Kentucky pygmy shrews with specific

locality information. These were taken at elevations above 600 msl in

Harlan and Letcher counties (Fig. IE). The habitat was cool, mesic

forests in the terraces of small streams where rock outcrops, boulders,

and fallen logs were numerous. We recently collected this shrew in ripar-

ian woodlands at low elevations. In eastern Kentucky, two were taken

in a narrow wooded corridor adjacent to cropland along the Little

Sandy River, Greenup County (elevation 150 msl). Another was col-

lected along Sinking Creek in west-central Kentucky, Breckinridge

County. It was found in a similar narrow (5-10 m) wooded corridor

next to a cornfield. At both locations, typical stream border trees such

as sycamore, silver maple, box elder, and river birch were the dominant

species. The soil in these riparian habitats was sandy, and periodic

flooding had prevented accumulation of a deep layer of organic litter.

Tree stumps, logs, and brushpiles provided cover. We also collected

pygmy shrews in rich mesophytic forests with a well-developed litter

layer in Ohio County along the Rough River, and in northwestern

Warren County on the edge of the Dripping Springs Escarpment. The

overstory of these mature forest habitats was dominated by beech, sugar

maple, and yellow poplar. Soils were loamy in texture and were not

periodically flooded. The species was collected in both xeric and mesic

hardwood forest by biologists from the Department of Forestry at the

University of Kentucky's Robinson Forest, Breathitt County (J. Mor-

iarty and W. McComb, pers. comm.).

Sorex palustris Richardson, Water Shrew — This shrew has not

been taken in Kentucky in Recent times although it occurred here dur-

ing the Pleistocene (Guilday, et al. 1971). However, the species has been

taken in the highlands of Southern Appalachia to the east and south of

Kentucky (Conaway and Pfitzer 1952; Hooper 1942; Pagels and Tate

1976; Whitaker, et al. 1975). It is still possible that an isolated popula-

tion of the species may be found in the state.

Species Associations

We found several Kentucky sites inhabited by more than one spe-

96 Ronald S. Caldwell and Hal Bryan

cies of shrews. Five species, including four long-tailed species, were

taken in the talus slopes and rich mesic woodland at Bad Branch Falls,

Letcher County. These were Sorex dispar, S. fumeus, S. cinereus,

Microsorex hoyi, and Blarina brevicauda. This is the greatest number of

shrew species known from a single location in Kentucky. We also col-

lected S. fumeus and S. longirostris in the same pitfall traps in hard-

wood forest in Barren County. Both species were also collected with M.

hoyi in a mesic forest in Ohio County (Western Coal Field).

The three smallest species of long-tailed shrews in Kentucky —

Sorex cinereus, S. longirostris and Microsorex hoyi — are now known

from riparian areas within the Western Coal Field. Microsorex hoyi and

S. cinereus have been collected together in the Eastern Coal Field (Har-

lan and Letcher counties). Sorex longirostris and S. cinereus have yet to

be found together in Kentucky. Where they occur sympatrically in south-

ern Indiana, S. cinereus occupies lowland sites and S. longirostris

occurs primarily in upland habitats (T. French, pers. comm.; Hamilton

and Whitaker 1979). Rose (1980) reported that S. cinereus occurred

more often in forests while S. longirostris was captured more frequently

in old-fields.

KEY TO THE SORICIDAE OF KENTUCKY

To facilitate a more complete knowledge of soricid distribution within the

state, the following eclectic key is presented to aid collectors. General references

that can be used in conjunction with the key are Burt and Grossenheider (1976);

Hall (1981); Jackson (1928); Whitaker (1968, 1980); and Junge and Hoffman

(1981).

la. Less than 5 unicuspids visible from the side, or if 5, the third and fifth

greatly reduced (Figs. 2A, B, C); medial tine (Fig. 2F) of first upper

incisor present or absent 2

lb. Five unicuspids visible from side, fifth may be very reduced; medial

tine present 4

2a. Three or four unicuspids visible from the side (Figs. 2A, B); upper

incisors not possessing medial tine 3

2b. Unicuspids 3 and 5 greatly reduced, may not be readily apparent in

side view (Fig. 2C); fifth unicuspid peglike, third unicuspid platelike;

medial tine of upper incisors well developed (Fig. 2F) Microsorex

hoyi

3a. Three unicuspids visible in side view (Fig. 2A); fourth unicuspid hid-

den from view; total number of teeth 30 Cryptotis parva

3b. Four unicuspids visible in side view (Fig. 2B); first and second upper

unicuspids large, third and fourth smaller and subequal; total number

of teeth 32 {Blarina) 8

4a. Total length usually greater than 140 mm; fringe of hairs between toes;

maxillary breadth 6.0 mm or greater Sorex palustris

Notes on Kentucky Shrews

97

medial tine

Fig. 2. Dental characters of Kentucky Soricidae. A, Cryptotis parva. B, Blar-

ina. C, Microsorex hoyi, with first unicuspid shown on occlusional view. D,

Sorex longirostris. E, Sorex cinereus. F, medial tine of genera Sorex and Micro-

sorex.

98 Ronald S. Caldwell and Hal Bryan

4b. Total length usually less than 140 mm; no fringe of hairs between toes;

maxillary breadth less than 6.0 mm 5

5a. Infraorbital foramen with posterior border lying behind space between

first and second upper molars (Fig. 3A); tail longer than 50 mm

Sorex dispar

5b. Infraorbital foramen with posterior border lying ahead of the space

(Fig. 3B); tail less than 50 mm 6

6a. Third unicuspid usually smaller than fourth (Fig. 2D); tail relatively

shorter, generally 32-38 percent of total length; rostrum short and

wide; length of posterior palate to anterior end of first incisors usually

less than twice the greatest width across outside of first large molari-

form tooth Sorex longirostris

6b. Not with above combination of characters 7

7a. Ventral color distinctly lighter than dorsal color; midventral hairs just

anterior to axillary region light from midshaft to tip; maxillary breadth

narrower than 4.6 mm Sorex cinereus

7b. Ventral color not distinctly lighter than dorsal color; midventral hairs

just anterior to axillary region dark tipped; maxillary breadth greater

than 4.6 mm Sorex fumeus

8a. Total length greater than 105 mm, or hind foot 13 mm or longer; condy-

lobasal length greater than 20 mm Blarina brevicauda

8b. Total length 105 mm or less, and hind foot shorter than 13 mm; condy-

lobasal length less than 20 mm Blarina carolinensis

^jj&*^

(B)

Fig. 3. Infraorbital foramen position. A, Sorex dispar; B, Sorex fumeus. Ml is

first molar.

Notes on Kentucky Shrews 99

ACKNOWLEDGMENTS. —Some information presented was

obtained while performing field investigations with the biological staffs

of the Division of Environmental Analysis, Kentucky Department of

Transportation (Floyd Hughes, Director), and the Kentucky Nature

Preserves Commission. John Moriarty, William McComb, and Floyd

Scott allowed use of unpublished data. Thanks are extended to John O.

Whitaker, Jr., Wayne H. Davis, Roger W. Barbour, and John R. Mac-

Gregor for reviewing the manuscript. Special thanks to Cathy Caldwell,

Ann Barkley, and Cecilia Hayden for typing and proofreading earlier

drafts. Joyce Bryan provided the illustrations.

LITERATURE CITED

Bailey, Vernon. 1933. Cave life of Kentucky. Am. Midi. Nat. 74:385-635.

Barbour, Roger W., and W. H, Davis. 1974. Mammals of Kentucky. Univ.

Press Kentucky, Lexington. 322 pp.

Blair, W. Frank, A. P. Blair, P. Brodkorb, F. R. Cagle and G. A. Moore. 1968.

Vertebrates of the United States. McGraw-Hill Book Co., New York. 616

pp.

Briese, Linda A., and M. H. Smith. 1974. Seasonal abundance and movement

of nine species of small mammals. J. Mammal. 55:615-629.

Brown, Larry N. 1967. Ecological distribution of six species of shrews and

comparison of sampling methods in the central Rocky Mountains. J.

Mammal. 48:611-623.

Bryan, Hal D. 1979. The occurrence of two species of shrews in central Ken-

tucky. Trans. Ky. Acad. Sci.40:41-42.

Buckner, Charles H. 1966. Populations and ecological relationships of shrews in

tamarack bogs in southeastern Manitoba. J. Mammal. 47:191-194.

Burt, William H., and R. P. Grossenheider. 1976. A Field Guide to the Mam-

mals. Houghton Mifflin Co., Boston. 289 pp.

Caldwell, Ronald S. 1980. First records of Sorex dispar and Microsorex

thompsoni in Kentucky with distributional notes on associated species.

Trans. Ky. Acad. Sci. 41:46-47.

Conaway, Clinton H., and D. W. Pfitzer. 1952. Sorex palustris and Sorex dis-

par from the Great Smoky Mountains National Park. J. Mammal.

55:106-108.

Dicken, Samuel N. 1935. The Kentucky Barrens. Bull. Geogr. Soc. Phila.

55:42-51.

Diersing, Victor E. 1980. Systematics and evolution of the pygmy shrews (sub-

genus Microsorex) of North America. J. Mammal. 67:76-101.

French, Thomas W. 1978. First record of the masked shrew in western Ken-

tucky. Trans. Ky. Acad. Sci. 59:78-79.

. 1980. Natural history of the southeastern shrew, Sorex longirostris

Bachman. Am. Midi. Nat. 704:13-31.

Getz, Lowell L. 1961. Factors influencing the local distribution of shrews. Am.

Midi. Nat. 65:67-88.

100 Ronald S. Caldwell and Hal Bryan

Guilday, John E., H. W. Hamilton and A. D. McGrady. 1971. The Welsh Cave

peccaries (Platygonus) and associated fauna, Kentucky Pleistocene. Ann.

Carnegie Mus. ¥5:249-320.

Hall, E. Raymond. 1981. The Mammals of North America. John Wiley and

Sons, New York. 2 vols. 1181 pp.

Hamilton, William Jr., Jr. 1940. The biology of the smoky shrew (Sorex fumeus

fumeus Miller). Zoologica 25:473-492.

, and J. O. Whitaker, Jr., 1979. Mammals of the Eastern United States.

Cornell Univ. Press, Ithaca. 346 pp.

Hooper, Emmett T. 1942. The water shrew (Sorex palustris) of the southern

Allegheny Mountains. Occas. Pap. Mus. Zool. Univ. Mich. 463:1-4.

Hudson, G. E., and J. D. Solf. 1959. Control of small mammals with sunken

can pitfalls. J. Mammal. 40:455-457.

Jackson, Hartley H. T. 1928. A taxonomic review of the American long-tailed

shrews (genera Sorex and Microsorex). N. Am. Fauna 51. 238 pp.

Junge, J. A., and R. S. Hoffman. 1981. An annotated key to the long-tailed

shrews (genus Sorex) of the United States and Canada, with notes on

middle American Sorex. Occas. Pap. Mus. Nat. Hist. Univ. Kans. 94:1-48.

Long, Charles A. 1972 a. Taxonomic revision of the mammalian genus Micro-

sorex Coues. Trans. Kans. Acad. Sci. 74:181-196.

. 1972b. Notes on the habitat preference and reproduction in pigmy

shrews, Microsorex. Can. Field-Nat. #6:155-160.

. 1974. Microsorex hoyi and Microsorex thompsoni. Mamm. Spe-

cies 55:1-4.

MacLeod, C. F., and J. L. Lethiecq. 1963. A comparison of two trapping

procedures for Sorex cinereus. J. Mammal. 44:227-278.

Pagels, John R., and C. M. Tate. 1976. Shrews (Insectivora: Soricidae) of the

Paddy Knob-Little Back Creek area of western Virginia. Va. J. Sci.

27:202-203.

Rose, Robert K. 1980. The southeastern shrew, Sorex longirostris, in southern

Indiana. J. Mammal. (57:162-164.

Spencer, Albert W., and D. Pettus. 1966. Habitat preference of five sympatric

species of long-tailed shrews. Ecology 47:677-683.

Whitaker, John O., Jr. 1968. Key to the vertebrates of the eastern United States

excluding birds. Burgess Publ. Co., Minneapolis. 256 pp.

. 1980. The Audubon Society Field Guide to North American

Mammals. Chanticleer Press, New York. 745 pp.

, G. S. Jones and D. D. Pascal, Jr. 1975. Notes on the mammals of

the Fires Creek area, Nantahala Mountains, North Carolina, including

their ectoparasites. J Elisha Mitchell Sci. Soc. 97:13-17.

Wolfe, James L., and R. J. Esher. 1981. Relative abundance of the southeastern

shrew. J. Mammal. 62:649-650.

Accepted 24 August 1982

Unionid Mollusca (Bivalvia) from Little South Fork

Cumberland River, with Ecological and

Nomenclatural Notes

Lynn B. Starnes

Tennessee Valley Authority,

450 Evans Building, Knoxville, Tennessee 37902

AND

Arthur E. Bogan

Department of Malacology,

Academy of Natural Sciences of Philadelphia,

19th and the Parkway, Philadelphia, Pennsylvania 19103

ABSTRACT— A mollusk survey of the Little South Fork of the

Cumberland River in southern Kentucky from 1977 to 1981 yielded 24

species of Unionidae, one species of Corbiculidae, and 5 species of

aquatic gastropods. Pertinent taxonomic notes on unionids are made

herein. A series of quantitative surveys in riffles in the lower third of

the river (an area designated a Kentucky Wild River) revealed average

unionid densities ranging from 2.87 to 7.53 individuals per square

meter. Approximately five percent of the river contains optimal riffle

habitat. Average corbiculid densities ranged from 10.75 to 46.59 indi-

viduals per square meter.

INTRODUCTION

The Little South Fork of the Cumberland River (herein referred to

as Little South Fork) originates in Pickett County, Tennessee, meanders

through the Interior Low Plateau physiographic province, and conflu-

ences with the Big South Fork of the Cumberland River approximately

110 stream kilometers (64 air km) from its source (Fig. 1) (Fenneman

1938). Little South Fork changes from a high gradient stream south of

the Kentucky-Tennessee border to a moderate gradient stream with well

developed riffles and increasingly larger pools downstream.

Throughout its length little South Fork has eroded through Penn-

sylvanian shale and sandstone to Mississippian limestone. Water qual-

ity data reported by Harker et al. (1979, 1980) indicate that the Kidder

and Ste. Genevieve Limestone Members of the Monteagle Limestone,

exposed in the streambed, strongly influence water chemistry.

Development within the Little South Fork watershed is limited and

approximately 65% of the drainage area forested (Harker et al. 1980.)

Agriculture is primarily limited to floodplains. The watershed in the

vicinity of Mt. Pisgah and Parmleysville has historically been and

remains an area of oil production. Harker et al. (1980) reported oil

slicks, and we noted hydrogen sulfide odors, in the river in this area.

Surface mining of coal deposits associated with the Breathitt Formation

Brimleyana No. 8:101-119. December 1982 101

102

Lynn B. Starnes and Arthur E. Bogan

Fig. 1. Little South Fork of the Cumberland River, indicating major tributar-

ies, sampling stations, and approximate distances.

Mollusca From Little South Fork Cumberland River 103

beneath the Rockcastle Sandstone Member of the Lee Formation began

in the 1970s. As of March 1981 there were 11 active and 4 inactive

surface mines in the watershed.

While the fauna of Little South Fork has never been thoroughly

surveyed, historical records from the Cumberland River and its tributar-

ies provide information about which species had potential access to Lit-

tle South Fork. Wilson and Clark (1914) documented the distribution,

relative abundance and habitat of mussel resources in the Cumberland

River and the lower Big South Fork of the Cumberland River (Table 1)

from a commercial standpoint. Shoup and Peyton (1940) provided data

on unionids collected from the Tennessee section of Big South Fork.

Neel and Allen (1964) surveyed the upper Cumberland River in Ken-

tucky from 1947 to 1949, especially the area above Wolf Creek Dam

prior to impoundment. Two of the stations collected by Neel and Allen

were on Big South Fork — one above Burnside, which corresponds to a

Wilson and Clark locality, and the other at Yamacraw (Tables 1 and 2).

Ortmann (1924, 1925, 1926) provided taxonomic and distributional

information pertaining to unionids of the Cumberland River. William-

son (1905) reported a limited fauna in the Rockcastle River, a major

tributary located in the eastern headwaters of the Cumberland River.

Stansbery (1969) reviewed naiad faunal changes at Cumberland Falls

based on surveys of Wilson and Clark (1914) and Neel and Allen (1964).

Recent reports of the naiad fauna of Little South Fork include the

work on Pegias fabula by Starnes and Starnes (1980) and the species

lists provided by Harker et al. (1979, 1980). Their collection stations,

which are from sites that approximate our stations 4, 5, 6, 7, 8, 14, and

16, have been combined with our data in Table 3. B. Branson and G.

Schuster, Eastern Kentucky University (EKU), in 1980 surveyed the

lower part of Little South Fork from the Highway 92 crossing down-

stream to Freedom Church Ford at approximately our stations 8, 11,

12, 14, and 16. Their unpublished unionid data have been combined

with our data in Table 3.

MATERIALS AND METHODS

In 1977 we initiated qualitative surveys to establish a list of mussel

species. Our efforts in 1979 centered around collecting live specimens,

determining species assemblages, and estimating upstream distribution

limits of unionids. Independently, in 1978 the Kentucky Nature Pre-

serves Commission (KNPC) began comprehensive water quality and

biological surveys in the Little South Fork (Harker et al. 1979, 1980). In

1981 we conducted a quantitative and qualitative survey of the lower

Little South Fork. Ten-square foot (0.1 m2) samples were taken along

three transects at stations 8, 13, and 16, using a metal frame placed over

104

Lynn B. Starnes and Arthur E. Bogan

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the area to be sampled. To avoid wind ripples or sun glare, mask and

snorkel was used to observe the substrate while each rock was removed

from within the frame. All mollusks recovered from the square were

retained for identification.

Specimens collected by KNPC were identified by D. H. Stansbery

and deposited at Ohio State University. Collections by G. Schuster and

B. Branson are at EKU. Most of our specimens have been deposited in

the Department of Anthropology Zooarchaeology Collection, Univer-

sity of Tennessee (UT), Knoxville. Additional smaller collections have

been deposited at the Department of Malacology, Academy of Natural

Sciences of Philadelphia (ANSP), and in the senior author's collection.

TAXONOMY

Approximately 70% of the taxa recorded from Big South Fork,

confluent with Little South Fork, has undergone taxonomic revisions

since 1914; therefore, we feel that Table 1 and the following discussion

are essential to a general understanding of historical and modern union-

id taxonomy of Cumberland River tributaries. Table 1 compares our

nomenclature with that reported in Wilson and Clark (1914) and Neel

and Allen (1964). Synonyms were traced by using Bogan and Parmalee

(in press), Burch (1975), Clarke (1981), Haas (1969), Ortmann (1917,

1918), Ortmann and Walker (1922), and Simpson (1914). Of the 78 total

unionid species found in the Cumberland River, 24 have been docu-

mented in Little South Fork.

The Pleurobema clava of Wilson and Clark (1914) is here called the

P. oviforme "complex". Ortmann (1924) suggested that P. clava from

the Upper Cumberland and Big South Fork may be P. oviforme. We

have identified Fusconaia subrotunda reported in Starnes and Starnes

(1980) to be Pleurobema oviforme. Stansbery (OSU) identified KNPC

materials from the Rockcastle and Little South Fork rivers as P. ovi-

forme. Some specimens from Little South Fork approach P. clava in

shell shape, hence our use of "complex".

Wilson and Clark (1914) listed both Alasmidonta minor and A.

truncata from Big South Fork (Table 1). Clarke (1981) included both A.

minor and A. truncata as synonyms of A. calceolus, which he placed as

a junior synonym of A. viridis (Rafinesque 1820).

Taxa reported in the genera Carunculina, Truncilla, and Dysnomia

require clarification. Both Wilson and Clark (1914) and Neel and Allen

(1964) reported Carunculina, but not the taxon C. lividus, from Big

South Fork. Stansbery (1976) observed that Villosa vanuxemensis was

absent from the Rockcastle River and that the purple-nacred Toxo-

lasma was T. lividus. This suggests that the identification of V. vanuxe-

mensis by both Wilson and Clark (1914) and Neel and Allen (1964) was

Mollusca From Little South Fork Cumberland River 107

in fact a confusion with specimens of T. lividus. Morrison (1969) listed

Toxolasma Raf. 1831 as an earlier available name for Carunculina

Simpson in Baker, 1898. Johnson (1978) moved taxa from Epioblasma

(= Dysnomia), formerly placed in Truncilla, to the genus Plagiola, and

recognized the lectotype of Plagiola interrupta Raf., 1820 (Johnson and

Baker 1973). If the Poulson Collection of Rafinesque's types (see Vanatta

1916; Walker 1916) is accepted, then Johnson's revision based on the

type of Plagiola interrupta (= Dysnomia brevidens [Lea, 1934]) is accu-

rate. Our examination of the lectotype of P. interrupta confirms that it

is a female Epioblasma brevidens.

Rafinesque's types also affect two other taxa. Lampsilis ovata card-

ium Raf., 1820 (= L. o. ventricosa Barnes, 1823) is based on Raf-

inesque's type from the Poulson Collection (Johnson and Baker 1973;

Walker 1916; Vanatta 1916). The use of Potamilus (- Proptera) may be

argued on the basis of priority, i.e., Potamilus Raf., 1818 versus Pro-

ptera Raf., 1819.

RESULTS

In the combined 1977-1981 surveys, 24 species of Unionidae and

one species of Corbiculidae were collected (Table 3). This table repre-

sents the compilation of distribution data from our surveys, the KNPC

collections, and unpublished data from Schuster. Observed live speci-

mens of each species are indicated by an asterisk. There is a relatively

high correlation between species occurring historically in Big South

Fork and species currently occurring in Little South Fork. However, 15

species are conspicuously absent (Table 2). Some species, such as Ellip-

tic) crassidens, may have been excluded by preferences for a larger river

habitat. However, the majority of species, such as Fusconaia barnesiana

and Hemistena lata, are normally associated with other rivers the size of

Little South Fork. Comparable rivers, such as the Stones (Wilson and

Clark 1914), upper Powell (Ahlstedt and Brown 1980), middle Duck

(Ahlstedt 1981), upper Holston (Stansbery 1972; Stansbery and Clench

1974, 1975, 1978), and the Rockcastle (Williamson 1905) all contain

taxa absent from Little South Fork. It seems probable that the require-

ments of the 15 absent species are similar to those of the documented

species in Big South Fork and Little South Fork.

Table 3 is a compilation of molluscan collections at respective sta-

tions on Little South Fork. There is a pattern of longitudinal diversity,

with species being added as one moves downstream from Station 2.

From January 1980 to October 1981 there was a 37 cm shortage

from mean average rainfall. Due to this prolonged dryness, during our

quantitative sampling the entire river downstream from Station 13

(approximately 3 km above Ritner Ford) entered a sandy pool and fil-

108

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tered underground to enter subterranean channels in the Ste. Genevieve

limestone. The streambed was dry for approximately one kilometer,

after which the river emerged at the base of limestone bluffs. Below the

point of resurgence the water was cooler and had greater clarity, and

there were long, deep pools.

A sampling transect at Freedom Church Ford (Sta. 16) yielded an

average 2.87 unionid individual/ m2, compared with 7.53 and 7.17 indiv-

iduals/m2 at upstream stations 8 and 13, respectively (Table 4).

Decreases in abundance at Station 16 were probably related to de-

creases in optimal habitat. G. Schuster (pers. comm.) reported sedimen-

tation at Ritner Ford in spring 1981. Although these sediments were

gone by fall 1981, mussel populations are incapable of withstanding

repeated or extended siltation. Limestone outcrops and bedrock, which

increasingly dominated the substrate in the river below Station 13, may

also reduce the amount of optimal habitat. At Freedom Church (Sta.

16) the ford is a continuous sheet of limestone, 30 m wide and 20 m

long, and unionids are restricted to gravel accumulated above and

below the bedrock outcrops.

Within the river, certain general habitat preferences were apparent.

Without heavy spring rains in 1981, deep pools were covered with 2 to

10 cm of organic detritus. No live unionids were collected in these areas.

Neither unionids nor Corbicula occurred along stream margins where

water willow, Justicia americana, is abundant. However, in 1980 and

1981 these areas were exposed or in shallow water. In pool areas, heavi-

est concentrations of unionids, especially Potamilus alata, occurred

along current-swept banks. No live Corbicula were recorded in deep,

sluggish pools except at inflow areas, while shallow pools contained

occasional live specimens.

Unionids were not recorded in water less than 10 cm deep, but

Corbicula was found in water 3 cm in depth. Greatest unionid densities

occurred in water from 10 to 25 cm deep. With the exception of Lampsil-

is ovata and Pegias fabula, all unionids could be found with the anter-

ior end protruding slightly from the substrate. Lampsilis ovata was

uncovered in gravel with the anterior end approximately 5 cm below the

normal substrate surface. Pegias fabula, previously reported by Starnes

and Starnes (1980) at the interface between pool and riffle, was also

collected live from the shallower, current-swept areas in the riffle

proper. During low flow periods, Pegias was observed reclining upon

the substrate between gravel and cobbles, with a somewhat more hori-

zontal than vertical orientation. In higher spring flows, Pegias anchored

into the substrate with more typical unionid orientation. Ptychobran-

chus subtentum was ubiquitous in riffle areas in water 10 to 25 cm deep

and in all but the swiftest current. Medionidus conradicus was restricted

112

Lynn B. Starnes and Arthur E. Bogan

Mollusca From Little South Fork Cumberland River 113

tiree transects at stations 8, 13, and 16, Little South Fork Cumberland

001 002 012 024 20 30 72

Corbiculidae 43.73

114 Lynn B. Starnes and Arthur E. Bogan

to these deeper, swift areas, anchored between rocks or in sand-filled

cracks in bedrock.

Populations of Corbicula were widespread and four to six times as

dense as unionid populations, with 10.75, 43.73, and 46.59 individ-

uals/m2 at the three quantitatively sampled stations (Table 4). Relict

Corbicula shells were found in great numbers along banks and in pools.

While muskrats harvest both unionids and Corbicula, the ratio of Cor-

bicula to unionid shells in 1981 seemed disproportionately high, indicat-

ing possible changes in Corbicula populations.

Gastropods identified from Little South Fork included: Goniobasis

semicarinata (Say, 1829), G. ebenum (Lea, 1941), Pleurocera acuta

(Rafinesque, 1831), Physella sp., and Campeloma crassulum (Rafin-

esque, 1819). Campeloma rubrum is widely distributed, yet relatively

uncommon in the river outside of its preferred habitat. Live specimens

were restricted to mud (loamy) banks. Consequently, Campeloma was

not collected in quantitative surveys. The upstream limit for gastropods

was not determined; however, at the uppermost collecting locality their

numbers were significantly lower, ranging from 0 to 8 individuals/ m2.

Densities of gastropods ranged from 1 to 25 individuals/ m2 at Station 4,

from 5 to 38 individuals/ m2 at Station 8, and from 0 to 12 indivi-

duals/ m2 at Station 16.

DISCUSSION AND CONCLUSIONS

The Little South Fork of the Cumberland River contains perhaps

one of the last extant representative populations of the Cumberlandian

mollusk fauna in Kentucky. A total of 24 unionid, one corbiculid, and 5

gastropod species are reported in these surveys. After clarification of

unionid taxonomy, recent Little South Fork samples were compared

with historical Big South Fork records, revealing that approximately

one-third of Big South Fork species are absent from Little South Fork.

We can only speculate on the factor(s) responsible for the absence of

these 15 species. Along these lines, Stansbery and Clench (1975, 1978)

listed factors possibly limiting molluscan populations, including avail-

able nutrients, stable substrate or dissolved calcium. Unionid distribu-

tions in Little South Fork are most likely limited by a combination of

factors, including the possible absence of the proper fish host.

Unionid densities were greatest in current-swept substrates. Opti-

mal habitat was riffles with a relatively coarse substrate, in water 10 to

25 cm deep. Downstream density decreases were related to increases in

percentage of bedrock in pools and riffles, which reduced available

habitat.

Distribution of the naiad fauna is curtailed where the river gradient

increases from 1.2 m/km to 1.8 m/km before reaching the 3.8 m/km

Mollusca From Little South Fork Cumberland River

115

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116 Lynn B. Starnes and Arthur E. Bogan

gradient in the headwaters. Elevation changes in Little South Fork are

shown in Figure 2. Above Parmleysville (Sta. 4), where 12 species occur,

there is a relatively rapid decline in species diversity. The last species

encountered as we progressed upstream (Sta. 2) were Villosa taeniata

and V. trabalis, with all species absent at Station 1, 11 km above Sta-

tion 4. In the headwaters above Mt. Pisgah there is an average gradient

of 3.75 m/km; below this area it decreases to 1.2 m/km. The abrupt

change in stream gradient with associated physical changes is an effec-

tive faunal barrier (Fig. 2), as noted by Masnik (1975) for fish in the

upper Clinch River system. From the headwaters to the river area just

above Mt. Pisgah there is a faunal transition zone: gradient sharply

decreases, substrate shifts from boulder and bedrock to cobble and

sand, and river velocity decreases.

The section of the river at Mt. Pisgah is apparently a major physi-

cal barrier to naiad distribution farther into the headwaters. Similar

distributional limitations have been noted in other Tennesseee River

tributary streams. Apparently at and above this section of the river, the

unionids are not able to become established for any of several reasons:

change in fish fauna (lack of suitable host fish), lack of suitable sub-

strate, lack of nutrients, and thermal fluctuations. The observed faunal

barrier above Mt. Pisgah was anticipated by the work on unionid dis-

tribution in the headwaters of the Tennessee River by Stansbery (1972),

Stansbery and Clench (1974, 1975, 1978), Ahlstedt and Brown (1980),

Ahlstedt (1982), and the work on fish distribution by Masnik (1975). A

similar increase in stream gradient and corresponding faunal changes

was also observed in the Little River, Blount County, Tennessee (Bogan

and Starnes 1982). The small assemblage of species found in the rivers

just below the zone of increased gradient is one typical of small streams

(e.g., Lasmigona holstonia, Medionidus conradicus, Villosa iris, V. taen-

iata, Pleurobema oviforme, Alasmidonta viridis).

Current surface coal mining regulations (Kentucky Permanent

Regulatory Program, and the Surface Mining Control and Reclamation

Act of 1977) are designed to protect aquatic resources. However, even if

roads and silt control structures are properly designed, constructed, and

maintained, we are uncertain as to the survival of sensitive unionid

populations. Multiple mines, whose impacts on the watershed will be

combined or cumulative, characteristically will locate within coal-rich

watersheds. To preserve water quality and protect the unionid fauna,

the number of permits issued within both Little South Fork and indi-

vidual tributaries should be limited, and an annual monitoring program

to evaluate the status of the unionid populations should be initiated.

Survival of this river's unionid fauna possibly will be directly related to

compliance with and enforcement of the Act by inspection and

enforcement personnel of Kentucky and the Office of Surface Mining.

Mollusca From Little South Fork Cumberland River 117

ACKNOWLEDGMENTS. — We acknowledge the field, lab, and

editorial assistance of Wayne C. Starnes and Cindy Bogan. Special

appreciation is expressed to Jerry and Christine Louton for field assist-

ance, especially in an unexpected portage of canoes and samples. For

providing specimens, identifications, collection records, and observa-

tions we acknowledge the assistance of: B. A. Branson and G. A. Schus-

ter (EKU), Paul W. Parmalee (UT), David H. Stansbery (OSU), Melvin

L. Warren, Jr. (KNPC), and Sam M. Call (Kentucky Division of

Water). An anonymous reviewer provided helpful questions and

comments.

LITERATURE CITED

Ahlstedt, Steven A. 1981. The molluscan fauna of the Duck River between

Normandy and Columbia dams in central Tennessee. Bull. Am. Malacol.

Union Inc. for 1980:60-62.

1982. The molluscan fauna of Copper Creek (Clinch River System)

in southwestern Virginia. Bull. Am. Malacol. Union Inc. for 1981: 4-6

, and S. R. Brown. 1980. The naiad fauna of the Powell River in

Virginia and Tennessee (Bivalvia: Unionacea). Bull. Am. Malacol. Union

Inc. for 1979:40-43.

Bogan, Arthur E., and P. W. Parmalee. In press. Endangered and threatened

mollusks of Tennessee. Tenn. Wildl. Resour. Agency.

, and L. B. Starnes. 1982. The unionid fauna of the Little River and

some observations on unionid biogeography. Presentation annual meeting

American Malacological Union, New Orleans.

Burch, John B. 1975. Freshwater Unionacean clams (Mollusca: Pelecypoda) of

North America. Malacological Publications, Hamburg, MI. 204 pp.

Clarke, Arthur H. 1981. The Tribe Alasmidontini (Unionidae: Anodontinae),

Part I: Pegias, Alasmidonta, and Arcidens. Smithson. Contrib. Zool. No.

326. 101 pp.

Fenneman, Neville M. 1938. Physiography of Eastern United States. McGraw-

Hill, New York. 691 pp.

Haas, Fritz. 1969. Superfamilia Unionacea. Das Tierreich 55:1-663

Harker, Donald F., Jr., S. M. Call, M. L. Warren, Jr., K. E. Camburn and P.

Wigley. 1979. Aquatic biota and water quality survey of the Appalachian

Province, eastern Kentucky. Techn. Rep. Ky Nature Preserves Comm.,

Vol. 1. Frankfort. 522 pp.

, M. L. Warren, Jr., K.E. Camburn, S. M. Call, G. J. Fallo and P.

Wigley. 1980. Aquatic biota and water quality survey of the Upper Cum-

berland River Basin. Techn. Rep. Ky. Nature Preserves Comm., Vol. 2.

Frankfort. 409 pp.

Johnson, Richard I. 1978. Systematics and zoogeography of Plagiola (-Dys-

nomia-Epioblasma) an almost extinct genus of freshwater mussels (Bival-

via: Unionidae) from middle North America. Bull. Mus. Comp. Zool.

74S(6):239-321.

118 Lynn B. Starnes and Arthur E. Bogan

, and H. B. Baker. 1973. The types of Unionacea (Mollusca:

Bivalvia) in the Academy of Natural Sciences of Philadelphia. Proc. Acad.

Nat. Sci. Phila. 725(9): 145- 186.

Masnik, Michael T. 1975. Composition, longitudinal distribution, and zoogeo-

graphy of the fish fauna of the upper Clinch River system in Tennessee and

Virginia. Unpubl. Ph. D. dissert. Va. Polytech. Institute State Univ.,

Blacksburg. 401 pp.

Morrison, Joseph P. E. 1969. The earliest names for North American naiads.

Am. Malacol. Union Inc. Annu. Rep. 1969:22-24.

Neel, Joe K., and W. R. Allen. 1964. The mussel fauna of the upper Cumber-

land basin before its impoundment. Malacologia 7(3):427-459.

Ortmann, Arnold E. 1917. A new type of nayad-genus Fusconaia. Group of F.

barnesiana Lea. Nautilus 57(2):58-64.

. 1918. The naiades (freshwater mussels) of the upper Tennessee

drainage, with notes on synonymy. Proc. Am. Philos. Soc. 57:521-626.

1924. The naiad fauna of Duck River in Tennessee. Am. Midi. Nat.

9(1): 18-62.

. 1925. The naiad-fauna of the Tennessee River system below

Walden Gorge. Am. Midi. Nat. 9(8):321-372.

. 1926. The naiades of the Green River drainage in Kentucky. Ann.

Carnegie Mus. 77:167-188.

, and B. Walker. 1922. On the nomenclature of certain North

American naiades. Occas. Pap. Mus. Zool. Univ. Mich. No. 112. 75 pp.

Shoup, Charles S., and J. H. Peyton. 1940. Collections from the drainage of the

Big South Fork of the Cumberland River in Tennessee. J. Tenn. Acad. Sci.

75(1):106-116.

Simpson, Charles T. 1914. A descriptive catalogue of the naiades, or pearly

freshwater mussels. Bryant Walker, Detroit. 1540 pp.

Stansbery, David H. 1969. Changes in the naiad fauna of the Cumberland River

at Cumberland Falls in eastern Kentucky. Am. Malacol. Union Inc. Annu.

Rep. 1969:16-17.

. 1972. The mollusk fauna of the North Fork Holston River at

Saltville, Virginia. Bull. Am. Malacol. Union Inc. for 1971:45-46.

. 1976. Status of endangered fluviatile mollusks in central North

America: Toxolasma cylindrellus (Lea, 1868). Ohio State Univ. Res. Foundation.

Report to Bur. Sport Fish. Wildl., USFWS, USDI, Columbus. 7 pp.

, and W. J. Clench. 1974. The Pleuroceridae and Unionidae of the

North Fork Holston River above Saltville, Virginia. Bull. Am. Malacol.

Union Inc. for 1973:33-36.

, and 1975. The Pleuroceridae and Unionidae of the Middle

Fork Holston River in Virginia. Bull. Am. Malacol. Union Inc. for

1974:51-54.

, and 1978. The Pleuroceridae and Unionidae of the Upper

South Fork Holston River in Virginia. Bull. Am. Malacol. Union Inc. for

1977:75-78.

Starnes, Lynn B., and W. C. Starnes. 1980. Discovery of a new population of

Pegias fabula (Lea) (Unionidae). Nautilus 94{\):5-6.

Mollusca From Little South Fork Cumberland River 119

Vanatta, Edward G. 1916. Rafinesque's types of Unio. Proc. Acad. Nat. Sci.

Phila. (57:549-559.

Walker, Bryant. 1916. The Rafinesque-Poulson unios. Nautilus 50(4):43-47.

Williamson, E. B. 1905. Odonata, Astacidae and Unionidae collected along the

Rockcastle River at Livingston, Kentucky. Ohio Nat. 5(6):309-312.

Wilson, Charles B., and H. W. Clark. 1914. The mussels of the Cumberland

River and its tributaries. Bur. Fish. Doc. 781:1-63.

Accepted 6 December 1982

Response of Small Mammals to Forest Clearings

Created by Herbicides in the Central Appalachians

William C. McComb

and

Robert L. Rumsey '

Department of Forestry,

University of Kentucky, Lexington, Kentucky 40546

ABSTRACT. — Removal trapping was used to determine relative

abundance of six small mammals species on uncut, clearcut, and

herbicide-treated plots on ridgetop, southfacing and northfacing sites

in eastern Kentucky. Four rates of picloram-based herbicides were

tested. Twenty-nine microsite characteristics were measured at each

trap station to determine variables important to capture of each spe-

cies. Clearcuts supported 1.5 times as many small mammals as uncut

plots. Relative abundance of Sorex fumeus, Peromyscus leucopus,

Ochrotomys nuttalli, and Microtus pinetorum was higher on at least

one of the treatments than on untreated plots. Sorex fumeus was cap-

tured more frequently on northfacing slopes than on ridgetops. Rela-

tive abundances of Blarina brevicauda and Tamias striatus were unaf-

fected by treatment. Proximity to edge was found to be important to

capture of B. brevicauda, T striatus, and P. leucopus. Changes in

structure of overstory and understory, as well as in snag, log, stump

and /or rock characteristics, accounted for differential responses of

four species to treatments. Sorex fumeus, B. brevicauda, T. striatus,

and P. leucopus were more tolerant of a wide range of available habi-

tats than were O. nuttalli or M. pinetorum.

INTRODUCTION

Forest disturbance by clearcutting in small blocks is an accepted

method of increasing diversity and/ or abundance of those wildlife spe-

cies that benefit from two or more kinds of habitats (Kirkland 1977).

Effects of forest cutting on small mammal communities have been inves-

tigated by Gentry et al. (1968), and Hahn and Michael (1980). The

structure and composition of understory were reported to be important

to soricids, sciurids, cricetids, and microtines by Dueser and Shugart

(1978) and Geier and Best (1980). Herbicides affect understory composi-

tion and structure, and are frequently used in forest management prac-

tices (Dewey 1980; Loftis 1978). McCaffery et al. (1974) found picloram

herbicide to be useful in maintaining wildlife clearings in Minnesota,

and such herbicides are considered more desirable than others for creat-

ing forest clearings because of low toxicity to many vertebrate species

(McCollister and Leng 1969). We know of no studies that compare wild-

life use of picloram-created forest clearings with clearcut or uncut areas.

1 Present address: Department of Agriculture, McNeese State University, Lake Charles,

Louisiana 70601.

Brimleyana No. 8: 1 2 1 - 1 34. December 1 982 121

122 William C. McComb and Robert L. Rumsey

Our objectives were to compare the relative abundance of small mam-

mals among herbicide-created forest clearings, clearcuts, and uncut

areas on ridgetop, northfacing, and southfacing sites, and to identify the

habitat characteristics selected by each species.

STUDY AREA AND METHODS

Snag Ridge Fork watershed, in the University of Kentucky's

Robinson Forest, Knott County, Kentucky, contains a second-growth

mixed mesophytic forest typical of much of the central Appalachians

(Carpenter and Rumsey 1976). Ridges are dominated by shortleaf pine,

Pinus echinata; pitch pine, P. rigida; chestnut oak, Quercus prinus; and

scarlet oak, Q. coccinea. Southfacing slopes are dominated by hickories,

Carya spp.; white oak, Q. alba; black oak, Q. velutina; and sourwood,

Oxydendrum arboreum. Northfacing slopes are dominated by northern

red oak, Q. rubra; cucumbertree, Magnolia acuminata; and yellow-

poplar, Liriodendron tulipifera.

Eighteen 0.4-ha square plots in the watershed were sampled. Four

plots on each of a northfacing slope, southfacing slope, and ridgetop

randomly received one of the following hand-broadcast herbicide treat-

ments in May 1976: 23 kg/ ha TORDON 10K (T23), 46 kg/ ha TOR-

DON 10K (T46), 68 kg/ha TORDON 10K (T68), or 91 kg/ha M-3864

(M91) (mention of trade names is for identification and does not imply

endorsement by the Kentucky Agricultural Experiment Station, Lexing-

ton, Kentucky). TORDON 10K is a pelletized picloram-based (4-amino-

3, 5, 6-trichloropicolinic acid) herbicide, and M-3864 is a 5% picloram

pellet. A fifth plot on each aspect was clearcut; felled trees were not

removed. A sixth plot on each aspect was established in the untreated

forest at least 75 m from any treated plot. Treated plots were 15 to 50 m

apart.

Fifteen stations were established 4.3 m apart perpendicular to the

contour through the center of each plot. Two Museum Special snap-

traps, baited with peanut butter, were set within 2 m of each station for

one night each month on each plot, January to June 1980. Treatments

on each aspect were sampled simultaneously, and aspects were sampled

on consecutive nights. Stations were grouped into upper edge (5), plot

center (5), and lower edge (5). Analysis of variance and Duncan's New

Multiple Range Test were used to compare mean captures per station of

each species among treatments, aspects, and plot edges versus center.

The reciprocal of Simpson's Index (I/SPi2 , where Pi = proportion of

captures in the ith treatment) was used as an index to each species toler-

ance (TI) to habitat changes (Geier and Best 1980). The maximum TI

for treatments was 6.00, and for aspects was 3.00.

Twenty-one of twenty-nine microsite characteristics, chosen on the

basis of previous studies (Dueser and Shugart 1978; Geier and Best

Small Mammal Response to Clearings

123

Table 1. Habitat variables used in correlation and regression analyses, small

mammal site preference on clearcut, herbicide-treated and uncut plots,

January to June 1980, Robinson Forest, Knott County, Kentucky.

Acronym

Description

NTR Number of trees > 10 cm dbh within 2 m of a station.

DMT Diameter of nearest tree (cm).

DST Distance to nearest tree (m).

BA Basal area of living stems (m2 /ha).

NSN Number of snags> 10 cm dbh, > 1 .8 m tall, within 2 m

of a station.

DMS Diameter of nearest snag (cm).

DSS Distance to nearest snag (m).

NSP Number of stumps > 10 cm in diameter and < 1.8 m tall, within

2 m of a station.

DSP Distance to nearest stump (m).

NLG Number of logs > 10 cm diameter, > 1 .8 m long, within

2 m of a station.

DML Maximum diameter of nearest log (cm).

LGL Length of nearest log (m).

DSL Distance to nearest log (m).

PCL Percent of ground covered by logs within 2 m of a station.

DSR Distance to nearest rock > 5 cm above ground.

PCR Percent of ground covered by rocks.

CRN Percent crown cover above 6.1 m at a station.

MID Percent vegetation cover between 1.8 and 6.1 m at a station.

LFC Percent of ground covered by fallen leaves.

CV1, CV2 Percent understory cover < 1.8 m tall per 4m2 in Jan. and Apr.

DN1, DN2 Number of understory stems per 4m2 in Jan. and Apr.

Rl, R2 Number of understory taxa per 4m2 in Jan. and Apr.

DIV1, DIV2 Understory species diversity per 4m2 in Jan. and Apr.

DWT Distance to water (m).

SLP Slope (percent).

1980), were quantified at each station, May 1980 (Table 1). Estimates of

cover by rocks, logs, leaves, canopy, and midstory followed methods

described by James and Shugart (1971). Understory vegetation charac-

teristics (8) were quantified on 4 m2 circular plots 2 m away from each

station along the contour in January and April 1980. A modified

Aldous method was used similar to that described by Murphy and

Noble (1972). Percent cover and stem density were determined for each

124 William C. McComb and Robert L. Rumsey

plant taxon on each 4 m2 plot. Total cover, total density, plant species

richness, and Shannon-Weaver plant species diversity were calculated

for each station during both sampling periods. Habitat characteristics

on treatments and aspects are described in detail by McComb and

Rumsey (1981). Correlation and stepwise regression were used to iden-

tify microsite characteristics important to the capture of each species.

Since stepwise regression selects variables in a progressive series,

some characteristics potentially important to small mammal captures

may not have been selected if they were highly correlated with a pre-

viously selected variable. Correlation used in conjunction with stepwise

regression ensured that any highly correlated microsite characteristics

important to small mammal capture were not overlooked. A compari-

son of PRESS (predicted residual sum of squares) statistics was used to

objectively select the best model produced by the stepwise procedure.

Each model was entered into the General Linear Models (GLM) proce-

dure with the print (P) and confidence limits (CLM) options specified

within the Statistical Analysis System (SAS 79) to provide the PRESS

statistic (Helwig and Council 1979). The strongest model was assumed

to be the one with the lowest PRESS statistic. Microsite characteristics

that were distributed linearly with respect to small mammal capture

were identified with this method.

RESULTS

Small Mammal Captures

We captured 385 mammals in 3,250 trapnights (Table 2). Peromys-

cus leucopus accounted for 76.4% of the captures, followed by Blarina

brevicauda (6.8%), Sorex fumeus (6.5%), Microtus pinetorum (3.9%),

Ochrotomys nuttalli (3.6%), and Tamias striatus (2.9%). Species rich-

ness and species diversity were similar among treatments (x=4.3, 0.467,

respectively) and among aspects (x=3.0, 0.833, respectively) (Z^O.05).

With the exception of T68, more mammals were captured on clearcut

and herbicide-treated plots than on control plots (/><0.05) (Table 2),

and more mammals were captured on northfacing slopes than on south-

facing slopes (P<0.05) (Table 3).

More P. leucopus, O. nuttalli, and S. fumeus were captured on

clearcuts than on control plots (/K0.05) (Table 2). Kirkland (1977)

reported an increase in S. fumeus abundance, and only a slight increase

in P. leucopus abundance, after clearcutting at West Virginia sites. We

found no differences in relative abundance of B. brevicauda or S.

fumeus between herbicide-treated plots and control plots; however, with

the exception of T46, 100% more P. leucopus were caught on herbicide-

treated plots than on control plots (Table 2). More M. pinetorum were

caught on M91 plots than on any other except T23 plots (/K0.05).

Small Mammal Response to Clearings

125

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126 William C. McComb and Robert L. Rumsey

Table 3. Number of small mammals captured, January to June 1980, from

plots with south, north and ridgetop exposures in Robinson Forest,

Knott County, Kentucky. (TN = trap-nights; values within each row

with different letters vary significantly (P<0.05); Duncan's New Multi-

ple Range Test).

Sorex fumeus was more abundant on north slopes than on ridges, and

M. pinetorum was more abundant on north slopes than on ridges or

south slopes (/<0.05) (Table 3).

Blarina brevicauda (TI=5.24) and P. leucopus (TI=5. 12) were least

sensitive to habitat disturbances due to herbicides or cutting. Sorex

fumeus (TI=4.03) and T. striatus (TI=3.90) were moderately tolerant of

habitat disturbances due to herbicides or cutting, and M. pinetorum

(TI=2.03) and O. nuttalli (TI=1.00) were the most sensitive to habitat

differences. Peromyscus leucopus (TI=2.99), T. striatus (TI=2.8 1) and B.

brevicauda (TI=2.75) were tolerant of different aspects, followed by S.

fumeus (TI=2.38), O. nuttalli (TI=2.18), and M. pinetorum (TI=1.14).

Geier and Best (1980) reported T. striatus tolerant (TI=4.48) and B. bre-

vicauda moderately tolerant (TI=2.73) of habitat change in Iowa

(TImax-6.00). The relative tolerance of each species depends upon the

small mammal community composition and/ or geographic location.

Proximity of trapping station to plot edge was important to the

capture of B. brevicauda, T. striatus, and P. leucopus (/K0.05). These

species benefitted more from edge presence than did S. fumeus, O. nut-

talli, or M. pinetorum.

Habitat Selection

Results of habitat selection analyses are constrained by the number

of microsite characteristics quantified and by the range of values for

each characteristic. The results are useful for determining which habitat

characteristic within these constraints were important to small mammal

captures.

Small Mammal Response to Clearings 127

As habitat heterogeneity increased, the capture site for each species

became less predictable. Models derived from captures from all plots

(habitat-wide) had low R2 values (5.6 to 26.1). These models selected

the habitat characteristics that described the plot(s) with the most cap-

tures. Models derived from captures on separate treatments or aspects

(habitat-specific) had higher R2 values (25.6 to 62.1) than habitat-wide

models. These models selected characteristics useful in predicting cap-

ture sites within a treatment or aspect. In the following discussions,

standardized partial regression coefficients are indicated parenthetically.

Sorex fumeus. — A habitat-wide model selected number of logs

(SPRC = +0.18) and distance to a rock (-0.12) as important to capture

of S. fumeus, and both characteristics were correlated (/><0.01) with

capture (r = +0.20 and -0.16, respectively) (Table 4). Captures were pre-

dictable on T46 (R2 = 47.0), clearcuts (R2 = 25.6), and southfacing slopes

(R2 = 27.2). Log characteristics (number, diameter, and length) were

significant (P<0.05) in habitat-wide and in habitat-specific correlation

and multiple regression analyses. Sorex fumeus was most apt to be

caught on clearcuts or heavily treated plots on northfacing slopes with

sparse understory (> 8 understory species per 4m2, < 45 understory

stems per 4 m2), within 4.4 m of a rock, and within 1 .7 m of an 8 to 14

m long log.

Blarina brevicauda. — A habitat-wide model selected seven charac-

teristics as important to capture of B. brevicauda and four of these vari-

ables were also correlated (P<0.05) with B. brevicauda capture: number

of logs (SPRC=+0.31), number of trees (SPRC= -0.16), slope (SPRC =

+0.21), and basal area (SPRC = -0.09) (R2 = 26.1). Relatively many logs,

high understory density, and high understory cover were important in

habitat-specific models. Gottschalk and Shure (1979) reported high leaf

decomposition rates and high microarthropod populations on herbicide-

treated forest floors, but Getz (1961) suggested that leaf cover is impor-

tant in maintaining high humidity within soricid tunnels. Logs may pro-

vide moisture-maintaining cover for B. brevicauda and S. fumeus in

areas with low leaf cover and high food availability. Blarina brevicauda

was most likely captured at the edge of a plot on a northfacing slope >

47%, with > 21ogs per 4 m2, and< 1 tree per 4 m2 (R2 = 27.5). Geier and

Best (1980) also reported low plant species richness as important to B.

brevicauda abundance.

Tamias striatus. — Capture sites of T. striatus were the least pre-

dictable (R2 = 5.1) of those for the mammals captured. Tamias striatus

was most likely caught on the edge of a plot within 1.4 m of a log in an

area with basal area > 21 m2/ha, < 8 understory species per 4 m2 , and

> 55 understory stems per 4 m2 . Dueser and Shugart (1978) and Geier

and Best (1980) reported high understory density or cover as important

128

William C. McComb and Robert L. Rumsey

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130 William C. McComb and Robert L. Rumsey

to captures of T. striatus, but they also found high plant species richness

a characteristic important to this species.

Peromyscus leucopus. — The habitat-wide model selected crown

cover (-0.28) and log diameter (+0.17) as important to capture of P.

leucopus (/K0.05). Captures were most strongly correlated (P<0.01)

with crown cover (-0.31), log diameter (+0.23), basal area (-0.22), dis-

tance to log (-0.21), log length (+0.20), and distance to a tree (+0.20)

(Table 4). Understory structure and log characteristics were correlated

(ZK0.05) with capture on four and three of the treatments, and three

and two of the aspects, respectively. The importance of each microsite

characteristic to capture of P. leucopus varied among treatments and

probably varies geographically. Dueser and Shugart (1978) reported a

deciduous overstory, wide tree dispersion, high species richness, and

large stumps important to P. leucopus in Tennessee. Geier and Best

(1980) found logs, brushpiles, and low plant species richness important

to P. leucopus abundance in Iowa. The most likely place to capture P.

leucopus in our plots was on the edge of a clearcut with an understory

of > 8 species and > 50 stems per 4 m2, and within 2.8 m of a log over

20 cm in diameter, on a site covered more than 4% by rocks (R2=47.3).

Ochrotomys nuttalli. — This species selected sites on clearcuts with

slopes > 21%, < 146 m from water, with < 25% leaf cover, > 50%

understory cover, > 1.1 m from a log and > 5.4 m from a rock

(R2=62.1). Dueser and Shugart (1978) and Linzey (1968) identified dense

understory cover as important to O. nuttalli in Tennessee, but they did

not report leaf cover (SPRC = -0.86), logs (SPRC = +0.29), or rocks

(SPRC = +0.52) as important to this species.

Microtus pinetorum. — A habitat-wide model selected understory

diversity (-0.98), understory density (+0.90), understory richness (+0.76),

basal area (+0.81), and tree density (-0.69) as important to capture M.

pinetorum. Kirkland (1978) also reported microtines responding to

changes in vegetative structure at West Virginia sites. Microtus pineto-

rum was most likely captured on the M91 plot on northfacing slopes

with a basal area > 24 m2/ha, near a snag < 28 cm dbh, on a 4 m2 site

with < 1 tree, and with logs < 5.5 m long (R2=32.6).

DISCUSSION

Differential response of small mammals to treatments and aspects

is a function of differences in the relative abundance of individuals, and

of habitat components among sites. Results of studies of habitat prefer-

ences and relative abundances of small mammals may differ from ours

if conducted in a contiguous rather than a discontiguous forest. Four

species responded to treatments by change in relative abundance and

two species responded to aspects by the same change. Biotic microsite

characteristics (vegetative cover, logs, stumps, leaf cover, etc.) were

Small Mammal Response to Clearings 131

important to capture of all six species, but abiotic factors (rocks, slope,

water) were also important to capture of S. fumeus, B. brevicauda, P.

leucopus, and O. nuttalli. Habitat characteristics selected by most spe-

cies were similar to those reported by previous investigators, but there

were a few notable exceptions. The similar relative abundance of S.

fumeus and B. brevicauda on southfacing and northfacing slopes would

be unexpected based on the findings of Manville (1949), Pruitt (1953),

Wetzel (1958), Getz (1961), and Barbour and Davis (1974) who reported

moisture as important to the occurrence of shrews. Contrary to Getz

(1961), we did not find leaf cover correlated with soricid capture. Gott-

schalk and Shure (1979) and Bormann and Likens (1979) reported that

herbicides and clearcutting, respectively, will increase leaf litter decom-

position rates and increase microarthropod populations; microarthro-

pods are important food items in soricid diets. Log cover was important

to capture of both species; logs may serve as moisture-maintaining cover

and/ or feeding sites for shrews. Several authors have reported B. brevi-

cauda to be tolerant of a wide range of habitats (Getz 1961; Briese and

Smith 1974; Geier and Best 1980), but no similar estimate of tolerance is

available for S. fumeus. We found S. fumeus not as tolerant of site

differences as B. brevicauda, but it was relatively tolerant of site change

within the small mammal community we sampled. Sorex fumeus and B.

brevicauda were similar in abundance on each aspect and treatment,

and both selected similar microhabitats. The number of logs per 4 m2

was the most useful variable in predicting captures of both species, and

log cover was also important to both species. Occurrence of S. fumeus

was positively associated (r = +0.31; SPRC = +0.26) with understory

species richness on northfacing slopes, while B. brevicauda occurrence

was associated negatively (SPRC = -0.12) with understory species rich-

ness in the habitat-wide regression model. Geier and Best (1980) also

reported low plant species richness as important to occurrence of B.

brevicauda. Coexistence of these shrews seems to be allowed by subtle

differences in vegetative structure. Additional research is needed to

determine differences in foods, microhabitats, and microclimates of

these two sympatric soricids.

Dueser and Shugart (1978) compared microhabitat characteristics

among P. leucopus, O. nuttalli, and T. striatus. Our results support

their findings: P. leucopus and T. striatus are more tolerant of a wide

range of habitat characteristics than are O. nuttalli. We found O. nut-

talli associated (r>0.40) with high log cover and canopy openings

(+DST, +DSS, and -BA) on ridges and north slopes. Peromyscus leu-

copus was associated to that degree (r>0.40) with low crown cover on

ridges, but otherwise was not strongly associated with any particular

habitat characteristics, and T. striatus occurrence was not correlated

132 William C. McComb and Robert L. Rumsey

(P<0.05) with any habitat characteristic. Dueser and Shugart (1978)

suggested that since T. striatus is diurnal and P. leucopus is nocturnal,

and since they differ morphologically, they can coexist with little com-

petition. Microtus pinetorum was found most frequently on M91 on

northfacing slopes, and was intolerant of other sites. Understory and

overstory structure was important to capture of M. pinetorum, and

these characteristics differed from characteristics important to the other

rodents sampled. Variation in vegetative structure seemed to allow coex-

istence of M. pinetorum with other rodents on northfacing slopes. If

competition is occurring within the small mammal community on our

study areas, then it is probably most intense on northfacing slopes and

least intense on ridgetops.

Differences in relative abundance and habitat selection among dif-

ferent physiographic sites may allow a more complete understanding of

small mammal species distribution within a watershed, and may increase

our understanding of niche overlap and competitive exclusion among

sympatric small mammals.

Management Implications

An increase in small mammal abundance may be desired as a non-

game management practice or to produce small mammal biomass for

predatory game or furbearers. If nongame management is an objective,

then plots should be small and widely spaced to maximize edge. Habitat

alterations should include providing habitat for species intolerant of

habitat variety, such as O. nuttalli (clearcut) and M. pinetorum (herbi-

cide, preferably similar to M-3864). Resulting increases in rock expo-

sure and microarthropod populations through increased leaf decompo-

sition on herbicide and clearcut plots (Bormann and Likens 1979;

Gottschalk and Shure 1979), and in log abundance, stump abundance,

and understory density, will benefit the six species studied.

Small mammals were 1.5 times more abundant on clearcuts than

on herbicide plots, and twice as abundant on herbicide-treated plots

than on uncut plots. Pelleted herbicide application is less expensive than

cutting (Dewey 1980), but the resulting increase in small mammal bio-

mass following herbicide application up to 91 kg/ ha is also less than

results from cutting. Limited accessibility, ruggedness of the terrain,

and/ or increasing fuel and labor costs may influence managers to use

herbicides to create clearings for wildlife.

ACKNOWLEDGMENTS.— We wish to thank R. L. Anderson, D.

M. Allen, and M. J. Immel for advice on statistical analyses; J. B. Davis

and G. M. Gigante for assistance in field work; Dow Chemical Co. for

providing herbicides; and B. A. Thielges, S. B. Carpenter, W. H. Davis,

Small Mammal Response to Clearings 133

R. S. Caldwell, G. L. Kirkland, Jr., and R. W. Barbour for reviewing an

early draft of this manuscript. The investigation reported in this manu-

script (No. 81-8-41) is in connection with the Kentucky Agricultural

Experiment Station Project No. 620 and is published with the approval

of the Director.

LITERATURE CITED

Barbour, Roger W., and W. H. Davis. 1974. Mammals of Kentucky. Univ.

Press Kentucky, Lexington. 322 pp.

Bormann, F. Herbert, and G. E. Likens. 1979. Patterns and process in a for-

ested ecosystem. Springer-Verlag, N.Y. 253 pp.

Briese, Linda A., and M. H. Smith. 1974. Seasonal abundance and movement

of small mammals. J. Mammal. 55:615-629.

Carpenter, Stanley B., and R. L. Rumsey. 1976. Trees and shrubs of Robinson

Forest, Breathitt County, Kentucky. Castanea ¥7:277-282.

Dewey, John B. 1980. "Gridball Pellets" — a new tool for brush control in

pines. Forest Farmer 59(20): 14-15, 34.

Dueser, Raymond D., and H. H. Shugart, Jr. 1978. Microhabitats in a forest-

floor small mammal fauna. Ecology 59:89-98.

Geier, Anthony R., and L. B. Best. 1980. Habitat selection by small mammals of

riparian communities: evaluating effects of habitat alterations. J. Wildl.

Manage. 44:16-24.

Gentry, John B., E. P. Odum, M. Mason, V. Nabholz, S. Marshall and J. T.

McGinnis. 1968. The effect of altitude and forest manipulation on relative

abundance of small mammals. J. Mammal. ¥9:539-541.

Getz, Lowell L. 1961. Factors influencing the local distribution of shrews. Am.

Midi. Nat. (55:67-88.

Gottschalk, Marlin R., and D. J. Shure. 1979. Herbicide effects on leaf litter

decomposition processes in an oak-hickory forest. Ecology (50:143-151.

Hahn, Brad L., and E. D. Michael. 1980. Responses of small mammals to whole

tree harvesting in central Appalachia. Trans. Northeast Fish and Wildl.

Conf. 37:32-44.

Helwig, Jane T., and K. A. Council (eds.). 1979. SAS users guide, 1979 edition.

SAS Institute, Raleigh. 494 pp.

James, Francis C, and H. H. Shugart, Jr. 1971. A quantitative method of

habitat description. Audubon Field Notes 24:721-736.

Kirkland, Gordon L., Jr. 1977. Responses of small mammals to the clearcutting

of northern Appalachian forests. J. Mammal. 55:600-609.

1978. Population and community responses of small mammals to

2,4,5,-T. U.S. Dep. Agric. For. Serv. Res. Note PNW-314. 7 pp.

Linzey, Donald W. 1968. An ecological study of the golden mouse, Ochrotomys

nuttalli, in the Great Smoky Mountains National Park. Am. Midi. Nat.

79:320-345.

Loftis, David L. 1978. Preharvest herbicide control of undesirable vegetation in

southern Appalachian hardwoods. South. J. Appl. For. 2:51-54.

Manville, Richard H. 1949. A study of small mammal populations in Northern

Michigan. Misc. Publ. Mus. Zool. Univ. Mich. 75:1-83

134 William C. McComb and Robert L. Rumsey

McCaffery, Keith R., F. L. Johnson and L. D. Martoglio. 1974. Maintaining

wildlife openings with pellets containing picloram. Wildl. Soc. Bull. 2:40-45.

McCollister, D. D., and M. L. Leng. 1969. Toxicology of picloram: safety eval-

uation of Tordon herbicides. Down-to-Earth 25(2):5-10.

McComb, William C, and R. L. Rumsey. 1981. Habitat characteristics of forest

clearings created by picloram herbicides and clearcutting. Proc. Annu.

Conf. Southeast. Assoc. Game Fish Comm. 55:(in press).

Murphy, Patrick K., and R. E. Noble. 1972. The monthly availability and use of

browse plants by deer on a bottomland hardwood area in Tensas Parish,

Louisiana. Proc. Annu. Conf. Southeast. Assoc. Game Fish Comm. 26:39-57.

Pruitt, William O. 1953. An analysis of some physical factors affecting the local

distribution of the shorttail shrew (Blarina brevicauda) in the northern part

of the Lower Peninsula of Michigan. Misc. Publ. Mus. Zool. Univ. Mich.

79:1-39.

Wetzel, Ralph M. 1958. Mammalian succession on midwestern flood plains.

Ecology 59:262-271.

Accepted 4 October 1982

Spawning Behavior in Seven

Species of Darters (Pisces: Percidae)

Lawrence M. Page and Michael E. Retzer

Illinois Natural History Survey,

607 E. Peabody, Champaign, Illinois 61820

AND

Robert A. Stiles

Department of Biology,

Samford University, Birmingham, Alabama 35209

ABSTRACT. — Recent observations reveal that Percina evides is an

egg-burier, Etheostoma duryi, E. asprigene, and E. chlorosomum are

egg-attachers, E. aquali and E. microlepidum are egg-clumpers, and E.

barbouri is an egg-clusterer. Etheostoma duryi attaches its eggs to

rocks; E. asprigene and E. chlorosomum attach their eggs to plants.

With documentation of egg-clumping in E. aquali and E. micro-

lepidum, three species of the subgenus Nothonotus now are known to

be egg-clumpers and three to be egg-buriers. Sites of egg-deposition

are known for 57 species of darters.

Darters spawn in four general ways: (1) they bury their eggs in the

substrate and abandon them, (2) attach them over a relatively large area

to objects (usually plants or rocks) above the substrate and abandon

them, (3) clump them in the interface between a slanted stone and the

stream substrate and guard them until they hatch, or (4) cluster them on

the underside of an object (stone or log) and guard them until they

hatch. Winn (1958) discussed these modes of behavior in darters. Page

(in press) summarizes information on 48 species for which spawning

behavior or at least the site of egg deposition had been reported in the

literature. Page et al. (1981) provided data on spawning in two addi-

tional species — Etheostoma longimanum Jordan and Etheostoma

obeyense Kirsch, and O'Neil (1981) on a third species — Etheostoma

coosae (Fowler). Herein are presented data on six more species —

Percina evides (Jordan and Copeland), Etheostoma duryi Henshall,

Etheostoma asprigene (Forbes), Etheostoma barbouri Kuehne and

Small, Etheostoma aquali Williams and Etnier, and Etheostoma micro-

lepidum Raney and Zorach — and more detailed information on

Etheostoma chlorosomum (Hay). All species of darters for which the

site of egg deposition now is known are categorized in Table 1.

Percina (Ericosma) evides. — As are all species of Percina reported

to date (Table 1), P. evides is an egg-burier. On 9 July 1971, 2 June

1972, and 17 June 1979, at water temperatures of 20°, 17°, and 18° C,

respectively, pairs of P. evides were observed spawning in Little River,

Blount County, Tennessee. All spawnings occurred over sand and gravel

interspersed with cobble and boulders in water 30 to 60 cm deep in the

upper parts of riffles. Current readings, taken on the bottom with a

Teledyne Gurley Pygmy Current Meter, were 0.20 to 0.61 m/sec.

Brimleyana No. 8:135-143. December 1982 135

136

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137

Fig. 1. Egg deposition sites of darters. A) Percina evides spawning in Little River,

Blount Co., TN, 17 June 1979. B) Etheostoma duryi spawning in Butler Creek,

Lawrence Co., TN, 5 April 1975. C) Etheostoma chlorosomum spawning in

aquarium; eggs are being deposited on dead leaf. D) E. asprigene spawning in

aquarium; eggs are being deposited on leaves of grass. E) E. barbouri male and

his nest of eggs on the underside of stone removed from Pettys Fork, Adair Co.,

KY, 29 May 1981. F) E. aquali male and his nest of eggs on underside of stone

removed from Buffalo River, Lewis Co., TN, 5 May 1981. The male E. barbouri

and E. aquali were guarding the eggs prior to removal.

138 Lawrence M. Page, Michael E. Retzer, Robert A. Stiles

Prespawning activities typically consisted of a female swimming

fairly rapidly over the bottom, examining the substrate to locate a

spawning site, closely followed by a male. When the female stopped, the

male mounted and, if the female did not swim away, vibrated his head

and body. If not ready to spawn, the female swam away. Once the

female selected the spawning location, vibrations of the male and female

became intense, and as eggs and sperm were released the female's genital

region pushed into the substrate displacing sand and gravel (Fig. 1A).

Once the spawning act was completed, the female rested on the bottom

for up to three minutes before seeking a new site.

Males twice were observed reversing their positions during mount-

ing. In the first instance, the female had stopped in a sand-gravel

depression among cobble with her head pointing downstream. The

courting male mounted with his head over hers and vibrated several

times. She quivered in response, then after about one minute he turned

and positioned himself with his head upstream over her tail. They

remained head-to-tail for about another minute, then the female turned

to face upstream for a few seconds before returning to a head-to-tail

position. The male turned to put his head over the female's, but soon

returned to a head-to-tail position. The female then turned to face

upstream, and head-to-head, spawned. Shortly thereafter the male was

displaced by a larger male and the female moved a short distance to a

new site between two rocks. The second male mounted with his head

over the female's, but soon turned to a head-to-tail position. After

about one minute he returned to the head-to-head position and then

back to head-to-tail. The female left the area with the male in pursuit.

By aligning head-to-tail, the males appeared to have been attempt-

ing to induce the female to turn around; alternatively, by positioning his

tail over the female's head, the male may have been stimulating her nape

with tubercles on his anal fin. The first explanation seems to be

supported by the fact that, of the four pairs of P. evides observed

courting, only two males, courting the same female, displayed this head-

to-tail maneuver. The female changed her positions, in one instance

turning around to spawn facing upstream, and in the other seeking

another spawning site. Additional observations may yield a more defini-

tive explanation for head-to-tail maneuvering.

Etheostoma (Nanostoma) duryi. — Etheostoma duryi inhabits runs

and pools of streams with moderate current and a rocky substrate. On

seven dates between early March and late May, E.duryi was observed

spawning in Butler and Factory creeks, Lauderdale County, Alabama,

and Lawrence County, Tennessee. The reproductive behavior of E.

duryi is essentially the same as that reported for the closely related

Etheostoma simoterum (Cope) (Page and Mayden 1981). Eggs usually

are deposited individually in slight depressions on the sides, or less often

on the tops, of rocks.

Prior to spawning, females swam slowly over the substrate, exam-

ining the sides of rocks for sites on which to attach eggs. Typically, as a

Darter Spawning Behavior 139

female moved about, a male followed closely and drove away approach-

ing males. Agonistic encounters between males usually consisted of the

larger male swimming at, or making a quick dash at, the smaller male.

Two males about the same size sometimes engaged in lateral displays

(with median fins held erect). Females accepted any male, and if a

male previously courting her was chasing away an intruding male, the

female sometimes spawned with a third male.

When a female selected a spawning site, she aligned her body over

it and the male mounted. Both vibrated with their bodies in an S-shaped

figure and their genital papillae pressed against or near the rock (Fig.

IB). Etheostoma duryi appeared to vibrate less intensely than E.

simoterum; however, verification of this will require cinematographic

analysis.

Etheostoma (Vaillantia) chlorosomum. — Winn (1958), para-

phrasing C. Hubbs, stated that in Texas E. chlorosomum lays its eggs

on plants or plant debris, but gave no other details. Three breeding male

and three ripe female E. chlorosomum, 40 to 50 mm SL, were captured

in East Fork Kaskaskia River, Marion County, Illinois, on 30 April

1981 in a sand-bottom pool at 22° C and a maximum depth of 80 cm.

At one end of the pool was a large accumulation of logs and other plant

debris. East Fork is a fairly slow stream composed mostly of long pools

but with some gravel-bottom riffles.

The E. chlorosomum were transferred at 0900 hr the following day

to a 40 1 aquarium (23° C) at the Illinois Natural History Survey

(INHS). One half of the aquarium bottom was covered with sand, the

other half with gravel. In one corner a large stone was propped on a

smaller stone so that its underside was accessible for egg-clustering or

clumping, and its top and sides were available for egg-attachment. A

small log (5 cm diameter) was placed across the aquarium, over the sand

and gravel, for attachment of eggs. An accumulation of filamentous

algae, dead leaves, and twigs was provided at the end of the log over

sand.

Little activity ensued until 1 145 hr when the darters were fed live

cladocerans. After feeding, the males became aggressive towards one

another and courted the females. Both aggressive and courting behavior

consisted mainly of lateral displays. Usually when displaying, the males

were swimming and quivering about 1 cm above the substrate. All such

activities occurred over the sand-covered half of the aquarium.

At 1230 hr, one male succeeded in chasing the other two out of the

sand-covered part of the aquarium and sequentially enticed all three

females to spawn. Spawning continued until 1700 hr when observations

were discontinued. As in other egg-attaching darters, the female selected

the site of egg-deposition, the male closely followed her to the site and

mounted her, the pair vibrated, and eggs and sperm were released.

Often the male mounted a female while she remained horizontal on the

substrate and beat her with his pectoral fins until she swam into the

plant material. Usually 1 to 3 eggs (1 mm diameter) were laid at a time,

140 Lawrence M. Page, Michael E. Retzer, Robert A. Stiles

and all were attached to plant material. Most were deposited on a twig

(3 mm diameter), but others were laid in the algae or on dead leaves

(Fig. 1C), and a few were laid on the log.

Etheostoma (Oligocephalus) asprigene. — Five breeding (2 males, 3

females) E. asprigene, 35 to 50 mm SL, were captured in Cane Creek

(Saline system), Gallatin County, Illinois, on 9 April 1981 in fairly fast

water just below an artifical riffle (of rip-rap) in 20° C water. For much

of its length Cane Creek is a sluggish stream with clay and sand

substrates. At 0900 hr the following day the darters were transferred to

a 40 1 aquarium at INHS. Because the darters were captured over gravel

in current, and because some consubgenerics of E. asprigene (Etheos-

toma spectabile (Agassiz) and Etheostoma caeruleum Storer) bury their

eggs in gravel, the substrate provided in the aquarium was fine gravel.

Two large stones, one propped on one end to create a space beneath,

were also placed in the aquarium to provide the opportunity for

attachment of the eggs to their sides or to cluster or clump them on the

underside.

By 1000 hr the males were courting the females, and by 1200 hr one

pair was spawning. Eggs were attached, 1 to 3 at a time, 15 to 25 cm

above the substrate, to the glass in the black corners of the metal-frame

aquarium. We assumed that the vertical position of the corner posts of

the aquarium simulated to the darters vertical shafts of vegetation, and

at 1300 hr a crabgrass plant (no aquatic vegetation was immediately

available) was rooted in the gravel. Spawning continued to be restricted

to the corners of the aquarium until about 1600 hr, and then one pair

moved to the crabgrass and began laying eggs on its blades (Fig. ID).

Once spawning on the grass began, no more eggs were laid in the

corners of the aquarium. Spawning continued for several hours, but

ended that day. Only one of the two males spawned, but with at least

two of the three females.

The spawning act in E. asprigene occurred as described above for

E. chlorosomum. Although both crabgrass and aquarium corners are

artificial spawning substrates, it is apparent from the aquarium obser-

vations that E. asprigene is an egg-attaching species. Eggs were not

buried in the substrate, or clustered or clumped on the underside of the

available stone. The fact that eggs were not laid on the tops or sides of

the large stones suggests that in nature eggs are attached to living or

dead plant material. Other species of the subgenus Oligocephalus that

lay their eggs on plants are Etheostoma lepidum (Baird and Girard),

Etheostoma grahami (Girard), and Etheostoma ditrema Ramsey and

Suttkus (Strawn 1956; Winn 1958; Seesock et al. 1978).

Etheostoma (Catonotus) barbouri. — Two nests of E. barbouri

eggs were found on the undersides of flat stones in a large slab-rock

pool in Pettys Fork (of Russell Creek), Adair County, Kentucky, on 29

May 1981. The first nest, found in water about 50 cm deep, contained

70 eggs (2.1 mm diameter) and was guarded by a breeding male (Fig.

IE). The second nest was found on a stone dragged up in a minnow

Darter Spawning Behavior 141

seine and contained 42 eggs. E. barbouri is the last of the ten

taxonomically described species of Catonotus to be confirmed as an

egg-clustering species.

Etheostoma (Nothonotus) aquali. — The subgenus Nothonotus of

Etheostoma is the only subgenus of darters in which some species bury

their eggs and others clump them (Table 1). From a study conducted in

Pennsylvania, Raney and Lachner (1939) first reported the egg-

deposition site of Etheostoma maculatum Kirtland. In Tennessee, Stiles

(1972) found eggs of E. maculatum on the undersides of stones held as

territories by males. To date, no other species of Nothonotus had been

reported as an egg-clumper.

On 5 May 1981 we found four nests of E. aquali in a large rubble

riffle in Buffalo River, Lewis County, Tennessee. The water was swift,

22° C, and averaged 30 cm deep where the nests were found. Eggs were

difficult to find in the swift water, and it was even more difficult to capture

the male in association with the nest. However, one 63 mm SL male was

captured beneath a stone to which a clump of eggs was attached. (Fig.

IF).

Nests of E. aquali were similar in construction to those of E.

maculatum as described by Raney and Lachner (1939). Unlike the

single-layer clusters of species of the subgenera Boleosoma and Catono-

tus, nests of E. maculatum and E. aquali contain a multi-layer clump of

eggs. Each nest contained eggs in various stages of development, which

obviously were the results of more than one spawn. One nest, removed

and preserved for an accurate egg count, contained 551 eggs averaging

1.8 mm in diameter.

Two male and four female E. aquali, 50 to 65 mm SL, were col-

lected in Buffalo River on 5 May 1981 and returned to INHS. On 8

May they were placed in an 80 1 aquarium, at 24° C, outfitted with a

current pump. Two stones were propped with the undersides accessible

as nesting sites. Observations were made through the next two days but

no spawning activities were seen. However, by 0800 hr on 11 May a

male had established a territory under the smaller of the two stones (the

other male had died) and was guarding three small clumps of eggs. The

clumps were separated by distances of at least 2 cm, but all were in the

interface between the nest stone and the gravel substrate. No further

spawning occurred.

The only notable behavior of aquarium-held E. aquali, other than

the constant attention given his territory by the breeding male, was that

of the females. On two occasions a female was seen to swim beneath the

nest stone and tightly wedge herself (more or less right-side up) in the

interface between the stone and gravel substrate. Although the male

ignored her, it appears that eggs are laid with the female wedged

142 Lawrence M. Page, Michael E. Retzer, Robert A. Stiles

between the rock and substrate. One of us (RAS) has seen similar

behavior in E. maculatum in the upper Tennessee River system. The

clumping of eggs results not from the stacking of eggs on top of one

another, but because the eggs are pumped into the interface between the

slanted stone and stream substrate. When the stone is lifted from the

water, the adhesive clump (or at least part of it) is attached.

Although most of the E. aquali eggs laid in the aquarium died, six

of a clump transferred to an aerated 600 ml glass dish hatched. Hatch-

ing time was three to four days at a fluctuating 24 to 28° C. Hatchlings

were 6.5 mm TL, had well-developed pectoral fins and jaws, and a

mid-dorsal series of bright gold flecks.

Etheostoma (Nothonotus) microlepidum. — One nest of E. micro-

lepidum eggs and an attendant male, 53 mm SL, were found in a mod-

erately fast riffle in East Fork Stones River, Rutherford County, Ten-

nessee, on 6 May 1981. The riffle averaged 15 cm deep and was

composed of large gravel and small rubble; water temperature was 21° C.

The nest contained an estimated 346 eggs, averaging 2.0 mm in diame-

ter, arranged in a multi-layer clump essentially identical to those of E.

maculatum and E. aquali.

Etheostoma maculatum, E. aquali, and E. microlepidum are closely

related, and we predicted that all would be egg-clumpers. Relationships

among other species of Nothonotus are less obvious, and it is difficult to

predict which clump eggs and which bury eggs. According to Zorach

(1972), other relatives of E. maculatum are Etheostoma acuticeps

Bailey, Etheostoma moorei Raney and Suttkus, and Etheostoma rubrum

Raney and Suttkus; confirmation of egg-clumping (the derived state) in

these species would confirm their recent shared ancestry with E.

maculatum.

Although three unrelated subgenera (i.e. nonsister-groups) of dar-

ters amass and guard their eggs, the manner in which this is accom-

plished in Nothonotus is unlike that in Boleosoma and Catonotus.

Among North American freshwater fishes, single-layer clusters are laid

by Boleosoma, Catonotus, and the minnow genus Pimephales (McMil-

lan and Smith 1974); multi-layer clumps of eggs are laid by Nothonotus,

Noturus (madtoms) (Mayden and Burr 1981), and some Cottus (scul-

pins) (Smith 1922). The selective factors separating these two types of

behavior remain to be determined.

ACKNOWLEDGMENTS. — We thank B. M. Burr, Southern Illi-

nois University at Carbondale, for helping search for nests of Etheos-

toma barbouri, and P. W. Smith, Illinois Natural History Survey, D. G.

Lindquist, University of North Carolina at Wilmington, and an ano-

nymous reviewer, for comments and suggestions on the manuscript.

Darter Spawning Behavior 143

LITERATURE CITED

Mayden, Richard L., and B. M. Burr. 1981. Life history of the slender madtom,

Noturus exilis, in southern Illinois (Pisces: Ictaluridae). Occas. Pap. Mus.

Nat. Hist. Univ. Kans. 95:1-64.

McMillan, Victoria E., and R. J. F. Smith. 1974. Agonistic and reproductive

behavior of the fathead minnow (Pimephales promelas Rafinesque). Z.

Tierpsychol. 54:25-58.

O'Neil, Patrick E. 1981. Life history of Etheostoma coosae (Pisces: Percidae) in

Barbaree Creek, Alabama. Tulane Stud. Zool. Bot. 23( l):75-83.

Page, Lawrence M. In press. Handbook of Darters. T.F.H., Inc., Nepture, NJ.

, and R. L. Mayden. 1981. The life history of the Tennessee

snubnose darter, Etheostoma simoterum in Brush Creek, Tennessee. 111.

Nat. Hist. Surv. Biol. Notes 117. 11 pp.

, W. L. Keller and L. E. Cordes. 1981. Etheostoma (Boleosoma)

longimanum and E. (Catonotus) obeyense, two more darters confirmed as

egg-clusterers. Trans. Ky. Acad. Sci. 47:35-36.

Raney, Edward C, and E. A. Lachner. 1939. Observations on the life history of

the spotted darter, Poecilichthys maculatus (Kirtland). Copeia 1939(3):

157-165.

Seesock, Wendy E., J. S. Ramsey and F. L. Seesock. 1978. Life and limitation

of the goldwater darter (Etheostoma ditrema) in Glencoe Spring, Alabama.

ASB (Assoc. Southeast. Biol.) Bull. 25:56.

Smith, Bertram G. 1922. Notes on the nesting habits of Cottus. Pap. Mich.

Acad. Sci. Arts Lett. 2:222-224.

Stiles, Robert A. 1972. The comparative ecology of three species of Notho-

notus (Percidae — Etheostoma) in Tennessee's Little River. Unpubl. Ph.D.

Dissert., Univ. Tennessee, Knoxville. 97 pp.

Strawn, Kirk. 1956. A method of breeding and raising three Texas darters. Part

II. Aquarium J. 27:12-14, 17,31-32.

Winn, Howard E. 1958. Comparative reproductive behavior and ecology of

fourteen species of darters (Pisces-Percidae). Ecol. Monogr. 25:155-191.

Zorach, Timothy. 1972. Systematics of the percid fishes, Etheostoma camurum

and E. chlorobranchium new species, with a discussion of the subgenus

Nothonotus. Copeia 1972(3):427-447.

Accepted 1 June 1982

Occurrence and Distribution of Land Snails of the Family

Polygyridae (Mollusca: Gastropoda: Pulmonata)

in West Virginia

Clement L. Counts, III1

Department of Biological Sciences,

Marshall University, Huntington, West Virginia 25701

ABSTRACT. — Twenty-four species in the family Polygyridae were

collected in West Virginia. Mesodon inflectus and Stenotrema barbiger-

um are reported from the state for the first time, and additional

records are also reported for other species of polygyrid snails. Results

of this study are compared with those of previously reported studies.

INTRODUCTION

The polygyrid gastropods of West Virginia have been discussed

with varying degrees of detail in several works. Pilsbry's (1940) major

study of North American land snails contained many county records for

West Virginia polygyrid species, although it provided little information

on specific collection sites. Brooks and MacMillan (1940) and MacMil-

lan (1949) presented more precise distributional lists. Many polygyrid

taxa have since been described from the state (Grimm 1971; Hubricht

1976), and new county and state distribution records have been pub-

lished (Grimm 1974; Taylor and Counts 1976). Many of the species

included in these works have been placed in synonymy (Burch 1962;

Grimm 1974; Hubricht 1974), creating some confusion as to which spe-

cies are present in the state. In addition, gaps still exist in our know-

ledge of West Virginia's polygyrid fauna.

This paper summarizes the occurrence and distribution of this

group of land snails in West Virginia, and presents new locality data

and species reports.

MATERIALS AND METHODS

Both live snails and empty shells were collected from several locali-

ties in each of the 55 counties of West Virginia, during the period Sep-

tember 1975 through December 1976. Animals were identified to species

or subspecies using Burch (1962) or Burch and Patterson (1966). Des-

criptions in these keys were compared with those of Pilsbry (1940) and

MacMillan (1949). Some identifications were made or confirmed by Mr.

Leslie Hubricht, Meridian, Mississippi.

Voucher specimens were deposited in the collections of the Museum

of Comparative Zoology, Harvard University; the Delaware Museum of

Natural History, Greenville; the Marshall University Malacological Col-

Present address: College of Marine Studies, University of Delaware, Lewes, Delaware

19958.

Brimleyana No. 8:145-157. December 1982 145

146 Clement L. Counts, III

lection (MUMC) of the N. Bayard Green Museum of Zoology; and Mr.

Leslie Hubricht's collection.

Locality data from the collections were compared with and added

to those in the available literature on West Virginia land snails, and the

results plotted on county maps of the state. Published localities that I

did not visit or where I failed to find the reported species were not

plotted. Species not previously reported from West Virginia in the liter-

ature are designated new state records.

RESULTS

Twenty-four polygyrid species were found in West Virginia and are

discussed in the following accounts. Previously unpublished localities

are provided by county. MUMC accession numbers are given for each

collection.

Triodopsis denotata (Deshayes, 1830). — Triodopsis denotata was

found in widely separated populations throughout the state (Fig. 1).

Mason Co.: McClintic Wildlife Refuge Station (MUMC 50).

Triodopsis tridentata (Say, 1816). — This species was found in all

but Berkeley, Hancock, Pleasants, Tyler, and Upshur counties (Fig. 2).

Boone Co.: West Virginia State Route (SR) 3, 3.4 km E of US 119

(MUMC 491); US 1 19, 1 km S of Kanawha Co. line (MUMC 485); SR

3, 4.6 km E of US 119 (MUMC 494). Cabell Co.: SR 2, 1.6 km E of

Huntington (MUMC 3); 8342 Big Seven Mile Rd. (MUMC 41, 55, 57);

Ona (MUMC 545). Fayette Co.: US 60, 7.5 km W of Gauley Bridge

(MUMC 217). Jackson Co.: SR 35, 3.5 km E of Putnam Co. line

(MUMC 127); US 33, 7.2 km E of Ripley (MUMC 153). Kanawha Co.:

Nitro (MUMC 120). Lincoln Co.: SR 10, 3.2 km W of West Hamlin

(MUMC 89); Alkol (MUMC 449). Logan Co.: Big Creek (MUMC 388);

Chief Logan State Park (MUMC 585). Mason Co.: McClintic Wildlife

Refuge Station (MUMC 60). McDowell Co.: US 52, 9.6 km W of

Welch (MUMC 703). Mercer Co.: Flat Top (MUMC 302). Mineral Co.:

US 50, 9.1 km E of Grant Co. line (MUMC 472, 479). Mingo Co.: Grey

Eagle (MUMC 536). Nicholas Co.: SR 39, 1.6 km W of Summersville

(MUMC 234); SR 39, 4 km W of Richwood (MUMC 418). Ohio Co.:

SR 2, 1.6 km E of Warwood (MUMC 259). Pendleton Co.: Mouth of

Seneca (MUMC 625). Pocahontas Co.: SR 39, east entrance to Monon-

gahela National Forest (MUMC 210). Putnam Co.: Hurricane (MUMC

71, 84); Clymer's Creek Rd. off SR 34, 3 km N of Kanawha River

(MUMC 152). Raleigh Co.: Slab Fork (MUMC 345); SR 3, 37 km N of

Daniels (MUMC 529); Beaver (MUMC 566). Randolph Co.: Glady

Fork (MUMC 19); US 33, 20.8 km W of Elkins (MUMC 459). Roane

Co.: US 33, 5.6 km E of Jackson Co. line (MUMC 137). Summers Co.:

SR 3, 2.6 km W of Jumping Branch (MUMC 511). Tucker Co.: Dolly

West Virginia Polygyrid Snails

147

Figs. 1-6. County distributions of Triodopsis in West Virginia. Horizontal

scale bar in all figures = 100 km. 1, T. denotata; 2, T. tridentata; 3, T. albolabris;

4, T. fraudulenta fraudulenta; 5, T.f. vulgata; 6, T. rugosa.

148 Clement L. Counts, III

Sods (MUMC 570); Lanesville (MUMC 596). Wayne Co.: Lavalette

(MUMC 98); Cabwaylingo State Park (MUMC 201); Shoals (MUMC

563). Wirt Co.: SR 14, 4 km N of Roane Co. line (MUMC 173). Wyom-

ing Co.: SR 10, 4.8 km S of Logan Co. line (MUMC 719).

Triodopsis albolabris (Say, 1816). — Triodopsis albolabris was also

widely distributed throughout West Virginia (Fig. 3). Braxton Co.:

Interstate (I) Highway 79, 1.6 km S of Burnsville exit (MUMC 600,

602). Cabell Co.: 8342 Big Seven Mile Rd.(MUMC 45, 53). Jackson

Co.: US 33, 7.2 km E of Ripley (MUMC 155). Lincoln Co.: Clymer's

Creek Rd. off SR 34 (MUMC 83). Mason Co.: McClintic Wildlife

Refuge Station (MUMC 61). Mercer Co.: Flat Top (MUMC 309). Min-

eral Co.: US 50, 9.1 km E of Grant Co. line (MUMC 466). Monroe Co.:

Red Sulphur Springs (MUMC 523); SR 12, 3.2 km S of Ballard

(MUMC 526). Nicholas Co.: SR 39, 1.6 km W of Summersville

(MUMC 228). Pendleton Co.: Mouth of Seneca (MUMC 626). Poca-

hontas Co.: Cranberry Glades (MUMC 620). Putnam Co.: SR 34, 2.2

km N of Kanawha River (MUMC 145). Summers Co.: SR 3, 2.7 km W

of Jumping Branch (MUMC 513). Randolph Co.: US 33, 22.8 km W of

Elkins (MUMC 460). Tucker Co.: Lanesville (MUMC 614). Wayne Co.:

SR 75, Lavalette (MUMC 105); Cabwaylingo State Park (MUMC 255,

296); Beech Fork (MUMC 640).

Triodopsis fraudulenta fraudulenta (Pilsbry, 1894). — Populations

of this snail occur in the northern panhandle and along the eastern

border of the state (Fig. 4). Fayette Co.: US 60, 7.5 km W of Gauley

Bridge (MUMC 215). Greenbrier Co.: SR 39, 9.6 km W of Cranberry

Glades (MUMC 623). Hardy Co.: US 220, 1.6 km N of Grant Co. line

(MUMC 583). Mineral Co.: US 50, 9.1 km E of Grant Co. line (MUMC

471). Ohio Co.: County Road (CR) 25 at Petes Creek (MUMC 251).

Pocahontas Co.: SR 39 at east entrance to Monongahela National

Forest (MUMC 205).

Triodopsis fraudulenta vulgata (Pilsbry, 1940). — Ohio Co.: SR 2,

1.6 km E of Warwood (MUMC 260). Wetzel Co.: SR 2, 0.8 km S of

Marshall Co. line (MUMC 271).

Triodopsis rugosa Brooks and MacMillan, 1940. — Triodopsis rugosa

populations are found only in the southern half of the state (Fig. 6).

Pocahontas Co.: Hills Creek Falls (MUMC 630).

Mesodon thyroidus (Say, 1816). — This species was found in all but

Fayette, Gilmer, Harrison, Morgan, Pendleton, Pleasants, Pocahontas,

Raleigh, and Upshur counties (Fig. 7). Boone Co.: SR 3, 3.4 km E of

US 1 19 (MUMC 433); SR 3, 4.6 km E of US 1 19 (MUMC 496). Brax-

ton Co.: Burnsville (MUMC 601). Brooke Co.: Bethany (MUMC 275).

Cabell Co.: 8342 Big Seven Mile Rd. (MUMC 38, 44, 52, 54, 56); SR

10, 7 km N of West Hamlin (MUMC 544). Clay Co.: Elkhurst (MUMC

West Virginia Polygyrid Snails

149

Figs. 7-12. County distributions of Mesodon in West Virginia. 7, M. thyroid-

us\ 8, M. say anus; 9, M. zaletus; 10, M. appressus; 11, M. perigraptus; 12, M.

dentiferus.

150 Clement L. Counts, III

115). Hancock Co.: SR 2, 3.8 km N of Weirton (MUMC 284). Jackson

Co.: SR 35, 3.5 km E of Putnam Co. line (MUMC 128); US 33, 5.6 km

E of Ripley (MUMC 154). Kanawha Co.: Nitro (MUMC 118). Lincoln

Co.: SR 34, 3.2 km S of Putnam Co. line (MUMC 1 12). Logan Co.: SR

10, 4.3 km E of Davin (MUMC 389); Chapmanville (MUMC 575); Big

Creek (MUMC 577): Chief Logan State Park (MUMC 594). Mason

Co.: McClintic Wildlife Refuge Station (MUMC 59). Mineral Co.: US

50, 9.1 km E of Grany Co. line (MUMC 567). Mingo Co.: Grey Eagle

(MUMC 535). Monroe Co.: SR 12, 0.2 km S of Red Sulphur Springs

(MUMC 522). McDowell Co.: US 52, 9.6 km W of Welch (MUMC

702). Ohio Co.: SR 2, 1.6 km E of Warwood (MUMC 256). Pendleton

Co.: Mouth of Seneca (MUMC 627). Putnam Co.: Hurricane (MUMC

72); SR 34 off US 60, Clymer's Creek Rd. (MUMC 81); SR 34, 3 km N

of Kanawha River (MUMC 144). Roane Co.: US 33, 5.6 km E of Jack-

son Co. line (MUMC 136). Summers Co.: SR 3, 2.6 km E of Nimitz

(MUMC 505, 507); SR 3, 2.6 km W of Jumping Branch (MUMC 517).

Wayne Co.: SR 75, Lavalette (MUMC 104); SR 75, Shoals (MUMC

562); Beech Fork (MUMC 638). Wirt Co.: SR 14, 4 km N of Roane Co.

line (MUMC 174).

Mesodon sayanus (Pilsbry, 1906). — Mesodon sayanus occurs prim-

arily in the eastern highlands, and in the Kanawha and Guyandotte

river valleys of the southwest (Fig. 8). A few populations are also found

in the floodplain of the Ohio River in the northern panhandle, and in

the southwest corner of the state. Cabell Co.: Ona (MUMC 541). Grant

Co.: SR 93, 11.3 km E of Tucker Co. line (MUMC 463). Logan Co.:

Chief Logan State Park (MUMC 586). Pocahontas Co.: Cranberry

Glades (MUMC 621). Putnam Co.: Hurricane (MUMC 70). Raleigh

Co.: Slab Fork (MUMC 375). Wayne Co.: Cabwaylingo State Park

(MUMC 289).

Mesodon zaletus (Binney, 1837). — This species was found in essen-

tially the same distribution pattern as M. sayanus (Fig. 9). Cabell Co.:

SR 2, 3.2 km E of Huntington (MUMC 1). Mercer Co.: Flat Top

(MUMC 310). Pocahontas Co.: Cranberry Glades (MUMC 619). Tucker

Co.: Dolly Sods (MUMC 569). Wyoming Co.: SR 10, 4.8 km S of

Logan Co. line (MUMC 718).

Mesodon appressus (Say, 1821). — Populations of M. appressus are

most heavily concentrated in the southeast, with a single northern popu-

lation in Monongalia County (Fig. 10). Boone Co.: SR 3, 3.4 km E of

US 119 (MUMC 424); Peytona (MUMC 489). Cabell Co.: 8342 Big

Seven Mile Rd. (MUMC 38, 51). Fayette Co.: US 60, 7.5 km W of

Gauley Bridge (MUMC 223). Lincoln Co.: SR 10, 3.2 km W of West

Hamlin (MUMC 88); Alkol (MUMC 450). Logan Co.: Chief Logan

State Park (MUMC 588). Mingo Co.: Grey Eagle (MUMC 534).

West Virginia Polygyrid Snails

151

Figs. 13-17. County distributions of Mesodon in West Virginia. 13, M. pen-

nsylvanicus; 14, M. mitchellianus; 15, M. clausus; 16, M. rugeli; 17, M. inflect-

us. Fig. 18. County distribution of Stenotrema hirsutum in West Virginia.

152 Clement L. Counts, III

Summers Co.: SR 3, 3.4 km E of Bluestone Dam (MUMC 500); SR 3,

2.6 km E of Nimitz (MUMC 504). Wayne Co.: Cabwaylingo State Park

(MUMC 292).

Mesodon perigraptus (Pilsbry, 1894). — Populations of this snail

were found in the southern half of West Virginia, and a single popula-

tion in the eastern panhandle in Grant County (Fig. 11). Mercer Co.:

Flat Top (MUMC 311).

Mesodon dentiferus (Binney, 1837). — Populations were found in

the eastern part of the state, with an isolated population in Wayne

County and another in Wirt County (Fig. 12). Grant Co.: SR 91, 11.4

km E of Tucker Co. line (MUMC 465). Greenbrier Co.: SR 39, 24 km E

of Richwood (MUMC 357). Nicholas Co.: SR 39, 1.6 km W of Sum-

mersville (MUMC 227). Pocahontas Co.: SR 39. west entrance to

Monongahela National Forest (MUMC 206); Beartown State Park

(MUMC 568); SR 39, 11.2 km W of Cranberry Glades (MUMC 617).

Wayne Co.: Cabwaylingo State Park (MUMC 290).

Mesodon pennsylvanicus (Green, 1827). — This species was found

most frequently in the northern part of the state, with two isolated

southern populations (one each in Monroe and Wayne counties) (Fig.

13). Ohio Co.: SR 2, 1.6 km E of Warwood (MUMC 262). Wayne Co.:

SR 75, Lavalette (MUMC 97).

Mesodon mitchellianus (Lea, 1839). — The populations of M. mitch-

ellianus were widely scattered throughout the state, with the heaviest

concentration in the northern panhandle (Fig. 14). Other scattered pop-

ulations were found in the southern and central counties, with one pop-

ulation in westernmost Wayne County. Boone Co.: SR 3, 4.6 km E of

US 119 (MUMC 431, 497). Clay Co.: Elkhurst (MUMC 116). Ohio Co.:

Wheeling (MUMC 245); CR 15, 1.6 km E of Wheeling at Petes Creek

(MUMC 252); SR 2, 1.6 km E of Warwood (MUMC 263). Summers

Co.: SR 3, 3.4 km E of Bluestone Dam (MUMC 498). Wayne Co.: SR

75, Lavalette (MUMC 106).

Mesodon clausus (Say, 1821). — Three widely separated populations

of M. clausus were found (Fig. 15). Wayne Co.: SR 75, Lavalette

(MUMC 195).

Mesodon rugeli (Shuttleworth, 1852). — Populations of M. rugeli

were found only in the southern counties (Fig. 16). Fayette Co.: US 60,

7.5 km W of Gauley Bridge (MUMC 218).

Mesodon inflectus (Say, 1821). — Only one M. inflectus population

has been found in West Virginia (Fig. 17), and represents a new state

record. Fayette Co.: US 60, 7.5 km W of Gauley Bridge (MUMC 219).

Stenotrema hirsutum (Say, 1817). — Stenotrema hirsutum is the

most widely distributed species of the genus in West Virginia (Fig. 18).

Barbour Co.: Audra State Park (MUMC 605). Boone Co.: Peytona

West Virginia Polygyrid Snails

153

Figs. 19-23. County distributions of Stenotrema in West Virginia. 19, S. fra-

ternum; 20, 5. stenotrema; 21, S. leai; 22, S. edvardsi; 23, S. barbigerum. Fig.

24. County distribution of A llogona profunda in West Virginia.

154 Clement L. Counts, III

(MUMC 487); SR 3, 4.6 km E of US 119 (MUMC 493). Brooke Co.:

Bethany (MUMC 247). Cabell Co.: 8342 Big Seven Mile Rd. (MUMC

42); Kiwanavista Roadside Park, Ona (MUMC 547). Jackson Co.:

Ripley (MUMC 156). Lincoln Co.: West Hamlin (MUMC 90). Logan

Co.: Chief Logan State Park (MUMC 591). Mason Co.: McClintic

Wildlife Refuge Station (MUMC 62); Chief Cornstalk Public Hunting

Area (MUMC 66). McDowell Co.: US 52, 9.6 km W of Welch (MUMC

706). Nicholas Co.: Summersville (MUMC 226). Ohio Co.: Wheeling

(MUMC 244). Pocahontas Co.: SR 39, east entrance to Monongahela

National Forest (MUMC 208). Putnam Co.: Hurricane (MUMC 80);

SR 34, 3 km N of Kanawha River (MUMC 143); Clymer's Creek Rd.,

4.8 km off SR 34 (MUMC 181). Tyler Co.: SR 2, 3.4 km S of Wetzel

Co. line (MUMC 266). Wayne Co.: Lavalette (MUMC 203); Cabway-

lingo State Park (MUMC 204). Wirt Co.: SR 14, 4 km N of Roane Co.

line (MUMC 179).

Stenotrema fraternum (Say, 1824). — This species was also widely

distributed across the state, but was not found in the northern panhan-

dle (Fig. 19). Harrison Co.: Saltwell (MUMC 608). Jackson Co.: SR 39,

2.4 km W of Putnam Co. line (MUMC 124). Kanawha Co.: Nitro

(MUMC 121). Lincoln Co.: SR 10, 3.2 km W of West Hamlin

(MUMC 87). Mercer Co.: Flat Top (MUMC 304).

Stenotrema stenotrema (Pfeiffer, 1842). — Most populations of S.

stenotrema occur in the southern counties, with a single population in

the eastern panhandle (Fig. 20). Pocahontas Co.: SR 39, 4.8 km E of

Cranberry Glades (MUMC 372).

Stenotrema leai (Binney, 1841). — Stenotrema leai populations were

generally distributed through the east central counties, with one popula-

tion in the northern panhandle and another along the west central

border in the floodplain of the Ohio River (Fig. 21). Mason Co.: Chief

Cornstalk Public Hunting Area (MUMC 64).

Stenotrema edvardsi (Bland, 1856). — Populations of S. edvardsi

are concentrated in the southern counties (Fig. 22). Boone Co.: SR 3,

3.4 km E of US 119 (MUMC 423). Logan Co.: SR 10, 3.4 km E of

Davin (MUMC 393); Chief Logan State Park (MUMC 592). Mercer

Co.: Flat Top (MUMC 318). McDowell Co.: US 52, 9.6 km W of

Welch (MUMC 707). Raleigh Co.: Slab Fork (MUMC 346).

Stenotrema barbigerum (Redfield, 1856). — A single population of

this species was found in southern West Virginia (Fig. 23) and is a new

state record. Mercer Co.: Flat Top (MUMC 303).

Allogona profunda (Say, 1821). — Populations of A. profunda are

widely distributed over the state (Fig. 24). Ohio Co.: SR 2, 1.6 km E of

Warwood (MUMC 258).

West Virginia Polygyrid Snails 155

DISCUSSION

Pilsbry (1940) reported 12 polygyrid species in four genera in West

Virginia: Stenotrema stenotrema, S. edvardsi, S. hirsutum, S. frater-

num, Mesodon thyroidus, Triodopsis tridentata, T. t. juxtidens, T pla-

tysayoides, T. rugosa, T. albolabris, T. dentifera, and Allogona pro-

funda. Four of these were described as new species from collections

made in West Virginia. Triodopsis platysayoides was described from

specimens collected by S. T. Brooks in 1933 at Cooper's Rock, Monon-

galia County. Hubricht (1972) placed this snail on his endangered spe-

cies list, and no populations could be located during this study. Its sta-

tus as endangered may be the result of a restricted range, or of the

spring droughts of the 1970s that reduced many land snail populations

and could have caused the extirpation of localized colonies (Hubricht

1972).

Triodopsis rugosa was first collected from a deep ravine, 1.6 km

southwest of Blair, Blair Mountain, Logan County (Brooks and Mac-

Millan 1940). It was originally described as T tridentata rugosa and the

taxonomy was changed by Pilsbry (1940). Triodopsis fraudulenta was

described by Pilsbry (1894) from Morgan County and Stenotrema

edvardsi was first collected from the mountains of Fayette or Greenbrier

counties by Bland, who described the species in 1858 (Pilsbry 1940).

MacMillan (1949) published the latest report to specifically discuss

the land snails of West Virginia. Twenty-seven Polygyridae, in three

genera, were reported: Mesodon albolabris albolabris, M. a. dentatus,

M. profundus, M. mitchellianus, M. thyroidus, M. zaletus, M. pennsyl-

vanicus, M. sayanus, M. dentiferus, M. clausus, M. appressus, M. a.

perigraptus, Triodopsis platysayoides, T. fraudulenta fraudulenta, Tf

vulgata, T rugeli, T denotata, T tridentata tridentata, T t. juxtidens,

T rugosa, T fallax, Stenotrema edvardsi, S. fraternum fraternum, and

S.f cavum. I was unable to find T platysayoides, M. a. dentatus, T. t.

juxtidens, T fallax, and S.f cavum. Other new taxa have been reported

from West Virginia since the work of MacMillan, and these new species

have added to the confusion as to which species are present in the state.

Grimm (1971) described Stenotrema simile which, although the

type locality was Maryland, was also reported to be found in Mononga-

lia, Nicholas, and Pocahontas counties, West Virginia. He noted that

the chief differences between S. simile and S. hirsutum were the larger

size of S. simile, its coarser granulations on the embryonic whorls, and

other shell sculpture characteristics best seen with the scanning electron

microscope. Grimm (1974) also found intermediate populations of T

juxtidens juxtidens and T j. discoidea near the New River Gorge at

Hinton, Summers County. Vagvolgyi (1969) found similar populations

156 Clement L. Counts, III

in Clay County. None of these taxa (some as subspecies) could be found

during my study.

Hubricht (1976) described Mesodon panselenus from specimens

collected at Scotford, Clay County; Hernshaw, Kanawha County; Blair,

Logan County; and Iaeger, McDowell County. He reported that the

new species was similar to, and easily confused with, M. perigraptus,

but differed from it in having a more depressed shell and shorter penis.

My visits to the type locality and other collection localities listed by

Hubricht for M. panselenus failed to reveal specimens referable to this

species.

Hubricht (1974) placed Mesodon burringtoni Hubricht in syno-

nymy with M. mitchellianus. He noted that the first specimens of M.

burringtoni were collected in southwest Virginia and at two localities in

the Kanawha-New River valleys of West Virginia. However, as more

material became available, especially larger shells, no apparent differen-

ces could be found between the two species. In the same report,

Hubricht noted that topotypes of Stenotrema hirsutum were indistin-

guishable from S. burringtoni Grimm. Such taxonomic gyrations and

the inability to clearly define some species in the family Polygyridae

may account for some of the difficulties in accurately assessing the poly-

gyrid fauna of West Virginia.

Briscoe (1963) reported 8 species of Polygyridae in Jefferson County,

West Virginia, and reported populations of Mesodon thyroidus buccu-

lenta (Gould, 1848) from 11 localities within the county. However, I

could not locate populations of this subspecies during my study. Pilsbry

(1940) described the zoogeographic range of M. t. bucculenta to extend

from Wilmington, North Carolina south to Georgia and west to Arkan-

sas, Oklahoma, and Texas. He also noted that specimens referable to

M. t. bucculenta from Pennsylvania and other northern states are better

referred to M. t. thyroidus.

Triodopsis multilineata is found only on Blennerhassett Island in

the Ohio River, Wood County, West Virginia (Taylor and Counts

1976). No specimens of this species were collected from the West Virgi-

nia shore of the river during my study.

ACKNOWLEDGMENTS.— \ thank Drs. Ralph W. Taylor, Dan

K. Evans, and Donald C. Tarter, Marshall University, for their gui-

dance during the course of this study. Thanks are also due Dr. R.

Tucker Abbott, Delaware Museum of Natural History; Dr. Kenneth J.

Boss and Mr. David McHenry, Museum of Comparative Zoology,

Harvard University; and Mr. Leslie Hubricht, Meridian, Mississippi, for

their assistance in identifying snails. Further thanks are due the many

students and faculty members of Marshall University who generously

West Virginia Polygyrid Snails 157

assisted in the collection of snails for this study. I am also grateful to

three anonymous reviewers for their valuable comments and advice.

This paper is from a thesis presented to the faculty of Marshall

University in partial fulfillment of the requirements for the Master of

Science degree.

LITERATURE CITED

Briscoe, M. S. 1963. A survey of land and freshwater snails in Jefferson County,

West Virginia. Sterkiana 6:41-48.

Brooks, Stanley T., and G. K. MacMillan. 1940. New Gastropoda from West

Virginia. Nautilus 53:95-97.

Burch, John B. 1962. How to Know the Eastern Land Snails. William C. Brown

Co., Dubuque, Iowa, v + 214 pp.

, and C. M. Patterson. 1966. Key to the genera of land gastropods

(snails and slugs) of Michigan. Univ. Mich. Mus. Zool. Circ. No. 5. 19 pp.

Grimm, F. Wayne. 1971. Annotated checklist of the land snails of Maryland

and the District of Columbia. Sterkiana 47:51-57.

. 1974. Speciation within the Triodopsis fallax group (Pulmonata:

Polygyridae). Bull. Am. Malacol. Union Inc. 1974:23-29.

Hubricht, Leslie. 1972. Endangered land snails of the eastern United States.

Sterkiana 45:33.

. 1974. A review of some land snails of the eastern United States.

Malacol. Rev. 7:33-34.

. 1976. Five new species of land snails from the eastern United

States. Malacol. Rev. 9:126-130.

MacMillan, Gordon K. 1949. The land snails of West Virginia. Ann. Carnegie

Mus. 57:89-238.

Pilsbry, Henry A. 1894. Critical list of mollusks collected in the Potomac Val-

ley. Proc. Acad. Nat. Sci. Phila. 46:11-24.

. 1940. The land Mollusca of North America (north of Mexico).

Acad. Nat. Sci. Phila., Monogr. No. 3, Vol. 1: 575-944.

Taylor, Ralph W., and Clement L. Counts, III. 1976. Note on some land snails

from Blennerhassett Island, West Virginia. Sterkiana 63-64:11.

Vagvolgyi, Joseph. 1969. Systematics and evolution of the genus Triodopsis

(Mollusca: Pulmonata: Polygyridae). Bull. Mus. Comp. Zool. 75:145-254.

Accepted 9 July 1982

159

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CONTENTS

Terrestrial Isopods (Crustacea: Isopoda: Oniscoidea) from North Caro-

lina. George A. Schultz 1

Aquatic Macroinvertebrates of the Upper French Broad River Basin.

David L. Penrose, David R. Lenat and K. W. Eagleson 27

Northern Limits of the Southeastern Shrew, Sorex longirostris Bach-

man (Insectivora: Soricidae), on the Atlantic Coast of the United

States. John F. Pagels, Carol S. Jones and Charles O. Handley 51

Use of Lepomis macrochirus Rafinesque Nests by Spawning Notemi-

gonus crysoleucas (Mitchill) (Pisces: Centrarchidae and Cyprinidae).

David J. DeMont 61

Systematics of the Troglobitic Caecidotea (Crustacea: Isopoda: Asellid-

ae) of the Southern Interior Low Plateaus. Julian J. Lewis 65

Aboriginal and Modern Freshwater Mussel Assemblages (Pelecypoda:

Unionidae) from the Chickamauga Reservoir, Tennessee. Paul W.

Parmalee, Walter E. Klippel and Arthur E. Bogan 75

Notes on Distribution and Habitats of Sorex and Microsorex (Insecti-

vora: Soricidae) in Kentucky. Ronald S. Caldwell and Hal Bryan .... 91

Unionid Mollusca (Bivalvia) from Little South Fork Cumberland River,

with Ecological and Nomenclatural Notes. Lynn B. Starnes and

Arthur E. Bogan 101

Response of Small Mammals to Forest Clearings Created by Herbicides

in the Central Appalachians. William C. McComb and Robert L.

Rumsey 121

Spawning Behavior in Seven Species of Darters (Pisces: Percidae).

Lawrence M. Page, Michael E. Retzer and Robert A. Stiles 135

Occurrence and Distribution of Land Snails of the Family Polygyridae

(Mollusca: Gastropoda: Pulmonata) in West Virginia. Clement L.

Counts III 145

Errata and Miscellany 1 59

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