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number 13

july 1987

«W 11111

EDITORIAL STAFF

Eloise F. Potter, Acting Editor

Eloise F. Potter, Managing Editor

Sheree Worrell, Production Manager

John B. Funderburg, Editor-in-Chief

Board

James W. Hardin

Department of Botany

N.C. State University

William M. Palmer

Curator of Lower Vertebrates

N. C. State Museum

David S. Lee

Curator of Birds

N.C. State Museum

Rowland M. Shelley

Curator of Invertebrates

N.C. State Museum

Brimleyana, the Journal of the North Carolina State Museum of Natural

Sciences, will appear at irregular intervals in consecutively numbered issues.

Contents will emphasize zoology of the southeastern United States, especially

North Carolina and adjacent areas. Geographic coverage will be limited to Ala-

bama, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Missis-

sippi, 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

invertebrate 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 Editor and, where neces-

sary, the Editorial Board. Appropriate specialists will review each manuscript

judged suitable, and final acceptability will be determined by the Editor.

Address manuscripts and all correspondence (except that relating to subscrip-

tions and exchanges) to Editor, Brimleyana, North Carolina State Museum of

Natural Sciences, P. O. Box 27647, Raleigh, NC 27611.

Address correspondence pertaining to subscriptions, back issues, and ex-

changes to Shelly Turner, Brimleyana secretary. North Carolina State Museum

of Natural Sciences, P.O. Box 27647, Raleigh, NC 2761 1.

In citations please use the full name — Brimleyana.

North Carolina State Museum of Natural Sciences

North Carolina Department of Agriculture

James A. Graham, Commissioner

CODN BRIMD 7

ISSN 0193-4406

Life History of the Pinewoods Darter,

Etheostoma mariae (Osteichthyes: Percidae), a Fish

Endemic to the Carolina Sandhills

Fred C. Rohde and Steve W. Ross

North Carolina Division of Marine Fisheries,

7225 Wrightsville Avenue, Wilmington, North Carolina 28403

ABSTRACT. — Etheostoma mariae is restricted to streams in the

Carolina Sandhills area of North and South Carolina below the Fall

Line. It is found primarily in small creeks with gravel substrate, often

in or near vegetation. Twenty-five fish-associates were taken with E.

mariae. Spawning occurred from April to July at water temperatures

ranging from 14 to 21 °C. A strong relationship existed between

mature ova number and standard length (Log Y = -2.8523 + 2.8710

Log SL, r = 0.89); mean mature ovum diameter was 1.6 mm. Growth

was rapid in the first year with maturity occurring at the end of the

year. Maximum age of E. mariae was 36 months. Males were longer

than females, with maximum lengths attained of 60 mm and 51 mm

SL, respectively. The sex ratio was 2.06 females: 1 male. Females were

slightly heavier than males at the same length. Commonly eaten foods

were dipteran larvae (in 86.8% of stomachs), ephemeropteran nymphs,

trichopteran larvae, and plecopteran nymphs. Chironomids were the

dominant food in all months except May, June, and October when

simuliids were most common. Chironomids were important foods for

smaller fish (< 40 mm) with simuliids gaining importance in larger E.

mariae.

The pinewoods darter, Etheostoma ( Belophlox ) mariae (Fowler), is

endemic to streams in the Carolina Sandhills where it is restricted to the

i

Lumber River (Little Pee Dee system) in North Carolina and to the

Little Pee Dee River in North and South Carolina. Contrary to the

statement made by Kuehne and Barbour (1983), this species is found

only below the Fall Line (Fig. 1). The single South Carolina record (two

specimens) is from Panther Creek (reported as Beaver Dam Creek by

Richards 1963). This darter is more commonly encountered in the

Lumber River headwaters than in those of the Little Pee Dee. Because

of its limited range, the presence of significant pollution in the upper

Lumber River, and several plans to channelize and drain parts of this

river, Bailey (1977) considered it to be a species of special concern in

North Carolina. Its status is also of concern to the U.S. Fish and Wild-

life Service (R. Biggins, pers. comm.).

Except for a brief discussion of habitat and species associates in

the systematic treatment of E. mariae and E. fricksium by Richards

Brimleyana No. 13:1-20, July 1987

1

2

Fred C. Rohde and Steve W. Ross

(1963), and the summary of biological information on E. okaloosae by

Yerger (1978), little published biological information is available on the

three members of the subgenus Belophlox. In order to add to our

knowledge of E. mariae, we studied a population in Naked Creek,

North Carolina, and report here on reproduction, sexual dimorphism,

age and growth, and food habits. The information gained can be used in

future management plans for the species and may be useful in predict-

ing environmental requirements for unstudied close relatives.

STUDY AREA

Naked Creek lies primarily in Richmond County, approximately 20

km west of Aberdeen, Moore County, North Carolina (Fig. 1). It origi-

nates in Montgomery County and flows 21 km southeast to join Drown-

ing Creek, a tributary of the Lumber River. The creek is typically bor-

dered by lowland forest of tag alder, Alnus serrulata\ holly, Ilex opaca\

red maple, Acer rubrum\ and leucothoe, Leucothoe axillaris . Turkey

oak, Quercus laevis, and scrubby post oak, Q. stellata var. margaretta,

occur in the higher areas.

Two locations were sampled for E. mariae (Fig. 1). Site 1 was

downstream of State Route 1003 bridge just above the confluence with

Drowning Creek. Most collections at this location were made about 1 to

20 m below the bridge over sand, gravel, and rubble where the creek

averaged 5.5 m wide and 0.5 m deep. Most of the E. mariae were col-

lected over gravel. Young were often found in vegetated areas with a

slow current. Dominant aquatic vegetation was filamentous algae;

water moss, Fontinalis sp.; pondweed, Potamogeton diver sifolius; tape-

grass, Vallisneria americana\ and spatter-dock, Nuphar luteum. Upstream

and downstream of this area the creek was deeper and narrower with a

silt bottom. Water temperatures ranged from 2.2 to 21.7 °C. Site 2 was

upstream of State Route 1321 bridge near the headwaters, where the

creek averaged 4 m wide and 0.3 m deep. A large gravel riffle (7 m x 4

m) profusely covered by tapegrass was about 3 m upstream of the

bridge. Because most E. mariae were located throughout this riffle,

sampling effort was concentrated there. Immediately below and above

the riffle were pools to 1.2 m deep with silt bottoms. A few specimens,

primarily young, were collected in these pools. Water temperatures were

similar to those recorded downstream at Site 1.

Twenty-five fish species were taken with E. mariae (number of

times in parentheses) in a total of 23 collections made in the sampling

areas: Anguilla rostrata (8), Esox americanus (5), E. niger (1), Nocomis

leptocephalus (1), Notropis cummingsae (16), Semotilus lumbee (3),

Erimyzon oblongus (1), Ictalurus natalis (4), Noturus gyrinus (5), N .

Life History of Pinewoods Darter

3

Fig. 1. Distribution of Etheostoma mariae. Circles represent localities where

species was collected; triangles represent study sites in Naked Creek.

4

Fred C. Rohde and Steve W. Ross

insignis (5), Chologaster cornuta (3), Aphredoderus sayanus (5), Fundu-

lus lineolatus (1), Gambusia affinis (1), Acantharchus pomotis (2), Elas-

soma zonatum (7), Enneacanthus chaetodon (3), E. gloriosus (3), Lepomis

auritus (4), L. marginatus (1), Micropterus salmoides (1), Etheostoma

fusiforme (1), E. olmstedi (14), E. serriferum (6), and Percina crassa (4).

Four other vertebrates were seined with E. mariae : Notophthalmus viri-

descens, Necturus punctatus, Desmognathus fuscus, and Sternotherus

odoratus.

MATERIALS AND METHODS

We collected 485 specimens of E. mariae with a 3.0 m x 1.2 m, 3.2

mm mesh nylon flat seine. Collection dates were 4 January; 18 Febru-

ary; 16 March; 2*, 9, and 23 April; 12 May; 11 June*; 21 July*; 22

August*; 24 September*; 28 October*; 26 November; and 9 December*

in 1978, and 13 April* in 1982 (* = both sites sampled this date). Collec-

tion times were between 0920 and 1700 hours. Unless noted, data refer

to 1978. Fish were preserved in 10% formalin and stored in 40% iso-

propyl alcohol until examined. For conservation purposes, specimens of

E. mariae in each collection made in June, July, and August were

placed alive in a water-filled plastic pan with a ruler and photographed.

Measurements from these photographs were later used to obtain length-

frequency data. Twenty fish were then selected and preserved for data

on food, reproduction, and age, and the remainder were released. Air

and water temperatures were recorded at time of capture.

Standard length (SL) was measured to the nearest mm and body

weight to the nearest 0.01 g. Left and right gonads were excised (both

sexes) and were weighed together to the nearest 0.001 g. Gonads < 0.001

g were not included. A gonosomatic ratio (gonad weight as a percentage

of total body weight) was calculated to examine seasonal development

of the gonads. Ovaries were teased apart under a dissecting microscope

and the somewhat irregularly shaped ova were measured across their

widest dimension to the nearest 0.1 mm with an ocular micrometer. Ova

from developing or ripe ovaries were grouped using the criteria defined

in Table 1. Ova classes were usually distributed randomly in the ovaries.

Fecundity was determined for 35 females collected in April (32) and

May (3) by counting all mature (Class I and II, > 1.3 mm diam.) ova in

both gonads.

Ages of 313 specimens were determined from scales removed from

just above the lateral line between the spinous and soft dorsal fins. Four

to seven unregenerated scales from each fish were cleaned, mounted on

a microfiche reader, and analyzed under 36X magnification. Annuli

distances and total scale sizes were measured along an axis from the

Life History of Pinewoods Darter 5

Table 1. Ova classification of Etheostoma mariae from Naked Creek, North

Carolina.

focus to the middle of the anterior field on one representative scale per

fish. Age (year class) was determined from a count of scale annuli. Since

peak spawning was in April, May was designated as month one (Page

1974). Age class 0 included young-of-the-year fish from 1 through 12

months of age, age class 1 those from 13 through 24 months, and age

class 2 those fish from 25 through 36 months. Because of the extended

spawning season, the possibility of an aging error of as much as three to

four months existed. However, since the vast majority of spawning

occurred in April, we discounted this possibility.

Based on examination of 321 stomachs, we determined the number

and kinds of food items in five size groups of fish (< 20 mm, 20-29 mm,

30-39 mm, 40-49 mm, > 49 mm), and the percent occurrence of food by

month.

Locality data for E. mariae were obtained from Richards (1963),

from the fish collections at the University of North Carolina at Char-

lotte, and from collections at the North Carolina State Museum of Nat-

ural History. During our study additional sites were sampled through-

out the known range of E. mariae to document distribution and

population sizes. Specimens from these collections were not used to

obtain life history data.

RESULTS AND DISCUSSION

Reproduction. — Spawning occurred from April through July with an

April and May peak (Fig. 2). Water temperature during the spawning

6

Fred C. Rohde and Steve W. Ross

period ranged from 14 to 18 °C in April to 21 °C in June and July. The

female gonosomatic ratio was low in January, increased rapidly in Feb-

ruary and March, and peaked in April and May at maximum spawning

(Fig. 2). Values dropped rapidly in June and July, reflecting the large

number of spent females, and decreased sharply until September, after

which the values increased as ova production began in maturing fish.

The monthly gonosomatic ratio pattern of males was similar to that of

females, but not as pronounced (Fig. 2).

Few females were in spawning condition at the same time; only 8 of

the 43 mature females collected on six dates from April through July

contained ripe ova (Class I). Two of the eight mature females (May,

June) were spent, as evidenced by the presence of but a single mature

ovum in the flacid ovary. Ripe ova were first found in two females

collected on 2 April and were last observed in two females taken on 21

July, indicating a long spawning period. On 13 April 1982, six of the

fourteen females taken contained ripe ova, while the rest contained

developing ova.

Other Southeastern members of the genus Etheostoma tend to have

long spawning periods: two months for E. simoterum (Page and Mayden

1981); three months for E. striatulum (Page 1980) and E. coosae (O’Neil

1981); and four months for E. ditrema (Ramsey and Suttkus 1965), E.

okaloosae (Yerger 1978), and E. perlongum (Lindquist et al. 1981).

Taber and Taber (1983) observed a similar protracted spawning period

(March-July) for E. tetrazonum in Missouri, combined with the produc-

tion of many small clutches of eggs. They suggested that these two fac-

tors enhance the reproductive success of this species in a fluctuating

riffle habitat. Hubbs (1985) suggested multiple spawnings and clutches

by a single female by a number of darters with protracted spawning

periods. We found no evidence to suggest that E. mariae has multiple

spawnings.

Fecundity was calculated only for peak spawning periods (April,

May). Mean mature (Class I and II) ova number per female (N = 35)

was 57.6 ± 3.4SE (range 21-109). In April 1982, mean ova number of 14

females was 59.7 ± 5.7SE (range 38-1 17). These estimates of mature ova

are similar to those observed in a number of darters, as summarized by

Page (1983). The regression equation for the relationship between SL

and mature ova number (Y) in 1978 is Log Y = -2.8523 + 2.8710 Log SL,

r = 0.89 (P < 0.001), indicating a significant increase in ova production

with increasing length.

The regression equation for the relationship of mature ova number

(Y) to body weight (W) is Log Y = -1.5248 + 0.9643 Log W, r = 0.90, and

the relationship of ova number to ovary weight (OW) is Log Y = 2.2735

+ 0.5994 Log OW, r = 0.89. These correlation coefficients are highly

Life History of Pinewoods Darter

7

Fig. 2. Mean (± SE) male and female gonosomatic ratios for Etheostoma

mariae from Naked Creek, North Carolina, January through December 1978.

Numbers refer to number of specimens examined.

8

Fred C. Rohde and Steve W. Ross

significant (P < 0.001), and indicate that either body weight or ovary

weight is a good predictor of fecundity. However, these parameters are

more variable than standard length and, unless measured just prior to

spawning, will vary seasonally. Because ovary weight was included in

the body weight, some autocorrelation was possible.

Females reached sexual maturity by the end of the first year of life.

Of the 35 gravid females examined, 14 were completing the first year of

life, 19 were age class 1, and 2 were age class 2. Nine other young-of-

the-year darters were immature. Males also reached sexual maturity at

the end of the first year of life based on the presence of large, white and

spongy testes.

Breeding tubercles were never noted. The genital papilla of mature

females is a long, prominent tube slightly crenulate at the tip and about

two-thirds the length of the anal spine. The conical structure of the

papilla and diameter of ripe eggs suggest that this species is an egg-

burier (Page 1983). The male genital papilla is a small projection and

does not enlarge at spawning.

Sexual dimorphism in spawning colors is pronounced. Body colo-

ration of the male is gold dorsally to yellow-tan ventrally. Dark blotches

occur along the sides and tend to merge with each other. The lateral line

is bright gold and bisects these blotches. The. margin of the spinous

dorsal fin is clear to white, and immediately below is a wide orange-red

band. The interspinous membranes below this band are dark, particu-

larly obvious in the anterior three. Fin rays in the soft dorsal, caudal,

anal, and pelvic fins are outlined in yellow-orange. The interradial

membranes of these fins, plus the pectoral fin rays, are dusky blue.

Brown spots on the soft dorsal and caudal fins form four slightly irregu-

lar bands. The procurrent rays of the caudal fin are bright blue-green,

and this coloration continues anteriorly onto the dorsal and ventral

edges of the caudal peduncle. The iris of the eye is bright orange. The

male coloration in this study differed slightly from that reported by

Kuehne and Barbour (1983) and Page (1983) in that all the fins were

darkened, the marginal band was clear to white, and the venter was

yellow-tan. Neither mentioned the bright blue-green edging on the cau-

dal peduncle and fin.

Breeding colors in the female are poorly developed. An orange-red

band develops in the spinous dorsal fin, but the white marginal band

and darkened interspinous membranes are not as pronounced as in the

male. Kuehne and Barbour (1983) noted that the spiny dorsal fin was

more spotted than banded. Other fin pigmentation is similar to the

male, except for a lack of darkening of the interradial membranes.

Life History of Pinewoods Darter

9

Age and Growth. — Apparently one mark (annulus) was formed on

scales per year. Minimum marginal increments (Bagenal and Tesch

1978) in 1-year-old fish occurred during March and April (Table 2).

Although sample sizes were small, 2- and 3-year-old fish also had small

scale marginal increments during spring (Table 2). Consistency of time

of mark formation between age classes, strong relationship between fish

length and scale size (r = 0.97), and increasing fish size with increasing

number of scale marks (Table 3) indicated that marks on scales were

valid annuli. Annuli exhibited many of the characteristics described by

Lachner et al. (1950) for other darters, and annuli were apparent in both

the anterior and lateral fields (Fig. 3). Although some annuli were faint,

only one of the 313 fish had unreadable scales.

Because of the extended spawning season and the rapid growth of

young, growth by month of the various year classes was difficult to

distinguish in a length-frequency graph (Fig. 4). Earliest young-of-the-

year (age class 0) recruits were first collected in June and July and

“recruitment” continued into September (Fig. 4). Fish <15 mm SL were

not efficiently collected by our gear. Late-spawned young-of-the-year

appeared as a minor component from October through December. In

any given month each age class exhibited a wide range of sizes, presum-

ably reflecting the long spawning season (Fig. 5). Our data also indi-

cated a broad overlap in sizes of age classes, particularly age classes 1

and 2 in October and all three age classes in January (Fig. 5). The two

largest males (60 mm SL) were collected in September but differed

greatly in age (17 and 29 months). The largest female was 51 mm SL

and 36 months old.

Growth was most rapid during the first year. Males and females

attained 58% and 66%, respectively, of their observed average maximum

lengths (Table 3). During this year there were no significant sexual dif-

ferences in growth rate (t = 1.32, df = 173, P > 0.05). However, in age

classes 1 and 2 males grew significantly longer (t = 3.66, df = 91, P <

0.005 for age class 1; t = 3.22, df = 33, P < 0.005 for age class 2) (Fig. 6).

In most collections males were larger than females of the same age (Fig.

5). Significant differences also existed between males and females in

weight-length relationships as indicated by analysis of covariance (F =

12.38, df = 1,318, P < 0.05). Females were slightly heavier than males of

the same length, with a weight-length relationship of WT = 8.1 x 10~6

SL3 23, r = 0.98. Male weight-length relationship was WT = 12.6 x 10'6

SL3 09, r = 0.98. Because all fish were used to develop the weight-length

relationships, some of the intersexual differences could be attributed to

the females’ having a greater increase in body weight than do males

during spawning (Fig. 2).

10

Fred C. Rohde and Steve W. Ross

Table 2. Monthly mean scale marginal increments (MI), standard deviation

(SD), and ranges for Etheostoma mariae from Naked Creek, North

Carolina. Fish with 2 and 3 marks were combined because of the

small sample size.

Life History of Pinewoods Darter

11

Fig. 3. (Left) Scale from 60 mm SL male E. mariae collected 24 September 1978. Two annuli on this 29-month-old fish are

illustrated. (Right) Scale from 48 mm SL female collected 23 April 1978. Three annuli are apparent on this 36-month-old fish.

12

Fred C. Rohde and Steve W. Ross

3-

1-

M ar

n=12

Standard Length (mm)

Fig. 4. Monthly length frequencies of Etheostoma mariae from Naked Creek,

North Carolina, January through December 1978.

Life History of Pinewoods Darter 13

Fig. 5. Length-age relationship by sex of Etheostoma mariae from Naked Creek, North Carolina, January through December 1978.

14

Fred C. Rohde and Steve W. Ross

Fig. 6. Annual growth of Etheostoma mariae males and females from Naked

Creek, North Carolina. Solid line = range; triangle = mean; rectangle = 95%

confidence interval.

Regression of standard length onto scale radius (SR) yielded the

formula: SL = 7.47 + 0.83 SR, r = 0.97. This was used to back-calculate

to size at annulus formation by inserting measurement to annulus in

place of scale radius. Back-calculated sizes at age 1 from age classes 1

and 2 were in close agreement with observed standard lengths (Table 3)

with only a slight tendency toward Lee’s phenomenon, which we attrib-

uted to selective natural mortality favoring survival of younger fish

(Bagenal and Tesch 1978). Back-calculation indicated a small mean

growth increment between ages 1 and 2 of 13.7 mm for males and 11.2

mm for females (Table 3).

Rapid growth of young-of-the-year darters is typical (e.g. Raney

and Lachner 1943, Burr and Page 1978, Pflieger 1978, O’Neil 1981,

Shute et al. 1982). Growth differences by sex were not significant in

immature E. mariae', however, males were significantly larger when fish

were older (Fig. 6). Males of the related E. okaloosae are also larger

than females (Collette and Yerger 1962). Raney and Lachner (1943) and

Lachner et al. (1950) have shown that this is a common phenomenon

for species in which the male guards spawning areas. Page (1974, 1975)

and Page and Burr (1976) found that mature males were larger than

females in several members of the subgenus Catonotus ( E . squamiceps.

15

Life History of Pinewoods Darter

E. kennicotti, E. smithi), and males of these species were also strongly

territorial at spawning. Another tendency common in territorial percids

is that females usually outnumber males (Lachner et al. 1950), as also

noted in E. mariae (Table 4). The difference between number of males

(99) and females (204) was highly significant (chi-square P < 0.001,

expected ratio 1:1). Although we did not observe territoriality, the

larger size of mature males, preponderance of females, and sexual di-

chromatism during the breeding season suggest that such behavior

occurs.

The oldest E. mariae were 36 months (age class 2). Two fish had just

formed a third annulus. It is doubtful that E. mariae lives four years.

The majority (58.7%) were in age class 0, 30.1% in age class 1, and

11.2% in age class 2 (Table 4). Making the same assumptions as Page

(1974), we found that the survival values as based on Table 5 indicated a

low survival rate in the third year of life. Males had a 53% survival to

age class 1 and a 10% survival to age class 2. Females had a 54% survi-

val to age class 1, but a survival to age class 2 (25%) that was much

greater than that of males (Table 5). Male E. mariae exhibited a faster

growth rate and a higher mortality rate in the older ages than did

females.

Food Habits. — Based on stomach contents of 321 fish (11 to 59 mm

SL), 13 animal taxa were identified (Table 6). Only 25 (7.8%) stomachs

were empty. The majority of E. mariae ate larval dipterans (in 86.8% of

stomachs). Other commonly ingested invertebrates were ephemerop-

teran nymphs (28.0%), trichopteran larvae (26.0%), and plecopteran

nymphs (22.0%) (Table 6). Food habits of E. mariae were similar to

those of the few Southeastern darters studied (see references in Page

1983).

Seasonal variation in diet was evident (Table 6). Dipterans, primar-

ily Chironomidae and Simuliidae, were eaten in all months. Chirono-

mids were the most commonly occurring food in all months except May,

June, and October, when simuliids were most frequent. Ephemerop-

terans were important in May and June. Trichopterans, mainly Hydro-

psychidae, were common in April and plecopterans were commonly eaten

in January, March, and December. The number of taxa represented in

the stomachs per fish per month was lowest in the coldest months,

November through March (3-5 taxa) excepting December, and highest

during summer through mid-fall (6-8 taxa) (Table 6).

Prey taxa differed by fish size groups (Table 7). Chironomids were

most important in the diet of small fish (< 40 mm). Larger darters also

ate large numbers of chironomids but consumed even more of the larger

simuliids. Plecopterans, ephemeropterans, and trichopterans were eaten

16

Fred C. Rohde and Steve W. Ross

Table 4. Numbers of Etheostoma mariae by sex in each age class collected

from Naked Creek, North Carolina, and aged by scale reading.

* Nine of 312 fish aged were unsexed (immature).

Table 5. Relative survival values by sex of Etheostoma mariae year classes.

in increasingly greater numbers as fish size increased. Hydropsychids

were the second most important food of the largest darters (> 49 mm).

Although not important overall, ostracods were most evident in the diet

of young fish (< 30 mm). Such feeding on small crustaceans by young

darters, followed by an increse in use of larger aquatic insects as the fish

grows, is common among darters (e.g. Scalet 1972, Page 1983).

The diet of E. mariae reflects its habitat. Young darters may

inhabit quiet waters near shore and in deep pools, where they consume

large numbers of ostracods that also prefer such areas (Pennak 1978).

Adult darters were more common in the swift portions of the stream.

They fed primarily on insects either attached to, crawling on (e.g. dip-

terans, trichopterans, ephemeropterans), or beneath (e.g. plecopterans)

the gravel/ rubble substrate. The proportion in which these insects are

consumed is dependent upon a combination of preference, availability,

and vulnerability of prey present.

Table 6. Percent occurrence by month of food items in stomachs of Etheostoma mariae from Naked Creek, North Carolina, in 1978.

Food organism Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total

Life History of Pinewoods Darter

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18

Fred C. Rohde and Steve W. Ross

Table 7. Mean number of food items by length groups in stomachs of Etheo-

stoma mariae from Naked Creek, North Carolina, in 1978.

Etheostoma SL (mm)

statistical matters. E. F. Menhinick, University of North Carolina at

Charlotte, and W. M. Palmer, North Carolina State Museum of Natu-

ral History, Raleigh, provided E. mariae locality data. R. G. Arndt,

Stockton State College, made helpful comments on drafts of the

manuscript. W. B. Harris, University of North Carolina at Wilmington,

provided information concerning the Fall Line. G. W. Link, Jr., Univer-

sity of North Carolina, gave photographic assistance. We appreciate the

efforts of Dee Willis in typing this manuscript.

LITERATURE CITED

Bagenal, Timothy B., and F. W. Tesch. 1978. Age and growth. Pages 101-136 in

T. Bagenal, editor. Methods for Assessment of Fish Production in Fresh

Waters. IBP Handbook No. 3. 3rd ed. Blackwell Sci. Publ., Oxford.

19

Life History of Pinewoods Darter

Bailey, Joseph R., and Committee. 1977. Freshwater fishes. Pages 265-298 in J.

E. Cooper, S. S. Robinson, and J. B. Funderburg, editors. Endangered and

Threatened Plants and Animals of North Carolina. N.C. State Mus. Nat.

Hist., Raleigh.

Burr, Brooks M., and L. M. Page. 1978. The life history of the cypress darter,

Etheostoma proeliare, in Max Creek, Illinois. 111. Nat. Hist. Surv. Biol.

Notes 106.

Collette, Bruce B., and R. W. Yerger. 1962. The American percid fishes of the

subgenus Villora. Tulane Stud. Zool. 9:213-230.

Hubbs, Clark. 1985. Darter reproductive seasons. Copeia 1985( 1 ): 56-68.

Kuehne, Robert A., and R. W. Barbour. 1983. The American Darters. Univ.

Press Kentucky, Lexington.

Lachner, Ernest A., E. F. Westlake, and P. S. Handwerk. 1950. Studies of the

biology of some percid fishes from western Pennsylvania. Am. Midi. Nat.

43:92-111.

Lindquist, David G., J. R. Shute, and P. W. Shute. 1981. Spawning and nesting

behavior of the Waccamaw darter, Etheostoma perlongum. Environ. Biol.

Fishes 6(2): 177-191.

O’Neil, Patrick E. 1981. Life history of Etheostoma coosae (Pisces: Percidae) in

Barbaree Creek, Alabama. Tulane Stud. Zool. 23:75-83.

Page, Lawrence M. 1974. The life history of the spottail darter, Etheostoma

squamiceps, in Big Creek, Illinois, and Ferguson Creek, Kentucky. 111. Nat.

Hist. Surv. Biol. Notes 89.

. 1975. The life history of the stripetail darter, Etheostoma kennicotti,

in Big Creek, Illinois. 111. Nat. Hist. Surv. Biol. Notes 93.

. 1980. The life histories of Etheostoma olivaceum and Etheostoma

striatulum, two species of darters in central Tennessee. 111. Nat. Hist. Surv.

Biol. Notes 1 13.

. 1983. Handbook of Darters. T. F. H. Publ., Neptune, N.J.

, and B. M. Burr. 1976. The life history of the slabrock darter,

Etheostoma smithi, in Ferguson Creek, Kentucky, 111. Nat. Hist. Surv. Biol.

Notes 99.

, 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 1 17.

Pennak, Robert W. 1978. Fresh-water Invertebrates of the United States. 2nd

ed. John Wiley, New York.

Pflieger, William L. 1978. Distribution, status, and life history of the Niangua

darter, Etheostoma nianguae. Missouri Dep. Conserv. Aquatic Series 16.

Ramsey, John S., and R. D. Suttkus. 1965. Etheostoma ditrema , a new darter

of the subgenus Oligocephalus (Percidae) from springs of the Alabama

River basin in Alabama and Georgia. Tulane Stud. Zool. 12(3):65-77.

Raney, Edward C., and E. A. Lachner. 1943. Age and growth of johnny darters,

Boleosoma nigrum (Storer) and B. longimanum (Jordan). Am. Midi. Nat.

29:229-238.

Richards, William J. 1963. Systematic studies of darters from southeastern

United States (Pisces, Percidae). Unpubl. Ph.D. Dissert., Cornell Univ.,

Ithaca.

20

Fred C. Rohde and Steve W. Ross

Scalet, Charles G. 1972. Food habits of the orangebelly darter, Etheostoma

radiosum cyanorum (Osteichthyes: Percidae). Am. Midi. Nat. 87:515-522.

Shute, Peggy W., J. R. Shute, and D. G. Lindquist. 1982. Age, growth and early

life history of the Waccamaw darter, Etheostoma perlongum. Copeia

1 982(3):56 1-567.

Taber, Charles A., and B. A. Taber. 1983. Reproductive biology and age and

growth of the Missouri saddled darter Etheostoma tetrazonum. Am. Midi.

Nat. 109:222-229.

Yerger, Ralph W. 1978. Okaloosa darter. Pages 2-4 in C. R. Gilbert, editor.

Rare and endangered biota of Florida. Vol. 4. Fishes. Univ. Presses Flor-

ida, Gainesville.

Accepted 13 August 1985

Batriasymmodes from Caves in the Virginias

(Coleoptera: Pselaphidae)

Thomas C. Barr, Jr.

School of Biological Sciences,

University of Kentucky, Lexington, Kentucky 40506

ABSTRACT. — Batriasymmodes parki, n. sp., is described and illus-

trated from a cave in Mercer County, West Virginia, and B. monstro-

sus (LeC.) is newly recorded from a cave in Lee County, Virginia.

The batrisine pselaphid beetles of the genus Batriasymmodes Park

differ from the more common species of Batrisodes in the absence of a

metatibial spur and in the large aedeagus with well-differentiated, spe-

cific diagnostic characters. Males typically bear a recurved spine on the

metatrochanter. Originally proposed as a subgenus of Batrisodes (Park

1951), the group was elevated to generic status (Park 1960) and eventu-

ally revised (Park 1965) to include 14 species. The majority of its species

occur in forests in the Unaka mountain province in eastern Tennessee

and western North Carolina or in caves of Alabama, Tennessee, and

southern Kentucky. Two species are known from central Florida, and

B. monstrosus (LeC.) is rather widely distributed in northeastern United

States (see Park 1965). The four cavernicole species are probably all

troglophiles rather than troglobites, and they exhibit no troglomorphic

features suggesting long-term adaptation to caves or regressive charac-

ters that might restrict them to caves. Furthermore, they usually retain

metathoracic wings.

Two male Batriasymmodes specimens sent to me for study by John

R. Holsinger extend the known range of cavernicole representatives of

the genus northward from Sullivan County, Tennessee (B. greeveri Park

1965), into southwest Virginia and southeast West Virginia.

Batriasymmodes monstrosus (LeConte)

One male of this widely distributed northeastern species, with meta-

thoracic wings, was taken by J. R. Holsinger and D. C. Culver in Poor

Farm Cave, Lee County, Virginia, 27 August 1971. The specimen is

slightly aberrant in obsolescence of the oblique vertexal carinae and the

weak but distinct median vertexal carina, but such irregularities are not

uncommon among species of the genus. The distinctly modified protibia

and mesofemur are highly diagnostic, as is the structure of the aedeagus.

This species has not been previously recorded from caves.

Brimleyana No. 13:21-24, July 1987

21

22

Thomas C. Barr, Jr.

Batriasymmodes (s. str.) parki, new species

Fig. 1

Etymology. — It gives me great pleasure to name this species in

honor of my late colleague and friend, Professor Orlando Park.

Diagnosis. — A species of Batriasymmodes, s. str., resembling quis-

namus Park in antennal characters but with the general aedeagal pat-

tern of monstrosus (LeC.) and greeveri Park; projections of ventral plate

and movable process elongate as in greeveri, but apex of movable pro-

cess very deeply bifid and bent at right angle to left.

Description. — Total length 2.96 mm (holotype). Head 0.66 mm long

x 0.54 mm wide, pronotum 0.60 mm long x 0.56 mm wide, elytra 0.86

mm long x 0.96 mm wide, abdomen 0.98 mm long x 0.82 mm wide.

Head with pair of nude vertexal foveae, median vertexal carina fine,

oblique vertexal carinae replaced by low, noncarinate ridges; eye ovoid,

rather strongly convex, slightly emarginate on posteroventral side, with

about 40 facets. Antenna with club formed by distal three segments;

VIII with large, triangular, laminoid spine directed distally from ventral

side; IX with very deep, transverse and slightly oblique ventral groove,

distal margin of which is fimbriate and overhangs the cavity as in quis-

namus, but much more strongly developed; X slightly knobbed beneath

and narrowed apically; VII and XI normal, without special modifica-

tions. Pronotum about as in suteri Park, with deep median fovea

flanked by an elongate, laminoid spine either side; lateral to each spine

a fovea which is continued apically as a shallow sulcus. Each elytron

with three nude antebasal foveae and flank with one nude subhumeral

fovea, as in suteri and greeveri. Metathoracic wings present. Abdomen

with five simple tergites and five sternites, second sternite with trans-

verse tumulus each side, fifth sternite with large, shallow concavity.

Legs with protibia bearing conspicuous arcuation on distal half of dor-

somesial face, all femora simple; each metatrochanter with short, re-

curved spine. Aedeagus with movable process and medial process of

ventral plate elongate, as in greeveri; apex of movable process deeply

bifid and bent laterad at right angle. Female unknown.

Holotype. — Male, a unique, Neely Farm Cave, just southeast of

Athens, Mercer Co., West Virginia, 31 May 1969, J. R. Holsinger and

Roger Baroody. Type deposited in Field Museum of Natural History,

Chicago.

Discussion. — This is the 15th species to be assigned to Batriasym-

modes and the fifth cavernicole in the genus. It is most closely allied to

quisnamus and greeveri; in Park’s (1965:186) key to species it will run

to greeveri, a species from a cave in Sullivan County, Tennessee, close

Batriasymmodes from Virginia Caves

23

>

0

0 .3 m m

Fig. 1. Batriasymmodes parki, n. sp., aedeagus of holotype, dorsal view.

24

Thomas C. Barr, Jr.

to the Virginia border and about 150 km southwest of Neely Farm

Cave. In comparison with greeveri, however, the concavity in antennal

segment IX is much more extensive, the oblique vertexal ridges are not

carinate, the pronotal spines are not continued forward, and the prox-

imal margin of the protibial groove is not cuspate. The aedeagi of the

two species are clearly similar in form, but in B. parki the arc formed by

the two processes of the ventral plate is more circular than oblong-oval,

and the movable process is straighter and sharply bent to the left at its

apex. Whether parki is primarily a cavernicole, or whether it will even-

tually be found in epigean habitats like monstrosus, remains to be

determined.

ACKNOWLEDGMENTS. — I thank J. R. Holsinger for the speci-

mens on which this paper is based and John Wagner for reviewing the

manuscript. This study was supported in part by NSF 82-02339.

LITERATURE CITED

Park, Orlando. 1951. Cavernicolous pselaphid beetles of Alabama and Ten-

nessee, with observations on the taxonomy of the family. Geol. Surv. Ala.

Mus. Pap. 31:1-107.

. 1960. Cavernicolous pselaphid beetles of the United States. Am.

Midi. Nat. 64:66-184.

. 1965. Revision of the genus Batriasymmodes (Coleoptera: Psela-

phidae). Trans. Am. Microsc. Soc. 84:184-204.

Accepted 17 June 1985

Zoogeography of the Freshwater Fish Fauna

of Southern Georgia and Peninsular Florida

Carter R. Gilbert

Florida State Museum,

University of Florida, Gainesville, Florida 3261 1

ABSTRACT. — The freshwater fish fauna of southern Georgia and

peninsular Florida is impoverished in comparison with those of other

areas of southeastern United States, but at the same time displays a

relatively high percentage of endemism at species, subspecies, or racial

levels. Historical factors relating to this are reviewed, from which it is

concluded that peninsular Florida had its origin in the middle Oligo-

cene epoch (about 30 million years before present), and that subse-

quent pre-Pleistocene fluctuations in sea level resulted in complete

insularization of the peninsula on at least two separate occasions. This

and various related zoogeographic phenomena are discussed.

Peninsular Florida is an important area of biological endemism

(Neill 1957). It is generally accepted that this endemism has, in most

cases, resulted from physical separation of the peninsula from the rest of

southeastern United States by a saltwater barrier located in the northern

part (“neck”) of the peninsula. Until recently, this insularization was

believed to have been strictly a Pleistocene event, resulting from fluctua-

tions in sea level brought about by cycles of glacial freezing and melting.

This explanation was unsatisfactory for a number of reasons, but par-

ticularly from the standpoint that the duration of periods of isolation

would have been insufficient to have permitted evolution to proceed to

its present state. In this paper I will discuss recent findings pertaining to

pre-Pleistocene fluctuations in sea level that provide a more logical

explanation for the above. Related problems considered include (a)

composition, relationships and origins of the fish fauna; (b) factors con-

tributing to faunal distinctiveness of the area, including discussion of

the origin and biogeographic effects of Okefenokee Swamp; (c) origin of

the St. Johns River drainage and its fauna; (d) influence of the early

Pliocene “Cody Scarp” on the distributions of certain species; and (g)

species evolution in southern Georgia and northern Florida, including

the enigmatic absence of amblyopsid fishes or other cave fishes from the

area.

The acronym UF-FSU appearing in this paper refers to specimen

lots originally housed in the Florida State University ichthyological col-

lection, Tallahassee. This collection was subsequently combined in its

entirety with that in the Florida State Museum, University of Florida,

Gainesville.

Brimleyana No. 13:25-54, July 1987

25

26

Carter R. Gilbert

Some of the information contained herein appears, in condensed

form, in a chapter dealing with the zoogeography of the freshwater

fishes of southeastern United States, from the Savannah River to Lake

Pontchartrain (Swift et al. 1986). This in turn is included in a recently

published book on North American freshwater fish zoogeography

(Hocutt and Wiley, editors, 1986). The reader is referred to these refer-

ence sources for further information on the overall relationships of the

south Georgia-peninsular Florida fish fauna to those of other areas.

This study was supported in part by National Science Foundation

grant no. GB-1 1744.

COMPOSITION, RELATIONSHIPS AND ORIGINS

OF FISH FAUNA

The area south of the Altamaha River and east of the Apalachicola

River lies entirely below the Fall Line (Fig. 1), and may be termed the

south Georgia-peninsular Florida faunal complex.1 2 It is separated from

the neighboring Altamaha-Ogeechee-Savannah faunal complex at a

Jaccard coefficient of association level of about 0.42, and from the Apa-

lachicola River fauna at a coefficient level of 0.29 (Fig. 2). The south

Georgia-peninsular Florida complex may in turn be subdivided (at

about the 0.53 level) into northern and southern segments. The former

includes the St. Johns and Suwannee drainages, as well as other drain-

ages to the north, whereas the latter consists of the remaining rivers of

peninsular Florida. Although the St. Johns falls within the northern

segment of this faunal complex, this is somewhat misleading in terms of

the actual origin of the drainage and its fauna. The history of this

drainage is complex, and its fauna could just as logically be included

among those occurring in the drainages of southern peninsular Florida.

Although 126 native fish species are known to occur in fresh waters

of the area, only those species (72) that normally spend their entire lives

in fresh water were included in my analysis. The other 54 are species of

marine affinities, about half of which regularly enter fresh water, either

to spawn or to spend much of their lives, and the others of which occur

there on a more-or-less casual basis. This is by far the greatest number

of marine species entering fresh water for any area of comparable size in

North America, exclusive of Central America. The 72 species analyzed

1 The Fall Line is a zone of abrupt topographic change, characterized in many places by

waterfalls or rapids on the various streams, which marks the boundary between the

Coastal Plain and Piedmont areas and represents the maximal extent of past sea-level

incursion.

2 Jaccard coefficients of association between river drainages were calculated using species

presence and absence data (Sneath and Sokal 1973; Sepkoski and Rex 1974; Swift et al.

1986).

Freshwater Fish Zoogeography 27

Fig. 1. Major drainages of southern Atlantic and eastern Gulf slopes, includ-

ing peninsular Florida. Fall Line indicated by heavy dashed line extending diag-

onally across middle Alabama, Georgia and South Carolina. From Swift et al.

(1986).

constitute the smallest total for any of the faunal complexes (Fig. 2)

between the Savannah and Mississippi rivers (Swift et al. 1986), and this

is particularly significant when one considers the relatively large geo-

graphical area involved. Only 50 of the 72 species are primary-division

freshwater fishes, as defined by Myers (1938), and thus are essentially

intolerant of salt water. The remaining 22 species, which belong to

28

Carter R. Gilbert

secondary or peripheral (i.e., salt-tolerant) freshwater families, exhibit a

gradient in terms of their occurrence in fresh water. Some, such as

Labidesthes sicculus , Fundulus chrysotus, Fundulus cingulatus , Fundu-

lus escambiae and Fundulus lineolatus, seldom enter brackish waters.

Others, including Poecilia latipinna , Lucania parva , Rivulus marmora-

tus and Menidia beryllina, often are found there. Finally, species such

as Alosa alabamae, Alosa chrysochloris, Acipenser oxyrhynchus , Aci-

penser brevirostrum and Morone saxatilis are anadromous and regu-

larly enter fresh water to spawn.

The area south of the Suwannee and St. Johns river drainages con-

tains only about 50 of the 72 species analyzed, and represents still

further impoverishment. Despite this, the fishes of peninsular Florida

show a high percentage of endemism, as is true also of many plants and

other animals (Neill 1957). Species such as Micropterus salmoides,

Lepomis macrochirus, Notropis emiliae and probably Lepomis margi-

natus have distinctive subspecies restricted to peninsular Florida (Bailey

and Hubbs 1949; Felley 1980; Gilbert and Bailey 1972; Bruce H. Bauer,

pers. comm.), and for species such as Ictalurus natalis there is clearcut

differentiation at the racial level (Lodge 1974).

The south Georgia-peninsular Florida area is notable for the paucity

of species in three fish families that otherwise are dominant through-

out eastern North America: Cyprinidae (13 species), Percidae (5 species)

and Catostomidae (2 species). By comparison, there are 47, 34 and 13

species in each of these respective groups in the faunistically richer

Mobile Bay basin. The families Centrarchidae, Ictaluridae (primary-

division groups) and Cyprinodontidae (a secondary-division group),

along with the large number of marine forms, provide outstanding con-

tributions to the fauna in terms of numbers of species. A total of 17

centrarchid, 8 ictalurid and 1 1 cyprinodontid species occur in southern

Georgia and peninsular Florida, as compared with 16, 12 and 8, respec-

tively, for these groups in the Mobile Bay basin. The total of 17 repre-

sents exactly half the total number of described centrarchid species found

in eastern North America. The 8 ictalurids equal about one-fifth the

total number of species in eastern North America, and the 11 cyprino-

dontids represent about one-third of the freshwater members of this

family from the Mississippi basin eastward.

In summary, the south Georgia-peninsular Florida area may be

characterized by (a) a comparative paucity of species in the families

Cyprinidae, Percidae and Catostomidae; (b) an “average” number of

species of Ictaluridae; (c) a comparatively large number of species of

Centrarchidae and Cyprinodontidae; and (d) an unusually large number

of marine species that invade fresh waters on a regular or casual basis.

Freshwater Fish Zoogeography

29

1 .2 3 4 5 .6 7 .8 .9 VO

r t r i T f I I I I

SAVANNAH

OGEECHEE

ALTAMAHA

SATILLA

ST. MARYS

ST JOHNS

AUCILLA

SUWANNEE

OCHLOCKNEE

ST. MARKS

LAKE OKEECHOBEE

HILLSBOROUGH

WITHLACOOCHEE

LOWER EVERGLADES

PEACE

ALAFIA

MYAKKA

LITTLE MANATEE

MANATEE

WACCASASSA

APALACHICOLA

CHOCTAWHATCHEE

ESCAMBIA

BLACKWATER YELLOW

PEHDlDO

ESCATAWPA

ST. ANDREWS BAY

MOBILE BAY

PASCAGOULA

PEARL

LAKE PONTCHARTRAIN

J I I I L I 1 I 1 L

.1 2 3 .4 5 6 .7 8 9 1.0

Fig. 2. Phenetic clustering of 31 southeastern stream drainages, from the

Savannah River to Lake Pontchartrain (including peninsular Florida), based on

presence or absence of species (Jaccard coefficients of association). From Swift

et al. (1986).

30

Carter R. Gilbert

The unusual character of the south Georgia-peninsular Florida fish

fauna results from (a) the relatively unvarying lowland habitat of much

of the area, particularly the lower two-thirds of peninsular Florida; (b)

periodic inundation of much of the area during the late Cenozoic era;

and (c) the comparative isolation of this area from the main centers of

faunal diversity (thus presumed centers of evolution) in southeastern

United States, particularly the Tennessee uplands and Mobile Bay

basin.

Speculation on the sources of origin for various faunal elements is

especially complicated here in comparison to other areas of the

Southeast. The geographically intermediate position of the state in rela-

tion to the Gulf and Atlantic slopes is largely responsible, inasmuch as a

number of species are widespread on both slopes. In the cases of Esox

americanus and Aphredoderus sayanus, northern and peninsular Flor-

ida populations have been shown mostly to be subspecific intergrades

(Crossman 1966, 1980a; Lee 1980), and thus a dual origin is clearly

involved. Given this and related problems, only 50 of the 72 species

under consideration could be evaluated with reasonable certainty, with

the following results: (a) Gulf slope — 17, (b) Atlantic slope — 16, (c)

combined Gulf and Atlantic slopes — 2, (d) peninsular Florida — 9, (e)

southern Georgia and northern Florida — 4, and (f) marine — 2.

It is often assumed that peninsular Florida, because of its relatively

uniform habitat and obscure drainage divides, has an essentially homo-

geneously distributed fish fauna, but this is far from the truth. Analysis

of the fauna reveals distinctive distribution patterns that, in many cases,

reflect pathways of dispersal. For example, the distribution of Percina

nigrofasciata in the southern half of the peninsula is limited to the Kis-

simmee River, which flows in a southerly direction down the center of

the state and is the principal tributary emptying into Lake Okeechobee

(Burgess 1980b). Esox niger and Lepisosteus osseus show similar dis-

tributions, but differ in that both species are also in the Hillsborough

River (on the west coast), which is the largest tributary to Tampa Bay

(Crossman 1980b, Wiley 1980a). Notropis emiliae peninsularis occurs in

the Kissimmee River, but is absent from the Hillsborough; it also

occurs, on the lower west coast, in the Peace River system, which emp-

ties into Charlotte Harbor (Gilbert and Bailey 1972, Gilbert 1980b). All

of the above species are absent from the four geographically interme-

diate, and ecologically similar, river systems between the Peace and

Hillsborough rivers on the middle west coast (Myakka, Manatee, Little

Manatee and Alafia rivers).

Enneacanthus obesus and Elassoma okefenokee range southward

only as far as the middle of the peninsula, with neither entering the

Kissimmee system (Lee and Gilbert 1980b, Bohlke and Rohde 1980b).

31

Freshwater Fish Zoogeography

In both cases, however, their ecological preferences do not appear to

differ significantly from those of their closest respective relatives,

Enneacanthus gloriosus and Elassoma evergladei, which are widely and

uniformly distributed throughout the peninsula (Lee and Gilbert 1980a,

Bohlke and Rohde 1980a). It seems clear that drainage divides in penin-

sular Florida either are less easily breached than would first appear, or

unknown ecological factors prevent species such as Enneacanthus obe-

sus and Elassoma okefenokee from becoming established in the lower

part of peninsular Florida.

FACTORS CONTRIBUTING TO FAUNAL DISTINCTIVENESS

There are several explanations for the pronounced faunal break

between the Altamaha and Satilla river drainages, which show a Jac-

card coefficient of association level of only about 0.42 (Fig. 2). The

most obvious explanation is that much of the Altamaha drainage lies

above the Fall Line, whereas the Satilla and adjacent drainages to the

south are situated entirely below the Fall Line, on the Coastal Plain

(Fig. 1). The Altamaha thus has a much greater diversity of habitat

and a consequently more diverse fish fauna than the Satilla (62 vs. 44

species). This is not the entire answer, however, since there are four

species ( Notropis cummingsae , Ictalurus brunneus , Etheostoma olm-

stedi and Percina nigrofasciata ) that are absent from both the Satilla

and adjacent St. Marys rivers, but reappear in the St. Johns drainage

farther south (Gilbert and Burgess 1980a,c; Lee and McAllister 1980;

Burgess 1980b). This phenomenon appears to be related to the presence

of Okefenokee Swamp, which is situated about 35 m above present sea

level and forms the headwaters for the Suwannee, St. Marys and Satilla

rivers. The swamp originated as a marine embayment (Cooke 1925),

presumably during the Pliocene when sea levels were higher. It was sub-

sequently isolated from the ocean by Trail Ridge, which has a maxi-

mum elevation of nearly 50 m and which extends in a northeasterly

direction toward Jesup, Georgia, from a point west of the north-flowing

loop of the St. Marys River (between Georgia and Florida) (Fig. 3, in

part). The time of formation of Trail Ridge has been the subject of

debate, with estimates ranging from Miocene (Alt and Brooks 1965) to

Pleistocene (Cooke 1939). Opdyke et al. (1984), however, have firmly

placed its origin in the Pleistocene, a conclusion based partly on the

presence, at elevations of 42-49 m, of a marine fossil invertebrate fauna

consisting entirely of Recent species of gastropods, pelecypods, echi-

noids, bryozoans, barnacles and others. Later in the Pleistocene, epeiro-

genic uplift of the Florida peninsula raised Trail Ridge to its present

elevation, after which the Okefenokee Swamp assumed its present char-

acter. Since this provided an unsuitable habitat for most lotic fishes,

32

Carter R. Gilbert

one may assume that (a) it limited the southern distributions of many

species, and (b) the four species listed above by-passed the Satilla and

St. Marys via the upper Suwannee River, from where they subsequently

entered the lower (i.e., northern) St. Johns drainage by way of the Santa

Fe River system (Burgess and Franz 1978). Although some lotic spe-

cies, such as Noturus leptacanthus and Minytrema melanops, later

gained access to the St. Marys and/or Satilla, this does not negate the

present hypothesis.

The break between the Apalachicola River and those drainages

immediately to the east (Altamaha, Suwannee and Ochlockonee) is the

most significant faunal break in the area of study, showing a coefficient

level of only 0.29. This is particularly surprising in the case of the Apa-

lachicola and Ochlockonee, inasmuch as the Coastal Plain habitat, with

its poorly defined drainage divides, appears to offer easy passage in

both directions to a large number of lowland species. Furthermore, the

Ochlockonee River, in Florida, appears to have captured its western-

most tributary, Telogia Creek, from the Apalachicola River. Evidence

for this may be seen from the geographic appearance of Telogia Creek,

the upper section of which flows in a southwesterly direction toward the

Apalachicola, then doubles completely back on itself to take a northeast-

erly course to the Ochlockonee (Swift et al. 1977: Fig. 1-2). Several

species of western affinities that typically occupy flowing streams of

varying size (e.g., Ichthyomyzon gagei, Notropis t exanus, Notropis

venustus, Semotilus thoreauianus and Etheostoma swaini ) presumably

entered the Ochlockonee drainage via this route (Rohde and Lanteigne-

Courchene 1980, Swift 1980b, Gilbert and Burgess 1980b, Lee and Pla-

tania 1980, Starnes 1980). Overall, however, the fish faunas of the Apa-

lachicola and Ochlockonee are quite different, a situation that suggests

long-term physiographic independence of these drainages. Price and

Whetstone (1977) indicate that the Apalachicola River (which upstream

is called the Chattahoochee River) has maintained its basic integrity

throughout the latter half of the Cenozoic, and has remained deeply

entrenched in its southerly course since it was rapidly eroding down-

ward in a positive area of terrestrial rise. The long-term phys-

iographic integrity of the Apalachicola would thus tend to preserve its

biological integrity as well.

GEOGRAPHICAL ISOLATION AND ENDEMISM

IN PENINSULAR FLORIDA

Past discussion of the zoogeography of the south Georgia-peninsular

Florida area has been complicated by the fact that interpretations of

major sea-level cycles have frequently been modified with regard to both

Freshwater Fish Zoogeography

33

1 Suwannee River

2 Santa Fe River

3 Orange Creek

4 Little Orange Creek

5 Levy's Prairie

6 Oklawaha River

7 Deep Creek

8 Rice Creek

9 Clark Creek

10 Black Creek

11 St. Johns River

^ Northern Highlands

il Trail Ridge

@ Baywood Promontory

H Duval Upland

Fig. 3. Northeastern Florida, showing streams and topographic features asso-

ciated with Cody Scarp and Trail Ridge. Although not specifically labelled here,

Cody Scarp forms boundary of Northern Highlands, with Trail Ridge, Bay-

wood Promontory and Duval Upland being situated beyond scarp limits. From

Burgess and Franz (1978).

timing and scale. There is no question that, in the past, peninsular Flor-

ida was isolated by the sea from the rest of southeastern United States,

and that the major break occurred in the northern part of the peninsula

in the so-called “Suwannee Straits” area (Fig. 4). The principal evidence

for former sea-level stands consists of geomorphic indications of actual

shorelines (MacNeil 1949). There is also direct fossil evidence, since

marine organisms are represented well inland from where the sea lies

today (Alt and Brooks 1965, Robertson 1976). Furthermore, it is

obvious that this isolation must have existed for a rather long time, as

indicated by the level of genetic differentiation of many peninsular ele-

ments from their close relatives to the north. Fishes furnish several good

examples of this, as do other animal groups and various plants.

34

Carter R. Gilbert

Recent papers by Vail et al. (1977) and Vail and Hardenbol (1979),

which discuss sea-level changes during the Cenozoic, have done much to

clarify this situation. However, this picture has subsequently been modi-

fied somewhat by the findings of Opdyke et al. (1984), who presented

evidence to show that peninsular Florida and closely adjacent areas

have undergone epeirogenic uplift during the late Cenozoic. One impor-

tant consequence of this is that present elevations of the various marine

terraces may be exaggerated in terms of reflecting changes in sea level.

For example, the Cody Scarp (or Wicomico Shoreline) is at an eleva-

tion approximately 30 m above present mean sea level; however, the

actual drop in sea level may have been somewhat less than 30 m, with

the difference being accounted for by epeirogenic uplift. Unfortunately,

the exact relative contributions of the two events cannot be determined

with certainty. Despite this new complication, the picture presented

below regarding isolation and evolution of faunal elements in peninsular

Florida correlates well with the sequences of rise and fall in sea level

presented by Vail et al. (1977) and Vail and Hardenbol (1979). It corre-

lates so well, in fact, that I must conclude that epeirogenic uplift,

although an important consideration, does not significantly change the

basic scenario.

Cooke (1945) presented the first thorough summary of Florida sea-

level stands. He envisioned the periodic separation and rejoining of

peninsular Florida and the mainland to be essentially a Pleistocene

event. Alt and Brooks (1965) and Alt (1968) regarded the isolation of

these peninsular islands as having occurred earlier, with the last com-

plete separation being no later than the first (Aftonian) interglacial, if

then. They pointed out that pre-Pleistocene sea levels were higher than

subsequently, this presumably related to the fact that large amounts of

sea water were not then tied up in glacial ice. If so, this would suggest a

continuous pre-Pleistocene isolation of the peninsular islands from the

mainland. This assumption, however, is at odds with the fact that var-

ious freshwater fishes that are intolerant of sea water (Myers’ [1938]

primary-division groups) almost certainly had reached peninsular Flor-

ida earlier, where they remained isolated sufficiently long to permit well-

defined racial or subspecific differentiation to occur.

Peninsular Florida is thought to have had its origin during the

middle Oligocene epoch, probably around 30 million years before pres-

ent (B.P.). This event is correlated with an unusually severe drop in sea

level, perhaps as much as 250 m (Vail et al. 1977, Vail and Hardenbol

1979), which in turn appears to be correlated with formation of the

Antarctic ice cap (Miller et al. 1985, Prothero 1985, Savin and Douglas

1985). Major mid-Oligocene cooling and circulation events are sug-

gested by the benthic foraminiferal <5l80and planktonic foraminiferal

Freshwater Fish Zoogeography

Fig. 4. Early Pliocene high-sea stand in Florida, Georgia and South Carolina

(ca. 30 m above present mean sea level). Heavy solid line bordering continental

landmass indicates Cody Scarp. Open area separating mainland from offshore

islands indicates position of Suwannee Straits. From Swift et al. (1986).

<513C records, respectively (Cavelier et al. 1981, Prothero 1985). These

events apparently resulted from movement of the Antarctic plate and

the concomitant increased isolation of the Antarctic continent, which in

turn would have caused increased development of the circum-Antarctic

current and resulting refrigeration of the continent (Kennett et al. 1972,

Prothero 1985). Lowered sea levels also tended to accelerate the ero-

sion1 processes of the overlying clastic deposits and of the underlying

limestone substrate in Florida by increasing the speed of percolation of

surface water through the strata. This is thought to have reduced the

density of the carbonate rocks exposed near the surface, thus causing

36

Carter R. Gilbert

the peninsula to rise somewhat by epeirogenic adjustment. This same

mechanism, recurring throughout the rest of the Cenozoic, may partly

account for the pronounced elevational differences of successive shore-

lines (Opdyke et al. 1984). As indicated earlier, the Pliocene “Wicomico

Shoreline” (Cody Scarp) may have occupied a position closer to present

mean sea level than the current (30 m) elevation of this land feature

would seem to suggest.

Peninsular Florida is believed to have maintained a more-or-less

continuous connection to the adjacent mainland during the next 12 to

14 million years (i.e., into the early mid-Miocene), at which time a

gradual rise in sea level resulted in partial, or possibly complete, isola-

tion of the peninsula (Vail et al. 1977, Vail and Hardenbol 1979) (Fig.

5). In the late Miocene, ca. 10 million B.P., another sharp drop in sea

level occurred (ca. 200 m below present mean sea level), which again

caused peninsular Florida to be joined to the mainland. This connection

was broken in the latest Miocene, ca. 5 million B.P., when another pro-

nounced rise in sea level isolated the peninsula. This was the same

marked eustatic cycle that produced the desiccation and subsequent

refilling of the Mediterranean Sea by the Atlantic Ocean, an event Hsu

(1972, 1978) believed to have occurred bewteen 5 and 5.5 million B.P. High

sea levels persisted throughout the Pliocene, 5 to 2 million B.P., but

dropped as a result of Pleistocene glaciation, when the Florida penin-

sula was restored.

Endemic fishes of the Florida peninsula fall into two basic groups:

(a) well-defined insular populations (either subspecies or races) of wide-

ranging eastern North American species; and (b) endemic peninsular

species, mostly in the family Cyprinodontidae, that have no obviously

close relatives elsewhere in eastern United States. The first group is

represented by distinct subspecies of Notropis emiliae , Micropterus sal-

moides, Lepomis macrochirus and possibly Lepomis marginatus, and by

a racially distinct population of Ictalurus natalis. Subsequent study will

probably reveal other examples. The second group is represented by

Fundulus seminolis and Jordanella floridae (Gilbert 1980f, Gilbert and

Burgess 1980e), and probably also by Fundulus cingulatus , Leptoluca-

nia ommata, Lucania goodei and Heterandria formosa (Gilbert and

Burgess 1980d,f,g; Martin 1980). The last four species are found

throughout all or much of peninsular Florida ( L . ommata ranges about

halfway down the peninsula), but also range narrowly outside this area.

Fundulus cingulatus may be related to certain other freshwater species

in North America (Chen 1971), the genus Lucania apparently is most

closely related to certain Cuban and Jamaican genera (Hubbs and

Miller 1965), and the relationships of F. seminolis are uncertain. The

Freshwater Fish Zoogeography

37

1 i | i — i — i — i — ] — i — i — i — i — | — , — , — ( — i — | — i — i — , — i — | — | — i — i — | — | — i — i — ( — , — • -200

60 50 40 30 20 10 0

Fig. 5. Tertiary eustatic changes in sea level. Meters above or below sea level

are tentative. From Vail and Hardenbol (1979).

genus Jordanella has been considered to be most closely related to the

Yucatan genus Garmanella, but recent unpublished studies suggest that

this is not the case (J. M. Humphries, pers. comm.). The poeciliid genus

Heterandria, which contains nine species, occurs disjunctly in southeast-

ern United States (one species, H. formosa) (Martin 1980), and in both

lowland and upland areas from southeastern Mexico south to eastern

Nicaragua (Rosen 1979: Fig. 22). Movement of Heterandria stocks

between Mexico and peninsular Florida could have occurred by direct

migration through the ocean, perhaps aided by much lower sea levels

that existed throughout parts of the Oligocene and Miocene, or by

movement and subsequent elimination of coastal populations around

the northern rim of the Gulf of Mexico. Of these, the latter possibility

seems more likely, as indicated by (a) the absence of Heterandria from

Cuba, and (b) the continuous distribution of another poeciliid, Poecilia

latipinna (also absent from Cuba), around the northern rim of the Gulf

of Mexico from Yucatan to Florida and beyond (Burgess 1980a).

Based on a lower level of taxonomic differentiation (i.e., subspecific

or racial), it appears likely that the group composed entirely of primary-

division freshwater fishes evolved during the second (i.e., Pliocene) iso-

lation of peninsular Florida. However, the species-level differentiation

of those in the salt-tolerant, secondary-division freshwater families

Cyprinodontidae and Poeciliidae could be interpreted as more likely

indicating an evolutionary history dating back to the mid-Miocene iso-

lation of the peninsula. If the above scenarios are correct, we may ask

METERS

38

Carter R. Gilbert

why evolution of these primary and secondary freshwater groups was

temporally separated? One possible explanation is that during the mid-

Miocene permanent freshwater habitats were more limited in insular

Florida, which could have prevented the establishment of such primary-

division freshwater families as the Esocidae, Cyprinidae, Catostomidae,

Ictaluridae and Centrarchidae. Circumstantial evidence suggests that the

family Percidae may not have immigrated into North America by this

time (Gilbert 1976).

Isolation and subsequent differentiation of fishes in insular Florida

was not necessarily confined to freshwater forms, although certainly the

freshwater groups provide more numerous examples. Evolution of the

two species of the genus Chasmodes (of the marine family Blenniidae)

appears also to have occurred as a direct result of the insularization of

peninsular Florida. Chasmodes saburrae, which apparently evolved in

that area following isolation of part of the original Chasmodes stock,

ranges from the upper two-thirds of the east Florida coast around the

peninsula westward to just beyond Mobile Bay (Williams 1983: Fig. 5).

It effectively bisects the range of the closely related C. bosquianus ,

which displays an allopatric distributional relationship to C. saburrae

on the Atlantic coast, but a syntopic relationship to that species on the

middle Gulf coast, from the area of Pensacola west to just beyond

Mobile Bay. It is obvious that C. bosquianus once had a continuous

distribution, but has subsequently become divided as a result of intru-

sion of C. saburrae into the center of its range following geographic

reamalgamation of insular and mainland Florida. Although Williams

(1983) believed this to be a .pre-Pleistocene event, he did not indicate

exactly how much earlier this might have taken place. Based on the

discussion presented earlier, it seems likely that differentiation of the

two species of Chasmodes occurred during the Pliocene, an hypothesis

that is concordant with the essentially similar level of morphological

differentiation seen in the primary-division freshwater fish species cited

earlier.

Peninsular Florida contains a number of faunal and floral elements

of West Indian origin, with those groups displaying the highest levels of

mobility or vagility being best represented (e.g., birds, flying insects, and

many kinds of plants). A number of fish species whose ranges lie princi-

pally in the West Indies and Caribbean areas enter United States fresh

waters only in peninsular Florida. These include Rivulus marmoratus

(family Cyprinodontidae), Gambusia rhizophorae (family Poeciliidae),

Awaous tajasica and Gobionellus pseudofasciatus (family Gobiidae),

and three species of the genus Centropomus (family Centropomidae)

(Gilbert and Burgess 1980h; Getter 1980; Lindquist 1980; Hastings 1980;

Freshwater Fish Zoogeography

39

Burgess 1980c,d,e). Other freshwater fishes of similar origin range only

a short distance beyond Florida. All the above species are completely

salt tolerant, and thus are subject to continual recruitment from farther

south. Furthermore, in none of the above cases has the Florida popula-

tion been shown to differ taxonomically from those elsewhere.

ORIGIN OF ST. JOHNS RIVER DRAINAGE

Previous reference has been made to the complex origin of the St.

Johns River drainage and to the fact that, although it falls into the

faunal cluster with those drainages immediately to the north (Fig. 2), it

could equally well be included with those occurring in the southern part

of peninsular Florida. Partial evidence for this is based on the taxo-

nomic character of Micropterus salmoides, Lepomis macrochirus ,

Lepomis marginatus, Notropis emiliae and Ictalurus natalis, each of

which has undergone genetic differentiation in peninsular Florida, with

the peninsular form in each case being found in the St. Johns drainage.

The St. Johns River historically was an ocean estuary with a north-

south orientation determined by a series of barrier islands that parallel

the coast and separate the river from the Atlantic Ocean. These islands,

which collectively have been termed the “Atlantic Coastal Ridge,”

appear to have been formed during Pamlico time (Yarmouth intergla-

cial), when sea level was about 8 to 10 m highei than at present (White

1970: 86). Most of the drainage was inundated during the Pliocene, the

only areas not so covered being several tributaries flowing off higher

ridges in the Palatka area, to the west, and a relatively small section in

the upper (i.e., southern) part of the drainage (Burgess and Franz 1978)

(Fig. 4). The latter included part of the upper Oklawaha River system,

as well as that area included in the present-day Ocala National Forest

(the so-called “Ocala scrub”), which is situated between the Oklawaha

and St. Johns rivers. During the Pleistocene interglacial periods, partial

inundations of the peninsula occurred, with each successive inundation

covering less of the peninsula than before (Alt and Brooks 1965).

The only endemic fish in the St. Johns drainage is the taxonomi-

cally weakly defined Cyprinodon variegatus hubbsi, which occupies six

lakes in the upper Oklawaha River system (Weir, Harris, Eustis, Yale,

Griffin and Dora) (Johnson 1974, 1980). The precursor of this form

(presumably very similar to C. v. variegatus ) is believed to have reached

this area during either the Pliocene or early Pleistocene, when sea levels

were highest, and evolved to its present taxonomic level during the

Pleistocene. Cyprinodon v. variegatus , although tolerant of fresh water,

is basically an inhabitant of brackish water, and the progressively lower

sea stands during successive Pleistocene interglacial periods would have

40

Carter R. Gilbert

served to increase the degree of isolation of the upper Oklawaha popu-

lation. Johnson (1974) has shown that the present inland distribution of

C. variegatus variegatus is sharply limited by the Pamlico terrace, which

is the lowest (8 to 10 m) and most recent of the Pleistocene marine

terraces (Fig. 6). Why this fish failed to become established farther

inland during earlier marine high stands is unclear. An interesting point

is that the factor (lowered sea levels) leading to the evolution of C.

variegatus hubbsi in peninsular Florida is essentially the opposite of

that resulting in evolution of the other fish taxa discussed earlier.

The lower (i.e., northern) part of the St. Johns drainage had an

independent origin from the upper area. Several tributaries of the lower

St. Johns and Oklawaha rivers (the Oklawaha is the principal system

within the St. Johns drainage), including Orange, Deep, Rice, Etonia

and Black creeks (Fig. 3), were not inundated during Pleistocene inter-

glacial or pre-Pleistocene rises in sea level (Fig. 4). This area, which has

been called the ‘’Wicomico Peninsula” (Pirkle et al. 1974), is basically an

eastern extension of the Cody Scarp. Several major topographic fea-

tures were formed on the margins of this peninsula, including Trail

Ridge, which probably originated as a beach ridge during the early

Pleistocene, and the Duval Upland and Baywood Promontory, which

developed as regressional plains as sea levels dropped (Fig. 3). The

streams in this area served as refugia for various fishes and inverte-

brates, and their isolation was of sufficient duration to permit the evolu-

tion of several endemic species of invertebrates (Burgess and Franz

1978: Table 1). Although no endemic fishes are present here, four

( Notropis welaka , Notropis cummingsae, Ictalurus brunneus and

Etheostoma olmstedi ) have isolated relict populations centered in the

area. None of these species is found in the adjacent Suwannee or St.

Marys drainages (Gilbert 1980e; Gilbert and Burgess 1980a,c; Lee and

McAllister 1980), and only N. cummingsae occurs in any immediately

adjacent drainages. These four fishes are believed to have entered the

lower St. Johns drainage via the upper Santa Fe River (a major tribu-

tary of the Suwannee), which also was located on the Wicomico Penin-

sula (Burgess and Franz 1978) (Fig. 3). Burgess and Franz (1978) specu-

lated that current absence of these species from the Santa Fe system,

and elsewhere in the Suwannee drainage, could have been caused by

two related factors: reduction, during glacial maxima, in the overall

amount of surface water as a result of solution draining; and increased

interspecific competition. The ecological conditions existing today in the

area appear unsuitable for N. welaka , but seem favorable for the other

three species. Several other fishes ( Elassoma zonatum, Acantharchus

pomotis and Umbra pygmaea ) have similarly restricted distributions in

Freshwater Fish Zoogeography

41

Fig. 6. Distribution of Cyprinodon variegatus in Florida, in relation to Pleis-

tocene Pamlico shoreline (8-10 m above present mean sea level). Dots denote

collection localities for C. v. variegatus ; stars denote localities for C. v. hubbsi.

Modified from Johnson (1974).

the St. Johns drainage, but in contrast to the situation just described

they still occur in parts of the Suwannee drainage (Bohlke and Rohde

1980c, Cashner 1980, Gilbert 1980a).

INFLUENCE OF CODY SCARP ON FISH DISTRIBUTIONS

All but the shortest rivers of northern Florida drain two physio-

graphic areas, the Northern Highlands and the Coastal Lowlands (Puri and

Vernon 1964, White 1970). The section of the Northern Highlands east

of the Apalachicola River valley is called the Tallahassee Hills (Puri and

42

Carter R. Gilbert

Vernon 1964: Fig. 5). This upland area is limited on the south and east

by an out-facing scarp, a marine terrace about 30 m above present sea

level that is the most persistent topographic break in the state. In Flor-

ida this is called the Cody Scarp, a name that is synonymous with Alt’s

(1968) “Surry Scarp”; the same feature was referred to as the “Wico-

mico Shoreline” by Cooke (1945) and by Hoyt and Hails (1967). In

Florida the Cody Scarp extends almost straight eastward to a point in

the upper Suwannee River valley, just east of the Alapaha River,

beyond which it turns southward at about a 75-degree angle to a point

not far southwest of Gainesville, curves eastward to run just north of the

Oklawaha River, and then turns abruptly northward west of the St.

Johns River (Burgess and Franz 1978) (Fig. 3). The area so defined, the

“Wicomico Peninsula,” borders Trail Ridge, which is a beach ridge

formed along its eastern margin. The “Wicomico Peninsula” also borders

the Duval Upland and Baywood Promontory, which developed as

regressional plains just beyond the eastern margin of the scarp as sea

levels dropped (Fig. 3).

The continuity of the Cody Scarp is unbroken except by the valleys

of major streams, but its definition is variable. In many places it can be

delineated with unequivocal sharpness, whereas elsewhere it appears as

a gradual reduction of average elevation and a general flattening of ter-

rain as lower elevations are reached. Swift et al. (1977) indicated that in

the Ochlockonee River valley elevations above sea level range from 60

to 90 m in the highlands immediately surrounding the drainage, and are

30 m or less in the lowlands. These two areas are separated by the scarp.

Northward the highlands remain hilly but more rolling, and the tribu-

taries have more mature, meandering channels. To the south the land is

flat and streams are sluggish and swampy, but where streams flow over

the scarp they have steeper gradients.

The Cody Scarp forms an important zoogeographic boundary.

Swift et al. (1977: 61) cited 12 fish species whose distributions in the

Ochlockonee drainage seem to be affected by it, including Enneacanthus

obesus, Heterandria formosa , Fundulus chrysotus , Fundulus cingulatus

and Leptolucania ommata (all restricted to below the scarp); Notropis

petersoni (found only immediately above); and Ichthyomyzon gagei ,

Semotilus thoreauianus and Etheostoma swaini (all essentially con-

fined to the scarp area) (Lee and Gilbert 1980b; Martin 1980; Shute

1980; Gilbert and Burgess 1980d,f; Swift 1980a; Rohde and Lanteigne-

Courchene 1980; Lee and Platania 1980; Starnes 1980). The influence of

this topographic feature is best illustrated, however, by the distribu-

tional and systematic relationships of two closely related but easily dis-

tinguishable cyprinodontid fishes, Fundulus escambiae and Fundulus

lineolatus (Rivas 1966; Wiley 1977, 1980b, c) (Fig. 7). Fundulus escam-

biae is an eastern Gulf slope species that ranges from the Perdido River

Freshwater Fish Zoogeography

43

Fig. 7. Distributions of Fundulus escambiae (dots) and Fundulus lineolatus

(stars) in Florida, Alabama, Georgia and southeastern South Carolina, in rela-

tion to Cody Scarp (based on records in Florida State Museum and Tulane

University fish collections). Upper (inland) solid line denotes Fall Line; lower

(more peripheral) solid line indicates Cody Scarp. Distribution of F. escambiae

shown in entirety; distribution of F. lineolatus extends north to southeastern

Virginia.

44

Carter R. Gilbert

east to the Suwannee, whereas F. lineolatus is an Atlantic slope form

occurring from extreme southeastern Virginia south throughout most of

peninsular Florida and west to the Apalachicola drainage. Outside the

area of distributional overlap, these species regularly occur both above

and below the Cody Scarp, but within the area of overlap (i.e., the

Suwannee to the Apalachicola) their distributions usually appear to be

strongly influenced by this physiographic break, with most populations

of F. lineolatus occurring above and most populations of F. escambiae

below (Fig. 7). This correlation holds up especially well in the Ochlock-

onee and Suwannee drainages, except in Telogia Creek (the principal

western tributary of the Ochlockonee), from which F. lineolatus appears

to be absent. In the Ochlockonee River proper the two forms come in

close proximity, but have been collected together only twice (UF-FSU

323 and 6002), from a locality just below the Lake Talquin (= Jackson

Bluff) dam, which is situated along the scarp on the main river. In both

cases gene interchange appears to have occurred (pers. observ.), despite

previous information to the contrary (Rivas 1966, Swift et al. 1977). In

the Suwannee drainage the two are more widely separated, F. escambiae

occurring only in the lower river, upstream to the Santa Fe River, and

F. lineolatus in the upper reaches. Neither species has yet been encoun-

tered in the intervening 120-km stretch of the Suwannee River proper,

above the mouth of the Santa Fe. In the Santa Fe system, F. escambiae

ranges upstream to the point where the river emerges from under-

ground, whereas in the sections above the Santa Fe Sink only F. lineo-

latus occurs. The area where the Santa Fe descends underground is pre-

cisely on the edge of the scarp.

In the Suwannee drainage the distributions of several other species

also appear to be limited by this physiographic feature. This is best

demonstrated by Fundulus cingulatus , whose distribution in the Santa

Fe and upper Suwannee parallels almost exactly that of F. lineolatus

(Gilbert and Burgess 1980d). Other species limited to lower parts of the

Suwannee drainage include Fundulus seminolis (also absent from the

upper Santa Fe system), Micropterus notius , Fundulus chrysotus and

Notropis petersoni (Gilbert 1980f,g; Shute 1980; Swift 1980a). Species

confined to the upper sections of the Suwannee include Notropis leedsi,

Notropis texanus and Notropis venustus (Gilbert 1980d, Swift 1980b,

Gilbert and Burgess 1980b). Unlike the situation involving F. escambiae

and F. lineolatus , there is, in many cases, considerable overlap in ranges

of these “upstream” and “downstream” species.

The distributional break between F. escambiae and F. lineolatus

may be equally sharp in the St. Marks drainage, but as yet collections

from upsteam have failed to yield individuals of either species. There are

Freshwater Fish Zoogeography

45

a number of downstream records of F. escambiae, however, one of

which (UF-FSU 3042) is in close proximity to the scarp. In the Apala-

chicola River drainage, only F. escambiae is found in downstream

areas, but it also ranges up into the Flint River system as far north as

Albany, Georgia, where it is abruptly replaced by F. lineolatus .3 Exten-

sion of the range of F. escambiae into this area, well above the limits of

the Cody Scarp, may be attributable to the combination of natural ero-

sional and depositional processes often seen in the lower sections of

large rivers, which cause the lowland type of habitat to be extended well

upstream.

Elsewhere the influence of the Cody Scarp becomes less obvious.

Apparently only F. lineolatus occurs in the series of endorheic lakes

located, near Tallahassee, mostly below the scarp in an area midway

between the Ochlockonee and St. Marks drainages. In the Aucilla

drainage a dichotomous distribution pattern seems to emerge. There F.

escambiae is limited to the more westerly Wacissa system, situated

entirely below the scarp, with F. lineolatus the only species in the

Aucilla system proper (where collections exist from both upstream and

downstream areas).4 Those drainages situated to the east (Econfina,

Fenholloway, Steinhatchee), between the Aucilla and Suwannee, lie

entirely in the lowlands. They apparently are inhabited exclusively by F.

lineolatus , even though the presence of F. escambiae in the lower

Suwannee River would logically lead one to expect that species to

exhibit a continuous distribution along the northeastern Gulf coast of

Florida. Considering the coastal distribution pattern of F. escambiae ,

together with the presence of (and thus probably in competition with) F.

lineolatus upstream, it seems unlikely that F. escambiae would have had

the opportunity to enter the Suwannee via headwater stream capture, as

is believed to have been the case with a number of other species (see

subsequent discussion). Possibly those small drainages lying between the

Aucilla and Suwannee were endorheic at certain critical times during

the past, which thus would have made them unavailable to colonization

by F. escambiae.

Despite certain inconsistencies, it seems clear that the overall dis-

tributions of Fundulus escambiae and F. lineolatus , as well as those of

3 A single individual of Fundulus lineolatus (UF-FSU 4521), which is presumed to be a

bait-fish release, has been collected in the Apalachicola River proper below Jim Wood-

ruff (= Lake Seminole) dam.

4 A series of eight specimens from the headwaters of the Wacissa River (UF-FSU 939)

contains seven Fundulus escambiae and one F. lineolatus (all adults). This is thought to

have resulted from mixing of collections; but if not, it represents the only record of F.

lineolatus from the Wacissa system and one of only three cases of known sympatry

between the two species.

46

Carter R. Gilbert

certain other species^ are strongly influenced by the Cody Scarp, which

in turn delineates an ecological break between the Northern Highlands

and Coastal Lowlands. These ecological differences alone do not appear

sufficiently great to limit the distributions of either F. escambiae or F.

lineolatus in those drainages in which only one of them is present. In

areas containing both species, however, ecological competition presum-

ably is involved and the upstream-downstream pattern is manifested, as

for example in the Ochlockonee and Apalachicola river drainages. The

same may not hold true in the Suwannee drainage, however, since the

ranges of the two fishes are widely separated either by an impenetrable

physical barrier on the Santa Fe River (i.e., the Santa Fe Sink) or by an

extensive (120 km) stretch of river that for some reason appears not to

be inhabited by either species.

I theorize that Fundulus lineolatus was present in the Tallahassee

Hills and Wicomico Peninsula during the “Wicomico high stand” (i.e.,

in the area above the present-day Cody Scarp). At the same time, F.

escambiae is presumed to have occupied the Western Highlands to the

west (Puri and Vernon 1964: Fig. 5; Burgess and Franz 1978: Fig. 1). As

the sea retreated, F. escambiae presumably moved eastward to occupy

many of the Coastal Lowland streams, but in those streams colonized

was prevented from moving farther upstream because of ecological com-

petition with the previously established F. lineolatus. The latter species,

as well as many others, was present in those tributaries draining the

eastern side of the Wicomico Peninsula, and thus had access to the

remainder of the St. Johns River when it subsequently assumed its pres-

ent form. It is also possible that this species was present, during the

Pliocene, on the insular areas of peninsular Florida.

SPECIES EVOLUTION IN NORTHERN FLORIDA

AND SOUTHERN GEORGIA

Northern Florida and southern Georgia are characterized by two

endemic fishes, Micropterus notius and Ictalurus serracanthus. The first

is restricted to the Suwannee and Ochlockonee rivers (Gilbert 1980g),

whereas the latter occurs in these two drainages, as well as in the Apala-

chicola and St. Marks rivers and in Econfina Creek to the west (Yerger

1980). (Econfina Creek should not be confused with Econfina River,

which is to the east, just beyond the Aucilla River). The present range of

M. notius is mostly confined to areas below the Cody Scarp, whereas /.

serracanthus is slightly more ubiquitous in its distribution. Although no

other fishes share these same distribution patterns, two other species of

wider occurrence ( Notropis harperi and Etheostoma edwini ) have dis-

tributions that are more or less centered in the same area (Gilbert 1980c,

Stauffer and Hocutt 1980), thus suggesting a similar distributional

history.

47

Freshwater Fish Zoogeography

Both Micropterus notius and Ictalurus serracanthus are well defined

species, with the former regarded as the most generalized form of black

bass (Bailey and Hubbs 1949). Both have achieved a higher level of

taxonomic differentiation from their closest congeners than have the

endemic peninsular populations of Micropterus salmoides and at least

four other species. Inasmuch as peninsular endemism in the last five

species is believed to have been achieved during the Pliocene, it may be

assumed that M. notius and /. serracanthus evolved earlier, and that

species differentiation was well under way at least by the end of the

Miocene.

As previously indicated, average Pliocene sea levels were higher than

those in the latter half of the Miocene. At its maximum incursion, about

5 million years ago, the sea stood at or near the present Cody Scarp, as

evidenced both by the presence of the scarp itself and by the existence of

marine deposits from the upper Suwannee River, in Hamilton County,

Florida. Later, as Pliocene sea levels gradually dropped, there was a

corresponding increase in the amount of land exposed (Robertson

1976). Considering that much of the freshwater habitat present today

and during pre-Pliocene times in northern Florida and southern Geor-

gia was absent during the Pliocene, the question may be asked as to

how and where Micropterus notius and Ictalurus serracanthus (as well

as other freshwater fishes) survived.

It is reasonable to assume that both of the above-named species

occupied much of their present ranges during the latter half of the Mio-

cene. During the Pliocene, however, a considerable amount of dis-

placement must have occurred. One possibility is that these species

essentially remained in place during the marine invasion, and found

refuge in the many freshwater springs located throughout the Suwannee

River valley. It appears likely, in fact, that the populations of troglobitic

crayfishes living in the region survived in this way, although Franz and

Lee (1982) did not address this particular point in their discussion of

Florida cave crayfish zoogeography. It seems, however, that the physi-

cal and ecological resources of these springs would be much too limited

for M. notius and /. serracanthus, particularly considering that neither

is today intimately associated with such areas. Furthermore, neither

species is known to go underground. A more likely possibility is that

they simply occupied more confined ranges upstream during the Plio-

cene, and at times may have been found only above the scarp. Although

upstream habitat today seems marginal for these species (particularly

for M. notius ), this may not always have been the case. The distributions

of both species in the Suwannee and Ochlockonee rivers, together with

their near absence from all intervening drainages (/. serracanthus is

present in the St. Marks), strongly suggest that headwater stream

transfer was involved (Gilbert 1980g, Yerger 1980). This presumably

48

Carter R. Gilbert

would have been in the area around Moultrie, Georgia, where these two

drainages come in close proximity (Fig. 1).

Reference was made earlier to the caves and springs that are so

prevalent throughout the northern half of peninsular Florida (Rosenau

et al. 1977). These result from the highly soluble limestone substrate

that underlies the region, and they support a distinctive and highly

endemic invertebrate fauna, particularly the crayfishes (Franz and Lee

1982). Many vertebrate animals inhabit the springs and spring runs,

although only a few move very far into the underground aquifer (Relyea

and Sutton 1973). One of these, the cyprinid fish Notropis harperi,

probably is more closely associated with a spring habitat than any other

fish in eastern North America (Gilbert 1980c), and in fact this extremely

close ecological association was an important consideration in hypothe-

sizing a north Florida-south Georgia origin for the species. Despite this,

no troglobitic fish is known from either this area or the independent

Marianna cave system in southeastern Alabama, southwestern Georgia

and closely adjacent areas in Florida. No other troglobitic vertebrates

are known from the north Florida-south Georgia area, although a

cavernicolous salamander, Haideotriton wallacei , is known from caves

in the Marianna area and from one cave in the adjacent Dougherty

Plain of southwestern Georgia.

Notropis harperi and Ictalurus natalis move considerable distances

underground, and are the only fish species in the north Florida area that

have naturally reached isolated sink holes and springs in this way

(Hubbs and Crowe 1956, Relyea and Sutton 1973). Hubbs and Crowe

(1956) described the isolated sinkhole populations of N. harperi as a

new subspecies, N. h. subterranea, but Howell (1960) showed this not to

be valid.

This brings us to the question of why troglobitic fishes apparently

have not evolved in the Florida caves, especially considering that obli-

gate cavernicoles are numerous among the crustaceans and other inver-

tebrates of peninsular Florida. This situation becomes even more enig-

matic in view of the presence of Chologaster cornuta , a species of the

cavefish family Amblyopsidae, on the Atlantic slope as far south as cen-

tral Georgia (Cooper and Rohde 1980). A congener, Chologaster agas-

sizi, inhabits springs and caves of the central Mississippi valley (Cooper

1980). Chologaster cornuta occurs in quiet, dark, well-protected situa-

tions that in some respects resemble the cave environment (Poulson

1963). Based on habitat preferences and present geographic distribu-

tions, it is virtually certain that this amblyopsid group reached the

Atlantic slope via movement along the coastal plain, presumably prior*

to the Pliocene, and thus should have had access to Florida caves. It is

Freshwater Fish Zoogeography

49

particularly difficult to understand the absence of amblyopsids from the

Marianna caves, considering that this area lies above the Cody Scarp,

and thus should not have been affected by higher sea levels during the

Pliocene. Since these fishes apparently did not reach the Marianna

caves, it is even less likely that they ever reached those caves to the east.

Mention should also be made of the absence of Florida troglobitic

fishes of marine derivation. Two genera, comprising four species, of the

otherwise exclusively marine family Bythitidae occur in fresh and brack-

ish waters of caves in the Bahamas, Cuba and the Yucatan peninsula

(Hubbs 1938; Cohen and Robins 1970; Burgess 1983a, b,c). Considering

this, it is reasonable to think that species of this family might also occur

in Florida caves. Various reasons for their absence could be advanced,

involving a combination of geographical distance, opportunity, and tim-

ing of evolutionary and geological events. For the moment, at least,

there are no obvious answers to this question.

ACKNOWLEDGMENTS.— I wish to thank S. David Webb and

Gary S. Morgan, Florida State Museum, and Anthony F. Randazzo,

Department of Geology, University of Florida, for their helpful advice

and comments pertaining to Florida geology, and also for reading and

commenting upon an earlier draft of this paper.

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Freshwater Fish Zoogeography

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Accepted 4 November 1985

Evolution of the Intercalary Cartilage

in Chorus Frogs, Genus Pseudacris

(Salientia: Hylidae)

Gary L. Paukstis

Department of Zoology,

Colorado State University, Fort Collins, Colorado 80523

AND

Lauren E. Brown

Department of Biological Sciences,

Illinois State University, Normal, Illinois 61761

ABSTRACT. — A comparison was made of the structure of interca-

lary cartilages of the digits of three groups of hylid frogs: (1) arboreal

Hyla; (2) terrestrial Pseudacris not known to burrow; and (3) forelimb

burrowing Pseudacris. In the arboreal Hyla the biconcave intercalary

cartilage has an apex shorter than the base and a height greater than

the length. A nearly cuboidal intercalary cartilage characterizes terres-

trial Pseudacris not known to burrow, whereas a reduced, wafer-

shaped intercalary is found in the forelimbs of forward-burrowing

Pseudacris. In the arboreal Hyla and Pseudacris not known to bur-

row, the cartilage offsets the positions of the terminal and subterminal

phalanges so that the terminal phalanx is displaced ventrally relative to

the other phalanges. Forward-burrowing Pseudacris do not exhibit

this offsetting of digital elements in the forelimb. In the arboreal Hyla

the shape and spatial relationship of the intercalary cartilage to the

other phalanges probably facilitate flexion of the end of the digit and

enhance arborealism. In contrast, the wafer-shaped intercalary carti-

lage in forward-burrowing Pseudacris could be considered a degenera-

tive condition resulting from the evolution of forelimb burrowing.

In spite of the predominance of arborealism in the frog family

Hylidae, several lines have evolved terrestrial habits. Of considerable

interest among the terrestrial frogs of the Hylidae are the seven chorus

frog species of the genus Pseudacris, of which five species are found in

the southeastern United States. Two species, Pseudacris ornata and P.

streckeri, are highly unusual in that they burrow forward into the sub-

strate using their forelimbs (Brown et al. 1972, Brown and Means 1984).

Important morphological adaptations for this type of burrowing include

thick forelimbs, enlarged tubercles on the palms of the hands, stout fin-

gers, and the absence (or great reduction) of toe pads. In contrast, the

arboreal hylids have long, narrow limbs and digits, and enlarged, adhe-

sive toe pads. The vast majority of other fossorial species of salientians

Brimleyana No. 13:55-61, July 1987

55

56

Gary L. Paukstis and Lauren E. Brown

in other genera dig backwards by scraping soil with their hind feet

(Emerson 1976). The most conspicuous morphological adaptations for

this type of burrowing are well-developed metatarsal tubercles on the

undersides of the hind feet. Thus, P. ornata and P. streckeri are quite

different from their relatives, the hylid treefrogs, and also from other

burrowing salientians in more distantly related groups.

Brown et al. (1972) showed experimentally that P. streckeri had a

strong preference for digging in sand and that the species was unable to

burrow in more resistant black prairie sod because of its unusual

method of digging. This helps explain the restriction of P. streckeri in

the Midwest to areas with sandy soils (Smith 1966, Brown and Brown

1973, Axtell and Haskell 1977). Brown and Means (1984) also found

that P. ornata is associated with easily penetrated sandy soils. An exper-

iment conducted by Brown (1978) demonstrated that P. streckeri can

feed while underground, and it is probable that this is one of the main

reasons why forward burrowing evolved in P. streckeri instead of back-

ward burrowing. No other species of salientian is known to feed under-

ground. Pseudacris ornata and P. streckeri have rarely been found on

the surface except during their breeding seasons, and both species emit

vocalizations underground that may be of an aversive nature (Brown

1978 and unpubl. data, Brown and Means 1984, Smith 1966).

The other species of Pseudacris — P. brachyphona, P. brimleyi, P.

clarki, P. nigrita, and P. triseriata — live on the surface of the ground

and occasionally climb on low vegetation. These species, which have not

been observed to burrow, have narrow limbs and thin fingers with small

toe pads. Thus, the subjects of our research are three groups of frogs in

the family Hylidae that differ considerably in behavior, habitat, and

gross morphology.

An extra element termed the intercalary cartilage occurs between

the terminal and subterminal phalanges of the digits of members of the

family Hylidae. Although the intercalary cartilage is probably of con-

siderable importance for arborealism, relatively little has been published

on its structure. Publications of greatest significance that deal with this

subject include Howes and Davies (1888), Noble and Jaeckle (1928),

Noble (1931), Savage and De Carvalho (1953), and Tyler and Davies

(1978). These studies examined a number of species of frogs in the fam-

ily Hylidae, but none considered the intercalary cartilage of Pseudacris .

The objective of our study was to compare the intercalary cartilages

of the digits of the hylid treefrogs, the more terrestrial species of Pseu-

dacris that are not known to burrow, and the forward-burrowing spe-

cies of Pseudacris. Primary emphasis was placed on determining how

the specialized intercalary cartilage in the family Hylidae was modified

through evolution to facilitate forward burrowing.

MATERIALS AND METHODS

Histological preparations were made of the digits of 122 alcohol-

preserved specimens of Pseudacris: 16 P. brachyphona, 8 P. brimleyi, 8

Intercalary Cartilages of Chorus Frogs

57

P. clarki, 18 P. nigrita, 24 P. ornata, 20 P. streckeri, and 28 P. triseriata.

Skeletons of an additional 12 specimens were examined: 2 P. nigrita, 1

P. ornata, 5 P. streckeri, and 4 P. triseriata. Histological preparations

from the following representative arboreal hylids were also examined: 1

Hyla andersoni, 1 H. chrysoscelis, 1 H. cinerea, 1 H. femoralis, 4 H.

squirella, and 2 H. versicolor. Specimens were obtained from the Amer-

ican Museum of Natural History, Charleston Museum, Florida State

Museum, Illinois State University, Texas Memorial Museum, and Uni-

versity of Southeastern Louisiana, as well as from private collections of

L. E. Brown, R. S. Funk, and G. L. Paukstis.

From 1 to 18 digits (usually 2) per preserved specimen were

removed and individually labeled with letters and numbers according to

the location of the digit on the body. Slide preparation was based pri-

marily on the techniques of Humason (1962). Digits were moved

through a 50% to 100% ethanol dehydration series and were vacuum

embedded in paraffin. Serial, sagittal sections (7 microns thick) were

prepared using a rotary microtome. Slides were stained in either Dela-

field’s hematoxylin and eosin or Mallory’s triple stain. Representative

whole specimens and individual digits were also cleared and stained for

bone and cartilage following the methods of Hardaway and Williams

(1975) and Dingerkus and Uhler (1977). Photomicrographs were taken

with a Zeiss Photomicroscope II using Kodak High Contrast Copy Film

at an ASA film speed of 18. Comparisons of length (distance between

articulating surfaces) and height (determined at a right angle through

the midpoint of the length distance) were made in the center of the

mid-sagittal section of the intercalary cartilages.

Morphological terminology follows Paukstis (1978).

RESULTS

Three major types of intercalary cartilages were found in the spec-

imens examined. The first type was found in Hyla andersoni (Fig. 1A),

H. chrysoscelis, H. cinerea, H. femoralis, H. squirella, H. versicolor,

and numerous other species of arboreal Hyla examined by Paukstis

(1978). In these species the intercalary cartilage is characterized as fol-

lows: height greater than length; distal articulating surface generally

more concave than proximal articulating surface; apex shorter than

base. This type of intercalary cartilage always offsets the positions of the

terminal and subterminal phalanges so that the terminal phalanx is dis-

placed vertically and ventrally relative to other phalanges of the digit

(Fig. 1A).

The second type of intercalary cartilage was found in P. brachy-

phona, P. brimleyi, P. clarki, P. nigrita, and P. triseriata. Their interca-

lary cartilages are nearly cuboidal (Fig. IB). The terminal phalanx is

offset from the other phalanges by the intercalary cartilage in the same

manner as in the arboreal hylids examined.

The third type of intercalary cartilage is quite reduced, with the

curvatures of articulating surfaces nearly equal and very shallow. This

58

Gary L. Paukstis and Lauren E. Brown

reduced or wafer-shaped cartilage was found in the digits of the fore-

limbs of P. ornata and P. streckeri (Fig. 1C). The intercalary cartilages

of digits of the hind limbs are variable. They may be nearly wafer-

shaped or cuboidal or of various intermediate shapes. Pseudacris ornata

and P. streckeri show no vertical offsetting of the digital elements in the

forelimb by the intercalary cartilage. This condition is not as well de-

veloped in digits of the hind limbs in these species.

DISCUSSION

The intercalary cartilage of frogs that climb probably facilitates

flexion of the end of the digit so that the toe pad may be more effi-

ciently used for climbing and more readily applied to a vertical surface

(Paukstis 1978). This type of flexion would seem disadvantageous and

inefficient for forelimb burrowing because the terminal phalanx would

move upwards when the digit is pushed against the substrate. In P.

ornata and P. streckeri the problem of upward flexion has been resolved

by the reduction in size of the intercalary cartilage and elimination of

vertical offsetting of the terminal phalanx. These morphological charac-

teristics have evolved to provide a relatively efficient mechanism of fore-

limb burrowing in these species. It is probably appropriate to consider

the thin, wafer-shaped intercalary cartilage of P. ornata and P. streckeri

to be a degenerative condition, and further evolution in these species

might result in loss of the cartilage.

Paukstis (1978) noted that the nearly cuboidal intercalary cartilage

characteristic of the species of Pseudacris that are not known to burrow

is also found in some other semiterrestrial species in the family Hylidae,

such as Acris crepitans, A. gryllus, Hyla crucifer, and Limnaeodus ocu-

laris. It is possible that the semiterrestrial groups of frogs in all four of

the aforementioned genera are independent lines that have separately

evolved cuboidal intercalary cartilages. Thus, this may be an example of

convergent evolution. On the other hand, the similar cuboidal interca-

lary cartilages in these different genera may have descended relatively

unchanged from an ancient common ancestor that was semiterrestrial in

habits.

The nearly cuboidal intercalary cartilage of the terrestrial species of

Pseudacris that are not known to burrow may represent an intermediate

evolutionary stage between those cartilages in the arboreal species of

Hyla and those in the forelimbs of the fossorial P. ornata and P. streck-

eri. The occasional occurrence of cuboidal intercalary cartilages in the

hind limbs of the last two species supports this suggestion. However, the

cuboidal intercalary cartilage is longer (relative to width) than either of

the other two types of intercalary cartilages. Thus there would have had

to have been selection for increase in length and then selection for

marked decrease in length if the cuboidal intercalary cartilage is the

intermediate stage. Another interpretation is also possible. The cuboidal

Intercalary Cartilages of Chorus Frogs

59

Fig. 1. Diagrams of mid-sagittal sections of three types of intercalary cartilages

(IC) from the digits of forelimbs of frogs in the family Hylidae: A. Hyla ander-

soni, showing the type found in numerous arboreal hylids; B. Pseudacris nigrita,

showing the cuboidal type found in terrestrial chorus frogs not known to bur-

row; C. Pseudacris streckeri, showing the reduced or wafer-shaped type found in

the forelimbs of forward-burrowing chorus frogs.

60

Gary L. Paukstis and Lauren E. Brown

intercalary cartilage may represent an ancestral condition from which

both of the other types were independently derived. This can explain the

occasional occurrence of cuboidal intercalary cartilages in the hind

limbs of P. ornata and P. streckeri, while not requiring complex shifts

in selection on this level of evolution. We do not mean to advocate that

the genus Hyla was derived from Pseudacris with cuboidal intercalary

cartilages (our comments here are restricted to the evolution of the

intercalary cartilage). The cuboidal intercalary cartilages of the extant

Pseudacris and the intercalary cartilages of the extant arboreal Hyla

may both be derived from a cuboidal intercalary cartilage of a semiter-

restrial ancestral form. There are at least seven other possible evolution-

ary histories involving various combinations of the three types of inter-

calary cartilages. However, these schemes lack a middle stage involving

a cuboidal intercalary cartilage that would indicate an intermediate

nonburrowing, terrestrial frog, or they involve a reversal of the direction

of selection in regard to the length of the intercalary cartilage; some

encounter both problems. Even if it is assumed that the cuboidal inter-

calary is the ancestral stage, there are still problems of interpretation in

regard to reversal in direction of selection. Presumably, the ancestral

condition from which the cuboidal intercalary cartilage was derived

could have been absence of the cartilage. Thus, the cuboidal intercalary

cartilage would have passed through a stage that was shorter in length

during the course of its evolutionary history. Consequently, both ances-

tors and derivatives of the cuboidal intercalary cartilage would have

been shorter, if this interpretation is followed.

ACKNOWLEDGMENTS. — We thank J. Brown for preparing slides

in the early phases of this study; W. Auffenberg (Florida State Museum),

R. Funk, E. Keiser (University of Southeastern Louisiana), R. Martin

(Texas Memorial Museum), A. Sanders (Charleston Museum), and R.

Zweifel (American Museum of Natural History) for the loan of speci-

mens and materials; R. Anderson, J. Brown, and E. Mockford for criti-

cally reading the manuscript; and Jane Bucci for drawing Figure 1.

LITERATURE CITED

Axtell, Ralph W., and N. Haskell. 1977. An interhiatal population of Pseu-

dacris streckeri from Illinois, with an assessment of its postglacial disper-

sion history. Nat. Hist. Misc. No. 202:1-8.

Brown, Lauren E. 1978. Subterranean feeding by the chorus frog, Pseudacris

streckeri (Anura: Hylidae). Herpetologica 34(2):21 2-2 1 6.

, and J. R. Brown. 1973. Notes on breeding choruses of two anurans

( Scaphiopus holbrookii, Pseudacris streckeri) in southern Illinois. Nat.

Hist. Misc. No. 192:1-3.

Intercalary Cartilages of Chorus Frogs

61

, H. O. Jackson, and J. R. Brown. 1972. Burrowing behavior of the

chorus frog, Pseudacris streckeri. Herpetologica 28(4): 325-328.

, and D. B. Means. 1984(1985). Fossorial behavior and ecology of

the chorus frog Pseudacris ornata. Amph.-Rept. 5(3-4):261-273.

Dingerkus, Guido, and L. D. Uhler. 1977. Enzyme clearing of alcian blue

stained whole small vertebrates for demonstration of cartilage. Stain

Technol. 52(4):229-232.

Emerson, Sharon B. 1976. Burrowing in frogs. J. Morphol. 149(4):437-458.

Hardaway, Thomas E., and K. L. Williams. 1975. A procedure for double

staining cartilage and bone. Br. J. Herpetol. 5(4):473-474.

Howes, G. B., and A. M. Davies. 1888. Observations upon the morphology

and genesis of supernumerary phalanges, with especial reference to those of

the Amphibia. Proc. Zool. Soc. London 1888:495-511.

Humason, Gretchen L. 1962. Animal Tissue Techniques. W. H. Freeman and

Co., San Francisco and London.

Noble, Gladwyn Kingsley. 1931. The Biology of the Amphibia. McGraw-Hill

Book Co., New York; reprinted 1954, Dover Publications, New York.

, and M. E. Jaeckle. 1928. The digital pads of the tree frogs. A

study of the phylogenesis of an adaptive structure. J. Morphol. Physiol.

45(l):259-292.

Paukstis, Gary L. 1978. Phylogenetic significance of interspecific variation in

the morphology of the intercalary cartilage of certain salientians. M.S.

thesis, Illinois State University, Normal.

Savage, Jay M., and A. L. De Carvalho. 1953. The family position of Neo-

tropical frogs currently referred to the genus Pseudis. Zoologica (NY)

38(17): 193-200.

Smith, Philip W. 1966. Pseudacris streckeri. Cat. Amer. Amphib. Rept.

27.1-27.2.

Tyler, Michael J., and M. Davies. 1978. Species-groups within the Australo-

papuan hylid frog genus Litoria Tschudi. Aust. J. Zool., Suppl. Ser. No.

63:1-47.

Accepted 17 January 1986

62

THE SEASIDE SPARROW,

ITS BIOLOGY AND MANAGEMENT

Edited by

Thomas L. Quay, John B. Funderburg, Jr., David S. Lee,

Eloise F. Potter, and Chandler S. Robbins

The proceedings of a symposium held at Raleigh, North Carolina,

in October 1981, this book presents the keynote address of F. Eugene

Hester, Deputy Director of the U. S. Fish and Wildlife Service, a bibli-

ography of publications on the Seaside Sparrow, and 16 major papers

on the species. Authors include Arthur W. Cooper, Oliver L. Austin, Jr.,

Herbert W. Kale II, William Post, Harold W. Werner, Glen E. Wool-

fenden, Mary Victoria McDonald, Jon S. Greenlaw, Michael F. Delany,

James A. Mosher, Thomas L. Merriam, James A. Kushlan, Oron L.

Bass, Jr., Dale L. Taylor, Thomas A. Webber, and George F. Gee. A

full-color frontispiece by John Henry Dick illustrates the nine races of

the Seaside Sparrow, and a recording prepared by J. W. Hardy supple-

ments two papers on vocalizations.

“The Seaside Sparrow, with its extensive but exceedingly narrow

breeding range in the coastal salt marshes, is a fascinating species. All

the authors emphasize that the salt marsh habitat is at peril. . . . The

collection is well worth reading.” — George A. Hall , Wilson Bulletin.

1983 174 pages Softbound

Price: $15, postpaid. North Carolina residents add 4*4% sales tax. Please make

checks payable in U. S. currency to NCDA Museum Extension Fund.

Send to SEASIDE SPARROW, N. C. State Museum of Natural History,

P. O. Box 27647, Raleigh, NC 27611.

Abundance and Distribution of Shrews

in Western South Carolina

Michael T. Mengak, David C. Guynn, Jr., J. Kenneth

Edwards, Diane L. Sanders, and Stanlee M. Miller1

Department of Forestry , Lehotsky Hall

and

1 Department of Biological Sciences, Long Hall,

Clemson University, Clemson, South Carolina 29631.

ABSTRACT. — New information on distributions of six species of

shrews in western South Carolina is presented. Blarina carolinensis

was the most abundant shrew in our sample. Blarina brevicauda was

collected less often than expected in hardwood forests (P < 0.05).

Sorex fumeus and S. longirostris were caught more often than expected

in floodplain habitats (P < 0.01). The first records of Sorex hoyi are

reported from South Carolina.

Shrews are often poorly represented in surveys of small mammals,

and the relative abundance of various species or even their within-state

distributions are frequently unclear (Golley 1966, Wolfe and Esher

1981). The southeastern shrew ( Sorex longirostris ) is considered rare in

the Southeast (French 1980a) and is not known to have a wide distribu-

tion in South Carolina (Golley 1966, French 1980a). The pygmy shrew

{Sorex hoyi ) is suspected to occur in South Carolina (Diersing 1980) but

has never been reported (Golley 1966, Thompson 1982). The least shrew

{Cryptotis parva) was the least abundant shrew in a Mississippi study

(Wolfe and Esher 1981). Taxonomic work on the short-tailed shrews

{Blarina spp.) was reported by French (1981) and Braun and Kennedy

(1983), but little is known of the habitat requirements of these species

(see, for example, Wolfe and Esher 1981). Our objective is to update the

information on the abundance and distribution of shrews in western

South Carolina.

From 1981 through 1985, small mammals were collected through-

out the year from eight counties in the Piedmont and Mountain regions

of South Carolina. The counties sampled are in the western portion of

the state, along or near the Georgia border including (proceeding

approximately from southeast to northwest): Saluda, Edgefield,

McCormick, Greenwood, Abbeville, Anderson, Oconee, and Pickens.

Habitats sampled were loblolly pine {Pinus taeda ) stands (0 to 25 years

old), hardwood forests (30 to 90 years old), and flood plains along first-

and second-order streams. Pine stands were sampled in all eight coun-

ties, whereas flood plains and hardwood stands were sampled in the

three northernmost counties.

Brimleyana No. 13:63-66, July 1987

63

64

Michael T. Mengak et al.

Trapping consisted of snap trapping with Victor mouse traps and

Museum Specials and using pitfall traps with drift fences. Snap traps

were placed in lines with a minimum of 20 m between lines and 10 m

between stations on a line. Trap lines had five or eight stations depend-

ing on size of the study area. Three traps were set at each station and

baited with peanut butter. Snap trap types were not mixed within a

study site, and each site had a series of pitfalls and drift fences. On pine

and hardwood sites, pitfalls with drift fences were constructed as de-

scribed in Campbell and Christman (1982). At floodplain sites, fences

were parallel to the stream and 1 to 4 m from the water’s edge. Water

was maintained in all pitfalls so that captured animals drowned. All

specimens are currently deposited in the Vertebrate Collections of

Clemson University and The Charleston Museum.

Combined trapping effort with snap traps and pitfalls equals 61,404

trap nights. Effort was not distributed evenly among habitat types. A

total of 367 shrews were collected from all habitats: 243 (66%) from pine

stands, 21 (6%) from hardwood stands, and 103 (28%) from flood plains

(Table 1). Southern short-tailed shrews were captured most frequently

(42% of total catch), followed by southeastern shrews (32%), least

shrews (11%), northern short-tailed shrews (10%), smoky shrews (5%),

and pygmy shrews (1%). Chi-square analysis was used to test the

hypothesis that catchability was equal for all habitat types.

The catchability of five species of shrews was compared across hab-

itat types (Table 1); southern short-tailed shrews were not considered

because, within their range, only one habitat type was sampled. South-

eastern shrews were captured more often than expected in flood plains

(P < 0.01) and less often than expected in pine stands (P < 0.01) (Table

1). Wolfe and Esher (1981) felt that this species is not habitat specific in

Mississippi. However, our data support French (1980a), who compiled

habitat information provided by collectors on specimen labels and in

field work and noted that most specimens came from moist areas. The

northern short-tailed shrew was found less often than expected in hard-

wood stands (P < 0.05) and more often than expected in pine stands (P

< 0.05). Smoky shrews were taken in the mountainous areas of Pickens

and Oconee counties. This species was collected less often than expected

in pine stands (P< 0.01) and more often than expected in flood plains

(PC0.01).

Five specimens of Sorex hoyi were collected in 1984 and 1985.

These are the first known records of the pygmy shrew for South Caro-

lina. Two females were collected 2.1 km SW of Whetstone Corner,

Oconee County, on 21 June and 28 November 1984. The first male was

collected on 22 June 1984, 1.9 km SE of Round Mountain, Oconee

County. Another male was collected 0.5 km SW of the first collection

Shrews in Western South Carolina

65

Table 1. Relative abundance (percent of total catch) of shrews collected from

three habitats in eight western South Carolina counties from 1981

through 1985.

site on 26 June 1985. The third was collected 48.3 km to the east at

Laurel Fork Creek, above Lake Jocassee in Pickens County on 29

October 1985. All five specimens were captured in pitfalls. The nearest

known locations for this species are 32 km to the north of the Laurel

Fork Creek site at Cedar Mountain, Transylvania County, North Caro-

lina (CUSC 532), and 64 km to the west of the Round Mountain site in

Townes County, Georgia (Wharton 1968).

The two females were collected at an elevation of 503 m in a 75-

year-old mixed pine-hardwood forest. The first two males were collected

at 488 and 491 m in 17-year-old loblolly pine plantations. The Laurel

Fork Creek site is a 100-year-old hemlock ( Tsuga canadensis) forest at

66 Michael T. Mengak et al.

402 m. All specimens were identified by their small third and fifth uni-

cuspid teeth.

The occurrence of shrews is not well documented in South Carolina.

Although Golley (1966) felt the least shrew was distributed statewide,

only five specimens were reported from the western part of the state.

Similarly, the southeastern shrew was reported only in the Coastal Plain

and Aiken County. Prior to our investigation the southern short-tailed

shrew was not known from Greenwood County (French 1981). The

occurrence of shrews in eight western South Carolina counties is given

in Table 2. We captured more species and documented a different dis-

tribution of relative abundance than Wolfe and Esher (1981) reported

from similar habitats in Mississippi. Our results reflect the presence of

several northern species, such as Blarina brevicauda and Sorex fumeus ,

whose ranges extend southward along the Appalachian Mountains into

western South Carolina. We agree with French (1980b) that when using

appropriate trapping techniques in favorable habitats, the southeastern

shrew is easily captured and frequently found to be an abundant

member of the local fauna.

LITERATURE CITED

Braun, Janet K., and M. L. Kennedy. 1983. Systematics of the genus Blarina in

Tennessee and adjacent areas. J. Mammal. 64:414-425.

Campbell, Howard W., and S. P. Christman. 1982. Field techniques for herpe-

tological community analysis. Pages 193-200 in Herpetological Communi-

ties, Norman J. Scott, Jr., editor. USDI Fish and Wildl. Ser., Wildl. Res.

Rep. 13.

Diersing, Victor E. 1980. Systematics and evolution of the pygmy shrews

(Subgenus Microsorex) of North America. J. Mammal. 61:76-101.

French, Thomas W. 1980a. Sorex longirostirs. Mamm. Species 143:1-3.

. 1980b. Natural history of the southeastern shrew, Sorex longiros-

tris Bachman. Am. Midi. Nat. 104:13-31.

. 1981. Notes on the distribution and taxonomy of short-tailed

shrews (genus Blarina) in the Southeast. Brimleyana 6.T0 1-1 10.

Golley, Frank B. 1966. South Carolina Mammals. Contrib. Charleston Mus.

Charleston.

Thompson, E. F. 1982. A Guide to the Amphibians, Reptiles and Mammals of

South Carolina. State Printing Co., Inc., Columbia.

Wharton, Charles H. 1968. First records of Microsorex hoyi and Sorex cine-

reus from Georgia. J. Mammal. 49:158.

Wolfe, James L., and R. J. Esher. 1981. Relative abundance of the southeast-

ern shrew. J. Mammal. 62:649-650.

Accepted 14 May 1986

Unionid Mollusks from the Upper Cape Fear River

Basin, North Carolina, with a Comparison of the

Faunas of the Neuse, Tar, and Cape Fear Drainages

(Bivalvia: Unionacea)

Rowland M. Shelley

North Carolina State Museum of Natural Sciences,

P.O. Box 27647, Raleigh, North Carolina 27611

ABSTRACT. — The unionid molluscan fauna of the piedmont portion

of the Cape Fear River System consists of 10 genera and 15 species.

Six additional species in the Coastal Plain part of the basin are known

from literature records, and the reported occurrence of Alasmidonta

heterodon (Lea 1830) cannot be verified. Elliptio complanata (Light-

foot 1786), Uniomerus tetralasmas (Say 1831), Anodonta cataracta

Say 1817, and Villosa delumbis (Conrad 1834) are numerical domi-

nants, while Fusconaia masoni (Conrad 1834), Carunculina pulla

(Conrad 1838), Lampsilis cariosa (Say 1817), Anodonta imbecilis Say

1829, and Lasmigona subviridis (Conrad 1835) are least abundant.

Photographic plates and diagnostic comments are provided to facili-

tate identifications of unionids in the Atlantic drainages of North

Carolina.

Thirty years have elapsed since Walter (1956) surveyed the mol-

lusks, of the upper Neuse River basin and provided the first detailed

study on these invertebrates in a North Carolina drainage. Nine years

later Dawley (1965) published an uncritical listing of freshwater mol-

lusks for the entire state, based in part on Walter’s paper, the author’s

personal collection, and materials at the Academy of Natural Sciences,

Philadelphia, and the Museum of Zoology at the University of Michi-

gan. Clarke (1983) published a comprehensive report on the mollusks of

the Tar River system as part of an assessment of the status of the

endemic spiny mussel, Elliptio ( Canthyria ) steinstansana Johnson and

Clarke 1983. This study also included less intensive sampling in parts of

the Roanoke (Roanoke and Cashie rivers) and Neuse (Neuse, Little, and

Trent rivers) watersheds. Porter (1985) conducted a detailed sampling of

the mollusks of Lake Waccamaw and the Waccamaw River system.

These four works are the major ones dealing exclusively with aquatic

mollusks of North Carolina, and additional records are available in

Johnson (1967, 1970, 1984), Johnson and Clarke (1983), Fuller (1972,

1973, 1977), Burch (1975), Porter and Horn (1980, 1983, 1984), Horn

and Porter (1981), Clarke (1981, 1983, 1985), and Shelley (1983). Earlier

literature pertinent to the state is summarized by Walter (1956).

From 1971 to 1978 I sampled unionids non-quantitatively in the

Piedmont Plateau section of the Cape Fear River basin, mostly in lower

Brimleyana No. 13:67-89, July 1987

67

68

Rowland M. Shelley

reaches of the Haw and Deep rivers, which join along the Chatham-Lee

counties line to form the Cape Fear River proper. Sampling of the live

individuals and empty valves on the banks took place at public access

points, and smaller creeks were waded for most of their lengths. In addi-

tion, Dr. Charlotte Dawley donated to the State Museum her collection

of mollusks, most of which came from headwater areas of the Haw

River in Guilford County and formed the basis for her checklist (1965).

Thus her material plus mine spans the upper part of the basin, and this

contribution supplements the works of Walter and Clarke in detailing

the unionid fauna of central North Carolina. For the sake of comple-

tion, I include records and species from the lower or Coastal Plain sec-

tion of the basin and also summarize reports from other North Carolina

drainages. The paper concludes with a discussion and comparison

(Table 1) of the faunas of the Tar, Neuse, and Cape Fear basins, the

three major drainages located wholly within the state.

THE CAPE FEAR RIVER BASIN1

The Cape Fear River basin, the largest wholly within North Caro-

lina, drains about 18% of the state (14,624 km2) and all or part of 26

counties (Fig. 1,2). The basin is roughly 322 km long by a maximum of

96 km wide, extending from northwest of Greensboro to southeast of

Wilmington. About 6 billion gallons of water a day flow into the Atlan-

tic Ocean through the Cape Fear estuary, the only major one in North

Carolina with direct access to the sea. Three locks and dams in Bladen

County control flow of the Cape Fear River, and tidal influences are felt

to the lowest of these, approximately 59 river km above Wilmington or

just above site L (Fig. 2). The upper third of the watershed is in the

Piedmont Plateau Physiographic Province and is characterized by

undulating terrain with relatively deep valleys and narrow flood plains.

Most of the basin, however, lies in the Sandhills and Coastal Plain prov-

inces, where flat terrain causes relatively sluggish streams. Between the

Piedmont Plateau and Coastal Plain is the Fall Zone, so named because

of the small, discontinous rapids formed as water passes from the con-

solidated rock substrates of the Piedmont onto the unconsolidated sed-

iments of the Coastal Plain. Elevations in the basin range from mean

sea level to 90 to 120 m in the Fall Zone to around 300 m in the

headwaters.

The Cape Fear River begins on the inner edge of the Fall Zone

near the town of Moncure, and the area of study was entirely in the

Haw and Deep river sub-basins, of which the former, draining an area

of 2,712 km2, is slightly larger. This sub-basin is about 1 12 km long and

1 All information in this section is from the 1983 report titled “Status of Water

Resources in the Cape Fear River Basin,” North Carolina Department of Natu-

ral Resources and Community Development, Raleigh.

Unionid Mollusks

69

Table 1. Comparison of the native unionid faunas of the Tar, Neuse, and Cape

Fear river systems.

1 Clarke (1983) also reported a live specimen of Anodontoides ferussacianus

(Lea) from the upper reaches of the Tar in Granville County, but Johnson

(1970) does not record this species from any Atlantic drainage, even as a junior

synonym. Burch (1975) gives the range as the interior basins of the United

States and Canada, the Great Lakes, and the St. Lawrence River drainages.

2 Taken from Johnson (1970).

3 Based on Johnson (1970), which is in accordance with Burch (1975), who

reports the range as being from the Altamaha River system of Georgia to the

Chowan of North Carolina. However Clarke (1983) questioned the concept of

U. tetralasmus and considered it absent from the Roanoke, Tar, and Neuse

systems.

70

Rowland M. Shelley

Fig. 1. Map of North Carolina showing the location and proportionate size

of the Cape Fear Basin.

Fig. 2. PP, Piedmont Plateau; FZ, Fall Zone.

Unionid Mollusks

71

Fig. 2. The Cape Fear Basin. The dashed lines indicate the approximate

boundaries of the Fall Zone. PP, Piedmont Plateau; FZ, Fall Zone; CP, Coastal

Plain. Collecting sites are as follows (numbers and dots indicate new sites

sampled in this study, letters and squares indicate ones cited in the literature):

Piedmont Plateau Physiographic Province

Haw River Sub-basin

Rockingham County

A. Haw R. ca. 2.0 km NE Benaja (Johnson 1970)

Guilford County

1. Lake Brandt

2. Lake at Greensboro Country Park

3. Lake at Boy Scout Camp

4. Lake Philadelphia

5. Lake at Hamilton Lakes subdivision

6. Lake on UNC-Greensboro campus

7. Alamance Cr., precise location unknown

8. Stinking Quarter Cr., precise location unknown

9. Lake at Kimesville

B. Buffalo Cr., 1.6 km E Greensboro (Johnson 1970)

C. Travis Cr.rca. 2.4 km N Gibsonville (Johnson 1970)

Alamance County

10. Alamance Cr., precise location unknown

Orange County

1 1. University Lake, Chapel Hill

12. Booker Cr. at crossing of US 15-501, Chapel Hill

D. Morgan Cr., ca. 1.6 km SE Chapel Hill

Durham County

E. New Hope Cr., Duke Forest, Durham vicinity (Johnson 1970)

Chatham County

13. Terrell Cr. ca. 0.2 km above confluence with Haw R., ca.

13.6 km ENE Pittsboro

14. Haw R. at crossing of US 64, ca. 6.4 km ENE Pittsboro

Deep River Sub-basin

Randolph County

15. Deep R., precise location unknown

F. Sandy Cr., precise location unknown (Johnson 1970)

Moore County

16. Deep R. at Highfalls

Chatham County

17. Rocky R. at crossing of secondary road 2170, ca. 8.5 km SE Siler City

18. Rocky R. at crossing of NC 902, ca. 1 1.2 km SW Pittsboro

19. Rocky R. at crossing of secondary road 1010, ca. 17.6 km N

Sanford and 9. 1 km SW Pittsboro

20. Rocky R. ca. 0.2 km above confluence with Deep R., ca. 8.0 km S

Pittsboro

21. Bear Cr. at crossing of NC 902, ca. 14.6 km S Siler City

22. Bear Cr. at crossing of secondary road 2187, ca. 15.7 km SW Pittsboro

23. Bear Cr. at crossing of secondary road 1010, ca. 9.1 km SW Pittsboro

24. Bear Cr. at crossing or secondary road 2155, ca. 12.6 km SW Pittsboro

Chatham-Lee Counties

25. Deep R. below crossing of US 15-501 near disjunct

White Pine community, ca. 13.4 km S Pittsboro

26. Deep R. at crossing of US 1, ca. 17.9 km NE Sanford

G. Deep R. at Gulf (Johnson 1970)

72

Rowland M. Shelley

Fig. 2. CP, Coastal Plain

Unionid Mollusks

73

Coastal Plain Physiographic Province

Cape Fear River Sub-basin

Cumberland County

H. Cape Fear R. near Carlos (Johnson 1970)

I. Cape Fear R. ca. 4.8 km SW Slocomb and 9.6 km NNE

Fayetteville (Fuller 1973, 1977)

Bladen County

J. Cape Fear R. ca. 4.8 km ENE Tobermoy (Fuller 1972)

K. Cape Fear R. ca. 2.0 km ESE Duart (Fuller (1972)

L. Cape Fear R. ca. 3.2 km S Kings Bluff (Johnson 1970)

Columbus-Brunswick Counties

M. Livingston Cr., precise location unknown (Johnson 1970,

Dawley 1965)

New Hanover County

N. Greenfield Lake (formerly Greenfield Mill Pond) and outlet

stream, Wilmington (Johnson 1970)

South River Sub-basin

Sampson County

O. Six Runs Cr., precise location unknown (Johnson 1970)

Northeast Cape Fear River Sub-basin

Duplin County

P. Northeast Cape Fear R., precise location unknown

(Johnson 1970)

Pender County

Q. Ashes Cr., precise location unknown (Johnson 1970)

an average of 38 km wide. Flows are generally in a southeasterly direc-

tion from the headwaters in eastern Forsyth County to Moncure.

Stream gradients fall uniformly by 1.2 to 2.1 m per mile in the upper

two-thirds of the sub-basin, but in the lower third they vary from 0.2 to

5.4 m per mile. The Haw River sub-basin, the most densely populated

part of the Cape Fear watershed, includes Greensboro, Burlington,

Graham, Chapel Hill, Pittsboro, and parts of Reidsville and Durham.

Major tributaries of the upper Haw River include Reedy Branch, Trou-

blesome Cr^ek, and Alamance Creek. The New Hope River, arising in

central Orange County and joining the Haw a few kilometers above its

confluence with the Deep River, is the major tributary of the lower

Haw. However, B. Everett Jordan Dam, recently constructed on the

Haw below its confluence with the New Hope, impounds the latter into

Durham County.

The Deep River sub-basin drains about 2,304 km2. Headwaters

arise in eastern Forsyth County and flow southward to northern Moore

County, then northeastward to the confluence with the Haw River.

Stream gradients fall at a relatively uniform rate of 1.5 m per mile in the

upper two-thirds of the sub-basin before leveling off to about 0.5 m per

mile in the lower third. High Point and Asheboro, both on headwater

tributaries, are the only major communities in the sub-basin. The only

major tributary is the Rocky River, which arises in northeastern Ran-

dolph County and flows southeastward through Chatham County. No

74

Rowland M. Shelley

major impoundments exist in the Deep River sub-basin, but Randleman

and Howards Mill lakes were authorized for construction in the mid-

1970s (Shelley 1972). Work may soon begin on the former, but the latter

is considered economically unjustifiable for the foreseeable future.

KEY TO PIEDMONT CAPE FEAR BASIN UNIONIDAE

Two regional keys are pertinent to North Carolina’s Atlantic unio-

nids, in addition to the continental publication by Burch (1975). That by

Johnson (1970) to all southern Atlantic species is more detailed, but it is

mechanically flawed and of limited utility. Fuller’s key (1971) to the

Savannah River basin fauna is more useful but contains species absent

from this state. The following key relies mostly on conchological fea-

tures and was devised to facilitate determinations in any Piedmont

North Carolina drainage when used in combination with the diagnoses

and figures herein and in Johnson (1970).

1. Valves with lateral teeth 2

Without this character 11

2. Left valve with small interdental projection

Lasmigona subviridis (Conrad)

Without this character 3

3. Post-basal mantle margin (before or ventral to incurrent

aperture) with caruncle . . . Carunculina pulla (Conrad)

Caruncle absent 4

4. Post-basal mantle margin with ribbon-like flap of tissue ....

Lampsilis cariosa (Say)

Without this character 5

5. Mantle and viscera pink or reddish in color

Fusconaia masoni (Conrad)

Mantle and viscera white or cream colored 6

6. Shells sexually dimorphic, females swollen ventrad to accom-

modate marsupium; post-basal margin with long papil-

late projections 7

Shells not sexually dimorphic; post-basal mantle margin not

modified 8

7. Valves small, periostracum dark green to black

Villosa constricta (Conrad)

Valves moderately-large, periostracum yellow

Villosa delumbis (Conrad)

8. Valves over three times as long as high

Elliptio folliculata (Lea)

Valves about twice as long as high 9

9. Dendritic branchial papillae present

Uniomerus tetralasmus (Say)

Without this character 10

Unionid Mollusks

75

10. Post-basal mantle margin heavily pigmented; periostra-

cum dull Elliptio complanata (Lightfoot)

Post-basal mantle margin not heavily pigmented; periostra-

cum glossy Elliptio raveneli (Conrad)

11. Valves without pseudocardinal teeth 12

Valves with prominent or vestigial pseudocardinal teeth .... 13

12. Umbos extending above dorsal margin

Anodonta cataracta (Say)

Umbos flat, not protruding above dorsal margin

Anodonta imbecilis (Say)

13. Pseudocardinal teeth slight, barely detectable

Strophitus undulatus (Say)

Pseudocardinal teeth prominent 14

14. Posterior slope corrugated, with radial undulations or wrin-

kles; umbos relatively low

Alasmidonta varicosa (Lamarck)

Posterior slope smooth; umbos high, inflated

Alasmidonta undulata (Say)

SPECIES COMPOSITION

Unionid species collected in the upper Cape Fear system are listed

below along with synonyms that have been used to refer to North Caro-

lina records. Localities refer to symbols in Figure 2. Previous North

Carolina records from the Cape Fear and other drainages, observations

on preferred habitat, and the statewide conservation status reported by

Fuller (1977) are variously included under “remarks.” Subfamilies and

tribes follow the arrangement of Burch (1975).

Subfamily Amleminae

Fusconaia masoni (Conrad 1834) [= Pleurobema brimleyi (Wright)

(Walter 1956); Elliptio merus (Lea) (Dawley 1965); and Pleuro-

bema (Lexingtonia) masoni (Conrad) (Johnson 1970)] — Fig. 3.

Diagnosis'. Valves rhomboidal in outline; mantle and viscera pink or

reddish in color.

Localities'. 19, I.

Remarks'. Walter (1956) encountered F. masoni below dams in tributar-

ies of the Neuse River near Raleigh, and Clarke (1983) found it at 13

stations throughout the length of the Tar River. Johnson (1970) cited

specific localities in the Roanoke, Tar-Pamlico, Neuse, Yadkin, and

Catawba systms but recorded the Cape Fear River without further data.

Fuller (1973) placed this species in the genus Fusconaia on anatomical

grounds and reported it from the Coastal Plain section of the Cape Fear

River at site I. Fuller (1977) alluded to its occurrence in the Rocky

River and assigned F. masoni to “Threatened” status in North Carolina.

76

Rowland M. Shelley

Subfamily Unioninae

Tribe Lampsilini

Villosa constricta (Conrad 1838) [= Lignumia (Micromya) constricta

(Conrad) (Walter 1956); and Ligumia constricta (Conrad) and V.

lienosa (Conrad) (Dawley 1965)] — Fig. 4-5.

Diagnosis'. Valves small, sexually dimorphic, periostracum dark green

to black; post-basal mantal margin papillate.

Localities'. 2, 4, 19, 26, G.

Remarks'. Walter (1956) collected V. constricta at nine unspecified sites

in the upper Neuse basin; Johnston (1970) listed these along with two in

the Tar, four in the Catawba, and two in the Cape Fear systems. In the

Tar drainage, Clarke (1983) found living specimens at seven stations

and empty valves at nine. Fuller (1977) considered V. constricta to be

“Of Special Concern” to North Carolina.

Villosa delumbis (Conrad 1834) [= Ligumia ( Micromya ) delumbis (Con-

rad) (Walter 1956); Ligumia delumbis (Conrad), V. ogeecheensis

(Conrad), and V. tenera (Lea) (Dawley 1965); V. ogeecheensis

(Conrad) (Porter and Horn 1980); and V. delumbus (Conrad)

(Johnson 1984)] — Fig. 7-8.

Diagnosis'. Valves moderately large, sexually dimorphic, periostracum

yellow; post-basal mantle margin papillate.

Localities'. 9, 10, 11, 15, 16, 18, 19, 24, 26, N, P.

Remarks'. Villosa delumbis is the most common lampsiline mollusk in

the Cape Fear basin. Walter (1956) found it at only one site in the upper

Neuse, but Johnson (1970) listed two localities in the Neuse, four in the

Cape Fear, one in the Waccamaw, one in the Yadkin, and six in the

Catawba basins. Porter and Horn (1980) and Johnson (1984) reported

the species from Lake Waccamaw. The Neuse system is the northern

range limit (Johnson 1970, Burch 1975); thus, Clarke (1983) did not

encounter V. delumbis during his survey of the Tar River.

Carunculina pulla (Conrad 1838) [= Toxolasma pullus (Conrad) (Porter

and Horn 1980)]— Fig. 6

Diagnosis'. Post-basal mantal margin with prominent caruncle.

Localities'. 11, 17.

Remarks'. Johnson (1967, 1970) and Burch (1975) give the range of this

species as being from the Neuse River to Georgia. Clarke (1983) found

one questionable specimen in the Tar drainage, and C. pulla has since

been collected in the Tar River at Spring Hope, Nash County (Gerbe-

rich, pers. comm.). University Lake at Chapel Hill (site 1 1) is the most

commonly recorded North Carolina locality, but the species was also

found in this study at site 17 in the Rocky River. Johnson (1970)

recorded C. pulla from the Neuse River near Raleigh, but Walter failed

Fig. 3-10. Ambleminae and Lampsilini (Unioninae), localities in parentheses.

“M” and “F” refer to male and female shells, respectively. 3, Fusconaia masoni

(19). 4, Villosa constricta M (24). 5, V. constricta F (24). 6, Carunculina pulla

(17). 7, Villosa delumbis M (18). 8, V. delumbis F (24). 9, Lampsilis cariosa M

(14). 10, L. cariosa F (15). Scale line= 10 cm.

to find it in his study. Johnson (1970) also reported four localities near

Charlotte (Catawba drainage), and Porter and Horn (1980) listed it

from the Waccamaw system. According to Fuller (1977) the only sizea-

ble population is in the Savannah River, and because of its rarity and

the decline in recorded occurrences, he considered C. pulla “Endan-

gered” in North Carolina.

78

Rowland M. Shelley

Lampsilis cariosa (Say 1817) Fig. 9-10.

Diagnosis : Valves subovoid in outline, sexually dimorphic, periostra-

cum smooth and yellow; post-basal mantle margin with ribbon-like flap.

Localities : 14, 15.

Remarks : Lampsilis cariosa is widely distributed along the eastern sea-

board from Nova Scotia to Georgia (Johnson 1970, Burch 1975), but

only five North Carolina records are cited — one each in the Tar and

Cape Fear rivers, and three in the Neuse River. In the Tar, Clarke

(1983) found living individuals and empty valves at five and six stations,

respectively, from Franklin County in the Piedmont Plateau to Edge-

combe County in the Coastal Plain. The present study records L. cari-

osa from both the Haw and Deep river sub-basins, and as in the Tar, L.

cariosa occurs in both physiographic provinces in the Cape Fear drain-

age.

Tribe Anodontini

Andonta cataract a Say 1817 [= A. hallenbeckii Lea and A. doliaris Lea

(Dawley 1965); and A. ( Pyganodon ) cataracta cataracta Say

(Johnson 1970, 1984)]— Fig. 11-14.

Diagnosis’. Valves moderately thick and heavy, without lateral or pseu-

docardinal teeth, umbos extending beyond hinge line.

Localities : 1, 2, 3, 4, 5, 6, 11, 13, 14, 17, 19, 26, C, M.

Remarks: Anondonta cataracta, the most common anondontine mussel

in North Carolina, is a variable species sometimes reaching large size,

especially in impoundments. Figures 11 through 14 show four forms

encountered in the Cape Fear basin. The species occurs from the St.

Lawrence to the Alabama-Coosa river systems (Johnson 1970, Burch

1975). In North Carolina, Johnson (1970, 1984) reported specific locali-

ties in the Roanoke, Neuse, Cape Fear, Waccamaw, Yadkin, and

Catawba drainages. Although A. cataracta is not known from the Tar

watershed and was not encountered by Clarke (1983), it should be

expected there and in any North Carolina drainage in headwater

streams, farm ponds, and small impoundments. Both Walter (1956) and

Johnson (1970) cite only one collection from the Piedmont portion of

the Neuse basin, but A. cataracta was common in the adjacent part of

the Cape Fear. As explained by Shelley 1983, it was erroneously

reported from the Rocky River as A. implicata Say by Fuller (1977).

Anodonta imbecilis Say 1829 — Fig. 15-16.

Diagnosis’. Valves relatively thin and fragile, without lateral or pseudo-

cardinal teeth, umbos flat, not protruding above hinge line.

Localities’. 26, N.

Remarks: Anodonta imbecilis occurs sporadically from the St. Law-

rence to the Rio Grande river systems (Burch 1975). Walter (1956)

Unionid Mollusks

79

11

Fig. 11-16. Anodontini (Unioninae), localities in parentheses. 11-14, Anodonta

cataracta: 11 (17), 12 (14), 13 (1), 14 (6). 15-16, A. imbecilis (both 26). Scale line

= 10 cm.

reported two localities in the upper Neuse, but Johnson (1970) did not

include any from this basin among his three North Carolina records —

one each from the Yadkin, Catawba, and Cape Fear watersheds. In the

Tar, Clarke (1983) encountered living specimens and empty valves at

two and three stations, respectively, and found a thriving population in

the impoundment upstream from Rocky Mount, Nash County. Ano-

donta imbecilis may be confused with A. couperiana, which also has

flattened umbos and is known from Greenfield Lake (site N), but the

former has a straight ventral margin and the latter a broadly curved

profile.

80 Rowland M. Shelley

Strophitus undulatus (Say 1817) [= 5. rugosus (Swainson) (Dawley 1965)]

—Fig. 17-18.

Diagnosis'. Valves without lateral teeth, pseudocardinal teeth vestigial.

Localities : 14, 17, 19, 21, 26.

Remarks'. Widespread in small creeks in both the Mississippi and Atlan-

tic drainages from Canada to Texas, S. undulatus is known only from

the Tar, Neuse, and Cape Fear systems in North Carolina. Walter

(1956) found it at four localities in the Piedmont portion of the Neuse,

and these were later listed by Johnson (1970). In the Tar (Clarke 1983),

S. undulatus was found alive at five stations, and empty valves were

encountered at ten. The species is prevalent in the lower reaches of the

Haw and Deep sub-basins and has not been collected outside of Chat-

ham County.

Alasmidonta varicosa (Lamarck 1819) [= A. marginata varicosa

(Lamarck) (Dawley 1965)] — Fig. 19-20.

Diagnosis'. Valves without lateral teeth, pseudocardinal teeth prominent;

posterior slope with radial wrinkles.

Localities'. 14, 19, 24, 25, 26.

Remarks'. Alasmidonta varicosa ranges from the St. Lawrence to the

Savannah river systems. It is comparatively rare south of the Potomac

(Johnson 1970, Burch 1975, Clarke 1981) and was not encountered in

the Tar by Clarke (1983). The species was found at five sites in the lower

reaches of the Haw and Deep sub-basins and is also known from the

Uwharrie (Yadkin system) and the Catawba rivers in North Carolina. In

general A. varicosa is rare in Coastal Plain sections and is more com-

mon in Piedmont habitats and other areas above the Fall Zone.

Alasmidonta undulata (Say 1817) — Fig. 21.

Diagnosis’. Valves without lateral teeth, pseudocardinal teeth prominent;

posterior slope smooth; umbos high and rounded.

Localities: 11, 19, 25.

Remarks: Ranging from the St. Lawrence to the Chattahoochee river

systems, A. undulata is known from the Catawba, Yadkin, Cape Fear,

Neuse, and Tar systems in North Carolina (Walter 1956; Johnson 1970;

Clarke 1981, 1983). A fourth locality in the Cape Fear is the vicinity of

Fayetteville, where Fuller (1977) tentatively reported a juvenile of A.

triangulata (Lea), an apparent ecomorph that was placed in synonymy

under A. undulata by Clarke (1981).

Lasmigona subviridis (Conrad 1835) [= L. charlottensis (Lea) and L.

decor ata (Lea) (Dawley 1965)] — Fig. 22.

Diagnosis: Left valve with small interdental projection fitting into

groove in right valve.

Locality: 21, N.

Unionid Mollusks

81

Fig. 17-22. Anodontini (Unioninae), localities in parentheses. 17-18, Strophitus

undulatus: 17 (21), 18 (14). 19-20, Alasmidonta varicosu: 19 (14), 20 (24). 21, A.

undulata (25). 22, Lasmigonia subviridis (21). Scale line = 10 cm

Remarks'. This species is less common in the upper Cape Fear than in

the Tar system and Piedmont part of the Neuse system, where it was

taken at 13 and 18 sites, respectively (Walter 1956, Clarke 1983). In this

study only one valve was taken, in the headwaters of Bear Creek. John-

son (1970) reported localities in the Neuse, Tar, Yadkin, and Catawba

basins, along with one from the Cape Fear River at Kinnon, near Fay-

etteville. Clarke (1985), however, considered the Catawba population as

representing L. decorata (Lea) and did not record either species from

the Yadkin. Thus, L. subviridis is widespread on the Atlantic slope from

New York to the Cape Fear basin, and it is also known from the New

and Greenbrier rivers of the Kanawha drainage in North Carolina and

West Virginia (Johnson 1970, Burch 1975). It prefers small streams and

tributaries, and tends to avoid large rivers (Clarke 1985).

82

Rowland M. Shelley

Tribe Pleurobemini

Elliptio folliculata (Lea 1838) [= E. perlatus (Lea) and E. perstriatus

(Lea) (Dawley 1965); and E. arctata (Conrad) (Johnson 1970)] —

Fig. 23.

Diagnosis’. Valves more than three times as long as high, ventral margin

with variable indentation near midlength.

Localities’. 14, 16, L.

Remarks'. Johnson (1984) agreed with Davis et al. (1981) that Unio per-

latus, assigned to the arcuate unionids in the Cape Fear system, was

probably a synonym of E. folliculata instead of E. arctata, where it had

been placed by Johnson (1970). In the 1970 paper Johnson reported a

large series from the lower Cape Fear River, and the present material

extends the range of E. folliculata into the Piedmont Plateau.

Elliptio raveneli (Conrad 1834) [= E. confertus (Lea), E. livingstonesis

(Lea), E. micans (Lea), and E. tuomeyi (Lea) (Dawley) (1965); and

E. icterina (Conrad) (Johnson 1970)] — Fig. 24-25.

Diagnosis’. Posterior slope of valves angled, caudoventral corner slightly

pointed; mantle margin slightly pigmented.

Localities : 17, 19, 22, 24, G, M, Q.

Remarks’. Elliptio raveneli is widely distributed from the Escambia

River system in Florida to the White Oak system of North Carolina

(Johnson 1970, Burch 1975). Besides the Cape Fear, it is known from

the Broad, Catawba, Yadkin, and Waccamaw drainages of North Caro-

lina (Johnson 1970, 1984).

Elliptio complanata (Lightfoot 1786) [= E. complanatus roanokensis

(Lea) (Walter 1956, Dawley 1965); E. burkensis (Lea), E. catawben-

sis (Lea), E. complanatus jejunus (Lea), E. complanatus quadrilate-

rus (Lea), E. errans (Lea), E. insulsus (Lea), E. purus (Lea), E.

spadiceus (Lea), and E. subinflatus (Conrad) (Dawley 1965)] — Fig.

32-38.

Diagnosis’. Valves rounded posteriorly; mantle margin heavily pigmented.

Localities: 1, 2, 4, 5, 7, 8, 9, 11, 12, 14, 15, 16, 18, 19, 20, 24, 25, 26, A,

B, D, E, F, G, H, N, Q.

Remarks'. The most widely distributed and abundant unionid in eastern

North America and one of the most poorly understood systematically,

E. complanata ranges from northern Canada to Georgia (Johnson 1970,

Burch 1975). Specimens have been collected in the Broad, Catawba,

Yadkin, Waccamaw, Neuse, Tar-Pamlico, and Roanoke drainages in

addition to the Cape Fear (Johnson 1970, 1984). Walter (1956) encoun-

tered this species at 46 localities in the upper Neuse basin, six of which

were medium to large rivers; in the Tar drainage, Clarke (1983) found

living specimens at 53 stations and empty valves at 40, mostly in the Tar

River itself. This species was also the only unionid Clarke (1983)

Unionid Mollusks

83

Fig. 23-31. Pleurobemini (Unioninae), localities in parentheses. 23, Elliptio

folliculata (16). 24-25, E. icterina (both 24). 26-31, Uniomerus tetralasmus: 26-28

(17), 29 (8), 30 (18), 31 (21). Scale line = 10 cm.

encountered in the Neuse River between Raleigh and Lenoir County, a

reduction in diversity that he attributed to pollution. As expected, E.

complanata is widespread in the upper Cape Fear, and was collected at

24 stations. Elliptio complanata is a highly variable species and subject

to extreme ecophenotypic variation, which has generated the large

number of synonyms. Seven forms are shown in Figures 32 through 38.

The species can be easily confused with other representatives of the gen-

era Elliptio and Uniomerus, and close attention should be paid to the

illustrations here, in Johnson (1970), and in Burch (1975) when making

determinations.

84

Rowland M. Shelley

Uniomerus tetralasmus (Say 1831) [= Elliptio obesus (Lea) and U.

obesus (Lea) (Dawley 1965); and U. obesus (Lea) (Johnson

1984)]— Fig. 26-31.

Diagnosis’. Periostracum coarse; mantle margin darkly pigmented.

Localities: 4, 6, 7, 8, 12, 15, 17, 18, 19, 21, 22, 23, 24, N.

Remarks : The second most abundant species in the area of study, U.

tetralasmus is widespread in the central and eastern United States. On

the Atlantic slope it ranges from the Chowan River system in Virginia

to the Altamaha in Georgia (Johnson 1970, Burch 1975), and Johnson

(1970), Burch (1975), and Johnson (1970, 1984) reported it from the

Neuse, Roanoke, Tar, Cape Fear, Waccamaw, Yadkin, and Catawba

basins of North Carolina. These views contrast with that of Clarke

(1983), who considers U. tetralasmus a poorly understood species that is

absent from the Roanoke, Tar, and Neuse systems.

LOWER CAPE FEAR SPECIES

Listed below are unionids that were not encountered in the upper

Cape Fear River basin but have been reported from sites in the Coastal

Plain.

Villosa vibex (Conrad 1834) [= V. modioliformis (Lea) (Dawley 1965)].

Localities : N and Sprunt’s Pond, precise location unknown (Johnson

1970).

Lampsilis radiata radiata (Gmelin 1791) [= L. conspicua (Lea) and L.

radiata conspicua (Lea) (Dawley 1965)].

Locality : N (Johnson 1970).

Remarks : Fuller (1977:166, Fig. 4) showed photographs of male and

female shells of what he called the “L. radiata complex” from Orton

Pond, Brunswick County, which empties into the Cape Fear estuary

below Wilmington.

Anodonta couperiana (Lea 1842).

Locality : N (Johnson 1970).

Remarks: Fuller (1977) considered A. couperiana to be “Of Special

Concern” to North Carolina. Its current existence at this site, Greenfield

Lake in Wilmington, New Hanover County, is questionable, as recent

attempts to collect it have failed (William Adams, pers. comm.).

Elliptio congaraea (Lea 1831) [= E. dorsatus (Lea), E. forbesiana (Lea),

E. sordidis (Lea), and E. strumosus (Lea), (Dawley 1965)].

Locality: Cape Fear River without further specification (Johnson 1970).

The White Oak system is its northern range limit.

Elliptio lanceolata (Lea 1828) [= E. productus (Conrad) (Walter 1956,

Dawley 1965); E. fisherianus (Lea) and E. viridulus (Lea) (Dawley

1965)].

Unionid Mollusks

85

Fig. 32-38. Elliptio complanata, localities in parentheses. 32 (14), 33 (26), 34

(15), 35 (18), 36 (7), 37 (4), 38 (12). Scale line = 10 cm.

Localities : M, and an unspecified site in Wilmington, New Hanover

County (Johnson 1970).

Remarks'. Published concepts of this species vary greatly, and a diversity

of forms are currently assigned to this binomial by different malacolo-

gists. Only a comprehensive taxonomic revision can clarify this

confusion.

Elliptio marsupiobesa Fuller 1972.

Localities : I, J, K.

Remarks'. The small size, wedge shape, and yellow periostracum distin-

guish the valves of E. marsupiobesa, which is named for its “obese”

marsupium (Fuller 1972). It is known definitely from only these three

86

Rowland M. Shelley

sites (Fuller 1972, 1977); however, Fuller (1977) mentioned a possible

specimen from an unspecified locality in the Northeast Cape Fear River.

He also listed E. marsupiobesa as “Endangered” within North Carolina

because of proposed impoundments above Fayetteville and pollution

and sedimentation in the river. As it is similar conchologically to E.

complanata and E. raveneli, many collections doubtlessly contain misi-

dentified specimens of this species. Elliptio marsupiobesa may be widely

distributed in the lower Cape Fear and perhaps in other coastal drain-

ages. “Undetermined” status is more appropriate than “Endangered”

until this matter is resolved.

DELETION

Alasmidonta heterodon (Lea 1830).

Remarks’. Dawley (1965) reported A. heterodon from an unspecified

lake in Guilford County, but this determination is suspect because the

unionid is strictly a riverine species. The specimen(s) were supposedly in

her collection, but they were not among the materials donated to the

State Museum of Natural Sciences. Thus without verification, this

report has to be considered erroneous. Considered an “Endangered”

species both within North Carolina and nationally because of the dis-

junct distribution and small population sizes (Fuller 1977), A. hete-

rodon has been authentically reported from the Neuse and Tar systems

(Walter 1956, Johnson 1970, Clarke 1981), the known southern distribu-

tional limit. However, it has not been collected in the Neuse since Wal-

ter’s study; in his survey of the Tar River, Clarke (1983) found no empty

valves and only one living individual, in the Piedmont section in Gran-

ville County. The Granville record is unusual as A. heterodon is primar-

ily a Coastal Plain species.

DISCUSSION

With this report, the known unionid faunas of the three major

drainages lying entirely within North Carolina — the Tar, Neuse, and

Cape Fear — have been discussed. Surveys in the Neuse and Cape Fear

were not quantitative; they concentrated on the upper or Piedmont sec-

tions, and included relatively small tributaries. Clarke’s (1983) survey in

the Tar was quantitative; it included both Piedmont and coastal sec-

tions, but was largely limited to the Tar River itself with little sampling

in tributaries. Although the three studies are not equivalent, they never-

theless reveal the native faunas, which can be compared and contrasted

with enhancement from other published records. Comparable sampling

may not be done for years, and investigations in such lesser-known sys-

tems as the New (Atlantic), White Oak, and North Carolina sections of

the Chowan, Lumber, Yadkin/ Pee Dee, Catawba, and Broad drainages

deserve equal priority.

Unionid faunas also change over time as human alterations and

Unionid Mollusks

87

pollution exert negative influences over the resident species. For exam-

ple, two impoundments, Falls Lake on the Neuse near Raleigh and B.

Everett Jordan Lake on the Haw, have been constructed since the most

recent samplings in their areas, and a dam creating Randleman Lake on

the Deep River in Randolph County may soon be built. Clarke (1983)

found a diverse fauna in the Little River tributary of the Neuse River in

Wake and Johnston counties, but except for E. complanata, the natural

molluscan fauna of the Neuse itself had been eliminated, apparently by

pollution, from Raleigh to Jenny Lind in Lenoir County, a distance of

more than 136 river km. Similarly, the coastal Trent River tributary

below Trenton was devoid of mollusks, again probably the result of

pollution. The situation in the Roanoke system was similar. Clarke

(1983) found a few empty valves of Anodonta implicata Say and Lamp-

silis ochracea (Say) in rapids between Lake Gaston dam and Weldon,

only a few kilometers downstream, but from there to the river mouth no

unionids were seen. In contrast, the coastal Cashie River tributary con-

tained living E. complanata and Ligumia nasuta (Say). Clarke attrib-

uted the depauperate Roanoke River fauna to siltation complicated by

pollution and fluctuating water releases from Lake Gaston dam. Thus,

in contrast to the adjacent Neuse and Roanoke Rivers, coastal reaches

of the Tar River still support a diverse unionid fauna, which includes

the nationally endangered endemic species Elliptio ( Canthyria ) stein-

stansana. Every effort should therefore be made to maintain current

water quality levels in the Tar River, for much of North Carolina’s

diverse coastal unionid fauna is in imminent danger of disappearing.

The faunas of the three drainages are compared in Table 1. The Tar

and Neuse have the same number of species, but the Cape Fear has 50%

more. Discounting the presence of U. tetralasmus, since malacologists

disagree on its concept, 10 species are common to all three basins. The

Tar shares or shared Anodonta imbecilis with the Cape Fear and Alas-

midonta heterodon with the Neuse, and the Neuse and Cape Fear share

Anodonta cataracta and perhaps Uniomerus tetralasmus. Lampsilis

ochracea and Elliptio steinstansana occur only in the Tar, and eight

species occur solely in the Cape Fear, four of which represent literature

records from the lower basin. In contrast, no species are recorded solely

from the Neuse, a possible reflection of its intermediate geographical

position. The greater fauna of the Cape Fear is partly the result of

southern elements (V. vibex, A. couperiana, E. icterina, and E. conga-

raea) that occur northward to this or the proximal White Oak drainage.

Investigations are now needed in the latter and such drainages as the

New (Atlantic) and Newport, where no one has looked for unionids.

Likewise, areas in many watersheds have been sampled incompletely

and/or insufficiently. Additional undescribed species may well occur in

the river basins of North Carolina, and resolution of the perplexing

taxonomic enigmas can come only with additional and intensive empha-

sis here and in other Southeastern States.

88

Rowland M. Shelley

ACKNOWLEDGMENTS.— I thank Richard I. Johnson, Museum

of Comparative Zoology, for assistance with the determinations, and

Hugh J. Porter, University of North Carolina Institute of Marine Scien-

ces, and Andrew Gerberich, National Museum of Natural History,

Smithsonian Institution, for comments on preliminary drafts of the

manuscript. The photographic plates were prepared with the assistance

of Daniel J. Lyons, North Carolina State Museum Exhibits Section.

LITERATURE CITED

Burch, John B. 1975. Freshwater Unionacean Clams (Mollusca: Pelecypoda)

of North America. Malacological Publications, Hamburg, Mich.

Clarke, Arthur H. 1981. The tribe Alasmidontini (Unionidae: Anodontinae),

Part I: Pegias, Alasmidonta, and Arcidens. Smithsonian Cont. Zool. No.

326.

. 1983. Status survey of the Tar River spiny mussel. Final Project

Report for the U.S. Fish and Wildlife Service, Asheville, N.C.

. 1985. The tribe Alasmidontini (Unionidae: Anodontinae), Part II:

Lasmigonia and Simpsonaias. Smith. Cont. Zool. No. 399.

Davis, George M., W. H. Heard, S. L. H. Fuller, and C. Hesterman. 1981.

Molecular genetics and speciation in Elliptio and its relationship to other

taxa of North American Unionidae (Bivalvia). Biol. J. Linn. Soc.,

15:131-150.

Dawley, Charlotte. 1965. Checklist of freshwater mollusks of North Carolina.

Sterkiana 19:35-39.

Fuller, Samuel L. H. 1971. A brief field guide to the fresh-water mussels

(Mollusca: Bivalvia: Unionacea) of the Savannah River System. Assoc.

Southeastern Biol. Bull. 18:137-146.

. 1972. Elliptio marsupiobesa, a new fresh-water mussel (Mollusca:

Bivalvia: Unionidae) from the Cape Fear River, North Carolina. Proc.

Acad. Nat. Sci. Phila. 124:1-10.

. 1973. Fusconaia masoni (Conrad 1834) (Bivalvia: Unionacea) in

the Atlantic drainage of the southeastern United States. Malacological

Rev. 6:105-117.

. 1977. Freshwater and terrestrial mollusks. Pages 143-194 in

Endangered and Threatened Plants and Animals of North Carolina, J. E.

Cooper, S. S. Robinson, and J. B. Funderburg, editors. N.C. State Mus.

Nat. Hist., Raleigh.

Horn, Karen J., and Hugh J. Porter. 1981. Correlations of shell shape of

Elliptio waccamawensis, Leptodea ochracea and Lampsilis sp. (Bivalvia,

Unionidae) with environmental factors in Lake Waccamaw, Columbus

County, North Carolina. Bull. Amer. Malacol. Union Inc. for 1981:1-4.

Johnson, Richard I. 1967. Carunculina pulla (Conrad), an overlooked Atlantic

drainage unionid. Nautilus 80:127-131.

. 1970. The systematics and zoogeography of the Unionidae (Mol-

lusca: Bivalvia) of the southern Atlantic slope region. Bull. Mus. Comp.

Zool. 140:263-449.

Unionid Mollusks

89

. 1984. A new mussel, Lampsilis ( Lampsilis ) fullerkati (Bivalvia:

Unionidae) from Lake Waccamaw, Columbus County, North Carolina,

with a list of the other unionid species of the Waccamaw River System.

Mus. Comp. Zool. Occ. Pap. Mollusks 4:305-319.

, and Arthur H. Clarke. 1983. A new spiny mussel, Elliptio ( Can -

thyria) steinstansana (Bivalvia: Unionidae) from the Tar River, North

Carolina. Mus. Comp. Zool. Occ. Pap. Mollusks 4:289-298.

Porter, Hugh J. 1985. Molluscan census and ecological interrelationships. Rare

and endangered fauna of Lake Waccamaw, North Carolina, watershed

system. Vols. 1 and 2. Final report, North Carolina endangered species

restoration, 1978-1981. N.C. Wildl. Res. Comm, and Inst. Marine Sci.,

Univ. North Carolina, Chapel Hill.

, and Karen J. Horn. 1980. Freshwater mussel glochidia from Lake

Waccamaw, Columbus County, North Carolina. Bull. Amer. Malacol.

Union Inc. for 1980:13-17.

, and 1983. Habitat distribution of sympatric popluations

of selected lampsiline species (Bivalvia: Unionoida) in the Waccamaw

drainage of eastern North and South Carolina. Bull. Amer. Malacol. Union

Inc. 1:61-68.

, and 1984. Freshwater Mollusca of upper Waccamaw

River, North and South Carolina. J. Elisha Mitchell Sci. Soc. 97:270.

Shelley, Rowland M. 1972. In defense of naiades. Wildlife in North Carolina

36:4-8, 26-27.

. 1983. Occurrence of the unionid mollusk Anodonta implicata Say

in North Carolina. Nautilus 97:145-146.

Walter, Waldemar M. 1956. Mollusks of the upper Neuse River Basin, North

Carolina. J. Elisha Mitchell Sci. Soc. 72:262-274.

Accepted 27 May 1986

90

FISHERMAN’S GUIDE

FISHES OF THE SOUTHEASTERN UNITED STATES

by

Charles S. Manooch, III

Illustrated by Duane Raver, Jr.

Remarkable for its breadth of coverage, this book details the hab-

its, range, and appearance of more than 250 species of fish, 150 of which

are illustrated in color. Each account also includes tips on catching the

fish and preparing it for the table.

Manooch is experienced in the field of Fisheries Management and

Technology, and Raver is nationally known for his paintings of wildlife.

“An excellent general reference book for scientific or non-scientific

audiences. ... It contains information not easily found in any other

source.” — Carter R. Gilbert , Curator of Fishes , Florida State Museum.

1984 376 pages Index Bibliography ISBN 0-917134-07-9

Price: $24.95, plus $1.25 for shipping. North Carolina residents add Ax/i% sales

tax. Please make checks payable in U. S. currency to NCDA Museum

Extension Fund.

Send to FISHERMAN’S GUIDE, N. C. State Museum of Natural

History, P. O. Box 27647, Raleigh, NC 27611.

Drainagewide Occurrence

of the Freshwater Jellyfish,

Craspedacusta sowerbyi Lankester 1880,

in the Tennessee River System

Bruce L. Yeager

Office of Natural Resources and Economic Development,

Division of Services and Field Operations,

Field Operations, Eastern Area,

Tennessee Valley Authority,

Norris, Tennessee 37828

ABSTRACT. — Historical records and 42 new collections at 13 sites

along 673 miles of the Tennessee River system are reviewed and pre-

sented for the freshwater jellyfish, Craspedacusta sowerbyi. The exten-

sive distribution pattern of C. sowerbyi in the Tennessee River system

represents one of the few examples in North America of watershed

colonization, rather than isolated occurrence of this organism.

Although some researchers (Hargitt 1919, Schmitt 1939, Lytle

1960) have discussed the general distribution of the freshwater jellyfish,

Craspedacusta sowerbyi Lankester 1880, in North America, almost all

reports are of single, isolated occurrences with no watershed or drain-

agewide observations. Nine records of the freshwater jellyfish in the

state of Tennessee have been published. Powers (1938) documented the

species’ presence in east Tennessee from a bloom on Andrew Jackson

Lake, a small private lake in Knox County. Isom and Sinclair (1962)

published five records from Center Hill and Old Hickory reservoirs on

the Cumberland River. One collection of medusae (Chadwick and

Houston 1953) was recorded from the most downstream portion of the

Tennessee River on Kentucky Reservoir. Pennington and Fletcher

(1980) collected C. sowerbyi in the Tennessee River system, from Ken-

tucky Reservoir in Tennessee and Guntersville Reservoir in Alabama.

Summarizing the relatively few records of freshwater medusae from the

southeastern United States, Lytle (1962) reported an observation from

Wilson Reservoir, another mainstream impoundment of the Tennessee

River in Alabama. The present report summarizes historical and new

records in the Tennessee River system and firmly establishes the drain-

agewide presence of the medusoid stage of Craspedacusta sowerbyi.

Brimleyana No. 13:91-98, July 1987

91

92

Bruce L. Yeager

METHODS

Except for two collections from tributary reservoirs and one from

Wilson Reservoir, medusae were taken in a 0.5-m-square beam net

towed obliquely through vertically integrated strata at approximately

1.0 m/s for 10 minutes (Graser 1977, Tuberville 1979). The method was

used as a standard ichthyoplankton sampling procedure throughout the

Tennessee River Valley.

Samples were preserved in 10% formalin. In the laboratory, speci-

mens of C. sowerbyi were sorted from the ichthyoplankton samples and

transferred to 5% formalin buffered to ph 7.0. Specimens were donated

to the Department of Zoology at the University of Tennessee.

RESULTS AND DISCUSSION

Incidental collections of medusae of freshwater jellyfish in ichthyo-

plankton samples taken between 1978 and 1985 in the vicinity of Ten-

nessee Valley Authority power projects yielded numerous additional

records in Mississippi, Alabama, and Tennessee, and expanded the

range of collections in the Tennessee River system (Table 1). Collection

localities for medusae reflect efforts directed to specific sampling sites

and do not necessarily define habitat for C. sowerbyi within the Tennes-

see River system.

Craspedacusta sowerbyi was present in seven mainstream impound-

ments of the Tennessee River and in at least two tributary impound-

ments, Norris and Douglas reservoirs. Historical records (Table 2) and

those reported here (Fig. 1) extend over 394 miles of the Tennessee

River, between river miles 136 and 530, and include all mainstream

impoundments except Fort Loudon Reservoir, which has not been

sampled with the types of gear likely to capture medusae. Collections

from tributary rivers (exclusive of the Cumberland River localities)

extend the records to include 673 river miles from the most downstream

site on the Tennessee River to the most upstream sites on the French

Broad and Clinch rivers.

Craspedacusta sowerbyi is probably an exotic species introduced

from the Orient (Kramp 1950) where it occurs in a system of standing

and running waters along 1500 miles of the Yangtze River basin in

China. First found in the United States in 1908, the medusae have been

reported here only sporadically. Hydroids are associated with lotic, or

running water, habitats (Hutchinson 1967), whereas medusae are most

often found in lentic or standing waters, particularly artificially con-

structed impoundments (Lytle 1960). The system of alternating riverine

habitats and impoundments available in the Tennessee River system is

conducive to the establishment or formation of both the hydroid and

medusoid stages.

Freshwater Jellyfish Occurrence

93

Evidence of this animal’s successful spread throughout other drain-

ages in North America is limited. The only watersheds in North Amer-

ica where reports indicate a widespread distribution of the medusae are

the Kentucky River in Kentucky (Garman 1916, 1922, 1924; Payne

1925, 1926) and in the Potomac and James rivers in Virginia, Maryland,

and the District of Columbia (Lytle 1960). The distribution pattern of

C. sowerbyi in the Tennessee River system resembles the pattern in

these North American rivers, as well as that of the Yangtze River in

Asia, and represents one of the few examples in North America of

watershed colonization, rather than isolated occurrence, of C. sowerbyi.

LITERATURE CITED

Chadwick, C. S., and H. Houston. 1953. A “bloom” of freshwater medusae

Craspedacusta ryderi (Potts) in Kentucky Lake, Tennessee. J. Tenn. Acad.

Sci. 28:36-37.

Garman, H. 1916. The sudden appearance of great numbers of freshwater

medusae in a Kentucky creek. Science 44(1 146):858-860.

. 1922. Freshwater coelenterata in Kentucky. Science 56(1458):664.

. 1924. The freshwater jellyfish ( Craspedacusta sowerbyi) in Ken-

tucky again. Science 60:477-478.

Graser, L. F. 1977. Selectivity of larval fish gear and some new techniques for

entrainment and open water larval fish sampling. Page, 56-71 in Proceed-

ings of First Symposium on Freshwater Larval Fish, 1977, L. L. Olmsted,

editor. Duke Power Company, Huntersville, N.C.

Hargitt, C. W. 1919. Distribution of the freshwater medusa, Craspedacusta, in

the United States. Science 50:413-415.

Hutchinson, G. E. 1967. A treatise on limnology, introduction to lake biology

and the limnoplankton. Vol. 2. John Wiley and Sons, Inc., New York.

Isom, B. G., and R. M. Sinclair. 1962. Freshwater coelenterates in Tennessee,

New distribution records for: I. Craspedacusta sowerbyi Lankester 1880. II.

Cordylophora lacustris Allman 1871. Publ. No. 9 Tenn. Stream Pollution

Control Board.

Kramp, P. L. 1950. Freshwater medusae in China. Proc. Zool. Soc. Lond. 120

(Part I): 165-1 84.

Lytle, C. F. 1960. A note on the distribution patterns in Crapedacusta. Trans.

Am. Microsc. Soc. 79:461-469.

. 1962. Craspedacusta in the southeastern United States. Tulane

Stud. Zool. 95:309-314.

Payne, F. 1925. The hydroid of Craspedacusta ryderi in Kentucky. Science

62( 161 0):42 1 .

. 1926. Further studies on the life history of Craspedacusta ryderi, a

freshwater hydromedusan. Biol. Bull. 50(6):433-443.

Pennington, W. L., and J. W. Fletcher. 1980. Two additional records of Cras-

pedacusta sowerbyi Lankester in the Tennessee river system. J. Tenn. Acad.

Sci. 55:31-34.

Table 1. Collections of the freshwater jellyfish, Craspedacusta sowerbyi, from the Tennessee River system between 1973 and 1985.

94

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Table 2. Historical collections of freshwater jellyfish, Craspedacusta sower by i,

from the Tennessee-Cumberland River system.

KENTUCKY

Freshwater Jellyfish Occurrence

97

Historical and present (1973-1985) collection localities for Craspedacusta sower by i

in the Tennessee and Cumberland River systems.

98

Bruce L. Yeager

Powers, E. B. 1938. Freshwater medusae in Tennessee. Science 88:498-499.

Schmitt, W. L. 1939. Freshwater jellyfish records since 1932. Am. Nat.

73:83-89.

Tuberville, J. D. 1979. Vertical distribution of ichthyoplankton in upper Nick-

ajack Reservoir, Tennessee, with a comparison of three sampling method-

ologies. Pages 185-203 in Proceedings of Third Symposium on Larval Fish,

1979, R. D. Hoyt, editor. Western Kentucky Univ., Bowling Green.

Accepted 29 May 1986

Results of Fish Surveys

in the Tennessee River Drainage, 1979-1981

Joe C. Feeman, Jr.

Office of Natural Resources,

Tennessee Valley Authority,

Norris, Tennessee 37828

ABSTRACT. — From 1979 through 1981 the Tennessee Valley

Authority (TV A) conducted extensive fish surveys on nine streams in

the Tennessee River drainage. A total of 122 species of fish, represent-

ing 18 families, were identified from more than 107,000 specimens. One

hundred eighteen samples were taken at 32 localities. Findings include:

new occurrences for Notropis heterolepis, Noturus elegans, Etheo-

stoma cinereum, and Ammocrypta clara\ range extensions for Hybop-

sis cahni, Pimephales vigilax, Ammocrypta clara, and Etheostoma tip-

pecanoe\ numerous species with both state and federal protected

status; and relative abundance estimates for selected species.

In 1978 the Tennessee Valley Authority (TV A) was denied a U.S.

Army Corps of Engineers 404 permit for the Columbia Dam Project.

The permit was denied because of U.S. Fish and Wildlife Service con-

cern about the probable impact on the endangered birdwing pearly

mussel, Conradilla caelata (Conrad). During subsequent negotiations,

TVA and the U.S.F.W.S. agreed upon a program to conserve this and

other Cumberlandian mollusk species by establishing new populations

in the Duck and other Tennessee valley streams. Success of the program

would then constitute a basis for permission for TVA to be issued the

permit.

The life cycle of a freshwater mussel includes a parasitic stage in

which the larval mussel (glochidium) has a fish host. When released

from the gravid mussel, the glochidia attach to the gill filaments or fins

of a passing fish and eventually embed themselves into the membrane.

Two to four weeks later the glochidia drop off to the substrate, where

they remain for the rest of their adult lives.

Some species of mussel have a single species of fish as their host

(Howard 1912, Surber 1912). Thus, a requirement for a suitable trans-

plant site for this mussel is the presence of its fish host species. Fish

populations in areas known to support, or to have supported, popula-

tions of the endangerd mussel were sampled to aid in identification of

the fish species most likely to serve as hosts.

Streams of the Tennessee River drainage have been popular collect-

ing sites since the beginning of American ichthyology. Their diverse

faunas have attracted countless collectors for more than 150 years.

Brimleyana No. 13:99-121, July 1987

99

100

Joe C. Feeman, Jr.

Fig. 1. Map of the Tennessee River drainage showing sampling stations on the nine study streams.

Tennessee River Fish Surveys

101

From 1979 to 1981 TV A conducted extensive fish surveys on nine

streams in the Tennessee River drainage. A total of 118 samples were

taken at 32 localities using various collecting techniques. This paper

reports the findings of these samples, including new occurrences, range

extensions, and relative abundance estimates for some species. Parts of

this report are based on literature searches and may not reflect all the

efforts by many other collectors whose data have not been published.

Previous studies and surveys have been reported on all of our nine

study streams except Copper Creek and the Paint Rock River. The pub-

lications and data considered in this report are: Duck River — Whitney

(1957), Anonymous (1975), Nieland (unpubl. data); Powell River —

Woolman (1892), Anonymous (1970), Masnik (1975); Clinch River —

Masnik (1975); Nolichucky River — Ward (1960); North Fork Holston

River — Cope (1868), Jordan (1889), Ross and Carico (1963), Feeman

(1980), Feeman (1986); Buffalo River — Anonymous (1973); Elk

River — Jandebeur (1972).

DESCRIPTION OF STUDY AREA

The general study area of this project was the Tennessee River

drainage (Fig. 1). Five of the study streams are in the Great Valley

physiographic region of southwest Virginia and east Tennessee: Clinch,

Powell, North Fork Holston, and Nolichucky rivers; Copper Creek. The

other four streams are in the Highland Rim physiographic region of

middle Tennessee and northern Alabama: Duck, Buffalo, Paint Rock,

and Elk rivers. Table 1 gives specific locality information with river

mile, county, state, and map coordinates.

METHODS AND MATERIALS

The sampling scheme of this study consisted of two stages. Da.ta

collected in the first stage were used to determine the host species; three

streams with known populations of Conradilla caelata (Duck, Clinch,

and Powell rivers) were sampled once in 1979 and three times in 1980

(Table 1). Data collected in the second stage were used to determine

which streams had suitable populations of the host species and could be

considered as transplant sites; nine streams were sampled three times in

1981 (Table 1).

Three collecting techniques were used: (1) seine-snorkeling, (2)

backpack electrofishing (day and night), and (3) seine hauling. Both

quantitative and qualitative data were obtained. Transects were run

over a known area, using seine-snorkel and backpack electrofishing

techniques to collect quantitative data. Six transects were taken in each

habitat type present (e.g., sand and gravel, rubble, vegetation) to get a

representative sample of the site. Relative abundance estimates were

calculated from these data.

Sites were also sampled at night for nocturnal species and others

difficult to collect in the day. Electrofishing transects were repeated in

102

Joe C. Feeman, Jr.

Table 1. Sampling station locality descriptions for sites in nine study streams in the

Tennessee River drainage.

Buffalo River

Tennessee River Fish Surveys 103

Table 1. Continued.

Mile 91.4

104

Joe C. Feeman, Jr.

Table 1. Continued.

Nolichucky River

Tennessee River Fish Surveys

105

Table 1. Continued.

Mile 117.9

Both sides of island 0.6 mile upstream of Fletcher Ford at

Fletcher Cliff; Lee County, Virginia; TVA map 161 SE, 36°

36' 16"N, 83° 17'13"W. Collection dates: 8 August 1979, 13 April

and 21 April 1980, 18 May 1980, 18 June 1980.

Mile 120.6

At old ford, 0.4 river mile above unnamed county road bridge

(county road intersects with county road 661, 0.3 mile northwest

of bridge); Lee County, Virginia; TVA map 161 SE, 36°37'16"N,

83°16'50"W. Collection dates: 29 June 1979, 23 April 1980, 16

May 1980, 16 June 1980.

the same areas. Qualitative collections were also taken with seines and

backpack electrofishing units.

The seine-snorkel technique (Hickman and Saylor 1984) involved

four snorkelers in a line, moving downstream into a stationary 6-m

seine with 5-mm bar-mesh. Snorkelers swam abreast of each other while

bumping and pushing 1.1 -m fiberglass bars along the bottom. Fish were

thus herded downstream into the seine.

To insure a standard sampling area (4.9 m x 15.2 m), the snorkel-

ers’ bars were attached by 16.6-cm lengths of chain. The transect length

was determined by using a 15.2-m anchored rope. At the start of each

transect the rope was anchored at the upstream end of the transect, and

the seine was set at the downstream end.

Relative abundance estimates were also calculated from transects

taken with backpack electrofishing gear. The method was similar to that

used for seine-snorkel transects.

Fish caught were identified to species, enumerated, and released. A

voucher specimen of each species, along with any specimens of uncer-

tain identity, was preserved in 10% formalin and returned to the labora-

tory for verification. After identification, specimens were stored in 70%

ethanol and deposited in the TVA reference collection or in one of the

following museums: Northeast Louisiana University Museum, Florida

State Museum, Ohio State University Museum of Zoology, U.S.

National Museum, University of Tennessee Museum, University of Ala-

bama Ichthyological Collection, or Roanoke College Museum.

Nomenclature used in this paper follows Robins et al. (1980).

RESULTS AND DISCUSSION

A total of 122 species of fish, representing 18 families, were identi-

fied from more than 107,000 specimens collected in the nine study

streams (Table 2). The results will be discussed for each river, with

comments on noteworthy species collected.

106

Joe C. Feeman, Jr.

Duck River

With 86 species (Table 2), the Duck River produced the greatest

species diversity found in the nine study streams. Fishes were taken dur-

ing 41 collections at 10 sites from RM 153.1 to RM 243.1. The most

productive site in the study was also on the Duck River, where 72 spe-

cies were found at RM 179.1. Several significant species were taken,

including one species listed as threatened and five listed as in need of

management in Tennessee.

One of the most important finds was the collection of relatively

large numbers of the Tippecanoe darter, Etheostoma tippecanoe Jordan

and Evermann, a species listed as in need of management in Tennessee

(Starnes and Etnier 1980). This species was not known from the Duck

River until 1975 when two specimens were collected at Anchor Mill

(RM 206.6; C. F. Saylor, unpubl. data). During our study, 1,640 speci-

mens were collected at 9 of the 10 Duck River sites; the exception was

the RM 243.1 site. Three sites yielded most of the specimens: RM 159.4

(395), RM 176.9 (411), and RM 179.1 (461). Numbers diminished

upstream from RM 179.1, with only 38 specimens taken at RM 186.8

and 18 at RM 202.2. Of the darter species in the Duck River samples,

only the redline darter, Etheostoma rufilineatum (Cope), and the banded

darter, Etheostoma zonale (Cope), were more abundant.

In July samples, large numbers of E. tippecanoe in spawning condi-

tion were taken in shallow (15-30 cm), swift shoals with sand and gravel

substrate. One 25-foot transect at RM 176.9 into a seine yielded more

than 100 specimens. Spawning males exhibited billiant yellow and blue

coloration and were free-milting, indicating peak spawning condition.

During earlier and later samples, both sexes were most often taken in

deeper water with less current.

The small size of E. tippecanoe (35 mm SL, maximum; Zorach

1969) is probably the main reason previous collectors failed to collect it

in the Duck River. A TVA survey in 1972 (Anonymous 1975) included

three samples in areas where E. tippecanoe should occur. Sampling by

Whitney (1957) also included two sites where E. tippecanoe was found

to be common in our study. Samples in both of these surveys were

taken using rotenone, and any specimens of this small darter could have

washed through the block net.

The blacknose shiner, Notropis heterolepis Eigenmann and Eigen-

mann, was collected for the first time in the Duck River system during

this study. This species is listed as in need of management in Tennessee

(Starnes and Etnier 1980). Three specimens were collected at RM 243.1

in June 1981. This is also the first record of this species from the Ten-

nessee River drainage. A northerly species, it was previously known

from Tennessee only in tributaries to the Cumberland River in Smith,

Sumner, and Wilson counties (Starnes and Etnier 1980). The collection

site on the Duck River (RM 243.1) is adjacent to the Cumberland River

system and represents the southernmost occurrence of N. heterolepis.

Tennessee River Fish Surveys

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Tennessee River Fish Surveys

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110

Joe C. Feeman, Jr.

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Tennessee River Fish Surveys

111

Results of our study indicate an abundant population of the cop-

percheek darter, Etheostoma aquali Williams and Etnier, in the Duck

River. Because of its restricted range, E. aquali is considered a threat-

ened species in Tennessee (Starnes and Etnier 1980). A total of 554 spec-

imens were collected from 1979 through 1981. This species, which is

endemic to the Duck and Buffalo rivers (Williams and Etnier 1978), was

common at six sites on the Duck River from RM 159.4 to RM 179.1.

Above this point, E. aquali was less abundant and was absent from

three samples at RM 243.1. The decrease in numbers upstream is prob-

ably caused by the tailwater influence from Normandy Dam. TVA col-

lected seven specimens (reported as spotted darter) at RM 245.0 in 1972

(Anonymous 1975), prior to completion of Normandy.

A single specimen of the striated darter, Etheostoma striatulum

Page and Braasch, was taken at RM 179.1. It is listed as in need of

management in Tennessee (Starnes and Etnier 1980). Page (1983) con-

siders this species “rare and limited in distribution to tributaries of the

Duck River.” Although it is mainly a small-stream species, there are

several records from the main Duck River (TVA, regional Natural Her-

itage Project, unpubl. data). The collection of a single specimen at RM

153.1 in 1980 (TVA, unpubl. data) represents the downstream limit in

the main Duck River for this species.

Two other species collected have the status of in need of manage-

ment in Tennessee: redband darter, Etheostoma luteovinctum Gilbert

and Swain, and slenderhead darter, Percina phoxocephala (Nelson).

Clinch River

A total of 63 species were collected from five sites on the Clinch

River (Table 2). The most significant find was the western sand darter,

Ammocrypta clara Jordan and Meek, which had not been found pre-

viously in the Clinch River. A single specimen was taken at each of two

sites in June 1980: Kyles Ford (RM 189.5), Hancock County, Tennes-

see, and Hwy. 645 bridge (RM 219.4), Scott County, Virginia. The lack

of previous collections of this species is probably the result of several

conditions: (1) only a small population exists in the Clinch River, (2) the

preferred habitat is scarce, and (3) the preferred habitat is in deep pools,

making collection difficult. Starnes et al. (1977) discussed the annual

population density fluctuations and their effect on the success of collect-

ing species that inhabit sand and gravel substrate, especially A. clara.

They further speculated that this species was a part of the Clinch fauna

(Starnes et al. 1977), which our study proved correct. Although A. clara

is not listed on the official Tennessee list, it is considered endangered by

the Tennessee Heritage Program (Starnes and Etnier 1980). Since our

collections produced the first specimens from Virginia, no status has yet

been given to A. clara.

The slender chub, Hybopsis cahni Hubbs and Crowe, was taken at

two sites (RM 183.7 and RM 189.5). May and June collections pro-

duced the largest number of specimens (15 and 52, respectively) of this

112

Joe C. Feeman, Jr.

species, which is listed as threatened by the U.S. Fish and Wildlife

Service. H. cahni was common in this reach of the Clinch River in moder-

ate current over sand and pea-sized gravel substrate, similar to that de-

scribed by Masnik (1974) and Burkhead and Jenkins (1982).

A single collection (two specimens) of the pygmy madtom, Noturus

stanauli Etnier and Jenkins, was taken in August 1979 at Brooks Island

(RM 183.7). This site is composed mainly of medium gravel substrate

and moderate flow, similar to the type locality described by Etnier and

Jenkins (1980). Two subsequent collecting trips in 1980 at this site pro-

duced no more specimens. Noturus stanauli , which is restricted to this

section of the upper Clinch River and to the lower Duck River, is con-

sidered threatened in Tennessee (Starnes and Etnier 1980).

The undescribed “duskytail” darter, Etheostoma ( Catonotus ) sp.,

had not been collected from the main Clinch River prior to our study.

We collected a single specimen at RM 211.1 in May 1980. This site is

only 0.5 mile below the mouth of Copper Creek (CRM 21 1.6), which is

known to have a stable population of the duskytail darter. Thus, the

Clinch River specimen was most likely a transient individual from

Copper Creek.

Masnik (1974) reported a total of 75 specimens of the Tippecanoe

darter, Etheostoma tippecanoe , from six localities in the Clinch River.

He regarded the apparent rarity of this species in the Clinch River as

possibly resulting from use of collecting gear (seines) that is ineffective

for small-size specimens (Masnik 1974). (This theory is also discussed in

a previous section on E. tippecanoe in the Duck River.) Our study in

the same general area of the Clinch River produced a total of 170 spec-

imens at four sites (RM 183.7 to RM 219.4). These results indicate that

E. tippecanoe is not rare in the Clinch River and that Masnik’s assump-

tion about the ineffectiveness of collecting gear for this species may be

correct.

The sampling site at RM 189.5 (Kyles Ford) yielded the most spec-

imens of E. tippecanoe , 125; RM 211.1, 183.7, and 219.4 produced 24,

12, and 9 specimens, respectively. A total of 115 specimens were taken

in June samples, whereas August, May, and July samples produced 29,

19, and 7 specimens, respectively. As discussed previously, E. tippeca-

noe was most abundant in July samples from the Duck River.

The only populations of E. tippecanoe in Virginia are in the Clinch

River and Copper Creek. Owing to its restricted range in Virginia, this

species is considered threatened in that state (Jenkins and Musick 1980).

Several other species taken from the Clinch River in our study have

special status. Species listed by Jenkins and Musick (1980) as being of

special concern in Virginia are: popeye shiner, Notropis ariommus

(Cope); steelcolor shiner, Notropis whipplei (Girard); river redhorse,

Moxostoma carinatum (Cope); bluebreast darter, Etheostoma camurum

(Cope); blueside darter, E. jessiae (Jordan and Brayton); blotchside log-

perch, Percina burtoni Fowler; and channel darter, P. copelandi (Jor-

dan). Tangerine darter, Percina aurantiaca (Cope), is listed as in need of

Tennessee River Fish Surveys

113

management in Tennessee (Starnes and Etnier 1980) and of special con-

cern in Virginia (Jenkins and Musick 1980).

Powell River

A total of 64 species were taken at six sites on the Powell River

(Table 2). The most significant find was the yellowfin madtom, Noturus

flavipinnis Taylor, listed by the U.S. Fish and Wildlife Service as a

threatened species. Our collection of a single specimen at Buchanan

Ford (RM 99.2) represents only the second occurrence for this rare spe-

cies in the Powell River. In 1968, TVA collected a single specimen at

McDowell Ford (RM 106.6; Anonymous 1970). Since our collection,

another specimen was collected in 1983 by P. Shute and several other

University of Tennessee graduate students, also at Buchanan Ford

(Shute 1984).

Our specimen was taken from the back of a small island, in a pool

(1.5 m) over silty, bedrock substrate. The collection was taken at night

using backpack electrofishing methods. During 1980 and 1981, seven

additional sampling trips at this site failed to produce any additional

specimens. A 1976 rotenone sample by TVA in the same area produced

no specimens. Also, four samples in 1979 and 1980 at RM 106.7 (the

1968 TVA collection site) yielded no specimens.

Collections of such small numbers of N. flavipinnis indicate a

limited population of this species in the Powell River. Shute (1984) and

Burkhead and Jenkins (1982) suggest that this species prefers smaller

streams. Shute (1984) indicates the Powell River specimens may repres-

ent “strays” from nearby tributary populations. Several tributaries

sampled with rotenone during two TVA studies, 1968 (Anonymous

1970) and 1976 (TVA, unpubl. data) yielded no specimens of this spe-

cies. But owing to the difficulty of collecting the yellowfin madtom,

these samples are inconclusive. These data indicate that at least a relict

population still exists in the Powell River or in one of its tributaries.

Ammocrypta was first reported from the Powell River by Wool-

man (1892) as Ammocrypta pellucidum (- Ammocrypta pellucida).

Woolman reported it as the “most abundant of the darters” from the

“Powell River 8 miles south of Cumberland Gap, Tennessee.” In 1976,

W. C. and L. B. Starnes collected 16 specimens of Ammocrypta clara at

US 25E bridge, which is near the Woolman site (Starnes et al. 1977).

The validity of the Woolman identification of A. pellucida (Putnam) is

questionable, but no specimens are extant (Williams 1975). Starnes et

al. (1977) speculate that Woolman’s specimens were A. clara , and that

the occurrence of both species is unlikely.

In July 1979, five specimens of A. clara were taken in our samples

at Buchanan Ford (RM 99.2). The following day, five more specimens

were collected at Brooks Bridge (RM 95.3). During the next two years,

specimens were taken at three additional study sites: RM 106.6, RM

117.4, and RM 120.6, with a total of 55 specimens taken in this study.

River miles 99.2 and 120.6 produced the most individuals, 27 and 14,

respectively. These two sites had numerous areas of sand and fine

114

Joe C. Feeman, Jr.

gravel, the preferred habitiat of A. clara (Williams 1975). Most individ-

uals were collected from pools (0. 7-2.0 m) with slight current.

The greatest number of specimens of A. clara taken in a single

collecting trip was 20 on 26 May 1981 at Buchanan Ford (RM 99.2).

These individuals were in spawning condition, as indicated by faint, iri-

descent blue coloration on their opercles. Most of these individuals were

taken in a shallow (30 cm), narrow chute with moderate current over

clean-swept gravel substrate with small pockets of sand, which suggests

that A. clara moves up on shallow riffles to spawn. The presence of such

a large number of individuals displaying coloration indicates that this

species was very near spawning at the time of collection. This is some-

what earlier than Williams (1975) found (July and early August) for the

breeding season of A. clara.

The results of our collections indicate a healthy population of A.

clara in the Powell River. Population size is apparently limited by avail-

ability of suitable habitat. Other areas of suitable habitat are present,

but are mainly in pools too deep for effective collection.

The slender chub, Hybopsis cahni, which is presently endemic to

the Clinch and Powell rivers (a single specimen was found in a preim-

poundment study of the Holston River, 1941; Etnier et al. 1979), had

not previously been collected in Virginia. We collected this species at

RM 117.4 (37 specimens) and RM 117.9 (5 specimens). Both of these

sites are in Lee County, Virginia. This species was also taken at three

lower sites: RM 95.3 (3), RM 99.2 (1 1), and RM 106.7 (9).

The bullhead minnow, Pimephales vigilax (Baird and Girard), also

had not been collected from the state of Virginia prior to this study. Six

specimens were collected from the Powell River, RM 1 17.3 (Lee County,

Virginia) in June 1979. In 1980, a single specimen was collected in the

Clinch River (RM 211.1). This represents the northernmost occurrence

of P. vigilax in the Tennessee River drainage (Lee et al. 1980). Later in

the study, five specimens were also collected at RM 6.2 in the North

Fork Holston River. The last two sites are in Scott County, Virginia.

Other species collected in the Powell River have special concern

status in Virginia, according to Jenkins and Musick (1980). They were:

paddlefish, Polyodon spathula (Walbaum); popeye shiner, Notropis ari-

ommus\ river redhorse, Moxostoma carinatum\ bluebreast darter,

Etheostoma camurum\ blueside darter, E. jessiae\ channel darter, Per -

cina copelandi ; and tangerine darter, P. aurantiaca. Tangerine darter is

also listed as in need of management in Tennessee (Starnes and Etnier

1980).

North Fork Elolston River

Forty-two species were taken from the two sites on the North Fork

Holston River (Table 2). The two sites were dissimilar in gradient, sub-

strate type, and average width. The upstream site was characterized by

moderately steep gradient and swift current, boulder-rubble substrate,

and an average width of approximately 30 m. The lower site had a less

steep gradient and reduced water velocity, a more diverse substrate with

115

Tennessee River Fish Surveys

large areas of sand and gravel, and an average width of approximately

55 m. Faunal compostion reflects these differences. The upstream site

(RM 85.3) produced 32 species, whereas the lower site (RM 6.2) had 38.

Fishes considered to be large-river species and absent from the upstream

site were: popeye shiner, Notropis ariommus\ bullhead minnow, Pime-

p hales vigilax\ mountain madtom, Noturus eleutherus Jordan; and gilt

darter, Percina evides (Jordan and Copeland). Although only three

specimens of the banded darter, Etheostoma zonale, were taken

upstream, 73 were collected at the lower site.

The most significant species collected in North Fork samples was

the spotfin chub, Hybopsis monacha (Cope). A single specimen was

taken over sand and gravel substrate at RM 6.2. This rare chub (listed

as threatened by the U.S. Fish and Wildlife Service) is known from this

site by several previous collections by Jenkins and Burkhead (1984) and

Feeman (1986).

Several other species taken in our samples are considered of special

concern in Virginia (Jenkins and Musick 1980): popeye shiner, Notropis

ariommus\ bluebreast darter, Etheostoma camurum ; and tangerine dar-

ter, Percina aurantiaca.

Copper Creek

Copper Creek is a medium-sized tributary to the upper Clinch

Rfver (CRM 211.6). Visual examination of this stream is misleading. It

appears to be a typical tributary of this region and would not be

expected to have an unusual fauna. Yet, compared to other Clinch trib-

utaries this stream has a much more diverse fish fauna, including several

rare and sensitive species. A total of 43 species were identified from our

three samples at river mile 1.9 (Table 2).

The species of most significance collected in our samples was the

yellowfin madtom, Noturus flavipinnis, which is listed as threatened by

the U.S. Fish and Wildlife Service. Owing to the status of this fish,

Copper Creek is designated as critical habitat for N . flavipinnis (Federal

Register 1977). We collected three specimens in the May 1981 sample,

but two subsequent collecting trips in June and July failed to produce

additional specimens. The three specimens collected were taken along

the bank, in pool and backwater areas similar to habitat described for

this species by Jenkins (1975) and Taylor et al. (1971).

Recent collections by Burkhead and Jenkins (1982) have indicated

a decline in the Copper Creek population of this species. It was taken in

numerous collections from 1969 through 1972, but recently has been

difficult to collect (Burkhead and Jenkins 1982). The cause of this sus-

pected decline has not been determined.

Another sensitive species collected in our Copper Creek samples

was the undescribed “duskytail” darter, Etheostoma ( Catonotus ) sp.

Because of its restricted range and vulnerability to siltation, the dusky-

tail darter is listed as threatened in both Virginia and Tennessee (Jen-

kins and Musick 1980, Starnes and Etnier 1980). A total of 24 speci-

mens (13 in May, 9 in June, and 2 in July) were collected during the

116

Joe C. Feeman, Jr.

three samples in 1981, with night samples being more successful. The

present known range of this species includes Copper Creek, Citico

Creek, and Little River of the Tennessee River drainage, and the Big

South Fork of the Cumberland River drainage.

Several other species collected in Copper Creek are considered to

be of special concern in Virginia (Jenkins and Burkhead 1980): blueside

darter ( Etheostoma jessiae ), tangerine darter ( Percina aurantiaca), and

blotchside logperch ( P . burtoni).

Nolichucky River

The Nolichucky River site (RM 27.8) produced the lowest number

of species (38) of the nine study streams (Table 2). At this site the Noli-

chucky is a large river with a diversity of habitats. However, our three

samples were taken from the narrow channel behind Hale Island, with

the majority of flow directed toward the opposite side of the island. This

restriction of sampling area may account for the low number of species

collected.

The most significant species taken in the Nolichucky River was the

sharphead darter, Etheostoma acuticeps Bailey. This species was thought

to be extinct until 1972, when Jenkins and Burkhead collected three

specimens in the South Fork Holston River, just above South Holston

Reservoir (Jenkins and Burkhead 1975). Three years later, TVA col-

lected several specimens in the Nolichucky River at RM 18.0 (Saylor

and Etnier 1976). The sharphead darter has since been collected through-

out the Nolichucky and portions of its headwaters (Bryant et al. 1979,

Haxo and Neves 1984).

A total of 121 specimens of E. acuticeps were collected in our three

samples on the Nolichucky River. The July sample yielded 64 speci-

mens, many of which exhibited breeding condition. Males displayed

brilliant blue coloration and body forms that were more robust than

usual. This collection corresponds to the spawning period determined

by Bryant (1979). Specimens in May and June (39 and 18, respectively)

did not display spawning coloration.

Most individuals were taken in swift riffles with boulder and rubble

substrate, which agrees with findings of earlier workers (Bailey 1959,

Bryant 1979). The most successful areas of collection were the swiftest

(and most difficult to sample) riffles. Only a few specimens were taken

in other habitats with less current.

Paint Rock River

The fauna of this direct tributary to the Tennessee River was rela-

tively unknown until recent years. Although no major work has been

published on the fauna of this stream, Tom Jandebeur (pers. comm.) is

currently preparing a manuscript on the fishes of the Paint Rock River.

Our sample site (RM 60.0) was just below the confluence of the

Estill Fork and Hurricane Creek, which form the river proper (Fig. 1)

Average width at this site was approximately 20 to 25 m. Although the

Paint Rock is a small stream, the number and kinds of species collected

are indicative of a larger stream.

Tennessee River Fish Surveys

117

A total of 49 species were collected during three samples in 1981

(Table 2). The most significant species collected was the undescribed

“paleband” shiner, listed as threatened in Alabama (Ramsey 1984). A

rare species, it was previously known only from the Little South Fork of

the Cumberland River, Kentucky, and from a single preimpoundment

collection from Cove Creek, Tennessee (tributary to Norris Reservoir,

Clinch River drainage; Starnes and Etnier 1980). T. S. Janderbeur first

collected this little-known shiner in the Paint Rock River in November

1980. He collected specimens in six samples taken from RM 45.0 to RM

60.0. In our study, a total of 15 specimens were collected in two of the

three samples.

The elegant madtom, Noturus elegans Taylor, was collected for the

first time in the Paint Rock River during our study. Two specimens

were collected in the April 1981 sample. Owing to its limited range in

Alabama, N. elegans is listed as rare by Ramsey (1984). In recent years,

researchers have been able to describe the range of N. elegans in Ala-

bama more thoroughly. Its range has been reported recently by Ramsey

(1976), Lee et al. (1980), and Ramsey (1984). Earlier workers reported

no occurrences in Alabama (Smith-Vaniz 1968, Wall 1968), and Taylor

(1969) reported only one questionable record. Ramsey (1984) gives the

most recent distributional account for this species in Alabama as “one

stream locality each in Franklin, Jackson, and Limestone Counties.” He

does not, however, mention a 1969 TVA collection from Madison

County, Alabama (Anonymous 1971). All the known occurrences in

Alabama are from the Tennessee River drainage.

A collection of an American eel, Anguilla rostrata (LeSueur), at

this site is interesting. This fish has been found in the Tennessee River

proper and in its larger tributaries (Lee et al. 1980), but is generally

considered to inhabit only moderate to large streams (Pflieger 1975,

Trautman 1981). The site where the eel was collected is 60 miles above

the mouth of the Paint Rock, an unusual occurrence for this species.

Elk River

The one Elk River sample site (RM 91.4) is heavily influenced by

tailwater releases from Tims Ford Reservoir (RM 133.3). The water

fluctuates up to 1 m per day and is quite cold. Despite these adverse

conditions, 44 species were collected in three sampling trips in 1981

(Table 2).

The most significant find was the ashy darter, Etheostoma cine -

reum Storer, which is listed as in need of management in Tennessee

(Starnes and Etnier 1980). A single specimen, taken in April, represents

the first collection of this species from the Elk River. The discovery of a

potential new population of E. cinereum in the Elk River is reassuring,

for it appears to be declining in many other parts of its limited range

(Shepard and Burr ‘1984). The specimen was collected at night, in a pool

with moderate current, near a submerged log, by electrofishing into a

seine.

Another species taken in our samples and apparently rare in the

118

Joe C. Feeman, Jr.

Elk River is the popeye shiner, Notropis ariommus. Only a few previous

collections had found this species in the Elk River system, and our study

yielded only two specimens. Jandebeur (1972) reported 10 individuals

from two fifth-order streams in the Elk River system. The popeye shiner

is rather rare in most of its range (Starnes and Etnier 1980); however,

our results indicate that abundant populations still occur in portions of

the Duck, Buffalo, Powell, and North Fork Holston rivers.

Buffalo River

The Buffalo River, largest tributary of the Duck River, remains a

fairly constant size from the mouth up to our sampling site at RM 79.9.

This site has areas of shifting sand, gravel, and silt substrate, intermixed

with rubble, boulders, and bedrock. The flow is sluggish with several

short riffles. Three samples during 1981 produced 59 species (Table 2).

The most significant species collected in our samples on the Buffalo

River was the coppercheek darter, Etheostoma aquali, which is threat-

ened in Tennessee (Starnes and Etnier 1980). A total of 67 specimens of

this darter were collected with May and June samples producing all but

three of the total number collected. The large number of specimens

taken during the May-June period could indicate the beginning of the

spawning season, but little is known about the reproductive period of E.

aquali. The similar spotted darter, Etheostoma maculatum Kirtland,

spawns during this period on the underside of rocks in boulder riffle

areas (Starnes and Etnier 1980).

Another species that warrants mention is the blotchside logperch,

Percina burtoni. A single specimen was taken in the June sample.

Although the range of this species is rather widespread in the Cumber-

land and Tennessee river drainages, it is not abundant at any locality

(Starnes and Etnier 1980). Most workers consider silt and overall water

quality degradation to be the limiting perturbations for this species

(Starnes and Etnier 1980, Jenkins and Musick 1980). Because of these

habitat requirements, P. burtoni is listed as in need of management in

Tennessee (Starnes and Etnier 1980).

In addition to our samples, P. Jolly caught a single specimen of the

ashy darter, Etheostoma cinereum, on a hook and line, using a worm as

bait. The fish was caught between day and night samples, just upstream

from the sampling site. As noted earlier, E. cinereum is listed as in need

of management in Tennessee (Starnes and Etnier 1980).

ACKNOWLEDGMENTS. — This study was conducted with person-

nel from TV A Field Operations, and the names of all those involved are

too numerous to mention here. Gary Hickman was responsible for the

fish collections and also provided manuscript review. Charles F. Saylor

was a crew leader throughout the study and also assisted in identifica-

tion of specimens in the laboratory.

I thank D. A. Etnier, of The University of Tennessee, for his assist-

ance in identification of difficult specimens and for his review of the

119

Tennessee River Fish Surveys

manuscript. Mike Aaron also provided valuable assistance in manu-

script preparation. And finally, 1 am eternally grateful to Michael

Keene, of The University of Tennessee, for his instruction and motiva-

tion in technical writing and for his review of this manuscript.

LITERATURE CITED

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aquatic habitat. Powell River drainage basin, 1968. TVA, Norris, Tenn.

. 1971. Tennessee valley streams: Their fish, bottom fauna, and

aquatic habitat. Flint River drainage basin, 1969. TVA, Norris, Tenn.

. 1973. Tennessee Valley streams: Their fish, bottom fauna, and

aquatic habitat. Buffalo River drainage basin, September-October, 1974.

TVA, Norris, Tenn.

. 1975. Tennessee valley streams: Their fish, bottom fauna, and

aquatic habitat. Duck River drainage basin, September-October, 1972.

TVA, Norris, Tenn.

Bailey, R. M. 1959. Etheostoma acuticeps, a new darter from the Tennessee

River system, with remarks on the subgenus Nothonotus. Occas. Pap.

Mus. Zool. Univ. Mich. 603:1-10.

Bryant, R. T. 1979. The life history and comparative ecology of the sharphead

darter, Etheostoma acuticeps. Tenn. Wildl. Res. Agen. Tech. Rept. 79-50.

, J. P. Beets, and M. G. Ryon. 1979. Rediscovery of the sharphead

darter, Etheostoma acuticeps, in North Carolina (Pisces: Percidae). Brim-

leyana. 2:137-140.

Burkhead, N. M., and R. E. Jenkins. 1982. Five year status review of the

slender chub, Hybopsis cahni, a threatened cyprinid fish of the upper Ten-

nessee drainage. Unpubl. report to U.S. Fish and Wildlife Service.

, and R. E. Jenkins. 1982. Five-year status review of the yellowfin

madtom, Noturus flavipinnis, a threatened ictalurid catfish of the Tennes-

see drainage. Unpubl. report to the U.S. Fish and Wildlife Service.

Cope, E. D. 1896. On the distribution of freshwater fishes in the Allegheny

region of southwestern Virginia. J. Acad. Nat. Sci. Phil., Ser. 2.6:207-247.

Etnier, D. A., W. C. Starnes, and B. H. Bauer. 1979. Whatever happened to

the silvery minnow ( Hybognathus nuchalis ) in the Tennessee River? Proc.

Southeastern Fishes Council 2:1-3.

Federal Register. 1977. Threatened and endangered plants and animals. Fed.

Reg. 42:2507-2515.

Feeman, J. C. 1980. A Quantitative Survey of Fish and Macroinvertebrates of

the Holston River Basin: August-September, 1973. Report No. WR(FO)40-

4-80.1. TVA, Norris, Tenn.

. 1986. Fishes of the North Fork Holston River system, Virginia

and Tennessee. Proc. Southeastern Fishes Council 4(4).

Haxo, W. H., and R. J. Neves. 1984. A status survey of the sharphead darter

( Etheostoma acuticeps ). Unpubl. report to U.S. Fish and Wildlife Service,

Asheville, N.C.

120

Joe C. Feeman, Jr.

Hickman, G. D., and C. F. Saylor. 1984. Snorkeling and scuba diving. Pages

(5-l)-(5-8) in Warmwater Streams Techniques Manual, C. F. Bryan, editor.

Warmwater Streams Com., Southeastern Div., Am. Fish. Soc.

Howard, A. D. 1912. Experiments in Propagation of Freshwater Mussels of

the Quadrula group. U.S. Bur. Fish. Doc. 801.

Jandebeur, T. S. 1972. A study of the fishes of the Elk River drainage system

in Tennessee and Alabama. Unpubl. M.S. thesis, Univ. Ala., Tuscaloosa.

Jenkins, R. E. 1975. Status of the yellowfin madtom, Noturus flavipinnis.

Unpubl. report to U.S. Off. Endang. Spec. Internat. Activities, Washington.

, and N. M. Burkhead. 1975. Recent capture and analysis of the

sharphead darter, Etheostoma acuticeps, an endangered percid fish of the

upper Tennessee River drainage. Copeia 1975:731-740.

, and 1984. Description, biology, and distribution of the

spotfin chub, Hybopsis monacha, a threatened cyprinid fish of the Ten-

nessee River drainage. Bull. Ala. Mus. Nat. His. 8:1-30.

, and J. A. Musick. 1980. Freshwater and marine fishes. Pages

319-373 in Endangered and Threatened Plants and Animals of Virginia, D.

W. Linzey, editor. Va. Polytech. Inst. St. Univ. Blacksburg.

Jordan, D. S. 1889. Report of explorations made during the summer and

autumn of 1888, in the Allegheny region of Virginia, North Carolina, and

Tennessee, and in western Indiana, with an account of the fishes found in

each of the river basins of those areas. Bull. U.S. Fish. Comm. 8:97-173.

Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J.

R. Stauffer, Jr. 1980. Atlas of North American Freshwater Fishes. N.C.

Biol. Surv., N.C. State Mus. Nat. Hist., Raleigh.

Masnik, M. T. 1974. Composition, longitudinal distribution, and zoogeo-

graphy of the upper Clinch system in Tennessee and Virginia. Ph.D. dissert.

Va. Polytech. Inst. St. Univ., Blacksburg.

Page, L. M. 1983. Handbook of Darters. TFI Publications, Inc., Neptune City,

N.J.

Pflieger, W. L. 1975. The Fishes of Missouri. Missouri Dept. Consv.,

Columbia.

Ramsey, J. S. 1976. Freshwater fishes. Pages 53-65 in Endangered and

Threatened Plants and Animals of Alabama, H. T. Boschung, editor. Bull.

Ala. Mus. Nat. Hist. No. 2.

. 1984. Freshwater fishes. Pages 1-14 in Vertebrate Wildlife of

Alabama, G. Buchanan, dir. Ala. Agric. Expr. Sta., Auburn Univ.,

Auburn.

Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N.

Lea, and W. B. Scott. 1980. A list of common and scientific names of

fishes from the United States and Canada. Am. Fish. Soc. Spec. Publ.

12:1-174.

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ginia. Va. Agric. Expr. Sta. Tech. Bull. 161.

Tennessee River Fish Surveys

121

Saylor, C. F., and D. A. Etnier. 1976. Rediscovery of the sharphead darter,

Etheostoma acuticeps Bailey, in the lower Nolichucky River, Tennessee.

Bull. Assoc. Southeast. Biol. 23:93.

Shepard, T. E., and B. M. Burr. 1984. Systematics, status, and life history

aspects of the ashy darter, Etheostoma cinereum (Pisces: Percidae). Proc.

Biol. Soc. Wash. 97:693-715.

Shute, P. W. 1984. Ecology of the rare yellowfin madtom, Noturus flavipinnis

Taylor, in Citico Creek, Tennessee. Unpubl. M.S. thesis, Univ. Tenn., Knoxville.

Smith-Vaniz, W. F. 1968. Freshwater Fishes of Alabama. Auburn Univ. Expr.

Sta., Auburn.

Starnes, W. C., D. A. Etnier, L. B. Starnes, and N. H. Douglas. 1977. Zoo-

geographic implications of the rediscovery of the percid fish genus Ammo-

crypta in the Tennessee River drainage. Copeia 1977:783-786.

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tors. Tenn. Wildl. Res. Agency, Nashville.

Surber, T. 1912. Identification of the glochidia of freshwater mussels. Rept.

U.S. Comm. Fish, for 1912 and Spec. Pap.: 1-10. Separately issued as Bur.

Fish. Doc. No. 771.

Taylor, W. R. 1969. A Revision of the Catfish Genus Noturus Rafinesque with

an Analysis of Higher Groups in the Ictaluridae. Bull. U.S. Nat. Mus. 282.

, R. E. Jenkins, and E. A. Lachner. 1971. Rediscovery and de-

scription of the ictalurid catfish, Noturus flavipinnis. Proc. Biol. Soc.

Wash. 83:469-476.

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Columbus.

Wall, B. W. 1968. Studies on the fishes of the Bear Creek drainage of the

Tennessee River system. Unpubl. M.S. thesis, Univ. Ala., Tuscaloosa.

Ward, C. 1960. The fishes of the Nolichucky River, a contribution to a pollu-

tion survey. Special report, Tenn. Game and Fish Comm., Nashville.

Whitney, E. W. 1957. Final report, Duck River survey, a contribution to a

pollution survey. Tenn. Game and Fish Comm.

Williams, J. D. 1975. Systematics of the percid fishes of the subgenus Ammo-

crypt a, genus Ammocrypta, with description of two new species. Bull. Ala.

Mus. Nat. His. 1:1-56.

Woolman, A. J. 1892. Report of an examination of the fishes of Kentucky,

with lists of fishes obtained. Bull. U.S. Fish Comm. 10:249-288.

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nus Nothonotus (Pisces: Percidae). Am. Midi. Nat. 81:412-434.

Accepted 12 June 1986

122

ATLAS OF NORTH AMERICAN FRESHWATER FISHES

by

D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins,

D. E. McAllister, J. R. Stauffer, Jr., and many collaborators

This timely book provides accounts for all 777 species of fish

known to occur in fresh waters in the United States and Canada. Each

account gives a distribution map and illustration of the species, along

with information on systematics, distribution, habitat, abundance, size,

and general biology.

“. . . represents the most important contribution to freshwater

fishes of this continent since Jordan and Evermann’s ‘Fishes of North

and Middle America’ over 80 years ago.” — Southeastern Fishes Coun-

cil Proceedings.

1 980 825 pages Indexed Softbound ISBN 0-9 1 7 1 34-03-6

Price: $25, postpaid. North Carolina residents add 4!^% sales tax. Please make

checks payable in U. S. currency to NCDA Museum Extension Fund.

Send to FISH ATLAS, N. C. State Museum of Natural History,

P. O. Box 27647, Raleigh, NC 27611.

ATLAS OF NORTH AMERICAN FRESHWATER FISHES

1983 SUPPLEMENT

by

D. S. Lee, S. P. Platania, and G. H. Burgess

The 1983 supplement to the 1980 Atlas of North American Fresh-

water Fishes treats the freshwater ichthyofauna of the Greater Antilles.

In addition to this bound supplement, there are 19 accounts, mostly

species not described in 1980, in looseleaf form to be added to the 1980

volume. Illustrated by Renaldo Kuhler.

1983 67 pages Indexed Softbound

Price: $5, postpaid. North Carolina residents add 4*4% sales tax. Please make

checks payable in U. S. currency to NCDA Museum Extension Fund.

Send to FISH ATLAS, N. C. State Museum of Natural History,

P. O. Box 27647, Raleigh, NC 27611.

The Eastern Box Turtle,

Terrapene c. Carolina (Testudines: Emydidae),

in North Carolina1

Michael D. Stuart2 and Grover C. Miller

Department of Zoology,

North Carolina State University,

Raleigh, North Carolina 27695-7617

ABSTRACT. — One hundred and four eastern box turtles, Terrapene

Carolina Carolina , were collected in North Carolina, primarily from the

Piedmont. Baseline data were collected on weight, population sex and

age structure, seasonal distribution, reproduction, and food habits.

These data add to the information on the ecology and life history of

this common reptile in North Carolina.

The eastern box turtle, Terrapene Carolina (L.) (Testudines: Emy-

didae), is a terrestrial turtle found throughout the eastern half of the

United States. Four of the six recognized subspecies occur within the

United States with the subspecies T. c. Carolina reported to range from

southern Maine to Georgia and west to Tennessee and Michigan (Ernst

and Barbour 1972). The box turtle is widespread in North Carolina, but

little statewide information has been reported. As part of a dissertation

project on helminth distribution and life cycles, 104 eastern box turtles

were collected in North Carolina and surveyed for parasites. Data were

compiled on these hosts insofar as these factors might affect the type or

intensity of parasitism. This procedure revealed ecological information

on the life history of the box turtle in North Carolina. The number of

individuals removed from the wild was minimized by utilizing road-

killed specimens. Intact eggs were artificially incubated, and hatchlings

were released. We report here on sex and size distribution, seasonal

distribution, habitat, diet, reproduction, and egg incubation of T. c.

Carolina in North Carolina. The parasites of this host will be published

in a separate paper.

MATERIALS AND METHODS

Box turtles were collected from 10 counties in North Carolina

between June 1982 and December 1984. Collecting was done primarily

on the Piedmont Plateau, with small comparative samples taken from

1 Paper number 10214 of the Journal Series of the North Carolina Agricultural

Research Service, Raleigh, North Carolina 27695-7601.

2 Present address: Department of Biology, University of North Carolina at

Asheville, Asheville, North Carolina 28804.

Brimleyana No. 13:123-131, July 1987

123

124

Michael D. Stuart and Grover C. Miller

the Blue Ridge and Coastal Plain. Turtles were sexed or grouped as

juveniles if immature, weighed, and measured. Collection data, includ-

ing locality, habitat, and date, were recorded. Gut contents were identi-

fied during examination for parasites. Eighty-four specimens were col-

lected as road kills or while they were trying to cross highways. An

additional 20 were collected in their natural habitat, in part with the aid

of a border terrier that was trained by being encouraged to play with

captive turtles and then rewarded for finding hidden turtles.

RESULTS AND DISCUSSION

Adult box turtles may be sexed by a series of secondary sexual

characteristics. Males were identified by their slightly concave posterior

plastral plate, heavier and more strongly curved hindclaws, and longer

and thicker precloacal portion of the tail. The carapace of the males

tended to be flatter and lower than that of the females, but the ratio of

carapace length to plastron length was almost identical for both sexes

(Fig. 1 and 2). Most of the males had a light red color to the iris of the

eye while the females were usually brownish, but this characteristic was

not completely reliable.

Stickel (1950) classed specimens of T. c. Carolina from Maryland as

juveniles if they had a straight-line carapace length of less than 107 mm,

and Schwartz and Schwartz (1974) classed juvenile T. c. triunguis from

Missouri as having a carapace length of less than 110 mm. Auffenberg

and Iverson (1979) listed male size at sexual maturity as a carapace

length of 100 mm and females at 130 mm. The smallest sexually mature

individual taken in this study, as determined by condition of the gonads,

was a female with a carapace curve length of 124 mm and a plastron

length of 100 mm. Consequently, this plastron length was selected for

separating juveniles and adults.

Means and ranges for weight, plastron length, and curved carapace

length for males, females, and juveniles are in Table 1. Stickel (1950)

reported a male:female sex ratio of 1.00:1.09 (N = 245) on 42.6 acres in

Maryland with juveniles forming less than 10% of the population, and

Dolbeer (1969) gave a ratio 1.61:1.00 on 22 acres in Tennessee. Schwartz

and Schwartz gave a ratio of 55 males to 45 females (N = 698) in T. c.

triunguis on a 55-acre plot in Missouri; juveniles composed 18 to 25% of

this population. The total ratio of males to females to juveniles collected

in this North Carolina study was 37:43:24, which agrees most closely

with Stickel’s data except in the greater percentage of juveniles (Fig. 3).

Ernst and Barbour (1972) state that Terrapene Carolina is found

predominantly in open woodlands but may be found in pastures and

marshy meadows in the Northeast, and Martof et al. (1980) reported

Eastern Box Turtle

125

Carapace

Length

(mm)

■ m —

100 105 110 115 120 125 130 135 140 145 150

Plastron Length (mm)

Fig. 2. Shell measurements of adult female T. c. Carolina from North Carolina.

126

Michael D. Stuart and Grover C. Miller

this species in forested habitat throughout North Carolina, South Caro-

lina, and Virginia up to an elevation of about 1220 m. All specimens

collected in this study were adjacent to or in mixed hardwood or mixed

hardwood-pine forests.

During the hot, dry period from late June through early Sep-

tember, turtles were often found active immediately after a rainstorm or

very early in the morning. During these dry spells they were found rest-

ing in or quite near standing pools of water. It seemed that the seasonal

activity of the box turtle is temperature dependent. Ernst and Barbour

(1972) described T. Carolina as emerging from winter hibernation in

April and returning in October or November. Schwartz and Schwartz

(1974) stated that T. c. triunguis in Missouri usually emerged from hi-

bernation between the end of March and the end of April. This gener-

ally coincided with the last severe frost in the spring. Return to hiberna-

tion in mid-September to early November also coincided with the first

killing frost of the fall. The earliest date of collection in our study was

28 April with the latest date being 1 1 December (Fig. 4).

Box turtles are opportunistic omnivores. Principal foods are mush-

rooms, snails, and insects (Table 2). Of the 104 specimens examined in

our survey, 72 contained recognizable food items. The following were

identified, in order of percent of frequency of occurrence (Fig. 5): snails

59%, insects 43%, Armadillidium vulgare 40%, plants material (mainly

fungus) 32%, slugs 7%, rodents 5.5%, earthworms 3%, and millipeds

3%. The low level of earthworm consumption agrees with Klimstra and

Newsome (1960) in contrast to most generalized reports of feeding hab-

its. The vertebrate material in all these reports is almost certainly the

result of scavenging carrion, although Legler (1960) reported Terrapene

ornata capturing and killing small chicks.

Ernst and Barbour (1972) stated that juvenile box turtles were

mainly carnivorous but became more herbivorous with age. However,

no such difference in dietary preference was noted between juveniles and

adults or between adult males and females in the 72 North Carolina

specimens that contained recognizable food items. Both Stickel (1950)

and Klimstra and Newsome (1960) reported seasonality and availability

as important factors in determining the use of food items. The dietary

differences between juveniles and adults reported by Ernst and Barbour

(1972) may have been influenced by seasonality or availability of certain

food items.

Box turtles are reported to mate in the spring or occasionally in the

fall, laying two to eight eggs in each of two or three clutches between

late May and July (Smith 1961, Huheey and Stupka 1967, Mount 1975).

Of the 43 females examined in this study, 14 contained shelled eggs.

Eastern Box Turtle

127

Table 1. Weight and size measurements of North Carolina box turtles showing mean (x),

standard error (S.E.), number measured (N), and range of measurements (r).

23.1%

35.6%

41.3%

■ Adult Males

0 Adult Females

d Juveniles

Fig. 3. Population distribution by age/ sex class of T. c. Carolina trom North

Carolina.

128

Michael D. Stuart and Grover C. Miller

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Eastern Box Turtle

129

Month

Fig. 4. Seasonal activity patterns of T. c. Carolina from North Carolina

as measured by the number of specimens collected each month.

Two egg-bearing females were collected in May, five in June, and seven

in July. A total of 50 intact eggs were recovered. The number of eggs

per female ranged from two to seven, with a mode of three. The clutch

of seven eggs and another of three eggs were infertile. The remaining 40

eggs were incubated at room temperature in a tray of moistened ver-

miculite. The tray was enclosed in a plastic garbage bag to maintain a

high humidity. Thirty-six turtles hatched, with an average incubation

time of 78.8 days (range 77-80 days). These hatchings were released in

woodland areas where adults had been collected.

LITERATURE CITED

Auffenberg, W., and J. B. Iverson 1979. Demography of terrestrial turtles.

Pages 541-570 in Turtles, Perspectives and Research, M. Harless and H.

Morlock, editors. John Wiley & Sons, New York

Barbour, R. W. 1950. The reptiles of Big Black Mountain, Harlan County,

Kentucky. Copeia 1950(2): 100-107.

Bush, F. M. 1959. Foods of some Kentucky herptiles. Herpetologica 15:73-77.

130

Michael D. Stuart and Grover C. Miller

Food

Items

Plant Material

Earthworms

Millipeds

Scwbugs

Rodents

Insects

Snails

Slugs

59

0 10 20 30 40 50 60 70 80 90 100

Frequency (%)

Fig. 5. Food items in the stomachs of T. c. Carolina from North Carolina.

Dolbeer, R. A. 1969. A study of population density, seasonal movements and

weight changes of the Eastern Box Turtle ( Terrapene c. Carolina ) in eastern

Tennessee. M.S. thesis, Univ. Tennessee.

Ernst, C. H., and R. W. Barbour. 1972. Turtles of the United States. Univ.

Press Kentucky, Lexington.

Huheey, J. E., and A. Stupka. 1967. Amphibians and Reptiles of Great Smoky

Mountains National Park. Univ. Tennessee Press, Knoxville.

Klimstra, W. D., and F. Newsome. 1960. Some observations on the food

coactions of the common box turtle, Terrapene c. Carolina. Ecology

41:639-647.

Legler, J. M. 1960. Natural history of the ornate box turtle, Terrapene ornata

ornata Agassiz. Univ. Kansas Publ. Mus. Nat. Hist. 11:527-669.

Martof, B. S., W. M. Palmer, J. R. Bailey, and J. R. Harrison III. 1980.

Amphibians and Reptiles of the Carolinas and Virginia. Univ. North Caro-

lina Press, Chapel Hill.

Mount, R. H. 1975. The Reptiles and Amphibians of Alabama. Agri. Exper.

Sta., Auburn Univ.

Schwartz, C. W., and E. R. Schwartz. 1974. The three-toed box turtle in

central Missouri: its population, home range, and movements. Terr. Ser. 5,

Missouri Dep. Conserv.

Eastern Box Turtle

131

Smith, P. W. 1961. The amphibians and reptiles of Illinois. Illinois Nat. Hist.

Surv. Bull. 28:1-298.

Stickel, L. F. 1950. Population and home range relationships of the box turtle,

Terrapene c. Carolina (Linnaeus). Ecol. Monogr. 20:351-378.

Strang, C. A. 1983. Spatial and temporal activity patterns in two terrestrial

turtles. J. Herpetol. 17:43-47.

Surface, H. A. 1908. First report on the economic features of the turtles of

Pennsylvania. Zool. Bull. Div. Zool. Pennsylvania Dep. Agri. 6:105-196.

Accepted 23 June 1986

132

A DISTRIBUTIONAL SURVEY

OF NORTH CAROLINA MAMMALS

by

David S. Lee, John B. Funderburg, Jr., and Mary K. Clark

This book lists all the mammals of North Carolina and offers spe-

cies accounts and range maps for all of the non-marine species. Intro-

ductory chapters describe the plant communities of the state as they

relate to mammal distribution and discuss local zoogeographic patterns.

1982 72 pages Softbound

Price: $5, postpaid. North Carolina residents add sales tax. Please make

checks payable in U. S. currency to NCDA Museum Extension Fund.

Send to MAMMAL BOOK, N. C. State Museum of Natural History,

P. O. Box 27647, Raleigh, NC 27611.

Rediscovery of the Crystal Darter,

Ammocrypta asprella, in the Ohio River Basin

D. A. ClNCOTTA1 AND M. E. HOEFT2

Wildlife Resources Division,

West Virginia Department of Natural Resources,

Charleston, West Virginia

ABSTRACT. — The collection of Ammocrypta asprella is reported

from the Elk River, Kanawha River system, West Virginia. This is the

first record of the crystal darter from the Ohio River basin in 41 years

and the first for West Virginia. The status of the population is

unknown; repeated attempts to collect additional specimens failed.

Mining of natural gas and coal, along with urban and industrial

development, poses a threat to this population. Because West Virginia

has no state “endangered species” laws, the potential for protection

and management of this population is limited.

Ammocrypta asprella (Jordan), the crystal darter, is found in cer-

tain Gulf slope drainages and widely throughout the Mississippi River

basin of North America (Page 1983). It is the largest representative of

the genus Ammocrypta and is the only member of the subgenus Crystal -

laria. This darter occurs primarily in large streams and rivers, usually

over or buried in clean sand and gravel, and is most frequently taken in

waters where a constant current is maintained (Kuehne and Barbour

1983). Its distribution and abundance have been significantly reduced in

recent years owing to dam building and pollution (Kuehne and Barbour

1983, Page 1983). Because of this apparent jeopardy, Deacon et al.

(1979) regarded the species to be of Special Concern (i.e., potentially

threatened at the national level).

On 13 November 1980, we collected a single specimen (55.6 mm

SL) of A. asprella while boat electroshocking (240 volts AC) at night on

the Elk River of the lower Kanawha River system (ca. 1.6 km below

Mink Shoals or Elk Hills, Kanawha County, West Virginia). The spec-

imen was deposited in the West Virginia Department of Natural

Resources, Wildlife Resources Division Fish Museum at Elkins (catalog

no. WVWR 363). This account marks the rediscovery of the crystal dar-

ter in the Ohio River basin, where it is considered extirpated from all

states in which it was previously recorded (i.e., Tennessee, D. A. Etnier,

1 Present address: P.O. Box 67, Elkins, West Virginia 26241.

2 Present address: McClinic Wildlife Station, Point Pleasant, West Virginia 25550.

Brimleyana No. 13:133-136, July 1987

133

134

D. A. Cincotta and M. E. Hoeft

per. comm.; Illinois, Smith 1979; Indiana, Gerking 1945; Kentucky,

Burr 1980; Ohio, Trautman 1981). Apparently the last collection from

the basin was in 1939 from the Cumberland River drainage of Tennes-

see (UMMZ 125780, 125159; D. A. Etnier, pers. comm.). Its occurrence

in West Virginia represents an addition to the state ichthyofauna

(Denoncourt et. al. 1975, Cincotta et al. 1986).

The specimen was taken adjacent to a submerged log in a sandy

run at a depth of ca. 150 cm. The river width was about 30 m, and the

substrate was estimated to contain 25% rubble, 50% gravel, 20% sand,

and 5% silt. Water quality values recorded during this effort were pH

6.5, Fe 0.2mg/l, alkalinity 15mg/l as CaC03, hardness 8.0mg/ 1 as

CaC03, conductivity 87 micromhos /cm, and water temperature 25C.

Water clarity of the river was considered excellent (1.52 m Secchi dish

reading). Although Steele and McCoy (1980) indicated that the water

quality of the Elk River sub-basin is generally good, the area where the

specimen was collected is often reported for violations of state standards

for organic phenols and total coliform concentrations. Species collected

concurrently with the crystal darter were Notropis atherinoides, Hybop-

sis dissimilis, Hypentelium nigricans, Moxostoma anisurum, M. ery-

thrurum, M. macrolepidotum, Ictiobus sp., Ictalurus punctatus, Notu-

rus flavus, Pylodictis olivaris, Micropterus punctulatus, Lepomis

macrochirus, Etheostoma blennioides, E. zonale, Percina caprodes, P.

sciera, Stizostedion canadense, S. vitreum, and Aplodinotus grunniens.

The survey site was resampled by Hoeft at night in the same

manner and season in 1981 and 1983, but no additional specimens were

taken. It is interesting to note that the species was also not captured in

daytime surveys conducted by WVWR personnel in the subject area

during 1978, 1979, and 1984. The absence of A. asprella in past collec-

tions within the state is attributed to the lack of sampling in large rivers

under ideal conditions (low, clear flows) and its general rarity. The type

of collecting gear used and time of day sampling occurred may also

have contributed to its past omission, because electrofishing equipment

is not particularly efficient for collecting darters (pers. obs.) and this

species is apparently more active at night (J. D. Williams, pers. comm.).

The crystal darter was described in 1878 by D. S. Jordan from a

small, rocky tributary of the Mississippi River, Hancock County, Illi-

nois (Page 1980). No systematic review is available for this distinctive

percid, which is regarded by some investigators (e.g., Moore in Blair et

al. 1968) as constituting the monotypic genus Crystallaria. J. D. Wil-

liams (pers. comm.; in Smith 1979) believes that the species as presently

recognized may actually represent more than one subspecies or species.

The diagnostic characters of our specimen are within those reported by

Page (1983).

Crystal Darter in Ohio River Basin

135

Owing to the rarity of this species in the northeast, the lack of data

relative to its geographic variation, urban and industrial development in

the collection area, and extensive coal and natural gas exploration in

the lower Elk River sub-basin, the crystal darter was listed by the West

Virginia Department of Natural Resources as a species of Special Con-

cern, i.e., endangered or threatened at the state level (Cincotta, in

review). However, because West Virginia does not have a state “endan-

gered species” law, this designation will provide limited protection to

the Kanawha River population. This species is presently in Category II

(evaluation for potential listing) under the Endangered Species Act of

1973 (Fed. Reg. 1985). If the crystal darter is indeed rare nationally,

then Federal listing pursuant to the Act would probably be the only

legal mechanism available that would assist the protection and man-

agement of the population in West Virginia.

ACKNOWLEDGMENTS. — We thank Reeve M. Bailey, University

of Michigan, for confirming our identification, and providing UMMZ

collection records from the Ohio River basin. Bailey and R. E. Jenkins

critically reviewed this manuscript and made recommendations for its

improvement. D. A. Etnier, University of Tennessee, and J. D. Wil-

liams, U.S. Fish and Wildlife Service, graciously provided us with cer-

tain data. For field assistance we are indebted to C. Doerfer, Wildlife

Resources Division, and J. Fisher and D. Fisher, Water Resources Di-

vision. Special thanks are extended to K. Cash and B. Knapp for typing

the manuscript. All Wildlife Resources Division surveys cited herein

were funded under D-J Federal Aid Project F-10-R.

LITERATURE CITED

Blair, W. Frank, A. P. Blair, P. Brodkorb, F. R. Cagle, and G. A. Moore.

1968. Vertebrates of the United States. McGraw-Hill, New York.

Burr, Brooks M. 1980. A distributional checklist of the fishes of Kentucky.

Brimleyana 3:53-84.

Cincotta, Dan A. In Review. Fishes. In Endangered, Threatened or Special

Concern Vertebrates of West Virginia: 1986, K. B. Knight and L. B.

McArthur, editors. W.Va. Dep. Nat. Resour., Charleston.

Cincotta, Dan A., R. L. Miles, M. E. Hoeft, and G. E. Lewis. 1986. Discsovery

of Noturus eleutherus, Noturus stigmosus, and Percina peltata in West

Virginia, with discussions of other additions and records of fishes. Brim-

leyana 12:101-121.

Deacon, James E., G. Kobetich, J. D. Williams, S. Contreras, and committee.

1979. Fishes of North America endangered, threatened, or of special con-

cern: 1979. Fisheries 4(2):29-44.

136

D. A. Cincotta and M. E. Hoeft

Denoncourt, Robert F., E. C. Raney, C. H. Hocutt, and J. R. Stauffer, Jr.

1975. A checklist of the fishes of West Virginia. Va. Sci. 26(3): 1 17-120.

Federal Register. 1985. Endangered and threatened wildlife and plants: Review

of vertebrate wildlife. U.S. Dep. Interior, Fish Wildl. Serv. 50( 1 8 1):37598-

37967 (18 September).

Gerking, Shelby D. 1945. The distribution of the fishes of Indiana. Investiga-

tions of Indiana Lakes and Streams. 3:1-137.

Kuehne, Robert A., and R. W. Barbour 1983. The American Darters. Univ.

Kentucky Press, Lexington.

Page, Lawrence M. 1980. Ammocrypta asprella (Jordan), Crystal darter. Page

615 in Atlas of North American Freshwater Fishes, D. S. Lee et al., editors.

N.C. State Mus. Nat. Hist., Raleigh

. 1983. Handbook of Darters. T.F.H. Publications, Inc. Neptune

City, N.J.

Smith, Philip W. 1979. The Fishes of Illinois. Univ. Illinois Press, Urbana.

Trautman, Milton B. 1981. The Fishes of Ohio. Ohio State Univ. Press,

Columbus.

Steele, B. Douglas, and L. E. McCoy. 1980. Water Quality Status Assessment

1977-1979. W.Va. Dep. Nat. Resour., Div. Water Resour., Charleston.

Accepted 14 July 1986

Big-eared Bat, Plecotus townsendii ,

in Western North Carolina

Mary Kay Clark and David S. Lee

North Carolina State Museum of Natural Sciences,

P.O. Box 27647, Raleigh, North Carolina 27611

ABSTRACT. — This paper presents information on the occurrence

and biology of the Virginia big-eared bat, Plecotus townsendii

virginianus , the easternmost race of the species, in the mountains of

North Carolina. The four sites in Avery County represent a southward

range extension of 1 18 km.

Although Plecotus townsendii has the largest geographical

distribution of the three Recent Plecotus species occurring in North

America, its distribution in the eastern United States is restricted and

fragmented. Relict populations in the East appear to have been isolated

as a result of the influence of post-Pleistocene climates (Handley 1959,

Humphrey and Kunz 1976). Plecotus townsendii virginianus , the

easternmost race, has been designated endangered in federal listings

(Federal Register 44FR69208, 30 Nov. 1979). The species is a cave obli-

gate previously known from only a few scattered karst areas in eastern

Kentucky, eastern West Virginia, and extreme western Virginia. Here

we present information on the occurrence and biology of the Virginia

big-eared bat, Plecotus t. virginianus , in North Carolina, a range exten-

sion of 118 km south of the nearest reported colony in Burkes Garden

(elevation 975 m), Tazewell County, Virginia (Handley 1959). This col-

ony is still extant. Figure 1 depicts the distribution of this bat as it is

now understood.

In 1982 we examined a bat identified as Plecotus rafinesquii

donated to the North Carolina State Museum of Natural History

(NCSM 3578) from the teaching collection of Applachian State

University, Boone, N.C. This specimen actually represents Plecotus

townsendii virginianus . At our request Charles O. Handley, Jr., National

Museum of Natural History, examined the specimen and confirmed our

identification. Prior to this discovery it was assumed that all big-eared

bats in North Carolina were P. rafinesquii. We therefore decided to

re-examine Plecotus populations reported from adjacent areas. On 25

March 1984, with the assistance of Robert Currie, Office of Endangered

Species, U.S. Fish and Wildlife Service (Asheville Office), and Cato

Holler, Jr., from the Flittermouse Grotto of the National Speleological

Society, Old Fort, N.C., we visited a small cave (ca. 95 m in length) in

Avery County and found about 20 wintering Plecotus townsendii. On

the same day cavers informed us of a nearby cave that also housed

big-eared bats. Lee and Currie visited this cave in late September 1984

Brimleyana No. 13:137-140, July 1987

137

138

Mary Kay Clark and David S. Lee

and found several P. townsendii present at that time. Both caves are

close to the 1524-m elevation contour and in a transition zone between a

hardwood and a spruce-fir forest. Unlike most other caves inhabited by

Plecotus in the East, these caves are formed in Montezuma Schist

(Grandfather Mountain formation) rather than Greenbrier Limestone.

In North Carolina, Montezuma Schist is restricted to small areas in

Avery and adjacent counties, and this formation is the southermost site

of volcanic rocks of the late Precambarin in the Blue Ridge.

Subsequently we learned that Sturgis McKeever had collected two

Plecotus in a rock outcropping near the previously mentioned Avery

County caves on 22 September 1968. We examined the specimens,

which are in McKeever’s private collection, and found them also to be

P. t. virginianus . Thus, records of this bat have been obtained from four

North Carolina sites (three in extremely close proximity) over a period

of nearly 20 years and from both sides of the Eastern Continental

Divide.

Our visits to the Avery County sites in March, June, July, August,

and September and reports by cavers of big-eared bats in these same

caves in November and January indicate that the bats are present

throughout the year. Apparently they are few in number; we have never

encountered more than 20 on a single visit, although Currie reported to

us the presence of more than 60 bats in one cave in 1986. The Avery

County caves have multiple entrances that allow enough turnover of air

to produce the low temperatures necessary for hibernation. Because of

the high elevation of the caves, summer cave temperatures are appar-

ently too cool for summering bats or maternity colonies. We were

unable to locate bats of any species in these caves in July and August.

Mist-netting at the entrances, however, showed that modest numbers of

Plecotus were visiting the caves after dark during these months; so we

suspect there is at least one other Plecotus cave in the immediate vicin-

ity. All bats netted in summer were caught as they entered the cave. In

netting these bats we actually captured and handled only a few, but

perhaps as many as a dozen were seen on each of several nights during a

1-hour period. On 29 September 1984 a juvenile with an attendant

female was found in a group of 10 to 20 bats roosting in one Avery

County cave. Late in the afternoon of 11 July 1984, Clark accidently

startled a roosting Plecotus from a rhododendron, Rhodo-

dendron catawbiense, near a cave entrance. The bat flew directly into

the cave.

Plecotus began roosting in the high-elevation Avery County caves

as early as early September and remained in them until at least late

March. Within the caves the bats used several distinct roosting sites, one

in the twilight zone and others high in the back part of the cave; several

roosted in side passages. Roosting sites appear to shift with changing air

temperatures, and perhaps also with human disturbance. Main-passage

cave temperatures during our visits varied from 0 °C (March) to 13 °C

Big-eared Bat in North Carolina

139

Fig. 1. Specific locality records for Plecotus townsendii virginianus in West

Virginia, Virginia, Kentucky, and North Carolina.

(July-September). The only other bats found in the Avery County caves

inhabited by Plecotus were wintering Pipistrellus subflavus, a single

Myotis leibii (NCSM 4578) netted in the entrance to one cave in July

1984, and a single Myotis keenii netted in September 1986. Two

rodents, Neotoma floridana and Clethrionomys gapperi , were also

found in these caves.

The closest P. rafinesquii locality known to us is Taylorsville,

Alexander County, N.C., 66 km to the southeast. To date, P. townsendii

has been found locally only in a geographic area, forest type, and eleva-

tion zone not known to be occupied by P. rafinesquii , whereas in east-

ern Kentucky the two species are known to occur sympatrically (Rippy

and Harvey 1965). Summer mist-netting of streams near the Avery

County caves (but at an elevation 300 m below the forest type surround-

ing caves) yielded no Plecotus.

The locations of the sites at which bats were encountered are on file

at the North Carolina State Museum of Natural History and the Office

of Endangered Species, USFWS, Asheville.

140

Mary Kay Clark and David S. Lee

ACKNOWLEDGMENTS. — We thank Appalachian State University,

Boone, N.C., for donation of the specimen that alerted us to the pres-

ence of Plecotus townsendii in North Carolina. We also appreciate the

assistance of Charles O. Handley, Jr., National Museum of Natural His-

tory; Robert Currie, Office of Endangered Species, U.S. Fish and Wild-

life Service, Asheville, N.C.; Cato Holler, Jr., Flittermouse Grotto,

National Speleological Society, Old Fort, N.C.; and Sturgis McKeever,

who permitted us to examine specimens in his private collection. The

staff of the Tennessee Valley Authority was most cooperative in helping

us locate caves. This study was conducted under a special endangered

species permit (PRT-686498) provided to the N.C. State Museum by the

U.S. Fish and Wildlife Service. Since 1986 cooperative funding from the

N.C. Wildlife Resources Commission’s Nongame and Endangered

Wildlife Fund has provided assistance with ongoing field work.

LITERATURE CITED

Handley, Charles O. Jr. 1959. A revision of American bats of the genera

Euderma and Plecotus. Proc. U.S. Nat. Mus. 110:95-246.

Humphrey, Steven R., and Thomas H. Kunz. 1976. Ecology of a Pleistocene

relict, the western big-eared bat ( Plecotus townsendii ), in the southern

Great Plains. J. Mammal 57(3):470-494.

Rippy, Charles L., and Michael J. Harvey. 1965. Notes on Plecotus townsendii

virginianus in Kentucky. J. Mammal 46(3):499.

Accepted 4 December 1986

141

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Brimleyana No. 12 was mailed on 29 September 1986.

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BRIMLEYANA NO. 13, JULY 1987

CONTENTS

Life History of the Pinewoods Darter, Etheostoma mariae (Osteichthyes:

Percidae), a Fish Endemic to the Carolina Sandhills. Fred C. Rohde and

Steve W. Ross 1

Batriasymmodes from Caves in the Virginias (Coleoptera: Pselaphidae).

Thomas C. Barr, Jr 21

Zoogeography of the Freshwater Fish Fauna of Southern Georgia and Peninsular

Florida. Carter R. Gilbert 25

Evolution of the Intercalary Cartilage in Chorus Frogs, Genus Pseudacris

(Salientia: Hylidae). Gary L. Paukstis and Lauren E. Brown 55

Abundance and Distribution of Shrews in Western South Carolina. Michael T.

Mengak, David C. Guynn , Jr., J. Kenneth Edwards, Diane L. Sanders, and

Stanlee M. Miller 63

Unionid Mollusks from the Upper Cape Fear River Basin, North Carolina, with

a Comparison of the Faunas of the Neuse, Tar, and Cape Fear Drainages

(Bivalvia: Unionacea). Rowland M. Shelley 67

Drainagewide Occurrence of the Freshwater Jellyfish, Craspedacusta sowerbyi

Lankester 1880, in the Tennessee River System. Bruce L. Yeager 91

Results of Fish Surveys in the Tennessee River Drainage, 1979-1981. Joe C.

Feeman, Jr 99

The Eastern Box Turtle, Terrapene c. Carolina (Testudines: Emydidae), in North

Carolina. Michael D. Sturart and Grover C. Miller 123

Rediscovery of the Crystal Darter, Ammocrypta asprella, in the Ohio River

Basin. D. A. Cincotta and M. E. Hoeft 133

Big-eared Bat, Plecotus townsendii, in Western North Carolina. Mary Kay

Clark and David S. Lee 137

Miscellany 141