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North Carolina State Library
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N. r
1ST
B2
number 6 december 1981
EDITORIAL STAFF
John E. Cooper, Editor
Alexa C. Williams, Managing Editor
John B. Funderburg, Editor-in-Chief
Board
Alvin L. Braswell, Curator of David S. Lee, Chief Curator
Lower Vertebrates, N.C. of Birds and Mammals, N.C.
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(Invertebrates), N.C. of Lower Vertebrates, N.C.
State Museum State Museum
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of Botany, N.C. State Curator of Invertebrates, N.C.
University State Museum
Brimleyana, the Journal of the North Carolina State Museum of Natural His-
tory, will appear at irregular intervals in consecutively numbered issues. Contents
will emphasize zoology of the southeastern United States, especially North Caro-
lina and adjacent areas. Geographic coverage will be limited to Alabama, Dela-
ware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North
Carolina, South Carolina, Tennessee, Virginia, and West Virginia.
Subject matter will focus on taxonomy and systematics, ecology, zoogeog-
raphy, evolution, and behavior. Subdiscipline areas will include general inverte-
brate zoology, ichthyology, herpetology, ornithology, mammalogy, and paleon-
tology. Papers will stress the results of original empirical field studies, but
synthesizing reviews and papers of significant historical interest to southeastern
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In citations please use the full name — Brimleyana.
North Carolina State Museum of Natural History
North Carolina Department of Agriculture
James A. Graham, Commissioner
CODN BRIMD 7
ISSN 0193-4406
Fishes of the Waccamaw River Drainage
John R. Shute \ Peggy W. Shute l and David G. Lindquist
Department of Biology,
University of North Carolina at Wilmington,
Wilmington, North Carolina 28406
ABSTRACT.— From 1976 through February 1981, we made 827 col-
lections of fishes from 75 stations in Lake Waccamaw and the Wac-
camaw River and tributaries; they yielded a total of 56 species from 18
families. Additional records increased the probable total to 62 species.
At least five of the Waccamaw species are endemic or exclusively
shared with one other drainage. These aspects of the Waccamaw indi-
cate that it is unique among Atlantic Coastal Plain drainage systems.
Geological and zoogeographical evidence suggest that the Waccamaw
River once drained a larger area extending into the inner Coastal Plain
and Piedmont. Uplifting of the Cape Fear Fault resulted in piracy of
the Waccamaw headwaters, creating the present Cape Fear drainage.
Faunal resemblances between the drainages lend support to this theory.
INTRODUCTION
Since the description of the Waccamaw killifish, Fundulus
waccamensis, silverside, Menidia externa, and darter, Etheostoma per-
longum, by Hubbs and Raney (1946), Lake Waccamaw, North Carol-
ina, has been the subject of both biological and physiographical studies
(Frey 1948a,b, 1949, 1951) alluding to the relatively high level of fish
diversity and endemism. Louder (1962a) provided a checklist of fishes
and Hueske (1948) discussed fishery resources. Four species from the
lake have been subjects of biological studies: Notropis petersoni (Davis
and Louder 1971); F. waccamensis (Shute et al., ms.); M. extensa (Davis
and Louder 1969); and E. perlongum (Lindquist et al. 198 1 ; Shute et al., in
press).
Apart from those of the lake, the fishes of the Waccamaw drainage
have received little attention. Louder (1962b) included the Waccamaw
drainage in a survey of the Lumber and Shallotte River drainage in
North Carolina, and the major purpose of his survey was to evaluate the
recreational fishery potential. Fowler (1935) reported on several collec-
tions from the Waccamaw drainage in South Carolina.
The Waccamaw is unique among Atlantic coastal drainages. Its
overall fish diversity is quite high and includes a number of endemics
and forms shared with but one other drainage. Presently two undes-
cribed species of fish are known to occur within the Waccamaw drain-
1 Present address: Department of Zoology and Entomology, University of Ten-
nessee, Knoxville, TN 37916.
Direct reprint requests to DGL.
Brimleyana No. 6: 1-24. December 1981. 1
2 John R. Shute, Peggy W. Shute, David G. Lindquist
age. Seventeen species of plants and animals considered of special con-
cern by biologists were listed from in and around Lake Waccamaw by
Teulings and Cooper (1977). Parts of the Waccamaw River's upper
reaches have been proposed for inclusion in the National and Scenic
Rivers System (Anonymous 1978) because of their relatively undis-
turbed nature and the river's unique assemblage of flora and fauna.
Geological evidence suggests that the Waccamaw River once
drained a much larger area, extending into the inner Coastal Plain and
Piedmont. As discussed later, zoogeographical evidence supports this
theory, which may explain the high species diversity.
This study was intended to provide a baseline of information on the
overall distribution of fishes within the drainage, with special emphasis
on the endemic and undescribed forms, because habitat alteration could
present a definite problem for most of the unique fishes. There is limited
suitable habitat for the species with upland affinities, and impoundment
or channelization projects could prove disastrous.
STUDY AREA
The Waccamaw River drainage lies entirely within the low Coastal
Plain of North and South Carolina, draining an area of approximately
4000 km" (Fig. 1). It is a relatively young system, believed to have been
formed during the Late Pleistocene, 32,000 to 75,000 years ago (Zullo
and Harris 1979). Sediments consist of Pleistocene sands underlain by
the fossiliferous Waccamaw Formation, a limestone formation that is
exposed in some areas of the river. The Surry Scarp forms the western
border of the system, and to the north the Cape Fear Fault forms a
barrier separating the Cape Fear Basin from the Pee Dee Basin (Zullo
and Harris 1979).
Friar Swamp
Friar Swamp is the principal feeder system to Lake Waccamaw.
Originating from Council Mill Pond (approximately 15 km north-
northwest of Lake Waccamaw), this small stream flows south and con-
verges with Slap Swamp, Buckhead Branch, and Gum Swamp to form
Big Creek (Fig. 1). Big Creek is a typical black water stream with a
sandy, muck bottom and an abundance of aquatic vegetation along its
shoreline.
Lake Waccamaw
Lake Waccamaw is the largest of the Carolina Bays, with a total
area of 3618 ha. Most of its 22.9 km shoreline is characterized by sandy,
low-gradient beaches. Vast beds of maidencane, Panicum hemitomum,
extend offshore along the eastern, southern, and western shores of the
lake. Cape Fear spatterdock, Nuphar luteum sagitifollium, grows in
Waccamaw Drainage Fishes
PEE DEE BIVEP,
SCALE IN KM.
WACCAMAW RIVER DRAINAGE
WINY AH v
BAY
Fig. 1. Map of the Waccamaw River drainage, North and South Carolina,
showing fish sampling localities.
dense beds off the northern and northeastern shores. The bottom is
mainly sand and fibrous peat. Over the peat bottom, generally toward
the middle of the lake, thick stands of bushy-pondweed, Najas guadalu-
pensis, occur and a green alga, Nitella sp., is seasonally abundant.
Average depth of the lake is 2.3 m, and maximum depth is 3.3 m.
In addition to Big Creek, the lake is fed by three smaller streams:
Little, Second, and Third creeks. Acid water from these streams is neu-
tralized by the calcareous Waccamaw limestone formation, which
underlies the lake and is exposed along the north shore (Frey 1951).
4 John R. Shute, Peggy W. Shute, David G. Lindquist
Man-made canals surround much of Lake Waccamaw. They are
characterized by dense vegetation (mainly alligator-weed, Alternanthera
philoxeroides; duckweed, Lemna perpusilla; and beggar tick, Bidens
laevis) and steep, grassy roadside banks, and are open to the lake and
the Waccamaw River.
Waccamaw River
The Waccamaw River originates at the south shore of Lake Wac-
camaw and flows southward approximately 225 km to its confluence
with the Pee Dee River at Winyah Bay, South Carolina. The river is
sluggish and meandering with an average gradient of only 6.44 cm/ km.
Above Juniper Creek (Fig. 1), much of the river is less than 15 m wide
with the exception of an area known locally as "The Fishponds,"
located just below Bogue Swamp. Here, for a distance of several
hundred meters, the river widens to approximately 50 to 60 m. Below
Juniper Creek the river widens considerably, and before crossing the
North Carolina/ South Carolina state line averages around 75 m wide.
Seasonally, water levels in the river fluctuate considerably. High
water occurs in late winter and throughout the spring. During summer
and fall the waters recede, creating shallow, sandy stretches.
Bogue Swamp
Bogue Swamp is the first tributary to the Waccamaw River, origi-
nating approximately 6.5 km northwest of Lake Waccamaw and flow-
ing 13 km southeast before entering the river (Fig. 1). This is a small,
sand-bottom stream, often intermittent during dry months.
White Marsh
Brown Marsh, Elkton Marsh, and Red Hill Swamp flow from the
upper part of White Marsh, approximately 23 km northwest of Lake
Waccamaw. White Marsh is a sluggish, muck-bottom stream that flows
southeast for 33 km before entering the Waccamaw River. Stations
sampled during our survey were in Red Hill Swamp and the main
stream of White Marsh (Fig. 1). Aquatic vegetation was dense in the
areas collected, and station 15 was heavily obstructed with roots,
stumps and water-logged branches.
Juniper Creek
Juniper Creek is the largest tributary to the Waccamaw River in
North Carolina and is the first to enter the river from the east (Fig. 1).
It is formed by the confluence of Muddy Branch and Bear Pen Island,
Honey Island and Alligator swamps. Juniper Creek and its tributaries
form the major drainage system for Green Swamp. Many interconnec-
ting man-made canals east of Lake Waccamaw join Honey Island
Swamp with Dans Creek of the adjacent Cape Fear Drainage. These
Waccamaw Drainage Fishes 5
canals are normally 3 to 5 m wide and 1 to 2 m deep.
Upper Juniper Creek is generally narrow (6 to 8 m) and shallow,
and flows year-round. The channel widens to approximately 25 m
downstream and becomes sluggish. Before reaching the river it again
narrows and flow increases. Much of this stream is characterized by
sand bottom and patches of dense aquatic vegetation. The wide, slug-
gish areas are generally richer in organic debris, and sphagnum moss,
Sphagnum sp., grows in dense mats along the shoreline. Juniper Creek
flows west approximately 35 km to its junction with the Waccamaw
River.
Seven Creeks
The headwaters of Seven Creeks are formed by Toms Fork, Mill
Branch, and Juniper, Brissett, Gum, Beaver Dam, and Monie swamps.
This predominantly muck-bottom stream flows approximately 16 km
southeast until reaching the Waccamaw River 13 km above the North
Carolina/ South Carolina state line. Areas sampled were clogged with a
tangle of old tree stumps, roots, and waterlogged branches. Aquatic
vegetation was present but not dense.
Many smaller tributaries not discussed also flow into the Wac-
camaw River. During our survey, no South Carolina tributaries were
sampled. Kingston Lake Swamp forms the largest of the Waccamaw
River tributaries above the Atlantic Intracoastal Waterway (AIWW).
Several small streams and canals connect the Waccamaw and Pee Dee
rivers before their confluence at Winyah Bay.
METHODS
From January 1979 through February 1981 a total of 827 collec-
tions was made from 75 stations within the Waccamaw River drainage
(Table 1). Stations la through m, 2a,b,c, 3, 4, 5, 6, and 7 were sampled
monthly during this period. The remaining stations were sampled on an
irregular basis, sometimes only once. Six of the mid-lake stations in
Lake Waccamaw are treated as one throughout this paper (Station li).
Most collections were made with seines varying in size from 3 m X
1.2 m to 15.2 m X 1.8 m, typically with 3 mm mesh. Offshore stations in
Lake Waccamaw (and one Waccamaw River station) were sampled with
a small otter trawl measuring 2.8 m X 1.3 m at the mouth and lined
with 3 mm mesh. Dip nets of various sizes were also used. A representa-
tive sample of fishes was usually field preserved in 10% formalin and
later stored in 70% ethanol. Large or extremely common species were
occasionally returned to the water (some were photographed before
release) and records are based on field identifications. Museum speci-
mens and literature records were verified where possible.
Nomenclature follows that used by Robins et al. (1980). Most spec-
imens are housed in the University of North Carolina at Wilmington
Fish Collection (UNCW).
John R. Shute, Peggy W. Shute, David G. Lindquist
Table 1. Sampling stations. US = U.S. highway, CR = county road; NC = N.C.
highway.
NORTH CAROLINA
Columbus County
la. Lake Waccamaw, north shore public beach on NC 214.
lb. Lake Waccamaw, northeast shore at Hobbs Harbor on NC 214.
lc. Lake Waccamaw, northeast shore just west Big Creek.
Id. Lake Waccamaw, southeast shore at northern boundary state park.
le. Lake Waccamaw, south shore at southern boundary state park.
If. Lake Waccamaw, south shore just east dam.
lg. Lake Waccamaw, south shore just above dam.
lh. Lake Waccamaw, southwest shore.
li. Lake Waccamaw, mid-lake.
lj. Lake Waccamaw, north offshore transect.
Ik. Lake Waccamaw. north offshore transect.
1 1. Lake Waccamaw, northeast offshore transect.
lm. Lake Waccamaw, south offshore transect.
2a. Big Creek Wildlife Access Area on SR 1947.
2b. Big Cr. trib. at second bridge going east on SR 1947.
2c. Big Cr. at first bridge going east on SR 1947.
3. Waccamaw canal, northeast shore lake on SR 1947.
4. Waccamaw canal, north shore lake on NC 214.
5. Waccamaw canal, northeast shore lake at junction NC 214 and
SR 1957.
6. Waccamaw canal, southwest shore lake.
7. Waccamaw River just below dam on south shore lake.
8a. Waccamaw R. from below station 7 to approx. 1.6 river km below
lake.
8b. Waccamaw R. between 1.6 and 3.2 river km below lake.
8c. Waccamaw R. between 3.2 and 4.8 river km below lake.
8d. Waccamaw R. between 4.8 and 6.4 river km below lake.
9. Big Cr. at bridge on US 74-76, 4.4 km W Lake Waccamaw.
10. Bogue Swamp, 1.6 km ESE Hallsboro on CR 1736.
1 1. Bogue Swamp, 1.2 km E Hallsboro on US 74-76.
12. Red Hill Swamp, 16.1 air km N Whiteville on CR 1700.
13. White Marsh trib., 6.4 air km NNE Whiteville on CR 1700.
14. White Marsh, 3.2 km E Whiteville on US 74-76.
15. White Marsh, 8 air km S Hallsboro on CR 1001.
16. drainage canal, 2.4 km S Bolton on NC 21 1.
17. drainage canal, 9.2 km S Bolton on NC 21 1.
18. Waccamaw R. at Crusoe Island, 4 air km NE Old Dock.
19. Waccamaw R., 2.4 air km SE Old Dock on CR 1928.
20. Brunswick Co. line; Juniper Cr., oxbow just above confluence
Waccamaw R., 4.8 air km ESE Old Dock.
Waccamaw Drainage Fishes
21. Juniper Cr., overflow pond on CR 1928, 5.6 air km ESE Old
Dock.
31. Waccamaw R., 7.2. air km NE Pireway, "Reeve's Ferry."
32. Seven Crs., approx. 1.6 km N Pireway on NC 905.
33. Seven Crs., 8 air km NW Pireway on CR 1 108.
34. Juniper Swamp, 13.7 air km NW Pireway on CR 1 1 18.
35. Toms Fork, 12.9 air km SE Tabor City on CR 1 1 19.
37. Brunswick Co. line; Waccamaw R., 1.6 air km SE Pireway on NC
904.
38. Brunswick Co. line; Waccamaw R., 1 .6-9.7 river km S of NC 904
bridge.
Brunswick County
22. Aligator Swamp, just N Exum on CR 1335.
23. Big Swamp, 17.7 km S Bolton on NC 211.
24. Juniper Cr., 1.6 air km E Makatoka on CR 1340.
25. Juniper Cr. trib., 3.5 air km SE Makatoka on Cr 1342.
26. Juniper Cr., 29.8 km S Bolton on NC 21 1.
27a. Columbus Co. line; Waccamaw R. just below confluence Juniper
Cr., 5.2 air km SE Old Dock.
27b. Columbus Co. line; Waccamaw R., approx. 1.6 river km below
confluence Juniper Cr., 6 air km SE Old Dock.
28. Columbus Co. line; Waccamaw R., 6.4. air km SE Old Dock,
approx. 6.4 river km above NC 130.
29a. Columbus Co. line; Waccamaw R., 7.2 air km SE Old Dock,
approx. 2.8 river km above NC 130.
29b. Columbus Co. line; Waccamaw R., 8 air km SE Old Dock,
approx. 1.6 river km above NC 130.
29c. Columbus Co. line; Waccamaw R. at NC 130 bridge.
30. Wet Ash Swamp, 5.3 air km NE Longwood on NC 130.
36. Scippeo Swamp, 3.2 air km WNW Longwood on CR 1300.
SOUTH CAROLINA
Horry County (all stations on Waccamaw River)
39. approx. 3.2 air km SE Longs. 0.8 km N to 2.4 km S of SC 9
bridges.
40. 0.8 km S Red Bluff on SC 31.
41. approx. 4.8 river km W Red Bluff.
42. 4.8 air km N Nixonville.
43. 3.2 air km NNW Nixonville.
44. 2.4 air km NW Nixonville at SC 105.
45. 2.4 air km WNW Nixonville.
46. 0.8 air km NW Grahamville.
47. 7.2 air km SE Hickory Grove.
48. 2.4 air km SE Hickory Grove.
49. 5.6 air km W Conway (Hardee's Ferry).
50. 4 air km W Conway.
8 John R. Shute, Peggy W. Shute, David G. Lindquist
ANNOTATED SPECIES LIST
Lepisosteidae — gars
Lepisosteus osseus (Linnaeus), longnose gar. The longnose gar
occurs throughout the entire system. Most specimens are from Lake
Waccamaw and the main channel of the Waccamaw River. Stations:
la,c,d,e,f,g, 2b, c, 7, 8a,b,c,d.
Amiidae — bowfins
Amia calva Linnaeus, bowfin. This species appears to be uncom-
mon in Lake Waccamaw, but specimens have been taken throughout
the canal system and upper parts of the Waccamaw River. Tightly
packed schools of very young bowfin were occasionally observed, and
large adults were commonly seen in the river during low water. Stations:
Id, 3,4, 5, 8a,b,c,d, 9.
Anguillidae — freshwater eels
Anguilla rostrata (Lesueur), American eel. The American eel is
common throughout the entire system, in all habitat types sampled
where adequate cover exists. Stations: la,g,h,i, 2a, c, 7, 8a,b,c,d, 14, 19,
27b, 28, 29a,b,c, 33, 37, 38, 43, 44, 49.
Clupeidae — herrings
Alosa pseudoharengus (Wilson), alewife. Nine juveniles of this
anadromous species were collected from a single locality in the main-
stream of the river. Station: 42.
Alosa sapidissima (Wilson), American shad. Baker (1968) indicated
that the American shad ran upstream in the Waccamaw River as far
north as Juniper Creek. No specimens were collected during our survey.
Dorosoma cepedianum (Lesueur), gizzard shad. All of our collec-
tions of this species are from Lake Waccamaw. Most specimens were
seined from open shoreline areas at night, but small schools were occa-
sionally encountered while trawling at mid-lake stations. R. H. Moore
(pers. comm.) reported gizzard shad from lower sections of the Wac-
camaw River in South Carolina, Stations: la,c,f,i.
Dorsoma petenense (Gunther), threadfin shad. Threadfin shad were
introduced into Lake Waccamaw to provide forage for game species
(Nichols 1975). There is no record of successful overwintering and no
specimens were taken during our survey.
Umbridae — mudminnows
Umbra pygmaea (DeKay), eastern mudminnow. The mudminnow
occurs throughout the system where suitable habitat exists — standing
waters with dense growth of aquatic vegetation — and was common in
the canals around Lake Waccamaw. Louder (1962a) reported several
Waccamaw Drainage Fishes 9
specimens collected during a half-acre rotenone sample along the north-
east shoreline of the lake. We collected specimens from the lake on one
occasion. Stations: Id, 2b,c, 3, 5, 7, 8d, 9, 12, 21, 24, 25.
Esocidae — pikes
Esox americanus Gemlin, redfin pickerel. The redfin pickerel is
common throughout most of the system, except Lake Waccamaw where
it was rarely taken. Louder (1962a) reported nine specimens from near
the mouth of Second Creek along the northeast shore of the lake. Our
only collection from the lake was from an overflow area on the south-
eastern shore; most of our specimens were taken from standing (often
stagnant), weed-choked waters of small streams, canals, and occasionally
the main river channel. Stations: Id, 2a,b,c, 3, 5, 6, 8c, d, 9, 10, 12, 13, 15,
24, 25, 29a,c, 49, 50.
Esox niger Lesueur, chain pickerel. This species is also common
throughout much of the area surveyed. Frey (1951) and Louder (1962a)
reported its presence in the lake. Our only specimen from the lake is a
large adult (400-500mm TL) found in a gill net set off the southeastern
shore. The main channel of the Waccamaw River and some of its larger
tributaries appear to support the best populations of chain pickerel. Sta-
tions: le, 2a,b,c, 3, 6, 7, 8c,d, 12, 19, 29a,c, 38, 40, 42, 46, 47, 50.
Cyprinidae — minnows and carps
Cyprinus carpio Linnaeus, carp. Carp are reported by local residents
to be abundant in Lake Waccamaw. Indeed, the 1979 annual bow-fishing
tournament held at the lake produced 2,860 pounds of carp and longnose
gar. Despite these reports, we collected only three carp during our survey
and sighted several large adults at one of the canal stations. Louder
(1962a) listed this species for the first time from Lake Waccamaw, and
(1962b) also reported it from Bogue Swamp and White Marsh Swamp,
both tributaries to the Waccamaw River. In addition, R. H. Moore (pers.
comm.) reported carp from the lower sections of the river in South Carol-
ina. Stations: lb,i, 6, 7.
Hybognathus regius Girard, eastern silvery minnow. Although the
eastern silvery minnow is common in the main channel of Waccamaw
River in South Carolina, our survey produced the first specimens from
the river in North Carolina. Specimens were collected in open sluggish
waters devoid of aquatic vegetation. Stations: 29a, 38, 39, 40, 42, 44, 46.
Notemigonus crysoleucas (Mitchill), golden shiner. The golden
shiner occurs in a wide variety of habitats throughout the system, but
collections usually consist of only a few individuals. It was most often
encountered in standing water, and occasionally in the mainstream of the
river. Stations: la,b,c,e,g,h, 2a,c, 4, 6, 7, 12, 28, 38, 44.
Notropis chalybaeus (Cope), ironcolor shiner. Ironcolor shiners
occur throughout most of the main channel of the Waccamaw River.
10 John R. Shute, Peggy W. Shute, David G. Lindquist
Louder (1962a) reported two specimens from the northeast shore of Lake
Waccamaw, and we collected one specimen from the lake. Specimens
identified by Fowler (1935) as Erogalaformosa (Putnam) {-N. hypselopter-
us) from the Waccamaw drainage were determined to be a mixture of N.
chalybaeus and N. cummingsae. Notropis chalybaeus was often found in
association with N. petersoni, although it was never as numerous. Sta-
tions: lg, 7, 19, 28, 29a,c, 39, 49.
Notropis cummingsae Myers, dusky shiner. The dusky shiner also
appears to be largely confined to the main channel of the river. Small
schools were often encountered in open parts of the river, generally those
lacking dense vegetation. There are no reports of the species from Lake
Waccamaw. Stations: 7, 8a,d, 19, 27a, 28, 29a,c, 50.
Notropis hudsonius (Clinton), spottail shiner. Louder (1962b)
reported seven specimens of the spottail shiner from Big Creek, the prin-
cipal feeder stream to Lake Waccamaw. Records also exist from the Wac-
camaw River in North Carolina (Menhinick, ms.; Gilbert and Burgess
1980). According to Gilbert and Burgess (1980), this species inhabits
large, sluggish coastal rivers and brackish waters on the Atlantic slope.
The Big Creek locality does not fit this habitat description. The pre-
viously mentioned records for N. hudsonius, especially Big Creek, may be
based on misidentifications. However, museum specimens for these
records in the Waccamaw drainage could not be located for examination.
During our survey, no spottail shiners were collected.
Notropis maculatus (Hay), taillight shiner. This cyprinid is distrib-
uted throughout much of the Waccamaw system. Louder (1962b) found
it in Bogue Swamp, but it does not appear to be present in Lake Wac-
camaw. However, one population was located in a canal on the southwest
shore of the lake, and another in Big Creek not far from its mouth. Other
populations exist in the mainstream of the Waccamaw River. Stations:
2a, 6, 29a, 37, 38.
Notropis petersoni Fowler, coastal shiner. This is perhaps the most
widespread and abundant cyprinid throughout the system. The Lake
Waccamaw population was originally described at Notropis waccamanus
by Fowler (1942) and later placed in the synonymy of N. petersoni by
Frey (1951). Davis and Louder ( 1 97 1 ) discussed the biology of the species
in North Carolina waters (including Lake Waccamaw). We encountered
it in all habitat types sampled, with the notable exception of the canals
around Lake Waccamaw. Stations: l,a,b,c,d,e,f,g,h,i,l,m, 2a, c, 7, 8a, b,
19, 27b,c, 31, 37, 38, 39, 40, 42, 44, 47, 49, 50.
Catostomidae — suckers
Erimyzon oblongus (Mitchill), creek chubsucker. This species is
common throughout the system in a wide variety of habitats. Frey
(1951) first reported it from Lake Waccamaw, and apparently our single
specimen represents the only other published record from the lake. We
Waccamaw Drainage Fishes 11
encountered difficulties in identifying certain individuals, especially
juveniles, which often appear to be intermediate between E. oblongus
and E. sucetta. Rohde et al. (1979) also experienced similar difficulties
with specimens from southeastern North Carolina. Hanley (1976) con-
cluded that hybrids between E. oblongus and E. sucetta do occur, and
that these hybrids may backcross with both parental stocks. Stations:
le, 2a,c, 6, 7, 14, 19,21,28,29c.
Erimyzon sucetta (Lacepede), lake chubsucker. Very few lake chub-
suckers were taken during our survey and none from Lake Waccamaw,
although Louder (1962a) reported four specimens from the northeast
shore of the lake. Most of our specimens were taken in heavily vege-
tated areas of standing or slow-moving water. Stations: 3, 7, 21, 30.
Minytrema melanops (Rafinesque), spotted sucker. Our specimens
came from the main channel of the Waccamaw River and were taken
from open, moving waters with little or no cover. Stations: 19, 38.
Ictaluridae — freshwater catfishes
Ictalurus catus (Linnaeus), white catfish. The white catfish is wide-
spread throughout Lake Waccamaw and the main channel of the Wac-
camaw River. Several adults exceeding 400 mm TL were trawled from
mid-lake stations. Stations: la,b,c,e,h,i, 7, 8a,b,c, 28, 29a, 38, 44, 47.
Ictalurus melas (Rafinesque), black bullhead. Louder (1962b)
reported the black bullhead from Red Hill Swamp (White Marsh tribu-
tary) and Shingletree Swamp (Waccamaw River tributary). We col-
lected no specimens and found no museum specimens, and its occur-
rence in the Waccamaw drainage is doubtful. Menhinick et al. (1974)
suggested that Louder's records probably referred to /. nebulosus.
Ictalurus natalis (Lesueur), yellow bullhead. No specimens of the
yellow bullhead were taken from Lake Waccamaw during our survey,
but several were collected from the main channel of the Waccamaw
River. Louder (1962a) reported one small specimen from the northeast
shore of the lake. Stations: 8a,b,d, 19, 29a,47.
Ictalurus nebulosus (Lesueur), brown bullhead. The brown bull-
head has not previously been reported from Lake Waccamaw, but we
collected four specimens there during our survey. E. F. Menhinick (pers.
comm.) collected two specimens from Toms Fork Creek, a tributary to
Seven Creeks. Stations: lb,h.
Ictalurus platycephalus (Girard), flat bullhead. We discovered one
adult flat bullhead in a gill net set off the southeastern shore of Lake
Waccamaw, which represents only the second published report of this
species from the lake. Louder (1962a) reported a specimen from along
the northeast shore of the lake. The species is also present in the Wac-
camaw River below Conway, South Carolina (R. H. Moore, pers.
comm.). Station: le.
12 John R. Shute, Peggy W. Shute, David G. Lindquist
Ictalurus punctatus (Rafinesque), channel catfish. Although no
specimens of the channel catfish were collected during our survey, R. H.
Moore (pers. comm.) reported its presence in the lower reaches of the
Waccamaw River.
Noturus gyrinus (Mitchill), tadpole madtom. The tadpole madtom
has been taken in a variety of habitats throughout much of the system.
Most of our specimens came from the lake and the main channel of the
Waccamaw River. This madtom generally avoids swifter sections of the
river and was usually associated with thick vegetation or debris. Frey
(1951) discussed variation between populations from the North Carolina
Bay Lakes (including Lake Waccamaw). Stations: la,c,e,f,g,h,i, 2c, 5, 7,
8a, 38, 40, 46.
Noturus insignis (Richardson), margined madtom. This species was
taken exclusively from areas of flowing water and abundant cover in the
main channel of the Waccamaw River. It appeared to be the dominant
ictalurid species captured in the river. Stations: 7, 8a, d, 19, 28, 29a, b,
38.
Noturus species, broadtail madtom. Two distinct populations of
this undescribed madtom exist in the Waccamaw drainage. The form
found in the main channel of the Waccamaw River, taken by us from
Station 19 and downstream sites, is also found in the adjacent Cape
Fear drainage (Jenkins and Palmer 1978). It often occurred with N.
insignis. Specimens from Lake Waccamaw clearly differ from the river
specimens, but the degree of differentiation has not yet been determined
(R. E. Jenkins, pers. comm.). The closest relative appears to be N. lep-
tacanthus (Jenkins and Palmer 1978). The Lake Waccamaw form
(found throughout the lake and directly below the dam) was often
found in cans and bottles as well as under tiles placed as experimental
spawning sites for the Waccamaw darter. Broadtail madtoms appear to
be relatively common in the lake and may outnumber N. gyrinus. Sta-
tions: la,c,d,f,i,j,k,l,m, 7, 19, 28, 29a, 38, 50.
Amblyopsidae — cavefishes
Chologaster cornuta Agassiz, swampfish. The swampfish was never
common at any locality. It was encountered in standing or sluggish
waters, usually choked with aquatic vegetation or debris. No record of
this species from Lake Waccamaw exists and we found none there dur-
ing our survey. Stations: 2b, c, 6, 7, 8a,b,c,d, 24, 25, 26.
Aphredoderidae — pirate perches
Aphredoderus sayanus (Gilliams), pirate perch. Although Louder
(1962a) reported the pirate perch from Lake Waccamaw, and E. F.
Menhinick (pers. comm.) also collected one lake specimen, we collected
none from the lake. Pirate perch do occur in standing or sluggish and
Waccamaw Drainage Fishes 13
heavily vegetated waters elsewhere throughout the system. Stations:
2a,b,c, 3, 4, 6, 7, 8a,b,c,d, 10, 12, 13, 15, 17, 19, 20, 21, 22, 24, 25, 26,
27a, 29b,c, 30, 38, 39, 43.
Cyprinodontidae — killifishes
Fundulus chrysotus (Gunther), golden topminnow. The range of
the golden topminnow was originally thought to extend along the
Atlantic coast only as far north as the Santee drainage, South Carolina
(Shute 1980). Recently, however, specimens were collected by R. H.
Moore (pers. comm.) from the Waccamaw River at Bucksville, South
Carolina, 12.5 air km south-southeast of Conway. We examined the
specimens and concur with Moore's identification. In addition, speci-
mens from Waverly Mills, South Carolina (Waccamaw drainage) identi-
fied as this species by Fowler (1935) have been verified.
This species prefers river backwaters, slow-moving streams, or
ditches, and is usually associated with dense growths of aquatic vegeta-
tion (Shute 1980). Ample habitat certainly exists throughout most of the
drainage and additional populations quite likely exist.
Fundulus diaphanus (Lesueur), banded killifish. Fowler (1935)
reported the banded killifish from Waverly Mills (presumably on the
Waccamaw River), South Carolina. These specimens were examined by
Hubbs and Raney (1946) and re-examined by us, and are typically F.
diaphanus. This represents the southernmost extent of the species' range
(Gilbert and Shute 1980). No banded killifish were collected during our
survey.
Fundulus lineolatus (Agassiz), lined topminnow. We collected this
species from Lake Waccamaw for the first time. The species is rare in
the lake but common throughout the swamps and canals of the system.
Specimens are usually encountered in standing, heavily-vegetated, dark-
stained waters. Stations: lc,g,h, 2a,b,c, 3, 6, 7, 13, 14, 17, 20, 21, 22, 24,
27a, 28, 29c, 30, 37, 38, 40.
Fundulus waccamensis Hubbs and Raney, Waccamaw killifish. The
Waccamaw killifish was originally described as a Lake Waccamaw
endemic by Hubbs and Raney (1946). Recently, however, Bailey (1977)
reported specimens believed to be F. waccamensis from Lake Phelps,
most of which lies in Washington County in northeastern North Caro-
lina. Specimens from Lake Phelps examined by us and E. F. Menhinick
(pers. comm.) were found to differ slightly from F. waccamensis in
respect to head length, interorbital width, and caudal peduncle length.
This slight differentiation might tend to lessen the possibility that F.
waccamensis was accidentally introduced into Lake Phelps.
In the Waccamaw system, this killifish occurred at nearly all lake
stations sampled. In addition, it was found (especially during winter
months) throughout the lower parts of Big Creek, the canals around the
14 John R. Shute, Peggy W. Shute, David G. Lindquist
lake, and in the headwaters of the river just below the dam. The Wac-
camaw killifish typically inhabits the shallow, sandy shoreline of the
lake where it is often associated with dense stands of Panicum hemito-
mum. No specimens have been taken in the river farther than 100 m
below the lake. Stations: la,b,c,d,e,f,g,h,i,l, 2a,b,c, 3, 4, 5, 6, 7.
Poeciliidae — livebearers
Gambusia affinis (Baird and Girard), mosquitofish. The mosquito-
fish was collected throughout the entire system in nearly every habitat
type sampled, but was never collected while trawling at mid-lake sta-
tions and was only rarely taken at other stations within the lake. Sta-
tions. la,b,c,d,e,f,g,h, 2a,b,c, 3, 4, 5, 6, 7, 8a,b,c,d, 9, 10, 12, 14, 19, 20,
21, 22, 27a,b, 28, 29a,b,c, 30, 31, 33, 34, 36, 37, 38, 39, 40, 42, 43, 44, 49,
50.
Heterandria formosa Agassiz, least killifish. This diminutive poecil-
iid was only collected from one station in the South Carolina section of
the Waccamaw River, and is probably more abundant in the extreme
lower reaches of the river. Fowler (1935) reported it from Waverly
Mills, South Carolina (Waccamaw drainage). Station: 42.
Atherinidae — silversides
Menidia extensa Hubbs and Raney, Waccamaw silverside. The
Waccamaw silverside is possibly the most abundant fish in Lake Wac-
camaw. It also has the most limited distribution of the described
endemic species in the lake. Specimens have never been collected in Big
Creek or the canals surrounding the lake. A few stragglers (washovers)
are occasionally taken below the dam, but never more than 30 or 40 m
downstream from the lake. This species inhabits open, non-vegetated
waters along the shoreline of the lake and is occasionally taken in off-
shore waters. Stations: la,b,c,d,e,f,g,h,i, 7.
Percichthyidae — temperate basses
Morone americana (Gmelin), white perch. The white perch is
common in Lake Waccamaw and is considered to be the predominant
game species there. It was encountered, often in large numbers, at mid-
lake trawl stations. Except for one adult from Big Creek and another
from below the dam, the species was not collected outside the lake dur-
ing our survey. The specimen from Big Creek was injured and may have
been released by a fisherman. R. H. Moore (pers. comm.) reported
white perch from the lower Waccamaw River and Winyah Bay in South
Carolina. Stations: la,b,c,e,f,g,i, 2a, 7.
Morone saxatilis (Walbaum), striped bass. Baker (1968) indicated
that this anadromous species runs up the main channel of the Wac-
camaw River almost as far north as Juniper Creek. No specimens were
collected during our survey.
Waccamaw Drainage Fishes 15
Centrarchidae — sunfishes
Acantharchus pomotis (Baird), mud sunfish. This secretive species
was rarely encountered during our survey and was never collected from
Lake Waccamaw. Louder (1962a) reported it from a rotenone station
along the northeast shore of the lake, an area where feeder streams enter
and aquatic vegetation is abundant. We usually found it in areas of
standing water where submergent vegetation was extremely dense. Sta-
tions: 2b, 3, 8d, 9, 13, 15.
Centrarchus macropterus (Lacepede), flier. Louder (1962a) re-
ported the flier from Lake Waccamaw. Bruce B. Collette (pers. comm.)
also collected an adult from the north shore of Lake Waccamaw in
1958. We collected no specimens from the lake, but found the species
throughout much of the rest of the drainage, where it preferred standing
or sluggish water, usually with an abundance of aquatic vegetation. Sta-
tions: 2b,c, 4, 5, 6, 9, 10, 12, 28, 34, 38, 43.
Elassoma evergladei Jordan, Everglades pygmy sunfish. Three spec-
imens of the Everglades pygmy sunfish, the first to be reported from
Lake Waccamaw, were collected from a swampy area on the southeast-
ern shore of the lake. Major populations appear to be confined mainly
to Juniper Creek and tributaries where it is often associated with an
undescribed pygmy sunfish. Specimens were usually collected from
weedy, shallow backwaters of small streams. Stations: Id, 17, 21, 23, 24,
25, 26, 30.
Elassoma zonatum Jordan, banded pygmy sunfish. Louder (1962a)
reported this pygmy sunfish from Lake Waccamaw, where it was col-
lected with rotenone from dense vegetation along the northeast shore.
We did not find it in the lake, but specimens were commonly taken
from the surrounding canals and Big Creek. The species is common
throughout the system, except where replaced by E. evergladei and the
undescribed form. The habitat is similar to that of E. evergladei. Sta-
tions: 2a,b, 3, 4, 5, 6, 7, 8a,b,c,d, 9, 10, 12, 13, 15, 19, 20, 22, 27b, 29c,
30, 33, 34, 38.
Elassoma species, undescribed pygmy sunfish. This species, closely
related to E. zonatum (Bohlke and Rohde 1980), has only been collected
from two streams in the Waccamaw drainage. It is relatively common
throughout Juniper Creek, where its distribution and habitat closely
parallel those of E. evergladei. It is also collected regularly from one Big
Creek tributary (Station 2b), and has been taken once in the main
channel of Big Creek (Station 2c) and once at Station 2a. The absence
of this species from other streams within the system suggests a limited
distribution. Olmsted and Cloutman (1978) reported collecting an
undescribed Elassoma species from Black Creek (Pee Dee drainage) in
the Sandhills National Wildlife Refuge, South Carolina, but this report
appears to be based on misidentified E. zonatum (F. C. Rohde, pers.
comm.). Pygmy sunfish superficially resembling this species have been
16 John R. Shute, Peggy W. Shute, David G. Lindquist
collected by Rohde (pers. comm.) from Jasper County, South Carolina
(Savannah drainage). In addition to stations listed below, three speci-
mens were recently collected from a canal just east of Lake Waccamaw
(not mapped). Stations: 2a,b,c, 21, 24, 25, 26.
Enne acanthus chaetodon (Baird), blackbanded sunfish. This cen-
trarchid was collected from only six localities throughout the system,
where it occurs in standing, heavily vegetated waters. Aquatic pond-
weed, Potamogeton sp., was often present where specimens were col-
lected. Stations: 2a,b,c, 6, 8d, 21.
Enne acanthus gloriosus (Holbrook), bluespotted sunfish. The
bluespotted sunfish is distributed throughout the entire system but was
collected only once from Lake Waccamaw during our survey. It occurs
in quiet weedy backwaters of the Waccamaw River and tributaries. Sta-
tions: Id, 2a,b,c, 3, 4, 5, 6, 7, 10, 16, 21, 23, 24, 26, 29c, 38.
Enneacanthus obesus (Girard), banded sunfish. Louder (1962a)
reported this small species from Lake Waccamaw, and we collected it
there once. It was collected at scattered localities throughout the system,
and many specimens came from the Juniper Creek area. Habitat prefer-
ences are similar to those of the other Enneacanthus species. Stations:
Id, 2a,b,c, 3, 8, 17,21,23,26,30.
Lepomis auritis (Linnaeus), redbreast sunfish. Major populations
of this sunfish appear to be confined to the Waccamaw River; few spec-
imens were collected from Lake Waccamaw, and only two individuals
were taken from Big Creek. The redbreast sunfish was stocked in Lake
Waccamaw by the North Carolina Wildlife Resources Commission
(Nichols 1975). Stations: l,g,j, 2a, 7, 8a, 19, 27a, 28, 29a,b, 31, 37, 38,
39, 40, 42, 44, 49, 50.
Lepomis gibbosus (Linnaeus), pumpkinseed. The pumpkinseed was
commonly collected from Lake Waccamaw and the main channel of the
Waccamaw River, but was clearly absent from most of the smaller trib-
utaries. Adult specimens were often trawled from open waters of the
lake. Stations: la,d,g,h,i, 2a,b,c, 3, 4, 7, 8a,d, 21, 29c, 37.
Lepomis gulosus (Cuvier), warmouth. Although both Louder
(1962a) and Frey (1951) reported this species in Lake Waccamaw, we
took none from the lake during our survey. The species was common in
Big Creek, the canals around Lake Waccamaw, and many Waccamaw
River tributaries, where it occurs in quiet, weedy streams and river
backwaters. Stations: 2a,b,c, 4, 6, 7, 12, 14, 20, 21, 38, 40.
Lepomis macrochirus Rafinesque, bluegill. Bluegills are common
throughout the entire Waccamaw system, including Lake Waccamaw.
Specimens were associated with some type of cover, usually aquatic
vegetation or cypress stumps. Despite its abundance, large adults were
rarely taken. Stations: la,b,c,e,g,h,i, 2a,b,c, 3, 4, 6, 7, 8a,b,c, 12, 14, 21,
24, 29a,c, 31, 33, 34, 37, 38, 39, 40, 44, 49, 50.
Waccamaw Drainage Fishes 17
Lepomis marginatus (Holbrook), dollar sunfish. Two adult speci-
mens, taken on separate occasions, were collected from the south shore
of Lake Waccamaw above the dam. This is the first report of the species
from the lake. Throughout the system the dollar sunfish has been col-
lected in shallow, weedy backwaters of the river and tributaries, as well
as in borrow pits in the Green Swamp. Stations: lg, 2a, c, 7, 8b, c, 19, 21,
27b, 29a, c, 37, 38, 39, 40.
Lepomis microlophus (Gunther), redear sunfish. The redear sunfish
was introduced into the Waccamaw drainage to establish another suita-
ble game species (Louder 1962b; Nichols 1975, and pers. comm.). Dur-
ing our survey no specimens were collected from Lake Waccamaw and
only one was collected from the Waccamaw River in south Carolina.
Louder (1962b) reported specimens from Big Creek, Gum and Grey
swamps (White Marsh tributaries), Tabor City Run (Seven Creeks trib-
utary), and South Ash Swamp (direct tributary to Waccamaw River).
Station: 42.
Lepomis punctatus (Valenciennes), spotted sunfish. Frey (1951)
first reported the spotted sunfish in Lake Waccamaw. We failed to col-
lect any from the lake during our survey and took specimens only from
two localities on the Waccamaw River. Louder (1962b) reported the
species present throughout much of the system, and included several
specimens from Big Creek. Preferred habitat appears to be quiet, vege-
tated backwaters. Stations: 38, 40.
Micropterus salmoides (Lacepede), largemouth bass. Largemouth
bass are common throughout Lake Waccamaw and much of the Wac-
camaw River. Outside the lake the species appeared to be most common
in the main channel of the river and its larger tributaries. Stations:
la,c,d,e,g,h,l,k, 2a,c, 6, 7, 8a,b, 14, 29a, 37, 38, 39, 49, 42.
Pomoxis nigromaeulatus (Lesueur), black crappie. Although
Louder (1962a) reported this species to be one of the most important
game fishes in Lake Waccamaw, it was not often collected during our
survey. The species has been found in habitats ranging from open lake
waters to flood ponds of small swamps. Stations: la,e,i, 7, 12, 44.
Percidae — perches
Etheostoma fusiforme (Girard), swamp darter. The swamp darter is
the most widespread percid in the Waccamaw system, occurring in
nearly every habitat type sampled. It is particularly abundant in the
offshore waters of Lake Waccamaw. The northern subspecies, Etheos-
toma fusiforme fusiforme (Girard) reaches it southernmost limit in the
Waccamaw River (Collette 1962), but is replaced by the southern spe-
cies, E. f barratti (Holbrook), in the Pee Dee River (of which the Wac-
camaw is a tributary). All of our specimens were the nominate subspe-
cies, but extensive studies have not been conducted. Bailey and Frey
18 John R. Shute, Peggy W. Shute, David G. Lindquist
(1951) studied variation in darters of the subgenus Hololepis from some
natural lakes of North Carolina (including Lake Waccamaw). Stations:
la,b,c,d,e,f,g,h,i,m, 2a,c, 3, 4, 6, 7, 14, 21, 22, 27b, 28, 29c, 33, 37, 39,
44, 49.
Etheostoma olmstedi Storer, tessellated darter. With very few
exceptions, this darter is confined to the main channel of the Wac-
camaw River. Few other streams in the system offer suitable habitat,
which most often was shallow, moving water over sand or fine gravel
substrate. Stations: 7, 8a,b,c,d, 19, 27a,b, 28, 29a,b,c, 31, 33, 38, 39, 40,
41,42,43,44,47,49.
Etheostoma perlongum (Hubbs and Raney), Waccamaw darter.
We collected the Waccamaw darter from all localities sampled within
Lake Waccamaw. In spring and summer months it is common along the
shallow shoreline of the lake, sometimes in association with emergent
vegetation, but during colder months specimens are more often col-
lected in offshore waters. Etheostoma perlongum is sometimes taken
below the dam and in the upper headwaters of the Waccamaw River,
where it is found in association with E olmstedi and where specimens
exhibiting characters intermediate between the specie are often col-
lected. Stations: la,b,c,d,e,f,g,h,i,j,k,l,m, 7,8a.
Etheostoma serriferum (Hubbs and Cannon), sawcheek darter. The
sawcheek darter is widely distributed throughout the system, but no
specimens were collected from Lake Waccamaw. The species prefers
standing or sluggish and heavily vegetated water, often rich in organic
debris, and often occurs in the same habitat as pygmy sunfishes. Sta-
tions: 2b, 4, 6, 7, 8d, 13, 19, 21, 22, 24, 26, 27a, 28, 29a,c, 30, 33, 37, 38,
40, 42, 43, 44.
Perca flaveseens (Mitchill), yellow perch. The yellow perch is com-
mon in Lake Waccamaw and occasionally was collected in the Wac-
camaw River. It occurs in open waters with little cover, and is taken by
anglers, especially from areas around the dam. Stations: la,b,e,f,g,i,k,l,m,
2a, 7, 8d, 38, 42.
Soleidae — soles
Trinectes maculatus (Bloch and Schneider), hogchoker. We col-
lected the hogchoker only from the lower reaches of the Waccamaw
River, where specimens were taken in quiet, sluggish open waters over
mud bottoms. Stations: 40, 44, 48, 49, 50.
DISCUSSION
A drainage is defined by Jenkins et al. (1972) as "an interconnected
major group of streams, or systems entering the marine habitat "
Geographically, the Waccamaw River is a tributary in the Pee Dee
drainage, and the Waccamaw and Pee Dee rivers converge to form the
upper part of Winyah Bay, an estuarine habitat. We propose that
Waccamaw Drainage Fishes 19
Winyah Bay forms an effective barrier limiting faunal exchange between
the two rivers, at least for most of the freshwater forms. Therefore, we
argue that the Waccamaw should be viewed as a separate drainage that
limits dispersal of most primary freshwater fishes. A similar situation
exists between the Chowan and Roanoke Rivers, where the Chowan
enters the Roanoke drainage in the estuarine habitat of Albemarle
Sound. According to Jenkins et al. (1972) there is "some merit in con-
sidering it (the Chowan) a separate drainage."
Fifty-six species of freshwater and diadromous fishes were collected
from the Waccamaw drainage during our survey. These include four
families of secondary freshwater fishes (Lepisosteidae, Cyprinodontidae,
Poeciliidae and Atherinidae), and four families of diadromous fishes
(Anguillidae, Clupeidae, Percichthyidae, and Soleidae), with the remain-
ing ten families representing primary freshwater forms (mostly after
Myers 1938). Most of the secondary freshwater and diadromous species
behave as primary forms, and two — Fundulus waccamensis and Menid-
ia extensa — are known only from fresh water. Nine additional
species — Alosa sapidissima, Dorosoma petenense, Notropis hudsonius,
N. hypselopterus, Ictalurus melas, I. punctatus, Fundulus chrysotus, F.
diaphanus and Morone saxatilis — have been reported (various sources
listed in text) from the Waccamaw River and tributaries. Of these, Fun-
dulus chrysotus and F. diaphanus have been examined and verified by
us. Alosa sapidissima, Notropis hudsonius, Ictalurus punctatus and
Morone saxatilis were not verified, but probably do occur. It is doubtful
that the remaining three species — Dorosoma petenense, Notropis hyp-
selopterus, and Ictalurus melas — are found in this drainage. Compared
to other small Atlantic Coastal Plain drainages, the Waccamaw drain-
age has an unusually high species diversity (Table 2). This can be partly
attributed to the lentic habitats of Lake Waccamaw, from which 44 spe-
cies have been collected by us or otherwise reported.
The Waccamaw and Little Pee Dee (Big Swamp and Lumber sys-
tems) once occupied a much larger basin draining areas of the inner
Coastal Plain and Piedmont. Approximately 75,000 years ago the uplift
of the Cape Fear Fault (roughly paralleling the Cape Fear River)
resulted in elevation of land southwest of the Cape Fear River and sub-
sequent pirating of the upper parts of the Waccamaw and Little Pee
Dee systems by the Cape Fear (Zullo and Harris 1979). Stream flow was
diverted along this fault to form the Cape Fear River, leaving the Wac-
camaw and Little Pee Dee systems with greatly reduced drainage basins
confined largely to the Coastal Plain. Zoogeographic evidence also sug-
gests a close relationship between these drainages. Three species of fish
are shared exclusively by the Cape Fear and Pee Dee drainages: Semoti-
lus lumbee, Sandhills chub (Snelson 1980); Hybopsis species, thinlip
chub (Jenkins and Lachner 1980); and Noturus species, broadtail mad-
torn (Jenkins and Palmer 1978). However, only one of these, the broad-
20
John R. Shute, Peggy W. Shute, David G. Lindquist
Table 2. Number of species (genera) found in fresh waters of selected Atlantic
coastal drainages.
boides, and Dormitator maculatus not included.
Combined Ashepoo, Combahee, Broad, and New rivers. Data modified
from Swift et al. (1977) with additional species from Anderson (1964);
record of Chaetodipterus faber not included.
tail madtom, is known from the Waccamaw. The remaining two species
are known from the Lumber and Lynches rivers of the Pee Dee drain-
age. Therefore, it is not surprising that much of the ichthyofauna of the
lower Pee Dee (Waccamaw, Lumber and Big Swamp systems) is shared
with the Cape Fear drainage (Average Faunal Resemblance Index = 84;
Jenkins et al. 1972). Indeed, only two other rivers of the Central Atlan-
tic Slope (Neuse and Tar) show a greater degree of faunal resemblance
(Average Faunal Resemblance Index = 94; Jenkins et al. 1972).
Additionally, Etheostoma fusiforme fusiforme reaches the southern
terminus of its range in the Waccamaw River (Collette 1962). Etheos-
toma f. barratti is found from the Pee Dee drainage (to which the Wac-
camaw is an eastern tributary) southward. The presence of E. f fusi-
forme in both the Waccamaw and Cape Fear rivers and its absence
Waccamaw Drainage Fishes 21
from the Pee Dee provides further evidence of past connections between
the two drainages and suggests a faunal separation of the Waccamaw
from the Pee Dee. We suggest that populations of E. f fusiforme were
probably present in the Little Pee Dee system (as in the Cape Fear and
Waccamaw) before the uplift of the Cape Fear Fault. Etheostoma f
barratti from the Pee Dee then invaded the Lumber via the Little Pee
Dee, replacing the nominate subspecies. There was possibly little or no
opportunity for such upstream dispersal of E. f. barratti into the Wac-
camaw because of salinity barriers, and therefore the populations of E.
f fusiforme persisted.
Only one known connection between the Waccamaw and Cape
Fear rivers presently exists. A series of man-made canals, dug to
improve tree farm drainage, connects Honey Island Swamp (Juniper
Creek tributary) with Dans Creek of the Cape Fear drainage and Big
absence Creek which drains into Lake Waccamaw (Fig. 1). Only a
limited number of species should be able to negotiate the small, shallow
and often stagnant canals, thus limiting substantial faunal exchange.
The undescribed Elassoma was recently collected in one of these canals
just east of Lake Waccamaw, which suggests the possibility that it may
have gained access to the Big Creek system via the canals from Juniper
Creek. This may explain of the species from areas west of the Wac-
camaw River and other tributaries of the river (apart from Juniper
Creek and Big Creek) even where habitat appears suitable. Northward
expansion of its range would be possible through the canal system into
the Cape Fear drainage. Collecting in adjacent Cape Fear drainages has,
however, provided no specimens.
According to Jenkins et al. (1972), lowland endemics or exclusively
shared forms are not common on the Central Atlantic Slope. The Wac-
camaw has at least five of these. Etheostoma perlongum and Menidia
extensa are Waccamaw endemics. Fundulus waccamensis is either
endemic to the Waccamaw or shared with Lake Phelps (coastal Albe-
marle drainage), pending taxonomic decisions. The undescribed Elas-
soma is probably shared between the Waccamaw and Savannah drain-
ages (F. C. Rohde, pers. comm.). The undescribed Noturus is represented
by two forms within the Waccamaw drainage; specimens from Lake
Waccamaw represent a population superficially distinct from the Wac-
camaw River population and may represent another Waccamaw
endemic. The form present in the Waccamaw River is also found in
rivers of the lower Pee Dee and Cape Fear drainage (Jenkins and
Palmer 1978; Jenkins, pers. comm.).
In summary, the Waccamaw drainage is unique among small cen-
tral Atlantic coastal drainages in having a highly diversified fish fauna
including endemic and exclusively shared forms. The Waccamaw and
Little Pee Dee systems once extended farther north into the inner Coast-
al Plain and Piedmont. These streams were beheaded by the uplifting of
22 John R. Shute, Peggy W. Shute, David G. Lindquist
the Cape Fear Fault and subsequent formation of the Cape Fear River,
resulting in faunal similarities between the drainages.
ACKNOWLEDGMENTS. — This study was made possible by grant-
in-aid funds under Section 6 of the Endangered Species Act of 1973 (PL
93-205). We wish to thank Edward F. Menhinick, University of North
Carolina at Charlotte, for his assistance in locating questionable records
and permission to review his unpublished distribution maps of North
Carolina freshwater fishes. William M. Palmer and Alvin L. Braswell
provided access to state stream survey collections housed at the North
Carolina State Museum of Natural History. Joseph R. Bailey, Duke
University, provided information on specimens in his care. Peter S.
Coleman, Charleston Museum, was extremely helpful in locating and
verifying specimens housed in his care. We are grateful to Richard H.
Moore, Coastal Carolina College, for supplying us with information
and specimens from the Waccamaw River in South Carolina. In addi-
tion, we would like to thank all those who assisted in many hours of
field work during this survey. Dr. Bruce B. Collette, National Marine
Fisheries Service, provided helpful comments on the manuscript, and
copies of some of his field notes from Waccamaw studies. Fred C.
Rohde and Robert E. Jenkins also reviewed the manuscript. Finally, we
thank John A. McNeill, Sr., for use of his cabin at Lake Waccamaw.
LITERATURE CITED
Anderson, William D. 1964. Fishes of some South Carolina Coastal Plain
streams. J. Fla. Acad. Sci. 27(l):31-54.
Anonymous. 1978. Preliminary analysis of the potentials of the Waccamaw
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Systems. Compiled by Natural and Scenic Rivers study team, Columbus
Co., North Carolina. Unpublished.
Bailey, Joseph R. 1977. Freshwater Fishes, pp. 265-298 in J. E. Cooper, S. S.
Robinson and J. B. Funderburg (eds.). Endangered and Threatened Plants
and Animals of North Carolina. N. C. State Mus. Nat. Hist., Raleigh, xvi
+ 444 pp.
, and D. G. Frey. 1951. Darters of the genus Hololepis from some
natural lakes of North Carolina. J. Elisha Mitchell Sci. Soc. 67(2): 191-204.
Baker, W. Donald. 1968. A reconnaissance of anadromous fish runs into the
inland fishing waters of North Carolina. Fed. Aid Fish Restor. Proj. AFS-
3, N. C. Wildl. Resour. Com., Raleigh. 23 pp.
Bohlke, James E., and F. C. Rohde. 1980. Elassoma zonatum Jordan, banded
pygmy sunfish. p. 586 in D. S. Lee, et al. Atlas of North American Fresh-
water Fishes, N.C. State Mus. Nat. Hist., Raleigh, x + 867 pp.
Collette, Bruce B. 1962. The swamp darters of the subgenus Hololepis (Pisces,
Percidae). Tulane Stud. Zool.9(4): 115-21 1.
Davis, James R., and D. E. Louder. 1969. Life history and ecology of Menidia
externa. Trans. Am. Fish. Soc. 9#(3):466-472.
Waccamaw Drainage Fishes 23
, and 1971. Life history and ecology of the cyprinid fish
Notropis petersoni in North Carolina waters. Trans. Am. Fish Soc. W0(4)J26-733.
Fowler, Henry W. 1935. Notes on South Carolina freshwater fishes. Contrib.
Charleston Mus. 7. 28pp.
1942. Description of six new freshwater fishes (Cyprinidae and Per-
cidae) from the southeastern United States. Not. Nat. (Phila.) 107. 1 1 pp.
Frey, David G. 1948a'. North Carolina's bay lakes. Wildl. in N. C. 72(9):3-8.
1948b. A biological survey of Lake Waccamaw. Wildl. in N. C.
72(9): 17-20.
1949. Morphometry and hydrography of some natural lakes of the
North Carolina Coastal Plain. The bay lake as a morphometric type. J.
Elisha Mitchell Sci. Soc. (55(1): 1-37.
1951. The fishes of North Carolina's bay lakes and their intraspecific
variation. J. Elisha Mitchell Sci. Soc. (57(1): 1-44.
Gilbert, Carter R., and G. H. Burgess. 1980. Notropis hudsonius (Clinton),
spottail shiner, pp. 275-76 in D. S. Lee, et al. Atlas of North American
Freshwater Fishes. N. C. State Mus. Nat Hist., Raleigh, x + 867 pp.
, and J. R. Shute. 1980. Fundulus diaphanus (Lesueur), banded killif-
ish. p. 513 in D. S. Lee, et al. Atlas of North American Freshwater Fishes.
N. C. State Mus. Nat. History., Raleigh, x + 867 pp.
Hanley, Robert W. 1976. Population phenetics of chubsuckers in North Carol-
ina (Erimyzon: Catostomidae). Unpub. Master's thesis, Duke Univ., Durham.
182 pp.
Hubbs, Carl L., and E. C. Raney. 1946. Endemic fish fauna of Lake Wac-
camaw, North Carolina. Misc. Publ. Univ. Mich. Mus. Zool. (55:1-30.
Hueske, Edward E. 1948. Fish resources of the bay lakes. Wildl. in N. C.
/2(9):11-16.
Jenkins, Robert E., and E. A. Lachner. 1980. Hybopsis zanema (Jordan and
Brayton), Santee chub. p. 197 in D. S. Lee, et al. Atlas of North American
Freshwater Fishes. N. C. State Mus. Nat Hist., Raleigh, x + 867 pp.
, and F. J. Schwartz. 1972. Fishes of the central Appalach-
ian drainages: their distribution and dispersal, pp. 43-117 in P. C. Holt
(ed.). The distributional history of the biota of the southern Appalachians,
Part III: Vertebrates. Res. Div. Monogr. 4, Va. Polytech. Inst. State Univ.,
Blacksburg. 306 pp.
, and W. M. Palmer. 1978. A new species of madtom catfish (Ictalur-
idae) from the Coastal Plain of the Carolinas. ASB Bull. 25(2):57. Abstract.
Lindquist, David G., J. R. Shute and P. W. Shute. 1981. Spawning and nesting
behavior of the Waccamaw darter, Etheostoma perlongum. Environ. Biol.
Fishes (5(2): 177-191.
Louder, Darrell E. 1962a. An annotated checklist of the North Carolina
bay lakes fishes. J. Elisha Mitchell Sci. Soc. 7S(l):68-73.
1962b. Survey and classification of the Lumber River and Shallotte
River, North Carolina. Final Rep. Fed. Aid Fish Restor. Proj. F-14-R, N.
C. Wildl. Resour. Com., Raleigh. 12 pp.
24 John R. Shute, Peggy W. Shute, David G. Lindquist
Menhinick, Edward F. Manuscript. The Freshwater Fishes of North Carolina.
, T. M. Burton and J. R. Bailey. 1974. An annotated checklist of the
freshwater fishes of North Carolina. J. Elisha Mitchell Sci. Soc.
90(1):24-5O.
Myers, George S. 1938. Fresh-water fishes and west Indian zoogeography.
Smithson. Inst. Annu. Rep. (1937), Publ. 3465:339-64.
Nichols, Lacy E. 1975. Lake Waccamaw. N. C. Wildl. Resour. Com.: 1-8.
Olmsted, Larry L., and D. G. Cloutman. 1978. Fishes of the Carolina
Sandhills National Wildlife Refuge. Final Rep. U. S. Dept. Inter., Fish
Wildl. Serv., Carolina Sandhills Nat. Wildl. Refuge, McBee, SC. 26 pp.
Robins, C. Richard, 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 Scien-
tific Names of Fishes from the United States and Canada (4th ed.). Am.
Fish. Soc. Spec. Publ. 12. 174 pp.
Rohde, Fred C, G. H. Burgess. and G. W. Link. 1979. Freshwater fishes of
Croatan National Forest, North Carolina, with comments on the zoogeo-
graphy of Coastal Plain fishes. Brimleyana 2:97-1 18.
Shute, John R. 1980. Fundulus chrysotus (Giinther), golden topminnow. p. 510
in D. S. Lee, et al. Atlas of North American Freshwater Fishes. N. C. State
Mus. Nat. Hist., Raleigh, x + 867 pp.
Shute, Peggy W., D. G. Lindquist and J. R. Shute. Manuscript. Spawning
behavior, reproduction, fecundity, sexual dimorphism and early life history
stages of the Waccamaw killfish, Fundulus waccamensis. Submitted to
Environ. Biol. Fishes.
, J. R. Shute and D. G. Lindquist. 1979. Notes on the Spawning,
fecundity and diet of the Waccamaw killfish, Fundulus waccamensis. ASB
Bull. 2<5(2):49. Abstract.
, and In press. Age, growth and early life history of
the Waccamaw darter, Etheostoma perlongum. Copeia.
Snelson, Franklin F., Jr. 1980, Semotilus lumbee Snelson and Suttkus, Sand-
hills Chub. p. 363 in D. S. Lee, et al. Atlas of North American Freshwater
Fishes. N. C. State Mus. Nat. Hist., Raleigh x + 867 pp.
Swift, Camm, R. W. Yerger and P. R. Parrish. 1977. Distribution and natural
history of the fresh and brackish water fishes of the Ochlockonee River,
Florida and Georgia. Bull. Tall Timbers Res. Sta. 20:1-1 1 1.
Teulings, Robert P., and J. E. Cooper. 1977. Cluster areas, pp. 409-433 in J. E.
Cooper, S. S. Robinson and J. B. Funderburg (eds.). Endangered and
Threatened Plant and Animals of North Carolina, N. C. State Mus. Nat.
Hist., Raleigh, xvi + 444 pp.
Zullo, Victor A., and W. B. Harris. 1979. Plio-Pleistocene crustal warping in the
outer Coastal Plain of North Carolina, pp. 31-40 in G. R. Baum, W. B.
Harris and V. A. Zullo (eds.). Structural and stratigraphic framework for
the Coastal Plain of North Carolina. Carol. Geol. Soc, 1979 Field Trip
Guidebook. 1 1 1 pp.
Accepted 2 December 1981
A Taxonomic Analysis of Pseudemyd Turtles
(Testudines: Emydidae) from the New River,
and Phenetic Relationships in the Subgenus Pseudemys
Michael E. Seidel
Biological Sciences and N. Bayard Green Museum,
Marshall University, Huntington, West Virginia 25701
ABSTRACT. — The morphology and geographical origin of a disjunct
population of aquatic turtles, genus Pseudemys (- Chrysemys auct.) in
the New River of Virginia and West Virginia are analyzed. Previous
identification of these turtles as "Chrysemys" floridana is reappraised
by comparison of shell proportions, color patterns, and cranial morph-
ology to those of other species and subspecies of Pseudemys (sensu
stricto). Discriminant analysis of 28 cranial characters broadly separ-
ates redbelly turtles (P. rubriventris, P. nelsoni, P. alabamensis) from
New River Pseudemys and most populations of P. concinna and P.
floridana. Based on morphological similarities, New River Pseudemys
are identified as eastern river cooters, P. c. concinna. Natural history
information and recent extensions of the known range suggest that
Pseudemys in the New River represents a natural, established popula-
tion. A late Pleistocene dispersal of cooters from the Virginia-North
Carolina Piedmont Plateau into the New River is proposed.
INTRODUCTION
Bayless (1972) reported a population of cooter turtles in the New
River at Bluestone Reservoir, Summers County, West Virginia. This
locality is in the southern part of the state where the New River, a
Kanawha-Ohio River tributary, enters from Virginia (Fig. 1). He tenta-
tively identified the turtles in Bluestone Reservoir as "Chrysemys" flori-
dana, without assigning them to subspecies. This population is broadly
disjunct from the range of cooters in the Mississippi and lower Ohio
River valleys and is isolated by the Atlantic-Ohio divide from the near-
est populations in the Virginia and North Carolina Piedmont Plateau
(Fig. 1).
Despite a thorough analysis of key morphological characters, Bay-
less (1972) failed to clearly establish the taxonomic status and probable
geographical origin of the New River Pseudemys. This was unavoidable
due to the poorly understood systematic and distributional relationships
between Pseudemys floridana (LeConte) and Pseudemys concinna
(LeConte) in northern parts of their ranges (Crenshaw 1955; Minton
1972; Pritchard 1979; Martof et al. 1980). Further uncertainty deve-
loped when Bayless' voucher specimens at the National Museum of
Natural History (USNM 192635-7) were reidentified in 1973 as redbelly
turtles, Pseudemys rubriventris (LeConte) (Fran I. McCullough, pers.
Brimleyana No. 6: 25^44. December 1981. 25
Michael E. Seidel
Taxonomy of Pseudemyd Turtles 27
comm.). This species is known to occur only in drainage basins of the
Atlantic slope (Conant 1975). Based on external morphology of the
New River specimens, Carl H. Ernst (pers. comm.) suggested possible P.
rubriventris influence in the population and the possibility of a P. flori-
dana x rubriventris hybrid swarm similar to that reported in North
Carolina by Crenshaw (1965). All that remained evident was that the
turtles collected from the New River at Bluestone Reservoir belong to
the subgenus Pseudemys (sensu McDowell 1964; Vogt and McCoy
1980). This subgenus includes P. floridana, P. concinna and a P. rubri-
ventris series of P. nelsoni Carr, P. alabamensis Baur and P. rubriventris.
The present study, which analyzes cranial structure, shell morphol-
ogy, color patterns and incidental natural history data, was designed to
evaluate the taxonomic position and probable origin of Pseudemys in
the New River. Although a systematic analysis of the subgenus Pseude-
mys was not the original or primary objective of the study, problematic
levels of speciation and questionable validity of traditional key charac-
ters necessitated comparisons with all taxa in the subgenus, especially
those with more northerly distributions.
MATERIALS AND METHODS
Twelve adult Pseudemys from the New River were examined and
morphologically compared to other forms of the subgenus Pseudemys.
The New River sample includes a specimen collected in Giles County,
Virginia, in 1975; seven individuals collected in Bluestone Reservoir in
May 1980; and the four specimens from Bluestone Reservoir described
by Bayless (1972). Weaver and Rose (1967) found shell depth, nuchal ( =
cervical) scute underlap, and gular scute overlap especially useful in
separating P. floridana, P. concinna, and P. nelsoni. These measure-
ments were taken on New River Pseudemys and initially calculated as
ratios following the methods of Weaver and Rose (1967) to allow direct
comparison to their data. For subsequent shell comparisons among
adult (153-300 mm carapace length) New River Pseudemys, P. rubriven-
tris, and northern subspecies of P. concinna and P. floridana (Fig. 3):
shell depth was calculated as a ratio of height to carapace length; gular
scute overlap was calculated as 10X the ratio of gular scute length (dor-
sal surface) to plastron length; and nuchal scute underlap was calculated
10X the ratio of cervical scute length (ventral surface) to carapace
length. Measurements were made with Helios or bow outside calipers.
Characters traditionally used by Carr (1952) and Crenshaw (1955), such
as patterns and coloration of head, neck, plastron and carapace, were
also analyzed.
For phenetic analysis of adult cranial morphology, skulls from 4
Bluestone specimens were compared to skulls of 8 P. rubriventris, 4 P.
nelsoni, 3 P. alabamensis, 5 P. f. floridana, 6 P. f peninsularis, 1 3 P. f
hoyi, 5 P. c. concinna, 5 P. c. suwanniensis, 8 P. c. hieroglyphica, 9 P. c.
28 Michael E. Seidel
mobilensis and 6 P. c. texana. All specimens were adults and ranged
from 27 to 57 mm skull length. Although the sex of some of the skulls
examined was not identified, it was evident that none of the samples
was skewed heavily toward males or females. Twenty-nine measure-
ments were made on each cranium and mandible (± 0.1mm): condyl-
obasal length (midline length of skull, from posterior aspect of occipital
condyle to anteriormost point of premaxillae); interquadratal width;
supraoccipital length; pterygoid width (least); interpterygoid process
width (greatest); orbital height and orbital width; narial height and
width; prefrontal length (midline); interorbital width (least); postorbital
length (width of postorbital arch, least distance from orbit to superior
temporal fossa); postorbital-quadratojugal breadth (least distance be-
tween superior temporal fossa and ventral ridge of quadratojugal);
jugal-quadratojugal length (least distance between orbit and tympanic
cavity); maxillary alveolar width (least); foramen magnum width and
height; basisphenoid-basioccipital length; interforamina stapedio-
temporale width (distance between the temporal-stapedial foramina);
anterior skull width (at anterior rim of tympanic cavity); posterior skull
width (at posterior rim of tympanic cavity); intersquamosal breadth
(distance between posterior aspects of squamosals); premaxillary height
(midline); temporal arch width (least distance between rim of tympanic
cavity and superior temporal fossa); otic capsule length (from posterior
rim of tympanic cavity); dentary-coronoid height; dentary breadth (mid-
line); dentary alveolar width (lateral); and lingual alveolar width (prox-
imal to median ridge of dentary).
Due to intra- and interspecific size variation, regression analysis
was applied to all skull measurements to remove linearly-related effects
of size. Condylobasal length was used as the independent variable for
regression analysis of the other variables. The SAS General Linear
Models procedure produced residual values for each character; these
values were used as "size-free" variables. Using these 28 variables, the
SAS discriminant analysis procedure tested for homogeneity of within-
group covariance matrices. As the 28 characters measured showed no
evidence of heterogeneity of the within-group covariance matrices,
groups (taxa) were compared by step-wise discriminant analysis using
the computer program, BMD07M (Dixon 1974), which generates canon-
ical variates with maximum between-group variance relative to within-
group variance. The canonical variate means are plotted on the first two
axes, and analysis of variance describes significant differences between
groups (P < 0.05). Using canonical functions, the posterior probability
of each turtle belonging to its respective group is computed and classi-
fied accordingly.
Taxonomy of Pseudemyd Turtles 29
ABBREVIATIONS
ACE, United States Army Corps of Engineers, Huntington District.
AMNH, American Museum of Natural History.
CM, Carnegie Museum of Natural History.
FMNH, Field Museum of Natural History.
KU, University of Kansas Museum of Natural History.
MCZ, Museum of Comparative Zoology, Harvard University.
MES, Collection of Michael E. Seidel.
NCSM, North Carolina State Museum of Natural History.
UMMZ, University of Michigan Museum of Zoology.
USNM, National Museum of Natural History.
WVBS, West Virginia Biological Survey, Marshall University.
SPECIMENS EXAMINED
P. alabamensis: MCZ 1659-61, 1663, 1898; AMNH 107676. P. nelsoni: UMMZ
127059-60; AMNH 75640; MCZ 54131, 54684. P. rubriventris: AMNH 71276,
71280, 71282, 71293, 76175, 79132, 79134, 80218, 81869, 90641-42, 90644; CM
34409, 37272, 39672, 45188; FMNH 22137; MCZ 1666, 1671, 1674, 1677, 12877,
76679, 157828; MES 132. P. f. flohdana: FMNH 8222; MCZ 1635, 1651,
46221-22; USNM 25260; AMNH 50985, 75641; NCSM 5884-85, 5927-28, 5930,
8518. P. f. peninsularis: FMNH 22074; UMMZ 12937, 130081,44976; AMNH
64156, 69899, 110189; MCZ 19179. P. f. hoyi: KU 1176-79, 1185, 2221, 2800,
2830-31, 40166; MCZ 29080-81. P. c. concinna: USNM 8920, 15990, 60895,
92529-31; FMNH 22138; MCZ 1642, 1664, 12764, 54680; AMNH 75649; NCSM
10328, 17339, 20128, 20240, 20253. P. c. suwanniensis: UMMZ 127058, 129385;
AMNH 80233; FMNH 22473; MCZ 43030, 54675, 54679. P. c. mobilensis:
AMNH 69908; MCZ 1636-39, 1648-50, 1652, 42327. P. c. hieroglyphica:
USNM 9659, 79449, 86728, 102679, 104397; CM 60560; CM (field series) 38068-
69, 38071-72, 38077, 38107, 38132, 38134, 38155; UMMZ 101754, 128176,
133845; AMNH 69901-02, 69905-06; WVBS 3155, 3965. P. c. texana: USNM
26424, 26438, 78518; UMMZ 133836, 154982; MCZ 46483; KU 39986, 49630;
AMNH 111960; MES 75. New River Pseudemys: USNM 192635-37; WVBS
4093; MES 489, 526, 528-29, 863, 865; UMMZ 88488; two adults released; two
hatchlings MES uncataloged; shell bones ACE FS1-68, FS2-81, FS2-183.
RESULTS
Analysis of patterns and color in adult specimens from Bluestone
Reservoir (Fig. 2) showed strong similarities to P. concinna. In all 12
specimens the submarginals contain dark markings that are usually
open circles rather than the smudgelike or solid circles common in P.
rubriventris and P.floridana (Carr 1952; Mount 1975). In most individ-
uals the bridge has an extensive dark pattern that contacts the submar-
ginal markings. The pleurals have relatively thin yellow-tan lines and
some individuals show the clearly defined "C" figure on pleural II typi-
cal of P. concinna. Ventral and supratemporal stripes on the head and
30
Michael E. Seidel
Fig. 2. Ventral and dorsal views of hatchling (above) and adult (middle and
below) Pseudemys from Bluestone Reservoir, Summers County, West Virginia.
Taxonomy of Pseudemyd Turtles
31
0.5
0.4
0.3
0.2
0.1
4tt
■ffl-f
ii
•fr
R B
SHELL DEPTH
GULAR SCUTE
OVERLAP
NUCHAL SCUTE
UNDERLAP
Fig. 3. Measurements of 7 P. c. concinna (C) from Virginia and North Carolina;
1 1 P. c. hieroglyphica (H) from Indiana and Tennessee; 8 P. f. floridana (F)
from North Carolina, South Carolina and Georgia; 14 P. rubriventris (R) from
Massachusetts, New Jersey, Pennsylvania, Delaware, Virginia and North Carol-
ina; 12 Pseudemys (B) from New River, Virginia and West Virginia. Horizontal
lines are means, vertical lines are ranges, rectangles are one standard deviation.
Values are ratio calculations described in MATERIALS AND METHODS.
32 Michael E. Seidel
neck are broad whereas paramedian dorsal stripes are very narrow,
sometimes convergent or obscure. All seven New River specimens that I
collected have a yellow plastron that was lightly tinged with orange in
two individuals. Plastral markings typical of P. concinna are present on
each turtle, but less prominent in older specimens (Fig. 2). Slight fading
of markings and coloration was evident after 2 to 3 months in captivity.
The cutting edge (tomium) of the upper jaw in most individuals is
weakly emarginate and the lower jaw is moderately serrate (Fig. 2).
Four skulls which were analyzed show no evidence of a vomerine shelf
contributing to the crushing (alveolar) surface of the upper jaw, a uni-
que characteristic of species in the rubriventris series (McDowell 1964).
Twelve adult New River turtles examined have a shell depth range
of 31.1 to 39.4 (ratio calculations following Weaver and Rose 1967),
which broadly overlaps the range reported for P. concinna but falls
below the ranges of P. floridana and P. nelsoni. Nuchal scute underlap
ratios in New River Pseudemys, 16.0 to 23.4, are greater than in P.
concinna but are within the combined range of ratios for P. floridana
and P. nelsoni. The gular scute overlap ratio was highly variable (7.9-
18.8), falling within the combined ratio range reported for all three spe-
cies (Weaver and Rose 1967). Because most of the specimens examined
by Weaver and Rose were from Florida, shells of adult Pseudemys from
northern localities were measured and compared to New River speci-
mens. These ratios (Fig. 3) again indicate a shallow shell depth in New
River turtles (B) and P. concinna (C,H). Gular scute overlap and nuchal
scute underlap are greater in P. rubriventris (R), than in New River
specimens (B) and northern subspecies of P. floridana (F) and P. con-
cinna (C,H). However, these two characters are highly variable and
apparently not effective in separating P. concinna and P. floridana out-
side of Florida.
Results from the discriminant analysis of all Pseudemys skulls are
presented in Figure 4. On the first canonical axis (K i) plots of skulls in
the rubriventris series (A,N,R) are clearly disjunct from all forms except
P. c. texana (T). Broad separation is seen between P. rubriventris (R)
and New River Pseudemys (B). Also noteworthy is the separation, on
the second canonical axis (K2) of P. alabamensis (A) from P. rubriven-
tris (R) and P. nelsoni (N). The first two axes account for 51% and 12%
of the total dispersion, respectively. In order of increasing importance,
temporal arch width, jugal-quadratojugal length, interorbital width and
dentary alveolar width were the most influential characters providing
separation on the first axis (Table la). Anterior skull width, dentary
alveolar width, and lingual alveolar width contributed most to separa-
tion on the second axis (Table la). All individuals were classified into
their appropriate taxa, except one P. c. mobilensis that was placed in P.
f floridana and one P. f hoyi that was placed in P. c. hieroglyphic a.
The P. c. mobilensis specimen (MCZ 1651) was collected within the
Taxonomy of Pseudemyd Turtles
33
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2 s
.Is
8 J
•S 2.
O _i
1>
00
T3
E
3
C
E
Z £
h u-
cd
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c
Q
34 Michael E. Seidel
range of P. f floridana (Mobile, Alabama) and a resemblance to P.
floridana was noted prior to discriminant analysis. This skull was re-
identified as P. f floridana for subsequent comparison. The misclassifi-
cation of a P. f hoyi skull as P. c. hieroglyphica is not surprising con-
sidering the proximity of means (H,Y) for these forms (Fig. 4). This
skull was reassigned to P. c. hieroglyphica in further analyses. Based on
the 28 cranial characters measured, analysis of variance indicated signif-
icant differences comparing New River Pseudemys to all taxa (P < 0.01)
except P. c. concinna (P > 0.05), P. c. suwanniensis ( > 0.01), and P. c.
mobilensis (P>0.01).
To further analyze the relationships of New River Pseudemys to P.
floridana and P. concinna, discriminant analysis of skulls was applied
again, with the rubriventris series omitted. Results of this analysis are
similar to the first, but three clusters now become apparent (Fig. 5): a
western and Mississippi Valley group (P. c. texana, P. f hoyi, P. c.
hieroglyphica; TYH), an eastern P. concinna group (P. c. suwanniensis,
P. c. mobilensis, P. c. concinna, and New River Pseudemys; SMCB),
and an eastern P. floridana group (P. f floridana, P. f peninsularis;
FP). Noteworthy is the broad separation (on the first axis, Ki) of P. f
hoyi from other P. floridana skulls and its extensive overlap with P.
concinna (Fig. 5). The first two canonical axes account for 49% and
20% of the total dispersion, respectively. In order of increasing impor-
tance, maxillary alveolar width, anterior skull width, interorbital width
and dentary alveolar width are the most influential characters providing
separation on the first axis (Table lb). Temporal arch width, jugal-
quadratojugal length and lingual alveolar width contributed most to
separation on the second axis (Table lb). All individuals were classified
into their assigned groups. Results from analysis of variance show New
River skulls significantly different from all taxa (P < 0.05) except P. c.
concinna and P. c. suwanniensis (canonical means enclosed by broken
lines in Fig. 5).
A third discriminant analysis of skulls, with P. f floridana and P.f
peninsularis removed, was applied to further clarify relationships with
the races of P. concinna (Fig. 6). The first two canonical axes collec-
tively account for 77% of the total dispersion. Characters most respon-
sible for separation on the first two axes are presented in Table lc.
Results from this comparison indicate a very close phenetic relationship
between the eastern river cooter, P. c. concinna (C) and New River
Pseudemys (B) (Fig. 6).
Observations on reproduction of cooters recently collected from
Bluestone Reservoir, discovery of a previously overlooked museum
record, and examination of archeological material have also contributed
to a better understanding of Pseudemys in the New River. Three adult
females from Bluestone Reservoir, collected 15 May 1980 and X-rayed 7
June (following the method of Gibbons and Greene 1979) showed no
Taxonomy of Pseudemyd Turtles
35
+
0
• P. floridana
O P.concinna
* New River Pseudemys
A P. rubri ventris series
'• «TS7
00
-o**°
C? O Q O OH .Yj
TcP ^
K,-
- 0 +
Fig. 4. Variates for skulls of the subgenus Pseudemys plotted on the first (K i)
and second (K2) canonical axes. Individual skulls represented by symbols, and
letters represent canonical variate means for P. alabamensis (A), New River
Pseudemys (B), P. c. concinna (C), P. f. floridana (F), P. c. hieroglyphica (H),
P. c. mobilensis (M), P. nelsoni (N), P. f. peninsularis (P), P. rubriventris (R),
P. c. suwanniensis (S), P. c. texana (T), P. f. hoyi (Y). Values for New River
Pseudemys connected by solid lines. Broken lines connect the most dispersed
values for P. rubriventris and P. nelsoni collectively, and values for P. alabamen-
sis.
36 Michael E. Seidel
K, - + 0 -
Fig. 5. Variates for skulls of P. floridana and P. concinna plotted on the first
(Ki) and second (K2) canonical axes. Individual skulls represented by symbols,
and letters represent canonical variate means for New River Pseudemys (B), P.
c. concinna (C), P. f. floridana (F), P. c. hieroglyphica (H), P. c. mobilensis (M),
P. f. peninsularis (P), P. c. suwanniensis (S), P. c. texana (T), P. f. hoyi (Y).
Broken lines connect and enclose New River skulls and plots for the subspecies
P. c. concinna and P. c. suwanniensis, which show no significant difference (P >
0.05) from New River Pseudemys.
Taxonomy of Pseudemyd Turtles
37
Ki-
+ 0 -
Fig. 6. Variates for skulls of P. concinna plotted on the first (Ki) and second
(K2) canonical axes. Lines connect the most dispersed values about canonical
means, which are represented by letters.
38 Michael E. Seidel
evidence of ripe (shelled) eggs. One of these females, dissected on 9 July,
contained no oviducal eggs. A second female held in laboratory depos-
ited eight eggs 10-21 July, three of which were incubated at 30-35° C in
moist vermiculite. Two hatched on 4 September and the third was
apparently infertile. Unlike P. rubriventris the two hatchlings have yel-
low plastrons, and unlike P. floridana the plastrons are extensively pat-
terned (Fig. 2). Dark markings also appear on the ventral surface of each
marginal (submarginal) and throughout the bridge. These hatchlings
and adults, maintained in laboratory and offered a wide variety of food
for three months, were entirely herbivorous. A juvenile P. concinna
(UMMZ 88488, fluid preserved), collected 3.2 km east of Hinton in the
Greenbrier River (near its junction with the New River below Bluestone
Reservoir), Summers County, West Virginia, has markings nearly iden-
tical to the Bluestone hatchlings. This fluid preserved specimen, pre-
sumably overlooked by other investigators, was taken in 1934 (collector
unknown) and confirms the presence of cooters in the New River system
prior to the construction of Bluestone. Dam, 1942-1949. Thus, the
hypothesis presented by Bayless (1972) that the impoundment created
new habitat necessary for establishment of an introduced population
can be dismissed. Further evidence for a relatively long natural history
of Pseudemys in the New River comes from examination of turtle bones
recovered from an archeological site (46SU3, Fort Ancient Village 1 100-
1300 AD) at Bluestone Reservoir (Applegarth et al. 1978). A peripheral
ACE FS2-81, pleural ACE FS2-183, and hypoplastron FS1-68 were
identified as Pseudemys cf. P. concinna by Dale R. Jackson (pers.
comm.).
DISCUSSION
The cooter species P. concinna and P. floridana have had a long,
confused taxonomic history. Following LeConte's (1830) original des-
criptions, Carr (1935, 1952) considered P. concinna and P. floridana a
conspecific assemblage of eight subspecies under P. floridana. However,
Carr (1952) commented that at least the Florida races, P.f peninsularis
and P. f suwanniensis, were broadly sympatric and behaved as separate
biological species. Crenshaw (1955) partitioned the cooters, recognizing
both P. concinna and P. floridana. A major weakness of Crenshaw's
conclusion is that it was based primarily on relationships between Flor-
ida forms, without a thorough understanding of ecological and morpho-
logical relationships of populations elsewhere. Another criticism of
Crenshaw's work is that it relied heavily on highly variable characters
such as markings and pigmentation, which may show greater variation
within a population than between species. These phenotypic characters
may also be subject to strong environmental influence. Mount (1975)
reported that the color of markings fades rapidly in captive P. concinna
and concluded that pigmentation in cooters has little systematic value.
Taxonomy of Pseudemyd Turtles 39
Moreover, Ewert (1979) demonstrated that, at least in map turtles of the
genus Graptemys, diagnostic head markings may be altered by varying
incubation temperatures and therefore are not entirely under genetic
control. Nevertheless, the taxonomic arrangement of Crenshaw (1955),
although never published with full supporting documentation, has gen-
erally been followed (Conant 1961, 1975; Ernst and Barbour 1972;
Wermuthand Mertens 1961, 1977).
Fahey (1980) proposed that P. concinna once again be placed in the
synonymy of P. floridana, a conclusion based exclusively on examina-
tion of turtles from Louisiana. Although Fahey's results suggest the
presence of a single species in the restricted region of his study, they
certainly do not substantiate relationships throughout the ranges of P.
floridana and P. concinna. In addition to Fahey's report, there are
numerous published references to either weak morphological separation
or putative hybridization between P. floridana and P. concinna in the
Mississippi Valley and areas to the west (Brown 1950; Smith 1961; An-
derson 1965; Webb 1970; Barbour 1971; Minton 1972; Mount 1975).
Traditional key characters, such as plastral markings and a "C" figure
on the second pair of pleurals, which are used to distinguish P. concinna
from P. floridana, are not consistent for central and western popula-
tions that are seemingly convergent in some characters. Ward (1980)
found no cranial characters with which he could separate the midwes-
tern subspecies P. f. hoyi and P. c. hieroglyphica and placed them in
synonymy. My results from discriminant analysis of cranial morphology
(Figs. 4, 5, and 6) support that decision. However, conclusions regard-
ing conspecific status for all turtles assigned to P. concinna and P. flori-
dana, especially eastern forms, must await a comprehensive and geogra-
phically broad analysis.
Based on shell markings and proportions, overall pigmentation,
and cranial morphology, New River Pseudemys are clearly distinct from
P. rubriventris. The emarginate tomial surface of New River specimens
(Fig. 2) might be interpreted as a weakly developed notch bordered by
cusps, characteristic of species in the rubriventris series (Carr 1952;
Ernst and Barbour 1972). Weak cusps, however, have been reported in
several populations of cooters allopatric and sympatric with redbelly
turtles (Carr 1952; Crenshaw 1955; and personal examination of skulls:
CM 60560, UMMZ 127058, MCZ 54680). Furthermore, prominent
cusps are typical in P. c. texana (Ernst and Barbour 1972). Jackson
(1978) cautioned that little taxonomic weight should be given to trophic
structures in Pseudemys. The overall similarity in cranial morphology of
P. c. texana, P. nelsoni, and P. rubriventris (Fig. 4) may represent con-
vergence of character states resulting from similar feeding habits.
A shallow carapace with evidence in some individuals of a "C" on
pleural II, and extensive dark markings on plastral, axillary and ingui-
nal scutes, are characteristics of New River cooters that justify their
40 Michael E. Seidel
assignment to P. concinna. Further indication that these turtles are ref-
erable to P. concinna comes from discriminant analysis of cranial morph-
ology. Skulls of New River turtles are phenetically close to P. c. con-
cinna and P. c. suwanniensis, and broadly separated from all forms of
P. floridana (Fig. 5).
Recent evidence of fertile eggs, numerous sight records of basking
adults, discovery of bones 700 to 800 years old, rediscovery of a
museum specimen collected in 1934, and extensions of the known range
22 km south of Bluestone Reservoir to Giles County, Virginia and 2 km
northeast of Bluestone Reservoir in the Greenbrier River, West Virgi-
nia, support the hypothesis that cooters in the New River represent a
natural, established population. Assuming an origin not involving
human interference, there are two possible avenues for dispersal that
could have allowed P. concinna to enter the New River. The first is an
early Pleistocene invasion from the Mississippi Valley (embayment
region) through the old Teays River, which followed the course of the
present Kanawha and New Rivers (Fig. 1). An argument for this route
is supported by the occurrence of cooters referrable to the Mississippi
Valley subspecies P. c. hieroglyphica in the lower Kanawha River,
Mason County, and Mud River, Cabell County, West Virginia (Seidel
and Green, in press). To examine this relationship, twelve New River
specimens were compared to seven adult and two subadult P. c. hiero-
glyphica from Reelfoot Lake, Tennessee. New River P. concinna differ
from these turtles in having: a lower carapace with the highest point at
the middle; shell not usually constricted at the region of the sixth mar-
ginal; fewer concentric light lines on the carapace; dark markings on the
bridge that frequently contact submarginal blotches; alveolar surface of
lower jaw relatively narrow; and fewer and broader stripes on the head
and neck, especially ventrally. These characteristics are more typical of
the Piedmont subspecies, P. c. concinna (Carr 1952; Mount 1975).
Although the New River is a Kanawha River tributary, it is separated
from the Ohio and Mississippi River valleys (inhabited by P. c. hiero-
glyphica) by Kanawha Falls (Fig. 1). This falls is believed to be one of
the largest natural river barriers east of the Rocky Mountains (Jenkins
et al. 1971). Furthermore, rapids and cataracts in New River Gorge just
above Kanawha Falls provide an effective barrier to fish distribution
between the Kanawha and New Rivers (Hocutt et al. 1979). Therefore,
preglacial dispersal of cooters from the Mississippi Valley into the upper
Teays (New) River may not have been possible.
A second potential avenue for dispersal of P. concinna is over the
Atlantic-Ohio divide of Virginia and North Carolina. Wright (1934),
Thompson (1939), Dietrich (1959), and Ross (1969), reported late Pleis-
tocene stream captures of the New River by the James and Roanoke,
two rivers inhabited by P. c. concinna in the Piedmont (Martof et al.
1980; and Fig. 1). Comparisons of New River P. concinna to five adult,
Taxonomy of Pseudemyd Turtles 41
fluid preserved P. c. concinna from the Piedmont of North Carolina
indicate an overall greater similarity in markings and shell shape than
seen comparing New River cooters to P. c. hieroglyphica. This relation-
ship is also supported by cranial morphology. In Figures 4-6, New River
specimens clearly plot closer to P. c. concinna than to P. c. hierogly-
phica. Although the subspecies of river cooters are not well defined and
their distinguishing characteristics are inconsistent within and between
populations (Mount 1975; and pers. observ.), Pseudemys in the New
River of Virginia and West Virginia are most similar morphologically to
the eastern river cooter and are here assigned to P. c. concinna. There-
fore, I suggest that during the Pleistocene, P. c. concinna ranged farther
up Piedmont streams in Virginia and North Carolina and gained access
to the New River through stream capture. The presence of the eastern
painted turtle, Chrysemys picta picta, in the upper Tennessee (Ernst
1970) and New River systems offers additional evidence that this corri-
dor has been used in the dispersal of aquatic turtles. Chrysemys p. picta
is typically an Atlantic Slope subspecies that is replaced by the midland
painted turtle, C. p. marginata, in the Ohio River system. Two speci-
mens from Mercer County, West Virginia (WVBS 4238, 4415) and two
specimens from Summers County, Bluestone Reservoir (MES 866, 868)
are referrable to the eastern subspecies. Two additional specimens from
Bluestone Reservoir (MES 867, 869) have characteristics typical of C. p.
picta xp. marginata intergrades. River cooters are highly aquatic turtles
and less likely than painted turtles to enter a new drainage by terrestrial
migration (Ernst and Barbour 1972). However, the preference of P. c.
concinna for rocky, fast-running stream habitats (LeConte 1836; Carr
1952; Pritchard 1979) might have facilitated its dispersal through small-
stream captures.
ACKNOWLEDGMENTS.— I am grateful to Carl H. Ernst,
George Mason University, for directing my attention to the taxonomic
problem of Pseudemys in the New River. I thank Laurence E. Bayless,
Concord College; Michael Little, Marshall University; and Samuel L.
Reynolds, Memphis State University, for help in collecting specimens.
The following individuals made this study possible by loan of speci-
mens: C. J. McCoy, Carnegie Museum of Natural History; Arnold
Kluge, University of Michigan Museum of Zoology; Ernest E. Williams
and Jose P. O. Rosado, Museum of Comparative Zoology, Harvard
University; George Zug and Roy McDiarmid, National Museum of
Natural History; William M. Palmer, North Carolina State Museum of
Natural History; Joseph T. Collins, University of Kansas Museum of
Natural History; Hymen Marx, Field Museum of Natural History; N.
Bayard Green, West Virginia Biological Survey; Robert Maslowski, U.
S. Army Corps of Engineers, Huntington District; and Richard Zweifel
and Michael Klemens, American Museum of Natural History. Appreci-
42 Michael E. Seidel
ation is extended to Dale R. Jackson, University of South Florida, for
identification of bones from archeological remains, and to Vickie Crager
for typing the manuscript. Partial support for the study was provided by
a Marshall University faculty research grant.
LITERATURE CITED
Anderson, Paul. 1965. The Reptiles of Missouri. Univ. Missouri Press, Colum-
bia. 330 pp.
Applegarth, J. D., J. M. Adovasio and J. Donahue. 1978. 46SU3 Revisited.
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Barbour, Roger W. 1971. Amphibians and Reptiles of Kentucky. Univ. Press
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Virginia. J. Herpetol. (5:39-41.
Brown, Bryce C. 1950. An annotated check list of the reptiles and amphibians of
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1952. Handbook of Turtles. Cornell Univ. Press, Ithaca. 542 pp.
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, and R. W. Barbour. 1972. Turtles of the United States. Univ. Press
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22( l):49-66
Taxonomy of Pseudemyd Turtles 43
Gibbons, J. Whitfield, and J. L. Greene. 1979. X-ray photography: A tech-
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Gauley River, West Virginia. Brimleyana 1:47-80.
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Appalachians, Part III: Vertebrates. Res. Div. Monogr. 4, Va. Polytech.
Inst. State Univ., Blacksburg. 306 pp.
LeConte, John. 1830. Description of the species of North American tortoises.
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southern Appalachians of Virginia, pp. 277-292 in P. C. Holt (ed.). The
distributional history of the biota of the southern Appalachians, Part I:
Invertebrates. Res. Div. Monogr. 1, Va. Polytech. Inst., Blacksburg. 295
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Seidel, Michael E., and N. B. Green. In press. On the occurrence of cooter
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family Emydinae (Testudines: Emydidae). Part I, Chrysemys, Pseudemys
and Trachemys. Annual Meeting Am. Soc. Ichthyol. Herpetol., Fort
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44 Michael E. Seidel
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Accepted 2 November 1981
Small Mammals in Openings in Virginia's Dismal
Swamp
Robert K. Rose
1
Department of Biological Sciences,
Old Dominion University, Norfolk, Virginia 23508
ABSTRACT. — In a study of small mammals of openings in the
Dismal Swamp of Virginia, seven species were obtained using pitfall
traps. Samples included several species rarely caught in the Swamp
— seven specimens of the Dismal Swamp subspecies of the southern
bog lemming, Synaptomys cooperi helaletes, the first collected in this
century; two least shrews, Cryptotis parva; and 15 southeastern shrews,
Sorex longirostris fisheri. Results are compared to previous studies,
conducted primarily in forested habitats, in which the white-footed
mouse, Peromyscus leucopus, and the golden mouse, Ochrotomys
nuttalli, were numerically dominant.
INTRODUCTION
The Great Dismal Swamp, originally occupying much of the area
between Virginia's James River drainage system and North Carolina's
Albemarle Sound, has long been recognized as a vegetationally distinc-
tive region with many unusual features. It has been subjected to clear-
ing, burning, ditching, farming, and other land-use practices during the
past 250 years, has long experienced a dropping water table, and is now
approximately 850 km2 (85,000 ha) in extent (Carter 1979). In 1974 the
Great Dismal Swamp National Wildlife Refuge (GDSNWR) was estab-
lished. At the end of 1980 it was 41,026 ha in extent, 24 percent (9,866
ha) of it in North Carolina. Kirk (1979) provided an excellent summary
of the history and lore of the Swamp.
Although there are conflicting historical reports about the abun-
dance of wildlife in the Swamp (see Handley 1979), it is clear that the
survival of some species there has been aided by remoteness and limited
access to the public, as well as by the existence of large tracts of suitable
habitat. For example, the only population of the black bear, Ursus amer-
icanus americanus Pallas, on the Virginia Coastal Plain, and perhaps
the largest populations of the bobcat, Lynx rufus floridanus Rafinesque,
are found in the Refuge and environs. However, its remoteness and rela-
tive inaccessibility have also apparently contributed to the dearth of
studies of birds and mammals in the Swamp; apart from species lists,
comparatively little is known about the wildlife.
The first systematic studies of mammals in the Swamp were con-
ducted by the Bureau of Biological Surveys, U. S. Department of Agri-
culture, during the period 1895-1898. Handley (1979), in an exhaustive
review that included an examination of field notebooks and unpub-
Brimleyana No. 6: 45-50. December 1981. 45
46 Robert K. Rose
lished manuscripts, reported that collections of Dismal Swamp mam-
mals were made during a total of 23 weeks in that period. A number of
new species (now recognized as subspecies) were collected then, mostly
near Lake Drummond. The greater short-tailed shrew, Blarina brevi-
cauda telmalestes Merriam; southeastern shrew, Sorex longirostris
fisheri Merriam; southern bog lemming, Synaptomys cooperi helaletes
Merriam; and muskrat, Ondatra zibethicus macrodon (Merriam), were
collected and named then. During the same period Rhoads and Young
(1897) described the dark-colored meadow vole, Microtus pennsylvani-
cus nigrans Rhoads, from nearby northeastern North Carolina. In sum,
the Dismal Swamp and nearby coastal marshes have several mammals
that are morphologically distinguishable from other populations of
these species, strongly suggesting genetic and perhaps geographic isola-
tion of their populations in the past. One of Handley's (1980) concerns
was that man-induced changes in the Swamp may have removed the
ecological barriers between Swamp and upland subspecies. The likely
result of such an event would be Loss of the Dismal Swamp subspecies
through genetic "swamping out" of the smaller gene pool. Of course,
this is the equivalent of ecological extinction of the taxon.
C. S. Brimley (1897) was among the investigators who wrote about
Dismal Swamp mammals, for he included the results of small mammal
collections made between 1891-1894 by the Smithwick brothers near the
head of Albemarle Sound in his history of the mammals of Bertie
County, North Carolina. Brimley later (1905) summarized the findings
of several investigators, including collections from the northeastern
corner of North Carolina close to the Swamp. Both papers, while
including some Dismal Swamp information, were based mostly on small
mammals that Brimley and his brother collected near Raleigh, Wake
County, from 1888 to 1900.
After a hiatus of about 25 years, sporadic collecting in the Swamp
resumed in the 1930s. Handley's 1953 visits for a week each in February
and June seem to have been typical of the trapping efforts made there.
One of the longer mammal studies conducted in the Swamp was that of
F. E. Breidling, Old Dominion University (ODU), who in 1979-1980
trapped four study areas for one week during each of three seasons.
Handley (1979) reported that the entire known Dismal Swamp
fauna of mice and shrews consists of 12 species. Most investigators have
found the white-footed mouse, Peromyscus leucopus easti Paradiso, to
be the most common small mammal, and about half of them have also
caught numerous golden mice, Ochrotomys n. nuttalli (Harlan), and
short-tailed shrews, Blarina brevicauda. Five other species — the cotton
mouse, Peromyscus g. gossypinus (Harlan); eastern harvest mouse, Rie-
throdontomys h. humulis (Audubon and Bachman); southern bog lemm-
ing; and southeastern shrew — were found to be numerous by only one
or two collectors. Handley (1979) attributed this to spotty distributions
Dismal Swamp Small Mammals 47
and local abundances. Finally, four species — the woodland vole, Micro-
tus pine tor um scalopsoides (Audubon and Bachman); the house mouse,
Mus musculus domesticus Rutty; the least shrew, Cryptotis p. parva
(Say); and the meadow vole — were seldom caught by any collector,
which may mean that the habitats required by these species are rarely
found in the Swamp (Handley 1979).
On 23 February 1980, David Harrelson, a senior biology student at
ODU, and I began a study of the small mammals of openings in the
GDSNWR. The term "openings" refers to any area in which a signifi-
cant level of shading provided by tree canopy is absent. These habitats
are vegetated predominantly by cane, Arundinaria gigantea; softstem
rush, Juncus effusus; sedges; grasses; and herbaceous forbs. Many open-
ings also have small trees and shrubs, plus a number of woody vines; the
most common of these are red maple, Acer rubrum; blackberry, Rubus
allegheniensis; grape, Vitis spp.; and greenbriers, Smilax spp.
MATERIALS AND METHODS
Pitfall traps, made of No. 10 tin cans sunk into the ground flush
with the soil surface and half-filled with water, were used to collect
small mammals. Seven pitfall traps were dug and placed on 23 Febru-
ary, but adverse weather, including a record snowfall that covered the
area for the first two weeks in March, delayed until 20 March the set-
ting of twenty-eight additional traps. All 35 traps were set within 150 m
of Jericho Ditch, north of Williamson Ditch, under the 1 10 kv electrical
powerline in the northwestern corner of the GDSNWR.
On 10 April, 10 pitfall traps were placed 9 km away, under the
same powerline near East Ditch, also in an area dominated by cane,
grasses, and rushes. This area had a higher proportion of standing water
than did the Jericho Ditch site. All traps were removed from the ground
on 2 May 1980.
RESULTS
Only one small mammal, a Microtus pennsylvanicus, was captured
in the seven traps from 23 February to 20 March. However, a total of
43 small mammals of seven species was trapped during the study period
at the Jericho Ditch site (Table 1). At the East Ditch site, three small
mammals of three species were caught (Table 1).
Based on the number of small mammals captured in 100 trap-
nights, the relative density of small mammals appeared to be greater in
the Jericho Ditch area (2.43) than in the East Ditch area (1.36). (One
trap in place for one night equals one trap-night; relative density =
N/trap nights X 100.) This difference in density may be due in part to
the greater vegetational diversity of the Jericho Ditch site, and to the
greater proportion of standing water on the East Ditch site.
48
Robert K. Rose
Table 1. Number and species of small mammals trapped in the Dismal Swamp
between 23 February and 2 May (Jericho Ditch area) and 10 April and
2 May (East Ditch area) 1980. "Others" refers to the results of previous
investigations in the Dismal Swamp, mostly in the 1895-1906 period,
but including Handley in 1953 (from Handley 1979, Table 1).
43
Number mammals/ 100
trapnights
2.43
1.36
DISCUSSION
Compared to previous investigators, we caught few individuals of
the two most common species, Peromyscus leucopus and Ochrotomys
nuttalli. This is not unexpected, because they are predominantly forest-
dwellers and we restricted our trapping to openings dominated by her-
baceous vegetation. However, the 40 m wide powerline right-of-way was
bordered on both sides by maple-gum forest. Consequently, the proxim-
ity to nearby suitable habitat for these climbing species may explain
their presence in the openings. Handley's 1953 study, which produced 34
P. leucopus and 14 O. nuttalli out of a total of 56 specimens, showed
the typical numerical dominance of these two species. Breidling (1980)
caught 15 P. leucopus and 4 Ochrotomys using live traps on four for-
ested study plots.
We caught a relatively large number of meadow voles and southern
bog lemmings (Table 1). Only 29 individuals of these two microtine spe-
cies had previously been collected in the Swamp. Although he took one
meadow vole in 1953, Handley (1979) contended that Synaptomys had
not been collected there since November 1898. According to Handley
(1979, 1980) several investigators, including himself, have speculated on
Dismal Swamp Small Mammals 49
the likely extinction of the Dismal Swamp subspecies of the southern
bog lemming, Synaptomys cooperi helaletes. We took specimens from
both sides of Jericho Ditch, and one specimen near East Ditch. The
cane-grass-sedge vegetation type is dominant under the powerline, and
it is possible that S. c. helaletes occurs throughout this habitat. Starting
in 1895, Fisher caught 21 specimens of southern bog lemming in the
Swamp, mostly in cane patches near Lake Drummond. We took one
Synaptomys in cane, but the remainder were captured in mixed grass-
land in which softstem rush was abundant. Meadow voles were present
in the mixed grass habitat, but not in the cane.
By far our greatest success was in trapping shrews (Table 1). We
captured 15 Sorex longirostris fisheri, which is as many as had been
obtained by all previous investigators (Handley 1979). We caught 2
specimens of the least shrew, Cryptotis parva, compared to 1 taken by
previous investigators, and 15 Blarina, compared to 39 collected in ear-
lier studies. Our comparatively high success in capturing shrews is prob-
ably related to use of pitfall traps. An advantage of pitfall traps is that
they more readily capture certain species of small mammals than do
snap (or break-back) traps (Rose and McKean 1980). Rose (1980)
reported the capture of 18 southeastern shrews in pitfall traps and none
in snap traps. The conclusion that southeastern shrews are not effec-
tively taken by snap (or live) traps is borne out by published records
(reviewed by Rose 1980; French 1980).
Handley's (1979) fears that S. longirostris fisheri has been geneti-
cally "swamped out" through introgression with the smaller upland S. I.
longirostris Bachman may be unfounded, at least for populations in the
northwestern corner of the Swamp in 1980. With a mean total length of
95.8±2.3 mm, the 1980 Dismal Swamp southeastern shrews are much
longer than any of the upland subspecies (French, pers. comm.).
Whether these values are larger than the 1890s S. I. fisheri is uncertain,
for Handley (1979:310, Table 1) did not give standard measurements for
the 15 S.l. fisheri collected by Fisher and housed in the National
Museum, nor have I examined the specimens. Nevertheless, the large
size of the 1980 specimens suggests that S. I. fisheri has maintained
genetic isolation from S. I. longirostris.
Similarly, the Blarina were large and undoubtedly referrable to B.
brevicauda telmalestes, which Handley (1979) called the greater short-
tailed shrew. Jones et al. (1979) referred to the taxon as Blarina telma-
lestes, the Dismal Swamp short-tailed shrew. This disparity of usage
correctly indicates that the taxonomy of the genus Blarina is in flux.
According to Tate et al. (1980) the Dismal Swamp is one region in
which two distinctive sizes of Blarina occur, perhaps sympatrically; the
larger is B. brevicauda and the smaller B. carolinensis.
The total number of species trapped in our study — seven —
compares well with previous studies. Handley obtained the same
50 Robert K. Rose
number in 1953, and of the four other studies he summarized (1979,
Table 1), two obtained more species (8 and 9) and two obtained fewer
species (4 and 6). Considering that this study was conducted at the end
of winter, when mammal densities are usually at their lowest levels, it
seems likely that other seasonal studies of the openings in the Dismal
Swamp will provide additional useful information.
ACKNOWLEDGMENTS.— I gratefully acknowledge the field assist-
ance of David Harrelson, the cooperation of Refuge Manager R. Keel
and Wildlife Biologist M. K. Garrett, and the help of C. O. Handley,
Jr., primarily through his exhaustive review of studies of mammals in
the Dismal Swamp but in other ways as well.
LITERATURE CITED
Breidling, F. E. 1980. An evaluation of small rodent populations in four Dismal
Swamp plant communities. M. S. Thesis, Old Dominion Univ., Norfolk.
47 pp.
Brimley, C. S. 1897. An incomplete list ©f the mammals of Bertie Co., N.C. Am.
Nat. 57:237-238.
. 1905. A descriptive catalogue of the mammals of North Carolina,
exclusive of the Cetacea. J. Elisha Mitchell Sci. Soc. 27:1-32.
Carter, Virginia. 1979. Remote sensing applications to the Dismal Swamp.
pp. 80-100 in Paul W. Kirk, Jr. (ed.). The Great Dismal Swamp. Univ.
Press Virginia, Charlottesville. 427 pp.
French, Thomas W. 1980. Natural history of the southeastern shrew, Sorex
longirostris Bachman. Amer. Midi. Nat. 104(\): 13-31.
Handley, Charles O., Jr. 1979. Mammals of the Dismal Swamp: a historical
account, pp. 297-357 in Paul W. Kirk, Jr. (ed.). The Great Dismal Swamp.
Univ. Press Virginia, Charlottesville. 427 pp.
. 1980. Mammals, pp. 483-621 in Donald W. Linzey (ed.). Endangered
and Threatened Plants and Animals of Virginia. Center for Environmental
Studies, VPI & SU, Blacksburg, Virginia. 665 pp.
Jones, J. Knox, Jr., D. C. Carter and H. H. Genoways. 1979. Revised checklist
of North American mammals north of Mexico, 1979. Occas. Pap. Mus.
Texas Tech. Univ. 62:1-17.
Kirk, Paul W., Jr. (ed.). 1979. The Great Dismal Swamp. Univ. Press Virginia,
Charlottesville. 427 pp.
Rhoads, Samuel N., and R. T. Young. 1897. Notes on a collection of small
mammals from northeastern North Carolina. Proc. Acad. Nat. Sci. Phila.
(1897):303-312.
Rose, Robert K. 1980. The southeastern shrew, Sorex longirostris, in southern
Indiana. J. Mammal. 67:162-164.
, and R. McKean. 1980. Habitat associations of small mammals in
southwestern Indiana. Proc. Indiana Acad. Sci. #9:432-439.
Tate, Cathy M., J. F. Pagels and C. O. Handley, Jr. 1980. Distribution and
systematic relationship of two kinds of short-tailed shrews (Soricidae:
Blarina) in south-central Virginia. Proc. Biol. Soc. Wash. 9J(l):50-60.
Accepted 3 July 1981.
A New Milliped of the Genus Brevigonus from South Carolina,
with Comments on the Genus and B. shelfordi (Loomis)
(Polydesmida: Xystodesmidae)
Rowland M. Shelley
North Carolina State Museum of Natural History,
P.O. Box 27647, Raleigh, North Carolina 27611
ABSTRACT. — Brevigonus arcuatus, new species, is characterized by
a broadly curved acropodite in which the distal zone is moderately
long and possesses either simple or reflexed tips. Its congener, B. shel-
fordi (Loomis), is distinguished by absence of a distal zone and apical
curve, and its abbreviated acropodite terminates at the distal extremity
of the peak. An improved generic diagnosis is possible. Brevigonus is
characterized by the spine at the base of the acropodite, and by the
proximal location of the medial flange, which arises on the basal zone
and ends on the peak and separates Brevigonus from the related genus
Sigmoria.
In 1980 I attempted to dispose of a long standing taxonomic prob-
lem in the Xystodesmidae by erecting the new genus Brevigonus for
Cleptoria shelfordi Loomis. Hoffman (1967) removed this species from
Cleptoria but did not assign it to another genus. Before publication I
collected extensively in and near the range of shelfordi, the north side of
the Savannah River in the Piedmont Plateau of South Carolina, to try
to discover other species for this genus. Finding none, however, I reluc-
tantly concluded that there was no alternative to a monotypic taxon.
Two basic gonopodal variants of shelfordi were at hand, but they
clearly were not reproductively isolated. Consequently, I proposed the
genus Brevigonus to emphasize what I considered the most distinctive
feature of shelfordi, its shortened gonopodal acropodities.
I did not know in 1980, however, that a form I was referring to
Sigmoria was actually closely related to shelfordi. I had collected this
species several times in piedmont South Carolina but had assigned it to
Sigmoria because of the overall curvature of the acropodite. Not until I
was well into revising Sigmoria did I realize that this species was conge-
neric with shelfordi, and by that time the Brevigonus paper had been
published. The new species shares several features with shelfordi that
make for a sound generic diagnosis, but shortness of the gonopodal
acropodites is unfortunately not one of them. This trait is a specific
characteristic of shelfordi', the acropodites of the new species are long
and form a broadly curved arch, which is the basis for its specific name,
arcuatus. Brevigonus was thus a regrettable choice for a generic name,
since it is based on a derived character of only one species and not on a
trait shared by all components of the genus. A better name would have
emphasized the spines on the basal zones (see terminology of the acro-
Brimleyana No. 6: 51-60. December 1981. 51
52 Rowland M. Shelley
podite section in Shelley 1981a) or the proximal locations of the medial
flanges, which originate on the basal zones in shelfordi and arcuatus.
Although an inappropriate name, however, Brevigonus was validly pro-
posed and must be retained for the taxon encompassing these two
species.
There are positive aspects to the belated inclusion of arcuatus in
Brevigonus. Improved diagnoses of the genus and of shelfordi are now
possible, and certain anatomical features of shelfordi can now be more
accurately interpreted. Moreover, arcuatus evinces a close relationship
between Brevigonus and Sigmoria as opposed to one between Brevigo-
nus and Cleptoria as previously stated (Shelley 1980). The significance
of the flange on the medial surface of the acropodite, for example, was
not evident when shelfordi was studied alone, because the shortened
acropodite of this species terminates at the peak and lacks the distal
zones and apical curves present in arcuatus and species of Sigmoria.
Brevigonus shelfordi is thus a modified species that lacks the distal 1/3
to 1/2 of the normal apheloriine acropodite. Consequently, the medial
flange of shelfordi did not previously show any resemblance to the
flanges of certain species of Sigmoria, but the similarity is obvious in
arcuatus, since it possesses the distal sections of the acropodite. Brevi-
gonus can now be partly defined as an apheloriine genus with a flange
on the medial face of the acropodite, arising on the proximal portion of
the basal zone and terminating on the proximal portion of the peak. It
differs from Sigmoria in the more proximal location of the flange,
which is located on the peak (arising at the anterior bend) or the distal
zone in this genus. The flange is variable in Brevigonus and is reduced
or vestigial in individuals of both species. The margin is also irregular,
and the flange may extend straight across the anterior bend as shown in
Figure 3, or curve parallel to the acropodite stem as shown in Figures
8-9 of shelfordi in my 1980 paper. The apparent distal lobes on the
acropodites of the latter individuals can now be recognized as the ter-
mination points of the medial flanges, which end on the distal portions
of the acropodites in shelfordi only because this structure is shortened.
Thus by clarifying the significance of the medial flange, arcuatus reveals
a close phylogenetic affinity between Brevigonus and Sigmoria, which is
confirmed by one other character — the reflexed tips on males in the
southern part of the range of arcuatus. These individuals, from Abbe-
ville County, South Carolina, have reflexed tips identical to those of
certain species of Sigmoria, for instance S. latior (Brolemann). No other
apheloriine taxa in the southeastern Atlantic lowlands display this ter-
mination of the acropodite, and its presence in species of Sigmoria and
Brevigonus is strong evidence of a close relationship between the two
genera. No such evidence exists of a relationship between Brevigonus
and Cleptoria. My comments to this effect in 1980 were based on what I
Millipeds of Genus Brevigonus
53
Fig. 1-5. Brevigonus arcuatus, new species. 1, process of 4th sternum of holo-
type, caudal view. 2, gonopods in situ, ventral view of paratype. 3, telopodite of
left gonopod of holotype, medial view. 4, the same, lateral view. 5, telopodite of
left gonopod of male from 5.8 km. SE Lowndesville, Abbeville Co., medial
view. Scale line for Fig. 2= 1.00 mm; line for other Figures = 1.16 mm for 1,
1.40 mm for 3-4, and 1.00 mm for 5.
54 Rowland M. Shelley
now realize are only coincidental similarities between the gonopods of
shelf or di and those of certain forms of Cleptoria, based on the shor-
tened acropodites and other derived features of shelfordi, which just
happen to resemble aspects of Cleptoria gonopods. Consequently, these
statements about a relationship between Brevigonus and Cleptoria must
now be discounted.
This paper presents amended diagnosis of both Brevigonus and
shelf ordi, a description of arcuatus, and new information on generic and
specific ranges. All specimens of arcuatus included in this study are
deposited in the North Carolina State Museum (NCSM) collection, the
invertebrate catalog numbers of which are shown in parentheses. Other
materials are in the collection of Richard L. Hoffman (RLH), Radford
University, Radford, Virginia.
Brevigonus Shelley
Brevigonus Shelley 1980:32-34.
Type species. — Cleptoria shelf ordi Loomis 1944.
Description. — The following comments on gonopods are supplemental
to the somatic description in my 1980 account, and present a parallel
treatment to the description of Sigmoria (Shelley 1981a), thus facilitat-
ing comparisons.
Gonopods in situ either crossing in midline or extending forward in
subparallel arrangement over anterior edge of aperture and between 7th
legs. Coxae large, without apophysis, connected by membrane only, no
sternal remnant. Prefemur generally large, with or without large, cuneate
process arising on dorsal side. Acropodite thick and heavy, either curv-
ing broadly through flattened peak into moderate distal zone and form-
ing arc with variable diameter, or terminating abruptly at distal extrem-
ity of peak, with distal zone and apical curve absent; basal zone
relatively long; anterior bend broad, moderate to poorly defined; peak
flattened to gently curved; apical curve broad, smoothly continuous
with peak; distal zone moderately long, curving broadly into arch of
acropodite and directed toward basal zone, tapering smoothly apically.
Termination variable; in forms with distal zone, either narrowly rounded,
simple tip, or reflexed tip; in forms without distal zone, broad, blunt,
occasionally notched tip. Basal sone usually with prominent basal spine
on ventral surface and flange of variable width and configuration on
medial face, arising proximally and continuing to termination on peak;
latter with or without small acute spur at termination point of flange.
Distal zone with remnant of lateral flange extending from proximal por-
tion to about 2/3 length, usually represented by thickened rim near
outer margin. Prostatic groove arising in pit on prefemur, crossing to
lateral side of acropodite at anterior bend and continuing to terminal
opening on simple or reflexed tips, or on distal extremity of peak.
Millipeds of Genus Brevigonus
55
Fig. 6. Distribution of Brevigonus: triangles, arcuatus; dots, shelfordi.
Range.— Western Piedmont Plateau of South Carolina, from eastern
Pickens and Oconee counties to central McCormick County. The genus
is best represented in the Savannah River Valley of McCormick, Abbe-
ville, and Anderson counties.
Species. — Two.
Brevigonus shelfordi (Loomis)
Cleptoria shelfordi Loomis 1944:172-173, Fig. 4. Chamberlin and Hoff-
man 1958:28.
Brevigonus shelfordi Shelley 1980:35-41, Figs. 1-13.
Diagnosis. — Distinguished by following features of male gonopods:
acropodites crossing in situ; prefemoral process present, cuneate; acrop-
odite short, terminating abruptly at distal extremity of peak, distal zone
and apical curve absent; with or without sharply acute spur on medial
face of acropodite distal to flange.
Description. — As with the generic description, the following com-
ments on gonopods supplement my 1980 description and present an
account parallel to those of species of Sigmoria.
Gonopods in situ with acropodites crossing at midline of aperture,
extending over opposite sides of aperture and projecting slightly across
56 Rowland M. Shelley
anterior margin. Prefemoral process short and subtriangular, cuneate.
Acropodite thick and heavy, curving into peak and terminating abruptly
at distal extremity of peak; distal zone and apical curve absent; peak
overhanging and extending usually beyond level of prefemoral process,
directed subperpendicularly to basal zone; basal zone long, usually
about 2/3 of acropodite length, with or without broad, caudally directed
spine basally on ventral margin; anterior bend variable, broad and
poorly defined to sharp and well defined; peak flattened to slightly
curved, relatively short, no more than 1/3 of acropodite length; distal
extremity of peak (termination of acropodite) variable — blunt, slightly
rounded, or indented with hood-like lobe overhanging basal projection.
Medial flange usually present, occasionally reduced and vestigial, aris-
ing on basal zone distal to spine, extending beyond anterior bend and
terminating on peak, margin irregular. Peak with or without sharply
acute spur on medial face distal to flange. Prostatic groove running
along medial face of basal zone, crossing to lateral side at anterior bend,
terminating on inner corner of basal projection of peak.
Remarks. — The specimens from Oconee County are now referred to
arcuatus, as mentioned in the ensuing account. Otherwise, the range of
shelfordi, as represented by the available material, remains unchanged
from 1980.
Since it is now apparent that the medial flange is homologous to
that of Sigmoria, the question arises as to whether the spur might be
homologous to the tooth of latior and other species of Sigmoria. This
seems plausible, but the material at hand provides no clues to resolve
the issue.
As mentioned, the acropodite of shelfordi is merely an extremely
shortened one in which the distal zone and apical curve are absent.
Given the preponderance of long, curved acropodites in the tribe Aphe-
loriini, such an abbreviated structure can only be interpreted as a
derived character.
Brevigonus arcuatus, new species
Figs. 1-5
Type specimens. — Male holotype (NCSM A2075) and 4 M and 2 F
paratypes collected by R. M. Shelley and W. B. Jones, 12 June 1978,
from Pickens Co., SC, 13.6 km (8.5 mi.) E Pickens, along SC highway
192 at George's Creek. Three M and 2 F, and 7 M and 1 F, paratypes
collected at same locality by R. M. Shelley on 8 May 1977 and 2 August
1977, respectively. Male and female paratypes deposited in Florida
State Collection of Arthropods and private collection of R. L. Hoffman.
Diagnosis.— Characterized by following features of male gonopods:
in situ arrangement parallel; prefemoral process absent; acropodite
long, curving distally into broad arch, apical curve and distal zone pres-
ent, without spur on peak; tip of distal zone variable, simple or reflexed.
Millipeds of Genus Brevigonus 57
Holotype.— Length 49.6 mm, maximum width 11.5 mm, W/L ratio
23.2%, depth/ width ratio 62.2%. Segmental widths (in mm) as follows:
collum 7.8 10th-13th 11.5
2nd 8.9 14th 11.3
3rd 10.1 15th 11.0
4th 10.4 16th 10.6
5th 11.0 17th 9.4
6th-9th 11.4 18th 6.9
Color in life: paranota bright red, color indented slightly mediad
along caudal edges; metaterga and collum glossy black, without stripes.
Somatic features similar to shelfordi (see Shelley 1980), with fol-
lowing exceptions: Width across genal apices 5.2 mm, interantennal
isthmus broad, 1.8 mm. Antennae moderately long and slender, reach-
ing back to caudal edge of 3rd paranota, relative lengths of antenno-
meres 2>3>4 = 5>6>1>7. Facial setae as follows: epicranial, interan-
tennal, frontal, and genal absent, clypeal about 10-10, labral about
14-14.
Dorsum very glossy, slightly coriaceous on anterior portions of
paranota. Latter moderately depressed, subcontinuous with slope of
dorsum. Collum very broad, extending well beyond ends of following
tergite. Caudolateral corners of paranota rounded through segment 7,
becoming blunt and progressively more acute thereafter.
Process of 4th sternum (Fig. 1) enormous, much longer than widths
of adjacent coxae; processes of 5th sternum large, knobs between 4th
legs subequal to widths of adjacent coxae, flattened areas between 5th
legs produced into knobs, shorter than widths of adjacent coxae; ster-
num of segment 6 slightly elevated into two small lobes between 6th
legs, convexly recessed between 7th legs to accommodate curvatures of
acropodites, 7th legs set slightly farther apart than 6th. Postgonopodal
sterna bilobed on segments 8-10, with thick patches of stiff setae on
lobes, becoming flattened with varying shallow grooves and impressions
and fewer setae posteriorly. Coxae with low tubercles beginning on
segment 10, becoming spine-like on 14 and continuing posteriorly.
Gonopodal aperture elliptical, 4.1 mm wide and 2.2 mm long at
midpoint, strongly indented on anteriolateral edges, sides thickened and
elevated above metazonal surface. Gonopods in situ (Fig. 2, of para-
type) with acropodites in subparallel arrangement, extending forward
over anterior edge of aperture between 7th legs, not overlapping or
touching. Gonopod structure as follows (Fig. 3-4): prefemoral process
absent, with elevated setose ridge at location of process but without
sclerotized projection. Acropodite moderately thick and heavy, curving
distally into broad arch, flattened at peak, overhanging and extending
well beyond level of prefemur; basal zone with large, caudally directed
58
spine basally on ventral surface and small basal lobe on inner surface
above prefemur; anterior bend moderately sharp, located at nearly 1/3
length; peak relatively flat, about 1/4 of acropodite length; apical curve
present, broad, located at about 3/4 length; distal zone present, long,
curving broadly into arch; tip narrowly rounded, not reflexed, directed
toward basal zone. Medial flange arising on basal zone just distal to
spine, extending across anterior bend and terminating at midlength of
peak, edge curved inward proximally and outward distally, obscurring
short section of acropodite stem at anterior bend. Spur absent. Lateral
flange present but greatly reduced, forming short rim on distal zone.
Prostatic groove running along inner surface of acropodite basally,
crossing to lateral side at anterior bend and continuing to tip.
Male paratypes. — The male paratypes agree essentially with the holo-
type in all particulars.
Female par atype. — Length 48.0 mm, maximum width 11.4mm, W/L
ratio 23.8%, depth/ width ratio 67.5%. Agreeing closely with holotype in
somatic details except paranota more strongly depressed, creating
appearance of more highly arched body.
Cyphopods in situ with valves visible in aperture, receptacle situated
internally against coxae. Receptacle relatively small, located anteriad to
and not overlapping valves, surface finely granulate. Valves moderate,
inner one slightly larger, surface finely granulate.
Variation. — Several aspects of the gonopods vary. The size of the
spine changes, being generally smaller in material from Abbeville County
and greatly reduced in one male from Anderson County. The acropo-
dite tends to be thinner and the apical curve broader in the southern
specimens, and all males from Abbeville County posses a reflexed tip
(Fig. 5). The tip is simple on all other specimens. Specimens from And-
erson County have a sharp spine projecting mediad at the base of the
medial flange in addition to that at the base of the acropodite, but this
structure is absent from the Abbeville County males. All males have an
elevated ridge on the prefemur but lack a prefemoral process.
Ecology. — Brevigonus arcuatus occurs under thin leaf layers on rela-
tively hard substrates near rivers or creeks. It is sometimes found on the
vertical bank of streams and has rarely been taken more than 6 to 9 m
from a water source.
Distribution. — Piedmont Plateau physiographic province of central-
western South Carolina, from southeastern Pickens to southwestern
Abbeville counties. Specimens examined as follows (all collected by the
author unless otherwise stated):
SOUTH CAROLINA: Oconee Co.— Clemson vie, under dead
pig, 2M, F, 18 July 1962, J. A. Payne (RLH). Pickens Co.— 13.6 km.
(8.5 mi.) E Pickens, along SC hwy. 192 at George's Cr., 3M, 2F, 8 May
1977 (NCSM A 1559); 7M, F, 2 August 1977 (NCSM A1617); and 5M,
2F, 12 June 1978, R. M. Shelley and W. B. Jones (NCSM A2075)
Millipeds of Genus Brevigonus 59
TYPE LOCALITY. Anderson Co.— 12.6 km. (7.9 mi.) SE Anderson,
along SC hwy. 459 at Rocky R., M, F, 7 May 1977 (NCSM A1550);
10.1 km. (6.3 mi.) NE Iva, along SC hwy. 413 at Rocky R., 2M, 2F, 1 1
June 1978, R. M. Shelley and W. B. Jones (NCSM A2066); and 6.4 km.
(4 mi.) SW Iva, along unnumbered rd. off SC hwy. 187 at Generostee
Cr., M, 7 May 1977 (NCSM A1549). Abbeville Co.— 5.8 km. (3.6. mi.)
SE Lowndesville, along SC hwy. 232 at Deal Cr., M, F, 6 May 1977
(NCSM A1544); and 4.2 km. (2.6 mi.) SW Lowndesville, along SC hwy.
70. 0.5 km. (0.3 mi.) SW jet. SC hwy. 64, M, 6 May 1977 (NCSM
A 1545).
Remarks. — The unusually large process of the 4th sternum, the long-
est of any apheloriine milliped known to me, is one of the key features
of arcuatus. The structure is also longer than the widths of the adjacent
coxae in two species of Croatania (Shelley 1977), but in these forms it is
apically divided and bent anteriad. In arcuatus, the process projects
directly ventrad and is not divided, although there is a very slight apical
indentation. The process is shaped similarly in shelfordi but is shorter,
being subequal in length to the coxal widths. A long 4th sternal process
is thus characteristic of the genus.
Of interest is the fact that, except for Sigmoria tuberosa Shelley in
the mountains of Swain County, North Carolina, the longest 4th sternal
processes in the tribe Apheloriini are found in species in the Piedmont
Plateau, and mostly in South Carolina. Hoffman (1967, Fig. 4) showed
that the structure is longer than the adjacent coxal widths in Cleptoria
abbotti Hoffman, which occurs along the southern side of the Savannah
River in piedmont Georgia, but otherwise all species demonstrating this
condition occur north of the river. However, not all the apheloriine mil-
lipeds in South Carolina are so well endowed. The process is shorter
than the widths of the adjacent coxae in Sigmoria latior (Brolemann),
Cleptoria macra Chamberlin, and all three species of Furcillaria; and is
subequal in length in Sigmoria quadrata Shelley and S. laticurvosa
Shelley (Shelley 1981a, 1981b; Hoffman 1967). Consequently, enlarge-
ment of this process seems to have evolved independently in four dis-
tantly related genera (Cleptoria, Croatania, Sigmoria, and Brevigonus).
Or could this be an ancestral trait that has been retained by certain
species in these genera? At present we have insufficient information on
other apheloriine taxa to answer this question, since past authors have
largely ignored the configuration of the 4th sterna. Having observed
sternal variation in many species, however, I incline toward the latter
interpretation and suggest that the length, and possibly also the shape,
of the 4th sternal process might be indicators of distant phylogenetic
relationships. The 4th sternum certainly warrants more attention than it
has received, and other authors are encouraged to examine and illus-
trate it in their species so we will be better able to interpret its signifi-
cance through a more complete knowledge of variation.
60 Rowland M. Shelley
Another unusual feature of arcuatus is the presence of both simple
and reflexed tips. I know of no other apheloriine species possessing such
broad variation, and I consider the forms in Figures 3 and 5 to be con-
specific because material from intervening areas in Anderson County
displays intermediate conditions. The reflexed tips in Abbeville speci-
mens of arcuatus, coupled with the medial flanges in arcuatus and some
forms of shelf ordi, are evidence of affinity with Sigmoria. However, the
more proximal location of the flange on the basal zone in arcuatus and
shelfordi, plus the spine at the base of the acropodite in these two spe-
cies, justify generic separation from Sigmoria.
I include with arcuatus the geographically and anatomically dis-
junct males from Oconee County listed with shelfordi in my 1980 paper.
Lacking a prefemoral process, and possessing a stronger basal spine and
more broadly curved acropodite than other males of shelfordi, these
males conform more to the description of arcuatus. However, they also
differ from other specimens of arcuatus in having more massive gonop-
ods, broader medial flanges, reduced acropodal curvatures, and broader
tips. Perhaps there is a third species of Brevigonus in Oconee County,
which is unknown except for these specimens. Several trips to Clemson
and areas in Oconee County, however, have failed to produce more
individuals, and I therefore include this sample under arcuatus pending
discovery of additional material.
ACKNOWLEDGMENTS.— This study was supported in part by
NSF Grant DEB 7702596. Thanks are extended to Renaldo G. Kuhler,
NCSM scientific illustrator, for preparing Figure 2.
LITERATURE CITED
Chamberlin, Ralph V., and Richard L. Hoffman. 1958. Checklist of the
millipeds of North America. U. S. Natl. Mus. Bull. 2/2:1-236.
Hoffman, Richard L. 1967. Revision of the milliped genus Cleptoria (Polydes-
mida: Xystodesmidae). Proc. Biol. Soc. Wash. 724:1-27.
Loomis, Harold F. 1944. Millipeds principally collected by Professor V. E.
Shelford in the eastern and southeastern states. Psych 51: 166- 177.
Shelley, Rowland M. 1977. The milliped genus Croatania (Polydesmida: Xys-
todesmidae). Proc. Biol. Soc. Wash. 90:302-325.
1980. The status of Cleptoria shelfordi Loomis, with the proposal of
a new genus in the milliped family Xystodesmidae (Polydesmida).
Brimleyana 3:31-42.
1981a. Revision of the milliped genus Sigmoria (Polydesmida: Xys-
todesmidae). Mem. Am. Entomol. Soc, No. 33, 140 pp.
1981b. A new xystodesmid milliped genus and three new species from
piedmont South Carolina (Polydesmida). Proc. Biol. Soc. Wash. 94:949-961.
Accepted 17 November 1981
New Records of Marine Fishes from the Carolinas,
With Notes on Additional Species
Steve W. Ross
North Carolina Division of Marine Fisheries,
P.O. Box 769, Morehead City, North Carolina 28557
Garnett W. Link, Jr.
University of North Carolina Institute of Marine Sciences,
Morehead City, North Carolina 28557
AND
Kerry A. MacPherson
Brunswick Biological Laboratory,
Carolina Power & Light Company, Southport, North Carolina 28461
ABSTRACT. — The distributional status of fifteen marine fishes from
Carolina waters is discussed. Eleven are new records to the area, the
remainder being species previously or presently considered rare. The
second collected specimen of Daramattus americanus and the first
from the Carolinas is reported and illustrated. New maximum size
records are established for Paraconger caudilimbatus, Lepophidium
jeannae, and Hemanthias leptus. Some reproductive data are included
for Prionotus stearnsi.
Biological sampling in North and South Carolina marine waters
continues to yield fishes new to or once considered rare in the area (e.g.,
Graffe 1972; Ross and Fast 1977; Anderson et al. 1979; Burgess et al.
1979; Bbhlke and Ross 1981). Most of the species recently reported
from the Carolinas occurred in reef areas and had tropical affinities
(Anderson and Gutherz 1964; Ross and Fast 1977; Anderson et al.
1979). The fishes in this report represent a mixture of zoogeographic
affinities and habitat associations. Eleven of the fifteen species reported
are new records to this area and the remainder are species presently or
previously considered rare.
Specimens were collected by trawl, hook and line, gill net, seine,
and by sampling the traveling screens of Carolina Power & Light Com-
pany's Brunswick Steam Electric Plant (BSEP) on the lower Cape Fear
River at Southport, North Carolina. Standard lengths are given unless
otherwise noted. Nomenclature follows Robins et al. (1980). Specimens
are deposited at the Florida State Museum, University of Florida (UF)
and the University of North Carolina Institute of Marine Sciences
(UNC).
Brimleyana No. 6: 61-72. December 1981. 61
62 Steve W. Ross, Garnett W. Link, Jr., Kerry A. MacPherson
Congridae
Paraconger caudilimbatus (Poey). The margintail conger is a rarely
collected species previously reported from the Bahamas, southeastern
coast of Florida near St. Lucie Inlet, Cuba, the Gulf of Mexico, and
Guiana (Kanazawa 1961; Bohlke and Chaplin 1968; Randall et al.
1977). Manooch (1975) reported specimens tentatively identified as P.
caudilimbatus from Carolina waters. We add the following records off
of North Carolina: 33°07'N, 77°49'W, 58 m, 1 1 November 1978, night
trawl over live bottom (404 mm TL; UF 30590); 33° 57'N, 76° 28'W, 61
m, 8 December 1978, night hook and line over live bottom (542 mm TL;
UF 30591); and « 34°30'N, 75°50'W, » 61 m, February 1979, night
hook and line over live bottom (455 mm TL; UF 30592). The above
collections were during darkness and near live bottom, suggesting that
this species, like other congrids, is nocturnal and may seek shelter in
reefs during the day. All of the above specimens exceed the maximum
size (356 mm TL) recorded by Bohlke and Chaplin (1968).
Gadidae
Melanogrammus aeglefinus (Linnaeus). The haddock normally
occurs in northern waters on both sides of the Atlantic, ranging along
North America from Newfoundland to deep waters off Cape Hatteras
(Leim and Scott 1966). Bigelow and Schroeder (1953) mentioned that
haddock are seldom caught near shore and perhaps never in the littoral
zone or brackish waters. On 10 March 1979, two specimens were cap-
tured by gill net in the Neuse River, North Carolina, between Adams
Creek and South River (=* 35°00'N, 76°38'W, « 3 m). Both were
about the same size, but only one was retained (417 mm; UF 27969).
Water temperature and salinity recorded in the caputre vicinity on 9
March 1979 were 16° C and 4 0/00, respectively. This collection repre-
sents the southernmost and one of the most inshore records for haddock.
Ophididae
Brotula barbata (Schneider). One bearded brotula was collected
from the BSEP traveling screens, Southport, North Carolina (Cape
Fear River) on 18 July 1975 (192 mm; UF 30595). The previously
recorded range of B. barbata included one record from Bermuda (Beebe
and Tee-Van 1933) and other records from the Caribbean (Jamaica and
Cuba), Florida, and northern Gulf of Mexico (Hubbs 1944). Our record
represents the northernmost extent of this species, which appears to be
rare north of the Gulf of Mexico.
Lepophidium jeannae Fowler. Although L. jeannae was recorded
from Raleigh Bay, North Carolina (Silver Bay Station 1268, Bullis and
Thompson 1965), Hoese and Moore (1977) reported its northernmost
occurrence as Georgia. This species was also listed from the Cape Fear
Marine Fishes From Carolinas 63
area by Wenner et al. (1979b, 1980). Four mottled cusk eels were col-
lected off Cape Fear, North Carolina (33 °06'N, 77°5TW, 67.7 m) by
trawl in an area of sand and live bottom on 11 November 1978 (230
mm, 249 mm, 258 mm, 270 mm; UF 30599). Our specimens were much
larger than the size (200 mm) given by Hoese and Moore (1977) and
generally larger than those (238 mm max.) examined by Robins (1960).
HOLOCENTRIDAE
Myripristis jacobus Cuvier. The blackbar soldierfish was reported
from tropical Atlantic waters of Florida, the Bahamas, the northern
Gulf of Mexico and through the Caribbean to Brazil (Hoese and Moore
1977). Dahlberg (1975) mentioned that this species occurs in deeper
waters off the Georgia coast and Powles and Barans (1980) reported one
specimen off Charleston, South Carolina. Two specimens, both gravid
females (91 and 99mm; UF 30598), were trawled off Cape Fear, North
Carolina (33°03'N, 78°02'W, 42 m) during the early morning hours of
27 June 1978. Their occurrence during darkness is not surprising, con-
sidering that holocentrids are nocturnal feeders, typically hiding under
ledges or in caves during the day (Randall 1968; Greenfield 1974). This
behavior would make them practically inaccessible to trawl capture dur-
ing daylight and may result in underestimations of their occurrence.
Grammicolepidae
Daramattus americanus (Nichols and Firth). The grammicolepid
fishes are generally deep sea, widely scattered, and rarely collected.
Worldwide there are five recognized species, but there is considerable
confusion concerning validity and relationships, particularly in the
genus Daramattus. Lack of specimens for study and lack of understand-
ing of the effects of allometric growth contribute to this confusion.
Smith (1960) described Daramattus, including two species: one new, D.
armatus, and one originally described as Xenolepidichthys americanus
by Nichols and Firth (1939). Only four specimens were known to Smith,
three of which seemed to be D. armatus from Japan (1) and South
Africa (2) and the fourth (D. americanus) from Georges Bank in the
Western Atlantic. One of the two South African specimens of D. arma-
tus was later redescribed as D. barnardi (Smith 1968).
According to Hugh H. Dewitt (pers. comm.) the type of D. ameri-
canus has 13 gill rakers and 39 total dorsal elements, not 20 and 38,
respectively, as given by Nichols and Firth (1939). Considering this
change, our single specimen collected off North Carolina by trawl on 29
September 1979 at 33°32'N, 76°39'W, 232 m (56 mm; UF 30669)
seems referable to D. americanus. Meristic and morphometric data are
presented in Table 1. Fresh coloration was as follows: body generally
silvery, shading dorsally to a darker blue-gray; dark, spiny projections
64 Steve W. Ross, Garnett W. Link, Jr., Kerry A. MacPherson
Table 1. Meristic and morphometric data for Daramattus americanus (UF
30669). All measurements made to the nearest 0. 1 mm with dial calipers.
on body are included in the 14-15 dark, vertical stripes that vary in
length; dorsal and pectoral fins clear; pelvic and anal fins with some
black bands; caudal with 2-3 black bands (Fig. 1).
This is only the second specimen yet collected of this species; how-
ever, further study, especially on allometric growth effects, may reveal
that D. armatus and D. americanus are conspecific.
Gasterosteidae
Gasterosteus aculeatus Linnaeus. A threespine stickleback was col-
lected from the BSEP traveling screens on 21 February 1979 (52 mm;
UF 30594). It was previously known only as far south as Chesapeake
Bay (Burgess and Lee 1980).
Apeltes quadracus (Mitchill). The fourspine stickleback has pre-
viously been reported in North Carolina from a single specimen col-
lected in the Trent River, Craven County (Rhode et al. 1979). We report
two additional specimens from North Carolina: Stumpy Point Bay,
Pamlico Sound, 18 February 1975 (40 mm; UF 30593), and a gravid
Marine Fishes From Carolinas
65
Fig. 1. Daramattus americanus, UF 30669, 56.3 mm SL specimen collected off
North Carolina.
female from Shallowbag Bay, Roanoke Island,
1978 (40 mm; UF 30670).
Dare County, 17 March
Syngnathidae
Oostethus brachyurus (Bleeker). The opossum pipefish, O. b. line-
atus, is a relatively small tropical species commonly found in fresh and
brackish waters of Central America, but only rarely reported from
North America (see review in Gilmore 1977; Dawson 1979). Although
Dawson (1979) reported its occurrence as far north as New Jersey, this
species has not previously been recorded from North Carolina waters.
Six specimens were collected on five separate occasions from the
BSEP traveling screens: 14 September 1976 (151 mm; UF 30605); 3
October 1976 (164 mm; UF 30673); 23 October 1976 (153 mm; UF
30674); 20 November 1978 (62 mm, 85 mm; UF 30607); and 5 December
1978 (89 mm; UF 30606). Collection salinities and temperatures ranged
from 23.5 to 29.0 0/00 and 17.0 to 24.2° C, respectively. The three larg-
est specimens were adult males with developed but empty brood pouches.
Gilmore (1977) indicated that this species spawns from July through
November in fresh water of the Indian River, Florida area. Dawson
66 Steve W. Ross, Garnett W. Link, Jr., Kerry A. MacPherson
(1970) also suggested that spawning in Mississippi waters occurred dur-
ing the same period. Although adults seem to prefer fresh or estuarine
environments, juveniles and larvae apparently spend some time in
marine waters (Bbhlke and Chaplin 1968; Gilbert and Kelso 1971; Hast-
ings and Bortone 1976). Further sampling may reveal that O. b. lineatus
is more common in North Carolina and that a breeding population
exists.
Syngnathus elucens Poey. One shortfin pipefish was collected on 1
October 1968 at 34°53'N, 75°29'W (115 mm; UNC 9926). This repre-
sents a considerable range extension from the previously recorded distri-
bution of Bermuda, the Bahamas, southern Florida, the northeastern
Gulf of Mexico, and the Greater and Lesser Antilles to Surinam
(Herald 1942; Bbhlke and Chaplin 1968; Dawson 1972).
Serranidae
Hemanthias leptus (Ginsburg). One specimen of the longtail bass
was collected by hook and line in 168 m off Murrells Inlet, South Caro-
lina, on 20 August 1979 (456 mm; UF 27790). This species was pre-
viously known only from the northwestern Gulf of Mexico (Hoese and
Moore 1977), hence our record represents a significant range extension.
This specimen also greatly exceeds the previously reported maximum
size of 310 mm (Briggs et al. 1964). Although the specimen was poorly
preserved, it did retain a yellowish lateral bar from the eye to the poster-
ior margin of the opercle. The caudal fin was bright yellow, the soft anal
and dorsal pale yellow, and the other fins nearly clear. The body was
generally silver with a pale golden color dorsally. These colors, espe-
cially the yellow bar across the opercle, are similar to those of H. viva-
nus (Walls 1975; Hoese and Moore 1977). Meristics agreed well with
those of Ginsburg (1952, 1954), except that lateral line scales were 73
(left) and 59 (right), and Ginsburg gave a range of 78 to 86. Discrep-
ancies may be due to the poor condition of our specimen.
Haemulidae
Anisotremus surinamensis (Bloch). The black margate is known
from the Bahamas, southern Florida, throughout the Caribbean and
Gulf of Mexico to Brazil (Hoese and Moore 1977). Collections from the
Wrightsville Beach, North Carolina area were noted by Anderson et al.
(1979). A single specimen of this reef species was collected on the BSEP
traveling screens on 27 December 1979 (86 mm; UF 30597).
Microdesmidae
Microdesmus longipinnis (Weymouth). Pink wormfish reportedly
occur in shallow water from Charleston, South Carolina (Hammond
Marine Fishes From Carolinas 67
1973) through the northern Gulf of Mexico, Cayman Islands, and Ber-
muda (Dawson 1969). Two specimens were collected in the lower Cape
Fear River, North Carolina: one from the BSEP traveling screens (237
mm; UF 30596) on 23 March 1977, and the other from an unknown
locality in the lower Cape Fear River (67 mm; UF 30671). A third spec-
imen from the lower Cape Fear, unavailable for study, is deposited in
the UNC collections. Our collections represent the first records of
microdesmids in North Carolina.
Triglidae
Prionotus ophryas Jordan and Swain. The bandtail searobin was
first reported from North Carolina waters by Bullis and Thompson
(1965): 34°00.5'N, 7602TW, 54-60 m, 5 September 1979, and subse-
quently by Wilk and Silverman (1976): 34°20'N, 76°52'W, 25 m, 17
November 1971 (200 mm TL); 34° 11.5*N, 76°47'W, 31 m, 16 November
1971 (140 mm TL); 33° 27.5'N, 77° 24'W, 32 m, 24 May 1972 (190 mm
TL). We have compiled a number of additional records from R/V Dan
Moore surveys and other collections (Table 2), which include the north-
ernmost record (DM 3754) of this species. Most P. ophryas were col-
lected over sand bottoms; however, they are known to occur near live
bottoms (S. W. Ross, pers. obs.) and calico scallop beds (Schwartz and
Porter 1977; pers. obs.). Although not one of the most common of the
offshore searobins, this species is regularly encountered in a depth range
of 24 to 60 m (Table 2).
Prionotus stearnsi Jordan and Swain. Fourteen shortwing searob-
ins were collected by trawl on three occasions: 33041'N, 76°42'W, 152
m, 19 May 1978 (63 mm; UF 30601); 33°02'N, 77°53'W, 113 m, 27
June 1978 (76-105 mm; UF 30602); and 34°29'N, 75°58'W, 57 m, 13
December 1978 (92 mm; UF 30600). The June collection contained six
females with ripe gonads, data for which are included in Table 3. Very
ripe females (125 and 130 mm) were reported in August near the Tortu-
gas by Longley and Hildebrand (1941). Lewis and Yerger (1976) sug-
gested that both sexes reach maturity by 60 mm and found well devel-
oped ova (0.6 mm diameter) in October and December. The large
gonads and eggs from the June specimens (Table 3) indicated that these
fish were probably near spawning. Shortwing searobins had previously
been reported from off Cape Fear, North Carolina (Bullis and Thomp-
son 1965; Wenner et al. 1979a, 1979b), although Roberts-Goodwin
(1981) had listed them as far north as South Carolina and Hoese and
Moore (1977) only north to Georgia.
68
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Marine Fishes From Carolinas 69
Table 3. Length, weight, and sex data of Prionotus stearnsi collected on 27
June 1978 (UF 30602).
SL(mm) Wt.(g) Sex °™Z Gonad index* Egg diameter
wt. (g)+ size range (mm)
+ Both gonads combined.
* Gonad index = gonad weight + (body weight - gonad weight) X 100.
ACKNOWLEDGMENTS.— Thanks are extended to W. D. Ander-
son for verification of Hemanthias leptus, to G. H. Burgess for use of
Florida State Museum collections, to H. H. DeWitt for information on
and review of the Daramattus section, and to C. Benedict, G. F. Booth,
R. Cahoon, D. S. Cooke, L. L. Davidson, S. Edwards, C. Harvell, G.
R. Huntsman, B. Mahaffee, E. G. McGowan, D. Mumford, H. J. Port-
er, D. R. Sager, J. M. Swing, and M. T. Tyndall, for providing some of
the specimens. We are especially grateful to B. F. Holland and J. W.
Gillikin, Jr. for allowing us use of data collected by the R/V Dan
Moore. We also thank Ms. M. Fotch for typing the manuscript.
LITERATURE CITED
Anderson, H. M., T. H. Handsel and D. G. Lindquist. 1979. Additional notes
on tropical reef fishes in the Carolinas with zoogeographic considerations.
ASB Bull. 26(2):31.
Anderson, William D., Jr., and E. J. Gutherz. 1964. New Atlantic coast ranges
for fishes. Q. J. Fla. Acad. Sci. 27(4):299-306.
Beebe, William, and J. Tee- Van. 1933. Field Book of the Shore Fishes of Ber-
muda and the West Indies. G. P. Putnam's Sons, New York, 337 pp.
Bigelow, Henry B., and W. C. Schroeder. 1953. Fishes of the Gulf of Maine.
U.S. Fish Wildl. Serv. Fish. Bull. 74,55:1-577.
Bohlke, Eugenia B., and S. W. Ross. 1981. The occurrence of Muraena robusta
Osorio (Anguilliformes, Muraenidae) in the west Atlantic. Northeast Gulf
Sci. 4(2): 123-125.
70 Steve W. Ross, Garnett W. Link, Jr., Kerry A. MacPherson
Bohlke, James E., and C. C. G. Chaplin. 1968. Fishes of the Bahamas and
Adjacent Tropical Waters. Livingston Publishing Co., Wynnewood, Pa.
771 pp.
Briggs, John C, H. D. Hoese, W. F. Hadley and R. S. Jones. 1964. Twenty-two
new marine fish records for the northwestern Gulf of Mexico. Tex. J. Sci.
7tf(l):113-116.
Bullis, Harvey R., Jr., and J. R. Thompson. 1965. Collections by the explora-
tory fishing vessels Oregon, Silver Bay, Combat, and Pelican made during
1956-1960 in the southwestern north Atlantic. U.S. Fish Wildl. Serv. Spec.
Sci. Rep. Fish. 510. 130 pp.
Burgess, George H., and D. S. Lee. 1980. Gasterosteus aculeatus Linneaus,
threespine stickleback, p. 563 in D. S. Lee et al. Atlas of North American
Freshwater Fishes. N.C. State Mus. Nat. Hist., Raleigh, x + 867 pp.
, G. W. Link, Jr. and S. W. Ross. 1979. Additional marine fishes new
or rare to Carolina waters. Northeast Gulf Sci. 5(2):74-87.
Dahlerg, Michael D. 1975. Guide to the Coastal Fishes of Georgia and Nearby
States. Univ. Georgia Press, Athens. 186 pp.
Dawson, Charles E. 1969. Studies on the gobies of Mississippi Sound and
adjacent waters II. An illustrated key to the Goboid fishes. Publ. Gulf
Coast Res. Lab Mus. I. 59 pp.
1970. A Mississippi population of the opossum pipefish, Oostethus
lineatus (Syngnathidae). Copeia 1970(4):772-773.
1972. Nektonic pipefishes (Syngnathidae) from the Gulf of Mex-
ico off Mississippi. Copeia 1972(4):844-848.
1979. Review of the polytypic doryrhamphine pipefish Oostethus
brachyurus (Bleeker). Bull. Mar. Sci. 29(4):465-480.
Gilbert, Carter R., and D. P. Kelso. 1971. Fishes of the Tortuguero area,
Caribbean Costa Rica. Bull. Fla. State Mus. Biol. Sci. 76(1): 1-54.
Gilmore, R. Grant, Jr. 1977. Notes on the Opossum Pipefish, Oostethus line-
atus, from the Indian River Lagoon and vicinity, Florida. Copeia 1977(4):
781-783.
Ginsburg, Isaac. 1952. Eight new fishes from the gulf coast of the United States,
with two new genera and notes on geographic distribution. J. Wash. Acad.
Sci.42(3):84-101.
1954. Four new fishes and one little known species from the east
coast of the United States including the Gulf of Mexico. J. Wash. Acad.
Sci. **(8):256-264.
Graffe, Arthur J. 1972. A range extension of the callionymid fish Callionymus
pauciradiatus (Callionymidae). Chesapeake Sci. 13(2): 153.
Greenfield, David W. 1974. A revision of the squirrelfish genus Myripristis
Cuvier (Pisces: Holocentridae). Nat. Hist. Mus. Los Ang. Cty. Sci. Bull.
79:1-54.
Hammond, Donald L. 1973. A record of Micro desmus longipinnis (Weymouth)
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Hastings, Philip A., and S. A. Bortone. 1976. Additional notes on tropical
marine fishes in the Northern Gulf of Mexico. Fla. Sci. 39(2): 123-125.
Herald, Earl S. 1942. Three new pipefishes from the Atlantic coast of North and
South America, with a key to the Atlantic American species. Stanford
Ichthyol. Bull. 2(4):125-134.
Marine Fishes From Carolinas 71
Hoese, H. Dickson, and R. H. Moore. 1977. Fishes of the Gulf of Mexico,
Texas, Louisiana, and Adjacent Waters. Texas A & M Univ. Press, College
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Hubbs, Carl L. 1944. Species of the circumtropical fish genus Brotula. Copeia
1944(3): 162-178.
Kanazawa, Robert H. 1961. Paraconger, a new genus with three new species of
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Leim, A. H., and W. B. Scott. 1966. Fishes of the Atlantic coast of Canada.
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fishes of Tortugas, Florida. Carnegie Inst. Wash. Paps. Tortugas Lab. 34.
331 pp.
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History, Ecology, and Tagging of the Red Porgy, Pagrus pagrus Linnaeus,
off North Carolina and South Carolina. Unpub. Ph.D. dissert., N. C. State
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72 Steve W. Ross, Garnett W. Link, Jr., Kerry A. MacPherson
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159 pp.
Accepted 31 October 1981
Nesting and Management of the Atlantic Loggerhead,
Caretta caretta caretta (Linnaeus) (Testudines:
Cheloniidae) on Cape Island, South Carolina,
in 1979.
John B. Andre1 and Larry West
Cape Romain National Wildlife Refuge,
Route 1, Box 191, Awendaw, South Carolina 29429
ABSTRACT. — Nesting activity of the Atlantic loggerhead turtle, Caretta
caretta caretta, was monitored on Cape Island, Cape Romain National
Wildlife Refuge, South Carolina, in 1979. The nesting season encom-
passed 106 days, beginning in mid-May and continuing through August.
An estimated total of 1,093 clutches was laid on the island with an
average of 136.6 nests/ km. Seventy-one percent of all turtle emergences
on the beach were false crawls (non-nesting emergences). Three-hundred
seventy-nine nests were removed to an on-site, predator-proof hatchery,
which produced 10,185 hatchlings from 1 17 nests. Mean clutch size was
117.0 (SD = ± 4.31)and hatching success was 74.4%. Raccoons and
erosion destroyed most of the natural nests, but 714 of them produced
3.605 hatchlings. On 4 September, 17.8 cm of rainfall associated with
Hurricane David drowned or washed away all unhatched eggs in the
nesting areas and hatchery. To reduce predator-induced mortality, 82
raccoons were removed from Cape Island prior to and during the turtle
nesting season. This resulted in a 25% reduction in first night predation
from 1978 levels. An average of 2.2 (SD = ± 1.84) nests were destroyed
per night in 1979 compared to 7.5 nests per night in 1978.
INTRODUCTION
The major worldwide nesting areas of loggerhead turtles, Caretta
caretta (Linnaeus), are in the southeastern United States, southeastern
Africa, and eastern Australia. In the United States, C c. caretta nests
primarily on the beaches of North and South Carolina, Georgia, and
the Atlantic and Gulf coasts of Florida (U.S. Fish and Wildlife Service
1979). Cape Island, in Cape Romain National Wildlife Refuge (CRNWR),
probably has more nesting activity than any other South Carolina log-
gerhead rookery.
In light of the classification of loggerheads as Threatened (Federal
Register, 28 July 1978), it is imperative that rookeries be protected and
managed to maximize loggerhead productivity. Little is known of the
fates of hatchling loggerheads in the ocean, but probably very few reach
sexual maturity. This unknown also necessitates that management activ-
ities be directed at increasing the number of loggerhead hatchlings
reaching the ocean.
Factors adversely affecting loggerhead productivity on or near
Cape Island are predators, poachers, erosion and inundation, and
•Present address: U.S. Fish and Wildlife Service, P. O. Box 87, Kilauea, Kauai,
Hawaii 96754
Brimleyana No. 6: 73-82. December 1981. 73
74 John B. Andre and Larry West
commercial fishing. Raccoons, Procyon lotor, are major egg predators
(Stancyk et al. 1980; Hopkins et al. 1979; Davis and Whiting 1977; and
Gallagher et al. 1972) and, if not controlled, can reduce loggerhead pro-
ductivity to zero. Erosion and inundation are responsible for the loss of
many egg clutches on Cape Island. Shrimp trawling also reduces log-
gerhead productivity by drowning turtles caught in trawls (Ulrich 1978).
This paper documents and describes loggerhead nesting activity at
Cape Island in 1979 and management practices initiated at CRNWR to
increase loggerhead productivity, i.e. the number of hatchlings entering
the ocean. Management practices include the operation of a hatchery
and a predator control program.
The Study Area
Cape Island is approximately 8 km long and 2.4 to 0.1 km wide.
The barrier island is a part of the Santee River Delta complex (Price
1955). Cape Island owes some of its origin to sediments supplied by the
Santee River (Brown 1977), and diversion of a large part of the Santee's
flow into the Cooper River in 1941 decreased its sediment load. This
factor was responsible for the shift from a stable or depositional phase
to a destructive, erosional phase at Cape Island (Aburawi 1972). Since
1941 the island has eroded over 215 m at several points (Stephen et al.
1975), and erosion continues to move the steep, narrow front" beach
landward at a rate of about 10 m/yr (Christopher H. Ruby, pers.
comm.). This erosion, combined with the wind and high tides that
accompanied Hurricane David on 4 September 1979, breached the
island at two locations. The hurricane also removed some of the remain-
ing dune system, thus reducing loggerhead nesting habitat.
The front beach and primary dune system are used by nesting log-
gerheads. Landward of the steep, narrow beach are dunes and tidal
wash-over areas (see Caldwell 1959 for a complete description of Cape
Island beach types). Large elongate dunes lie parallel to the beach and
grade (abruptly to gradually) into small, isolated clumps of dunes.
MATERIALS AND METHODS
Surveys of the Cape Island beach were conducted from 18 May to
28 August to document loggerhead nesting activity. Each survey began
about one hour after dark and ended at dawn. Data recorded during
each survey included the numbers of freshly laid nests, false crawls, and
nests destroyed by predators. An average of 4.2 (range 1-7) surveys was
conducted each week during the 15 week nesting season. Estimates of
total weekly nesting activity (Table 1) were obtained by summing the
numbers of nests and false crawls observed on surveys during that week,
dividing the sums by the number of surveys conducted, and multiplying
these means by seven. Data reported from 1978 were extracted from
Atlantic Loggerhead Nesting
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76 John B. Andre and Larry West
CRNWR report files and measurements of variation were not available.
To increase loggerhead productivity, nests were moved to a predator-proof
exclosure (hatchery) located behind a 2 m high primary dune that pro-
tected the eggs from erosion and saltwater inundation. Nests were found
by following crawl tracks. The site was then carefully probed with a
metal pole to locate the eggs, which were excavated by hand, removed
from the nest, and placed in a plastic bucket containing about 5 cm of
moist sand. The eggs were then covered with moist sand from the nest
cavity to reduce evaporation and temperature fluctuation. In probing
for nests, 1-3 eggs at the top of the egg mass were broken in approxi-
mately 10 clutches during the nesting season. The in-nest orientation of
eggs was not maintained when transferred to the hatchery. Clutches that
were partly destroyed by raccoons received the same treatment, except
that eggs from different clutches were combined so that each trans-
planted group contained at least 60 eggs. Exhumed eggs were usually
less than four, never more than twelve, hours old, and were placed in
the hatchery within two hours of collection.
Hatchery clutches, spaced 0.6 m apart, were reburied in holes
approximating the dimensions of natural nest cavities. Eggs were placed
in the holes until they held a complete clutch or at least 60 eggs. A
shallow layer of moist sand was placed over the egg mass and lightly
packed by hand, then additional sand layers were placed and packed
until the cavity was filled. Each nest in the hatchery was marked with a
flag indicating the nest number, number of eggs and date laid. Hatch-
ling turtles were removed from the hatchery before daylight, released on
the beach berm, and allowed to crawl down the beach and enter the
ocean unaided.
The hatchery consisted of two pens (6.1 X 12.2 m and 6.1 X 18.3
m) placed together. Both enclosures were 1.8 m tall and the sides and
tops were enclosed with 5.1 X 10.2 cm welded wire. A 91 cm width of
chicken wire was buried around the perimeter of the pens to prevent
predators from digging under the sides. Fiberglass panels, 76 cm tall,
were placed around the pens to prevent wind-blown sand from covering
nests and to keep ghost crabs, Ocypode quadrata, and rats, Rattus rat-
tus and R. norvegicus, from entering the hatchery. The sparse vegeta-
tion within the hatchery was removed by hand to protect the eggs from
penetration and entanglement by plant roots.
An egg and hatchling predator control program was conducted
from April 1979 through the loggerhead nesting season. Raccoons were
captured with live and leg-hold traps. Methods of raccoon population
management were outlined by Ehrhart (1979).
Atlantic Loggerhead Nesting 77
RESULTS AND DISCUSSION
Nesting Activity
The 1979 loggerhead nesting season at Cape Island encompassed 10
days, from the first clutch on 15 May to the last clutch on 28 August.
Results of 63 surveys of the Cape Island beach are depicted in Figures 1
and 2. Table 1 shows the estimated total numbers and the mean (±
standard deviation) numbers of nests and false crawls per week. Since
the surveys included almost 60% of the nesting season, we believe our
estimates reflect actual total nesting activity.
Loggerheads on Cape Island laid 1,093 clutches in 1979 compared
to 1,451 clutches in 1978. This apparent 25% decline may have been
caused by rapid erosion of the beach and dune system, but past records
at CRN WR indicate considerable fluctuation in annual nesting activity
at Cape Island. Another possible explanation for variation in number of
nests per season is the two to three year breeding cycle of female logger-
heads. Kaufman (1975) and Davis and Whiting (1977) reported that
female loggerheads demonstrated a two year breeding cycle in Colum-
bia, South America, and Everglades National Park, Florida, respec-
tively. Davis and Whiting (1977) also found that the even-year breeding
population was about twice the size of the odd-year breeding popula-
tion, resulting in different levels of nesting activity each year.
The frequency of false crawls or non-nesting emergences may be
related to quality of nesting habitat, since Davis and Whiting (1977)
observed more false crawls on poor quality beaches. Comparison of our
data with other studies supports this suggestion. In 1979, 71% of all
loggerhead emergences on the Cape Island beach were false crawls. The
mean number per night was 25.8 (SD = ± 22.21), compared to 5.3 per
night at Cape Island in 1939 (Caldwell 1959), a five-fold increase. Dif-
ferent data collecting methods between our study and Caldwell's proba-
bly account for part of the increase. Also, Talbert et al. (1980) found
that the frequency of false crawls at Kiawah Island, South Carolina,
ranged from 28.7% to 47.7%, with a mean of 40.5%, from 1972 to 1976.
Kiawah Island, approximately 83 km southwest of Cape Island, has
experienced much less erosion (Hayes et al. 1977).
Even though the frequency of false crawls is high, Cape Island is
heavily used as a loggerhead nesting area. We found a mean of 136.6
nests/ km of beach in 1979 based on the total estimated number of nests.
Ehrhart (1979) reported 44.2 to 79.2 nests/ km of beach at the Kennedy
Space Center on the east coast of Florida, while Talbert et al. (1980)
found an average of 9.5 nests/ km for five nesting seasons at Kiawah
Island. The nesting concentration at Cape Island indicates the impor-
tance of this island as a loggerhead rookery.
Figure 1 depicts the seasonal nesting activity of female loggerheads
at Cape Island in 1979. Numbers of nests per night increased rapidly
78
John B. Andre and Larry West
30
G> 20
u 10
0)
E
3
I
ill
20 30 10 20 30 10 20 30 10 20 30
May
June
July
August
Fig. I . Actual number of nests laid on each survey night on Cape Island in 1979.
Asterisks indicate completed surveys with no nests laid; surveys were not made
on other nights represented by zero nests.
20 30 10 20 30 10 20 30 10 20 30
May June July August
Fig. 2. Actual number of false crawls observed on each survey night on Cape
Island in I979. Asterisks indicate completed surveys with no false crawls
observed; surveys were not made on other nights represented by zero false crawls.
Atlantic Loggerhead Nesting 79
from 18 May to the first week of June. From June through most of July
nesting activity was roughly at the same level, but the numbers of nests
laid per night varied considerably. In the last week of July nesting activity
started to decrease until the end of the season in late August.
S. R. Hopkins of the South Carolina Wildlife and Marine Resour-
ces Department, in cooperation with the U.S. Fish and Wildlife Service,
conducted a loggerhead study from 1977 (Hopkins et al. 1979) to 1979
on a 3 km section of the Cape Island beach. She found that 5.8% of 209
natural nests (nests not placed in the hatchery) hatched successfully in
1979 (Hopkins, pers.comm.). With this percentage of natural nests
hatching, we can calculate natural loggerhead productivity for 1979 at
Cape Island. We determined that the mean clutch size was 117,
(SD = ± 4.31, N = 393), and that an average of 74.4% (N=10) of the
eggs in each clutch hatched. With 5.8% of 714 nests (379 of the total
1,093 nests were transplanted) hatching successfully, the number of
hatchlings produced from natural nests was estimated to be 3,605.
Hatchery Program
Loggerhead turtle hatcheries have been used since at least 1965.
The early hatcheries were designed to protect eggs and hatchlings from
raccoons and crabs (Richardson 1978). In addition to this function,
though, the Cape Island hatchery included eggs from nests that would
have been destroyed by beach erosion or saltwater inundation produced
by winds and tides.
At the end of the 1979 nesting season the hatchery contained 379
nests, the first six placed there on 29 May and the last three on 16
August. The first hatchlings were released from the enclosure on 28 July
and the last of the season on 3 September. The incubation period in the
hatchery was about 65 days, but could not be precisely determined since
tracks of the hatchlings made it difficult to determine from which nest
they emerged. Hopkins (unpublished data) observed that the incubation
period for natural nests on Cape Island averaged 65.3 days in 1979.
The hatchery and dune system of Cape Island were severely dam-
aged on 4 September by 50 to 60 mph winds produced by Hurricane
David and the occurrence of a 2.1 m spring tide. Prior to the hurricane,
1 17 nests hatched successfully in the enclosure. An average of 87.0 hatch-
lings emerged from each nest, resulting in the production and release of
10,185 hatchlings. From 1965 to 1976, the hatchery at Little Cumber-
land Island, Georgia, produced an average of 65 hatchlings /nest (Richardson
1978). We hand-released about 8,000 turtles from the hatchery, and
about 2,200 hatchlings were self-released at night through a tunnel con-
structed from hatchery to beach.
Morning releases of hatchlings near the hatchery site were quickly
discovered by Laughing Gulls, Larus atricilla, and Herring Gulls, L.
80 John B. Andre and Larry West
argentatus, and approximately 200 turtles were taken by these birds dur-
ing the season. Losses were minimized by releasing the turtles each day
at a point on the beach where gulls were not present.
We checked the condition of the hatchery on 6 September, after the
hurricane had passed, and found that the unhatched eggs had drowned,
regardless of the degree of embryo development. Loggerhead egg mor-
tality from excessive rainfall and the condition of drowned eggs has
been reported by Ragotzkie (1959), and our observations concur. A ser-
ies of exhumed nests showed that the last that hatched had been placed
in the hatchery on 28 June. Groundwater level was at or near the tops
of all exhumed nests.
Most of the natural nests on Cape Island were also destroyed by
the hurricane (Hopkins, pers. comm.). Of 51 unhatched nests remaining
in her 3 km study area, all were destroyed by erosion or inundation
produced by the hurricane. Our inspection of nesting habitat on the
island after the hurricane revealed that no natural nesting sites had been
effectively protected.
Predation
The major problem for management of the Atlantic population log-
gerhead rookeries is the raccoon (Ehrhart 1979). Destruction of nests by
raccoons is the most important factor limiting turtle productivity on
Cape Island. First night predation (nests destroyed on the same night
laid) is almost 100% in some rookeries (Ehrhart 1979). In 1972 and
1973, raccoons took 85% and 75%, respectively, of the loggerhead nests
at Cape Sable, Florida, and first night predation accounted for 87% of
the destroyed nests (Davis and Whiting 1977). To reduce egg and hatch-
ling losses, 82 raccoons were removed from the island during April
through August 1979. The effect of the trapping effort is clear. In 1978,
first night predation by raccoons destroyed 47% of the nests compared
to 22% in 1979. The greatest number of nests destroyed on the night laid
was 18 in 1978 and 7 in 1979, and the mean number of nests destroyed
per night was 7.5 in 1978 and 2.2 (SD = ± 1.84) in 1979. On a 3 km
section of Cape Island beach, Hopkins (unpublished data and pers.
comm.) found that raccoons destroyed 95.8% of the marked nests in
1978 and 59.3% in 1979. The 1979 predation percentage would have
been somewhat higher, but nests were not available to predators after
Hurricane David (Hopkins, pers. comm.). Klukas (1967) and Davis and
Whiting (1977) reported similar findings after initiating a raccoon con-
trol program at Cape Sable, Florida. However, trapping programs are
labor intensive and must be conducted yearly to maintain effectiveness.
Reproduction by raccoons, and immigration from nearby areas, quickly
replace those removed by trapping.
Atlantic Loggerhead Nesting 81
ACKNOWLEDGMENTS.— The U.S. Fish and Wildlife Service, Cape
Romain National Wildlife Refuge, provided funds, materials, and per-
sonnel for this refuge project. We thank the refuge manager, George
Garris, and the staff for their many nights of work, and George Balazs
for commenting on an earlier draft of the manuscript. We also thank
two anonymous referees for providing excellent reviews and John E.
Cooper for editorial assistance, all of which greatly improved the
manuscript.
LITERATURE CITED
Aburawi, R. 1972. Sediment facies of the Holocene Santee delta. Unpubl. M. S.
thesis, Univ. South Carolina, Columbia. 96 pp.
Brown, P. J. 1977. Variations in South Carolina coastal morphology. South-
east. Geol. 18:249-264.
Caldwell, D. K. 1959. The loggerhead turtles of Cape Romain, South Carolina.
Bull. Fla. State Mus. Biol. Sci. 4:3 19-348.
Davis, G. E., and M. C. Whiting. 1977. Loggerhead sea turtles nesting in Ever-
glades National Park, Florida, USA. Herpetologica 55.18-28.
Ehrhart, L. M. 1979. Reproductive characteristics and management potential of
the sea turtle rookery at Canaveral National Seashore, Florida, pp. 397-399
in R. M. Linn (ed.). Proceedings of the first conference on scientific
research in the national parks, 9-17 November 1976, New Orleans, La. NPS
Trans. Proc. Ser. No. 5.
Gallagher, Robert M., M. L. Hollinger, R. M. Ingle and C. R. Futch. 1972.
Marine turtle nesting on Hutchinson Island, Florida, in 1971. Fla. Dep.
Nat. Resour. Mar. Res. Lab. Spec. Sci. Rep. No. 37. 1 1 pp.
Hayes, M. O., T. F. Moslow and D. K. Hubbard. 1977. Beach erosion in South
Carolina. Coastal Res. Div., Dep. Geol., Univ. South Carolina, Columbia.
Hopkins, S. R., T. M. Murphy, Jr., K. B. Stansell and P. M. Wilkinson.
1979. Biotic and abiotic factors affecting nest mortality in the Atlantic
loggerhead turtle, Proc. 32nd Annu. Cong. Southeast. Assoc. Fish Wildl.
Agencies, pp. 213-223.
Kaufman, Reinhard. 1975. Studies of the loggerhead sea turtle, Caretta caretta
caretta (Linne') in Colombia, South America. Herpetologica 57.323-326.
Klukas, R. W. 1967. Factors affecting nesting success of Loggerhead turtles at
Cape Sable, Everglades National Park. File No. N1415, Natl. Park Serv.,
P. O. Box 279, Homestead, Fla. 33030. 58 pp., mimeo.
Price, W. A. 1955. Correlations of shoreline type with offshore bottom condi-
tions. Project 53, Dep. Oceanography, Texas A&M Univ., Bryan.
Ragotzkie, Robert A. 1959. Mortality of loggerhead turtle eggs from excessive
rainfall. Ecology 40:303-305.
Richardson, James I. 1978. Results of a hatchery for incubating loggerhead sea
turtles (Caretta caretta) (Linne') eggs on Little Cumberland Island, Geor-
gia. Fla. Dep. Nat. Resour. Mar. Res. Publ. No. 33. Abstract.
Stancyk, Stephen E., O. R. Talbert, Jr. and J. M. Dean. 1980. Nesting activity
of the loggerhead turtle (Caretta caretta) in South Carolina. II. Protection
of nests from raccoon predation by transplantation. Biol. Conserv. 75:289-298.
82 John B. Andre and Larry West
Stephen, M. F., P. J. Brown, C. M. Fitzgerald, D. K. Hubbard and M. O.
Hayes. 1975. Beach erosion inventory of Charleston County, South Caro-
lina: a preliminary report. S. C. Sea Grant Tech. Rep. No. 4. 79 pp.
Talbert, O. Rhett, Jr., S. E. Stancyk, J. M. Dean and J. M. Will. 1980. Nesting
activity of the loggerhead turtle (Caretta caretta) in South Carolina I. A
rookery in transition. Copeia 1980 (4):709-718.
Ulrich, G. F. 1978. Incidental catch of loggerhead turtles by South Carolina
commercial fisheries. Report to NMFS, Contract No. 01-7-042-35151 and
03-7-042-35121.
U. S. Fish and Wildlife Service. 1979. Endangered and threatened species of the
southeast United States. U. S. Dep. Inter., Fish Wild. Service, Atlanta.
Accepted 10 August 1981
Distribution, Morphology and Life History
of the Least Brook Lamprey,
Lampetra aepyptera (Pisces: Petromyzontidae), in Kentucky
Stephen J. Walsh and Brooks M. Burr
Department of Zoology,
Southern Illinois University at Carbondale,
Carbondale, Illinois 62901
ABSTRACT.— Kentucky distribution records for the least brook
lamprey, Lampetra aepyptera (Abbott), show that it is the most
abundant and widespread lamprey in the state, inhabiting most major
drainages where suitable habitat is available. Morphological and denti-
tion data from over 220 specimens reveal that the recently described
Lethenteron meridionale, from Tennessee, Alabama and Georgia, is a
synonym of L. aepyptera. Interdrainage variation in meristic and mor-
phometric characters of L. aepyptera falls within the normal range of
variability for the species in the Ohio Valley.
INTRODUCTION
Although a considerable distributional and ecological literature
exists for the least brook lamprey, Lampetra aepyptera (Abbott), little
information has been published on its occurrence and life history in
Kentucky. A few distributional records for the species in Kentucky were
reported by Branson (1970), Burr and Mayden (1979), Burr (1980), and
Clay (1975). The only information on its natural history in the state was
summarized by Clay (1975), who included original observations on the
demise of a population in Knob Creek, southwest of Louisville. The
most complete studies on the life history of L. aepyptera were done in
Maryland (Seversmith 1953) and Delaware (Rohde et al. 1976). Branson
(1970), Clay (1975), and Clay and Carter (1957) briefly described its
morphological characteristics in Kentucky. In their description of
Lethenteron meridionale, Vladykov et al. (1975) gave the most complete
morphological description of L. aepyptera to date. Rohde et al. (1976)
summarized meristic and morphometric characteristics of the species in
Delaware and the literature prior to their study.
The purposes of this paper are to (1) map and describe the Ken-
tucky distribution of L. aepyptera, (2) analyze its morphological charac-
teristics in the state, and (3) supplement published ecological informa-
tion with our observations of what appears to be a neotenic population
in western Kentucky, a heretofore unreported phenomenon in this spe-
cies. The discovery of posterior circumoral teeth in a number of Ken-
tucky L. aepyptera, and the variability of their presence, leaves little
doubt that Lethenteron meridionale is a synonym of Lampetra
aepyptera.
Brimleyana No. 6: 83-100. December 1981. 83
84 Stephen J. Walsh and Brooks M. Burr
METHODS AND MATERIALS
Over 220 adults, from the institutions listed in Acknowledgments,
were used for morphological comparisons between major drainages or
ichthyofaunal blocks as designated by Burr (1980). All measurements
were made to the nearest 0.01 mm with dial calipers. Terminology of
dentition follows Hubbs and Potter (1971) and Potter (1980). The oral
discs of some specimens were lightly stained with alizarin red S to facili-
tate tooth counts. Methods of counting and measuring were those of
Vladykov and Follett (1958, 1965).
The life history of L. aepyptera was studied at Terrapin Creek,
Graves County, Kentucky, near the Tennessee border. Collections were
made from the state line north to 0.8 km south of Bell City, Kentucky.
Intermittent sampling from 23 March 1978 to 4 June 1980 resulted in
138 specimens, which were fixed in 10% formalin and later transferred
to 70% ethanol. Frequencyof occurrence versus lengths of ammocoetes
taken at Terrapin Creek on 4 June 1980 were plotted and the curve
smoothed using 7 mm sliding averages (Hardisty 1961; Hardisty and
Potter 1971; Rohde et al. 1976; Taylor 1965). Lampreys were blotted
dry and weighed to the nearest 0.1 mg on a Sartorius analytical bal-
ance. Egg diameters were measured with an ocular micrometer at a
magnification of 25-50X; no correction was made for shrinkage.
Collections of L. aepyptera examined, all from Kentucky, are listed
below by major drainage, county, catalog number, locality, and date.
The number of specimens is shown in parentheses.
Big Sandy River Drainage.— Martin Co.: UL 7945 (1) Rockhouse Fork of
Rockcastle Cr. at Stidham, 27 March 1956. Floyd Co.: UL 5946 (3) John's
Cr. below Dewey Dam, 27 March 1956. Morgan Co.: SIUC uncat. (4)
Open Fork of Paint Cr. at ford, 24 March 1973.
Licking River Drainage.— Montgomery Co.: INHS 78829 (2) creek, 1.6 km W
Means, 12 April 1968. Rowan Co.: UL 5228 (16) Licking R. at Cranston,
12 April 1941. Menifee Co.: UL 4983 (1) Beaver Cr., 8 April 1954.
Kentucky River Drainage. — Wolfe Co.: KNPC uncat. (2) Swift Camp Cr. just
upstream from confluence Red R., 3 May 1980; UL 1014 (1) Red R. near
Hazel Green, 24 November 1960.
Upper Cumberland River Drainage.— McCreary Co.: WCS 720-01 (9) Taylor
Branch, 8.8 km E Whitley City, 16 April 1977. Whitley Co.: INHS 79259
(1) Youngs Cr., 1.6 km W Clio, 19 March 1978. Rockcastle Co.: UL 2015
(1) Clear Cr., 3.2 km S Disputanta, 3 April 1980; EKU 168 (1) Clear Cr.,
3.2 km SE Disputanta, 20 March 1966.
Salt River Drainage.— Hardin Co.: KNPC uncat. (3) Clear Cr. downstream
from overpass E Colesburg, 24 March 1978. Bullitt Co.: USNM 164102
(15) Knob Cr., 28 March 1950; UL 4791 (20) Knob Cr. near Mitchell Hill
Road, 25 March 1950; UL 4794 (73) Knob Cr. 9.6 km NW Shepherdsville,
25 March 1950; INHS 78228 (1) creek, 9.6 km S Fairdale, 7 April 1968.
Least Brook Lamprey in Kentucky 85
Ohio River Drainage.— Hardin Co.: UL 5949 (1) trib. Otter Cr. near Vine
Grove, 11 March 1956; UL 4810 (6) Otter Cr., 3.2 km W Vine Grove, 13
March 1954; UL 1 1886 (3) Otter Cr., S Vine Grove, 6 May 1959.
Green-Barren River Drainage.— Allen Co.: SIUC uncat. (4) Long Hungry Cr.,
1.6 km SE Mt. Zion, 18 March 1980. Barren Co.: SIUC uncat. (4) Peter
Cr., 1.6 km SE Dry Fork, 18 March 1980. Edmonson Co.: UL 5229 (5)
trib. Beaverdam Cr. near Brownsville, 31 March 1955. Green Co.: UL
11001 (1) Caney Fork at Hwy. 61, 8 April 1978. Larue Co.: INHS 78477 (1)
Walters Cr., 6.4 km N Magnolia, 28 March 1964. Russell Co.: INHS 79126
(1) trib. Goose Cr., 6.4 km NE Russell Spring, 18 March 1978. Simpson
Co.: UL 12835 (1) West Fork Drakes Cr., 1.6 km above confluence Lick
Fork, 8 November 1964. Taylor Co.: KU 11612(6) Big Pitman Cr., 11.8 km
NW Campbellsville, 2 April 1966.
Rough River Drainage.— Hardin Co.: UL 11893 (19) Rough R., 25 April 1959;
UL 12905 (1) trib. Rough Cr., 0.8 km S Four Corners, 22 March 1959.
Grayson Co.: SIUC uncat. (1) Spring Fork, 1.6 km NW Tousey, 12 March
1979. Ohio Co.: SIUC uncat. (2) West Fork, 0.8 km N Fordsville, 11
March 1979; SIUC uncat. (2) Rocky Fork, 2.4 km SW Shreve, 12 March
1979; SIUC uncat. (1) Halls Cr., 13.3 km NE Hartford, 11 March 1979;
SIUC uncat. (1) Sixes Cr., 3.2 km S Renfrow, 12 March 1979.
Lower Cumberland River Drainage. — Trigg Co.: SIUC uncat. (1) Donaldson
Cr., 9.6 km SE Canton, 10 March 1979.
Tennessee River Drainage. — Calloway Co.: SIUC uncat. (1) West Fork Clarks
R. at Backusburg, 27 April 1969; SIUC uncat. (4) Beechy Cr., 1.6 km SE
New Concord, 23 March 1978.
Obion River Drainage. — Graves Co.: SIUC uncat. (138) Terrapin Cr., from
Tennessee state line to 0.8 km S Bell City, 23 March 1978 - 4 June 1980.
RESULTS AND DISCUSSION
Distribution and Habitat
Lampetra aepyptera is the most common lamprey in Kentucky,
occurring in all major drainages of the state except the Tradewater
River and lower Ohio and Mississippi River tributaries in the extreme
west (Burr 1980). The species is collected less frequently in the Kentucky
River system than in other major eastern Kentucky drainages (Fig. 1).
Its occurrence in direct, sandy tributaries of the Mississippi River (e.g.,
Obion River) in western Kentucky and Tennessee is somewhat unusual,
because this area is generally inhabited by fishes characteristic of the
Coastal Plain and lowlands.
Collections made in riffles and raceways of small to medium-size,
sand-gravel bottom creeks during March and April often yielded adults
of L. aepyptera. The lack of more records of this species for Kentucky is
almost surely the result of few collections being made in small streams
during early spring.
86
Stephen J. Walsh and Brooks M. Burr
Least Brook Lamprey in Kentucky 87
Taxonomic Status of lethenteron meridionale
Vladykov et al. (1975) described Lethenteron meridionale as a new
nonparasitic lamprey from Tennessee, Alabama and Georgia. In all
characters except dentition it is inseparable from L. aepyptera, despite
the fact that the authors concluded that it was a close relative of Lethen-
teron lamottenii (now Lampetra appendix; fide Bailey and Rohde in
Robins et al. 1980). As defined by Valdykov and his associates, mem-
bers of the genus Lethenteron always possess posterior circumoral teeth
and, partly because L. meridionale has posterior circumoral teeth, it was
accordingly assigned to the genus. However, Lampetra zanandreai,
which has been variously assigned to Lethenteron or Lampetra, occa-
sionally lacks posterior circumorals (Zanandrea 1957; Hubbs and Potter
1971).
We found small, unicuspid posterior circumoral disc teeth in spec-
imens of L. aepyptera from several drainages throughout Kentucky. The
mean number of posterior circumorals in specimens having them was
7.6 (R = 1 - 22, N = 15). In most, these teeth formed an incomplete row
(Fig. 2), but in two individuals there was a complete infraoral row
extending between the two posterior inner laterals. In another specimen
there were several small, degenerate teeth scattered throughout the pos-
terior field of the disc. Several collections have specimens with and
without posterior circumorals.
Fig. 2. Diagrammatic illustration of the oral disc of Lampetra aepyptera, show-
ing incomplete row of posterior circumoral teeth. Drawn from a composite of
several specimens.
We noted variation in several other disc characters used in the
diagnosis of L. meridionale. The following counts are means based on
17 specimens of L aepyptera: oral papillae 16.3 (R = 12 - 20); infraoral
cusps 9.8 (R = 5 - 13); intermediate teeth in anterior field 14.3 (R = 0 - 32);
inner lateral teeth 6.2 (R = 4 - 9). The counts for each of these characters
as reported by Vladykov et al. overlap between L. aepyptera and L.
88 Stephen J. Walsh and Brooks M. Burr
meridionale. Moreover, the means for number of infraoral cusps, inter-
mediate teeth in the anterior field, and inner lateral teeth in Kentucky
specimens of L. aepyptera, are closest to the means for these same char-
acters in L. meridionale (Vladykov et al. 1975, Table 11). Further, the
mean number of oral papillae and number of cusps (1 - 4) on the
median inner lateral teeth of Kentucky L. aepyptera are similar to the
means for L. aepyptera given by Vladykov et al. Thus, there is much
variation in the dentition of L. aepyptera. Individual variation, popula-
tional differences, and possibly ontogenetic changes in dentition, occur.
We therefore regard Lethenteron meridionale a synonym of Lampetra
aepyptera, a conclusion anticipated by Rohde (1980) and formally
stated by Bailey (1980). Okkelbergia (a monotypic genus including
aepyptera as its only species) and Lethenteron were recently down-
graded to subgenera of Lampetra (Bailey 1980), an action with which
we concur.
Morphology of lampetra aepyptera
The number of myomeres in L. aepyptera ranges from 50 to 62
(e.g., Branson 1970; Cook 1952; Raney 1941; Rohde 1976; Rohde et al.
1975, 1976; Seversmith 1953; Vladykov et al. 1975). Rohde et al. (1976)
found no significant difference in the number of myomeres between
ammocoetes and adults in Delaware. The number of myomeres of
adults in Kentucky ranged from 52 to 62 (Fig. 3), with means between
55 and 59 in all drainages except the lower Cumberland and Tennessee
rivers. The number of myomeres in 105 ammocoetes from Terrapin
Creek, Graves County, ranged from 51 to 59 (x = 55.0), slightly fewer
than for adults. However, the means of myomere counts are not signifi-
cantly different between ammocoetes and adults. Although visually
there appears to be an east to west decrease in the mean number of
myomeres (Fig. 3), one way analysis of variance indicates no significant
differences between the means of any of the drainages (a = 0.05). A tho-
rough study of specimens from throughout the entire range of L. aepyp-
tera is needed to determine possible significant variation in this or any
other morphological character.
The most significant differences in body proportions of L. aepyp-
tera involved the partially transformed (neotenic) individuals from Ter-
rapin Creek (Obion River drainage). The secondary sexual characteris-
tics of nuptial adults (e.g., disc length, eye diameter, second dorsal fin
height, and prebranchial length; Table 1) were poorly developed in the
neotenes, Because individuals in Terrapin Creek fail to fully transform,
the discordant body proportion values exhibited by the population were
expected. The Cumberland River population from above Cumberland
Falls exhibited a relatively low proportional value for second dorsal fin
height. This population is geographically isolated and the low fin value
Least Brook Lamprey in Kentucky
89
90 Stephen J. Walsh and Brooks M. Burr
may result from reproductive isolation from more typical L. aepyptera.
Sexual differences in body proportions (Table 2) coincide reasona-
bly well with those reported by other workers. Males have a greater disc
length and second dorsal fin height than females, as noted by Rohde et
al. (1976) and Vladykov et al. (1975). We confirm the observations of
Vladykov et al. that males have a relatively longer tail and shorter trunk
than females. Rohde et al. found that tail length increased with age in
males and decreased in females. Males also have a greater prebranchial
length. Kentucky specimens are substantially longer (mean TL, both
sexes) than those reported by most other authors, especially in Atlantic
Coast populations where a much shorter maximum and mean TL was
found (Rohde et al. 1976). The mean length of the male genital papilla
was 4.4 mm (R = 2.4 - 7.5, N = 57). Rohde et al. found a mean papilla
length of 4.8 mm for 10 males from Delaware.
Breeding coloration in L. aepyptera has not been adequately des-
cribed. Nest-building individuals from Peter Creek, Barren County,
Kentucky, were in full nuptial color orr 18 March 1980. Both sexes were
mottled gray-brown on the dorsum and had light silvery-yellow venters.
Black horizontal bands were present on the sides, through the eye and at
the base of the first dorsal fin. Each dorsal fin had a black, speckled
marginal band that was widest around the longest fin rays. A gold band
extended through the base of the caudal fin and the center of each dor-
sal fin. The posterior margin of the caudal fin was darkly pigmented.
Aspects of Life History
Terrapin Creek is a clear, low-to-moderate-gradient tributary of
Obion River. The stream consists of alternating shallow riffles and
deeper pools, with a substratum of sand and gravel. In most areas the
stream is 2 to 5 m wide and 0.2 to 1.0 m deep during normal water
levels, and is bordered by deciduous forest throughout most of its
course. Ammocoetes were generally found in mud banks along raceways
and in pools with a moderate current. Transformers and neotenic indi-
viduals were generally found in debris-ridden riffles and raceways.
Between 22 February and 4 June 1980 we sampled a backwater
slough of Terrapin Creek, 0.8 km south of Bell City along Hwy. 94. The
slough is in a wooded area, has been inundated through beaver activity,
and is fed by runoff and seepage. Thick mats of watercress and other
aquatic macrophytes were abundant in the channels and pools. A pool
2.5 m wide and 1.5 m deep, with considerable mud and silt deposits,
yielded large numbers of ammocoetes. Species taken with L. aepyptera
in this beaver slough are indicated by asterisks in Table 3.
Because of the paucity of specimens for most months, only ammo-
coetes from the 4 June 1980 collection were used for the length-
frequency analysis (Fig. 4). At least five, and possibly six, age clases are
Least Brook Lamprey in Kentucky
91
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Stephen J. Walsh and Brooks M. Burr
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Least Brook Lamprey in Kentucky 93
Table 3. Fishes collected with Lampetra aepyptera in Terrapin Creek, Ken-
tucky, arranged in approximate descending order of relative abun-
dance. Percentages are of total number of fishes collected between 23
March 1978 and 4 June 1980. Asterisks indicate species taken only
from ( * ), or primarily from ( ** ), beaver slough area.
represented. Young-of-the-year (age class 0) individuals had a mean TL
of 16 mm. Mean lengths for age classes I, II, and III were 38.6 mm, 65.0
mm, and 88.8 mm, respectively. The last curve on the frequency
polygon (Fig. 4) may include two additional age classes (IV, V). As
noted by other authors (Leach 1940; Hardisty 1961; Hardisty and Potter
94 Stephen J. Walsh and Brooks M. Burr
TOTAL LENGTH (mm)
Fig. 4. Length-frequency distribution of 45 ammocoetes of Lampetra aepyptera
from Terrapin Creek, Graves County, Kentucky, 4 June 1980. Curves were
smoothed using sliding averages of 7 mm. Age classes are represented by
Roman numerals.
1971; Potter and Bailey 1972; Rohde et al. 1976), difficulty arises in
discerning older age classes because there may be arrested growth of
larger ammocoetes, i.e., simultaneous occurrence of ammocoetes as
large or larger than transforming individuals in autumn collections.
Inadequate numbers of specimens from fall months prevented our test-
ing for a period of arrested growth for the Terrapin Creek population.
Transformers taken throughout the study period averaged 112 mm TL
(R = 95 - 125, N = 14); neotenic individuals averaged 103 mm TL (R =
75 - 129, N = 15). The largest ammocoetes from all collections averaged
111 mm TL (R = 102 - 135, N = 22). Assuming a March spawning
period, we estimate the duration of the larval stage in the Terrapin
Creek population to be 4.5 years. If a period of arrested growth occurs
in this population the length of the larval stage could be 5.5 years. A
larval life of 3 to 5.4 years for L. aepyptera has been suggested by others
(Hubbs 1971; Rohde et al. 1976; Seversmith 1953). Rohde et al. (1976)
and Rohde and Jenkins (1980) speculated that some individuals may
remain as larvae for over 6 years.
When lengths (L) and weights (W) of ammocoetes were trans-
formed to log values and the line of best fit determined by least squares,
the regression equation was Log W = 1.95 + 0.352 Log L, r = 0.99).
Rohde et al. (1976) found that the weights of ammocoetes from Dela-
ware increased as the 2.73 power of length. When we compared the
linear regression for transformers and neotenic individuals combined,
we found a slightly higher slope than that for ammocoetes (Log W =
1.96 + 0.248 Log L, r - 0.92).
Coefficient of condition values (K) were calculated for the Terrapin
Least Brook Lamprey in Kentucky 95
Creek specimens from the formula K = W X 105 / L3 (Carlander 1977).
Mean K values were as follows: ammocoetes, 0.15 (R - 0.10 - 0.23, N
= 107); transformers, 0.17 (R = 0.15 -0.19, N = 14); neotenes, 0. 13 (R =
0.10-0.17, N= 15). Rohdeet al. (1976) reported substantially higher K
values for L. aepyptera from Delaware, and found that the smallest
ammocoetes had the highest K values.
Nest building and spawning behavior of L. aepyptera have been
well documented by earlier workers (Brigham 1973; Rohde et al. 1976;
Pflieger 1975; Seversmith 1953). Nesting areas typically contain large
aggregations of lampreys and spawning usually involves pairs, or one
female accompanied by two males. Reported water temperatures at
which spawning occurs have ranged from 10 to 16° C (Brigham 1973;
Rohde et al. 1976; Seversmith 1953). Although spawning was not
observed in Terrapin Creek, we observed nest construction by three
individuals in the Barren River drainage (Peter Creek, 1.6 km SE Dry
Fork, Barren Co., 18 March 1980). Our observations are similar to
those of Rohde et al. (1976) and Seversmith (1953). Nest building
occurred at the crest of a shallow riffle (water 15 cm deep) approxi-
mately 1.8 m from shore, over sand and loose gravel; water temperature
was 12° C. Of the three individuals on the nest, one left when we
approached. We observed stone movement and lateral body undulations
by both sexes, as described by Rohde et al. and Seversmith. The male
appeared to be more active in attempting to excavate a depression.
Lampetra aepyptera in the upper Cumberland drainage (Taylor Branch,
8.8 km E Whitley City, McCreary Co.) were found spawning with no
apparent nest construction over a bedrock substratum on 16 April 1977
(W. C. Starnes, pers. comm.). As determined from preserved specimens,
individuals of L. aepyptera in spawning condition in Kentucky have
been taken as early as 22 February and as late as 1 June, with a peak of
spawning activity from mid-March to early April.
Reported male to female sex ratios of adults have ranged from 1:3
(Seversmith 1953) to 2.7:1 (Rohde et al. 1976). In the largest series of
adults from the Salt River drainage, Kentucky, this ratio was 2.3:1
(Knob Creek, Bullitt Co., 25-28 March 1950, N = 108).
Fecundity estimates for L. aepyptera have ranged from 610 to 2154
eggs produced per adult female (Rohde et al. 1976; Seversmith 1953;
Valdykov et al. 1975). Rohde et al. found a positive correlation between
TL of female and number of eggs produced. The number of mature ova
in three Kentucky specimens (115, 142, 142 mm TL) was 1802, 2596,
and 3816, respectively, and egg diameters ranged from 0.70 to 1.00 mm
(x = 0.86 mm, N = 43). The number of mature ova from five neotenic
females (105-1 19 mm TL, x = 1 12.4 mm) from Terrapin Creek was 572,
753, 1036, 1443, and 1541, and egg diameters ranged from 0.66 to 1.36
mm(x = 0.99 mm, N = 100). Our data for the neotenes show the trend
reported by Rohde et al. (1976), that a decrease in the number of eggs
96 Stephen J. Walsh and Brooks M. Burr
produced is correlated with an increase in egg size. Increased egg size
may result in a greater survival rate of fry, thereby countering the effects
of reduced fecundity. Adults maturing at smaller sizes apparently divert
a higher percentage of metabolic energy from growth to greater repro-
ductive efficiency.
Only two species of lamprey, both nonparasitic, have been documented
to have characteristics of neoteny — the European Lampetra zanan-
dreai (Zanandrea 1957, 1961), and Lampetra lethophaga (Hubbs 1971)
of northern California. During the spawning season, Zanandrea (1961)
found 12 female ammocoetes of L. zanandreai that were in an advanced
("third") stage of ovarian development. The type of neoteny described
by Hubbs (1971) for L. lethophaga involved "the maturing of appar-
ently all individuals of both sexes in the prenuptial condition."
In Terrapin Creek, from late February through early June (the
spawning season has variedgreatly in the last three years, because win-
ters have varied in severity), 7 neotenic males and 6 females were col-
lected. They had passed through partial transformation but, although
some were in full maturity (Fig. 5D shows a females turgid with ripe
ova) none had developed the ordinary nuptial attributes (e.g., eye
diameter, melanistic pigmentation, englargement of the two dorsal fins,
development of the anal pseudo-fin, and enlargement of the disc). Three
neotenic females from late February have large, ripe ova visible through
the transparent body wall, a distinctive characteristic of mature females.
Thus, a situation very similar to that reported by Hubbs (1971) for L.
lethophaga seems to also have developed in the Terrapin Creek popula-
tion of L. aepyptera. This is apparently the normal condition for the
Terrapin Creek population, inasmuch as numerous collections contain
no nuptial adults, which are the kind of adults normally collected dur-
ing the spring throughout the remainder of Kentucky (Fig. 5A, C).
Since temperatures in Terrapin Creek are similar to those of other
streams that contain normal adults of L. aepyptera, unusual tempera-
tures are ruled out as a factor suppressing nuptial development.
The low K values of L. aepyptera in Terrapin Creek, and overall
low numbers of fishes in a stream that otherwise maintains a diversity of
at least 30 species (Table 3), indicate that food may be scarce. The fail-
ure of this population to exhibit normal secondary sexual characters is
tentatively attributed to limited food during the ammocoete stage, with
a resultant lack of sufficient lipid accumulation to bring about the com-
pletion of transformation to fully nuptial adults.
Least Brook Lamprey in Kentucky
97
Fig. 5. Specimens of Lampetra aepyptera from Kentucky. A, Fully nuptial male
(WCS 720-01, 136.6 mm TL) from Taylor Branch, McCreary County, 16 April
1977. B, Partially neotenic male (SIUC uncat., 101.6 mm TL) from Terrapin
Creek, Graves County, 4 April 1980. C, Fully nuptial female, same data as A
(147.2 mm TL), with abdominal wall pinned aside to show coelom packed with
mature ova. D, Partially neotenic female, same data as B (1 14.8 mm TL), with
abdominal wall pinned aside to show coelom packed with mature ova.
98 Stephen J. Walsh and Brooks M. Burr
ACKNOWLEDGMENTS.— M. E. Braasch, K. S. Cummings,
S. L. Dewey, R. L. Mayden, L. M. Page and M. E. Retzer assisted with
field work. Karen A. Schmitt of the Scientific Photography Illustration
Facility, Southern Illinois University at Carbondale (SIUC) Graduate
School, assisted in the preparation of figures. Drs. Ronald A. Brandon,
SIUC, and John E. Cooper, North Carolina State Museum of Natural
History, greatly improved earlier versions of the manuscript.
We are grateful to the following individuals and institutions for
providing loans and/ or laboratory space: Branley A. Branson, Eastern
Kentucky University, Richmond (EKU); Lawrence M. Page, Illinois
Natural History Survey, Champaign (INHS); Melvin L. Warren, Jr.,
Kentucky Nature Preserves Commission, Frankfort (KNPC); Frank B.
Cross, University of Kansas, Lawrence (KU); Ernest A. Lachner, Na-
tional Museum of Natural History, Washington D.C. (USNM); William
D. Pearson, University of Louisville, Louisville (UL); and Wayne C.
Starnes, University of Tennessee, Knoxville (WCS).
LITERATURE CITED
Bailey, Reeve M. 1980. Comments on the classification and nomenclature of
lampreys — an alternative view. Can. J. Fish. Aquat. Sci. 57(1 1):.1626-1629.
Branson, Branley A. 1970. Measurements, counts, and observations on four
lamprey species from Kentucky (Ichthyomyzon, Lampetra and Entosphe-
nus). Am. Midi. Nat. M(l):243-247.
Brigham, Warren U. 1973. Nest construction of the lamprey, Lampetra aepyp-
tera. Copeia 1973( 1): 135-136.
Burr, Brooks M. 1980. A distributional checklist of the fishes of Kentucky.
Brimleyana No. 3:53-84.
, and R. L. Mayden. 1979. Records of fishes in western Kentucky with
additions to the known fauna. Trans. Ky. Acad. Sci. 40(l-2):58-67.
Carlander, Kenneth D. 1977. Handbook of Freshwater Fishery Biology. Vol. 2.
Iowa State Univ. Press, Ames. 431 pp.
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Frankfort. 416 pp.
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(Abbott), in Kentucky. Trans. Ky. Acad. Sci. 18{\)\ 12-1 3.
Cook, Fannye A. 1952. Occurrence of the lamprey Lampetra aepyptera in the
Tombigbee and Pascagoula drainages, Mississippi. Copeia 1952(4):268.
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, and I. C. Potter. 1971. The behaviour, ecology and growth of larval
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ogy of Lampreys. Academis Press, London. 423 pp.
Least Brook Lamprey in Kentucky 99
Hubbs, Carl L. 1971. Lampetra (Entosphenus) lethophaga , new species, the
nonparasitc derivative of the Pacific lamprey. Trans. San Diego Soc. Nat.
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, and I. C. Potter. 1971. Distribution, phylogeny and taxonomy, pp. 1-65
in M. W. Hardisty and I. C. Potter (eds.). The Biology of Lampreys. Aca-
demic Press, London. 423 pp.
Leach, W. James. 1940. Occurrence and life history of the northern brook
lamprey, Ichthyomyzon fossor , in Indiana. Copeia 1940(l):21-34.
Pflieger, William L. 1975. The Fishes of Missouri. Mo. Dep. Conserv., Jefferson
City. 343 pp.
Potter, Ian C. 1980. The Petromyzoniformes with particular reference to paired
species. Can. J. Fish. Aquat. Sci. 57(1 1): 1595-1615.
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Raney, Edward C. 1941. Records of the brook lamprey, Lampetra aepyptera
(Abbott), from the Atlantic drainage of North Carolina and Virginia. J.
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Robins, C. Richard, R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R.
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Fishes from the United States and Canada. 4th ed. Am. Fish. Soc. Spec.
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100 Stephen J. Walsh and Brooks M. Burr
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Accepted 11 September 1981
Notes on the Distribution and Taxonomy of Short-
tailed Shrews (Genus Blarina) in the Southeast
Thomas W. French '
Department of Life Sciences,
Indiana State University, Terre Haute, Indiana 47809
ABSTRACT. — Seven hundred twenty-nine skulls of short-tailed
shrews (genus Blarina) were examined from 152 counties in Alabama,
Florida, Mississippi, North Carolina, South Carolina and Tennessee.
One hundred ninety-three B. brevicauda and 265 B. carolinensis were
compared by a stepwise discriminant analysis. Twenty-one central
Tennessee specimens were compared to these two identified target
samples. Although specimens from central Tennessee are scarce, the
cranial measurements of some appear intermediate in size. Plots of the
first two canonical variables show specimens from Hickman, Putnam
and Warren counties, Tennessee as distinct from either B. brevicauda
or B. carolinensis target clusters. A partial distribution map defining
the ranges of B. brevicauda and B. carolinensis in the Southeast is
presented. A possible disjunct population of B. brevicauda is reported
from both sides of the Chattahoochee River in Alabama and Georgia.
INTRODUCTION
Short-tailed shrews of the genus Blarina are the most abundant and
ubiquitous soricids in the Southeast. The taxonomy of this genus is cur-
rently undergoing revision, but recent publications (Genoways and
Choate 1972; Ellis et al. 1978; Schmidley and Brown 1979; Tate et al.
1980) recognize a large northern form, Blarina brevicauda, and a small
southern form, Blarina carolinensis, as distinct species. Another large
phena, B. telmalestes, restricted to the vicinity of the Great Dismal
Swamp of Virginia and North Carolina, is also currently recognized as
distinct (Jones et al. 1979). Handley (1971) considered B. brevicauda
and B. carolinensis to be entirely allopatric but contiguous, and Graham
and Semken (1976) considered them to represent the parapatric coexis-
tence of sibling species. Some early workers reported a zone of intergra-
dation between the two phena which were then recognized as well dif-
ferentiated subspecies (Merriam 1895; Cockrum 1952; Jones and Findley
1954), while others were unable to recognize intergrades (Rippy 1967;
Schlitter and Bowles 1967). More recent workers have found individual
areas of sympatry between B. brevicauda and B. carolinensis, both in
Recent (Genoways and Choate 1972; Ellis et al. 1978; Tate et al. 1980)
and Pleistocene material (Graham and Semken 1976), with only isolated
cases of possible hybrids (Genoways and Choate 1972; Tate et al. 1980).
'Current address: Cooperative Wildlife Research Unit, Department of Natural
Resources, Cornell University, Ithaca, New York 14853
Brimleyana No. 6: 101-110. December 1981 101
102 Thomas W. French
Although these shrews are quite common, their exact ranges and
thus the location of possible zones of sympatry are still not well known.
The purpose of this paper is to more accurately define their ranges in
the Southeast, to point out the intermediate nature of some Tennessee
specimens, and to report a possible disjunct population of B. brevicauda
in the upper Coastal Plain of Alabama and Georgia.
MATERIALS AND METHODS
Seven hundred twenty-nine Blarina skulls from 152 counties in
Alabama, Florida, Mississippi, North Carolina, South Carolina, and
Tennessee were examined. Five cranial measurements (condylobasal
length, cranial breadth, interorbital breadth, maxillary breadth and
maxillary toothrow length) were made to the neareest 0.1 mm with a
vernier caliper following Jackson (1928). Specimens were measured
regardless of sex or age, but included no nestlings.
Although age dimorphism has been documented by some (Guilday
1957; Choate 1972) and not by others (Ellis et al. 1978) Blarina is consi-
dered to be essentially adult size by the time it enters the trappable
population (Guilday 1957; Dapson 1968; Ellis et al. 1978). The most
noticeable differences between juvenile and adult Blarina are an increase
in weight, total body length, and tooth wear, and a decrease in cranial
height, with age. None of these characters was used to differentiate B.
brevicauda and B. carolinensis in this study. Sexual dimorphism in Blar-
ina has been recognized as slight by most workers, with males averaging
slightly larger than females in some characters (Guilday 1957; Dapson
1968; Choate 1972; Ellis et al. 1978; Kirkland 1978). Others have
reported no detectable sexual dimorphism (Graham and Semken 1976;
Schmidley and Brown 1979). Neither age nor secondary sexual dimor-
phism of cranial characters appear to be significant when differentiating
specimens of B. brevicauda and B. carolinensis in the trappable popula-
tion. This presumption is supported by the near lack of overlap in cran-
ial measurements between these two taxa reported by recent authors.
Body measurements were not used because standard body meas-
urements were found to be variable, even within local populations, and
especially because of obvious discrepancies in measuring techniques
between various collectors. Guilday (1957) and Jones and Glass (1960)
stressed that external measurements of Blarina (unless made by the
same worker) should be used with caution in geographic studies. They
also stressed that cranial measurements are much more constant and
can be more accurately measured than body measurements. Sample
body measurements of 50 B. brevicauda from Alabama and South
Carolina are: total length 114.6 (101-125), tail length 25.2 (21.0-29.5),
and hindfoot length 14.1 (12.5-15.5). Sample body measurements of 50
B. carolinensis from these same two states are: total length 92.2 (85-
Distribution and Taxonomy of Blarina 103
104), tail length 19.7 (14-27), and hind foot length 11.2(10.0-12.5).
Although specimens from central Tennessee are scarce, preliminary
analysis of cranial measurements indicated that specimens from this
area might be intermediate in size. A stepwise discriminant analysis was
conducted on 265 B. carolinensis and 193 B. brevicauda with complete
measurement data. The five previously described cranial measurements
were used in this analysis and the specimens represent localities
throughout the Southeast. Twenty-one central Tennessee specimens
were then compared to these target samples. Seven of the central Ten-
nessee specimens lacked condylobasal length and maxillary breadth
measurements and two others lacked only maxillary breadth measure-
ments due to breakage. Missing data were estimated for the nine speci-
mens using the REGR option in the PAM subroutine of the Biomedical
Computer Programs (Brown and Dixon 1979). In order to obtain a
visual representation, the first two canonical variables were computed
and plotted as described by Rao (1952) and used by Lawrence and Bos-
sert (1967, 1969), Gipson et al. (1974), Kirkland and Van Deusen (1979),
Parkinson (1979), Diersing (1980), and others. The Biomedical program
PAM was used for these calculations.
RESULTS
The number of specimens examined from any one county varied
from one to seventy-five. Small ranges in cranial measurements from
large series suggest that small samples, other than from near the zone of
contact, can usually be considered representative of the local popula-
tion. Only slight overlap was found between the cranial measurements
of all B. brevicauda and B. carolinensis (Table 1). Cranial measurements
(mm) of two very recently weaned B. brevicauda from Alabama were
condylobasal length 21.0 (broken); cranial breadth 11.2, 11.9; interorbi-
tal breadth 5.5, 5.8; and maxillary toothrow length 8.6, 8.9, maxillary
breadth 7.6, 7. 9. The lower range of each of these measurements is as
great as or greater than the upper range of the same measurements from
a mixed age sample of B. carolinensis from the Southeast (Table 1).
Perimeters of the extreme ranges of canonical variables for B. brev-
icauda and B. carolinensis are shown in Figure 1 to be nonoverlapping.
Individual specimens from central Tennessee are identified in this figure
by the first letter or letters of the county in which they were collected
(Anderson, Davidson, Franklin, Hickman, Marion, Putnam, Warren
and Wayne).
The most striking pattern is the clustering of the Hickman, Putnam
and Warren county specimens well outside the range of both B. brevi-
cauda and B. carolinensis canonical clusters. The Marion County spec-
imen appears properly identified as a B. carolinensis, although it is
located on the edge of this distribution. Seven Anderson and one Frank-
104 Thomas W. French
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Distribution and Taxonomy of Blarina 105
B. brevicauda B. carolinensis
Wr
-3
canonical variable l
Fig. 1. Relationships of 479 Blarina from the Southeast as plotted by discrimi-
nant analysis (BMD P7M). Stars represent group means for B. brevicauda and
B. carolinensis. Letters represent individual specimens from central Tennessee
and are the first letter or letters of the counties in which the shrews were col-
lected (Anderson, Davidson, Franklin, Hickman, Marion, Putnam, Warren and
Wayne).
lin County specimens fall within the B. brevicauda cluster, but the
remaining specimens are located outside the limits of this cluster and
intermediate to the two Blarina species.
Blarina from Anderson, Davidson, Franklin, Lincoln and Wayne
counties are probably best referred to as B. brevicauda and the Marion
County specimen as B. carolinensis. The possibility of hybridization or
intergradation between the two forms of Blarina, however, should not
be ruled out.
Most Tennessee B. brevicauda are found in the mountainous parts
of the state on the eastern and northern borders. Wayne County is
located on the Highland Rim and is not particularly high in elevation,
but the shrews from Franklin and Lincoln counties were collected on
the Cumberland Plateau at about 2000 feet (610 m) elevation. These
large shrews might represent a relict population of B. brevicauda, but it
seems more likely that they are joined to other B. brevicauda popula-
tions along the length of the Cumberland Plateau. If this is true, large
specimens of Blarina should be looked for at higher elevations in coun-
ties such as Bledsoe, Cumberland, Grundy and Sequatchie.
Many authors (Lawrence and Bossert 1967, 1969; Gipson et al.
1974; Kirkland and Van Deusen 1979; Parkinson 1979; and others) have
106 Thomas W. French
attributed the magnitude of difference between the Hickman, Putnam,
and Warren counties canonical cluster and either B. brevicauda or B.
carolinensis target clusters as representative of distinct species or species
hybrids. Although these results suggest that Blarina in central Tennessee
might be distinct from both B. brevicauda and B. carolinensis, their
correct identity will remain uncertain until more comparative material is
available.
I produced (Fig. 2) a partial distribution map of Blarina using the
five cranial characters as species criteria. The most notable difference
between this and other distribution maps (Hall and Kelson 1959; Hand-
ley 1971; Graham and Semken 1976; Tate et al. 1980; and others) is the
south and westward extension of B. brevicauda (synonym = B. brevi-
cauda churchi) approximately 195 miles (314 km) from the mountains
of Georgia into the Piedmont of Alabama. Also indicated is a possible
disjunct population of B. brevicauda on both sides of the Chattahoochee
River, Barbour County, Alabama, and in Quitman and Stewart coun-
ties, Georgia. In Alabama the largest form was overlooked by Howell
(1921) because none of his specimens of Blarina came from the
Piedmont.
Much of the southeastern distribution of these shrews approxi-
mates well established physiographic boundaries (see Fenneman 1938
and Hunt 1964 for descriptions of physiographic provinces). In North
Carolina the boundary between the two species roughly follows the
eastern edge of the mountains, but in South Carolina it extends south-
ward through the Piedmont and meets the Savannah River near the
center of this physiographic province. In Georgia and Alabama the
boundary closely follows the Fall Line, and in Alabama it swings north-
eastward along the south edge of the Great Valley (between the Pied-
mont and Ridge and Valley physiographic provinces). In Tennessee it
follows the western edge of the Smoky Mountains and appears to swing
around the southern edge of the Cumberland Mountains and south of
the Cumberland River. Although habitat features are often considerably
different in adjacent physiographic provinces, I know of no reason why
these features should limit the distribution of either form of Blarina.
Blarina brevicauda was discovered at three localities south of the
Fall Line in Georgia and Alabama. There are few specimens (10), but
the three localities appear to represent a disjunct population separated
from the Piedmont populations by only about 25 air miles (40 km). The
only other known disjunct populations of large Coastal Plain Blarina
are B. brevicauda shermani on the Gulf coast of Florida and B. telma-
lestes in the Great Dismal Swamp of Virginia and adjacent North Carol-
ina. Two large specimens (U.S. National Museum #268977 and North
Carolina State Museum #2575) were also examined from the Coastal
Plain of North Carolina (Sampson and Columbus counties). These are
the only individuals examined from each of these two counties. The
Distribution and Taxonomy of Blarina
107
108 Thomas W. French
specimens were collected at least 110 miles (177 km) east of B. brevi-
cauda populations of the North Carolina Piedmont and 125 miles (201
km) southwest of the nearest known B. telmalestes populations. Addi-
tional work is needed in the North Carolina Coastal Plain.
ACKNOWLEDGMENTS.— Appreciation is extended to the follow-
ing persons who allowed me to examine specimens under their care: R.
J. Baker, Texas Tech University; R. Bauer, Cornell University; G.
Breece, Georgia Power Co., Atlanta; J. E. Cadle, University of Georgia;
C. Carter, Mississippi Museum of Natural Science, Jackson; J. L. Dusi,
Auburn University; R. D. Fisher, National Museum of Natural History;
P. Goldstein, American Museum of Natural History; D. C. Holliman,
Birmingham Southern College; M. Kennedy, Memphis State University;
D. S. Lee, North Carolina State Museum of Natural History; J. P.
O'Neill, Louisiana State University; J. F. Parnell, University of North
Carolina at Wilmington; C. Ruckdeschel, Cumberland Island, GA; A.
E. Sanders, Charleston Museum; S. Scudder, Florida State Museum,
University of Florida; C. H. Wharton, Georgia State University; W. K.
Willard, Tennessee Technological University; and H. C. Yeatman, Uni-
versity of the South.
Special thanks are due A. E. Sanders and J. L. Dusi for their help,
and J. O. Whitaker, Jr., Indiana State University, for reading the
manuscript. The comments of two anonymous reviewers were also
greatly appreciated. Computer funds and typing were supplied by the
Cooperative Wildlife Research Unit, Cornell University.
LITERATURE CITED
Brown, M. B., and W. J. Dixon (eds.). 1979. Biomedical computer programs P
series. Univ. Calif. Press, Berkeley. 880 pp.
Choate, Jerry R. 1972. Variation within and among populations of the short-
tailed shrew in Connecticut. J. Mammal. 53:1 16-128.
Cockrum, E. Lendell. 1952. Mammals of Kansas. Univ. Kansas Publ. Mus. Nat.
Hist. 7:1-303.
Dapson, Richard W. 1968. Growth patterns in a post-juvenile population of
shorttailed shrews (Blarina brevicauda). Am. Midi. Nat. 79:118-129.
Diersing, Victor E. 1980. Systematics and evolution of the pygmy shrews (sub-
genus Microsorex) of North America. J. Mammal. <5/:76-101.
Ellis, L. Scott, V. E. Diersing and D. F. Hoffmeister. 1978. Taxonomic status of
short-tailed shrews (Blarina) in Illinois, J. Mammal. 59:305-31 1.
Fenneman, Nevin M. 1938. Physiography of Eastern United States. McGraw-
Hill Co., Inc., New York. 689 pp.
Genoways, Hugh H., and J. R. Choate. 1972. A multivariate analysis of system-
atic relationships among populations of the short-tailed shrew (genus
Blarina) in Nebraska. Syst. Zool. 27:106-1 16.
Distribution and Taxonomy of Blarina 109
Gipson, Philip S., J. A. Sealander and J. E. Dunn. 1974. The taxonomic status
of wild Canis in Arkansas. Syst. Zool. 23:1-1 1.
Graham, Russell W., and H. A. Semken. 1976. Paleoecological significance of
the short-tailed shrew (Blarina), with a systematic discussion of Blarina
ozarkensis. J. Mammal. 57:433-449.
Guilday, John E. 1957. Individual and geographic variations in Blarina brevi-
cauda from Pennsylvania. Ann. Carnegie Mus. 55:41-68.
Hall, E. Raymond, and K. R. Kelson. 1959. The Mammals of North America.
Ronald Press, New York. 546 pp.
Handley, Charles O., Jr. 1971. Appalachian mammalian geography -- Recent
Epoch, pp. 263-303 in P. C. Holt (ed.). The distributional history of the
biota of the southern Appalachians, Part III: Vertebrates. Res. Div.
Monogr. 4, Va. Polytech. Inst. State Univ., Blacksburg. 306 pp.
Howell, Arthur H. 1921. Mammals of Alabama. N. Am. Fauna 45. 88 pp.
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Co., San Francisco. 480 pp.
Jackson, Hartley H. T. 1928. A taxonomic review of the American long-tailed
shrews. N. Am. Fauna 51. 238 pp.
Jones, J. Knox, Jr., and J. S. Findley. 1954. Geographic distribution of the
short-tailed shrew, Blarina brevicauda, in the Great Plains. Trans. Kans.
Acad. Sci. 57:208-211.
, and B. P. Glass. 1960. The short-tailed shrew, Blarina brevicauda,
in Oklahoma. Southwest. Nat. 5:136-142.
, D. C. Carter and H. H. Genoways. 1979. Revised checklist of
North American mammals north of Mexico, 1979. Occas. Pap. Mus. Texas
Tech. Univ. 62:1-17.
Kirkland, Gordon L. 1978. The short-tailed shrew, Blarina brevicauda (Say), in
the central mountains of West Virginia. Proc. Pa. Acad. Sci. 52:126-130.
, and H. M. Van Duesen. 1979. The shrews of the Sorex dispar
group: Sorex dispar Batchelder and Sorex gaspensis Anthony and Good-
win. Am. Mus. Novit. 2675:1-21.
Lawrence, Barbara, and W. H. Bossert. 1967. Multiple character analysis of
Canis lupus, latrans and familiaris , with a discussion of the relationships of
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Rippy, Charles L. 1967. The taxonomy and distribution of the short-tailed
shrew, Blarina brevicauda, in Kentucky. Unpubl. M.S. thesis, Dept. Zool.,
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of some mammals in Iowa. Trans. Kans. Acad. Sci. 70:525-529.
110 Thomas W. French
Schmidley, David J., and W. A. Brown. 1979. Systematics of short-tailed shrews
(genus Blarina) in Texas. Southwest. Nat. 24:39-48.
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Accepted 20 March 1981
Observations of a Small Population
of Estuarine-Inhabiting Alligators
Near Southport, North Carolina
William S. Birkhead
Division of Science and Mathematics,
Columbus College, Columbus, Georgia 31993
AND
Charles R. Bennett
National Marine Fisheries Service,
Environmental Assessment Branch, Portland, Oregon 97208
ABSTRACT. — Field observations of the American Alligator, Alliga-
tor mississippiensis, were made incidental to a five-year study of the
nursery use of Dutchman Creek estuary near Southport, North Carol-
ina, by estuarine dependent fishes and shell fishes. Alligators were
most frequently seen between April and July and occurred over a wide
range of salinities. Some individuals that inhabited the lower reaches
of the estuary appeared to have shifted their distribution in response to
a major alteration of this habitat. After the headwaters and principal
tributaries of Dutchman Creek were bisected by the discharge canal of
a nuclear power plant, alligators appeared to move out of the creek
and into the lower reaches of its severed tributaries adjacent to a
drainage canal that received their diverted flow.
INTRODUCTION
Numerous sightings of alligators were made incidental to a five-
year study of Dutchman Creek estuary, a tidal creek and salt marsh
habitat located approximately 2 km west of Southport, Brunswick
County, North Carolina (Birkhead et al. 1977). Since the alligator is an
endangered species, and so little is known about the biology of northern
populations, we felt that these observations were of interest. The fact
that the habitat involved underwent a pronounced change after our
study began made the situation even more interesting.
In its unaltered state, most of the freshwater flow into Dutchman
Creek came from its headwaters and from three principal tributaries to
the north and west. Tidal water flushed in and out twice a day by enter-
ing the creek from the Intracoastal Waterway, which bisected its lower
reaches. Thus, a typical estuarine condition was established, with a
salinity gradient ranging from an average of approximately 2 ppt in the
upper reaches to an average of 20 ppt near its mouth (Birkhead et al.
1977). Marshes adjacent to the lower reaches of the creek and tributar-
ies are flooded regularly with the tides. Spartina alterniflora is the dom-
Brimleyana No. 6: 1 1 1-1 17. December 1981. Ill
112
William S. Birkhead and Charles R. Bennett
inant vegetation in these areas. As ground elevation increases slightly in
the upper reaches of the estuary, the regularly flooded marsh grades
into irregularly flooded marsh dominated by Juncus roemerianus (Seneca
etal. 1976).
Carolina Power and Light Company began constructing a nuclear-
fueled, steam electric generating plant just north of Southport approxi-
mately two years before our study began. This facility was designed to
employ a once-through cooling system. The discharge canal for the
cooling system was constructed during the summer and fall of 1972,
along the northwestern and western edge of the salt marsh bordering
Dutchman Creek. This was between 9 and 16 months after our study
began. Freshwater runoff from the upper reaches of the bisected tribu-
taries and headwaters of Dutchman Creek was diverted into a drainage
canal dredged adjacent to the discharge canal (Fig. 1). The drainage
canal emptied into the Intracoastal Waterway approximately 1.5 km
west of the mouth of Dutchman Creek. Although the hydrographic
regime of the lower reaches of the Dutchman Creek estuary was not
appreciably affected by the diversion of the freshwater input, salinity
increases of between 13 and 15 ppt were recorded in the upper reaches
of the estuary below the canal right-of-way and at the mouths of the
severed headwaters and tributaries above these canals (Birkhead et al.
1977).
The primary purpose of this paper is to denote the distribution of
alligators within the Dutchman Creek estuary before, during, and after
its alteration.
Fig. 1. Map of Dutchman Creek estuary with sampling stations.
Estuarine-Inhabiting Alligators 113
RESULTS AND DISCUSSION
Although alligators were observed during every month of the year,
the majority of sightings (76%) occurred between April and July (Table
1). The few fall and winter sightings were probably of a single alligator
that inhabited the upper reaches of Dutchman Creek below the canal
rights-of-way in the vicinity of station 2.
Prior to the diversion of the headwaters and principal tributaries of
Dutchman Creek (sampling year 1971-72), all alligators sighted were
downstream from what were then the proposed canal rights-of-way
(Table 2). After the freshwater input was diverted (sampling years 1972-
76), alligators virtually disappeared from the lower reaches of Dutch-
man Creek itself, and the majority of sightings were made in or along
the northwestern and western side of the freshwater drainage canal, par-
ticularly in the vicinity of the severed headwaters and tributaries. De-
spite these spatial shifts, alligators appeared to be randomly distributed
within the salinity regimes that were present in the estuary before and
after alterations were made (Table 3).
Alligator sightings were bimodal with respect to water temperature
(Table 3). Although most alligators (67%) were sighted when water
temperatures were between 22°C and 30°C, several (26%) were seen
when water temperatures were in the teens. The latter observations fre-
quently involved animals that were basking during the spring, a season
when air temperatures were undoubtedly higher than water temperatures.
Most of the alligators whose size could be estimated (Table 4)
appeared to be between 1 and 2 m total length. The smallest individual
seen was about 0.6 m long, while the largest appeared to be in excess of
2.4 m.
Since alligators observed during the course of this study were not
marked, long term movements of individual animals could not be ascer-
tained. Nevertheless, our sightings provide evidence for spatial shifts in
the population inhabiting the Dutchman Creek estuary following block-
age and diversion of its headwaters and principal tributaries. We suggest
these reptiles may have moved because they required access to fresh
water.
It is well known that tidal creeks and their associated salt marshes
are highly productive habitats that serve as nursery areas for large
numbers of fishes and shellfishes (Weinstein 1979). However, Chabreck
(1971) and Joanen et al. (1971) noted that immature and hatchling alli-
gators that remained under saline conditions for extended periods of
time experienced significant weight losses despite the fact that food was,
or appeared to be, readily available. Both authors attributed weight loss
to reduced food intake. It is entirely possible that estuarine-inhabiting
alligators can only make use of this productivity by either allocating
their feeding activities to periods of low tide or by hunting in the brack-
ish parts of the estuary where salinities are low and concentrations of
114
William S. Birkhead and Charles R. Bennett
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Estuarine-Inhabiting Alligators
115
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116 William S. Birkhead and Charles R. Bennett
Table 4. Estimated lengths of alligators sighted in Dutchman Creek estuary
from September 1971 through August 1976.
fish and shellfish are relatively high. In keeping with this hypothesis, the
alligators that we observed in the lower reaches of the Dutchman Creek
estuary in the first year of study (1971-72) may have been only tempor-
ary inhabitants whose presence was due to a plentiful food supply.
Another reason for the distributional shift may be related to nest-
ing requirements. Joanen (1969) found that most alligators on the
Rockefeller Refuge in coastal Louisiana nested in natural marsh consist-
ing primarily of wiregrass, Spartina patens. Salinities in these marshes
averaged 3.8 ppt. No nests were transected in the more saline marsh
type (Joanen et al. 1971). When questioned as to where estuarine-
inhabiting alligators in the vicinity of Southport nested, long-time resi-
dents invariably stated that nesting occurred along the upper reaches of
estuarine tributaries in a narrow zone characterized by freshwater marsh
vegetation. Because this type of habitat was not sampled in our study in
the Dutchman Creek estuary, such statements could not be verified for
North Carolina. This kind of marsh, although limited in extent, would
seem to be a more favorable nesting habitat than the salt marsh proper,
where the danger of inundation during storm tides would undoubtedly
be greater. In support of this belief is the observation that nearly all
juvenile (< 1 m) alligators seen in the vicinity of Southport were in
freshwater habitats. At one locality in particular, one of us (WSB)
repeatedly saw several alligators as small as 0.5 m long in a shallow
freshwater impoundment created by diking off approximately 10 ha of
saltmarsh adjacent to a tidal creek about 6 km north-northeast of
Southport.
The number of alligators sighted in the Dutchman Creek estuary
during the 5-year study remained relatively constant from year to year
despite a major alteration in their habitat. However, continued survival
of individuals now inhabiting the freshwater drainage canal could be
jeopardized because this canal has made their habitat more accessible to
humans.
Estuarine-Inhabiting Alligators 117
ACKNOWLEDGMENTS.— We sincerely appreciate the efforts of
several technicians, particularly J. Lee, D. Herring, L. Hall and J.
Schneider who assisted us in the field, and Thomas Sanders who
reviewed an early draft of the manuscript. Our studies of the Dutchman
Creek estuary were supported by Carolina Power and Light Company,
Raleigh, North Carolina.
LITERATURE CITED
Birkhead, William S., C. R. Bennett, E. C. Pendleton and B. J. Copeland. 1977.
Nursery utilization of the Dutchman Creek Estuary, N. C. 1971-1976.
Report of Carolina Power and Light Co., Raleigh. 258 pp.
Chabreck, Robert H. 1971 . The foods and feeding habits of alligators from fresh
and saline environments in Louisiana. Proc. Southeast. Assoc. Game Fish
Commissioners Conf. 25:1 17-124.
Joanen, Ted. 1969. Nesting ecology of alligators in Louisiana. Proc. Southeast.
Assoc. Fish Game Commissioners Conf. 23: 141-151.
, L. McNease, H. Dupuie and W. G. Perry. 1971. Louisiana
Wildlife and Fisheries Commission Fourteenth Biennial Report, 1970-1971.
Seneca, Ernest D., L. Stroud, U. Blum and G. R. Noggle. 1976. An analysis
of the effects of the Brunswick Nuclear Power Plant on the productivity of
Spartina alterniflora (smooth cordgrass) in the Dutchman Creek, Oak
Island, Snow's Marsh, and Walden Creek Marshes, Brunswick County,
North Carolina, 1975-1976. Third Annual Report to Carolina Power and
Light Co., Raleigh. 335 pp.
Weinstein, Michael P. 1979. Shallow marsh habitats as primary nurseries
for fishes and shellfish. Cape Fear River, North Carolina. Fish. Bull.
77:339-357.
Accepted 29 July 1 98 1
A Key to the Tadpoles of North Carolina
Joseph Travis1
Department of Zoology,
Duke University, Durham, North Carolina 27706
ABSTRACT.— A dichotomous key for identifying the tadpoles of
North Carolina and a guide for their field identification with a hand
lens are offered. Problems in identifying tadpoles are discussed. The
key should be useful throughout the southeastern United States,
because it treats most of the species present in this area.
INTRODUCTION
Although anuran larvae are excellent subjects for various types of
field and laboratory research, the difficulty of correctly identifying tad-
poles is well known. Early keys were either difficult to use (e.g., Wright
and Wright 1949) or restricted in taxonomic (e.g., Orton 1952) or geo-
graphic (e.g., Smith 1934) coverage. Altig (1970) provided a key to all of
the tadpoles found in the continental United States and Canada, and
proposed a standard terminology for use in identifying tadpoles. Later
references to Altig will be to this 1970 paper. A complete key to variable
animals such as anuran larvae can prove difficult to use. Geographic
variation can produce ambiguities in identification, and characters that
may be diagnostical at a local level may prove impossible to integrate
into a more thorough treatment.
North Carolina, with 30 species of anurans (Martof et al. 1980),
provides an excellent situation for the development of a more restricted
key. The extensive phenotypic variability seen in some species, such as
Hyla crucifer and Rana clamitans, often precludes the effective use of
Altig's key in the state. A North Carolina key should also be useful in
the southeastern United States, because it includes most of the species
found in this area.
My local key was constructed from Altig's and others in the litera-
ture, specimens collected by me and others during my four years in
North Carolina, and laboratory rearings of unidentified tadpoles. Some
key characters used by other workers subsequently proved unreliable
and have been deleted. I examined many specimens of 27 of the 30
North Carolina species, including living individuals. Pseudacris brim-
leyi, P. brachyphona, and Rana heckscheri were not personally exam-
ined. Hyla versicolor was obtained from Giles County, Virginia, and
specific identity was verified by karyotype analysis. The occurrence of
H. versicolor in North Carolina remains problematic (Martof et al.
1980).
Present address: Department of Biological Science, Florida State University,
Tallahassee, Florida 32306
Brimleyana No. 6: 1 19-127. December 1981. 119
120 Joseph Travis
USE OF THE KEY
This contribution should prove useful for live or preserved animals
in development stages 25 through 40 (Gosner 1960). The key is not
arranged phylogenetically. General arrangement and terminology follow
Altig's.
Altig discussed some of the major sources of difficulty in tadpole
identification, beginning with the problem of poor preservation. Colora-
tion, useful in identifying live material, will fade in preserved specimens.
Labial teeth can fall out, and keratinized skin layers can be dislodged
from their underlying structures.
Larval anurans are highly susceptible to environmental influences
on morphology. For example, food type can affect mouthpart mor-
phology, causing some distortions of the normal appearance, as is evi-
dent in a comparison of laboratory-reared and field-collected tadpoles.
Ambient temperatures will affect development rates, and may influence
allometric growth patterns (see below). The tails of many tadpoles are
damaged by predators, particularly nymphal dragonflies (Caldwell et al.
1980). This may affect comparisons based on length ratios, either
because of the tail's abbreviated length or because of a change in its
overall shape as regeneration occurs. In addition, a regenerated tail
often has a blackened tip or large blotches or large blotches on the fin,
marks that may not be part of the normal pattern. Color of a live
animal can vary with background. Rana clamitans larvae developing in
a pond that contains a high level of gray clay in suspension (as in some
ponds in the Sandhills region) will be very pale, while larvae in other
situations may range from green to dark brown.
Many characters vary ontogenetically. The most obvious of these is
color. The clear belly of young Rana clamitans larvae will become an
opaque, cream color as the animals develop. The number of rows of
labial teeth and the length of the rows change with tadpole age and size.
The most subtle ontogenetic variations are the allometric shape changes
exhibited by some species. The notable flagellum of a Hyla femoralis
tadpole is not present in a young larva, but becomes increasingly well
developed as the tadpole grows. Many species with broad fins, like Hyla
gratiosa and H. chrysoscelis, have more streamlined profiles as small
larvae. All these sources of phenotypic variation should be kept in mind
when using any key to tadpoles.
North Carolina Tadpole Key 121
KEY TO THE TADPOLES OF NORTH CAROLINA
1. Jaws without keratinized sheaths; oral disc and labial teeth absent
(Microhylidae) Gastrophryne carolinensis
Jaws with keratinized sheaths; oral disc and labial teeth present
(Fig. 2) 2
2. Anus medial (Fig. 4A) 3
Anus dextral (Fig. 4B) 4
3. Oral disc emarginate (Fig. 2); tooth row formula 1-2/3; spiracle distinctly on left
side of body (Fig. 3A) (Bufonidae) 5
Oral disc not emarginate (Fig. 2); tooth row formula 2/4 or more; spiracle
ventrolateral (Pelobatidae) Scaphiopus holbrooki
4. Oral disc emarginate (Ranidae) 33
Oral disc not emarginate (Hylidae) 8
5. P-2 (see Fig. 2) with distinct median gap; P-3 less than 0.50 P-l; papillary
border extends to lateral tips of P-2;
light color in life Bufo quercicus
P-l with no median gap; P-3 greater than 0.50 P-l; papillary border extends
distinctly around P-2; dark color in life 6
6. Substantial submarginal papillae (Fig. 2), particularly around emarginate areas
of oral disc; dorsal tail fin height equal to musculature height (Fig. 1); dorsal fin
may be higher than ventral fin Bufo terrestris
Few if any submarginal papillae; dorsal tail fin height lower than musculature
height; fins subequal in height 7
7. Dorsum unicolored; snout sloping in lateral view; tail musculature distinctly
bicolored; tail fin height/ musculature height 2.0 or
less Bufo americanus
Dorsum often slightly mottled in life; snout rounded in lateral view; tail
musculature often not distinctly bicolored; tail fin height/ musculature height
greater than 2.0 Bufo woodhousei fowleri
8. Two rows of posterior labial teeth (Fig. 2) 9
Three or more rows of posterial labial teeth 13
9. A-2 gap (Fig. 2) wide; spiracular tube at least partly free from body wall; body
slightly depressed; tail tip often solid black (Acris) 10
A-2 gap narrow to moderate; spiracular tube fully attached to body wall; body
globular; tail tip, if black, with mottle or blotched black edges 11
10. Free section of spiracular tube long, almost entire length of tube; throat dark;
tail musculature finely flecked; Coastal Plain Acris gryllus
Free section of spiracular tube short, less than or equal to half the length of
tube; throat light; tail musculature mottled or reticulated; Piedmont and moun-
tain valleys Acris crepitans
11. Tail musculature distinctly striped in lateral view; light stripe extends from
dorsal tail musculature stripe to eye; throat and chest may be mottled; dorsum
of tail musculature often banded or with
saddles Limnaoedus ocularis (part)
Tail musculature not or only faintly striped, but without extension to eye;
throat and chest light; dorsum of tail musculature not banded 12
VENTRAL FIN
SPIRACLE
Fig. 1. Left lateral aspect of a tadpole. TL = total length; BL - body length;
TaL = tail length; MH = musculature height; TH = tail height. Redrafted from
Altig (1970).
1— A-2 GAP
, ANTERIOR
LABIUM
POSTERIOR
LABIUM
Fig. 2. Tadpole mouth parts, schematic. UJ = upper jaw (mandible); LJ = lower
jaw (mandible); A-1,2 = first and second anterior tooth rows; P-1,2,3 = first,
second and third posterior tooth rows; SP = submarginal papilla; MP = mar-
ginal papilla; OD = oral disc (shown emarginate on viewer's left, marginate on
right). Redrafted from Altig (1970).
12. Tail musculature unicolored or bicolored; fins clear or with stellate melano-
phores; small black dots often present on body; A-2 subequal to A-l; one row of
marginal papillae (see Fig. 2); snout round when viewed
dorsally Pseudacris triseriata (part)
Tail musculature mottled; fins clear or with large blotches; no dots on body; A-2
longer than A-l; two rows of marginal papillae; snout square when viewed
dorsally Hyla crucifer (part)
North Carolina Tadpole Key 123
13. Posterior gap in papillary border 14
No posterior gap in papillary border 16
14. Tail musculature distinctly striped and stripe may extend forward to eye; snout
rounded or pointed in dorsal view 15
Tail musculature mottled or indistinctly striped, but in no case does a light
stripe extend forward to eye; snout square in dorsal
view Hyla crucifer (part)
15. Tail musculature stripe extends to eye; snout round when viewed dorsally;
posterior gap in papillary border greater than or equal to length of P-3;
interocular distance wide, only slightly less than maximal head
width Limnaoedus ocularis (part)
No extension of tail musculature stripe to eye; snout tapered or slightly pointed
in dorsal view; posterior gap in papillary border less than length of P-3;
interocular distance narrow, substantially less than maximal head
width Hyla andersoni (part)
16. P-3 length 0.50 or more times length of P-2; P-3 longer than upper jaw 17
P-3 length very short, less than 0.50 times length of P-2; P-3 subequal to upper
jaw 20
17. Submarginal papillae absent or few; dorsum of tail musculature usually with
one black saddle slightly anterior to midlength Hyla gratiosa (part)
Substantial submarginal papillae; no black saddle on dorsum of tail
musculature , 18
18. Tail musculature distinctly striped; well developed flagellum at tip of tail;
reddish color in life Hyla femoralis
Tail musculature not striped; flagellum absent; golden to brown or bluish color
in life 19
19. Dorsal fin height equal to or greater than musculature height; throat seldom
pigmented; dorsal fin never extends anterior to midway between spiracle and
eye Hyla chrysoscelis, Hyla versicolor
Dorsal fin height less than musculature height; throat pigmented in life; dorsal
fin extends to posterior border of the
eye Hyla squirella
20. Tail musculature striped 21
Tail musculature not striped 26
2 1 . A-2 gap wide Pseudacris brimleyi
A-2 gap narrow 22
22. Light dorsal stripe on tail extends to eye; fins clear or with a few stellate
melanophores; dorsum of tail musculature banded or marked with
saddles Limnaoedus ocularis (part)
Dorsal stripe does not extend to eye; fins clear or mottled; dorsum of tail
musculature not banded 23
23. Tail stripe distinct; snout rounded when viewed dorsally; body slightly de-
pressed 24
Tail stripe may or may not be distinct; snout squarish or tapering in dorsal view;
body not depressed 25
24. Dorsal fin originates anterior to spiracle Pseudacris ornata (part)
Dorsal fin originates posterior to spiracle Pseudacris triseriata (part)
124
Joseph Travis
Fig. 3. Eye positions, dorsal aspect. A. Lateral eyes (and spiracle). B. Dorsal
eyes. Redrafted from Altig (1970).
ANUS
Fig. 4. Anus positions, ventral aspect. A. medial. B. dextral. Redrafted from
Altig (1970).
25. Tail stripe faint; snout square when viewed dorsally; snout-spiracle distance/
body length greater than 0.60; spiracle just below eye level; dorsal fin higher
than ventral fin; interocular distance only slightly less than maximum head
width Hyla crucifer (part)
Tail stripe distinct; snout tapering in dorsal view; snout-spiracle distance/ body
length less than 0.60; spiracle well below eye level; fins equal in height;
interocular distance markedly less than maximum head
width Hyla andersoni (part)
26. Total length (Fig. 1) greater than 45 27
Total length less than 45 mm 28
North Carolina Tadpole Key 125
27. Jaws wide and rounded; tail musculature unicolored Hyla gratiosa (part)
Jaws narrow and angular; tail musculature bicolored
Pseudacris ornata (part)
28. Dorsal fin originates anterior to spiracle 29
Dorsal fin originates at or posterior to spiracle 31
29. Fins clear or with few stellate melanophores; fins rounded; dorsal fin higher
than ventral fin 30
Fins and tail musculature mottled or reticulated; fins tapering toward the tail
tip; dorsal and ventral fins equal in height Hyla cinerea
30. Jaws wide and rounded; dorsum of tail musculature usually with black saddle
slightly anterior to midlength; tail musculature
unicolored Hyla gratiosa (part)
Jaws narrow and angled; no black saddle on dorsum of tail musculature; tail
musculature bicolored Pseudacris ornata (part)
31. Fins and tail musculature mottled or reticulated; body somewhat globular;
snout square when viewed dorsally Hyla crucifer (part)
Fins and tail musculature clear or with a few stellate melanophores; body
somewhat depressed; snout round when viewed dorsally 32
32. Body dark brassy in life; dorsal fin terminates far posterior to
spiracle Pseudacris brachyphona
Body color not dark brassy; dorsal fin terminates at or slightly posterior to
spiracle 33
33. Dorsal fin higher than ventral fin and equal to tail musculature
height Pseudacris triseriata (part)
Fins subequal and both lower than musculature
height Pseudacris nigrita
34. Four or more rows of teeth on anterior or posterior
labium Rana sylvatica (part)
Less than four rows of teeth on both anterior and posterior labium 35
35. A-2 gap ratio greater than 1.5; dorsal fin originates at or only slightly posterior
to spiracle Rana sylvatica (part)
A-2 gap ratio variable; dorsal fin originates far posterior to spiracle 36
36. Lower jaw wide; nostrils medium to large; skin thin, gut visible, with weakly
pigmented belly in larger specimens; small animals uniform in color, even when
preserved Rana pipiens group: 37
Lower jaw narrow; nostrils small; skin thick; gut usually not visible, with
strongly pigmented belly in larger animals; small animals with gold transverse
bands on anterior part of body, appearing unevenly pigmented when
preserved Rana catesbeiana group: 39
37. A-2 gap ratio 2 or more; P-l/P-3 length ratio 1.3 or greater. . . Rana palustris
A-2 gap ratio less than 2; P-l / P-3 length ratio less than 1.5 38
38. No keratinized areas at medial tips of P-l; A-2 gap ratio often greater than 1.0;
color variable Rana sphenocephala
Keratinized areas present at medial tips of P-l; A-2 gap ratio always less than
1 .0; color usually dark Rana areolata
39. Tail musculature unicolored or mottled, but not striped; fins clear or mottled,
but not in any particular pattern 41
Tail musculature distinctly bicolored or striped; fins either striped (or with a
row of dots) or marked around edges 40
126 Joseph Travis
40. Tail musculature distinctly bicolored; fins without stripe; larger specimens have
prominent black edging around a clear or speckled fin Rana heckscheri
Tail musculature distinctly striped; stripe or row of dots (formed by pigment
around the lateral line pores) present on dorsal fin; no black edging on tail
fins Rana virgatipes
41. A-2 gap ratio greater than 0.50; body and tail patterned with distinct black dots;
belly light green, white, or yellow in life Rana catesbeiana
A-2 gap ratio less than 0.50; body and tail lacking distinct black dots; belly of
larger individuals is cream or white in life Rana clamitans
NOTES ON FIELD IDENTIFICATION
Many animals can be diagnosed to genus or species in the field with
the use of a hand lens. Small tadpoles are always difficult to identify,
but the following notes should allow larger individuals to be placed into
one of five principal groups.
Hylidae: body square in dorsal view, eyes lateral; nostrils small com-
pared to eyes; dextral anus; oral disc not emarginate;
never black in color, but can range from bluish to brown.
Rana: body oval or round in dorsal view, eyes dorsal or dorsolateral;
nostrils small compared to eyes; dextral anus; oral disc
emarginate; color diagnostically unreliable.
Bufo: body round or oval in dorsal view, eyes dorsal and with a "cross-
eyed" aspect; nostrils large, and head appears to have a
"snout"; median anus; oral disc emarginate; color may be
dark or light (Bufo quercicus).
Scaphiopus holbrooki: body round or oval in dorsal view, eyes close-
set and dorsal; head wide relative to body width; entire
body moves from side to side while swimming; median
anus; oral disc not emarginate; color black.
Gastrophryne carolinensis: body round in dorsal view, distinctly de-
pressed; eyes wide-set and lateral; median anus; no oral
disc; color dark, although larger individuals have mottled
venters and a stripe on the tail musculature.
ACKNOWLEDGMENTS.— Many people helped collect tadpoles,
but I owe a particular debt of gratitude to Mr. Jack Longino, Mr. Peter
Morin, and Dr. Henry Wilbur. Professor Joseph Bailey allowed me to
use the Duke Vertebrate Collection, and Alvin Braswell, North Carolina
State Museum of Natural History, graciously provided specimens of
preserved tadpoles. Professor Henry Wilbur generously provided me
with the time, funding, and encouragement to construct this key and to
refine it from 1977 to 1980 while I was at Duke University. Drs. Ronald
North Carolina Tadpole Key 127
Altig and John Cooper, and an anonymous reviewer, improved the
manuscript considerably through their conscientious suggestions, and
Dr. Cooper drafted the figures.
The work that gave rise to this key was supported by NSF grants
DEB 76-82620 and DEB 79-11539, both to H. M. Wilbur, and by a
postdoctoral fellowship to the author from the University of Virginia
through the Mountain Lake Biological Station. Final revisions were
made while I was supported by NSF grant DEB 81-82620.
LITERATURE CITED
Altig, Ronald. 1970. A key to the tadpoles of the continental United States and
Canada. Herpetologica 26: 180-207.
Caldwell, Janalee P., J. H. Thorp and T. O. Jervey. 1980. Predator-prey rela-
tionships among larval dragonflies, salamanders, and frogs. Oecologia
46:285-289.
Gosner, Kenneth L. 1960. A simplified table for staging anuran embryos and
larvae with notes on identification. Herpetologica 76:183-190.
Martof, Bernard S., W. M. Palmer, J. R. Bailey and J. R. Harrison. 1980.
Amphibians and Reptiles of the Carolinas and Virginia. Univ. North Caro-
lina Press, Chapel Hill. 264 p.
Orton, Grace L. 1952. Key to the genera of tadpoles in the United States and
Canada. Am. Midi. Nat. ¥7:382-395.
Smith, Hobart M. 1934. The amphibians of Kansas. Am. Midi. Nat. 75:377-528.
Wright, Albert H., and A. A. Wright. 1949. Handbook of Frogs and Toads of
the United States and Canada. Comstock Publ. Co., Ithaca, New York. 640 p.
Accepted 16 March 1981
New Distributional Records
of Eastern Kentucky Fishes
Melvin L. Warren, Jr.
Kentucky Nature Preserves Commission,
Frankfort, Kentucky 40601
ABSTRACT. — A two year survey of the major river drainages of
eastern Kentucky resulted in new records and range extensions of sev-
eral rare or poorly known fish species. Ichthyomyzon fossor is reported
for the first time in the Little Sandy River and South Fork of the
Kentucky River, and Notropis galacturus from the Big Sandy, Laurel
and upper Cumberland (above the falls) rivers. The range of Phoxinus
cumber landensis is extended to include Laurel and Rockcastle rivers.
Etheostoma tippecanoe is noted for the first time in the Cumberland
River within Kentucky, and the most upstream record of Percina
phoxocephala in the Ohio River valley of the state is reported. Addi-
tional localities and range extensions are noted for eight rare or poorly
known Kentucky fishes. Notes on distribution, habitat, and associates
are included.
INTRODUCTION
During 1978 and 1979, the Kentucky Nature Preserves Commission
conducted aquatic biota surveys of selected streams in four major river
drainages of eastern Kentucky (Harker et al. 1979, 1980). Considering
past and projected growth of the coal mining industry in this region and
the concomitant impacts on stream ichthyofauna, it is apparent that
documentation of the existing fauna is both necessary and timely. Much
of the distributional information concerning this region is from scat-
tered collections without proper documentation (e.g., locality data,
voucher material). The stream fishes of the Big Sandy, Licking, and
upper Cumberland rivers are particularly poorly known in terms of sub-
stantiated collections (Burr 1980). Collecting efforts of the Kentucky
Nature Preserves Commission resulted in distributional information for
several poorly known or rare species in the state and suggest that our
knowledge of the eastern Kentucky ichthyofauna is far from complete.
Several of the records reported are included in both Burr (1980)
and Lee et al. (1980), although not in the detail covered here. The fol-
lowing accounts of species are presented for the purpose of enriching
and elucidating knowledge of the varied Kentucky ichthyofauna.
SPECIES ACCOUNTS
The following accounts extend the range of several fish species
within and across drainages of eastern Kentucky. The majority of
reported specimens were collected during field work conducted by the
Kentucky Nature Preserves Commission, although several records were
taken from collections examined by the author at the Kentucky
Brimleyana No. 6: 129-140. December 1981. 129
130 Melvin L. Warren, Jr.
Department of Fish and Wildlife Resources (KFW). A detailed account
of methodology and a discussion of most collecting stations, including
macrobenthic, periphyton, substrate, and water quality analyses, was
presented in Harker et al. (1979, 1980). The bulk of the collections are
housed at the Kentucky Nature Preserves Commission (KNP). Other
material as noted is deposited at Auburn University (AU), Eastern Ken-
tucky University (EKU), Tulane University (TU), University of New
Orleans (UNO), University of Tennessee (UT), and in the collection of
Wayne C. Starnes (WCS).
Species accounts include the catalog numbers, followed in paren-
theses by the number of specimens, the stream and major drainage, the
locality, county, and date of collection. All scientific and common
names follow Robins et al. (1980), except in the case of undescribed
taxa.
Ichthyomyzon fossor Reighard and Cummins. Northern brook
lamprey. KNP SOICAR (2), UT 2.81 (2), WCS 1010-01 (2), Big Sinking
Cr. (Little Sandy R. dr.), 1.7 km above mouth, Carter Co., 31 May
1978; KNP K01CLA (1), Goose Cr. (S. Fk. Kentucky R. dr.), at Lipps,
Clay Co., 9 May 1978.
Bauer and Branson (1979) recently reported this nonparasitic lam-
prey from the Middle Fork of the Kentucky River. Previously it was
reported from the upper Big Sandy and Barren rivers (Clay 1975), but
Burr (1980) did not recognize the latter record. The present collections
constitute a new record for the Little Sandy River and an extension of
the known distribution in the Kentucky River to the South Fork. All
specimens were adults and were taken from areas of swift current over
substrates of rubble interspersed with sand and gravel. Both the Ozark-
ian and Ohio Valley populations of /. fossor are relatively isolated
from the widely distributed orthern populations (Pflieger 1971; Rohde
and Lanteigne-Courchene 1980). The Kentucky distribution of /. fossor
strongly suggests that former tributaries of the ancient Teays River (e.g.,
Big Sandy, Little Sandy and Kentucky rivers) (Hocutt et al. 1978)
served as refugia and redispersal points during and after Pleistocene
glaciation. Although its rarity in Kentucky is partly attributable to the
difficulty of collecting adults in preferred habitats (Bauer and Branson
1979), the Kentucky Academy of Science considers the species to be
threatened (Branson et al. 1981).
Notropis galacturus (Cope). Whitetail shiner. UT 44. 1757(3), Russell
Fk. (Big Sandy R. dr.), below Chesapeake and Ohio Railroad bridge at
KY 80, Pike Co., 23 May 1978; KFW 1526 (38), Russell Fk. (Big Sandy R.
dr.), at mouth of Grassy Br., Pike Co., 30 August 1961 ; KNP BOl PIK (70),
Elkhorn Cr. (Russell Fk. Big Sandy R. dr.), 3.3 km W of jet KY 80 and KY
197, Pike Co., 1 1 October 1978; KFW 1521 (7), same locality, 29 August
1961; KFW 1643 (37), Clover Fk. (Cumberland R. dr.), KY 38 bridge,
Harlan Co., 27 September 196 1 ; KFW 1 508 (4), Laurel R. (Cumberland R.
New Records Kentucky Fishes 131
dr.), at mouth of Spruce Cr., Whitley Co., 23 August 1961 ; KNP C07LAU
(13), Laurel R. (Cumberland R. dr.), 3.1 km above the mouth of Adams
Br., Laurel Co., 1 1 October 1979.
In Kentucky this distinctive cyprinid is relatively common in clear
streams in the Cumberland River below Cumberland Falls. Although
noted by Clay (1975) from the Big Sandy River and upper Cumberland
River (above the falls), locality data were not given. Gilbert and Burgess
(1980a) did not include the Big Sandy, upper Cumberland, or Laurel rivers
in their depiction of the Kentucky range. The collections noted here are
apparently the only formally published localities of the whitetail shiner in
these drainages. It is also known from the upper Tennessee and New rivers
of Virginia (Gibbs 1961), which closely abut the headwaters of the Big
Sandy and Cumberland rivers. The presence of the species in the four
adjacent drainages suggests stream capture as the means of dispersal, and
Gibbs (1961) interpreted the presence of N. galacturus in the New River as
the result of piracies between the New and upper Tennessee rivers.
Subsequent workers regarded the New River populations as probably
native (Jenkins et al. 1971) or introduced (Gilbert and Burgess 1980a).
Limited distribution such as that observed in the New and Big Sandy rivers
may be the result of first entry during recent times, reentry after
extirpation, or introduction rather than natural factors (Jenkins et al.
1971). In light of the apparent absence of other species indicative of stream
capture with adjacent drainages, populations of N. galacturus in the Big
Sandy River are most likely the result of introduction.
The only historical reference to N. galacturus in the Cumberland
River above the falls was by Evermann (1918), who apparently erroneously
cited Woolman's (1892) Rockcastle River locality. The rarity of N.
galacturus in surveys above the falls may be in part related to the extensive
habitat destruction associated with coal mining. Its apparent rarity in the
Laurel River is attributable to habitat destruction and the lack of
systematic surveys before the impoundment of Laurel River Reservoir.
Interpretation of the native or non-native status above Cumberland Falls
invokes reasoning similar to the interpretation of the populations in the
New and Big Sandy rivers, although there is strong faunal evidence of
lateral stream transfer between the Cumberland and upper Tennessee
(Clinch-Powell) rivers (Starnes et al. 1977). Additional collections,
comparison, and analyses of populations in the Big Sandy, Cumber-
land, New, and Tennessee rivers may further enlighten the enigmatic
dispersal history and distribution of N. galacturus.
Notropis sp. Undescribed. Sawfin shiner. AU 18680 (4), KNP
COIMCY (13), EKU uncat. (4), Rock Cr. (Big S. Fk. Cumberland R.
dr.), 6.7 km SW of Bell Farm at Great Meadows Camp Site, McCreary
Co., 19 September 1978; AU uncat. (1), KNP uncat. (4), Pitman Cr.
(Cumberland R. dr.), Co. Rd. 1247 bridge in Somerset, Pulaski Co., 25
October 1979.
132 Melvin L. Warren, Jr.
This undescribed relative of the mirror shiner, Notropis spectruncu-
lus, was previously known in Kentucky from a single record in the Little
South Fork of the Cumberland River (Comiskey and Etnier 1972; John
S. Ramsey, pers. comm.). The two additional localities noted above
indicate a wider but sporadic distribution in the Big South Fork and
middle Cumberland rivers of eastern Kentucky. The rarity of the sawfin
shiner in Kentucky may be related in part to lack of recognition by early
workers and to the embayment of tributaries by Cumberland River
Reservoir. Additional Kentucky collections are anticipated in other
medium-to-large streams of the drainage. The species is considered
threatened in Kentucky by the Kentucky Academy of Science (Branson
etal. 1981).
Phoxinus cumberlandensis Starnes and Starnes. Blackside dace.
WCS 883-01 (1), Marsh Cr. (Cumberland R. dr.), 1.8 km S of Co. Rd.
1470 on Marsh Cr. Rd., McCreary Co., 4 May 1978; KNP C02MCY
(3), same locality, 19 September 1978; WCS 1163-01 (1), Craig Cr.
(Laurel R. dr.), at KY 312 bridge, Laurel Co., 9 October 1979; KNP
C02LAU (2), Ned Branch (Rockcastle R. dr.), 0.6 km N of terminus of
Co. Rd. 1193 and 50 m above the impounded mouth, Laurel Co., 25
July 1979; KNP COILET (1), Colliers Branch (Poor Fk. Cumberland
R. dr.), 4.2 km E of jet US 119 and Colliers Br. Rd., Letcher Co., 22
April 1980; KNP uncat. (2), Poor Fk. (Cumberland R. dr.), 5.5 km
ENE of jet US 119 and KY 932, Letcher Co., 1 June 1979; KNP
C02WHI (15), Bunches Cr. (Cumberland R. dr.), 1.5 km above the
mouth, Whitley Co., 22 August 1979.
Previously, P. cumberlandensis was known in Kentucky from 12
extant and 2 apparently extirpated populations (Starnes and Starnes
1978). The addition of the six localities reported above indicates the
species occurs from the extreme headwaters of the Poor Fork of the
Cumberland River downstream to and including the Laurel River,
lower Rockcastle River, and Beaver Creek systems. The species was col-
lected in pool areas of headwater streams in association with undercut
banks and/ or rubble, boulder, and sand substrates. Most seine hauls
yielded only one or two individuals. The general habitat description
presented by Starnes and Starnes (1978) agrees well with my obser-
ations.
Phoxinus cumberlandensis apparently evolved in isolation in the
Cumberland River drainage above Cumberland Falls, which represents
the major part of the known range (Starnes and Starnes 1978). The
discovery of populations in the Laurel and lower Rockcastle rivers
below Cumberland Falls represents the first records for these drainages
and raises questions concerning the circumvention of the falls. In order
to explain the presence of P. cumberlandensis immediately below Cum-
berland Falls, Starnes and Starnes (1978) postulated lateral stream cap-
ture or the relatively rapid regression of the falls in recent geologic time.
New Records Kentucky Fishes 133
McGrain (1966) placed the downstream origin of the falls near the pres-
ent location of Burnside, Kentucky, which is well downstream of the
present mouths of both the Laurel and Rockcastle rivers. The presence
of P. cumberlandensis in these river systems suggests the relatively rapid
regression of Cumberland Falls as the most likely explanation for the
present distribution. Further faunal evidence is implied by the Cumber-
land River distribution of Etheostoma kennicotti as presented by Page
and Smith (1976). Unfortunately, the dispersal and distribution of P.
cumberlandensis is obscured and fragmented by man's activities in the
area (i.e., mining, impoundments, etc.). In addition, there is apparently
no geological record of the regression of Cumberland Falls (McGrain
1966). The blackside dace is listed as threatened by the Kentucky
Academy of Science (Branson et al. 1981).
Percopsis omiscomaycus (Walbaum). Trout-perch. KNP BO 1 LAW
(3), Little Blaine Cr. (Big Sandy R. dr.), 3.5 km NW of the jet KY 32 and
Little Blaine Cr. Rd., Lawrence Co., 3 October 1978; UT 79.4 (4), KNP
SOI CAR (2), Big Sinking Cr. (Little Sandy R. dr.), 1.7 km above mouth,
Carter Co., 13 September 1978.
Although primarily a northern species, the trout-perch is widely but
discontinuously distributed in Kentucky, with most records from the
extreme northeastern section of the state (Clay 1975; Burr 1980). The Little
Blaine Creek collections were believed to represent the most upstream
locality in the Big Sandy River; however, material recently examined from
Right Fork Beaver Creek (Levisa Fork Big Sandy R. dr.) indicates a much
wider distribution in the Big Sandy than was previously known. The
specimens are housed at the Kentucky Department of Transportation,
Frankfort, Kentucky (Steve P. Rice, pers. comm.). Additional records in
the middle and upper reaches of the Big Sandy River may be expected.
Percopsis omiscomaycus is listed as of special concern in Kentucky by the
Kentucky Academy of Science (Branson et al. 1981).
Ammocrypta pellucida (Putnam). Eastern sand darter. KNP uncat.
(1), N. Fk. Red R. (Kentucky R. dr.), below KY 715 bridge at Menifee-
Wolfe county line, 17 June 1978; KFW 1745 (4), Greasy Cr. (Middle Fk.
Ky. R. dr.), no locality, Leslie Co., 15 August 1962; KFW 1 160 (1), Middle
Fk. (Kentucky R. dr.), no locality, Leslie Co., 15 June 1960.
Ammocrypta pellucida is known from few localities in the Kentucky
River drainage (Williams 1975; Hocutt 1980). The species was previously
reported from localities in the lower reaches of the Red River (Branson and
Batch 1974) and in the North and South Forks of the Kentucky River
(Williams 1975). The above collection extends the known range in the Red
River approximately 33 km upstream and indicates a broader distribution
in this system than was previously reported. The specimens from Middle
Fork of the Kentucky River constitute a new record for this drainage and
indicate that A. pellucida occurred throughout the upper Kentucky River.
The Red River specimen was taken at the margin of a deep (1.0 m), gently
134 Melvin L. Warren, Jr.
flowing pool underlain by clean sand. Repeated efforts to secure addi-
tional specimens were unsuccessful. The specimen was taken with Etheos-
toma nigrum, another psammophilic species. Burr (1980) observed that
the once relatively common eastern sand darter is rapidly declining in
numbers in Kentucky. The Kentucky Academy of Science lists the species
as threatened (Branson et al. 1981).
Etheostoma cinereum Storer. Ashy darter. KNP C05ROC (2),
Rockcastle R. (Cumberland R. dr.), at mouth of Eagle Cr., Rockcastle
Co., 23 October 1979; KNP C09MCY (1), Big S. Fk. (Cumberland R. dr.),
3.0 km N of Tennessee state line at mouth of Troublesome Cr., McCreary
Co., 24 October 1979.
The ashy darter is confined to the Cumberland River in Kentucky and
is known from six substantiated collections, including those shown above
(Burr 1980 and pers. comm.). Although Saylor (1980) and Branson (1977)
noted the species in the Rockcastle River, no exact localities were given.
The above collection is considered the first formal report of the species
from this river. Subsequent collections in upstream segments of the Rock-
castle River have yielded a number of additional specimens (Brooks M. Burr,
pers. comm.). At both sites, E. cinereum was collected in sluggish current
adjacent to swift shoals over rubble-gravel substrate mixed with detritus
and/ or dead Justicia americana in areas 0.45-0.75 m deep. No fishes were
associated with E. cinereum on the substrate; however, Notropis ariom-
mus, N. chrysocephalus, and N. rubellus occurred in the water column
directly above.
Etheostoma nigrum susanae (Jordan and Swain). Johnny darter.
KNP C05MCY (2), Bridge Fk. Laurel Cr. (Cumberland R. dr.), directly
above mouth on KY 478, McCreary Co., 12 September 1979; KNP
C02WHI (5), Bunches Cr. (Cumberland R. dr.), 1.5 km above mouth,
Whitley Co., 22 August 1979.
In a recent taxonomic evaluation of this rare subspecies, Starnes and
Starnes (1979) reported extant populations above Cumberland Falls
within Whitley and McCreary counties. The collections reported above
represent new localities from the same general area. Both collections were
made in small streams with well-forested watersheds and excellent water
quality. Individuals were generally collected over clean-swept sand and
bedrock at the base of gentle riffles or in shallow pools. Etheostoma
nigrum susanae is the only endemic fish above Cumberland Falls and is
currently listed by the Kentucky Academy of Science as threatened (Bran-
son et al. 1981).
Etheostoma tippecanoe Jordan and Evermann. Tippecanoe darter.
KNP uncat. (1), Big S. Fk. (Cumberland R. dr.), 3.2 km N of Tennessee
state line at mouth of Oilwell Br., McCreary Co., 24 October 1979.
In Kentucky, the Tippecanoe darter was formerly known from local-
ized populations in the Licking, Green, and Kentucky rivers (Burr 1980;
Clay 1975; Hocutt 1980). Although known from the Big South Fork of the
New Records Kentucky Fishes 135
Cumberland River in Tennessee (Comiskey and Etnier 1972), this record,
included in Burr (1980), is the first reported occurrence in the Cumberland
River of Kentucky. One adult female was collected in a large, swift,
rubble-gravel shoal approximately 0.3-0.5 m deep. Other members of the
subgenus Nothonotus associated with E. tippecanoe were E. maculatum
sanguifluwn and E. camurum. The collection of one individual precludes
evaluation of the status of the species in this segment of the river; however,
E. tippecanoe is listed by the Kentucky Academy of Science as endangered
(Branson et al. 1981).
Percina copelandi (Jordan). Channel darter. TU 120014 (3), Buck-
horn Cr. (N. Fk. Kentucky R. dr.), 0.7 km NE of KY 476, Breathitt Co., 19
June 1978; KNP C09MCY (4), Big S. Fk. (Cumberland R. dr.), 3.0 km N
of Tennessee state line at mouth of Troublesome Cr., McCreary Co., 29
August 1979; UT 91. 1790 (22), WCS 1009-02 (5), KNP uncat. (6), Russell
Fk. (Big Sandy R. dr.), below Chesapeake and Ohio Railroad bridge at KY
80, Pike Co., 23 May 1978.
Burr (1980) regarded the channel darter as uncommon in Kentucky.
The presence of the species in Buckhorn Creek represents the first formal
record from the North Fork of the Kentucky River. Its occurrence in this
relatively small system is surprising in light of its reported preference for
big river habitats (Gilbert and Burgess 1980b). The population in the Big
South Fork of the Cumberland River apparently represents the second
reported locality from this system in Kentucky, although others have
noted it from the same drainage in Tennessee (Comiskey and Etnier 1972;
Page 1974; Gilbert and Burgess 1980b). An examination of the University
of Louisville and KFW museum records revealed that all the collections
reported by Clay (1975) from the Russell and Levisa Forks of the Big
Sandy River pre-date 1960. The channel darter is apparently persisting in
good numbers in Russell Fork as indicated by the present collections.
Habitat and species associates are presented under the Percina oxyrhyn-
cha account. The channel darter is considered of special concern in Ken-
tucky by the Kentucky Academy of Science (Branson et al. 1981).
Percina oxyrhyncha (Hubbs and Raney). Sharpnose darter. KNP
B0 1 JOH (5), Levisa Fk. (Big Sandy R. dr.), 1 km N of River, Johnson Co.,
2 October 1 978; WCS 1009-03 (4), UT 9 1 . 1 789 (7), Russell Fk. (Big Sandy
R. dr.), below Chesapeake and Ohio Railroad bridge at KY 80, Pike Co.,
23 May 1978; KFW 1533 (49), Russell Fk. (Big Sandy R. dr.), mouth of
Grassy Br. at Kentucky-Virginia line, Pike Co., 30 August 1961; KFW
1225 (5), Levisa Fk. (Big Sandy R. dr.), mouth of Morgans Cr., Pike Co.,
27 September 1960; KFW 1803 (34), N. Fk. (Kentucky R. dr.), Rocklick,
Breathitt Co., 19 September 1972.
The darter subgenus Swainia is represented in Kentucky by three
morphologically similar species, P. oxyrhyncha, P. squamata, and P.
phoxocephala (see following accounts). Because of morphological similari-
ties, much confusion has resulted concerning assignment to species within
136 Melvin L. Warren, Jr.
the subgenus. Percina oxyrhyncha was unknown in Kentucky until
Denoncourt et al. (1977) reported specimens from the upper Kentucky
River, and others later noted the species from the upper Big Sandy,
Licking, and Green rivers (Thompson 1978; Bauer and Branson 1979; Burr
1980; Thompson 1980a). Thompson (1980a) depicted the range in the Big
Sandy River as the extreme headwaters of Levisa and Russell Forks near
the Kentucky-Virginia line. The collections of my report indicate that the
species is common in Russell Fork, and that the range extends downstream
in Levisa Fork at least 80 km from the Kentucky-Virginia line.
The Levisa Fork specimens were adults and were taken in a swift,
boulder strewn shoal (0.8-1.0 m deep) directly adjacent to a dense bed of
Justicia americana. The only directly associated percid was Percina sclera.
In contrast, the series of juveniles from Russell Fork was collected from a
shallow (0.15 m), rubble-gravel shoreline area with moderate current.
Species associates included juvenile Percina ex ides, P. copelandi, P. sciera,
and P. caprodes. Denoncourt et al. ( 1 977) noted a correlation of specimen
size with both substrate and gradient; the above observations support their
findings. The rarity of P. oxyrhyncha in Kentucky is no doubt partly
attributable to taxonomic confusion and the difficulty of collecting adults
in the preferred big river habitats. The status of the species is currently
listed as undetermined by the Kentucky Academy of Science (Branson
etal. 1981).
Percina phoxocephala (Nelson). Slenderhead darter. UNO 3346 (1),
Tygarts Cr. (Ohio R. dr.), Bennetts Covered Bridge at jet K Y 7 and Co. Rd.
1215, Greenup Co., 31 May 1978.
Percina phoxocephala is most easily confused with P. oxyrhyncha. It
differs in having lower meristics, a more robust body, and less elongate
head and snout (Bruce A. Thompson, pers. comm). Hubbs and Raney
(1939) regarded the snout length as diagnostic in separating the two
species. The slenderhead darter is the most widely distributed member of
the subgenus Swalnia and has been previously reported in Kentucky from
the Green, lower Kentucky (Eagle Creek), Tennessee and Cumberland
rivers (Burr 1980; Thompson 1980b). The specimen from Tygarts Creek
represents the most upstream record in the Ohio River valley of Ken-
tucky. Its presence in this stream is not unexpected in light of the prox-
imity to populations in northern tributaries of the Ohio River (e.g., Sci-
oto River).
The single adult male was near breeding condition, supporting the
April to early June spawning period postulated by Thompson (1980b). The
specimen was taken in a moderately fast sand and gravel riffle that sup-
ported a dense growth of Justicia americana. Personal observations in this
and Green River collections suggest that adult P. phoxocephala occur
New Records Kentucky Fishes 137
most often over gravel and/ or finer substrates, whereas adult P. oxyr-
hyncha prefer coarser material such as rubble and boulders. Page and Smith
(1971) and Denoncourt et al. (1977) made similar observations on sub-
strate preferences of P. phoxocephala and P. oxyrhyncha, respectively.
Acquisition of additional material may change current views of the
complex distributional patterns of both P. phoxocephala and P. oxyr-
hyncha (Bruce A. Thompson, pers. comm.). Previous comments con-
cerning the status of the sharpnose darter in Kentucky are also applica-
ble to the slenderhead darter. The Kentucky Academy of Science
presently lists P. phoxocephala as of special concern in Kentucky (Bran-
son et al. 1981).
Percina squamata (Gilbert and Swain). Olive darter. KNP C04JAC
(2), Middle Fk. (Rockcastle R. dr.), 4.5 km W of jet KY 89 and Co. Rd.
2002, Jackson Co., 23 August 1979.
According to Burr (1980), P. squamata is known in Kentucky only
from the Rockcastle and Big South Fork Cumberland rivers. Although
previously reported from the Rockcastle River (Bauer and Branson
1979; Thompson 1980c), the above collections represent the most up-
stream occurrence. The specimens were secured from below a swift,
deep (0.8-1.0 m) riffle over a rubble and boulder substrate. Thompson
(1978) stated that until recently the olive darter was the poorest known
member of the subgenus Swainia, and one of the least known members
of the genus Percina. The olive darter is listed by the Kentucky
Academy of Science as endangered (Branson et al. 1981).
ACKNOWLEDGMENTS.— The author gratefully acknowledges the
assistance of B. A. Branson (Eastern Kentucky University), B. M. Burr
(Southern Illinois University), D. A. Etnier and W. C. Starnes (University
of Tennessee), R. A. Kuehne (University of Kentucky), J. S. Ramsey
(Auburn University), R. D. Suttkus (Tulane University), and B. A.
Thompson (Louisiana State University) for offering information and con-
firming original identifications of various species. L. E. Schaaf of the
Kentucky Department of Fish and Wildlife Resources generously allowed
examination of collections under his care. K. E. Camburn, R. R. Cicerello,
and M. A. Phillippi made helpful comments on the manuscript. This
publication was made possible as a result of studies conducted by the
Kentucky Nature Preserves Commission under the direction of D. F.
Harker, Jr.
138 Melvin L. Warren, Jr.
LITERATURE CITED
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Wigley. 1979. Aquatic biota and water quality survey of the Appalachian
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State Mus. Nat. Hist., Raleigh, x + 867 pp.
Hubbs, Carl L., and E. C. Raney. 1939. Hadropterus oxyrhynchus, a new percid
fish from Virginia and West Virginia. Occas. Pap. Mus. Zool. Univ. Mich.
595:1-9.
New Records Kentucky Fishes 139
Jenkins, Robert E., E. A. Lachner and F. J. Schwartz. 1971. Fishes of the central
Appalachian drainages: Their distribution and dispersal, pp. 43-1 17 in P. C.
Holt (ed.). The distributional history of the biota of the southern appalach-
ians, Part III: Vertebrates. Res. Div. Monogr. 4, Va. Polytech. Inst. State
Univ., Blacksburg. 306 pp.
Lee, David S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister and
J. R. Stauffer, Jr. 1980 et seq. Atlas of North American Freshwater Fishes.
N. C. State Mus. Nat. Hist., Raleigh, x + 867 pp.
McGrain, Preston 1966. Geology of the Cumberland Falls State Park area. Ky.
Geol. Surv. Series X, Spec. Publ. 1 1.
Page, Lawrence M. 1974. The subgenera of Percina (Percidae: Etheostomatini).
Copeia 1974(l):66-86.
, and P. W. Smith. 1971. The life history of the slenderhead darter, Percina
phoxocephala, in the Embarras River, Illinois. 111. Nat. Hist. Surv. Biol. Notes
74. 14 pp.
, and . 1976. Variation and systematics of the stripetail darter,
Etheostoma kennicotti. Copeia 1976(3):532-541.
Pfleiger, William L. 1971. A distributional study of Missouri fishes. Univ. Kan.
Publ. Mus. Nat. Hist. 20:225-570.
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. 4th ed. Am. Fish. Soc. Spec. Publ. No. 12. 174
pp.
Rohde, Fred C, and J. Lanteigne-Courchene. 1980. Ichthyomyzon fossor
Reighard and Cummins, Northern brook lamprey, p. 17 in D. S. Lee, et al.
Atlas of North American Freshwater Fishes. N.C. State Mus. Nat. Hist.,
Raleigh, x + 867 pp.
Starnes, Wayne C, and L. B. Starnes. 1978. A new cyprinid of the genus
Phoxinus endemic to the upper Cumberland River drainage. Copeia 1978(3)
:508-516.
, and 1979. Taxonomic status of the percid fish Etheostoma
nigrum susanae. Copeia 1979(3):426-430.
, D. A. Etnier, L. B. Starnes and N. H. Douglas. 1977. Zoogeographic
implications of the rediscovery of the percid genus Ammocrypta in the Ten-
nessee River drainage. Copeia 1977(4):783-786.
Thompson, Bruce A. 1978. An analysis of three subgenera (Hypohomus, Odon-
topholis and Swainia) of the genus Percina (Tribe Etheostomatini, Family
Percidae). Diss. Abstr. Int. B Sci. Eng. 38(9)A\ 14B-41 15B.
1980a. Percina oxyrhynca (Hubbs and Raney), Sharpnose darter, p. 733
in D. S. Lee, et al. Atlas of North American Freshwater Fishes. N.C. State
Mus. Nat. Hist., Raleigh, x + 867 pp.
1980b. Percina phoxocephala (Nelson), Slenderhead darter, p. 737 in D.
S. Lee, et al. Atlas of North American Freshwater Fishes. N.C. State Mus.
Nat. Hist., Raleigh, x + 867 pp.
1980c. Percina squamata (Gilbert and Swain), Olive darter, p. 742 in D.
S. Lee, et al. Atlas of North American Freshwater Fishes. N.C. State Mus.
Nat. Hist., Raleigh, x + 867 pp.
140 Melvin L. Warren, Jr.
Williams, James D. 1975. Systematics of the percid fishes of the subgenus Ammo-
crypta, genus Ammocrypta with descriptions of two new species. Bull. Ala-
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Accepted 14 September 1981
Systematic Status of the Cumberland Island
Pocket Gopher, Geomys cumberlandius
Joshua Laerm
Museum of Natural History and Department of Zoology,
University of Georgia, A then, Georgia 30602
ABSTRACT. — The Cumberland Island pocket gopher, Geomys cum-
berlandius, is known only from its type locality on Cumberland Island,
Camden County, Georgia. Statistical analyses of 21 morphometric
characters of G. cumberlandius and 5 mainland populations of G.
pinetis indicate that coastal populations of G pinetis are more similar
to G cumberlandius than they are to more inland populations of G
pinetis. These data, coupled with the Recent connection of Cumber-
land Island to the mainland, argue against taxonomic recognition of
G. cumberlandius, which is therefore regarded as a synonym of G
pinetis.
INTRODUCTION
Until the recent work of Williams and Genoways (1980), Geomys
cumberlandius Bangs was recognized as one of four nominal species of
pocket gophers occurring in Georgia (Hall and Kelson 1959; see also
Hall 1981). It has been considered monotypic and known only from its
type locality on Cumberland Island, Camden County, Georgia (Fig. 1).
Geomys colonus Bangs and Geomys fontanelus Sherman were also con-
sidered monotypic and known only from their type localities in Camden
and Chatham counties, Georgia, respectively. Geomys pinetis Rafinesque
was considered polytypic and widespread throughout Alabama, Florida
and Georgia.
Geomys cumberlandius was described in 1898. Specimens were
taken on Cumberland Island as late as 1956 but no subsequent speci-
mens were found and the species had been thought extinct. Recently,
however, a small population has been reported on the island (H. Neu-
hauser, pers. comm).
Williams and Genoways (1980) reviewed the systematics of south-
eastern pocket gophers and concluded, based on morphometries, that of
the four named species only G. pinetis is valid. They recognized only
two subspecies, G. p. pinetis and G. p. fontanelus, and synontmized G.
cumberlandius and G. colonus with G. p. pinetis.
This manuscript was in preparation when the Williams and Geno-
ways results were published. Because they employed only a portion
(77%) of available G. cumberlandius specimens, and did not include in
their study several characters upon which cumberlandius was originally
described, an independent corroboration of the systematic status of the
species is appropriate. The characters they omitted were: width of
nasals, breadth of ascending ramus of maxillary, and measurements of
Brimleyana No. 6: 141-151. December 1981. 141
142
Joshua Laerm
Fig. 1. Location of populations of Georgia pocket gophers examined. 1 = Cum-
berland Island; 2 = Scotchville, Camden Co.; 3 = Kingsland, Camden Co.; 4 =
Hursman's Lake, Screven Co.; 5 = Adam, Richmond Co.; 6= Augusta, Rich-
mond Co.
the auditory bullae. Another reason for corroborating the taxonomic
status of G. cumberlandius is the current political sensitivity of the
Cumberland Island population, which is a possible candidate for state
and/ or federal endandered status.
MATERIALS AND METHODS
All available specimens of G. cumberlandius (N = 73) were exam-
ined: Cumberland Island (8, AMNH; 5, CMNH; 1, CU; 13, DMNH; 26,
MCZ; 11, NMNH; 9, UGAMNH). These were compared to 157 speci-
mens of mainland G. pinetis from the following five populations in east
Georgia: Adam (41, MCZ), and Augusta (4, FSM; 11, NMNH), Rich-
mond County; Hursman's Lake (25, MCZ), Screven County; Kingsland
(6, FSM; 35, NMNH), and Scotchville (8, AMNH; 10, MCZ; 17,
NMNH), Camden County. Acronyms are defined in ACKNOWL-
EDGMENTS. The Scotchville population, previously recognized as G.
Status of Geomys cumberlandius 143
colonus has been shown by Laerm et al. (in press) and Williams and
Genoways (1980), on the basis of morphometries, electrophoresis,
karyology, and mitochondrial DNA sequence relatedness, to be syn-
onymous with G. pinetis. Twenty-one body and cranial measurements
were taken to the nearest 0.01 mm with dial calipers. These included: (1)
total body length, (2) tail length, (3) hind foot length, (4) condylobasilar
length, (5) zygomatic breadth, (6) mastoid breadth, (7) palatal length,
(8) palatal depth, (9) rostral breadth, (10) maxillary tooth row length,
(11) least interorbital constriction, (12) braincase breadth, (13) nasal
length, (14) greatest anterior nasal breadth, (15) breadth of nasals at
narrowest point, (16) greatest posterior nasal breadth, (17) interptery-
goid fossa length, (18) auditory bulla length, (19) breadth of ascending
ramus of maxillary, (20) anterior palatal breadth, and (21) posterior
palatal breadth. All measurements except variable 20 were made using
the methods of Williams and Genoways (1977) and DeBlase and Martin
(1974). Variable 20 was measured across the greatest width of the
ascending arm of the maxillary.
It has been well established that body and cranial measurements
change during the growth of an individual, but usually not at a constant
rate. It is, therefore, frequently desirable to compensate for size varia-
tion due to sex and age before comparisons are made. This is particu-
larly true in cases where small sample sizes limit the value of assigning
each individual to separate sex and age classes as in the present case.
This has commonly been done with proportions or transformation of
proportions; however, controversy has recently arisen over the use of
these techniques (Atchley et al. 1976; Albrecht 1978; Atchley and An-
derson 1978; Dodson 1978; Hill 1978). Fortunately, a number of statis-
tical techniques are available that permit compensation for the effects of
size without using proportions. We chose Analysis of Covariance using
SAS procedures (Barr et al. 1976) to determine if significant differences
could be detected between G. cumberlandius and widely separated pop-
ulations of G. pinetis. Males and females were treated separately
because of obvious sexual dimorphism. A Multiple Discriminant Func-
tion Analysis (SAS) was performed on raw data, separated into male
and female groups to obtain generalized distances between populations.
These were then clustered by UPGMA (Sokal and Sneath 1973) into
distance phenograms to graphically illustrate phenetic distances between
populations.
RESULTS
Initial tests of equal slope in the Analysis of Covariance used the
following model:
V4 (condylobasilar length)
population
V4 X population.
144 Joshua Laerm
Results indicate that the interaction term was not significant for the
characters examined. Therefore, the intercept (the differences between
populations) for the covariate was tested under the following model;
V4
population.
The results of this model (Table 1) indicate significant differences
between populations for most characters. For these characters the dif-
ference in least squares adjusted means for each population was deter-
mined by the Scheffe Test (Morrison 1967) (Table 2).
No significant differences for any of the characters examined can
be seen between females of G. cumberlandius and other populations of
female G. pinetis. Males of G. cumberlandius can be distinquished from
males of other G. pinetis populations on the basis of a single character —
total length.
Results of the Multiple Discriminant Function Analysis (Fig. 2)
similarly indicate low levels of morphological distinction between G.
cumberlandius and mainland populations of G. pinetis. Two assemb-
lages are indicated in both males and females: an upland assemblage
consisting of the populations from Adam, Augusta, and Hursman's
Lake, and a coastal assemblage consisting of the two Camden County
populations and G. cumberlandius.The only inconsistency in clustering
in both males and females occurs in the apparent relatedness of the coast-
al Camden County assemblage. Female G. cumberlandius and Kings-
land G. pinetis appear more closely related than either is to the Scotch-
ville G. pinetis population, while males from both Camden County
populations appear more closely related to each other than either does
to the Cumberland Island population. The important point is, of course,
that G. cumberlandius appears more closely related to coastal Camden
County G. pinetis than do these G. pinetis to their upland conspecifics.
DISCUSSION
Bangs' (1898) description of the insular G. cumberlandius was
based on a small series of specimens (N = 13) collected at "Stafford
Place." He distinquished it from adjacent mainland Georgia and Florida
populations of G. pinetis on the basis of the very large size and slight
pelage and cranial differences. I find that his pelage and cranial features
are generally unsatisfactory to permit the distinction of G. cumberlan-
dius from other populations of southeastern pocket gophers. Pelage in
the G. pinetis complex is extremely variable and tends to be correlated
with local soil color (Williams and Genoways 1980; Laerm et al., in
press). Hence, it has little value in taxonomy. The results of cranial
morphometry reported by Williams and Genoways (1980) and herein
indicate that cranial differences between G. cumberlandius and main-
land populations of G. pinetis are not sufficient to warrant species level
recognition for G. cumberlandius.
Status of Geomys cumberlandius
145
i3 > o
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B >
u
146
Joshua Laerm
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WO
CM
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oo
no r^
cm
co
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WO
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wo
wo ^
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r-
CM ^
en
o
CO
CO
OO
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CM
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WO
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CO
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WO
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CM
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CO
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o
Status of Geomys cumberlandius
147
sO vo
^" so
rsj
™ rsi
oo
<n
>n
m
NO sd
»n so
o
2
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SO
cm r^j
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rsj
rsj ^
«n
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6« -C
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= g
148
Joshua Laerm
Status of Geomys cumberlandius
149
q
T
60
100
80
40
T
20
&
80
60
T"
40
20
100
0
Fig. 2. Distance phenogram for males (A) and females (B). Generalized squared
Euclidean distances derived from Discriminant Function Analysis of raw data.
Ordinate numbers refer to population locations given in Figure 1; abscissa
numbers are generalized squared distances.
The very large size of Cumberland Island gophers in comparison to
mainland forms was the chief criterion for Bangs' recognition of G.
cumberlandius. The data in Table 2 support his observation only in
part, because only total length of males is seen to differ significantly in
comparison to other populations. However, significant size difference
between mainland and insular populations of the same species is not an
uncommon phenomenon in vertebrates, particularly mammals (see Case
1978; Heaney 1978). While insular populations of rodents and other
small mammals are generally larger than mainland forms, the insular
populations are rarely regarded as representing distinct taxa. The large
body size of Cumberland Island gophers is consistent with this general
observation in other small mammals. Thus, body size alone would not
be strong support for species level recognition for Cumberland Island
pocket gophers.
Other indirect evidence also argues against such recognition. First,
Avise et al. (1979) and Laerm et al. (in press) have shown that popula-
tions of pocket gophers throughout eastern Georgia and northeastern
Florida show no detectable protein heterozygosity for 25 loci examined,
150 Joshua Laerm
and no karyological differences; based on mitochondrial DNA sequence
relatedness, they share a common lineage. Second, based on archaeolog-
ical evidence it is believed that Cumberland Island was connected to
mainland Georgia as recently as 5000-7000 years Before Present (R.
Frey, pers. comm.).
Most species and subspecies groups of Recent mammals are, at the
youngest, of Pleistocene origin (Hibbard et al. 1965). Russel (1968) sug-
gested that G. pinetis differentiated from Geomys bursahus by the Sang-
amon, and G. pinetis is recorded from Irvingtonian to Recent deposits
in Florida (Webb 1974). Isolation of a population of pocket gophers on
Cumberland Island for 5000-7000 years would, in general, be recognized
as too short a period for speciation.
I conclude that the most parsimonious interpretation consistent
with available data is that G. cumberlandius cannot be shown to be
distinct from mainland populations of G. pinetis. I therefore agree with
the conclusions of Williams and Genoways (1980) that pocket gophers
on Cumberland Island be synonymized with adjacent mainland G. pine-
tis.
ACKNOWLEDGMENTS.— I wish to thank the curatorial staff of
the American Museum of Natural History (ANMH), Carnegie Museum
of Natural History (CMNH), Cornell University Museum (CU), Dela-
ware Museum of Natural History (DMNH), Florida State Museum
(FSM), Museum of Comparative Zoology (MCZ), and U.S. National
Museum of Natural History (NMNH) for access to their respective col-
lections. Additional specimens from the University of Georgia Museum
of Natural History (UGAMNH) were included in this study. The coop-
eration of Ron Odom of the Georgia Department of Natural Resources
is much appreciated, as was the statistical advice of T. Richardson. This
work was financed largely by grant-in-aid funds under Section 6 of the
Endangered Species Act of 1973, the Theodore Roosevelt Fund of the
American Museum, and the Department of Zoology, University of
Georgia. This is a contribution of the University of Georgia Museum of
Natural History.
LITERATURE CITED
Albrecht, Gene H. 1978. Some comments on the use of ratios. Syst. Zool.
27:67-71.
Atchley, William R., and D. Anderson. 1978. Ratios and statistical analysis of
biological data. Syst. Zool. 27:71-78.
C. T. Gaskins and D. Anderson. 1976. Statistical properties of ratios.
I. Empirical results. Syst. Zool. 25:137-148.
Avise, John C, C. Giblin-Davidson, J. Laerm, J. C. Patton and R. A. Lans-
man. 1979. Mitochondrial DNA clones and matriarchal phylogeny within
and among geographic populations of the pocket gopher, Geomys pinetis.
Proc. Natl. Acad. Sci. U.S.A. 7(5:6694-6698.
Status of Geomys cumberlandius 151
Bangs, O. 1898. The land mammals of peninsular Florida and coast region of
Georgia. Proc. Bost. Soc. Nat. Hist. 25:157-235.
Barr, A. J., J. H. Goodnight, S. P. Sail and J. T. Helwig. 1976. A user's guide to
SAS 76. SAS Institute, Inc. 329 pp.
Case, Ted C. 1978. A general explanation for insular body size trends in terres-
trial vertebrates. Ecology 59(1): 1-18.
DeBlase, Anthony F., and R. E. Martin. 1974. A manual of mammalogy. Wm.
C. Brown, Dubuque. 329 pp.
Dodson, Peter. 1978. On the use of ratios in growth studies. Syst. Zool.
27:62-67.
Foster, J. B. 1965. The evolution of the mammals of the Queen Charlotte
Islands, British Columbia. Occas. Pap. British Columbia Prov. Mus.
74:1-130.
Hall, E. Rayond. 1981. The Mammals of North America. 2nd Ed. John Wiley
and Sons, New York. 1181 pp.
, and K. R. Kelson. 1959. The Mammals of North America. Ronald
Press, New York. 1083 pp.
Heaney, Lawrence R. 1978. Island area and body size of insular animals; evi-
dence from the tricolored squirrel (Callosciurus prevosti) of southwest Asia.
Evolution 32(l):29-44.
Hibbard, Claude W., W. D. Ray, D. E. Savage, D. W. Taylor and J. E. Guil-
day. 1965. Quarternary mammals of North America, pp. 509-525 in H. E.
Wright, Jr. and D. G. Frey (eds.). The Quarternary of the United States.
Princeton Univ. Press, Princeton. 922 pp.
Hill, Michael. 1978. On ratios--a reply to Atchley, Gaskins, and Anderson. Syst.
Zool. 27:61-62.
Laerm, Joshua, J. C. Avise, J. C. Patton and R. A. Lansman. In press. The genetic
determination of the status of an endangered species of pocket gopher in
Georgia. J. Wildl. Mgmt.
Morrison, Donald. 1967. Multivariate Statistical Methods. McGraw Hill, New
York. 338 pp.
Russell, R. J. 1968. Evolution and classification of the pocket gophers of the
subfamily Geomyinae. Univ. Kans. Publ. Mus. Nat. Hist. 76:473-579.
Sneath, P. H. A., and R. R. Sokal. 1973. Numerical Taxonomy. W. H. Free-
man Co., San Francisco. 573 pp.
Webb, S. David (ed.) 1974. Pleistocene mammals of Florida. Univ. Presses Fl.,
Gainesville. 270 pp.
Williams, Stephen L., and H. H. Genoways. 1977. Morphological variation in
the tropical pocket gopher {Geomys tropicalis). Ann. Carnegie Mus. 46: 245-264.
, and 1980. Morphological variation in the southeastern pocket
gopher, Geomys pinetis (Mammalia: Rodentia). Ann. Carnegie Mus. 49:405-453.
Accepted 31 October 1981
New Records and Habitat Observations of
Hyla andersoni Baird (Anura: Hylidae) in
Chesterfield and Marlboro Counties, South Carolina
Janice Heard Tardell, Richard C. Yates,
and
David H. Schiller
Carolina Power and Light Company,
Harris Energy and Environmental Center,
Route 1, Box 327, New Hill, North Carolina 27562
ABSTRACT. — Sixty-seven Hyla andersoni localities were found in
Marlboro (56) and Chesterfield (11) counties, South Carolina in 1978
(41) and 1979 (26). Eight sites were in electrical transmission line or
gas pipeline rights-of-way, fifteen in recent clearcuts, and forty-four in
shrub bog habitats. Hyla andersoni apparently is more widely
distributed in South Carolina than was previously known. There seems
to be potential for maintaining or increasing suitable habitat through
appropriate management techniques.
INTRODUCTION
The Pine Barrens Treefrog, Hyla andersoni, was described by Baird
(1854) on the basis of a specimen received at the U.S. National Museum
from Anderson, South Carolina. The origin of the type specimen has
been questioned by several authors (Neill 1947, 1957; Wright and
Wright 1949; Brown 1980). After the original South Carolina record, H.
andersoni was not found again in the state until 1950, when specimens
were collected in Chesterfield County near the town of Patrick and a
population was discovered in Kershaw County (Brown 1980). A 1975
survey of the Carolina Sandhills National Wildlife Refuge in Chesterfield
County found H. andersoni to be present at 16 localities (Garton and
Sill 1979).
The breeding habitat of H. andersoni was described by Means and
Longden (1976) as occurring where water seeps laterally from sand-
capped hills, causing a sharp separation between the moisture requiring
shrubby-herbaceous communities in stream bottoms and the dry, pine-
oak-wiregrass communities of the hills and slopes. The plant com-
munities developing in such seepages have been described variously as
pocosins (Wells 1928), dismals (Kerr 1875), and bays or evergreen shrub
bogs (Kologiski 1977). Means and Moler (1978) differentiated between
adult and larval habitat of H. andersoni; shrub bogs constitute adult
habitat and herb bogs are appropriate larval habitat. They reported that
herb bogs typically occur on sandy soils upland of shrub bogs and are
dominated by grasses, sedges, and forbs. Herb bogs are usually wetter
on the surface than their shrub bog counterparts and are a transitional
Brimleyana No. 6: 153-158. December 1981. 153
154 Janice Heard Tardell, Richard C. Yates, David H. Schiller
stage to shrub bogs. Apparently, H. andersoni populations thrive where
these two habitat components occur together.
In this paper we report new information on the distribution and
habitat of H. andersoni in Chesterfield and Marlboro counties. During
the study we found the species inhabiting seepages in shrub and herb
bogs, a predictable habitat, but also found it in seeps of electrical
transmission line and gas pipeline rights-of-way, clearcut areas, and in a
broomsedge field.
METHODS AND MATERIALS
Study Area
We discovered Hyla andersoni in Marlboro County during a power
plant feasibility study for Carolina Power and Light Company. We then
investigated its occurrence along the drainages of Whites, Wolfs, Hicks,
Marks, and Phils creeks in northwestern Marlboro County and along
the Juniper Creek drainage in Chesterfield County.
Most of the area investigated lies in the northern part of South
Carolina's sandhill region and is located east of the dividing line
between the Piedmont and Coastal Plain physiographic provinces. The
dominant vegetation of the area is the longleaf pine-turkey oak-
wiregrass (Pinus palustris-Quercus laevis- Aristida stricta) association.
Loblolly pine, P. taeda, becomes codominant with longleaf, especially in
moister areas. In the drier areas post oak, Quercus stellata; and black-
jack oak, Q. marilandica; become more numerous. In several large areas
south of Whites Creek the natural forests have been logged, cleared, and
replaced with monocultures of slash pine, P. elliottii, or left to regener-
ate naturally. Some of these areas were in the first or second season of
growth. Along the small stream basins that drain the sandhills, the vege-
tation is composed of "pocosin" or "evergreen shrub" species. The dom-
inants are sweet bay, Magnolia virginiana\ titi, Cyrilla racemiflora; var-
ious members of the heath family (Ericaceae); and other evergreen
shrubs, along with poison sumac, Rhus vernix. In many places the vege-
tation is covered with a tangle of greenbriers, Smilax spp.
Methods
United States Geological Survey 7.5 minute series topographic
maps were used to locate stream headwaters and intermittent streams.
These and other likely areas were inspected during the day to identify
appropriate habitat. We returned to the preidentified sites during the
evening to listen for the call of H. andersoni. Voice identifications were
verified by following the sounds to their source and observing calling
males, whose numbers were then estimated. For our purposes, a locality
where H. andersoni was found is one that is separated from another
locality by an area where the frogs do not call. The localities were
Hyla andersoni in South Carolina 155
widely separated and generally were located along different tributary
streams or seepages.
During 1978 we searched appropriate habitats from 11 through 22
July and from 25 through 27 July. Most effort at that time was directed
at the shrub bog habitat. We began our 1979 searches in May and con-
tinued them through the summer. During this time we discovered that
H. andersoni occurred in some clearcuts and transmission line rights-of-
way, so emphasis was placed on searching for and surveying these areas.
However, this did not preclude searches in the typical shrub bogs. All
Marlboro County localities found in 1978 were rechecked in 1979 to
investigate the continued presence of H. andersoni, but the Chesterfield
County locations were not revisited.
RESULTS AND DISCUSSION
Locations
Hyla andersoni was first found in Marlboro County on 20 June
1978 in a shrub bog located in the headwaters of a tributary of Whites
Creek. In all, 67 localities were identified in Marlboro (56) and Chester-
field (11) counties during the summer of 1978 (41) and 1979 (26) (Figure
1). The continued presence in 1979 of H. andersoni was verified at 24 of
the 30 1978 localities in Marlboro County. At the six sites (all well-
developed shrub bogs) where H. andersoni was not found again in 1979,
the habitat was undisturbed. No human activities were evident.
Habitat
Plant species composition at most of the 67 sites was similar to that
reported by Means and Moler (1978). The dominant species are listed in
Table 1.
Of the 26 localities found in 1979, 8 were located along transmis-
sion line or gas pipeline rights-of-way. These areas were similar in struc-
ture to the shrub bog-herb bog habitat depicted by Means and Moler.
The herb bogs developed where the actual clearing for the rights-of-way
crossed a wet area, and the shrub bogs developed to both sides where
canopy trees had been removed. Since the rights-of-way are periodically
maintained, they would not undergo succession to a closed canopy
forest that would preclude H. andersoni.
Fifteen of the localities found in 1979 were in clearcuts to the east
of U.S. 1 and south of Whites Creek. These areas had been cut little
more than a year prior to our finding H. andersoni in them. We do not
know if the species inhabited these areas before the logging occurred,
but cursory checks in 1978 did not reveal their presence. The clearcuts
had herbaceous seepages surrounded by small shrubby stands at slightly
higher elevations; the shrub species were typical of older shrub bogs.
156
Janice Heard Tardell, Richard C. Yates, David H. Schiller
2
c/5
J*
u
•o
o
Hyla andersoni in South Carolina 157
The herbaceous species consisted of various grasses, sedges and forbs,
and in most cases Sphagnum spp. were present. One large clearcut had
developed into a broomsedge field uncharacteristic of the other clear-
cuts in the area. However, it had much relief and seepage and we found
H. andersoni at two localities there.
Table 1. Dominant plant species in herb and shrub bogs where Pine Barrens
Treefrogs were found.
Clethra alnifolia, sweet pepperbush Osmunda cinnamomea, cinnamon fern
Smilax laurifolia, bamboo smilax Cyrilla racemiflora, titi (cyrilla)
Magnolia virginiana, sweet bay Sphagnum spp., sphagnum moss
Pinus taeda, loblolly pine Nyssa sylvatica, black gum
Rhus vernix, poison sumac Liriodendron tulipifera, yellow poplar
Arundinaria gigantea, switch cane Oxydendrum arboreum, sourwood
Rhus copallina, winged sumac Acer rubrum, red maple
Persea borbonia, red bay Ilex coriacea, sweet gallberry
Calling Males
In 1978 the number of calling males ranged from 1 to an estimated
100, with an estimated mean of 12 per locality. However, of the 41
localities, only 4 were estimated to have greater than 15 calling males.
During 1979 the range at the 26 sites was from 1 to an estimated 25,
with an estimated mean of 6 per locality.
CONCLUSIONS
Apparently H. andersoni, although restricted in range, is more
widely distributed in South Carolina than previously known. The area
we searched was only a small part of the total area of similar habitat in
South Carolina, and further research may reveal the presence of H.
andersoni in other counties where it is now unknown.
Our observations suggest that H. andersoni quickly colonizes areas
of appropriate new habitat (clearcuts with seepages). Means and Moler
(1978) reported that clearcuts constitute good H. andersoni habitat, as
long as succession to a closed canopy is prohibited and pine monocul-
tures are not planted. Thus, as they suggested, there seems to be a
potential for maintaining, or even increasing suitable habitat for H.
andersoni in areas where suitable soil, topography, and groundwater
seepage exists. By applying habitat management practices such as pre-
scribed burning and selective logging and clearing, it is probable that
the open shrubby vegetative communities required by H. andersoni
could be sustained where currently existing, or restored where succes-
sion has progressed beyond the favorable stage.
158 Janice Heard Tardell, Richard C. Yates, David H. Schiller
ACKNOWLEDGMENTS.— Many biologists at Carolina Power and
Light Company participated in this study. We would like to especially
thank Dr. B. J. Ward, Ms. C. W. Anderson and Mr. R. L. Corpening
for their help in the field work. Our thanks also are extended to Dr.
Paul Feaver and to two anonymous reviewers for their helpful com-
ments on this manuscript.
LITERATURE CITED
Baird, Spencer F. 1854. Descriptions of new genera and species of North
American frogs. Proc. Acad. Nat. Sci. Phila. 7:59-62.
Brown, E. E. 1980. Some historical data bearing on the pine barrens treefrog,
Hyla andersoni, in South Carolina. Brimleyana 5:1 13-1 18.
Garton, John S., and B. L. Sill. 1979. The status of the pine barrens treefrog,
Hyla andersonii Baird, in South Carolina, pp. 131-132 in D. M. Forsyth
and W. B. Ezell, Jr. (eds). Proc. First S. C. Endangered Species Symp. The
Citadel, Charleston. 201 pp.
Kerr, Washington C. 1875. Physical geography, resume, economical geography.
Vol. 1. N. C. Geol. Sur. Rep., Raleigh. 120 pp.
Kologiski, Russell L. 1977. The phytosociology of the Green Swamp, North
Carolina. N. C. State Univ. Agric. Exp. Stn. Tech. Bull. No. 250. 101 pp.
Means, D. Bruce, and C. J. Longden. 1976. Aspects of the biology and zoo-
geography of the pine barrens treefrog (Hyla andersonii) in northern Flor-
ida. Herpetologica. 32(2): 117-130.
, and P. E. Moler, 1978. The pine barrens treefrog; fire, seepage
bogs, and management implications, pp. 77-83 in R. R. Odom and L.
Landers (eds). Proc. Rare Endangered Wild. Symp., Ga. Dep. Nat. Res-
our., Athens. 184 pp.
Neill, Wilfred T. 1947. Doubtful type localities in South Carolina. Herpetolog-
ica 4(2):75-76.
1957. Objections to wholesale revision of type localities. Copeia 1957
(2): 140-141.
Wells, B. W. 1928. Plant communities of the coastal plain of North Carolina
and their successional relations. Ecology 9:230-242.
Wright, Albert H., and A. A. Wright. 1949. Handbook of frogs and toads of the
United States and Canada. Comstock Publ. Co., Inc., Ithaca. 640 pp.
Accepted 23 March 1981
Observations on Some Maritime Forest Spiders
of Four South Carolina Barrier Islands
L.L. Gaddy
Route 1, Box 223, Walhalla, South Carolina 29691
ABSTRACT. — Quantitative observations on the seasonal abundance
of 22 species of araneid orb weavers and general observations on five
non-orb weaver species were made on four South Carolina barrier
islands. Data collected along transects revealed that 5 of 22 orb weaver
species matured in spring, 8 in early summer, 7 in late summer, and 2
in autumn. The greatest number of mature individuals of all orb
weaver species was found in early summer, but the greatest number of
species matured in late summer. Araneus bicentenarius, thought to be
rare, was found on all four islands.
INTRODUCTION
The spider fauna of the outer coastal plain of the Carolinas is rela-
tively unknown. Barnes (1953) and Barnes and Barnes (1954) studied
the ecology and species composition of spider communities in non-
forested maritime communities near Beaufort, North Carolina but did
not deal with the maritime forest. Berry (1971) discussed the seasonal
distribution of many of the species found on barrier islands, but his field
work was done in the Piedmont of North Carolina, an area climatically
different from the barrier islands of South Carolina.
Maritime forest is the dominant vegetation cover type on the
barrier islands of South Carolina, where live oak, Quercus virginiana;
laurel oak, Quercus laurifolia; palmetto, Sabal palmetto; southern magno-
lia, Magnolia grandiflora; and various pines (notably Pinus taeda and
Pinus elliottii) dominate the canopy. Red bay, Persea borbonia; yaupon
holly, Ilex vomitoria; American holly, Ilex opaca; and palmetto are the
most commonly encountered species in the understory and shrub layers.
The canopy and understory are extremely dense, with nearly 100
percent coverage; the shrub layer's average coverage, on the other hand,
is low, varying from 10 to 50 percent. The coverage in the herbaceous
layer is usually less than 10 percent.
From February to November 1979, monthly observations and
collections were made along transects on each of four South Carolina
barrier islands — Bulls, Kiawah, Capers, and Hunting islands. These tran-
sects, which averaged approximately one kilometer long, were walked at
least once a month between the hours of 0800 and 1700. All orb webs
and retreats between ground level and 2.7 meters high were checked for
adult spiders. The total number of individuals of each sex was recorded.
Brimleyana No. 6: 159-162. December 1981. 159
160 L. L. Gaddy
The data here are based primarily on field identification. Because of the
large number of spiders handled, only taxonomically difficult species
and representative specimens of common species were collected. Voucher
specimens are in the author's personal collection. General notes were
taken on the presence of non-araneid orb weavers and non-orb weavers.
RESULTS AND DISCUSSION
Twenty-two species of araneids were found on the four islands.
Table 1 lists the number of adults recorded per transect in winter (Feb-
ruary), spring (March and April), early summer (May and June), late
summer (July and August), and autumn (September and October). No
perceptible differences in population or species number were found
among the four islands. Most species, however, did exhibit some degree
of seasonality, as seen in Table 1.
Of the 22 araneid species, the genus Araneus was represented by 4
species, M angora by 3, and Neoscona, Micrathena, and Argiope by 2
species each. As seen in Table 1, only Araneus pegnia was found to be
in the adult stage in winter. Araneus miniatus, another small Araneus,
was observed in the spring, along with additional A. pegnia. Araneus
pratensis, the third small Araneus found on the islands, occurred in
forest openings only in late summer.
The spring dominants were M angora placida and Acanthepeira
spp. (Because of the difficulties of field identification of the large
number of Acanthepeira individuals encountered, these spiders were
identified only to genus.) Mangora maculata and M. gibberosa did not
mature until early summer, confirming the observations of Berry (1971).
The third most frequent species found mature in the spring was Araneus
bicentenarius. Levi (1971) thought this species rare in North America,
but its giant webs were seen frequently from early March through May
on the four islands studied. One individual seen in March was over 20
mm long, possibly having overwintered as an adult (however, no sub-
adults were seen on the transects in February). The retreat of A. bicen-
tenarius was usually made in Spanish moss, Tillandsia usneoides, on the
four study islands.
In early summer, Acanthepeira individuals became the dominant
adults, reaching a peak in early June. Neoscona arabesca matured in
early June and continued to be common into October. Leucage venusta
individuals were common from late May to July. Argiope trifasciata
peaked in abundance in late summer in shrubby areas. In late July,
females of Nephila clavipes were undergoing their final molt with males
beginning to appear in their webs. Mating frequently took place during
the final molt while the female hung defenseless in her web (see Robin-
son and Robinson 1973, 1976). Nephila, however, did not become the
most abundant spider until August. In late summer and autumn, Neo-
scona domiciliorum began to appear in the wetter areas of the maritime
Barrier Island Spiders
161
Table 1. Frequency (number of adults per transect) of orb weavers on four South
Carolina barrier islands.
Totals
.3
21.2
57.5
31.4
43.0
forests. By this time, Nephila clavipes was the overwhelmingly dominant
species (see Table 1).
The spiny-bodied orb weavers, Micrathena gracilis, Micrathena
sagittata, and Gasteracantha cancriformis, were not as common in the
maritime forest as they are on the adjacent mainland. It must be pointed
out, however, that G. cancriformis was more common than Table 1
indicates, being frequently seen in its web above 2.7 meters (spiders in
webs more than 2.7 meters above the ground were not counted due to
the difficulty of collecting and identifying these individuals).
Nephila clavipes, Acanthepeira spp., and Neoscona arabesca were
the most numerous of the 22 orb weaver species. More orb weaver indi-
viduals matured in early summer; however, more species of orb weavers
162 L. L. Gaddy
matured in late summer (see Table 1). The autumn totals in Table 1 are
artificially high because of the large number of Nephila clavipes, a visu-
ally and numerically dominant species.
General observations on non-orb weavers indicate that Latrodectus
mactans is common under debris in dunes dominated by sea oats,
Uniola paniculata. Large males (7 mm body length) of L. mactans were
found in the maritime forest. Three species of the inquilinous Argyrodes
were found on the four islands: Argyrodes fictilium (Hentz) [-Rhom-
phaea lacerta (Walckenaer)], Argyrodes furcatus (O. P. -Cambridge), and
Argyrodes nephilae Taczanowski. Argyrodes nephilae was found in the
webs of Acanthepeira spp. and Tidarren sisyphoides, as well as those of
Nephila clavipes.
Carico (1973) noted that the genus Dolomedes was absent from
"most islands off the coast of the southeastern United States," salt water
being a barrier to their dispersal. However, during my study Dolomedes
triton (Walckenaer) was found in a freshwater wetland less than 50 m
from the beach front.
LITERATURE CITED
Barnes, Robert D. 1953. The ecological distribution of spiders in non-forested
maritime communities at Beaufort, North Carolina. Ecol. Monogr.
2i:315-377.
Barnes, Betty M., and R. D. Barnes. 1954. The ecology of the spiders of mari-
time drift lines. Ecology 55:25-35.
Berry, J. W. 1971. Seasonal distribution of common spiders in the North
Carolina piedmont. Am. Midi. Nat. 55:526-531.
Carico, James E. 1973. The Nearctic species of the genus Dolomedes (Araneae,
Pisauridae). Bull. Mus. Comp. Zool. 744:435-488.
Levi, Herbert W. 1971. The Diadematus group of the orb weaver genus Araneus
north of Mexico (Araneae, Araneidae). Bull. Mus. Comp. Zool. 747:131-179.
Robinson, Michael H., and B. Robinson. 1973. Ecology and behavior of the
giant wood spider Nephila maculata (Fabricius). Smithson. Contrib. Zool.
749:1-76.
, and 1976. The ecology and behavior of Nephila maculata: a
supplement. Smithson. Contrib. Zool. 275:1-22.
Accepted 17 July 1981
Foods of Two Species of Plethodon (Caudata:
Plethodontidae) from Georgia and Alabama
Carlos D. Camp
Department of Zoology,
University of Georgia, Athens, Georgia 30602
AND
Luke L. Bozeman
Department of Zoology- Entomology,
Auburn University, Auburn, Alabama 36830
ABSTRACT.— The stomachs of 34 Plethodon websteri and 55 P. ser-
ratus from Georgia and Alabama were examined for food. Acarines
and Collembola were major food items in stomachs of smaller P. web-
steri whereas ants were predominant in the stomachs of larger individ-
uals. Ants were the dominant food item in the stomachs of P. serratus.
Very little is known concerning the ecology of the Southern Redback
Salamander, Plethodon serratus Grobman, which was recently taxo-
nomically separated from the Redback Salamander, Plethodon cinereus
(Highton and Webster 1976). Several reports exist on the foods of P.
cinereus (Blanchard 1928; Hamilton 1932; Jameson 1944; Jaeger 1972;
Caldwell and Jones 1973; Fraser 1976), but the only report concerning
the prey of P. serratus is a single statement by Johnson (1977) that it
feeds on arthropods. The foods of the recently described Webster's
Salamander, Plethodon websteri Highton (formerly considered Plethodon
dorsalis), have also not been reported, although Holman (1955) dis-
cussed the foods of P. dorsalis in Indiana. We present here an account
of the foods of P. serratus and P. websteri in Georgia and Alabama.
Fifty-five P. serratus were collected from Fulton and Harris coun-
ties, Georgia, during March and April of 1980. During the same period,
34 P. websteri were collected from Upson County, Georgia, and Lee
County, Alabama. Specimens were sacrificed in the field in chloretone
and preserved in 10% formalin to terminate digestion. In the laboratory,
stomach contents were placed in a petri dish lined with a paper grid (2.5
mm x 2.5 mm) and examined under a dissecting microscope. In order to
determine relative prey proportion in the diet, visual estimates of rela-
tive area occupied by prey items were made by comparing each prey
item to the grid and estimating the number of grid squares it occupied.
Snout-vent length (SVL) was recorded for each specimen. The foods of
different size classes were compared when appropriate.
Nineteen specimens of P. websteri had SVLs of 22-27 mm; the
remaining 15 specimens each has a SVL greater than 30 mm. Smaller
Brimleyana No. 6: 163-166. December 1981. 163
164
Carlos D. Camp and Luke L. Bozeman
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Foods of Plethodon 165
individuals differed noticeably from larger ones in prey selected (Table
1). Acarines (mites) were the dominant prey in the stomachs of small P.
websteri, both in numbers and percent area. Their next most important
food item was Collembola (springtails). Acarines were frequently
encountered in the stomachs of the larger P. websteri but did not con-
tribute much to percent area. Ants, however, were important in their
diet. Isoptera (termites) were important in both area and numbers in
larger P. websteri; however, they were not preyed upon frequently as
shown by the low percentage of stomachs containing them. These
results show that larger salamanders feed on larger prey (e.g., ants and
termites), and smaller salamanders feed on smaller prey (e.g., springtails
and mites).
Only one P. serratus had a SVL less than 30 mm, the remaining 54
being comparable in size to the larger P. websteri. Therefore, the food
data for P. serratus are not reported for separate size groups. Ants were
the most important prey of P. serratus in area and total numbers. Acar-
ines were common in the stomachs but contributed little to percent area.
Beetles, spiders, and isopods also were frequently eaten by this species.
Most of the apparent differences in feeding between the two species
of salamanders can be attributed to the large number of smaller P. web-
steri in the samples and the predominance of larger specimens in the P.
serratus samples. Although the foods of both species appear to be sim-
ilar, further work is needed on the feeding as well as other aspects of the
ecology of these salamanders before conclusions can be drawn concern-
ing their respective feeding strategies.
ACKNOWLEDGMENTS.— We would like to thank William E.
Garrett, Jr., Auburn University, for identification of some of the prey
items, and George W. Folkerts, Auburn University, for critically review-
ing the manuscript.
LITERATURE CITED
Blanchard, F. N. 1928. Topics from the life history and habits of the red-backed
salamander in southern Michigan. Am. Nat. 62(619): 156-164.
Caldwell, Ronald S., and G. S. Jones. 1973. Winter congregations of Plethodon
cinereus in ant mounds, with notes on their food habits. Am. Midi. Nat.
90(2):482-485.
Fraser, D. F. 1976. Empirical evaluation of the hypothesis of food competition
in salamanders of the genus Plethodon. Ecology 57(3):459-471.
Hamilton, W. J., Jr. 1932. The food and feeding habits of some eastern sala-
manders. Copeia 1932(2):83-86.
Highton, Richard, and T. P. Webster. 1976. Geographic protein variation and
divergence in populations of the salamander Plethodon cinereus. Evolution
50(l):33-45.
Holman, J. A. 1955. Fall and winter food of Plethodon dorsalis in Johnson
County, Indiana. Copeia 1955(2): 143.
166 Carlos D. Camp and Luke L. Bozeman
Jeager, R. G. 1972. Food as a limited resource in competition between two
species of terrestrial salamanders. Ecology 53(3):535-546.
Jameson, E. W., Jr. 1944. Food of the red-backed salamander. Copeia
1944(3): 145-147.
Johnson, T. R. 1977. The Amphibians of Missouri. Univ. Kans. Mus. Nat.
Hist., Lawrence. 134 pp.
Accepted 24 July 1981
167
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DATE OF MAILING
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ACKNOWLEDGMENT OF REVIEWERS
We are indebted to many reviewers who served as referees for manuscripts
for Numbers 3, 4, and 5. Their names and institutional affiliations will be pro-
vided in Number 7.
TABLE OF CONTENTS AND INDEX
An index of scientific names and a table of contents for 1981 issues will
appear in Number 7.
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CONTENTS
Fishes of the Waccamaw River Drainage. John R. Shute, Peggy W.
Shute and David G. Lindquist 1
A Taxonomic Analysis of Pseudemyd Turtles (Testudines: Emydidae)
from the tyew River, and Phenetic Relationships in the Subgenus
Pseudemys. Michael E. Seidel 25
Small Mammals in Openings in Virginia's Dismal Swamp. Robert K.
Rose 45
A New Milliped of the Genus Brevigonus from South Carolina, with
Comments on the Genus and B. shelfordi (Loomis) (Polydesmida:
Xystodesmidae). Rowland M. Shelley 51
New Records of Marine Fishes from the Carolinas, with Notes on
Additional Species. Steve W. Ross, Garnett W. Link, Jr. and Kerry A.
MacPherson 61
Nesting and Management of the Atlantic Loggerhead, Caretta caretta
caretta (Linnaeus) (Testudines: Cheloniidae) on Cape Island, South
Carolina, in 1979. John B. Andre and Larry West 73
Distribution, Morphology and Life History of the Least Brook Lam-
prey, Lampetra aepyptera (Pisces: Petromyzontidae), in Kentucky.
Stephen J. Walsh and Brooks M. Burr 83
Notes on the Distribution and Taxonomy of Short-tailed Shrews
(Genus Blarina) in the Southeast. Thomas W. French 101
Observations of a Small Population of Estuarine-inhabiting Alligators
near Southport, North Carolina. William S. Birkhead and Charles R.
Bennett Ill
A Key to the Tadpoles of North Carolina. Joseph Travis 119
New Distributional Records of Eastern Kentucky Fishes. Melvin L.
Warren, Jr 129
Systematic Status of the Cumberland Island Pocket Gopher, Geomys
cumberlandius . Joshua Laerm 141
New Records and Habitat Observations of Hyla andersoni Baird (Anura:
Hylidae) in Chesterfield and Marlboro Counties, South Caro-
lina. Janice Heard Tardell, Richard C. Yates and David H.
Schiller 153
Observations on Some Spiders of Maritime Forests on Four South
Carolina Barrier Islands. L. L. Gaddy 159
Foods of Two Species of Plethodon (Caudata: Plethodontidae) from
Georgia and Alabama. Carlos D. Camp and Luke L. Bozeman 163
Miscellany 1 67