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number 22

June 1995

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of Natural Sciences

Brimleyana, the Zoological Journal of the North Carolina State
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CODN BRIMD 7
ISSN 0193-4406

Aspects of the Feeding Ecology of the

Little Grass Frog, Pseudacris ocularis

(Anura: Hylidae)

Jeremy L. Marshall1 and Carlos D. Camp

Department of Biology, Piedmont College

Demorest, Georgia 30535

ABSTRACT— We report on the foods of the little grass frog,
Pseudacris ocularis, from Georgia. Fifty specimens were collected
from two isolated wetlands located in Evans and Grady counties,
Georgia, during late spring and summer 1993. Analysis of stomach
contents determined that the most abundant food items were
small arthropods associated with leaf litter and soil. Almost
50% of the food items were collembolans, followed by hymenopterans
(17%), acarines (9%), homopterans (8%), and coleopterans (8%).
We compared foods of adult males with those of newly meta-
morphosed juveniles collected at the same time from the Grady
County site. Juvenile frogs ate more individual food items and
a greater diversity of prey species than did adult males. This
difference could be due to adult Pseudacris selecting larger,
more profitable prey than juveniles select. Lower feeding activity
exhibited by breeding males might also be a contributing factor.

Little is known about the feeding ecology of many amphibians,
especially intraspecific variability in foods and foraging (Duellman
and Trueb 1986). Variation in dietary preferences among population
subgroups (e.g., breeding males, non-breeding females, subadults,
juveniles, larvae, etc.) has been reported to reflect differences in
habitat preference (Lamb 1984), gape (Toft 1980), developmental
condition (Brophy 1980, Davie 1991), and other factors.

The little grass frog, Pseudacris ocularis (Bosc and Daudin), is
the smallest North American anuran (Conant and Collins 1991). It
occurs in a wide variety of ephemeral and semi-permanent wetlands
in the southeastern Coastal Plain and favors grassy areas in and around
cypress ponds and similar sites (Harper 1939, Mount 1975). In spite
of its relative abundance in many of these areas, virtually nothing
is known of the feeding ecology of this frog. The purpose of our
study was to describe the diet of P. ocularis and to investigate any
potential differences between the feeding of adult frogs and newly
metamorphosed juveniles.

1 Present address: Biology Department, University of Mississippi, University,
Mississippi 38677.

Brimleyana 22:1-7, June 1995 1

Jeremy L. Marshall and Carlos D. Camp

MATERIALS AND METHODS

We collected 50 P. ocularis for stomach analysis from two
localities during May-July 1993. Both sites were ephemeral wetlands
in the lower Coastal Plain of Georgia. The first site, located in Grady
County, was dominated by black gum (Nyssa sylvatica) and was situ-
ated in a low pine flatwood having a canopy of slash pine (Pinus
elliottii) and an understory of saw palmetto (Serenoa repens) and
gallberry (Ilex glabra). The second, in Evans County, was a dome of
pond cypress (Taxodium ascendens) surrounded by sandhills domi-
nated by longleaf pine (P. palustris) and turkey oak (Quercus laevis).
Areas similar to both sites were described in detail by Wharton (1978).

After collection, all specimens were preserved in 10% formalin,
and stored in 35% isopropanol. Each frog was measured for snout-
vent length (SVL) and dissected for stomach analysis. Individual food
items were counted and identified. Because prey items were too small
to use volumetric displacement, relative importance of prey was de-
termined by comparing each individual prey item to a paper grid and
visually estimating the number of grid squares occupied (Camp and
Bozeman 1981).

Twenty of the Grady County frogs were collected between 2200
and 2400 EDT on 5 June. This sample consisted of 10 mature males
and 10 juveniles that had just completed metamorphosis. We used
this sample to make comparisons between feeding of adults and juve-
niles. Because we did not independently test for prey availability,
other collections were not used for comparisons because of possible
complications arising from temporal or between-site differences in
available prey items. In addition, although adult females were in-
cluded in these samples, small numbers (n = 4) precluded between-
sex comparisons. Correlation between the number of prey items eaten
and body size was tested using the procedure described by Zar
(1984). A comparison of diversity between adult male and juvenile
prey species was made using the Shannon-Wiener Index of Diversity
(H') (Zar 1984).

RESULTS

One hundred-forty individual prey items were identified and
consisted entirely of arthropods, mainly insects (Table 1). Springtails
(Collembola) were the most numerous group, making up 47% of the
food items eaten and found in 56% of the stomachs. Because they are
so small, however, they contributed less than 20% of the area occu-
pied by all prey items. Hymenopterans, especially ants (Formicidae)

Little Grass Frog Feeding Ecology

Table 1. Stomach contents of 50 Pseudacris ocularis from the Coastal
Plain of Georgia. Unless otherwise indicated, the smallest taxon in
each order is represented by a single species; "i" repreesnts immature
instars. Numbers (n) for higher taxa also include unidentified food
items.

Percentage of

Food Item

INSECTA
Collembola

Isotomidae

Poduridae

Sminthuridae
Coleoptera

Carabidae

Cleridae

Coccinellidae

Staphylinidae (3)

larvae
Dictyoptera

Blattidae
Diptera (larvae) (3)
Homoptera

Delphacidae (i)
Hymenoptera

Diapriidae

Dryinidae

Encyrtidae

Evaniidae

Formicidae

Scelionidae
Orthoptera

Acrididae (i)
Phasmida

Phasmatidae
Siphonaptera
Thysanoptera

Phlaeothripidae
ARACHNIDA
ACARINA

Mesostismata

Oribatei (2)
Araneida

Anyphaenidae
Palpigradi

(n) Total Number Total Area Frequency

122
66
26

2

37

11

1

1

1

5

1

2

2

3

11

10

24

1

1

1

2

7

7

2

2

1

1

1

1

1

18

13

1

12

4

1

1

87.1
47.1
18.6
1.4
26.4
7.9
0.7
0.7
0.7
3.6
0.7
1.4
1.4
2.1
7.9
7.1
17.1
0.7
0.7
0.7
1.4
5.0
5.0
1.4
1.4
0.7
0.7
0.7
0.7
0.7
12.9
9.3
0.7
8.6
2.9
0.7
0.7

90.1
19.5
5.6
0.5
13.4
9.2
1.6
0.3
1.1
4.3
0.4
19.1
19.1
1.2
11.1
10.9
14.6
2.2
0.8
0.1
6.4
1.2
1.9
4.8
4.8
9.6
9.6
0.1
0.1
0.1
9.9
5.5
0.1
5.4
4.2
1.6
0.2

82.0

56.0

20.0

4.0

26.0

18.0

2.0

2.0

2.0

6.0

2.0

2.0

2.0

6.0

16.0

14.0

32.0

2.0

2.0

2.0

2.0

6.0

6.0

4.0

4.0

2.0

2.0

2.0

2.0

2.0

22.0

16.0

2.0

14.0

6.0

2.0

2.0

Jeremy L. Marshall and Carlos D. Camp

and parasitic wasps (Scelionidae), were the second most important
group, making up 17% numerically and occurring in 32% of the
stomachs. Hymenopterans made up 15% of the relative area. Other
important insect groups represented were coleopterans, particularly
rove beetles (Staphylinidae), making up 8% of numbers, 9% of area,
and occurring in 19% of the stomachs, and delphacid homopterans
(7, 11, and 14%, respectively). Although found only occasionally,
relatively large roaches (Dictyoptera) and walking sticks (Phasmida)
made up a considerable amount of the total quantity of food eaten
(19 and 10% of total area, respectively). The only non-insect food
items found were arachnids. These consisted primarily of mites
(Acarina), which made up 9% numerically, 6% of the area, and occurr-
ed in 16% of stomachs.

Juvenile frogs from the 5 June, Grady County sample had a
mean SVL of 8.80 mm with a standard error (SE) of 0.21 mm. Adult
males from this sample had a mean SVL of 14.87 mm and a SE of
0.23 mm. Food items eaten by these frogs are shown in Table 2. The
Shannon-Wiener Index for juvenile prey species diversity (//' = 1.062)
was significantly larger than that for adults {H' = 0.739; t = 3.27, df
= 45, P < 0.01). There was a negative correlation between number
of food items eaten and frog size (r2 = 0.25, t = 2.18, df = 14, P <
0.05).

DISCUSSION

Pseudacris ocularis is commonly found on lower tree trunks and
foliage up to a height of 1 m or more (Harper 1939); males prefer
these sites as calling perches (Harper 1939, Mount 1975). However,
the majority of food items we found were arthropods that are associ-
ated with leaf litter and/or soil (e.g., springtails, mites, dipteran lar-
vae, staphylinids, ants, thrips, palpigrades, etc.). In addition, we found
a large number of frogs on the ground, particularly during daytime
collections. It is apparent, then, that P. ocularis spends a consider-
able amount of its foraging time on the ground.

According to optimal foraging theory (Pyke et al. 1977, Krebs
1978), a predator should choose prey that represent the greatest net
energy gain and forage in areas where profitable prey are most fre-
quently encountered. Considering the small size of these frogs, small
abundant leaf litter arthropods such as springtails and mites might
represent a relatively stable, predictable source of profitable prey.
However, amphibians might find larger arthropod prey to be more
profitable than small ones due to a proportionately smaller exoskel-
eton (Jaeger and Barnard 1981). Therefore, little grass frogs should

Little Grass Frog Feeding Ecology

Table 2. Stomach contents of 10 juvenile and 10 adult P. ocularis collected
5 June 1993, Grady County, Georgia; "i" represents immature instars;
* indicates two species represented. Numbers for higher taxa also include
unidentified food items.

r

Juveniles

Adults

Food Item

i %

n %

Collembola

9

27.3

10

45.5

Isotomidae

4

12.1

9

40.9

Sminthuridae

4

12.1

1

4.5

Coleoptera

2

6.1

2

9.1

Carabidae

1

3.0

0

0.0

Cleridae

1

3.0

0

0.0

Staphylinidae

0

0.0

2*

9.1

Diptera (larvae)

1

3.0

1

4.5

Homoptera

3

9.1

5

22.7

Delphacidae (i)

2

6.1

5

22.7

Hymenoptera

5

15.2

1

4.5

Formicidae

0

0.0

1

4.5

Scelionidae

4

12.1

0

0.0

Orthoptera

0

0.0

2

9.1

Acrididae (1)

2

9.1

Siphonaptera

1

3.0

0

0.0

Thysanoptera

1

3.0

0

0.0

Phlaeothripidae

1

3.0

Acarina

9

27.3

0

0.0

Meostigmata

1

3.0

Oribatei

8*

24.2

Araneida

1

3.0

1

4.5

Anyphaenidae

0

0.0

1

4.5

Palpigradi

1

3.0

0

0.0

feed more on larger prey when available. Our data would, in part,
appear to confirm this hypothesis. For instance, relatively large im-
mature delphacids made up < 7% of total food items. However, in the
5 June, Grady County sample of adults (Table 2), delphacids made up
23%, indicating these food items were probably more available at
that time, although we do not have independent confirmation of prey
abundance.

Newly metamorphosed P. ocularis ate more individual food
items and a greater diversity of prey species than did adult males.
Two factors may explain these results. First, there may be an ontogenetic
shift in foraging strategy during post-metamorphic growth of P. ocularis.
Such a shift has been inferred in P. triseriata (Christian 1982) where

Jeremy L. Marshall and Carlos D. Camp

adults select more optimal (i.e., large) prey than do juveniles, which
indiscriminately feed on prey they encounter. This may be the result
of larger animals being able to choose from a greater range of prey
sizes, whereas smaller individuals are largely restricted to small prey,
as is apparent in P. crucifer (Oplinger 1967). This shift would account
for the lower diversity of prey species taken by adult P. ocularis.
Second, the adult sample used in our comparisons consisted entirely
of males. Several authors have reported a sharp decline in feeding
activity by adult male frogs during the breeding season (Jenssen and
Klimstra 1966, Lamb 1984). The males in our study were not breeding
(the pond was completely dry) and only sporadically calling at the
time of collection (5 June), although breeding had been previously
observed at this site in March. Mount (1975), however, reported breeding
congregations of P. ocularis as late as 29 July in nearby Houston
County, Alabama, and Harper (1939) recorded vigorous chorusing in
the Okefenokee during August and September. Therefore, since P.
ocularis does breed throughout the summer, we cannot rule out the
possibility of lower feeding activity in adult males during the time of
our collections.

ACKNOWLEDGMENTS— Thanks are extended to Tyler Lee
(Cairo, Georgia) and Ben Cash (Georgia Southern University) for
help in collecting specimens, as well as Rob Wainberg (Piedmont
College) for assistance in identifying prey items. In addition, Rob
Wainberg and Rick Austin (University of Mississippi) critically read
the manuscript.

LITERATURE CITED
Brophy, T. E. 1980. Food habits of sympatric larval Ambystoma

tigrinum and Notophthalmus viridescens. Journal of Herpetology

14:1-6.
Camp, C. D., and L. L. Bozeman. 1981. Foods of two species of

Plethodon (Caudata: Plethodontidae) from Georgia and Alabama.

Brimleyana 6:163-166.
Christian, K. A. 1982. Changes in the food niche during post-

metamorphic ontogeny of the frog Pseudacris triseriata. Copeia

1982:73-80.
Conant, R., and J. T. Collins. 1991. Reptiles and amphibians of

eastern/central North America. Houghton-Mifflin, Boston, Massa-
chusetts.

Little Grass Frog Feeding Ecology

Davie, R. D. 1991. Ontogenetic shift in diet of Desmognathus quadramacula-
tus. Journal of Herpetology 25:108-111.

Duellman, W. E., and L. Trueb. 1986. Biology of amphibians. McGraw/
Hill, New York, New York.

Harper, F. 1939. Distribution, taxonomy, nomenclature and habits of
the little grass frog {Hyla ocularis). American Midland Naturalist
22:134-149.

Jaeger, R. G., and D. E. Barnard. 1981. Foraging tactics of a
terrestrial salamander: choice of diet in structurally simple envi-
ronments. American Naturalist 117:639-664.

Jenssen, T. A., and W. D. Klimstra. 1966. Food habits of the green
frog, Rana clamitans, in southern Illinois. American Midland Naturalist
76:169-182.

Krebs, J. R. 1978. Optimal foraging: decision rules for predators
Pages 23-63 in Behavioral ecology: an evolutionary approach
(J. R. Krebs and N. B. Davies, editors). Sinauer Publications,
Sunderland, Massachusetts.

Lamb, T. 1984. The influence of sex and breeding condition on
microhabitat selection and diet in the pig frog Rana grylio. American
Midland Naturalist 111:311-318.

Mount, R. H. 1975. The reptiles and amphibians of Alabama. Ala-
bama Agricultural Experiment Station, Auburn University, Auburn,
Alabama.

Oplinger, C. S. 1967. Food habits and feeding activity of recently
transformed and adult Hyla crucifer crucifer Wied. Herpetologica
23:209-217.

Pyke, G. H., H. R. Pulliam, and E. L. Charnov. 1977. Optimal
foraging: a selective review of theory and tests. Quarterly Re-
view of Biology 52:137-154.

Toft, C. A. 1980. Feeding ecology of thirteen syntopic species of
anurans in a seasonal tropical environment. Oecologia 45:131-141.

Wharton, C. H. 1978. The natural environments of Georgia. Georgia
Department of Natural Resources, Atlanta.

Zar, J. H. 1984. Biostatistical analysis, Second edition. Prentice-
Hall, Englewood Cliffs, New Jersey.

Received 28 June 1994
Accepted 10 November 1994

Digitized by the Internet Archive
in 2013

http://archive.org/details/brimleyana19nort_11

Corrections of Records of Occurrence

of Peromyscus polionotus (Wagner) and P. gossypinus

(LeConte) (Rodentia: Muridae) in the Blue Ridge

Province of Georgia

Joshua Laerm and James L. Boone

Museum of Natural History and Institute of Ecology

University of Georgia, Athens 30602

ABSTRACT — Reexamination of marginal records of Peromyscus
polionotus and P. gossypinus previously reported from the Blue
Ridge Province of Georgia indicate specimens were misidentified.
Neither species occurs in the Blue Ridge Province. The distribution
of P. polionotus is restricted to south of a line from Greenville
and Spartanburg counties, South Carolina, southeast to Clarke,
County, Georgia and west to Dawson and Cherokee counties,
Georgia. The distributional limit of P. gossypinus is south of
a line from Lincoln and Wilkes counties, Georgia west to Dekalb
and Fulton counties and then west and north to Polk, Floyd,
and Dade counties.

Mice of the genus Peromyscus Gloger are among the most
common and broadly studied small mammals in North America. Yet
they are often difficult to distinguish on the basis of traditional
morphological features, and consequently limits to their distribution
are difficult to delineate. This is particularly true within Osgood's
(1909) P. leucopus and P. maniculatus species groups (Hooper 1968,
Laerm and Boone 1994). Frequently biologists depend upon range
maps to rule out certain species; nevertheless, the correct identifica-
tion of a taxon should be based upon morphological characteristics.

Numerous regional studies have been undertaken to provide
mensural discrimination between species of Peromyscus (Choate 1973,
Linzey et al. 1976, Choate et al. 1979, Stromberg 1979, Engstrom et
al. 1982, Feldhamer et al. 1983, McDaniel et al. 1983). Laerm and
Boone (1994) recently used discriminant analysis to maximally
distinguish between the four Peromyscus species that occur in the
southeastern United States: P. gossypinus, leucopus, maniculatus, and
polionotus. Based upon reexamination and correct identification of
specimens representing marginal records of P. gossypinus and P.
polionotus with this discriminant analysis model, we questioned the
accuracy of existing range maps and marginal records for P.
polionotus (Wagner) and P. gossypinus (LeConte).

Brimleyana 22:9-14, June 1995

10 Joshua Laerm and James L. Boone

Peromyscus polionotus — A black-eyed, white Peromyscus was
reported from Tallulah Falls, Rabun County, Georgia by Dice (1934:246)
who identified the specimen as P. polionotus polionotus. This specimen
was noteworthy because the pelage was entirely white, and the feet,
toes, and nails lacked any pigment. However, as Dice noted, the eyes,
eye-ring, and outer parts of the ears were dark. Tallulah Falls, in the
Tallulah River Gorge, is at the southern edge of the Blue Ridge
Province in Georgia. While not specifically indicated by Dice, this
would have been the northernmost record of the species, from a
locality well outside the previously described range (Osgood 1909)
and in habitat from which the species had never been reported. Typically,
P. polionotus is restricted to sandy soils and does not occur north of
the middle Piedmont of Georgia.

Schwartz (1954) revised the Peromyscus polionotus complex and
described several new subspecies. He provided external and cranial
measurements of the six subspecies of polionotus he recognized and
referred populations of polionotus in the northern portion of Georgia
and South Carolina (essentially north of the Fall Line) to P. p. colemani.
Schwartz (1954:568) commented on the Dice specimen which he ". . .
presumed, on geographical grounds, to be assignable to P. p. colemani"
Had Schwartz actually examined the Dice specimen, he probably would
not have referred it to P. polionotus. However, Schwartz did not
include the specimen in his mensural analysis. Thus, following Dice
(1934) and Schwartz (1954), Hall and Kelson (1959), Golley (1962),
and Hall (1981) continued to include the specimen as a marginal
record for the species.

We questioned the identification of the Dice specimen because
(1) it was collected in quartzite sheer rock walls and talus of the
Tallulah River Gorge in the Blue Ridge Province and (2) at a locality
some 100 km north of any other specimen record. We compared our
measurements of the Dice specimen (University of Michigan Museum
of Zoology 68496) to measurements provided by Schwartz (1954) for
P. p. colemani and subjected it to our discriminant function (Laerm
and Boone 1994). For 8 of the 11 characters examined by Schwartz
(1954:565), the Dice specimen was larger than the range of the com-
parable measurements made on 11 P. p. colemani by Schwartz.

Visual comparison of means and ranges of measurements of 110
P. polionotus and 107 P. leucopus, which were used to develop a
discriminant function for mensural discrimination between these and
other southeastern Peromyscus spp. (measurements provided in Laerm
and Boone 1994), to the Dice specimen indicates that it generally
falls inside the range of measurements available for both polionotus

Occurrence of Peromyscus

11

and leucopus. Our discriminant analysis, however, strongly suggest the
Dice specimen to be P. leucopus (P = 0.939).

Concern over the correct identification of other northern
marginal range records of P. polionotus in Georgia, South Carolina,
and Alabama (see Schwartz 1954, Golley 1966, Wolfe and Rogers
1969, Hall 1981) prompted an examination of these specimens as
well. We are satisfied that specimens reported from Jackson County,
in extreme northeastern Alabama (Schwartz 1954, Hall and Kelson
1959, Hall 1981) are P. polionotus. Similarly, specimens from Greenville
and Spartanburg counties South Carolina referred to by Schwartz
(1954), Golley (1966), and Hall (1981) are correctly identified as P.
polionotus.

Review of currently available distributional records for P.
polionotus in Georgia now indicates the northernmost limit of its
range should be amended to extend from Spartanburg and Greenville
counties, South Carolina, southwest to Clarke County, Georgia, and
west to Dawson and Cherokee counties, Georgia. Unfortunately, there
are no records available for the Ridge and Valley or Cumberland
Plateau provinces of Georgia west of Dawson and Cherokee counties
(Fig. 1). The next records to the northwest are in Jackson County,
Alabama (Hall 1981).

Fig. 1 Distribution of Peromyscus gossypinus (a) and P. polionotus
(b) in Georgia. Dots represent northernmost distributional records only.

12 Joshua Laerm and James L. Boone

Peromyscus gossypinus — Hall and Kelson (1959), apparently
following Osgood (1909), indicated the range of P. gossypinus in
Georgia to be restricted to the Coastal Plain. At that time no speci-
mens were known from the Piedmont, Blue Ridge, Ridge and Valley,
or Cumberland Plateau provinces. Golley (1962) published new dis-
tributional records for Georgia and indicated (Golley 1962:124)
specimens occurring in the southeastern portion of the Piedmont
(including Columbia, Lincoln, McDuffie, and Wilkes counties),
Ridge and Valley (Floyd, Gordon, and Polk counties), Cumberland
Plateau (Dade County), extreme upper Piedmont (Habesham County)
as well as the Blue Ridge Province (Rabun County). Subsequently,
Wolfe and Linzey (1977) mapped the distribution of the species,
apparently following Hall and Kelson (1959). Wolfe and Linzey (1977)
do not cite Golley (1962); however, Hall (1981) does cite the Golley
(1962) records and maps them accordingly, indicating their range
extends into the extreme northeastern Piedmont and Blue Ridge
provinces.

We used our discriminant function to examine virtually all P.
gossypinus records from Georgia. Specimens from Dade County on
the Cumberland Plateau and those from Floyd and Polk counties in
the Ridge and Valley are P. gossypinus as noted by Golley (1962).
We were unable to locate any museum specimens referred to by
Golley (1962) from Habersham or Rabun counties either in University
of Georgia Museum of Natural History collections or those of all
other North American mammal collections housing specimens from
Georgia. Examination of skin tags and records has not indicated any
P. gossypinus from those counties to have been re-identified and/or
relabeled. We are confident that the specimens of P. gossypinus from
Ruban and Habersham counties in Golley (1962) were erroneously
mapped because, in his verbal description of the range, Golley (1962:128)
indicated P. gossypinus to be found "... on the coastal plain, but
extending into the Piedmont on the eastern margin of the state and
into the ridge and valley province on the west." He makes no mention
of any records in the extreme northern Piedmont or Blue Ridge. Thus,
the range of P. gossypinus in Georgia should be amended to extend
across the middle portion of the Piedmont from Lincoln and Wilkes
counties west to Dekalb and Fulton counties and then west and north
into Polk, Floyd, and Dade counties in the Ridge and Valley and
Cumberland Plateau (Fig. 1). This is essentially the range as previously
depicted by Hall and Kelson (1959) and Wolfe and Linzey (1977).

Occurrence of Peromyscus 13

LITERATURE CITED

Choate, J. R. 1973. Identification and recent distribution of white-
footed mice {Peromyscus) in New England. Journal of Mammalogy
54:41-49.

Choate, J. R., R. C. Dowler, and J. E. Krause. 1979. Mensural
discrimination between Peromyscus leucopus and P. maniculatus
(Rodentia) in Kansas. Southwestern Naturalist 24:249-258.

Dice, L. R. 1934. A black-eyed white Peromyscus. Journal of Mammalogy
15:246.

Engstrom, M. D., D. J. Schmidly, and P. K. Fox. 1982. Nongeographic
variation and discrimination of species within the Peromyscus leucopus
species group (Mammalia: Cricetinae) in eastern Texas. Texas Journal
of Science 34:149-162.

Feldhamer, G. A., J. E. Gates, and J. H. Howard. 1983. Field iden-
tification of Peromyscus maniculatus and P. leucopus in Maryland:
Reliability of morphological characters. Acta Theriologica 28:417-423.

Golley, F. B. 1962. Mammals of Georgia: A study of their distribution
and functional role in the ecosystem. University of Georgia Press,
Athens.

Golley, F. B. 1966. South Carolina mammals. The Charleston Museum,
Charleston, South Carolina.

Hall, E. R. 1981. The Mammals of North America. Second edition.
John Wiley and Sons, New York, New York.

Hall, E. R., and K. R. Kelson. 1959. The Mammals of North America.
Ronald Press, New York, New York.

Hooper, E. T. 1968. Classification. Pages 27-74 in Biology of
Peromyscus (Rodentia) (J. A. King, editor). Special Publication
of the American Society of Mammalogists 2:1-593.

Laerm, J., and J. L. Boone. 1994. Mensural discrimination of four
species of Peromyscus (Rodentia: Muridae) in the southeastern
United States. Brimleyana 21:107-124.

Linzey, A. V., D. W. Linzey, and S. E. Perkins, Jr. 1976. The
Peromyscus leucopus group in Alabama. Journal of the Alabama
Academy of Science 47:109-113.

McDaniel, V. R., R. Tumlison, and P. McLarty. 1983. Mensural
discrimination of the skulls of Arkansas Peromyscus. Proceedings
of the Arkansas Academy of Science 37:50-53.

Osgood, W. H. 1909. Revision of the mice of the American genus
Peromyscus. North American Fauna 28:1-285.

Schwartz, A. 1954. Oldfield mice, Peromyscus polionotus, of South
Carolina. Journal of Mammalogy 35:561-569.

Stromberg, M. R. 1979. Field identification of Peromyscus leucopus
and P. maniculatus with discriminant analysis. Wisconsin Academy
of Science, Art and Letters 67:159-164.

14 Joshua Laerm and James L. Boone

Wolfe, J. L., and A. V. Linzey. 1977. Peromyscus gossypinus. Mammalian

Species 70:1-5.
Wolfe, J. L., and D. T. Rogers. 1969. Oldfield mammals in western

Alabama. Journal of Mammalogy 50:609-612.

Received 25 July 1994
Accepted 28 November 1994

The Masked Shrew, Sorex cinereus

(Insectivora: Soricidae), and Red-backed Vole,

Clethrionomys gapperi (Rodentia: Muridae), in the

Blue Ridge Province of South Carolina

Joshua Laerm

Museum of Natural History and Institute of Ecology

University of Georgia, Athens, Georgia 30602

Eric Brown, Michael A. Menzel
Daniel B. Warnell School of Forest Resources
University of Georgia, Athens, Georgia 30602

Amanda Wotjalik

Museum of Natural History

University of Georgia, Athens, Georgia 30602

William Mark Ford
Daniel B. Warnell School of Forest Resources
University of Georgia, Athens, Georgia 30602

AND

Mary Strayer

Nongame and Heritage Trust Program,

South Carolina Wildlife and Marine Resources Department

P.O. Box 1806, Clemson, South Carolina 29633

ABSTRACT — The first records of Sorex cinereus Kerr are documented
from South Carolina. Pitfall surveys throughout the Blue Ridge
Province resulted in captures from two localities in markedly
mesic, relict, boreal habitats. Additional records of Clethironomys
gapperi (Vigors) were documented including the most southeastern
record. Both S. cinereus and C. gapperi are rare in South
Carolina, largely because of limited areas of appropriate habitats.

The masked shrew, Sorex cinereus Kerr, has the largest range
and exhibits the greatest geographic variation of any North American
Sorex (Hall 1981, Junge and Hoffmann 1981, van Zyll de Jong and
Kirkland 1989). It ranges throughout the transcontinental coniferous
forests from the Canadian Arctic south to the extreme northern por-
tions of the United States with significant extensions south into the

Brimleyana 22:15-21, June 1995 15

16 Joshua Laerm, et al.

montane forests of the Rocky and Appalachian mountains. In the
southeastern United States including Virginia and West Virginia, eastern
Kentucky and Tennessee, North Carolina and Georgia, S. cinereus is
restricted primarily to high elevation montane communities of the
Appalachian Highlands (van Zyll de Jong and Kirkland 1989). To
date, however, there have been no records from South Carolina
(Golley 1966, Mengak et al. 1987).

Previously, the southernmost records of S. cinereus have been
reported from Georgia based upon three specimens reported by
Wharton (1968) from Beech Creek near its confluence with the Talulla
River, Towns County, at an elevation of 807 m. More recently, Ford
et al. (In press) have reported S. cinereus from numerous, widely
scattered localities throughout the Blue Ridge Province of Georgia,
including localities in close proximity to the South Carolina state
line. Similarly, S. cinereus has been reported from several Blue Ridge
Province counties in North Carolina (Polk, Henderson, Transylvania,
Jackson, Macon, and Clay) which are contiguous to South Carolina
(Lee et al. 1982 and unpublished University of Georgia, Museum of
Natural History records). Because S. cinereus is known to occur in
immediately adjacent areas of Georgia and North Carolina and
because seemingly appropriate areas of high elevation habitat exist in
the Blue Ridge Province of South Carolina, we surveyed the moun-
tainous portions of Oconee, Pickens, and Greenville counties specifi-
cally for S. cinereus.

METHODS

From 23 January to 1 May 1994 pitfall trap surveys were
conducted throughout the Blue Ridge Province of extreme northwest-
ern South Carolina including, from east to west, portions of
Greenville, Pickens, and Oconee counties. We totalled 14,000 trap
nights at 17 individual sites. At each site twenty, 32-ounce, plastic
containers (14-cm lip diameter and 17-cm depth) were placed below
ground level adjacent to forest floor debris including stumps, fallen
logs, rocks, etc, for a minimum of 60 days. Approximately 0.14L of
preservative was placed in the bottom of each pitfall. General habitat
features, including dominant overstory and understory vegetation,
aspect, and approximate stand age, of each site were recorded and
elevations estimated from topographic maps. Traps were checked on
a biweekly basis. Specimens were preserved in alcohol for subsequent
reproductive and gut content analysis. Standard body measurements
were taken, and skulls were prepared for confirmation of identifica-

Masked Shrew and Red-backed Vole 17

tion. All specimens were reposited in the mammal collections of the
University of Georgia Museum of Natural History.

RESULTS AND DISCUSSION

We recovered 15 S. cinereus at two of 17 Blue Ridge Province
sites. Both S. cinereus localities were in the northwestern portion of
Oconee County. Seven individuals were recovered from the grounds
of the Walhala Fish Hatchery in a hemlock (Tsuga canadensis) and
rhododendron (Rhododendron maximum) streamside community which
grades upslope into a yellow poplar (Liriodendron tulipifera), mixed
oak (Quercus spp.), hickory (Carya spp.), and white pine (Pinus strobus)
community. Elevation was approximately 760 m. The second S.
cinereus locality (eight captures) was approximately 1.3 km east of
the Walhala Fish Hatchery site in a moderate to mesic mixed oak
and yellow poplar hardwood community at approximately 800 m.

Sorex cinereus was the dominant small mammal recovered in
the Walhala Fish Hatchery site. Fifteen small mammals were recov-
ered in 1,960 trap nights: seven S. cinereus, two S. fumeus, one Sorex
hoyi, one Blarina brevicauda, two Peromyscus maniculatus, and one
Clethrionomys gapperi. The recovery of S. cinereus was fairly evenly
distributed over the trapping period with one or two captured during
each sampling period.

At the second site, also with 1,960 trap nights, 12 S. fumeus,
four S. hoyi, two Peromyscus leucopus, one P. maniculatus, two Blarina
brevicauda, and one Clethrionomys gapperi were captured in addition
to the eight S. cinereus. Here all the cinereus were captured between
20 March and 3 April; six of which were taken in a single pitfall trap
beneath a large, heavily rotted log.

The breadth and intensity of our collection efforts indicate a
restricted distribution of S. cinereus in South Carolina. Sorex cinereus
is regarded as having a boreomontane distribution (Junge and
Hoffman 1981). In the southern Appalachians it has been documented
by Odum (1949), Johnston (1967), Gentry et al. (1968), Linzey and
Linzey (1971), Whitaker et al. (1975), and Lee et al. (1982) in west-
ern North Carolina; Conaway and Howell (1953), Smith et al. (1974),
and Harvey et al. (1991, 1992), in the mountainous regions of eastern
Tennessee; and Pagels and Tate (1976), Pagels and Handley (1989),
Pagels (1991), Kalko and Handley (1993), and Pagels et al. (1994) in
southwestern Virginia. It has not been recorded from elevations be-
low 610 m in southwestern Virginia (Pagels and Handley 1989) or
North Carolina (Linzey and Linzey 1971, Lee et al. 1982). Similarly,

18 Joshua Laerm, et al.

in Georgia, S. cinereus is restricted to high elevation (790-1,370 m)
in markedly mesic habitats with northern affinities (Ford et al. 1994).

Kirkland (1985, 1991) indicated that soricids, in general, are
most diverse in regions characterized by cool moist forests, possibly
by supporting an abundant, stable, and diverse soil invertebrate fauna
upon which shrews depend. Pagels et al. (1994) have shown that the
presence of S. cinereus was significantly correlated with soil moisture
holding capacity and total understory vegetation, and that habitat features
that promote shaded, moist habitats were particularly important in
relict, boreal forest habitats throughout the southern Appalachians.

Although considerable areas of the Blue Ridge Province in South
Carolina meet or exceed the minimum elevations at which S. cinereus
is found elsewhere in the southern Appalachians, boreomontane
habitats are limited there. At the southern limit of the Appalachian
Mountains, much of the mountain habitat in South Carolina is charac-
terized by south-facing aspects with more xeric, mixed oak and pine
communities. Similar xeric south-facing or ridgeline habitats in Geor-
gia yielded few, if any, S. cinereus in recent studies (Ford et al.
1994). In Georgia we encountered S. cinereus primarily at very high
elevations (over 1200 m) or in rich, moist, streamside communities
dominated by hemlock and rhododendron on the Rabun Bald Massif.
West of the Little Tennessee River in Georgia, S. cinereus is
restricted to higher (over 1000 m) elevations, and then they only
occur in restricted habitats with marked northern affinities such as
those described by Wharton (1968).

Our collection site at the Walhala Fish Hatchery is located in
a relatively narrow, steep-walled gorge of the East Fork of the Chat-
tooga River. Wharton (1977) noted that similar streamside forest
communities on the Georgia side of the Chattooga were kept cool and
moist due to complete shading by the hemlock overstory and
rhododendron shrub layer as well as by steep-walled gorges. He noted
that such areas were refugia of more typical northern forest commu-
nities. The Walhala Fish Hatchery, and its associated upslope north-
ern aspect cove hardwood habitat, might represent a limited finger
or refugia in South Carolina for boreal species such as S. cinereus.
The region of the Walhala Fish Hatchery is one of the few sites in
South Carolina that has yielded other small mammals with a typical
boreal distribution including Clethrionomys gapperi (Pivorun et al.
1984) and Peromyscus maniculatus. Other high elevation sites in the
Blue Ridge Province including Sassafrass Mountain, Jones Gap at
Caesar's Head State Park, Saluda Mountain and Hogback Mountain
were trapped but yielded no S. cinereus. Peromyscus maniculatus has

Masked Shrew and Red-backed Vole 19

been recorded at many of these sites (Golley 1966), and we recovered
several specimens at most of these localities. However, Clethriono-
mys gapperi was not reported beyond the Walhalla Fish Hatchery
site until we recovered one in the region of Sassafrass Mountain
(Pickens County, US Hwy 178, 7.4 m north of State Hwy 11). This is
most southeastern record for the species and, like S. cinereus, it
apparently has a very limited distribution in South Carolina.

ACKNOWLEDGMENTS— We appreciate the field assistance of
numerous persons including E. Mitchell, J. Kimbrell, N. Lautenschlager,
B. Smith, L. Grassman, and S. Chalmers. Collecting activities were
conducted under South Carolina Permit #0016-94. We also thank
biologists with the Walhala Fish Hatchery, Sumpter National Forest,
and Caesar's Head State Park for permission to conduct surveys on
United States Forest Service and South Carolina State Park lands.

LITERATURE CITED

Conaway, C. H., and J. C. Howell. 1953. Observations on the mammals
of Johnston and Carter counties, Tennessee and Avery County,
Virginia. Journal of the Tennessee Academy of Science 28:53-61.

Ford, W. M., J. Laerm, D. C. Weinand, and K. G. Barker. 1994.
Small mammal surveys in the Chatahoochee National Forest of
Georgia. Proceedings of the Annual Conference of the Southeast-
ern Association of Fish and Wildlife Agencies (In press).

Gentry, J. B., E. P. Odum, M. Mason, V. Nabholtz, S. Marshall, and
J. T. McGinnis. 1968. The effect of altitude and forest manipulation
on relative abundance of small mammals. Journal of Mammalogy
49:539-541.

Golley, F. B. 1966. South Carolina mammals. Contributions from
the Charleston Museum, Charleston, South Carolina.

Hall, E. R. 1981. The mammals of North America. Second edition.
John Wiley & Sons, New York, New York.

Harvey, M. J., C. S. Chaney, and M. D. McGimsey. 1991. Distribution,
status, and ecology of small mammals of the Cherokee National
Forest, Tennessee (Southern Districts). Unpublished manuscript. Center
for the Management, Utilization, and Protection of Water Resources,
Tennessee Technological University, Cookeville.

Harvey, M. J., M. D. McGimsey, and C. S. Chaney. 1992. Distribution,
status, and ecology of small mammals of the Cherokee National
Forest, Tennessee (Northern Districts). Unpublished manuscript. Center
for the Management, Utilization, and Protection of Water Resources,
Tennessee Technological University, Cookeville.

20 Joshua Laerm, et al.

Johnston, D. W. 1967. Ecology and distribution of mammals at
Highlands, North Carolina. Journal of the Elisha Mitchell Scientific
Society 83:88-98.

Junge, J. A., and R. S. Hoffmann. 1981. An annotated key to the
long-tailed shrews (genus Sorex) of the United States and Canada,
with notes on middle American Sorex. Occasional Papers of the
Museum of Natural History, University of Kansas 94:1-48.

Kalko, E. K. V., and C. O. Handley, Jr. 1993. Comparative studies
of small mammal populations with transects of snap traps and
pitfall arrays in southwest Virginia. Virginia Journal of Science
44:3-18.

Kirkland, G. L., Jr. 1985. Small mammal communities in temperate
North American forests. Australian Mammalogy 8:137-144.

Kirkland, G. L., Jr. 1991. Competition and coexistence in shrews
(Insectivora: Soricidae). Pages 15-22 in The biology of the Soricidae
(J. S. Findley and T. L. Yates, editors). The Museum of Southwestern
Biology, Albuquerque, New Mexico.

Kirkland, G. L., Jr., and P. K. Sheppard. 1994. Proposed standard
protocol for pitfall sampling of small mammal communities. Pages
277-281 in Advances in the biology of shrews (J. F. Merritt, G.
L. Kirkland, Jr., and R. K. Rose, editors). Special Publication,
Carnegie Museum of Natural History 18:1-485.

Lee, D. S., J. B. Funderburg, and M. K. Clark. 1982. A distribu-
tional survey of North Carolina mammals. Occasional Papers of
the North Carolina Biological Survey, Raleigh.

Linzey, A. V., and D. W. Linzey. 1971. Mammals of the Great
Smoky Mountains National Park. University of Tennessee Press,
Knoxville.

Mengak, M. T., D. C. Guynn, Jr., J. K. Edwards, D. L. Sanders, and
S. M. Miller. 1987. Abundance and distribution of shrews in
western South Carolina. Brimelyana 13:63-66.

Odum, E. P. 1949. Small mammals of the Highlands (North Caro-
lina) Plateau. Journal of Mammalogy 30:179-192.

Pagels, J. F. 1987. The pygmy shrew, rock shrew and water shrew:
Virginia's rarest shrews (Mammalia: Soricidae). Virginia Journal
of Science 38:364-368.

Pagels, J. F. 1991. A high elevation record for the least shrew,
Cryptotis parva (Say). Virginia Journal of Science 42:361-362.

Pagels, J. F., and C. O. Handley, Jr. 1989. Distribution of the
southeastern shrew, Sorex longirostris Bachman, in western Vir-
ginia. Brimleyana 15:123-131.

Pagels, J. F., and C. M. Tate. 1976. Shrews (Insectivora: Soricidae)
of the Paddy Knob-Little Back Creek Area of western Virginia.
Virginia Journal of Science 27:202-203.

Masked Shrew and Red-backed Vole 21

Pagels, J. F., K. L. Uthus, and H. E. Duval. 1994. The masked
shrew, Sorex cinereus, in a relictual habitat of the southern Ap-
palachians. Pages 103-109 in Advances in the biology of shrews
(J. F. Merritt, G. L. Kirkland, Jr., and R. K. Rose, editors).
Special publication, Carnegie Museum of Natural History 18:1—
485.

Pivorun, E. B., D. H. Allen, and D. T. Sawyer. 1984. First record
of Clethrionomys gapperi (Mammalia: Rodentia) in South Caro-
lina. Journal of the Elisha Mitchell Society 100:33.

Smith, C. R., J. Giles, and M. E. Richmond. 1976. The mammals
of northeastern Tennessee. Journal of the Tennessee Academy of
Science 49:88-94.

van Zyll de Jong, C. G., and G. L. Kirkland, Jr. 1989. A morpho-
metric analysis of the Sorex cinereus group in central and east-
ern North America. Journal of Mammalogy 70:110-122.

Wharton, C. H. 1968. First records of Microsorex hoyi and Sorex
cinereus from Georgia. Journal of Mammalogy 49:158.

Wharton, C. H. 1977. The natural environments of Georgia. Georgia
Department of Natural Resources, Office of Planning and Re-
sources, Atlanta.

Whitaker, J. O., Jr., G. S. Jones, and D. D. Pascal. 1975. Notes on
mammals of the Fires Creek Area, Nantahala Mountains, North
Carolina, including their parasites. Journal of the Elisha Mitchell
Scientific Society 91:13-17.

Received 25 July 1994
Accepted 30 November 1994

Rediscovery of the Aquatic Gastropod

Helisoma eucosmium (Bartsch, 1908),

(Basommatophora: Planorbidae)

William F. Adams

Environmental Resources Branch

United States Army Corps of Engineers

P.O. Box 1890, Wilmington, North Carolina 28402

AND

Susan G. Brady

Department of Biology

University of North Carolina at Wilmington

601 South College Road, Wilmington, North Carolina 28403

ABSTRACT — A population of Helisoma eucosmium (Bartsch, 1908),
a small freshwater planorbid snail considered to be extinct by
some authors, has been discovered in Town Creek, a tidal
swamp stream in southeastern North Carolina. This rediscovery
will permit a definitive determination of the proper systematic
placement of the taxon. In the absence of live specimens, past
analysis of this taxon during systematic revisions of the Planorbidae
relied only on shell morphology of type material. Under this
circumstance, this taxon was variously placed in both tropical
and temperate genera and, compounding that problem, was treated
as a full species by some authors and a subspecies by others.
Its apparently limited range elevates concern for the conserva-
tion of planorbid snail diversity in southeastern North Carolina
because it is the second taxon with a severely restricted dis-
tribution to be found in this rapidly urbanizing region.

Helisoma eucosmium (Bartsch, 1908) is a small, distinctive planorbid
snail which was collected and described from Greenfield Lake, a
millpond constructed prior to 1750 (Adams 1990a). This site is located
within the City of Wilmington, New Hanover County, North Carolina,
and is on a tributary to the lower Cape Fear River. During the past
few decades, repeated attempts to recollect H. eucosmium in Greenfield
Lake and elsewhere within the region have been unsuccessful (Fuller
1977, Adams 1990a). This fact, combined with the water quality degradation
in many regional streams, has led many investigators to treat the
taxon as extinct (Opler 1976, Imlay 1977, Palmer 1985) or possibly
extinct (Fuller 1977, Adams 19906).

Brimleyana 22:23-29, June 1995 23

24 William F. Adams and Susan G. Brady

During the spring of 1994, while performing a survey of the
mollusks of Town Creek, a tidal swamp stream tributary to the lower
Cape Fear River in adjacent Brunswick County, SGB discovered a
population of Helisoma eucosmium approximately 14.5 km (9 mi)
SSW of the type locality (Fig. 1). Although much of this stream
system remains to be investigated, the taxon is known to occupy at
least a 1.75 km (1.1 mi) stretch of the main creek. Five specimens
were deposited in the invertebrate research collection at the North
Carolina State Museum of Natural Sciences, Raleigh (NCSM). All
conform precisely to the original shell description given by Bartsch
(1908), particularly in displaying the chestnut-colored bands (Figs. 2
and 3). The largest specimen (NCSM #P1207), collected on 30 April
1994, has a greater diameter of 6.0 mm, a lesser diameter of 4.75
mm, and a height of 3.0 mm. It was probably an adult that overwintered,
a likelihood suggested by its size and the minor pitting and corrosion
of the shell surface. Three specimens taken in early July 1994 (NCSM
#P1208) are smaller; average dimensions of three specimens are maximum
diameter 3.8 mm, minimum diameter 3.0 mm, and height 1.9 mm
(n = 3). Similarly-sized specimens taken from the same period laid
eggs in captivity indicating that sexual maturity had been attained.

The molluscan community in the freshwater part of Town Creek
is diverse; species encountered to date are listed in Table 1. Continuing
surveys may disclose additional species and most of those historically
documented (Adams 1990a) from the type locality of Helisoma eucosmium
may ultimately be found.

In his description, Bartsch (1908) assigned Helisoma eucosmium
and a subspecies from Louisiana, H. eucosmium vaughani, to the
genus Planorbis Miiller 1774. F. C. Baker (1931) restricted that genus
to species of European origin and, without examination of soft tissue
anatomy, provisionally placed eucosmium in the genus Helisoma Swainson
1840. In his subsequent monograph on the Planorbidae, Baker (1945)
retained this assignment, placing it alone with Helisoma anceps (Menke,
1830) in the subgenus Helisoma s.s.; however, he made no reference
to anatomical examinations supporting this placement. On the strength
of unpublished observations by J. P. E. Morrison of live individuals
of H. e. vaughani from Louisiana, Fuller (1977) tentatively assigned
the species to the Central American genus Taphius H. & A. Adams
1855. Taphius and other planorbid genera of tropical affinity (Afroplanorbis
Thiele 1931, Biomphalaria Preston 1910, Australorbis Pilsbry 1934,
Tropicorbis Pilsbry and Brown 1914, Planorbina Haldeman 1842,
Armigerus Clessin 1884, and Platytaphius Pilsbry 1924) have long
been the subjects of taxonomic debate and divergent systematic treatment,

Helisoma eucosmium

25

Table 1. Freshwater mollusks of upper Town Creek, Brunswick County,
North Carolina, 1994.

Gastropods

Bivalves

Viviparidae

Campeloma decision (Say)
Hydrobiidae

Amnicola limosus (Say)

Gillia altilis (Lea)
Physidae

Physella hendersoni (Clench)
Planorbidae

Micromenetus dilitatus (Gould)

Helisoma eucosmium (Bartsch)

Planorbella trivolvis (Say)
Ancylidae

Laevapex fuscus (C. B. Adams)

Unionidae

Pyganodon cataracta (Say)
Villosa delumbis (Conrad)
Ligumia nasuta (Say)
Unknown taxa of the
Elliptio complanata and
Elliptio icterina complexes

Corbiculidae

Corbicula fluminea (Muller)

Sphaeriidae

Eupera cubensis (Prime)
Musculium securis (Prime)
Unidentified sphaeriids

a problem generated and sustained by years of inconclusive taxonomic
studies resulting from the great similarity in chonchology and anatomy
existing within a genus complex of worldwide distribution (Hubendick
1955, H. B. Baker 1960). While taxonomic and systematic problems
are common in the freshwater gastropods, the need for nomenclatural
stability within the tropical planorbids is a matter of great medical
and economic importance because these snails are intermediate hosts
of the human parasite Schistosoma mansoni.

In response to a petition by Wright (1962), the International
Commission on Zoological Nomenclature (1965) issued Opinion 735,
ruling that Biomphalaria is to be given precedence over the generic
names Taphius, Planorbina, and Armigera when any or all of these
names are considered to apply to the same genus. Under this ruling,
Fuller's (1977) binomen would be valid only if Taphius is determined
to be separable from Biomphalaria at the genus level. Bypassing this
issue, Burch (1982, 1989) stated Helisoma eucosmium may be a "form
or juvenile" of Helisoma anceps (Menke, 1930) and, therefore, did
not grant it specific status. Based on shell morphology and the reproductive
maturity of the material collected from Town Creek, we find no evidence
to support this possibility. Although the very oblique aperture of H.
eucosmium is suggestive of the Biomphalaria group, the lack of vertically
depressed whorls is not; therefore, we provisionally accept Baker's

26

William F. Adams and Susan G. Brady

WILMINGTON

Fig. 1. Town Creek and vicinity (* = known range of Helisoma eucosmium,
♦ = known sites of Planorbella magnifica.

(1945) designation pending definitive studies of soft tissue anatomy.

Over much of its length, Town Creek is a tidal system approximately
30-50 m wide with maximum depths varying between 5-7 m. Whereas
the lower reaches are brackish, no salinity has been detected in the
area where Helisoma eucosmium occurs. Physical parameters of the
water are pH 5.1-7.0 (* = 6.1, n = 3), conductivity 217 //mhos/cm
(n = 2), and calcium concentration 107.9 ppm (n = 2). All specimens
of H. eucosmium came from a littoral community, occurring in less
than 3 m of water, consisting of dense continuous mats of Brazilian
elodea (Egeria densa Planchon), fanwort {Cabomba caroliniana Gray),
fragrant waterlily (Nymphaea odorata Aiton), spatterdock (Nuphar
luteum (Linnaeus)), and floating-heart (Nymphoides aquatica (Walter)).

Helisoma euc osmium

27

Fig. 2. Dorsal, ventral, and apertural
views of Helisoma eucosmium
(NCSM #P1208). Lip aperture is
incomplete.

Sis

Kh D

mm--- vwi

MHWj^^l^^^P^^^^^^'" "&•■''*'■■&'

1

1

>

mm ^

Fig. 3. Foraging Helisoma eucosmium.

28 William F. Adams and Susan G. Brady

Streambank forest cover, typical of southern swamps, consists of bald
cypress (Taxodium distichum (Linnaeus)), tupelo gum (Nyssa aquatica
Linnaeus), red maple {Acer rubrum Linnaeus), and water ash (Fraxinus
caroliniana Miller).

Our report of Helisoma eucosmium constitutes the second recent
recollection of a likely "extinct" planorbid snail in the Cape Fear
River drainage of southeastern North Carolina, the other being the
rediscovery of Planorbella magnifica (Pilsbry) in Orton Pond (Adams
1988) and in Sandhill Creek Pond (Adams 1993), approximately 14.5
km (9.0 mi) SE and 12 km (7.5 mi) ESE, respectively (Fig. 1). Because
of the heavy residential and industrial development that has occurred
in southeastern North Carolina within the past half-century, these
waterbodies might hold some of the few remaining populations of the
original freshwater molluscan fauna of the lower Cape Fear Basin. If
H. eucosmium is restricted to Town Creek and systematic research
determines that it warrants full species rank, protection under state
and federal conservation laws might be warranted. Although the Town
Creek watershed is still very rural, covered primarily in crop and
forest lands, the surrounding region is urbanizing rapidly, and the
stream will surely be impacted by this trend. Consequently, additional
surveys to determine the distribution of H. eucosmium and resolution
of outstanding systematic issues are urgently needed.

ACKNOWLEDGMENTS— Partial funding for this work has been
provided by the University of North Carolina Center for Marine Science
Research and the Wilmington District, United States Army Corps of
Engineers. We would like to thank Dr. Eric Bolen for his administrative
support, Mr. Shaun Cain for field assistance, Dr. Courtney Hackney
for his ideas and encouragement, and Dr. Rowland Shelley for his
constructive review of this manuscript. Photographs were taken by
Mr. David M. DuMond.

LITERATURE CITED

Adams, W. F. 1988. Rediscovery of Planorbella magnifica (Pilsbry)
in southeastern North Carolina. Nautilus 102(3): 125-126.

Adams, W. F. 1990a. Recent changes in the freshwater molluscan
fauna of the Greenfield Lake basin, North Carolina. Brimleyana
16:103-117.

Adams, W. F. 1990fc. Helisoma eucosmium (Bartsch, 1908). Pages
16-17 in A report on the conservation status of North Carolina's
freshwater and terrestrial molluscan fauna (W. F. Adams, editor).
North Carolina Wildlife Resources Commission, Raleigh.

Helisoma eucosmium 29

Adams, W. F. 1993. A status survey for the magnificent ramshorn
(Planorbella magnified), a freshwater snail endemic to North Carolina.
North Carolina Wildlife Resources Commission, Raleigh.

Baker, F. C. 1931. The classification of the large planorbid snails
of Europe and America. Proceedings of the Zoological Society of
London 1931(2):575-592.

Baker, F. C. 1945. The molluscan family Planorbidae. University of
Illinois Press, Urbana.

Baker, H. B. 1960. Planorbina (1843) vs. Australorbis (1934) vs.
Biomphalaria (1910) vs. Taphius (1854). Nautilus 74(l):35-37.

Bartsch, P. 1908. Notes on the fresh-water mollusk Planorbis magnificus
and descriptions of two new forms of the same genus from the
southern states. Proceedings of the U. S. National Museum 33:697-
700.

Burch, J. B. 1982. Freshwater snails (Mollusca: Gastropoda) of North
America. Environmental Protection Agency, EPA-600/3-82-026.

Burch, J. B. 1989. North American freshwater snails. Malacological
Publications, Hamburg, Michigan.

Fuller, S. L. H. 1977. Freshwater and terrestrial mollusks. Pages
143-194 in Endangered and threatened plants and animals of North
Carolina (J. E. Cooper, S. S. Robinson, and J. B. Funderburg,
editors). North Carolina State Museum of Natural History, Ra-
leigh.

Hubendick, B. 1955. Phylogeny in the Planorbidae. Transactions of
the Zoological Society of London 28(6):453-542.

Imlay, M. J. 1977. Competing for survival. Water Spectrum 9:7-14.

International Commission on Zoological Nomenclature. 1965. Opinion
735. Bulletin of Zoological Nomenclature 22(2):94-99.

Opler, P. A. 1976. The parade of passing species: a survey of
extinctions in the U.S. Science Teacher 43(9):30-34.

Palmer, S. 1985. Some extinct molluscs of the U.S.A. Atala 13(1): 1-7.

Wright, C. A. 1962. Planorbina Haldeman, 1852, Taphius Adams
and Adams, 1955 and Armigerus Clessin, 1884 (Mollusca, Gastropoda):
Proposed Suppression under the Plenary Powers. Z.N.(S.) 1392.
Bulletin of Zoological Nomenclature 19(1):39-41.

Received 27 September 1994
Accepted 19 January 1995

Effects of a Clearcut on the Herpetofauna of a
South Carolina Bottomland Swamp

Joseph P. Phelps1

North Carolina State University

Hardwood Research Cooperative

Raleigh, North Carolina 27695-8008

AND

Richard A. Lancia

Department of Forestry

North Carolina State University

Raleigh, North Carolina 27695-8002

ABSTRACT — Amphibians and reptiles were trapped during summer
in a South Carolina bottomland swamp to assess the impacts
of clearcut timber harvesting. Animals were captured using drift
fences with pitfall traps, coverboards, and polyvinyl chloride
pipes which simulated treefrog habitat. Twenty-nine species (10
amphibians and 19 reptiles) were detected on the site. Some
were captured frequently enough to infer microhabitat preferences.
Salamanders were much more frequent in the control area than
in the clearcut. Other species showing preferences for the control
were bronze frogs (Rana clamitans), gray treefrogs (Hyla chrysoscelis),
and box turtles (Terrapene Carolina). Reptiles generally preferred
the clearcut. This was especially true of lizards and large snakes.
Diversities showed no significant differences between the control
and clearcut. Small clearcuts done on long rotations are recommended.
Machinery impact should be kept to a minimum, and down
wood and snags should be left on the site.

Bottomland hardwood forests have been recognized for their
importance in floodwater and sediment retention, water quality protection,
timber production, and wildlife habitat (Brinson et al. 1981, Clark
and Benforado 1981, Harris and Gosselink 1986). At the same time,
these ecosystems are being lost and degraded rapidly (Turner et al.
1981, Rudis 1993) due in large part to fragmentation. In the South-
east, 75,000 acres of forested wetlands have been lost since 1982
(Cubbage and Flather 1993), not including acreage that was logged
and regenerated. These logged wetlands may temporarily lose some
functional value, and they contribute to fragmentation.

1 Present address: Weyerhaeuser Company, Southern Forest Research, P.O. Box 1060,
Hot Springs, Arkansas 71902.

Brimleyana 22:31-45, June 1995 31

32 J. P. Phelps and R. A. Lancia

The research results presented here were obtained concurrent
with other studies (Pavel 1993, Perison In Press) designed to document
the impact of timber harvesting on the functional value of a bottom-
land swamp. Amphibians and reptiles (herpetofauna) were chosen as
the appropriate wildlife groups to study because of their abundance in
the Southeast (Keister 1971, Vickers et al. 1985, Hairston 1987) and
because of their importance in food chains (Pough et al. 1987, Blaustein
and Wake 1990). Herpetofaunal species are also influenced by factors
that are affected by timber harvest, including hydrology, soil quality,
and vegetative structure. Herpetofaunal communities have been shown
to be altered by clearcutting (Enge and Marion 1986, Pough et al.

1987, Petranka et al. 1993), ditching of wetlands (Vickers et al. 1985,
Enge and Marion 1986), and changing forest cover (Bennett et al.
1980, Pough et al. 1987).

In addition, much attention has been paid to a possible world-
wide decline in amphibian diversity (Blaustein and Wake 1990, Pechmann
et al. 1991, Hairston and Wiley 1993). Logging has been identified
among the many possible causes of such a decline (Wake 1991, Hairston
and Wiley 1993). Amphibians may be a good indicator of general
environmental degradation, due to their exposure to terrestrial and
aquatic toxins, and their sensitivity to habitat changes (Beiswenger

1988, Blaustein and Wake 1990).

The objectives of our research were to evaluate whether clear-
cutting in a hardwood swamp had any effect on community diversity
or abundance of summer-active amphibians and reptiles. We also attempted
to identify habitat variables that may have been related to changes in
the herpetofaunal community.

METHODS

Study Site

The study site was on the South Fork of the Edisto River,
near Norway (Orangeburg County), South Carolina. "The site is re-
presentative of blackwater swamps in the Carolinas that have timber
management potential" (Perison et al. 1993). Predominant trees in the
swamp forest included tupelo gum (Nyssa spp.), sweetgum (Liquidambar
styraciflua), willow/water oak (Quercus phellos/nigra), and green ash
(Fraxinus pennsylvanica) (Pavel 1993). A clearcut of approximately
10 ha was completed in January 1991. Much of the clearcut area was
impacted by skidder tire ruts. The adjacent control area was upstream
of the clearcut and was a second growth stand approximately 45 years
old. Second growth forest surrounded the clearcut. The sampled area
of control was comparable to the size of the clearcut. The edge sampled
was 650 m long.

Clearcut Herpetofauna 33

Herpetofauna

Sampling — Amphibians and reptiles were captured using three
types of traps: drift fences, coverboards, and polyvinyl chloride (PVC)
pipe. Nine arrays of drift fences, measuring 270 m in total, were
constructed of aluminum flashing (Gibbons and Semlitsch 1981).
Each array consisted of two 15-m lengths placed at right angles to
one another. Three of the arrays were placed in the clearcut, three in
the control, and three along the edge between the clearcut and control.
Twelve pitfall traps (19-L plastic paint buckets) were placed along
each array, for a total of 108 traps. Arrays were centered in the
clearcut, about 75 m from the edge. Control arrays averaged about
75 m from the edge.

Forty-five coverboards were systematically placed across the study
site, 30 in the clearcut and 15 in the control. Each coverboard con-
sisted of a piece of plywood or particle-board (about 120 x 60 x 0.625
cm) placed flat on the ground to simulate the type of cover often used
by ground-dwelling herpetofauna (DeGraff and Yamasaki 1992, Fitch

1992, Mitchell et al. 1993). Coverboards were placed a minimum of
20 m apart.

Sixty 1.5-m lengths of PVC pipe were driven into the soil to
capture treefrogs. The diameters of the pipes were 2.5 and 5 cm.
Forty pipes were used in the clearcut, and 20 in the control. These
pipes served as refuges for treefrogs, which were easily captured at
the open top ends of the pipes. One nocturnal chorus survey (July
1993) was done to compare to habitat use trends indicated by the pipe
captures. Frog choruses were monitored from the clearcut, control,
and edge for 30 minutes each. Weather was humid, warm, clear, and
calm.

Each of the traps was checked daily during the summers of 1992
and 1993. All captured animals were released immediately. Adult
anurans were toe-clipped, but no capture-recapture data will be pre-
sented here (see below). In 1992, we trapped from 29 May until 13
August, with the exception of 14 days when the swamp was flooded.
On those days, only the treefrog pipes were checked. In 1993, four
drift fence arrays (one control, two edge, one clearcut) were checked
from 20 May to 12 August. The remaining traps were checked from
20 June to 12 August. Again, sampling was impeded due to flooding.

Sampling was replicated within the control, edge, and clearcut
site, but not replicated with additional sites because of constraints
imposed by the establishment of concurrent studies, which used small
replicated treatment blocks within the clearcut (Pavel 1993, Phelps

1993, Perison In Press). These blocks were the reason for unequal

34 J. P. Phelps and R. A. Lancia

sampling effort for coverboards and PVC pipes in the clearcut, edge,
and control. Due to the lack of replication, the results and conclusions
from this study apply to the particular site we studied. Generalization
to other sites is risky.

Data analysis — Capture data were analyzed using the Shannon-
Weaver Diversity Index (Poole 1974). This allowed statistical com-
parisons among drift fence arrays in the clearcut, on the edge, and in
the control; and between coverboards in the clearcut and in the control.
Variance for each treatment was calculated based on the number of
captures in each treatment, and was used to calculate the ^-statistic (a
= 0.05) for each comparison (Poole 1974). The formulas used are as
follows:

Var (//') = [rMPl ln2p,] - [g.lf>, lnp,]2 + s-1

N 2N2

H\-H\

[Var(/f ,) + Var(//'2)]1/2

d.f. = [Var(/f ,) + Var(/r2)]2
[VarCtf',)]2 + [Var(tf'2)]:

N, N2

where i represents each species in the sample, pi = proportion of
species i in the sample, s = number of species in the sample, N =
number of captures in the sample.

Assumptions of the Shannon-Weaver Index were that all species
present were sampled, and all were equally catchable. These assumptions
were not met because of the differing ability of the traps to catch
various species (Gibbons and Semlitsch 1981). Also, two species caught
by hand were never captured in a trap of any kind and were therefore
excluded from calculations of H' . These were the brown water snake
(Nerodia taxispilota) and the timber rattlesnake (Crotalus horridus).
Their size explains the lack of pitfall captures, as no larger snakes
were caught in pitfalls. Other large snakes were captured under
coverboards.

Poole (1974) states that "no great error" will occur in calculating
Shannon-Weaver as if all the species available are present in the sample,
even when, as in this case, some species are not represented. Unequal
catchability of species should still allow relative diversity comparisons.

Clearcut Herpetofauna 35

The lack of replication allowed only a rough comparison of
capture frequencies, not a statistical test of differences in abundance
(Hurlbert 1984). The original intent was to estimate abundances of
anurans using mark-recapture methods, but a lack of sufficient recapture
prevented this. To make direct comparisons of capture frequencies,
we assumed that capture probabilities for each species were the same
across habitats. This assumption could not be tested, so habitat
preferences should be interpreted with caution.

Habitat Characteristics

Sampling — In an attempt to relate herpetofaunal diversity to
microhabitat variability, the percent cover of midstory and overstory
trees was measured with a spherical densiometer at five points around
each drift fence array. One reading was taken in the center and one at
a distance of 15 m in each cardinal direction. Percent cover of veg-
etation less than 2 m tall was also measured, using a line intercept
method (Barbour et al. 1987). There were four 15-m transects at each
drift fence array, originating at the middle of the array and extending
in each cardinal direction. A complete description of the vegetation
of the site is given by Pavel (1993).

The cross-sectional area of coarse woody debris was measured
using the same transects. In this case, the transects were thought of
as vertical planes that extended from the ground to the highest piece
of downed wood. Each piece of wood greater than 8-cm diameter (at
the point of intersection) was measured to the nearest centimeter and
classified as "sound" or "rotten" (Brown 1974).

Surface soil temperatures and soil densities were measured by
Perison (In Press) in the control and clearcut areas. All habitat data
were analyzed using Mests at the 0.05 alpha level. Each measurement
was considered to be independent, and means and standard errors
were calculated for each variable. In the clearcut, soil temperature
and density were measured in areas with skidder traffic, but not in
skidder ruts. These variables were not measured at the edge.

RESULTS

Herpetofauna

Shannon-Weaver Diversity Indices calculated for the drift fence
and coverboard captures are given in Table 1. There was no significant
difference between clearcut and control diversity indices for either of
the two trapping methods. The diversity of the edge drift fences was
lower than either the control (t = 8.70, d.f. = 1,538) or clearcut drift
fences (t = 9.70, d.f. = 2,209). The reason for this difference was the

36

J. P. Phelps and R. A. Lancia

Table 1. Summary of herpetofauna capture data, Edisto River swamp,
South Carolina, 1992-1993. H' is the value of the Shannon-Weaver
diversity index. Values of H' with the same letter were not different
at 5% alpha level.

Captures Species
(N) (s)

H'

Control drift fences
Clearcut drift fences
Edge drift fences
All drift fences
Control coverboards
Clearcut coverboards
All coverboards

930

16

0.9161a

1,172

16

0.9109a

2,647

17

0.4738b

4,749

23

0.7153c

150

9

1.7077d

129

16

1.7710e

279

19

1.9832e

heavy weighting toward one species, the southern toad {Bufo terrestris),
on the edge. Clearcut and control drift fence diversities were not
different at the 0.05 alpha level. Coverboard diversities were higher
than drift fence diversities (f = 18.5, d.f. = 333). Drift fences captured
more species (23) and individuals (4,749) than coverboards (19 and
279, respectively). The inclusion of the clearcut increased the richness
of the capture sample. Twenty-three species were captured on the site
in drift fences, but only 16 species were captured in the clearcut, and
16 in the control. Nineteen species were captured under coverboards,
but only 16 were captured in the clearcut, and nine in the control.

Several species showed clear preferences for either the control
or the clearcut (Table 2). All types of salamanders were detected
more often in the control area. A total of 112 salamanders was captured
in pitfall traps and under coverboards. Ninety-two (82.1%) of these
were captured in the control area, with only 9 (8.0%) in the clearcut
(Table 2). Bronze frogs {Rana clamitans) were captured more frequently
in the control (104 control captures to 17 in the clearcut), while
southern leopard frogs (R. utricularia) were less common in the control
(33 drift fence captures compared to 52 on the edge and 51 in the
clearcut). Other frog species showing preferences were eastern narrow-
mouth toads {Gastrophryne carolinensis) and green treefrogs (Hyla
cinerea), which preferred the clearcut, and gray treefrogs (H. chrysoscelis),
which preferred the control. Southern toads, most of which were
juveniles, were captured most frequently in the edge pitfalls. Southern
toads were abundant in all three areas. The data on frog and toad
species were supported by the breeding chorus monitoring.

Clearcut Herpetofauna 37

Reptiles generally seemed to prefer the clearcut. Eastern mud
turtles (Kinosternon subrubrum) were captured 41 times in the clearcut,
35 times on the edge, and only 10 times in the control area. Common
musk turtles {Sternotherus odoratus) and eastern box turtles (Terrapene
Carolina) showed the opposite trend. Lizards (Eumeces fasciatus and
Anolis carolinensis) and large snakes (chiefly Agkistrodon piscivorus
and Nerodia spp.) were more common in the clearcut. Because they
were rarely captured in traps but often seen, habitat preferences of
large snakes, lizards, and box turtles are largely inferred from hand
captures, rather than trapping data.

Habitat Characteristics

Microhabitat variables measured in each area are given in Table
3. Over- and mid-story canopy cover was highest in the control (95%),
followed by the edge (74%), and clearcut (6%). Understory canopy
cover followed the reverse trend, being highest in the clearcut (95%),
followed by the edge (48%), and the control (5%).

Cross-sectional area of sound coarse woody debris was significantly
higher in the clearcut, and not significantly different between the
edge and control. The mount of rotten coarse woody debris was not
different among the three areas. Soil surface temperature and soil
compaction were not measured on the edge, but were not significantly
different between the clearcut and the control, or between rutted and
non-rutted areas in the clearcut.

DISCUSSION

Habitat Preferences

Fewer salamanders were captured in the clearcut, as compared
to the control. High temperatures and insolation, and low relative
humidity may all have contributed to this, as these factors increase
the risk of desiccation. Salamanders need to keep their skin moist for
gas exchange, and their large surface areas to volume ratios make
moisture retention difficult (Duellman and Trueb 1986). Moisture is a
key factor in determining where salamanders can live (Wyman 1988,
Petranka et al. 1993). Petranka et al. (1993) speculated that 75% to
80% of southern Appalachian salamanders die of physiological stress
due to desiccation following clearcuts.

Ecological differences may explain the varying preferences of
frog species. Southern toads are generalists, and their common
occurrence along the edge suggests that individuals were using both
habitats. Bronze frogs and southern leopard frogs showed opposing
preferences, for the control and clearcut respectively. Bronze frogs

38

J. P. Phelps and R. A. Lancia

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40 J. P. Phelps and R. A. Lancia

are a forest wetland species, whereas leopard frogs are more ubiquitous
(Mount 1975). Enge and Marion (1986) found fewer southern leopard
frogs in clearcuts than in control areas in southern pine flatwoods.

Eastern narrowmouth toads emerge from burrows only when ground
water is sufficiently pooled for breeding. Reduced evapotranspiration
in the clearcut (Perison et al. 1993) may have increased the likelihood
of these frogs receiving the cue to emerge. Skidder tire ruts may have
had a similar effect on these and the other frog species, as they were
a source of fish-free breeding pools (Phelps 1993).

Two species of treefrogs were captured, and these showed opposing
preferences. Green treefrogs may have predominated in the clearcut
because the traps (treefrog pipes) were level with their habitat (leafy
sprouting vegetation). Green treefrogs may have been equally abundant
in the control (as suggested by the number of control captures, Table
2), but may have been living higher than the traps were sampling.
Gray treefrogs were rarely captured, and the reason again may have
been that the traps were too low. The removal of trees in the clearcut
may have excluded gray treefrogs because their habitat (tree boles)
was removed. In the case of the green treefrogs in the clearcut, their
habitat quickly returned, but at a lower height. Overall, the treefrog
pipe method showed promise, as it was inexpensive and trapped
significant numbers of frogs while causing them no apparent stress.

Reptiles may have preferred the clearcut because of increased
temperature and insolation. A basking reptile would be at an advantage
in the direct sun of the clearcut because it could achieve active temperatures
faster and remain active longer through the day. Bury and Corn (1988),
working in the Pacific Northwest, found that "reptiles predominate in
clearcuts, most likely responding to increased ambient tempera-
tures in such areas." The presence of a large amount of slash in the
clearcut may have provided a valuable habitat component in the form
of cover for reptiles or their prey. Reptiles that seemed to prefer the
control included ringneck snakes (Diadophis punctatus), copperheads
(Agkistrodon contortix), common musk turtles, and eastern box turtles.
The reasons for these preferences are not clear, although ringneck
snakes were the smallest and most fossorial reptile species regularly
captured, and may have been less dependent on basking and more
dependent on an undisturbed forest floor.

Recommendations and Conclusions

Clearcut size is one factor to be considered when planning for
natural forest regeneration in forested wetlands. Amphibians and reptiles
have relatively small home range sizes (Duellman and Trueb 1986)
and, therefore, cannot disperse from or quickly recolonize impacted

Clearcut Herpetofauna 41

areas. Hairston (1987) found that salamanders return to their home
range even after disturbance and handling. Small clearcuts with undisturbed
sources of recolonization nearby were advocated by Buhlmann et al.
(1988) and Enge and Marion (1986). A mosaic of small clearcuts,
second growth, and undisturbed areas would likely create increased
landscape diversity, as compared to a single homogeneous stand.

Recovery times of about 60 years for salamander populations in
clearcut areas are given by Petranka et al. (1993) in the southern
Appalachian and Pough et al. (1987) in New York. This suggests that
long rotation times are needed to avoid a long-term decline of sala-
mander populations over several rotations. Enge and Marion (1986)
also recommended long rotation times that allow adequate recovery
of herpetofaunal populations.

Controlling certain aspects of the harvest operation can minimize
the adverse effects of a clearcut. Most importantly, snags and coarse
woody debris should be left on the site (Enge and Marion 1986).
Woody debris should be of large size and in an advanced state of
decay (Bury and Corn 1988, Welsh and Lind 1988). Aubry et al.
(1988) suggested that "the abundance levels of salamanders are more
likely a function of the availability of woody debris for cover than
age of the overstory." Since suitable woody debris is more abundant
in older stands, longer rotation times are important. Leaf litter on the
forest floor is another important component of herpetofaunal habitat
(Pough et al. 1987, DeGraaf and Rudis 1990, Petranka et al. 1993),
and can be destroyed by ground machinery such as skidders (Buhlmann
et al. 1988). Skidders should be restricted to small areas, and helicopters
should be used when practical. Buhlmann et al. (1988) recommended
harvesting in the season of inactivity for the local herpetofauna, but
some southern Coastal Plain species are active at all times of the
year.

Further studies in which capture probabilities can be estimated
would allow direct comparison of capture data among species in the
same habitat, and within species across habitats. Remaining research
opportunities include determining the fate of salamanders in the face
of clearcutting, and monitoring subsequent recovery of populations
through recolonization. Also, a study similar to ours, focusing on
winter-active amphibians, would be valuable. In addition, the habitat
value of skidder ruts should be studied. The possible benefit of extra
standing water (Phelps 1993) may mitigate the effects of soil degradation
(Buhlmann et al. 1988). Finally, additional work with PVC pipes for
capturing treefrogs should be done, including their use in various
habitats, with different species, and the possible effect of pipe height
on trap efficiency.

42 J. P. Phelps and R. A. Lancia

The key to expanding knowledge in the area of wildlife habitat/
forestry relationships is to replicate treatments. In this case, conclusions
could have been strengthened by having several clearcuts and several
control areas (Hurlbert 1984). Specific factors such as woody debris,
size of clearcuts, and skidder rut impact could be studied with such a
design. Ideally, each area would be sampled prior to the installation
of the clearcut (Buhlmann et al. 1988). This would allow comparison
of data from the clearcut before and after treatment, and from the
control. Effects of space, time, and the treatment itself could be separated.
Replication and the collection of baseline data could be achieved
more easily within the framework of Adaptive Resource Management
(Walters 1986). This is a system of research integrated with management,
wherein management decisions are treated as hypotheses and tested
with replicated trials. After several iterations of hypothesis and experiment,
predictions involved with policy can become prescriptions based on
hard data.

ACKNOWLEDGMENTS— funding for this project was provided
by the Hardwood Research Cooperative of North Carolina State University.
Harvest treatments were done by Georgia-Pacific Corporation, and
housing for researchers was provided by the South Carolina Forestry
Commission. Dr. Robert Kellison of the Hardwood Cooperative provided
valuable support and a review of the manuscript. Two anonymous
reviewers provided valuable comments.

LITERATURE CITED

Aubry, K. B., L. L. C. Jones, and P. A. Hall. 1988. Use of woody
debris by plethodontid salamanders in Douglas-fir forests in
Washington. Pages 32-37 in Management of amphibians, reptiles
and small mammals in North America (R. C. Szaro, K. E. Severson,
and D. R. Patton, editors). United States Department of Agricul-
ture Forest Service, Technical Report RM166.

Barbour, M. G., J. H. Furke, and W. D. Pitts. 1987. Terrestrial
plant ecology (second edition). The Benjamin/Cummings Publish-
ing Co., Inc., Menlo Park, California.

Beiswenger, R. E. 1988. Integrating anuran amphibian species into
environmental assessment programs. Pages 159-165 in Manage-
ment of amphiabians, reptiles and small mammals in North America
(R. C. Szaro, K. E. Severson, and D. R. Patton, editors). United
States Department of Agriculture Forest Service, Technical Report
RM166.

Clearcut Herpetofauna 43

Bennett, S. H., J. W. Gibbons, and J. Glanville. 1980. Terrestrial
activity, abundance, and diversity of amphibians in differently managed
forest types. American Midland Naturalist 103:412-416.

Blaustein, A. R., and D. B. Wake. 1990. Declining amphibian popu-
lations: a global phenomenon? Trends in Ecological Evolution 5:203-
204.

Brinson, M., B. Swift, R. Plantico, and J. Barclay. 1981. Riparian
ecosystems: their ecology and status. United States Department
of Interior Fish and Wildlife Service. FWS/OBS-81/17.

Brown, J. K. 1974. Handbook for inventoring downed woody mate-
rial. United States Department of Agriculture Forest Service, Technical
Report INT16.

Buhlmann, K. A., C. A. Payne, J. C. Mitchell, and R. B. Glasgow.
1988. Forestry operations and terrestrial salamanders: techniques
in a study of the Cow Knob salamanders, Plethodon punctatus.
Pages 38^4 in Management of amphibians, reptiles and small
mammals in North America (R. C. Szaro, K. E. Severson, and
D. R. Patton, editors). United States Department of Agriculture
Forest Service, Technical Report RM166.

Bury, R. B., and P. S. Corn. 1988. Douglas-fir forests in the
Oregon and Washington Cascades: relation of the herpetofauna to
stand age and moisture. Pages 11-22 in Management of amphib-
ians, reptiles and small mammals in North America (R. C. Szaro,
K. E. Severson, and D. R. Patton, editors). United States Depart-
ment of Agriculture Forest Service, Technical Report RM166.

Clark, J. R., and J. Benforado (editors). 1981. Wetlands of bottom-
land hardwood forests. Elsevier Publishing Company, Amsterdam,
Netherlands.

Cubbage, F. W., and C. H. Flather. 1993. Distribution of and trends
in southern forested wetlands. Page 13-19 in Proceedings of the
seventh biennial southern silvicultural research conference (J. C.
Brissett, editor). United States Department of Agriculture Forest
Service, General Technical Report SO-93.

DeGraaf, R. M., and D. D. Rudis. 1990. Herpetofaunal species com-
position and relative abundance among three New England forest
types. Forest Ecology and Management 32:155-165.

DeGraaf, R. M., and M. Yamasaki. 1992. A nondestructive tech-
nique to monitor the relative abundance of terrestrial salamanders.
Wildlife Society Bulletin 20:260-264.

Duellman, W. E., and L. Trueb. 1986. Biology of amphibians. McGraw-
Hill, Inc., New York, New York.

Enge, K. M., and W. R. Marion. 1986. Effects of clearcutting and
site preparation on herpetofauna of a north Florida flatwoods.
Forest Ecology and Management 14:177-192.

Fitch, H. S. 1992. Methods of sampling snake populations and their
relative success. Herpetological Review 23:17-19.

44 J. P. Phelps and R. A. Lancia

Gibbons, J. W., and R. D. Semlitsch. 1981. Terrestrial drift fences
with pitfall traps: an effective technique for quantitative sampling
of animal populations. Brimleyana 7:1-16.

Hairston, N. G. 1987. Community ecology and salamander guilds.
Cambridge University Press. Cambridge, England.

Hairston, N. G., and R. H. Wiley. 1993. No decline in salamander
(Amphibia: Caudata) populations: a twenty year study in the southern
Appalachians. Brimleyana 18:59-64.

Harris, L., and J. G. Gosselink. 1986. Cumulative impacts of bot-
tomland hardwood conversion on hydrology, water quality, and
terrestrial wildlife. United States Environmental Protection Agency.
Washington, D.C. Unpublished.

Hurlbert, S. H. 1984. Pseudoreplication and the design of ecological
field experiments. Ecological Monographs 54:187-211.

Keister, A. R. 1971. Species density of North American amphibians
and reptiles. Systematic Zoology 20:127-137.

Mitchell, J. C, S. Y. Erdle, and J. F. Pagels. 1993. Evaluation of
capture techniques for amphibian, reptile and small mammal communities
in saturated forested wetlands. Wetlands 13:130-136.

Mount, R. H. 1975. The reptiles and amphibians of Alabama. Au-
burn University Agricultural Experiment Station, Alabama.

Pavel, C. M. 1993. An assessment of timber harvest on the biomass,
species diversity, and stand structure of the vegetation in a South
Carolina bottomland hardwood forest. M.S. Thesis. North Caro-
lina State University, Raleigh.

Pechmann, J. H. K., D. E. Scott, R. D. Semlitsch, J. P. Caldwell, L.
J. Vitt, and J. W. Gibbons. 1991. Declining amphibian popula-
tions: the problem of separating human impacts from natural fluctuations
Science 253:892-895.

Perison, D. M. In Press. The effects of timber harvest and soil
disturbance on soil processes and water quality in a South Caro-
lina blackwater swamp. Ph.D. Thesis. North Carolina State Uni-
versity, Raleigh.

Perison, D. M., R. Lea, and R. Kellison. 1993. The response of
soil physical and chemical properties and water quality to timber
harvest and soil disturbance: preliminary results. Page 143-146 in
Proceedings of the seventh biennial southern silvicultural research
conference (J. C. Bissette, editor). United States Department of
Agriculture Forest Service, Technical Report SO-93.

Petranka, J. W., M. E. Eldridge, and K. E. Haley. 1993. Effects of
timber harvesting on southern Appalachians salamanders. Conser-
vation Biology 7:363-370.

Phelps, J. P. 1993. The effect of clearcutting on the herpetofauna
of a South Carolina blackwater bottomland. M.S. Thesis. North
Carolina State University, Raleigh.

Clearcut Herpetofauna 45

Poole, R. W. 1974. An introduction to quantitative ecology. McGraw-
Hill, Inc., New York, New York.

Pough, F. H., E. M. Smith, D. H. Rhodes, and A. Collazo. 1987.
The abundance of salamanders in forest stands with different his-
tories of disturbance. Forest Ecology and Management 20:1-9.

Rudis, V. A. 1993. Forest fragmentation of southern U.S. bottom-
land hardwoods. Page 35-46 in Proceedings of the seventh bien-
nial southern silvicultural research conference (J. C. Brissett, editor).
United States Department of Agriculture Forest Service, Technical
Report SO-93.

Turner, R. E., S. Forsythe, and N. Craig. 1981. Bottomand hard-
wood forest land resources of the southeastern U.S. Pages 13-28
in Wetlands of bottomland hardwood forests (J. R. Clark and J.
Benforado, editors). Elsevier Publishing Company, Amsterdam, Netherlands.

Vickers, C. R., L. D. Harris, and B. F. Swindel. 1985. Changes in
herpetofauna resulting from ditching of cypress ponds in coastal
plains flatwoods. Forest Ecology and Management 11:17-29.

Wake, D. B. 1991. Declining amphibian populations. Science 253:860.

Walters, C. J. 1986. Adaptive management of renewable resources.
Macmillian, New York, New York.

Welsh, H. H., Jr., and A. J. Lind. 1988. Old-growth forests and the
distribution of terrestrial herpetofauna. Pages 439-458 in Manage-
ment of amphibians, reptiles and small mammals in North America
(R. C. Szaro, K. E. Severson, and D. R. Patton, editors). United
States Department of Agriculture Forest Service, Technical Report
RM166.

Wyman, R. L. 1988. Soil acidity and moisture and the distribution
of amphibians in five forests of southcentral New York. Copeia
1988:394-399.

Received 16 November 1994
Accepted 19 January 1995

First Record of the Water Shrew, Sorex palustris

Richardson (Insectivora: Soricidae), in Georgia

with Comments on its Distribution and Status

in the Southern Appalachians

Joshua Laerm, Charles H. Wharton
Museum of Natural History and Institute of Ecology

AND

William Mark Ford
Daniel B. Warnell School of Forest Resources
University of Georgia, Athens, Georgia 30602

ABSTRACT— The first state record of Sorex palustris is
reported from Georgia, in a markedly boreal habitat in the
upper headwaters of the Tallulah River in Towns County. Records
in the southern Appalachians indicate the species to be rare
and its distribution characterized by a series of disjunct
populations.

On 30 May 1994 one adult male water shrew (Sorex palustris
Richardson) was recovered from a sunken pitfall trap adjacent to
Mare Cove Branch at its junction with Burnt Cabin Branch, a tributary
of the Tallulah River in extreme northern portion of Towns County,
Georgia, at an elevation of 808 m. The specimen was recovered under
a rotting log and other woody debris immediately adjacent to the base
of a 25 m waterfall in rocky talus. Standard body measurements were:
138-64-19. This is the first record of the species from Georgia and
represents an extension of its range approximately 25 km southeast
from its nearest reported locality along a short section of Fires Creek
in Clay County, North Carolina, at an elevation of 1,160 m (Whitaker
et al. 1975)

During the period 30 October 1993 through 30 May 1994 we
established a transect of 20 pitfall traps (a total of 5,420 trap nights)
along Burnt Cabin Branch. Pitfalls were 946 cm3 plastic cups
(11-cm lip diameter and 14-cm depth) filled with approximately
0.13-L formalin solution and set flush to the ground adjacent to
fallen logs, rocks stumps, or other forest floor debris within 5 m of
the stream edge. Traps were checked biweekly. The collection locality,
which was selectively logged in the past, is a mature, predominantly
northern hardwood forest community dominated by yellow birch

Brimleyana 22:47-51, June 1995 47

48 Joshua Laerm, C. H. Wharton, and W. M. Ford

(Betula luted), black birch (B. lenta), liden {J ilia heterophylla), and
hemlock (Tsuga canadensis), with a rhododendron (Rhododendron
maximum) understory.

The mammalian fauna of Burnt Cabin Branch has marked boreal
affinities. Wharton (1968) reported the first Georgia records of the
masked shrew (Sorex cinereus) and pygmy shrew (S. hoyi) from Beech
Creek, another tributary of the Tallulah River within 500 m of the
present locality, and Laerm (1992) reported the first Georgia record
of the hairy-tailed mole (Parascalops brewerii) from the present locality.
Other small mammals recovered in pitfalls and snap traps at Burnt
Cabin Branch include Sorex cinereus, S. fumeus, Blarina brevicauda,
Tamiasciurus hudsonicus, Peromyscus maniculatus, Napaeozapus
insignis, and Clethrionomys gapperi.

Sorex palustris is distributed in the transcontinental Canadian
boreal forest from Nova Scotia westward to southeastern Alaska and
southward throughout much of the Sierra Nevada and Rocky
Mountains in the western United States as well as the Appalachian
Mountains to Tennessee, North Carolina (Hall 1981, Beneski and
Stinson 1987), and now Georgia in the eastern United States. Populations
throughout the Appalachian Mountains from southwestern Pennsylvania
to Georgia are referable to S. palustris punctulatus Hooper 1942, the
West Virginia water shrew. Based on available published sources,
museum records, and personal communications it appears that S. p.
punctulatus is rare and its distribution characterized by a series of
apparently disjunct populations.

The northernmost record for S. p. punctulatus is a single specimen
from Cove Run in the Negro Mountains, Somerset County, Pennsyl-
vania (Doutt et al. 1966, Enders 1985). More recently, two additional
specimens have been obtained from Somerset County (C. Bier and
S. McLaren, personal communication; specimens in Carnegie Museum
of Natural History). Apparently, the distribution of this subspecies is
disjunct from that of S. p. albibarbis which is reported from central
and northeastern Pennsylvania and northward (Hall 1981, Beneski
and Stinson 1987, Merritt 1987).

Mansuetti (1958), Paradiso (1969), and Feldhamer et al. (1984)
discussed the questionable occurrence of S. palustris in Maryland.
However, seven individuals of S. p. punctulatus are now known from
seven sites in Maryland, all from Garrett County (E. Thompson,
Maryland Natural Heritage Program, personal communication). At least
12 individuals are known from five counties (Pendleton, Pocahantas,
Preston, Randolph, and Tucker) in West Virginia (Kellogg 1937,
Hooper 1942, McKeever 1952, and records on file with West Virginia

Water Shrew 49

Heritage Inventory Program). Three individuals are known from a
single locality in Bath County, Virginia, and five from three localities
in Highlands County, Virginia (Pagels and Tate 1976, Pagels 1987,
Handley 1991, and J. Pagels, personal communication). Thirteen
records are known from the Great Smoky Mountains National Park,
Sevier County, Tennessee (Conaway and Pfitzer 1952, Linzey and
Linzey 1968), and Harvey et al. (1991) report an additional 18 speci-
mens from four localities in Monroe County, Tennessee. In North
Carolina it is known from five individuals from Clay County and one
specimen from Great Smoky Mountains National Park in Swain
County (Whitaker et al. 1975, Linzey 1983, Webster 1987).

Compared to other soricids in the southeastern United States,
Sorex palustris appears to be rare. Sorex p. punctulatus is considered
a Category 2 taxon by the United States Fish and Wildlife Service, is
listed as endangered in Virginia (Virginia Department of Game and
Inland Fisheries), and is considered a species of special concern
in North Carolina (North Carolina Natural Heritage Program) and
Tennessee (Tennessee Wildlife Resource Agency). Although other
soricids, such a Sorex hoyi and Sorex dispar, that historically have
been considered extremely rare are now known to be more widely
distributed and more common than previously believed (Pagels 1987,
Handley 1991, Laerm et al 1994), the water shrew appears to be the
rarest and most localized shrew in the southeastern United States.
Additional surveys for the water shrew are required to assess its
true status.

ACKNOWLEDGMENTS— This study was supported through a
cooperative funding agreement between the United States Forest
Service, Chatahoochee National Forest, and The University of
Georgia Museum of Natural History.

LITERATURE CITED

Beneski, J. T., Jr., and D. W. Stinson. 1987. Sorex palustris. Mam-
malian Species 296:1-6.

Conaway, C. S., and D. W. Pfitzer. 1952. Sorex palustris and Sorex
dispar from the Great Smoky Mountains National Park. Journal
of Mammalogy 33:106-108.

Doutt, J. K., C. A. Heppenstall, and J. E. Guilday. 1966. Mammals
of Pennsylvania. The Pennsylvania Game Commission, Harrisburg.

50 Joshua Laerm, C. H. Wharton, and W. M. Ford

Enders, J. E. 1985. Water shrew Sorex palustris punctulatus (Hooper).
Pages 383-385 in Species of special concern in Pennsylvania (H.
H. Genoways and F. J. Brenner, editors). Special Publication of
the Carnegie Museum of Natural History, Number 11, Pittsburgh,
Pennsylvania.

Feldhamer, G. A., J. E. Gates, and J. A. Chapman. 1984. Rare,
threatened and extirpated mammals from Maryland. Pages 395-438
in Threatened and endangered plants and animals of Maryland
(A. W. Norden, D. C. Forester, and G. H. Fenwick, editors).
Maryland Natural Heritage Program Special Publication 84-1.

Hall, E. R. 1981. The mammals of North American. Second edition.
John Wiley & Sons, New York, New York.

Handley, C. O., Jr. 1991. Mammals. Pages 539-613 in Virginia's
endangered species (K. Terwilliger, editor). McDonald and Woodward,
Blacksburg, Virginia.

Harvey, M. J., C. S. Chaney, and M. D. McGimsey. 1991. Distribu-
tion, status, and ecology of small mammals of the Cherokee Na-
tional Forest, Tennessee (Southern Districts). Report to the United
States Forest Service. Manuscript on file, Center for the Manage-
ment, Utilization, and Protection of Water Resources, Tennessee
Technological University, Cookville.

Hooper, E. T. 1942. The water shrew {Sorex palustris) of the south-
ern Allegheny Mountains. Occasional Papers of the Museum Zo-
ology, University of Michigan 463:1-4.

Kellogg, R. 1937. Annotated list of West Virginia mammals. Pro-
ceedings of the United States National Museum 84:443-479.

Laerm, J. 1992. Georgia's rarest mammal. Georgia Wildlife 2(2):46-53.

Laerm, J., W. M. Ford, and D. W. Weinand. 1994. Additional
records of the pygmy shrew, Sorex hoyi winnemana Preble (In-
sectivora: Soricidae), in western North Carolina. Brimleyana 21:91-
96.

Linzey, D. W. 1983. Status and distribution of the northern water
shrew {Sorex palustris) and two subspecies of northern flying
squirrel {Glaucomys sabrinus coloratus and Glaucomys sabrinus
fuscus). Final report under United States Fish and Wildlife Ser-
vice Contract No. 14-16-005-79-068.

Linzey, D. W., and A. V. Linzey. 1968. Mammals of Great Smoky
Mountains National Park. Journal of the Elisha Mitchell Scien-
tific Society 84:384-414.

Mansueti, R. 1958. The Craneville Pine Swamp. Atlantic Naturalist
13:72-84.

McKeever, S. 1952. A survey of West Virginia mammals. West
Virginia Conservation Commission, Pitman-Roberstson Project 22-
R. Unpublished report on file, United States National Museum of
Natural History, Washington, D.C.

Water Shrew 51

Pagels, J. F. 1987. The pygmy shrew, rock shrew, and water shrew:

Virginia's rarest shrews (Mammalia: Soricidae). Virginia Journal

of Science 38:364-368.
Pagels, J. F., and C. M. Tate. 1986. Shrews (Insectivora: Soricidae)

of the Paddy Knob-Little Back Creek Area of western Virginia.

Virginia Journal of Science 27:202-203.
Paradiso, J. L. 1969. Mammals of Maryland. North American Fauna

66:1-193.
Webster, W. D. 1987. Sorex palustris punctulatus. Pages 36-38 in

Endangered, threatened and rare fauna of North Carolina. Part 1.

A reevaluation of the mammals (M. K. Clark, editor). Occasional

Papers of the North Carolina Biological Survey, Raleigh.
Wharton, C. H. 1968. First records of Microsorex hoyi and Sorex

cinereus from Georgia. Journal of Mammalogy 49:158.
Whitaker, J. O., Jr., G. S. Jones, and D. D. Pascal, Jr. 1975. Notes

on the mammals of the Fires Creek Area, Nantahala Mountains,

North Carolina, including their ectoparasites. Journal of the Elisha

Mitchell Scientific Society 91:13-17.

Received 25 July 1994
Accepted 26 January 1995

Florida Manatees, Trichechus manatus

(Sirenia: Trichechidae),

in North Carolina 1919-1994

Frank J. Schwartz

Institute of Marine Sciences

University of North Carolina

Morehead City, North Carolina 28557

ABSTRACT— Florida manatees, first reported in 1919 from North
Carolina, are now known to have frequented 59 sites (68 indi-
viduals) during the period of 1919-1994. All but two have
been subadults of about 1.8-2.4-m lengths. Only seven deaths
have been recorded. Eleven coastal counties have harbored manatees.
Four occurrences have been at inlets and six in the open ocean.
Pelletier Creek, a Carteret County tributary of Bogue Sound,
along with the Atlantic Ocean have been the most frequented
sites (6); eight manatees occurred at a lush vegetation site in
the Trent River (Craven County), a tributary of the Neuse
River. Four records came from Wrightsville Beach and Sound,
three manatees entered the state from Chesapeake Bay via the
canal and Intracoastal Waterway into Currituck Sound. Farthest
inland river penetrations have been 94.4 km, 6.4 km north of
Wilmington; 92 km, Neuse River at Fort Barnwell Bridge, 33
km northeast of New Bern (Craven County); and one each
penetrated the Tar River at Washington (58 km, Beaufort County)
and Greenville (88 km, Pitt County). The increased frequency
of occurrences in later years may be the result of an in-
creased public awareness of the federally-protected species rather
than a seemingly increasing population.

The Florida manatee, Trichechus manatus, can attain a size of
4.1 m, 1,620 kg, and ranges from Maryland (Chester River, Chesapeake
Bay) to Louisiana in the northern Gulf of Mexico (C. Beck, National
Biological Service, personal communication; Jefferson et al. 1993).
Northernmost manatee records in North Carolina have been in
Currituck Sound (Dare County) (Brimley 1905, 1946; Brimley 1931;
Caldwell and Golley 1965) (Table 1, Fig. 1). My study summarizes
early records, adds 44 new records (total 59) and comments on where,
when, and what size manatees have occurred in North Carolina.

Brimleyana 22:53-60, June 1995 53

54

Frank J. Schwartz

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57

Fig. 1. Locations for 68 manatee sightings in North Carolina during
the years 1919-1994.

EARLY LITERATURE OBSERVATIONS
Fifteen live and three dead manatees were reported (16 sites)
from North Carolina between 1919 and 1976 by Brimley (1931, 1946),
Lee (1976), Rathbun et al. (1981), and Smith et al. (1960). The specimen
from Currituck Sound near Duck (Brimley 1946) was about 3 m long.
None was weighed, although a Wrightsville Sound specimen and
the Currituck specimens were estimated to weigh 450 kg (Brimley
1931, 1946). Clark (National Marine Fisheries Service, personal
communication) recorded a live manatee near Cape Lookout 25 June
1975.

58 Frank J. Schwartz

RECENT NORTH CAROLINA OBSERVATIONS

Manatees are now known (68 individuals from 59 sites) to fre-
quent nearly all North Carolina ocean and inland waters (Table 1,
Fig. 1). From 1977 to 1994, C. Beck (National Biological Service,
personal communication) reports three live and three dead manatees
from six sites between 1982 and 1990; Clark adds two additional live
individuals in 1982 and 1987 and the remaining 42 specimens (41
sites) are from my study (Table 1). All but four occurrences have
been of live specimens. Recent manatee sightings have been of sub-
adults or young about 1.8-2.4 m long (Table 1). None has been weighed.
Manatees have been recorded from 11 coastal North Carolina coun-
ties (Table 1); 15 occurrences have been in Carteret County, nine
each in Dare and New Hanover, and eight in Craven County. Most
sightings occurred in 1983 and 1994 (nine), five in 1976, and two to
four sightings during most other years. Sightings have occurred dur-
ing nine months of the year, mostly in September (14), followed by
eight in August and October (Table 1). Most often frequented locali-
ties have been: Pelletier Creek (six), a tributary to Bogue Sound at
Morehead City (Carteret County), various localities in the Atlantic
Ocean (6), four at Wrightsville Beach and Sound (New Hanover County),
and the Trent River near New Bern, North Carolina (8) (Craven County).
While Rathbun et al. (1981) noted that open ocean habitat occur-
rences are rare, four North Carolina records have been from inlet-
ocean sites and six in the open ocean. Of the latter, three ocean
occurrences were off Shackleford Banks (Carteret County), one off
Wrightsville Beach (New Hanover County), and one each off the
Outer Banks off Avon and Kitty Hawk (Dare County) (Fig. 1). Most
sightings have been of single individuals; the largest groups sighted
for several days before disappearing were three at Minnesott Beach
Yacht Basin, a tributary to the Neuse River (Pamlico River) in Au-
gust 1980, and three in the Trent River at the Sheraton Yacht Harbor,
New Bern (Craven County) in September 1994.

Although six records are of manatees frequenting Pelletier Creek,
the area seems an unlikely manatee habitat as the short 0.8 km creek
is plied by many boats, pollution is heavy from numerous boat works
and marinas, and considerable runoff occurs from the densely built
houses and condominiums lining the shoreline. Yet manatees have
spent several days in the creek during each visit. In 1993 to 1994 two
(once) to six separate manatees frequented the same general area of
the coffee-colored Trent River, tributary to the Neuse River, 2 km
southwest of New Bern (Craven County) and fed on dense aquatic
vegetation. Three specimens frequented the river 0.2 km west of New

Manatees in North Carolina 59

Bern, at Lawson Park or the Sheraton Hotel Yacht Basin in 1994. The
1993 specimen was followed downstream in the Neuse along its
western shore for 13 km before vanishing. The July 1994 specimen
did not linger in the Trent River prior to its next (presumed) sight-
ing in the lower Neuse River opposite Oriental in late July. August-
September 1994 specimens lingered and fed on the lush vegetation of
the Trent River even as late as 24 September 1994. Bottom and surface
salinities 17 September 1994 were 14 and 7 ppt respectively.

INLAND PENETRATIONS

Manatees are known to penetrate inland freshwater such as the
St. John's River of Florida for 224 km to Lake Monroe (Volusia
County; D. Odell, Sea World Inc., personal communication). Farthest
inland river penetrations by manatees in North Carolina have been:
Cape Fear River (one) for 94.4 km-6.4 km above Route 210 and north
of Wilmington (Pender County) 11 July 1993, and one for 92 km in
the Neuse River to the Fort Barnwell Bridge (Craven County) in
October 1980, 33 km northeast of New Bern (Fig. 1). Two other
manatees occurred in the Tar River system of the upper Pamlico
River, a tributary of Pamlico Sound: one for 88 km to Greenville (Pitt
County) in November 1989 and the other for 58 km to just upstream
of Washington (Beaufort County) (Fig. 1) in September 1985. The
most peculiar movements have been of three manatees that traversed
south from the lower Chesapeake Bay in late summer of 1993 via the
canal and Intracoastal Waterway into Currituck Sound (Fig. 1). While
most manatee sightings have been at localities which could have been
reached via the Intracoastal Waterway and sounds, the four inlet and
six ocean occurrences also suggest travel north from Florida to North
Carolina may have been via those avenues. In any event, observations
seem to indicate many more young manatees are expanding their range
into North Carolina, perhaps as a result of an increased public aware-
ness of Florida's manatees rather than a real population increase.

ACKNOWLEDGMENTS— Thanks are extended to the many
interested people of North Carolina who called or commented on
manatee occurrences in North Carolina. Others, elsewhere, who were
also helpful in supplying information were: C. Beck, National Bio-
logical Service, Sirenia Project, Gainesville, FL (occurrences, sizes,
and Rathbun et al. 1981 reference); D. Odell, Sea World Inc., Or-
lando, FL, (St. John's River occurrence, file data, and review of manu-
script; R. Dove, Neuse River Keeper (1994 Trent River observations);
J. Wolfenberger, (Minnesott Beach Yacht Basin observations); New

60 Frank J. Schwartz

Hanover Public Library (Cape Fear River stories); J. Oakley, Carolina
Biological Lab, Burlington (early Brimley references); M. Nabinetts,
Washington Daily News (Pamlico River stories), N. Winfrey, Orien-
tal News (comments on lower Neuse River observations); M. Clark
provided the three North Carolina State Museum of Natural Sciences
records and checked the Museum overall occurrence records. D. Lee
of the North Carolina State Museum of Natural Sciences reviewed the
paper, as did C. Potter, National Marine Fisheries Service, Division
of Mammals (North Carolina data). D. Webster, University of North
Carolina, Wilmington, noted the 29 July 1992 record. L. White typed
the text; R. Barnes produced Figure 1.

LITERATURE CITED

Brimley, C. S. 1905 (1908). A descriptive catalogue of the mammals
of North Carolina, exclusive of the Cetacea, Journal of the Elisha
Mitchell Scientific Society 21:1-32.

Brimley, C. S. 1946. The mammals of North Carolina, installment
No. 17. Carolina Tips 9(l):2-3.

Brimley, H. H. 1931. Mammals of North Carolina. Journal of Mammalogy
12:320-321.

Caldwell, D. K., and F. B. Golley. 1965. Marine mammals from the
coast of Georgia to Cape Hatteras. Journal of the Elisha Mitchell
Scientific Society 81:24-32.

Campbell, H. W. 1977. Mammalia: Sirenia; Trichechidae {Tricheches
manatus Linnaeus). Pages 396-397 in Endangered and threatened
plants and animals of North Carolina (J. E. Cooper, S. S. Robinson,
and J. B. Funderberg, editors). North Carolina State Museum of
Natural History, Raleigh.

Jefferson, T. A., S. Leatherwood, and M. A. Webber. 1993. Marine
mammals of the world. FAO Species Identification Guide, Rome,
Italy.

Lee, D. S. 1976. Manatees: Mammals in distress. Wildlife in North
Carolina 40(7):8-9, 27.

Mitchell, K. B. 1993. My god, that's not a beaver. Wilmington Star
News, 20 July: la, 4a.

Rathbun, G. B., R. K. Bonde, and D. Clay. 1981. The status of the
West Indian manatee on the Atlantic Coast north of Florida. Pages
152-165 in Proceedings of the Symposium on nongame and en-
dangered wildlife (R. R. Odom and J. W. Guthrie, editors). Georgia
Department of Natural Resources, Game Fish Division Technical
Bulletin WL5, Atlanta.

Smith, E. R., J. B. Funderberg, and T. L. Quay. 1980. A checklist
of North Carolina mammals. North Carolina Wildlife Resource
Commission, Raleigh.

Received 13 October 1994
Accepted 10 February 1995

Recovery of the Cave Crayfish (Decapoda: Cambridae)
Population in Peacock Springs, Florida?

W. J. Streever

Department of Environmental Engineering Sciences,

University of Florida, Gainesville, Florida 32611

ABSTRACT— In 1991, a cave fauna kill was observed within
the fully-flooded cave system at Peacock Springs, Suwannee
County, Florida. Organisms affected by the kill included the
pallid cave crayfish (Procambarus pallidus), the yellow bull-
head (Ameiurus natalis), the American eel (Anguilla rostrata),
and the Asiatic clam (Corbicula fluminea). Following the kill,
no crayfish were present along transects, but recently collected
census data are not significantly different from data collected
before the 1991 kill (Paired Mest, P > 0.1). However, cray-
fish numbers have not returned to their pre- 1991 levels in the
cave passage where the highest crayfish density occurred
before the kill. Also, the scarcity of small (<1.5 cm total
length) crayfish suggests that the return to pre-1991 levels may
reflect dispersal of animals from inaccessible portions of the
cave and not replacement of crayfish through reproduction.

The troglobitic pallid cave crayfish, Procambarus pallidus,
inhabits flooded caves in the Suwannee River drainage in north
Florida (Franz and Lee 1982). These caves also are inhabited by other
troglobitic and troglophillic species, including several species of
catfish (Hale and Streever 1994), ostracods (Walton and Hobbs 1959),
the American eel Anguilla rostrata (Franz et al. 1994), the Asiatic
clam Corbicula fluminea (Streever 1992a), tubificidae worms (Streever
19926), and the colonial cnidarian Cordylophora lacustris (Streever
1992c). Franz et al. (1994) provide an extensive review of Florida's
cave fauna, emphasizing distributional records for all species report-
ed in association with caves. Little is known about the ecology or
population dynamics for most of these species.

The Peacock Springs cave system consists of over 3,300 m of
fully-flooded passage, with depths down to about 60 m. Eight openings
provide access to the cave system. A surface channel connects open-
ings commonly called Peacock I, II, and III to the Suwannee River.
Water from the Floridan aquifer normally flows out of the Peacock I
opening and drains into the Peacock III opening and the Suwannee

Brimleyana 22:61-65, June 1995 61

62 W. J. Streever

River. However, during periods of high river levels, water from the
Suwannee River overcomes the head of pressure generated by aquifer
water. Flow in the surface channel reverses, and Suwannee River
water enters the cave system. Following a flow reversal in February
1991, virtually all of the animals in accessible portions of the Peacock
Springs cave system were killed. Animals affected by the kill include
crayfish, catfish, and Asiatic clams (Streever 19926). The cause of
the kill is unknown.

This paper addresses two issues. First, I present 1994 P. pallidus
census data and subjective observations of other taxa, and second
I discuss a possible mechanism for the increase in crayfish population
density following the 1991 kill.

METHODS

In 1990, divers established eight belt transects (100 x 4 m) in
association with permanent guidelines in the Peacock Springs cave
system. Transects were numbered 1-8. Both before and after the fauna
kill in February 1991, crayfish were visually censused along transects
by divers carrying 50-watt lights. Because cave crayfish roam in the
open on the cave floor, visual censusing provides a simple method of
estimating population density. After the kill, two additional belt tran-
sects, called 2-a (240 m long) and 8-a (350 m long), were established
in a tunnel north of the Peacock I opening and north of the Peacock
III opening, respectively. Transect 2-a incorporated transect 2, and
transect 8-a incorporated transects 7 and 8. All crayfish that were
visible from the cave's permanent guidelines were included in censuses
along transects 2-a and 8-a, making their width dependent on water
clarity and size of the cave passage. In general, the width of transects
2-a and 8-a varied between 4 and 10 m. The large size of the new
transects was intended to allow censusing of the smaller crayfish
densities occurring after the 1991 kill. A map of the cave showing
transect locations was provided by Streever (19926).

The 1994 transect censuses were made between 9 January 1994
and 25 November 1994. During the 1994 censuses, crayfish were
recorded as belonging to a size class of less than or greater than 1.5
cm in overall length, estimated from the anterior edge of the rostrum
to the posterior edge of the telson. All censuses along each of the
original transects were averaged to yield values for each transect
before the kill and in 1994. The null hypothesis of no difference
between censuses taken before the 1991 kill and those taken in 1994
was tested by a paired Mest, with averages of before and after censuses
along individual transects comprising pairs (Zar 1984).

Crayfish Population 63

RESULTS AND DISCUSSION

Differences in censuses (Table 1) before the 1991 kill and in
1994 are not statistically significant (P > 0.1). The increase in crayfish
population density immediately after the kill and in 1994 suggests
that the population is recovering. However, 1994 censuses in the Peacock
III tunnel indicate that populations have not recovered to pre-kill
levels along transects 7 and 8. Transect 8a, which encompassed transects
7 and 8 as well as an additional 100 m of cave passage, contained
only a fraction of the number of crayfish that were present along
transects 7 and 8 before the kill. This indicates that the large crayfish
population found in the Peacock III tunnel before the kill is not
recovering, nor is the overall population in the cave system.

Of the 138 crayfish censused after the 1991 kill, only one
animal was smaller than 1.5 cm. Although animals smaller than 1.5
cm in total length may be missed during censusing, the scarcity of

Table 1. Census data for Procambarus pallidus in the Peacock Springs,
Florida, cave system before the 1991 kill, immediately after the kill,
and in 1994.

Numbe

r of

crayfish counted

Immediately after

Before 1991
x S.D.

kill
n

1991

X

kill

1994

Transects

n x

S.D.

n

1

1.0

1.0

3

0

1 1.0

1

2

6.5

0.5

2

0

1 1.0

2

2a1

not

counted

0

1 9.7

5.4

3

3

1.5

0.5

2

0

1 7.0

1

4

4.0

0.0

3

0

1 0.0

0.0

2

5

1.0

0.0

4

0

1 1.0

1

6

6.2

0.8

5

0

1 6.0

1.0

2

7

34.5

2.7

8

0

1 0.0

0.0

8

8

49.5

7.1

8

0

1 0.0

0.0

8

8a1

not

counted

0

1 10.1

6.6

8

These transects were established after the 1991 kill to cover a larger area than
original transects.

64 W. J. Streever

small animals suggests that recovery of crayfish population density is
not dependent on replacement through reproduction. Cave popula-
tions frequently display size distributions with many large individuals
and few small individuals, suggesting infrequent reproduction (Poulson
1963, Culver 1982). The fecundity of P. pallidus is unknown, but an
extensive study of the cave-adapted crayfish Orconectes australis
australis in Shelta Cave, Alabama, suggests that reproduction in cave
crayfishes may be infrequent and that clutch sizes may be small
(Cooper 1975). If a similar pattern occurs in P. pallidus, crayfish
population recovery that depends on reproduction could be slow. Be-
cause crayfish population densities in the Peacock Springs cave sys-
tem do not appear to be recovering through reproduction, the increase
in numbers along transects 1 through 6 may not represent a true
recovery of the crayfish population. Instead, crayfish density may be
the result of colonization by individuals that survived the kill and
are now moving into the portion of the cave where transects were
located.

Catfish were not counted as part of this study. However, they
were common along transects before the 1991 kill, they were absent
from transects immediately after the kill, and they were common
during the 1994 census dives. An Asiatic clam population near the
mouth of Peacock III had an estimated density of 161 individuals/m 2
before the 1991 kill (Streever 1992a), but no live Asiatic clams were
found immediately after the kill or in 1994. Empty clam shells, pre-
sumably remaining from before the kill, were abundant.

The causes of the kill and the factors affecting recovery are
unknown. However, intrusion of river water into caves typically full
of groundwater appears to be linked to cave kills in the Peacock
system. On 19 November 1994, two dead crayfish and a dead catfish
were found along transect 6, immediately after a period of flooding
during which Suwannee River water entered the cave. However, seven
live crayfish were found along the transect on the same day, so this
event did not have the intensity of the 1991 kill. Despite this apparent
association, the link between river water and crayfish mortality is not
clear. P. pallidus has been observed in two siphon-spring cave
systems flooded by Sante Fe River water throughout the year
(personal observation), so intolerance of low temperatures and other
environmental factors associated with river water may not be the
cause of mortality. Furthermore, kills have been observed in other
caves without flooding by river water (Franz et al. 1994). Establish-
ment of the cause of cave crayfish kills requires further investigation.

Crayfish Population 65

ACKNOWLEDGMENTS—! thank cave divers Jason Hale, Wendy
Short, Steve Brooker, and Rob Korn for their assistance with field
work. The State of Florida Department of Parks and Recreation allowed
access to the Peacock Springs cave system. This work was funded in
part by a National Science Foundation Graduate Research Fellowship.

LITERATURE CITED

Cooper, J. E. 1975. Ecological and behavioral studies in Shelta
Cave, Alabama, with emphasis on decapod crustaceans. Ph.D. Thesis.
University of Kentucky, Lexington.

Culver, D. C. 1982. Cave life, evolution and ecology. Harvard Uni-
versity Press, Cambridge, Massachusetts.

Franz, R., J. Bauer, and T. Morris. 1994. Review of biologically
significant caves and their faunas in Florida and south Georgia.
Brimleyana 20:1-109.

Franz, R., and D. S. Lee. 1982. Distribution and evolution of Florida's
troglobitic crayfishes. Bulletin of the Florida State Museum Bio-
logical Sciences 28:53-78.

Hale, J. A., and W. J. Streever. 1994. Cave fauna distribution
within fully-flooded cave systems in Florida. Journal of Freshwa-
ter Ecology 9:171-174.

Poulson, T. L. 1963. Cave adaptation in amblyopsid fishes. Ameri-
can Midland Naturalist. 70:257-290.

Streever, W. J. 1992a. First record of Corbicula clams within flooded
cave systems. Florida Scientist 55:35-37.

Streever, W. J. 1992/?. Report of a cave fauna kill at Peacock
Springs cave system, Suwannee County, Florida. Florida Scientist
55:125-128.

Streever, W. J. 1992c. First record of the colonial cnidarian Cordylophora
lacustris within a flooded cave system. NSS Bulletin 54:77-78.

Walton, M., and H. H. Hobbs, Jr. 1959. Two new eyeless Ostracods
of the genus Entocythere from Florida. Quarterly Journal of the
Florida Academy of Sciences 22:114-120.

Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Englewood
Cliffs, New Jersey.

Received 1 January 1995
Accepted 14 February 1995

Premolar Cementum and Noncementum Lengths

As Potential Indicators of Age for Beavers,

Castor canadensis (Rodentia: Castoridae)

Allan E. Houston

Ames Plantation, Agricultural Experiment Station, The University

of Tennessee, P. O. Box 389, Grand Junction, Tennessee 38039-0389

AND

Michael R. Pelton

Department Forestry, Wildlife, and Fisheries, The University

of Tennessee, Knoxville, Tennessee 37901-1071

ABSTRACT — Jaws from 28 beavers (Castor canadensis), >4 years
old as determined from cementum annuli on premolars and
molars, were collected in west Tennessee. An exponential model
predicting age was developed based on the independent vari-
ables cementum length and noncementum length as measured
along the estimated, maximum longitudinal centerline of cross-
sectioned premolars: AGE = 0 + 6.1784 * e (:o.io3i*noncemen) +

2 6513 * e (0.1U9*CEMENTUAf)

Several methods of aging beavers have been attempted with varying
degrees of accuracy, including pelt size (Buckley and Libby 1955),
tail dimensions, total body mass, skull measurements (Patric and Webb
1960), and baculum size and mass (Friley 1949). Probably the most
accurate method was developed by van Nostrand and Stephenson (1964)
using tooth eruption and closure of basal openings of premolars and
molars for beavers up to 3 years old, and cementum layering in premolars
and molars for specimens >4 years. Larson and van Nostrand (1968)
further refined this technique to include criteria dealing with cementum
deposition around basal openings. They noted that age estimations
may be complicated by multi-annual cementum layering, but that the
ratio of cementum to noncementum on molars and premolars might
be used to estimate age in older specimens.

Our objective was to use premolar cementum length (aggregate
of all annual cementum depositions) and noncementum length (remains
of the original tooth) as independent variables to evaluate a model for
estimating age of beavers >4 years.

Brimleyana 22:67-72, June 1995 67

68 Allan E. Houston and Michael R. Pelton

METHODS

This study was conducted in the upper headwater basin of the
North Fork of the Wolf River in west Tennessee, largely on the Ames
Plantation. From November 1984 through May 1985, beaver jaws
from 169 beavers were collected from legal trapping efforts on
approximately 1,619 ha of the watershed.

Molars and premolars from lower mandibles were extracted and
cleaned. If tooth basal openings indicated a specimen to be >4 years
old, age was determined by grinding the lingual surface of premolars
or molars with a 120-grit stone to expose longitudinal cross-sections.
Ground surfaces were polished with a 400-grit emery cloth and cementum
layers were carefully counted using hand-held magnification (Larson
and van Nostrand 1968). Twenty-eight beavers, >4 years old as aged
in this manner (Table 1), provided the samples (i.e., dependent variables)
for our study.

Premolar cementum length and noncementum length were measured
to the nearest millimeter along the estimated maximum longitudinal
cross-sectional centerline of one premolar per specimen (Fig. 1). An
exponential model predicting age in years was developed by combining
a growth curve for cementum length and a decay curve for noncementum,
using nonlinear regression (PROC NLIN) and the multivariate secant
method (DUD) to set initial parameters (SAS Institute, Inc. 1985).
Intercept was specified at zero to prevent predicted ages from
dipping below zero. An "r2 like" statistic was calculated by taking
[1 - (residual SS/corrected total SS)].

RESULTS AND DISCUSSION

The following model was developed using cementum length and
noncementum length of premolars as independent variables:

AGE = 0 + 6 1784 * e (-o.io37*noncemen) + 2.6513 * e (°ul9*CEMENTUM)t

The "r2 like" statistic was 0.93. The predicted curves are combined
to produce a response surface (Fig. 2).

Our data were obtained from specimens on the headwaters of a
single watershed. Although beavers can travel considerable distances,
we assumed our sample represented only a small region. The study
area may not fully represent variation existing range-wide or within
adjacent watersheds. It is possible that genetic differences and dietary
regimes will yield different tooth size, wear, and cementum accretion

Aging Beaver Teeth 69

Table 1. Age, as estimated by cementum annuli, cementum and noncementum
length, measured to the nearest millimeter along the maximum longitu-
dinal cross-section of one premolar per specimen, and predicted ages
and residuals for beavers captured in west Tennessee, November 1984
through May 1985.

Specimen

Age

Cementum

Noncementum

Predicted Age1

Residual

1

4

3

22

4.34

-0.34

2

4

3

22

4.34

-0.34

3

4

1

24

3.48

0.52

4

4

3

22

4.34

-0.34

5

4

2

21

4.02

-0.02

6

4

3

21

4.41

-0.41

7

4

3

19

4.57

-0.57

8

4

1

24

3.48

0.52

9

4

1

24

3.48

0.52

10

4

2

20

4.09

-0.09

11

4

1

22

3.60

0.40

12

4

2

22

3.95

0.05

13

4

3

21

4.41

-0.41

14

5

4

14

5.59

-0.59

15

5

6

16

6.36

-1.36

16

5

5

20

5.42

-0.42

17

7

6

19

6.05

0.95

18

8

7

13

7.41

0.59

19

8

9

11

9.23

-1.23

20

9

9

10

9.45

-0.45

21

9

8

13

8.10

0.90

22

9

9

8

9.95

-0.95

23

9

9

9

9.69

-0.68

24

10

8

9

8.92

1.08

25

10

9

11

9.23

0.77

26

10

8

11

8.46

1.54

27

11

10

10

10.31

0.69

28

12

12

10

12.35

-0.35

AGE = 6.174*e<-°-1037*"°"C£M£") + 2.6513V0

1U9*CEMENTUM)

70

Allan E. Houston and Michael R. Pelton

Fig. 1. Schematic representation of cementum length and noncementum
length as measured along longitudinal cross-sections of premolars for
beaver.

Aging Beaver Teeth

71

PREDICTEDAGE
(YEARS)

V 12.00

CEMENTUM LENGTH
67 (MM)

18.67

NONCEMENTUM LENGTH
(MM)

Fig. 2. Response surface of an exponential model predicting age of
west Tennessee beaver using cementum length and noncementum length
measured along the estimated maximum longitudinal cross-sections of
premolars as independent variables.

patterns. Comparative study is needed to document potential variation
of these criteria. Also, the method should be validated using known-
age specimens. However, these results suggest that this technique
could be developed as a reliable method to age beavers.

ACKNOWLEDGMENTS— -The project was funded by the Hobart
Ames Foundation and The University of Tennessee Agricultural Experiment
Station. J. Schneider gave invaluable assistance with statistical analyses.
R. Henry, J. Morrow, and T. Sain provided the majority of data collections.
R. Maxey contributed the drawing in Figure 1.

LITERATURE CITED

Buckley, J. J., and W. L. Libby. 1955. Growth rates and age determi-
nation in Alaska beaver. Transactions of the North American
Wildlife and Natural Resources Conference 20:495-505.

Friley, C. E., Jr. 1949. Use of the baculum in age determination of
Michigan beaver. Journal of Mammalogy 30(3):261-267.

Larson, J. S., and F. C. van Nostrand. 1968. An evaluation of beaver
aging techniques. The Journal of Wildlife Management 32(1):99-103.

72 Allan E. Houston and Michael R. Pelton

Patric, E. F., and W. L. Webb. 1960. An evaluation of three age

determination criteria in live beavers. The Journal of Wildlife

Management 24:37-44.
SAS Institute, Inc. 1985. SAS User's Guide: Statistics. SAS Institute,

Inc., Box 8000, Cary, North Carolina,
van Nostrand, F. C, and A. B. Stephenson. 1964. Age determination

for beavers by tooth development. The Journal of Wildlife

Management 28(3):430-434.

Received 11 January 1994
Accepted 8 March 1995

Record Clutch Size for Chelydra serpentina
(Testudines: Chelydridae) in Virginia

Joseph C. Mitchell

Department of Biology

University of Richmond,

Richmond, Virginia 23173

AND

Michael C. Odom
U.S. Fish and Wildlife Service

11110 Kimages Road
Charles City, Virginia 23030

ABSTRACT — A record clutch size of 67 is reported for a
common snapping turtle {Chelydra serpentina serpentina) from
Charles City County, Virginia, 12 larger than previously
reported for this chelonian. The new average for this species
in the Commonwealth is 29.9 ± 16.6 ( x ± SD).

The snapping turtle (Chelydra serpentina) occurs from
Florida north to southern Canada and Nova Scotia in eastern North
America and from the Mexican border north to extreme southern
Saskatchewan in western North America (Conant and Collins 1991,
Iverson 1992, Russell and Bauer 1993). Clutch size varies latitudinally,
with the largest sizes reported for northern populations in Canada,
South Dakota, and Nebraska (Zug 1993; J. B. Iverson, Earlham College,
personal communication). Ash (1951) reported an average clutch
size of 28.6 and a range of 13-48 for 85 specimens presumably caught
in the Virginia area. However, these data were presented only in
an abstract. A complete paper was never published, and no docu-
mentation is available for us to determine the source of his
specimens. Information provided by the late John T. Wood
(formerly a retired medical doctor in Victoria, British Columbia
and Virginia in the 1950s, personal communication) indicated that
Ash's specimens could have been from multiple locations in and
out of Virginia. Mitchell (1994) reported an average clutch size of
27.0 ± 13.2 (range = 7-55) for 13 females collected from various
locations in Virginia.

On 3 June 1994, a gravid female (271-mm carapace length,
201-mm plastron length) was inadvertently killed by a vehicle
at Harrison Lake National Fish Hatchery, Charles City County,
Virginia. Subsequent dissection revealed 67 shelled eggs. One egg

Brimleyana 22:73-75, June 1995 73

74 Joseph C. Mitchell and Michael C. Odom

was opened on the date of collection, and the rest were placed in a
closed plastic container with a 3:1 mixture of top soil and vermiculite.
The soil was kept moist during the incubation period so that the
relative humidity was maintained at nearly 100%. Incubation temperatures
varied from 25 to 30° C. During the course of incubation, 12 of the
66 eggs died and were discarded; 54 eggs survived. Hatching
commenced on 20 August (78 days incubation), and the last egg
pipped on 24 August (82 days). Hatchlings showed considerable
variation in development, with larger individuals bearing a small
yolk sac and smaller individuals bearing a large, cumbersome yolk
sac. Of the 54 hatchlings, four failed to survive through yolk sac
absorption. The remaining turtles began exogenous feeding as their
yolk sac diminished, taking a diet of chopped nightcrawlers and
miscellaneous live aquatic macroinvertebrates collected from
ponds. The surviving juvenile turtles were released the following
spring.

A documented clutch size of 67 eggs is 12 larger than the
largest reported for Virginia (Mitchell 1994). Incorporation of the
large clutch size reported here yields a new state average of 29.9 ±
16.6 (range = 7-67). The maximum known clutch size for Virginia
populations lies between those known for northern populations
(83: Quebec (Bleakney 1957); 73: New York and Wisconsin (Yntema
1970); 109: Nebraska (Packard et al. 1990)) and southern populations
(43: North Carolina (Brown 1992); 21: Florida (Punzo 1975)).

Reports of average clutch sizes for snapping turtles in different
geographic areas based on small sample sizes should be used with
caution. Such averages will almost always change with additional
data.

ACKNOWLEDGMENTS— Viz thank Kurt A. Buhlmann and John
B. Iverson for their comments on the manuscript, and Thomas F.
Burnett for assistance in the collection of the adult turtle and
maintenance of the eggs and hatchlings.

LITERATURE CITED

Ash, R. P. 1951. A preliminary report on the size, egg number, incu-
bation period, and hatching in the common snapping turtle, Chelydra
serpentina. Virginia Journal of Science 2:312 (abstract).

Bleakney, S. 1957. A snapping turtle, Chelydra serpentina serpentina,
containing eighty-three eggs. Copeia 1957:143.

Chelydra serpentina clutch 75

Brown, E. E. 1992. Notes on amphibians and reptiles of the western
Piedmont of North Carolina. Journal of the Elisha Mitchell Sci-
entific Society 108:38-54.

Conant, R., and J. T. Collins. 1991. A field guide to the reptiles
and amphibians of eastern and central North America. Houghton
Mifflin Company, Boston, Massachusetts.

Iverson, J. B. 1992. A revised checklist with distribution maps of
the turtles of the world. Privately Printed, Richmond, Indiana.

Mitchell, J. C. 1994. The reptiles of Virginia. Smithsonian Institution
Press, Washington, D.C.

Packard, G. C, M. J. Packard, and K. Miller. 1990. Life history:
Chelydra serpentina, fecundity. Herpetological Review 21:92.

Punzo, F. 1975. Studies on the feeding behavior, diet, nesting
habits and temperature relationships of Chelydra serpentina osceola
(Chelonia: Chelydridae). Journal of Herpetology 9:207-210.

Russell, A. P, and A. M Bauer. 1993. The amphibians and reptiles of
Alberta. University of Calgary Press, Calgary, Alberta, Canada.

Yntema, C. L. 1970. Observations on females and eggs of the com-
mon snapping turtle Chelydra serpentina. American Midland Naturalist
84:68-76.

Zug, G. R. 1993. Herpetology, an introductory biology of amphibians and
reptiles. Academic Press, San Diego, California.

Received 12 January 1995
Accepted 9 March 1995

New Distributional Records for the Star-nosed Mole,

Condylura cristata (Insectivora: Talpidae), in

North Carolina, with Comments on its Occurrence

in the Piedmont Region

Jeffrey C. Beane

North Carolina State Museum of Natural Sciences

P.O. Box 29555

Raleigh, North Carolina 27626-0555

ABSTRACT — The distribution of the star-nosed mole (Condylura
cristata) in North Carolina is updated, including specimen-supported
records for seven new counties and sight records for several
additional counties. The species is locally common in the Mountains,
spottily distributed throughout much of the Coastal Plain, and
apparently rare to absent in much of the Piedmont. A specimen
from just off the Blue Ridge escarpment in western Surry
County represents the first Piedmont record supported by a
specimen, although there are additional sight records and unverified
reports from that region. This mole is sympatric with the state's
other two mole species in many areas, but usually occurs in
wetter habitats than either.

INTRODUCTION

The star-nosed mole (Condylura cristata) ranges from eastern
Manitoba and Minnesota eastward to Labrador and Nova Scotia, and
southeastward along the Atlantic coast to southeastern Georgia (Paradiso
1959, Burt and Grossenheider 1976, Petersen and Yates 1980, Hall
1981, Yates and Pedersen 1982). In North Carolina, its distribution
has frequently been reported as consisting of two distinct popula-
tions— one in the Mountains, where the species is locally common,
and one in the Coastal Plain, where it is uncommon to rare — with a
conspicuous hiatus in the Piedmont (Lee et al. 1982, Webster et al.
1985, Lee 1987, Webster 1987). The scarcity of records from the
Coastal Plain and complete absence of Piedmont records resulted in
the species being recommended for listing as status Undetermined by
the Nongame Advisory Committee to the North Carolina Wildlife
Resources Commission (Webster 1987), and in 1990 the species was
granted protection as a species of Special Concern under the North
Carolina Endangered Species Act (G.S. 113-331 to 113-337).

I here report on the current known distribution of Condylura
cristata in North Carolina by providing a list of existing records from
the state, and a dot distribution map (Fig. 1) updated from maps
provided by Lee et al. (1982) and Webster (1987).

Brimleyana 22:77-86, June 1995 77

78

Jeffrey C. Beane

Fig. 1. Known distribution of Condylura cristata in North Carolina.
Solid dots indicate localities documented by specimens in curated collections.
Open circles indicate literature records or sight records not supported
by specimens. A single dot or circle may represent two or more records
in close proximity. A "C" indicates a literature record, not supported
by a specimen, for which only the county and no further information
was given. Question marks accompany records for which no specimen
was observed, or which are unverifiable or otherwise questionable.

RECORDS OF OCCURRENCE
The following North Carolina localities are supported by voucher
specimens, many of them in the North Carolina State Museum of
Natural Sciences (NCSM), and are represented by solid dots on the
distribution map:

Alleghany Co.: New River, "Sparta" (The New River is actually, at
its nearest point, 4 mi. NW of Sparta), 30 May 1968, H. M. Tyus
(NCSM 1762). Ashe Co.: "3A mi. from jet. NC 16 and Friendship
Church Rd." (= ca. 2 mi. ENE Jefferson), 4 Oct. 1991, E. Marsh
(Appalachian State University 16731). Avery Co.: 2.6 mi. NNE Heaton,
Pisgah National Forest, 27 May 1975, R. Jordan and J. Pentecost
(Auburn University Museum 1-281); 3A mi. SE Linville, 24 July 1984,
D. S. Lee and J. P. Kumhyr (NCSM 4875). Brunswick Co.: ca. 6 mi.
NNW Supply, Green Swamp Ecological Preserve off NC 211, "Moon
Island." 16 May 1993, T. M. Padgett (TMP 511; to be deposited at
NCSM). Buncombe Co.: W of Asheville (University of Michigan Mu-
seum of Zoology); Biltmore (Harvard Museum of Comparative Zool-
ogy 6635); ca. 2.6 mi. WNW Swannanoa, Warren Wilson College
campus, Aug. 1977, D. S. Lee and B. Lee (NCSM 7837); Weaverville,
6 Feb. 1896, Mrs. J. S. Cairns (Harvard Museum of Comparative
Zoology 5399). Dare Co.: ca. 7 mi. SW Manns Harbor, Alligator

Star-nosed Mole 79

River National Wildlife Refuge, Milltail Rd., 20 March 1991, P. R.
Trail, M. L. Dunn, et al. (NCSM 6710). Haywood Co.: 5 mi. SW
Waynesville on U.S. 23, 30 Aug. 1951, F. S. Barkalow, Jr. (North
Carolina State University 274). Henderson Co.: Hoopers Creek near
Fletcher, 8 Oct. 1974, B. A. Sanders (NCSM 4937); 1 mi. S Gerton,
Little Bearwallow Mtn., 8 July 1994, P. B. Spivey (to be deposited at
NCSM); 1 mi. W. NW Mills River (town), near jet. SR 1336 and SR
1338, 25 May 1993, K. A. Buhlmann, D. W. Herman, and J. L. Warner
(NCSM 7868). Hoke Co.: 9.2 mi. NE Ashley Heights, Ft. Bragg Mili-
tary Reservation, 29 May 1992, LCTA survey crew (NCSM 7213).
Jackson Co.: 0.7 mi. NE Balsam, Blue Ridge Parkway at Balsam
Gap, 22 May 1991, A. C. Boynton (NCSM 7528). Macon Co.: High-
lands, 1 June 1946 (University of Georgia Museum of Natural History
192); Highlands, ca. 1 June 1946, R. Bridges (NCSM 6575); ca. 0.5
mi. E center Highlands, Highlands Biological Station, 23 Oct. 1982,
S. Morrison (University of Georgia Museum of Natural History 7595);
1 mi. E.NE Highlands, SR 1603, 0.3 rdmi. E jet. SR 1604, 20 Sept.
1984, A. L. Braswell and J. E. Cooper (NCSM 7462). New Hanover
Co.: near Carolina Beach, 16 Nov. 1956, G. Tregembo (NCSM 3243).
Mitchell Co.: Roan Mtn. (Wake Forest University); Roan Mtn. near
Magnetic City, 15 March 1894 (U. S. National Museum 66300); Roan
Mtn. near Magnetic City, 15 Aug. 1895 (U. S. National museum
91519). Polk Co.: Green River Valley near Saluda, 28 May 1974,
G. G. Shaw and M. Bradley (NCSM 1237). Rutherford Co.: Chimney
Rock, Southside Drive, 19 Dec. 1989, M. Jaeger-Gale (NCSM 6336).
Richmond Co.: 3.7 mi. NE Hoffman, U.S. 1, 0.1 rdmi. S Lumber
River, 18 May 1980, A. L. Braswell (NCSM 3283). Sampson Co.: 5.5
km W Faison (Duplin Co.) off SR 1734, 9 June 1994, W. D. Webster
(University of North Carolina at Wilmington 3370). Scotland Co.:
"Laurinburg, Mr. Newton's farm" [the farm referred to was located
ca. 8 mi. N Laurinburg, or 4.5 mi. W Wagram (NCSM files, W. M.
Palmer, pers. comm.)], 12 April 1975, R. B. Julian (NCSM 3047).
Surry Co.: 0.8 mi. NNW Low Gap near jet. NC 89 and SR 1433,
A. B. Somers et al., 7 June 1994 (NCSM 7744). Swain Co.: Deep
Creek; Kephart Prong Hatchery; Smokemont; Charlies Bunion Mtn.
at Sevier Co., TN line; and Appalachian Trail between Newfound
Gap and Indian Gap at Sevier Co., TN line; all in Great Smoky
Mountains National Park (Great Smoky Mountains National Park
collection, Linzey and Linzey 1968); Great Smoky Mountains
National Park, Appalachian Trail, lA mi. S Boulevard Trail (= ca. 6.8
mi. NW Smokemont), 20 Aug. 1961, J. B. Westbrook (University of
Georgia Museum of Natural History 3112). Watauga Co.: Blowing

80 Jeffrey C. Beane

Rock (Wake Forest University); Boone (Appalachian State University
7606); Boone, Hidden Valley Circle, 14 March 1981, D. Notrichia
(Appalachian State University 7591); Boone, 1987, R. W. Van Devender
(Appalachian State University 14633); ca. 1 mi. WSW Boone, jet.
NC 105 and Poplar Grove Rd., 16 Sept. 1978 (Appalachian State
University 5125 ); ca. 3 mi. WSW Boone, Laurel Cr. along NC 105,
13 Sept. 1985, F. Authenreith (Appalachian State University 12086);
Blue Ridge TWP, Jakes Mtn. Rd. near jet. SR 1511, Dec. 1986, M. P.
Rowe (Appalachian State University 12786); Blue Ridge TWP, SR
1511 along Sand Spit Branch and Blue Ridge Parkway, 27 Sept. 1987,
R. W. Van Devender (Appalachian State University 14205); Sugar
Grove, jet. U.S. 321 and Edmiston Rd., 24 Nov. 1978, H. Rogers
(Appalachian State University 7601); Sugar Grove, Edmiston Rd., 5
Sept. 1985, R. N. Henson (Appalachian State University 12485); Watauga
Co., no further data (Appalachian State University 16751). Yancey
Co.\ Burnsville, 3 Feb. 1976, L. P. Hartis (NCSM 2409).

In addition to localities supported by voucher specimens, the
following literature and sight records are considered valid, and most
are included as open circles on the distribution map:
"Mountains near the border of South Carolina" (Audubon and Bachman
1851). Avery Co.: Elk Park (Lee 1987, NCSM files). Bladen Co.: ca.
2 mi. N.NE White Lake (town) off SR 1517 near jet. U.S. 701, 30
April 1992, T. M. Padgett and R. Rageot (Thomas M. Padgett, pers.
comm.). Buncombe Co.: 2-3 mi. N Black Mountain at Montreat, ca.
1915-1920, E. E. Brown (Elmer E. Brown, pers. comm.); Swannanoa,
ca. 20 April 1931, C. S. Clapp (Lee 1987, NCSM files). Cherokee
Co.: Topton, 13 May 1934, E. B. King (Brimley 1944-1946). Clay
Co.: Hayesville, 3 Aug. 1947, R. G. Vick (Paradiso 1959, NCSM
files; specimen formerly at NCSM but apparently lost). Macon Co.:
Highlands vicinity, specimens "not infrequently" taken by H. C. Harbison
prior to May 1908 (Brimley 1944-1946); Highlands vicinity, three
specimens reported by Johnston (1967); one of these, formerly in the
Highlands Museum, is now NCSM 6575; the same specimen was also
reported by Odum (1949). McDowell Co.: just N Ashford along U.S.
221, ca. 1800' elev., two specimens taken from the stomach of a
black rat snake (Elaphe obsoleta), 23 May 1960, E. E. Brown. These
specimens were reported by Brown (1979), but their specific locality
was not included in that paper (Elmer E. Brown, pers. comm.). Mecklenburg
Co.: Davidson (east side near golf course branch), ca. 1950-1955, E.
E. Brown (Elmer E. Brown, pers. comm.). New Hanover Co.: near
Carolina Beach, 7 June 1959, G. Tregembo (Lee 1987, NCSM files).
Pitt Co.: no further data (Lee et al. 1982, Clark et al. 1985, Lee 1987,

Star-nosed Mole 81

Webster 1987). The original source of this record seems uncertain,
and it is accompanied by a question mark on the distribution map in
Fig. 1. Robeson Co.: near Lumberton, early Dec. 1943, A. M. Ivey
(Brimley 1944-1946). Sampson Co.: near Garland, 13 May 1918, J. F.
Johnson (Brimley 1923, Brimley 1944-1946, Paradiso 1959). Washington
Co.: Wenona, early Nov. 1992, E. R. Rainey (Brimley 1923, Brimley
1944-1946, Paradiso 1959). Watauga Co.: Boone, four specimens, 19
May 1918, (Brimley 1944-1946). Yancey Co.: ca. 7 mi. W. Burnsville
along Lickskillet Branch, ca. 1981. J. McFee (Allen C. Boynton,
pers. comm.).

Lee (1987) and Webster (1987) reported C. cristata from Moyock
in Currituck County, based on a specimen from the digestive tract of
a mink obtained from a fur dealer in that town and reported by Wilson
(1954). However, Wilson (1954) indicated that some of the mink in
his study were taken in Camden and Dare counties, and no specific
locality was given for the specimen from which the mole was retrieved.
That it was from Currituck County is possibly an erroneous assumption.
It is therefore accompanied by a question mark on the distribution
map. Regardless, this record affirms the presence of Condylura in the
northeastern corner of the state.

Clark et al. (1985) suggested the occurrence of Condylura in
Bladen and Hoke counties (no specific localities given) based on the
presence of mole runs "almost certainly made by this species." Lee
(1987) reported C. cristata from McCain, Hoke County, citing Clark
et al. (1985) as the source. That paper does not, however, contain a
clear reference to the species' occurrence at McCain. Lee (1987)
further reported observation of "burrows and mounds of Condylura"
from West End, Moore County, in 1980. Since these records did not
involve the observation of actual specimens, they are accompanied by
question marks on the distribution map.

Lee (1987) also reported C. cristata from Green [sic] County,
"pre-1950," citing Clark et al. (1985) as the source, but that paper
contains no reference to the species occurring in Greene County. Neither
are there records from that county in the North Carolina State Museum's
files, hence, it is not included on the distribution map.

Two additional reports of C. cristata from the Piedmont of North
Carolina are as follows.

Randolph Co.: 2.1 mi. SSE Ulah, off SR 2843 < 0.1 rdmi. SE
jet. NC 159. John Schneider, a horticulturist with the North Carolina
Zoological Park, reported (personal communication) a specimen that
was captured and badly mangled by his dog in March or April, 1991.
I visited the site in July, 1994, and found no obvious signs of fresh

82 Jeffrey C. Beane

mole activity, but no serious collecting efforts have yet been undertaken
there.

Wake Co.: 1.25 mi. SW Millbrook (= 4.0 mi. N.NE center Raleigh),
4001 Quail Hollow Drive along Big Branch. Dr. Wesley E. Kloos, a
genetics professor at North Carolina State University, reported (personal
communication, NCSM files) having captured two specimens in his
back yard between 1989 and 1991, and released both in nearby Eastgate
City Park. A photograph sent to the North Carolina State Museum, of
Kloos holding one of the captured animals, was too small and blurred
for positive identification. Frequent pitfall trapping on the site during
1992-1993 yielded no specimens.

Although neither Kloos nor Schneider is a mammalogist, both
are scientifically oriented individuals, and each seemed certain of the
animals's identity. The star-nosed mole is certainly a difficult animal
to misidentify. Their reports are therefore included here, and are probably
valid, although they remain unverified, and are accompanied by question
marks on the distribution map. Except for the Surry County record,
and the Mecklenburg County sight record by Brown, they represent
the only reports from the Piedmont region of the state. However, the
records from Polk and Rutherford counties (NCSM 1237 and 6336,
respectively) are very near the eastern edge of the escarpment, and
the sight record of burrows and mounds from West End in Moore
County (Lee 1987) is at the extreme inner edge of the Coastal Plain.

The locality for the Surry County specimen (NCSM 7744) lies
just off the Blue Ridge escarpment in the extreme western Piedmont,
and is the first specimen-supported record for that geographic province
in the state. The specimen was found dead along a bog at the edge of
a hayfield. Evidence of considerable mole activity was apparent at
this site during 1994 and 1995 (personal observation), and the landowners
reported that these moles were commonly killed by house cats on the
property, as was possibly the case with this specimen (Ann B. Somers,
personal communication).

The specimens from Ashe, Brunswick, Dare, Hoke, Jackson, and
Rutherford counties also represent new county records. The sight records
for Bladen, McDowell, Mecklenburg, Randolph, and Wake counties,
though not supported by specimens, represent previously unpublished
county records as well (see Lee et al. 1982, Webster 1987).

DISCUSSION
The range of the star-nosed mole overlaps that of the eastern
mole (Scalopus aquaticus) throughout North Carolina, and that of the
hairy-tailed mole (Parascalops breweri) in the Mountains. The hairy-

Star-nosed Mole 83

tailed mole is currently known from Avery, Buncombe, Caldwell,
Jackson, Haywood, Macon, Mitchell, Swain, Transylvania, Watauga,
Wilkes, and Yancey counties in the Mountains (Lee et al. 1982, NCSM
files). The late Joseph M. Bauman (personal communication) also
reported specimens from Cherokee County. The eastern mole apparently
occurs throughout the state (Lee et al. 1982, Webster et al, 1985),
and is in most places the most common mole.

Differences in habitat preference by the three have been noted
by several authors. Star-nosed moles are known to prefer, if not require,
wet areas as habitat (Hamilton 1931, Burt and Grossenheider 1976,
Yates and Pedersen 1982, Webster et al. 1985). Nearly all North
Carolina specimens for which habitat information is available were
collected in such areas. A possible exception is the Bladen County
specimen listed above, which was found dead in a dry, sandy area
apparently not near a wetland (T. M. Padgett, personal communication).
The eastern mole has been reported to avoid "the wet soils preferred
by the Star-nosed Mole" (Lee et al. 1982). The hairy-tailed mole
reportedly occurs in similar habitats as the eastern mole, but usually
at higher elevations, with most specimens being taken at above 2,000
ft. (610 m) (Lee et al. 1982, Webster et al. 1985, NCSM files). Lee et
al. (1982) reported the hairy-tailed mole to be "considerably more
common than the Star-nosed Mole," but this may not hold true at all
montane localities. In Watauga County, for instance, Condylura apparently
is more frequently encountered than Parascalops (R. W. Van Devender,
personal communication), and there are overall more Condylura than
Parascalops from the North Carolina mountains in the State Museum's
collections.

The ecological niches of the three moles may thus be loosely
described as Parascalops and Scalopus frequenting well-drained soils,
often with an altitudinal separation between them, and Condylura
utilizing wet, low-lying areas at a wide range of elevations. However,
some syntopy may occur. All three species may occur at some montane
localities, as is the case in the vicinity of Highlands Biological Station
in Macon County at ca. 3,900 ft. (1,190 m) in elevation (NCSM
files). In Ashe County at ca. 2,880 ft. (878 m), I found a dead eastern
mole (NCSM 7251) in a wet sedge meadow bog, a site which appeared
more suitable for a star-nosed mole. Eadie (1939) reported having
trapped Condylura and Parascalops in the same tunnel at a New
Hampshire site. Undoubtably, the distributions and interspecific zrelationships
of North Carolina's three mole species merit further study.

The possibility that Condylura cristata occurs at low densities
or at scattered localities throughout the state, and that the Mountain

84 Jeffrey C. Beane

and Coastal Plain populations are not disjunct, should not be ignored.
Although Lee et al. (1982) stated that "absence of records from the
piedmont is certainly not an artifact of collecting," there have probably
been few serious efforts to collect C. cristata in the North Carolina
Piedmont, other than the recent trapping efforts at the Wake County
site. Moreover, specific attempts to collect Condylura in the Mountains
and Coastal Plain have seldom if ever proven successful, and Clark et
al. (1985) commented on the difficulty encountered in trapping the
species. Most available specimens for which the method of collection
is known were found dead on roads or otherwise accidentally encountered.
Even in the Mountains where the species may be fairly common,
there are still several counties for which specimens have not been
reported. Much of the Piedmont, particularly the western part, has
been largely overlooked or ignored by biologists, and its fauna remains
poorly documented. Small, fossorial or otherwise secretive vertebrates
may long elude detection in any region. As examples, the eastern
tiger salamander (Ambystoma tigrinum) was first discovered in the
Piedmont in Wake County by me in 1982 (NCSM files) after that area
had been heavily collected by herpetologists for nearly a century; and
the bog turtle (Clemmys muhlenbergii) was discovered in three new
Piedmont counties during 1992-1993 (Beane 1993). Although some
areas, such as the Raleigh vicinity, have been heavily collected by
mammalogists and other biologists for more than a century, little
effort has been specifically aimed at star-nosed mole trapping.

The North Carolina Piedmont has been heavily altered for agriculture
and urbanization — probably more so overall than either the Mountains
or the Coastal Plain, and many Piedmont wetlands have been drained
or otherwise destroyed in the process. It is possible that wetlands
alteration or other human activities may have eliminated the star-
nosed mole from many areas of the Piedmont in recent times. If
populations of this mole do occur throughout the Piedmont of North
Carolina, it is likely that they exist as scattered relicts (either Pleistocene
relicts or more recent anthropogenic relicts) and at low densities. It is
hoped that this paper will help stimulate biologists working in North
Carolina (and other southern states) to make every reasonable effort
to collect evidence of Condylura, and to photograph or collect specimens
wherever they are encountered, especially at undocumented localities.

ACKNOWLEDGMENTS—^ am grateful to Allen C. Boynton, Elmer
E. Brown, Kurt A. Buhlmann, Dennis W. Herman, Elizabeth McGhee,
Thomas M. Padgett, Philip B. Spivey, R. Wayne Van Devender, W.
David Webster, and David K. Woodward for their assistance in locating

Star-nosed Mole 85

specimens and providing sight records. The discovery of the Surry
County specimen was a result of the Surry County Natural Areas
Inventory, for which Ann Berry Somers and J. Richard Everhart (among
numerous others) deserve special credit. William M. Palmer and two
anonymous reviewers provided helpful comments on earlier drafts of
the manuscript.

LITERATURE CITED

Audubon, J. J., and J. Bachman. 1851. The quadrupeds of North
America. Volume II. New York, New York.

Beane, J. C. 1993. A survey of bog turtle (Clemmys muhlenbergii)
habitat in the western Piedmont of North Carolina. Bulletin of
the Chicago Herpetological Society 28(ll):240-242.

Brimley, C. S. 1923. The star-nosed mole in eastern North Carolina.
Journal of Mammalogy 4(3): 183-184.

Brimley, C. S. 1944-1946. The mammals of North Carolina. 18
installments. Carolina Tips. Carolina Biological Supply Co., Elon
College, North Carolina.

Brown, E. E. 1979. Some snake food records from the Carolinas.
Brimleyana 1:113-124.

Burt, W. H., and R. P. Grossenheider. 1976. A field guide to the
mammals. Third edition. Houghton Mifflin Company, Boston, Massachusetts.

Clark, M. K., D. S. Lee, and J. B. Funderburg, Jr. 1985. The
mammal fauna of Carolina bays, pocosins, and associated com-
munities in North Carolina: An overview. Brimleyana 11:1-38.

Eadie, W. R. 1939. A contribution to the biology of Parascalops
breweri. Journal of Mammalogy 20(2): 150-173.

Hall, E. R. 1981. The mammals of North America. John Wiley and
Sons, New York, New York.

Hamilton, W. J. 1931. Habits of the star-nosed mole, Condylura
cristata. Journal of Mammalogy 12(4):345-355.

Johnston, D. W. 1967. Ecology and distribution of mammals of
Highlands, North Carolina. Journal of the Elisha Mitchell Scien-
tific Society 83(2):88-98.

Lee, D. S. 1987. The star-nosed mole on the Delmarva Peninsula:
zoogeographic and systematic problems of a boreal species in the
South. The Maryland Naturalist 31(2):44-57.

Lee, D. S., J. B. Funderburg, Jr., and M. K. Clark. 1982. A distri-
butional survey of North Carolina mammals. Occasional Papers
of the North Carolina Biological Survey 1982-10. North Carolina
State Museum, Raleigh.

Linzey, D. W., and A. V. Linzey. 1968. Mammals of the Great
Smoky Mountains National Park. Journal of the Elisha Mitchell
Scientific Society 84(3):384-414.

86 Jeffrey C. Beane

Odum, E. P. 1949. Small mammals of the Highlands (North Caro-
lina) Plateau. Journal of Mammalogy 30(2): 179-192.

Paradiso, J. 1959. A new star-nosed mole (Condylura) from the
southeastern United States. Proceedings of the Biological Society
of Washington 72:103-108.

Petersen, K. E., and T. L. Yates. 1980. Condylura cristata. Mamma-
lian Species 129:1-4.

Webster, W. D. 1987. Condylura cristata parva Paradiso, southern
star-nosed mole. Pages 42-43 in Endangered, threatened, and rare
fauna of North Carolina, Part I. A re-evaluation of the mammals
(M. K. Clark, editor). Occasional Papers of the North Carolina
Biological Survey 1987-3. North Carolina State Museum, Raleigh.

Webster, W. D., J. F. Parnell, and W. C. Biggs, Jr. 1985. Mam-
mals of the Carolinas, Virginia, and Maryland. The University of
North Carolina Press, Chapel Hill.

Wilson, K. A. 1954. The role of the mink and otter as muskrat
predators in northeastern North Carolina. The Journal of Wildlife
Management 18(2):199-207.

Yates, T. L., and R. J. Pedersen. 1982. Moles (Talpidae). Pages 37-
51 in Wild mammals of North America. (J. A. Chapman and G.
A. Feldhamer, editors). John Hopkins University Press, Baltimore,
Maryland.

Received 20 October 1994
Accepted 11 April 1995

Observations on North Carolina Crayfishes
(Decapoda: Cambaridae)

John E. Cooper

418 Wayne Drive

Raleigh, North Carolina 27608

AND

Alvin L. Braswell

North Carolina State Museum of Natural Sciences

P.O. Box 29555, Raleigh, North Carolina 27626

ABSTRACT — Cambarus {Tubericambarus) acanthura, Orconectes
(Procericambarus) spinosus, and an apparently undescribed species
of Orconectes (O. sp. B), are reported for the first time from
North Carolina. Six additional specimens of Orconectes (Crockerinus)
virginiensis are reported from the Chowan and Roanoke basins,
and its range in North Carolina is clarified. Orconectes (C.)
erichsonianus, for which a North Carolina locality has been
published, is deleted from the State list. Cambarus {Jugicambarus)
asperimanus is reported from the Watauga and New rivers in
Watauga County, and localities are provided that expand its
known range in the Piedmont Plateau. New locality data, distributional
clarifications, or natural history notes are provided for Cambarus
(Depressicambarus) latimanus, C (D) reduncus, Cambarus {Jugicambarus)
carolinus, C (J) dubius, C. (/.) nodosus, Cambarus (Puncticambarus)
georgiae, C (P.) parrishi, C. (P) reburrus, Procambarus (Ortmannicus)
medialis, P. (O.) pearsei, and P. (O.) plumimanus. A blue
specimen of C. latimanus is reported from the Neuse River
basin. The North Carolina crayfish fauna is correlated with
the State's major river basins and physiographic provinces.

Thanks largely to the efforts of the late Horton H. Hobbs, Jr.,
U. S. National Museum of Natural History, Smithsonian Institution,
with whom the modern era in crayfish studies essentially began, the
composition of the crayfish fauna of North Carolina is relatively well
known. As indicated in his most recent checklist of the American
crayfishes (Hobbs 1989:89), the North Carolina fauna consists of 27
described native species, a putative subspecies of one of them, and
one introduced species. Hobbs, sometimes with coworkers, described
one of the four genera, six of the 11 native subgenera, and 11 of the
27 native species known to occur in the State at the time of his
checklist.

Brimleyana 22:87-132, June 1995 87

88 J. E. Cooper and A. L. Braswell

In addition to the described species, at least four and probably
five known but undescribed species from North Carolina await
description. Hobbs and Peters (1977:8-9) mentioned Cambarus (Cambarus)
sp. A, close to Cambarus (Cambarus) bartonii Fabricius; Cambarus
(Depressicambarus) sp. B, close to Cambarus (Depressicambarus)
reduncus Hobbs; and Cambarus (Puncticambarus) sp. C. The latter,
C. (P.) sp. C, refers not to a single undescribed species but to an
undiagnosed species complex that includes Cambarus (Puncticambarus)
acuminatus Faxon, which is one of the 27 species listed for North
Carolina. In fact, however, Hobbs (1969:135) said that C. (P.)
acuminatus (sensu stricto) may be "confined to the Saluda drainage"
of South Carolina, and later (Hobbs 1989:25) added, "Even in the
Santee Cooper basin (of which the Saluda River is a tributary), . . .
more than one rather distinct 'variant' is recognizable." We see no
reason in our paper to depart from using C. (P.) sp. C for this complex
until a diagnosis has been completed. Considering that the species in
the complex range from the mountains to the coast in North Carolina,
there probably are several awaiting description. Cooper and Cooper
(1977a: 198-199) and Cooper and Ashton (1985:9) commented on the
undescribed Orconectes sp. A, and North Carolina localities for another
apparently undescribed Orconectes (herein designated O. sp. B) are
presented in our paper. One of us (JEC) is investigating a number of
other new species, but further comment on them at this time would
be premature. It is a certainty that other undescribed species await
discovery in North Carolina.

About another widespread crayfish that occurs in parts of
North Carolina, Procambarus (Ortmannicus) acutus acutus (Girard),
Hobbs (1989:64) said, "With little doubt, the populations currently
assigned to this subspecies constitute a species complex." The
complex was under study by Hobbs and Hobbs (1990:608). Hobbs
(1989:24) also said that Cambarus (Lacunicambarus) diogenes Girard,
a broadly distributed crayfish whose range includes eastern North
Carolina, "is a species complex and needs considerable attention."
Jezerinac (1993:532) "concluded that the complex consists of two
subgenera Lacunicambarus and Tubericambarus, new subgenus, and
at least five additional species or subspecies." Another extremely
variable crayfish that occurs in parts of North Carolina, C. (C.)
bartonii, is also in dire need of revision. Hobbs (1969:146) referred
to "the depauperate state of our knowledge of the relationships of
those crayfishes which are currently being tentatively designated as
C. b. bartonii . . . ." Hobbs (1989:82, 89) included Cambarus bartonii
cavatus Hay in his list of crayfishes occurring in North Carolina, but

North Carolina Crayfishes 89

North Carolina was not included in his statement of the range of this
putative subspecies (Hobbs 1989:14). We have opted to omit it as a
member of the North Carolina cambarid fauna until specific evidence
appears to support its inclusion.

Four described species are North Carolina endemics: Cambarus
{Depressicambarus) catagius Hobbs and Perkins, of the upper Cape
Fear River basin; Cambarus (Puncticambarus) reburrus Prins, of the
French Broad and Savannah basins; Procambarus (Ortmannicus)
medialis Hobbs, of the Neuse and Tar-Pamlico basins; and Procambarus
{Ortmannicus) plumimanus Hobbs and Walton, of the Northeast
Cape Fear and New (White Oak) basins, which, as we explain later,
may be expanding its range into the lower Neuse basin. The undescribed
Orconectes sp. A is another North Carolina endemic, found only in
the Neuse and Tar-Pamlico basins.

This article adds two described and one undescribed native species
to the State list, provides additional distributional and natural history
information for them and 13 other species, deletes a species for which
a North Carolina locality has been published, correlates the State's
crayfish fauna with its major hydrologic units, and summarizes the
distribution of this fauna in the State's three major physiographic
provinces.

Abbreviations used are as follows. SR = secondary road (formerly
CR for county road); NC = North Carolina state highway; US = United
States highway (an A after the number means alternate highway); I =
interstate highway; FRS = United States Forest Service Road; cntr =
center of town or city; NCSM C- = crustacean collections, North
Carolina State Museum of Natural Sciences, Raleigh; USNM = collections,
U. S. National Museum of Natural History, Smithsonian Institution,
Washington, D. C. Collector's names are given in first usage, initials
thereafter.

Cambarus {Depressicambarus) latimanus (LeConte)
Hobbs (1981:120), reporting observations made in a thesis by
J. L. Boyce (1969), stated that in Yellow River, Georgia, C. latimanus
"becomes relatively inactive during December, January, and
February," Hobbs (1981:119) also provided monthly capture data for
a large number of specimens that suggested the same thing: 77 collected
in November, five in December, 21 in January, and none in February,
as opposed to 164 in March, 1,079 in April, and relatively
high numbers in most other months. These kinds of collecting data,
however, can be more a reflection of seasonal activities of the collectors
than of the collected. If we can equate "activity" with "catchability,"

90 J. E. Cooper and A. L. Braswell

and in the case of animals taken in traps we feel justified in so
doing, C. latimanus is not inactive during winter months in eastern
North Carolina. Intensive sampling was conducted in the Neuse River
basin from late December 1978 through May 1979, and in the Tar-
Pamlico basin from January through April 1980, most of it part of a
study of the distribution and ecology of the salamander, Necturus
lewisi (Brimley) (Braswell and Ashton 1985). This effort yielded
considerable winter and spring capture ("activity") data for 409 adult
and 279 juvenile C. latimanus, the majority of them caught in
traps (Table 1).

Bouchard (1978:37) remarked on a dearth of ovigerous females
in collections of this species, saying, "Of the numerous collections of
C. latimanus that have been made, only three specimens are females
with eggs (or young)." One of these females was collected in a
tributary of the Neuse River in Wake County, North Carolina, on 16
April 1977, the other two were collected in Alabama on 22 April
1973. Hobbs (1981:119), who examined 2,424 specimens of this species
from approximately 400 localities, remarked, "I have no records of
ovigerous females or of those carrying young anywhere within its
range .... This paucity of such females in collections almost certainly
reflects inadequate sampling of burrows in the banks of streams."
Most of the smallest juveniles collected in the Neuse and Tar-Pamlico
surveys were taken in late April and May. This, combined with the
few available dates of capture of ovigerous females, Bouchard's (1978:47)
report of a copulating pair found on 2 November 1974 in Alabama,
and the fact that nearly 50 percent of the males in the January sample
reported in Table 1 were form I, leads us to suggest that in eastern
North Carolina (1) mating and egg production occur in winter and
perhaps late fall, (2) incubation occurs during winter and early spring,

Table 1. Summary of winter and spring captures of C. latimanus in
the Neuse and Tar-Pamlico river basins, 1978-1980.

Month

SI

SU

9

juv

N

Jan

46

48

81

21

196

Feb

15

20

39

46

120

Mar

14

40

50

22

126

Apr

9

19

23

168

219

May

1

2

2

22

27

Totals

85

129

195

279

688

North Carolina Crayfishes 91

and (3) the young are produced later in the spring (a time of high
water, expanded habitat, and increased availability of food). This
supports Thorp's (1978:278) statement that in the lower Savannah
River basin of South Carolina this species "enters the reproductive
period as temperatures are dropping in fall and winter."

On 17 February 1982, a blue specimen of C. latimanus was
found by a worker clearing a muddy ditch at the south end of the
Wayne Community College campus in Goldsboro, Wayne County
(Neuse River basin). It was taken to Gary W. Woodyard of the
college biology faculty, who generously donated it to the N. C. State
Museum. The animal, a form II male (NCSM C-775), was generally
cobalt blue, with the pigment obviously in the exoskeleton since the
underlying chromatophore pattern of blotches and abdominal
stripes was clearly visible. The right cheliped was missing, but the
left cheliped and all pereiopods were whitish on the ventral surfaces,
with a pale bluish tint at the base of the coxa of each limb. The
lateral margin of the palm and propodus was very light blue, but
the entire dactyl was darker except at the ventral tip. The gonopods
(first pleopods), eyestalks, antennae, antennules, and antennal scales
were pale blue. There was a small white area on the anterolateral
carapace just below the suborbital angle and along part of the margin
of the carapace. The small tubercles and punctations of the carapace
were points of white, as were the small cervical spines. Most of
the ventral abdomen was clear, but the transverse ridges between
segments were blue.

This is the first blue individual reported for this species, and
the first blue crayfish reported for any non-blue species in North
Carolina. One North Carolina crayfish, Cambarus (Jugicambarus)
dubius Faxon, is known to have a cobalt blue color morph. In the area
where the blue C. latimanus was found, the species normally is light
tan or greenish, with dark brown or green carapace markings and
abdominal stripes. Hundreds of normal-colored specimens of this
species have been collected in the Neuse River and its tributaries,
including a number from Wayne County.

Fitzpatrick (1987) summarized most of the known records for
the "blue color phase" in six crayfish genera, and discussed environ-
mental (diet and illumination) versus genetic causes of such color
variations. Penn (1951) reported a blue color variant of Procambarus
(Scapulicambarus) clarkii (Girard), a normally red species, in
Louisiana. Penn later (1959:10) said, "In recent years anomalous bright
blue specimens have appeared in several parts of the state . . . ."
(That someone may be culturing this obviously genetic variant is

92 J. E. Cooper and A. L. Braswell

indicated by the fact that in April 1984 JEC examined several bright
blue specimens of this species, said to be from "Thailand," that
were for sale as exotic novelties in a Raleigh tropical fish shop. A
form I male was purchased and is in the NCSM collections, C-1295.)

Smiley and Miller (1971:221) estimated the frequency of blue
variants in normally non-blue P. a. acutus as 1 in 50,000. Black
(1975) experimentally demonstrated that blue color in this species is
a mutation, in which the gene controlling the chemical composition
of a carotenoid pigment in the exoskeleton behaves as a single
recessive allele, with complete penetrance of the dominant normal
allele. He estimated the ratio of blue to normal crayfish in the pond
of origin of his parental stocks at 1:5,600. Anthony D'Agostino has
bred cobalt blue Homarus americanus, and the Fx offspring inherit
this coloration as a homozygous recessive trait (Porterfield 1982:38),
which is what Black found in P. a. acutus.

With a nod to William of Occam, we find it considerably more
parsimonious to conclude that the Wayne County C. latimanus was a
genetically blue individual than that its abnormal color was produced
by diet or environmental conditions.

In North Carolina, C. latimanus is a common, widespread
inhabitant of the eastern Piedmont Plateau and much of the Coastal
Plain. It is possible, though, that it also occurs in the Hiwassee River
basin of the Blue Ridge, as we discuss later.

Cambarus (Depressicambarus) reduncus Hobbs
Bouchard (1978:40) gave the range of this species as "in the
Piedmont province from the Cape Fear River drainage in North
Carolina southward to the Santee River basin . . . .," and in the latest
American checklist Hobbs (1989:16) gave it as "Piedmont Province
from Orange County, North Carolina to Richland County, South
Carolina." Concerning the northern terminus of its range, these
statements almost certainly were based on specimens from the upper
Cape Fear basin in and around Chapel Hill, Orange County, earlier
reported by Hobbs (1956:66-67) and Hobbs and Peters (1977:50).
Although Cooper and Ashton (1985:9) had reported C. reduncus from
the Neuse and Tar-Pamlico basins, north of the Cape Fear, they gave
no particulars. The following localities extend the known range of
this species north into the Piedmont headwaters of these two rivers,
very close to the Roanoke River basin.

NEUSE RIVER BASIN. Durham Co.— (1) Lick Crk at SR 1905,
10 air mi (16 air km) E Durham; 1 6 I (NCSM C-520), 27 Jan 1979,
A. P. Capparella. Granville Co. — (2) burrow in roadside ditch along

North Carolina Crayfishes 93

SR 1721, 3.5 air mi (5.6 air km) SE cntr Creedmoor, near Wake Co
line; 1 6 I (NCSM C-860), 23 Apr 1978, R. E. Ashton, Jr.; (3)
swamp at E edge Lake Rogers (on Ledge Crk), 1 air mi (1.6 air km)
NW Creedmoor; 1 j ^ 9 Mar 1991, D. G. Cooper, JEC. Wake Co.—
(4) burrow in lawn, Morrisville; 1 9 (NCSM C-907), 27 Mar 1978, J.
Clayton; (5) alive on SR 1300 (Hemlock Bluffs Rd) N of bridge; 1 6
I (NCSM C-909), night 3 Nov 1977, R. W. Laney, D. F. Lockwood;
(6) alive on road, 0.5 mi (0.8 km) W jet SR 1379 & Kildaire Rd; 1 6
I (NCSM C-910), night 3 Nov 1977, RWL, DFL. TAR-PAMLICO
RIVER BASIN. Granville Co.— (7) alive on road, jet SR 1304 &
1307, 1.8 air mi (2.9 air km) W Hebron; 1 6 II (NCSM C-1262),
night 23 Apr 1983, J. P. Kumyhr, D. Smith. Person Co.— (8) Tar R at
SR 1565, 1.7 air mi (2.7 air km) WSW Dennys Store; 1 6 I, I 9
(NCSM C-665), 20 Feb 1980, trap, E. Rawls. Vance Co.— (9) Tabbs
Crk at SR 1101, 2.3 air mi (3.7 air km) W Kittrell; 1 6 I, 1 9
(NCSM C-655), 24 Jan 1980, trap, ER.

Hobbs and Peters (1977:18) reported C. reduncus from "Mont-
gomery County: (2) Hamer Creek, 3 mi (4.8 km) N Richmond Co line
on St Rte 73," Assigning the locality to the "CATAWBA BASIN."
This locality, though, like nearly all of Montgomery County, is in
the Yadkin-Pee Dee basin. The following collections of C. reduncus,
however, are known from the Catawba basin in North Carolina:
Gaston Co. — (1) small stream (trib South Crowders Crk) near office
Crowders Mountain State Park, off SR 1125, 0.6 rd mi (0.9 rd km)
SW jet SR 1106, 5.2 air mi (8.3 air km) S Bessemer City; 1 6 II, 1
j 6, 3 9, 4 j 9 (NCSM C-2300), 22 Apr 1985, ALB. Mecklenburg
Co.— (2) plowed field along dirt rd at E end SR 3629 (Sixmile Crk
dr), ca. 6.3 air mi (10.1 air km) SE cntr Pineville; 1 6 II, 4 j 8, 1 9 ,
2j 9 (NCSM C-1043), 19 Apr 1980, from burrows with chimneys,
N. L. Elliott; (3) ditch along SR 3629 (Sixmile Crk dr), ca. 5.6 air mi
(8.9 air km) SE cntr Pineville; 9 j 8, 1 9, 11 j 9 (NCSM C-1045),
19 Apr 1980, NLE. Union Co.— (4) along SR 1624, 0.5 rd mi (0.8 rd
km) NW jet NC 200, 2.0 air mi (3.2 air km) NNE Monroe; 1 8 I
(NCSM C-311), 15 Jul 1978, J. W. Braswell, Jr., ALB; (5) East Fk
Twelve Mile Crk at SR 1329, ca. 2.1 air mi (3.4 air km) SSW Wesley
Chapel; 1 j 8, 3 j 9 (NCSM C-1031), 18 Apr 1980, in roadside ditch
fed by woodland pool; 1 j 8, 3 j 9 (NCSM C-1047), 18 Apr 1980, in
shallow, temporary floodplain pools; 1 8 II, 1 j 9 (NCSM C-1027),
26 Apr 1980, dug from burrows with chimneys in old-field, NLE; (6)
small branch Little Twelve Mile Crk at SR 1329, ca. 2.9 air mi (4.6
air km) SSW Wesley Chapel; 4 j 8, 7 j 9 (NCSM C-1037), 18 Apr
1980, NLE; (7) woodland pool off Little Twelve Mile Crk at SR 1328,

94 J. E. Cooper and A. L. Braswell

ca. 3.1 air mi (5.0 air km) SW Wesley Chapel; 3 j 8, 1 j 9 (NCSM
C-1040), 18 Apr 1980, NLE; (8) damp ditch at East Fk Twelve Mile
Crk at SR 1336, ca. 1.6 air mi (2.6 air km) SW Wesley Chapel; 1 j 8,
1 j 9 (NCSM C-1038), 20 Apr 1980, from burrows, NLE.

Cambarus (Jugicambarus) asperimanus Faxon
This crayfish occurs in the Watauga River basin in Tennessee
(Hobbs 1989:20), but no localities for the species have been reported
from this system in North Carolina. Additionally, no one unequivocally
has reported its occurrence anywhere in the New River basin. With
respect to the latter, in a discussion of "Cambarus (Cambarus)
bartoni asperimanus" Ortmann (1931:138) stated, " I have a male (I)
and two females from Blowing Rock (in the headwaters of New River,
some distance northeast of Asheville), in which the inner
margin of the palm has the tubercles somewhat more strongly develop-
ed than is usual in bartoni. This indicates a transition toward
asperimanus. However, I prefer to leave these with bartoni; the
character being much less distinct here than in the specimens from
Asheville and Canton" (Buncombe and Haywood counties, respectively,
French Broad River basin). There is one other published locality for
C. asperimanus that could be in the New River basin, but it was not
reported as such. Hobbs and Peters (1977:57) recorded the species
from "Watauga County: (8) creek at Deep Gap," but placed this locality
in the Pee Dee (Yadkin) basin. This citation does not indicate whether
"creek at Deep Gap" refers to the community of Deep Gap, or the
mountain gap of the same name. The community is in eastern Watauga
County on Deep Gap Creek, a tributary that joins South Fork New
River in Ashe County. The mountain gap lies southeast of the Blue
Ridge, in the drainage of Stony Fork Creek, a headwater tributary
that enters the Yadkin River in western Wilkes County.

On 25 July 1984, we collected C. asperimanus in both the
Watauga and New River basins at the following localities. WATAUGA
RIVER BASIN. Watauga Co.— (1) small stream entering N bank
Watauga R at NC 105 bridge, jet SR 1112, ca. 2.4 air mi (3.8 air km)
SE Valle Crucis; 2 $ (NCSM C-1811), from under separate rocks at
mouth of stream. NEW RIVER BASIN. Watauga Co.— (2) small,
shallow creek (South Fk New River dr) in hardwood ravine and
meadow on E side Howard Knob, off Hidden Valley Circle, 0.3 rd mi
(0.5 rd km) from jet Chestnut Drive in north Boone; 1 cM, 5 j 8,1 9,
1 j 9 (NCSM C-1814), 1 9 with attached young (NCSM C-1815),
3 9, 1 j 9 (NCSM C-1816), with R. W. VanDevender.

North Carolina Crayfishes 95

At the Watauga River locality, the only other crayfish collected
was Cambarus (Puncticambarus) robustus Girard, all specimens of
which were taken from the river itself. At the New River locality, the
orangish-tan C. asperimanus were "associated" with cobalt blue C.
dubius. The two species were dug from burrows within a few meters
of each other, but their habitats differed. The C. asperimanus that
came from burrows were in simple, shallow burrows at or just above
water level along the banks of the creek, while the C. dubius were
dug from more complex burrows in a mucky seepage area near the
same creek. Some C. asperimanus were found under rocks in the
creek, but no C. dubius were in this habitat, and no C. asperimanus
were dug from the seep.

The limits of the range of C. asperimanus east of the Blue Ridge
escarpment in North Carolina are not yet clear, but we made a number
of collections of the species at the base of the eastern foothills and
in the Piedmont as far east as western Catawba County. Hobbs and
Peters (1977:57) first recorded the species in this area, from a locality
in the upper Catawba basin in northwestern Burke County. Our
collections at the following new localities extend the range of C.
asperimanus farther east in the Catawba basin, and south and east
into the Broad River basin.

CATAWBA RIVER BASIN. Burke Co.— (1) Jacob Fk nr office
South Mountain State Park, 3.4 air mi (5.4 air km) SW Pleasant
Grove; 1 S I, 3 S II, 1 j S, 4 9 (NCSM C-2281), 23 Apr 1985; (2)
Laurel Crk along NC 18, 2.0 rd mi (3.2 rd km) ESE jet SR 1929, 3.2
air mi (5.1 air km) N Pleasant Grove; 2 S I, 5 jS, 6 j 9 (NCSM C-
2286), 1 ovig 9 (NCSM C-2309), 26 Apr 1985. Catawba Co.— (3)
small stream (trib Henry Fk) along I 40 at milepost 120 near Burke
Co line, ca. 4.0 air mi (6.4 air km) SW cntr Hickory; IS II, 3 j S, 1
9, 1 j 9 (NCSM C-1257), 12 Apr 1983. McDowell Co.— (4) trib
Swannanoa Crk along SR 1400, 1.3 rd mi (2.1 rd km) W jet E end
SR 1407, 2.1 air mi (3.4 air km) WNW cntr Old Fort; 2 S I, 2 S II, 1
j 6 (NCSM C-807), 20 Aug 1977; (5) trib Catawba R along US 70,
3.1 mi (5.0 km) E Buncombe Co line, 2.3 air mi (3.7 air km) WSW
cntr Old Fort; 2 9 (NCSM C-949), 24 Nov 1978, with D. L. Stephan;
(6) small cascading stream (trib Swannanoa Crk) along SR 1407, ca.
0.4 rd mi (0.6 rd km) NE Buncombe Co line, ca. 5.0 air mi (8.0 air
km) W cntr Old Fort; 1 j 6,3 9, 1 j 9 (NCSM C-1251), 12 Apr
1983; (7) Long Br (trib Mill Crk) along SR 1407, 0.4 rd mi (0.6 rd
km) S railroad crossing, 1.5 air mi (2.4 air km) SE Graphite & 3.2
air mi (5.1 air km) WNW cntr Old Fort; IS II, 1 j 9 (NCSM
C-1258), 12 Apr 1983; (8) Buck Crk at NC 80 bridge, 0.4 rd mi

96 J. E. Cooper and A. L. Braswell

(0.6 rd km) N jet SR 1437, ca. 0.2 air mi (0.3 air km) E Sunny Vale;
1 9 (NCSM C-2200), 26 Sep 1984. BROAD RIVER BASIN. Cleveland
Co. — (9) small stream (Buffalo Crk dr) in hardwoods, 0.8 rd mi
(1.3 rd km) S Lincoln Co line, 2.3 air mi (3.7 air km) NNE Belwood;
1 8 II, 3 j 8, 3 j 9 (NCSM C-2283), 22 Apr 1985. McDowell Co.—
(10) intermittent creek (trib Cane Crk) in hardwood ravine along
NC 226, 0.1 rd mi (0.2 rd km) WNW Rutherford Co line, 3.4 air mi
(5.4 air km) SE Dysartville; 4 8 II (NCSM C-987), 25 Jun 1977, 2 8
I, 5 8 II, 11 j 8, 9 9, 14 j 9 (NCSM C-1821), 1 9 with attached
young (NCSM C-1823), 1 8 II, 1 j8, 2 $ (NCSM C-1824), 26 Jul
1984. Rutherford Co.— (11) trib Broad R at NC 9, ca. 0.3 air mi
(0.5 air km) S town Lake Lure; 1 8 II (NCSM C-64), 6 Aug 1976;
(12) small stream in steep ravine at NC 9, 0.8 rd mi (1.3 rd km) W
cntr town Chimney Rock; 1 9 (NCSM C-959), 3 8 II, 1 9 (NCSM
C-960), 5 Jun 1977, with E. Messersmith. Polk Co.— (13) unnamed
trib Green R near Raccoon Mountain; 1 8 I (NCSM C-2012), summer
1974, G. G. Shaw.

We conclude, based on the apparently broad temporal distribution
of ovigerous females and those with young, that C. asperimanus
either has an extended breeding season or has a long development
time for the ova and young. Hobbs (1981:190) reported an ovigerous
female taken in Rabun County, Georgia, in April 1977, and said that
"elsewhere" such females have been taken in April, June, and
December. The latter statement may have been based on Bouchard
(1972:47), who reported ovigerous females (presumably in Tennessee)
in "April to June and December." As indicated above, we collected
a female with attached young in Watauga County on 25 July 1984,
and another a day later in McDowell County. David G. Cooper collected
a female with three attached young in Jackson County on 10 August
1993. In addition, we found a number of individual females
closely associated with groups of tiny young in Macon County on 19
September 1984, and on this same date we found free-living young of
the same size.

The Jackson County female collected by DGC on 10 August
1993 is the largest C. asperimanus we have seen, with a total carapace
length (TCL) of 42.0 mm and a postorbital carapace length (PCL) of
37.0 mm. Another very large specimen, a form I male measuring 38.0
mm TCL (32.5 mm PCL), was collected by DGC at the same site on
the same date. We collected a form I male in Macon County that
measured 36.0 mm TCL (31.5 mm PCL). The only other size data we
know of for this species are those of Hobbs (1981:190). His largest

North Carolina Crayfishes 97

Georgia specimen, an ovigerous female, measured 30.5 mm TCL (26.7
mm PCL), and his largest form I male was 27.6 mm TCL (24.1 mm
PCL).

Cambarus (Jugicambarus) carolinus (Erichson)
Although Hobbs and Bouchard (1973:42), Bouchard (1976:594),
and Hobbs (1989:21) included southwestern North Carolina, south
and west of the French Broad River, within the range of this species,
no one has yet reported precise locality or natural history data for
this primary burrower in North Carolina. Faxon (1890:624) reported a
form I male of his Cambarus dubius, collected by James Mooney
from " 'Among the Cherokees,' Indian Territory," and said, "According
to the label accompanying the specimen it is called Tsisgdgili (red
crayfish) by the Cherokee Indians." The same author later (Faxon
1914:396) included this specimen (USNM 14314) under C. carolinus,
said "I am advised by Mr. Mooney that it was in reality obtained
in Swain Co. or in Jackson Co., N. C, " and added (Faxon 1914:397)
that "The living color . . . was red . . . ." Ortmann (1931:147) reported
Faxon's specimen under "Cambarus (Cambarus) carolinus ," and
inexplicably indicated that it was from "Cherokee Co.; North Carolina."
He added, "I have seen, at Murphy, Cherokee Co., crawfish-chimneys,
and tried unsuccessfully to dig out specimens .... The owner of the
place told me, that these were red crawfish, and thus probably this
species." Ortmann (1931:147) also recorded the species from "Swampy
ground near springs, Marion, McDowell Co. . . . Swamp, Ashville
[sic], Buncombe Co., . . .," and "Blowing Rock, Watauga Co., . . . ."
These three localities are outside the range of C. carolinus as
currently understood, but the last two are within the known range of
C. dubius (sensu lato). The McDowell County locality is in the western
Piedmont Plateau, in the headwaters of the Catawba River basin,
beyond the known ranges of both C. carolinus and C. dubius. Brimley
(1938) recorded C. carolinus at "Judson," and Hobbs and Bouchard
(1973:21) called for confirmation of this record. This would have
been a reasonable place to find C. carolinus, since Judson was a
community on the Little Tennessee River in southwestern Swain
County, but it is now beneath the waters of Fontana Lake (Powell
1968:259). The reference by Harris (1903:142) to "C. carolinus" in
the Tar-Pamlico River basin was clearly in error.

We made the following collections of specimens referred to
this species at the indicated localities, all within the basin of the
Little Tennessee River: Clay Co. — (1) boggy spring seep in Riley
Cove, off unnumbered dirt road reached from US 64, 3.3 rd mi (5.3 rd

98 J. E. Cooper and A. L. Braswell

km) S jet SR 1359 & 1.2 rd mi (1.9 rd km) W jet FSR 71 (old US 64),
Natahala National Forest; 1 8 II (NCSM C-948), 16 Aug 1977, 1 8
I, 1 8 II, 5 j 8, 4 9, 3 j 9 (NCSM C-2165), 1 <? II, 1 9 (NCSM
C-2166), 21 Sep 1984. Graham Co.— (2) dug from muck in Talulah
(Tulula) Bog, along US 129, 1.4 rd mi (2.2 rd km) E jet SR 1201, 6.7
air mi (10.7 air km) SE Robbinsville; 1 j 8,2 9, 1 j 9 (NCSM
C-2290), 25 Apr 1985; (3) small boggy area near Cheoah R along
SR 1147, 0.1 rd mi (0.2 rd km) S jet US 129; 1 ovig 9, 1 j 9 (NCSM
C-2310), 26 Apr 1985. Macon Co. — (4) boggy area around flood-
plain pond by Nantahala R, off US 19, ca. 0.5 rd mi (0.8 rd km) NE
Graham Co line & 5.2 air mi (8.3 air km) NW Kyle; 1 8 II, 1 j 8, 1
9 , 1 j 9 , remains of decomposed 8 I, pair of loose chelipeds (NCSM
C-2292), 24 Apr 1985, with J. Bauman; (5) dug from sphagnum
bog and margin murky stream along Little Choga Crk on SR 1402,
3.3 rd mi (5.3 rd km) SE jet SR 1401, 3.2 air mi (5.1 air km) WSW
Aquone; 1 8 II, 1 9 (NCSM C-2287), 25 Apr 1975, with JB;
(6) under rocks in wet roadside ditch along SR 1401, 2.9 rd mi (4.6 rd
km) NE jet SR 1402, 2.3 air mi (3.7 air km) WNW Aquone; 2 j 8
(NCSM C-2289), 25 Apr 1985, with JB.

All the specimens were brick red in color, and all except those
from locality (6) were dug from burrows, usually constructed among
soil and roots in boggy areas. The bog at locality (1) was about 8 m
from a small, rocky stream, tributary to Buck Creek (Nantahala River).
Only one other crayfish, a juvenile male C. b. bartonii, was dug from
this bog. In the nearby stream, under rocks and in substrate under
rocks, we found other C. b. bartonii, and a female of an unidentified
Cambarus. The stream was about 1- to 1.5-m wide, 7- to 8-cm deep,
and had a low gradient and low velocity current. At locality (4), an
unidentified Cambarus was found under a rock in wet sand near the
bog, and at locality (5) C. b. bartonii was found in the stream but
not in the boggy area.

Cambarus (Jugicambarus) dubius Faxon
There are a number of published localities for C. dubius in North
Carolina, including recent ones in Hobbs and Peters (1977:24, 50;
1989:324). The range of the species in North Carolina, however, is
poorly known, both because it is a primary burrower and difficult to
find and collect, and because of taxonomic uncertainties. In general,
the species occurs in "northwestern North Carolina" (Hobbs 1989:
22), "north and west of the French Broad River basin" (Bouchard
1976:594). Little has been published about its natural history in North
Carolina.

North Carolina Crayfishes 99

The following collections of C. dubius are of distributional
and natural history interest. NEW RIVER BASIN. Watauga Co.—
(1) seepage area near small, shallow creek (South Fk New R dr) in
hardwood ravine and meadow on E side Howard Knob, off Hidden
Valley Circle, 0.3 rd mi (0.5 rd km) from jet Chestnut Drive, in north
Boone; 3<?II, 8jd,2$,8j$ (NCSM C-1812), 2 9 with attached
young (NCSM C-1813), 2 j 9 (NCSM C-1817), 1 6 I, 5 6 II, 3 j 6,
2 9, 5 j 9 (NCSM C-1818), 25 Jul 1984, RWV, ALB, JEC. FRENCH
BROAD RIVER BASIN. Haywood Co.— {2) bog near small stream
(trib Cold Springs Crk, Pigeon R dr), in rhododendron thicket in steep
ravine along FSR 148, Pisgah National Forest, 3.6 rd mi (5.8 rd km)
SW jet SR 1334; 1 6 I, 2 6 II, 4 j 9 (NCSM C-2192), 23 Sep 1984,
ALB, JEC. YADKIN-PEE DEE RIVER BASIN. Surry Co.— (3) Home
Crk Off SR 2072, ca. 8.3 air mi (13.3 air km) S town Pilot Mountain;
1 S II (NCSM C-95), 20 Jul 1976, R. M. Shelley, ALB; (4) open
bog Schuyler pasture, ca. 0.6 mi (1.0 km) N Low Gap, between
E side NC 89 and Gulley Crk (Fisher R dr); 1 9, 19 Aug 1994, A.
B. Somers. Wilkes Co. — (5) Stone Mountain State Park; 1 6 I
(NCSM C-1106), 16 Jul 1975, D. S. Lee, P. Hertl; (6) Hunting Crk
at SR 2428, 0.3 rd mi (0.5 rd km) S NC 115, ca. 9 air mi (14.4 air
km) SE Wilkesboro; 1 9 (NCSM C-1234), 15 Jul 1976, F. D. Scott,
M. E. Filka.

At the Watauga County site the C. dubius, cobalt blue in
color, were dug from burrows in a mucky seep near the creek. One of
the females (NCSM C-1813) had young attached to its pleopods.
Thirteen specimens of C. asperimanus also were collected at this
site, but occupied a different habitat than the C. dubius (see the C.
asperimanus account). At the Haywood County site the C. dubius,
this time of a brick red color morph, also were dug from burrows in a
boggy area, about 3 to 4 m from a small stream. The form I male, one
of the form II males, and two of the juvenile females showed consider-
able exoskeleton decalcification, and the other form II male was soft.
In the stream near this bog, eight C. b. bartonii were found under
rocks or dug from the stream substrate. The female from site (4)
in Surry County was found walking in wet grass at about 1900
hours under rainy conditions.

Cambarus (Jugicambarus) nodosus Bouchard and Hobbs

Only two specimens of this burrowing crayfish, females from

two separate localities in Cherokee County, have been reported from

North Carolina (Bouchard and Hobbs 1976:13). We collected 45 specimens

of this species from two additional localities in the Hiwassee River

100 J. E. Cooper and A. L. Braswell

basin: Cherokee Co. — (1) seepage area on slope above SR 1322 and
Shuler Crk, 0.1 rd mi (0.2 rd km) E of Tennessee state line, ca. 4.3
air mi (6.9 air km) WSW Violet; 4 6 II, 12 j 6,2 9, 8 j 9, 1 j
unsexed (NCSM C-1786), 1 6 I, 1 9 (NCSM C-1787), 21 Jul 1984;
(2) seeps and shallow intermittent water on hillside above SR 1323
and trib Shuler Crk, 2.8 rd mi (4.5 rd km) W jet SR 1324, ca. 3.3 air
mi (5.3 air km) WSW Violet; l<JI,4<?n,5jd,39,3j9 (NCSM
C-1788), 21 Jul 1984.

All specimens were dug or scraped from shallow burrows in
dark muck. Some of the burrows were submerged in seepage water,
and some specimens were found under rocks in these areas. No other
crayfishes were found with C. nodosus.

In North Carolina, C. nodosus is limited to the Hiwassee River
basin, where it appears to be the ecological equivalent of C.
asperimanus. The two species have never been reported from the
same localities or even the same river basins anywhere, and C.
asperimanus, although common and widespread in other montane
(and western Piedmont) river systems, seems to be absent from the
Hiwassee River and its tributaries.

Cambarus (Tubericambarus) acanthura Hobbs
Although known from two localities in Fannin County, Georgia
(Hobbs 1981:219-220), this crayfish has never been reported
from North Carolina. Three localities in the Hiwassee River basin
of North Carolina are now known: Cherokee Co. — (1) floodplain
pond and ditches along Nottely R off end SR 1404, ca. 1.0 mi
(1.6 km) N US 64 and 5.8 air mi (8.0 air km) SW Murphy; 1 6 I, 2
j 9, 1 carapace (NCSM C-529), 24 Nov 1978, DLS ALB, 5 j 6\ 2
j 9 (NCSM C-2291), 24 Apr 1985, JB, ALB; (2) floodplain
Nottely R at US 64 Bridge, 0.8 mi (1.3 km) E Ranger; 5 j 8, 3 j
9 (NCSM C-496), 1 6 II (NCSM C-497), Z j 6 (NCSM C-498), 1
6 I (NCSM C-499), 17 Dec 1976, FDS, ALB; (3) small creek
(trib Nottely R) in rhododendron thicket off SR 1423, ca. 0.5 mi (0.8
km) N US 64, 5.0 air mi (8.0 air km) SW Murphy; 1 j 9 (NCSM
C-1790), 22 Jul 1984, ALB, JEC.

At locality (3), the only other crayfish found in the creek was
a male II Cambarus (Puncticambarus) hiwasseensis Hobbs. Speci-
mens C-497 and C-499 were kept alive in the laboratory after their
capture in December 1976; the former died on 13 May 1980, and the
latter, which was a juvenile when collected, died a form I male on 22
May 1981. The pond at locality (1) had extensive mats of floating

North Carolina Crayfishes 101

water shield (Brasenia sp.) when we visited it on 21-22 July 1984. A
trap set there overnight yielded no crayfish.

Cambarus (Puncticambarus) georgiae Hobbs

This crayfish has been reported from only three localities, one
in Rabun County, Georgia (the type locality), and two in Macon
County, North Carolina, all in the upper Little Tennessee River basin
(Hobbs 1981:255). The second North Carolina locality was given as
"Sugar Fork River 8 mi NE of Franklin on US Hwy 64 ... 26 June
1957, E. A. Crawford." In response to a query about this site, Hobbs
(in litt.) said it should be emended to Cullasaja River, US 64 at
Gneiss, 8 mi (12.8 km) SE of Franklin. According to Powell (1968:130),
"The Cherokee work, Kul-say-gee, means 'sugar' or 'sweet,' " and at
least one part of the river apparently was known in the 19th century
as Sugar Town Creek.

We collected 39 C. georgiae at three localities in the Cullasaja
River watershed: Macon Co. — (1) Buck Crk at culvert under US
64-NC 28, 0.2 rd mi (0.3 rd km) S jet SR 1535, ca. 0.6 air mi (1.0 air
km) S Gneiss; 1 6 I, 2 9, 2 j 9 (NCSM C-2155), 20 Sep 1984; (2)
Cullasaja R at bridge, jet SR 1667 & 1653, ca. 1.0 air mi (1.6 air km)
SSE airport in Franklin; 1 j 6 (NCSM C-2161), 20 Sep 1984;
(3) Cullasaja R along US 64-NC 28, 0.2 rd mi (0.3 rd km) N jet
SR 1678, ca. 0.2 air mi (0.3 air km) S Gneiss; 6 j 6,9 j 9 (NCSM
C-2158), 3 6 I, 1 6 II, 1 j 6, 5 9 (NCSM C-2159), 4 j 6, 4 j 9
(NCSM C-2167), 21 Sep 1984; this locality is either the same as,
or very close to, the second North Carolina locality provided by
Hobbs (1981:255).

At locality (1), a single female C. asperimanus and 22 C. b.
bartonii were also taken. At site (3), we also found 29 C. b. bartonii.

On 19 and 20 September 1984, we made collections at three
localities in the Cullasaja drainage above Cullasaja Falls, at elevations
near or above 900 m (3,000 ft). At the highest of these sites we found
only a large number of C. asperimanus, and at the others we found
only 46 C. asperimanus and 51 C. b. bartonii. Thus, C. georgiae
almost certainty is absent from higher elevation, high-gradient streams
in the Cullasaja watershed.

We can add a second Georgia locality to the range of the
species: Rabun Co. — Little Tennessee R at Hwy 246, ca. 0.3 mi (0.5
km) E jet US 441, NNE of Dillard; 3 6 I, 1 j 6 (NCSM C-626), 20
Oct 1979, REA, Jr, J. Perry.

102 J. E. Cooper and A. L. Braswell

Cambarus (Puncticambarus) parrishi Hobbs
This species, which is endemic to the Hiwassee River basin,
previously was known in North Carolina from two collections made
in Clay County in 1959 and 1960 (Hobbs 1981:267). On 22
September 1984, we collected 21 specimens at two additional
sites: Clay Co.— (1) Fires Crk along FSR 340, 2.3 rd mi (3.7 rd km)
NE of end SR 1344 (which becomes FSR 340), Nantahala National
Forest, ca. 4.3 air mi (6.9 air km) NNW cntr Hayesville; 1 6 I, 2 9,
1 j 9 (NCSM C-2174); (2) trib Fires Crk (likely Rockhouse Crk)
along FSR 340A, ca. 0.8 rd mi (1.3 rd km) N jet FSR 340 and 2.9
rd mi (4.6 rd km) NE SR 1344, Nantahala National Forest; 16 1,
1 6 II, 3 j 6, 4 9, 1 j 9 (NCSM C-2175), 2 j 9, part of adult
exuvium (NCSM C-2178), 2 j 6,3] 9 (NCSM C-2180).

Cambarus parrishi was the only crayfish found at site (1), but
at site (2) nine C. b. bartonii also were collected. Of the four C.
parrishi taken at site (1), one of the females exhibited exoskeleton
decalcification and the male was very soft. At site (2), two males and
one female showed decalcification, two of the females were
soft, and part of an adult exuvium was found.

Cambarus (Puncticambarus) reburrus Prins
This species is endemic to headwater streams of the Savannah
and French Broad river basins, in Buncombe, Henderson, Jackson,
and Transylvania counties in the Blue Ridge province of North
Carolina (Cooper and Cooper 19776:214). There is some confusion,
however, about the precise distribution of the species. Its type locality
is a "Small tributary to the Horsepasture River from Sapphire
(= Fairfield) Lake off U. S. 64, 5.5 miles east of Cashiers, Jackson
County, North Carolina. Sapphire Lake is an impoundment from
which the tributary . . . flows several hundred yards into the Horse-
pasture River (upper Savannah River drainage)" (Prins 1968:459). It
should be noted that Fairfield Lake, although also on a tributary of
the Horsepasture River (Long Branch), is about 1.0 air mi (1.6 air
km) northwest of Sapphire Lake.

The type locality was the only known site for the species at
the time of its description, but four years later Prins and Hobbs (1972:
412) added the French Broad River basin to its range and suggested
that the "population described by Prins in the headwaters of the
Savannah River is an introduced one." Ross (1971:29-32), however,
had earlier discussed evidence for a probable Pleistocene breach of
the Blue Ridge by Savannah basin headwaters, and "an obvious encroach-

North Carolina Crayfishes 103

ment of Toxaway River of the Savannah River system upon the head-
waters of the French Broad River of the Tennessee River basin." Ross
further commented that "Toxaway Creek has not only breached the
Blue Ridge but it has also begun to drain some of the 2900 to 3000-
foot surface around Lake Toxaway, which once must have been part
of the French Broad basin." Later, Hobbs and Peters (1977:11) provided
evidence, based on the distributions of fishes and entocytherid
ostracods, that "the possibility of a former connection between parts
of the two basins exists." Despite these analyses, however, current
evidence indicates that Prins and Hobbs (1972:412) may have been
correct in suggesting that C. reburrus is not native to the Savannah
River basin.

The following additional collections of this species are all
from the same locality in the general area of the type locality.
Jackson Co. — pond in floodplain Horsepasture R along US 64, 1.2
rd mi (1.9 rd km) W jet SR 1119; 1 9 with young (NCSM C-890),
1 ovig 2 (NCSM C-891), 5 Jun 1977, ALB, 3 $ with young
(NCSM C-253, 258, 259), 1 j $ , 1 S II (NCSM C-260), 4 6 II, 1 $
1 j 9 (NCSM C-912), 27 Jun 1977, DLS, ALB. Several collections
made elsewhere in the Horsepasture watershed yielded only C. b.
bartonii.

At all localities where C. reburrus has been taken, it has proved
to be a creature of slack or slow-moving waters of low gradient and
generally with considerable organic debris. To our knowledge, it has
not been taken in swift, high-gradient streams devoid of detritus.
Collections we made in September 1984 in two such headwater streams
of the Chatooga River south of Highlands, Macon County, streams
that in North Carolina are independent of the more eastern head-
waters of the Savannah, produced only C. asperimanus. Suitable C.
reburrus habitat is unlikely in the lower Horsepasture and Toxaway
rivers, and in the Thompson River. It almost certainly exists, however,
in those parts of the Horsepasture watershed between Sapphire and
Fairfield lakes and the northern limits of Cashiers, in the headwaters
of the Chatooga River in and around Cashiers, in the floodplain of the
Toxaway River around Lake Toxaway above Toxaway Falls, and in
the Whitewater watershed above Whitewater Falls. We can find no
evidence that C. reburrus has been taken from any of these areas, but
perhaps more field work will reveal its presence in one or more of
them.

Although fairly common and widespread in the upper reaches
of the French Broad River basin, C. reburrus has yet to be verified
any farther downriver in this system than eastcentral Buncombe

104 J. E. Cooper and A. L. Braswell

County (see Prins and Hobbs 1972:412 for localities near Black
Mountain and Swannanoa, and the additional Buncombe County
locality listed below). Prins and Hobbs (1972:412) reported the species
from "Madison County — 3.5 mi. west of Old Fort in Pisgah National
Forest on U. S. Hwy. 70, . . . 17 June 1940, J. C. Moore, coll.," and
added, "These specimens are somewhat atypical and are thus
tentatively assigned to this species." Hobbs (1989:27) included
Madison County in the range of the species. The locality in
question, however, is in McDowell County and in the headwaters of
the Catawba River basin, not the French Broad. Old Fort is over 5 air
mi (8 air km) east of the Buncombe County line and the nearest
headwater creeks of the Swannanoa River (French Broad). The
Madison County line is some 24 air mi (38 air km) west of Old Fort,
so Madison County is in error. Hobbs very kindly loaned us the
specimens collected over 50 years ago at this site (USNM 131904; 1
j 8, 3 j $). Comparison with C. reburrus of similar sizes and sexes
showed that the questionable specimens did not belong to this
species. Hobbs later pointed out (in litt.) that they may belong to an
undescribed Puncticambarus that occurs in the Catawba River
basin, one of those subsumed under C. (P.) sp. C.

Further doubt is cast on the occurrence of C. reburrus in lower
parts of the French Broad basin by collections we made in July and
September 1984. We collected over 300 specimens at 11 sites of
various kinds in the Pisgah National Forest in Madison County, and
78 specimens at 4 sites in the Pigeon River drainage in Haywood
County, but found no C. reburrus. Our current knowledge, then,
indicates that this species is absent from the Pigeon (and probably
Nolichucky) hydrologic units of the French Broad, and from the main
French Broad and its tributaries northwest of the Asheville area.

Hobbs and Peters (1977:36) reported the ostracod, Dactylocy there
megadactylus Hart and Hart, an ectocommensal of several crayfish
species, from 37 localities in North Carolina, "in which it is confined
to the Piedmont Plateau." They inadvertently listed C. reburrus among
the ostracod's hosts, but their distribution map for D. megadactylus
(Hobbs and Peters 1977:71) showed no localities within the range of
this Blue Ridge crayfish.

So many additional collections of C. reburrus have been
made from Henderson County, particularly from the French Broad
and Mills rivers, that we see no reason to list them. The following
localities, however, are of sufficient interest to be specified: Buncombe
Co.— (1) Swannanoa R at SR 2416, 0.1 rd mi (0.2 rd km) S jet
SR 2002, ca. 1.5 air mi (2.4 air km) NW Wilson; numbers and sexes

North Carolina Crayfishes 105

not available (NCSM C-2194), 25 Sep 1984, ALB, JEC. Transylvania
Co.— (2) French Broad R at River Mile (RM) 195.8 along Wilson Rd
at head Elm Bend; no other data available, M. Ford; (3) French
Broad R at mouth Williamston Crk (RM 192.8) at US 276; 1 9
(NCSM C-1686), 23 Jun 1977, MF; (4) West Fk French Broad R near
jet SR 1309 & 1312, ca. 4.2 air mi (6.7 air km) NW Rosman; 1 9
(NCSM C-2221), 18 Oct 1984, V. Schneider; (5) Davidson R at US
276, 0.2 rd mi (0.3 rd km) SE of FSR to Pisgah Forest National Fish
Hatchery, ca. 4.3 air mi (6.9 air km) NNW cntr Brevard; 1 9, 13 Aug
1993, J. Weems.

Orconectes {Crockerinus) erichsonianus (Faxon)
Hobbs (1981:263) listed this species and C. latimanus as
associates of C. hiwasseensis at a locality in the Hiwassee River
system of North Carolina. This is the only published record for O.
erichsonianus in North Carolina. Hobbs (in litt.) said, "the locality is
1.6 miles west of the junction of U. S. Hwy. 64 on State Route 60,
Cherokee County — Nottely River, collected on June 12, 1960, by K.
W. Simonds . . . ." He added, "the whereabouts of the specimens is
a puzzle. They were never catalogued and perhaps are still shelved
with the uncatalogued material here [at the U. S. National Museum
of Natural History]."

On 21 July 1984, we collected 47 crayfish at a site that must
have been the same as, or very close to, the one in question — the
Nottely River at NC 60, 2 air mi (3.2 air km) SSW of Ranger.
This collection contained only C. b. bartonii, C. hiwasseensis, and
an unidentified Cambarus. Collections that we made on several
occasions elsewhere in the Hiwassee River basin in Cherokee and
Clay counties also failed to yield any Orconectes, and the species is
not known from this system in Georgia (Hobbs 1981:21, 287). Hobbs
later (1989:35, 84, 89) did not include North Carolina in the general
range of O. erichsonianus, and we are convinced that the species does
not occur here. It could have been extirpated from the State, but its
historic range just west of North Carolina is limited to the Appalachian
Plateau and the Ridge and Valley physiographic provinces, no
elements of which are found in North Carolina.

Orconectes {Crockerinus) virginiensis Hobbs
This is one of only two species of Orconectes known to occur
on the Atlantic versant in North Carolina, where it reaches the southern
limits of its range. It is also the only member of subgenus Crockerinus

106 J. E. Cooper and A. L. Braswell

in the State. Its type locality is a tributary of the Nottoway River,
which is a major trunk of the Meherrin River, in southeastern Virginia
(Hobbs 1951:124-125). The Meherrin River joins the Chowan River
at the Gates County, North Carolina, line. Orconectes virginiensis
originally was known in the Chowan basin of North Carolina from
five specimens collected at a single locality in Hertford County:
Cutawhiskie Creek (Swamp), a tributary of Potecasi Creek, 1 air
mi (1.6 air km) southwest of Menola (Cooper and Cooper 1977c:
215). The collection, made on 13 August 1974 by Chris Ellis, consisted
of a form I and two form II males (NCSM C-33), and two females
(NCSM C-31, 32). A sixth Chowan basin specimen, an adult female,
was collected on 24 July 1985 from a submerged log in the Meherrin
River at CR 1175 (Parkers Ferry), downstream from the confluence of
Potecasi Creek, about 3.2 air mi (5.1 air km) north-northwest of the
center of Winton, Hertford County, by David R. Lenat.

Although Hobbs (1989:38) gave the general range of O.
virginiensis as "Chowan drainage system in North Carolina and
Virginia," Cooper and Cooper (1977c:215) had reported it earlier from
the lower Roanoke River basin in Martin County. A form I male
(USNM 116979) was collected in Ready Branch, a tributary of
Sweetwater Creek, 4.5 air mi (7.2 air km) south of Williamston, on
28 March 1949 by E. C. Raney. This species is now known from the
following additional localities in the Roanoke River basin: Bertie
Co.—(l) Roanoke R at NC 45/308 at Washington Co line, 5.6 air mi
(9.0 air km) ENE cntr Sans Souci; 1 9, 2 j 9, 6 Jul 1986, DRL.
Halifax Co.— (2) Roanoke R at NC 258 at Northampton Co line,
2.9 air mi (4.6 air km) E cntr Spring Hill; 1 j 6, 9 Jul 1987, DRL.
Martin Co. — (3) Ready Br at US 17 bridge, ca. 5.5 air mi (8.8 air
km) S Williamston; 1 j 9 (NCSM C-363), 17 Jun 1980, JP.

The adult female collected at the Bertie County site was found
in a submerged log. About this locality, which is not far west of
Batchelor Bay, a brackish estuary of Albemarle Sound, Lenat (in litt.)
said that the water there was about 200 m wide, slow-moving, and
"may be slightly brackish." He added, "The shore areas have abundant
growths of water lily, with many pieces of dead wood on the bottom.
Prolific algal growths suggested some enrichment this year. The fauna
is primarily freshwater, but also contains some estuarine taxa." At the
Martin County site, a form I male and two female P. a. acutus were
collected with the O. virginiensis.

Occurrence of this species in both the Chowan and lower
Roanoke basins is not surprising, since during the Pleistocene the
Chowan "would have been a tributary of the Greater Roanoke

North Carolina Crayfishes 107

River" (Lachner and Jenkins 1971:62). Little has been reported on the
distribution and natural history of this species, especially in
North Carolina, and it remains one of the State's least known and
ostensibly rarest crayfishes.

Orconectes (Procericambarus) spinosus (Bundy)
There are no published localities for the occurrence of any
member of subgenus Procericambarus, and specifically for O. (P.)
spinosus, in North Carolina. Hobbs (1981:297) indicated its general
range as "Streams in the Coosa and Tennessee river basins in
Alabama, Georgia, North Carolina, Tennessee, and Virginia," but later
(Hobbs 1989:50, 85, 89) did not include North Carolina within the
range of the species. Prior references to Cambarus (= Orconectes)
spinosus in the Tar River (Faxon 1890:632; Harris 1903: 130, 142;
Ortmann 1905:115, 1931:87, 88), all apparently based on specimens
reported by Bundy (1877) from Rocky Mount, refer to the undescribed
Orconectes sp. A.

North Carolina specimens assigned to O. spinosus have been
collected at the following localities in the Cheoah River watershed
of the Little Tennessee River basin: Graham Co. — (1) Tulula
(Talula, Talulah) Crk at SR 1211, ca. 0.3 air mi (0.5 air km)
E Robbinsville; 1 6 II, 1 9- (NCSM C-252), 25 Nov 1978, DLS,
ALB; (2) Cheoah R just upstream from sewage treatment plant
outfall Robbinsville; 5 6 II, 3 9 (NCSM C-389), 8 Oct 1980, J. H.
Davies; (3) Tulula Crk at NC 143, 0.3 air mi (0.5 air km) E cntr
Robbinsville; 2 6 I, 2 6 II, 4 9 (NCSM C-1213), 23 Oct 1982,
RWV, ALB, D. Sever, 5 ovig 9 (NCSM C-2302), 2 8 I, 1 6 II, 19
j 6, 9 j 9 (NCSM C-2303), 25 Apr 1985, ALB; (4) Tulula Crk at
SR 1138, just N Robbinsville; 7 6 I, 1 6 II, 5 j S, 3 j 9 (NCSM
C-2306), 2 ovig 9 (NCSM C-2312), 2 ovig 9 (NCSM C-2313), 4 6
I (NCSM C-2352), 26 Apr 1985, ALB.

At locality (2), three of the form II males showed considerable
exoskeleton decalcification, and one of them and another form II
male and one female were soft. The specimens taken at locality (3)
were found under rocks in the creek, and the species appeared to be
more abundant than the only other crayfish found there, C. b. bartonii.
The two species also occurred together at locality (4).

Orconectes (Procericambarus) sp. B
A large species of Orconectes, previously considered to be
introduced Orconectes juvenilis (Hagen) [= Orconectes (Proceri-

108 J. E. Cooper and A. L. Braswell

cambarus) rusticus (Girard)] by Hobbs and Walton (1966) and
Hobbs et al. (1967), occurs in the New-Kanawha basin of Virginia
(Hobbs, in litt.). This animal also has been taken in New River
headwaters at the following North Carolina localities: Alleghany Co. —
(1) small stream entering New R just E US 21-221, ca. 0.2 air mi
(0.3 air km) S Virginia state line, 3.0 air mi (4.8 air km) NNE jet US
21 & 221 at Twin Oaks; 1 ovig 9 (NCSM C-284), 20 May 1978,
REA, Jr, JEC. Ashe Co.— {2) South Fk New R at SR 1602, ca. 0.6 air
mi (1.0 air km) SE jet US 221, 2.6 air mi (4.2 air km) ENE town
Nathans Creek & 7.8 air mi (12.4 air km) NE cntr Jefferson; 3 8 II, 2
j 8, 4 9 (NCSM C-257), 20 Jul 1978, DSL; (3) South Fk New R at
E side NC 88 bridge and along SR 1588, ca. 0.5 rd mi (0.8 rd km)
NE Orion & 4.8 air mi (7.6 air km) E West Jefferson; 2 8 I, 1 8 II
(NCSM C-276), 1 ovig 9 (NCSM C-277), 1 8 II (NCSM C-256),
21 May 1978, REA, Jr, JEC; (4) South Fk New R along CR 1566,
0.3 rd mi (0.5 rd km) S jet SR 1567; 2 ovig 9 (NCSM C-803, 804),
4 8 II (NCSM C-805), 20 May 1978, REA, Jr, JEC; (5) South Fk
New R at NC 221, ca. 1.7 air mi (2.7 air km) WSW Scottsville; 1 j 8,
2 9 (NCSM C-2358), 27 May 1985, F. Winborne, DRL; (6)
Buffalo Crk near confluence Little Buffalo Crk, ca. 2.5 mi (4.0 km)
W cntr West Jefferson; 1 j 8 (NCSM C-2355), 30 May 1985,
DRL, FW; (7) Dog Crk at SR 1592, 4.0 mi (6.4 km) E Jefferson; 1 j
9 (NCSM C-508), 6 Aug 1978, J. R. Clamp, M. Dennis.

This species was very common at locality (3), where it occurred
in close association with another large crayfish, Cambarus {Hiati-
cambarus) chasmodactylus James. The two often were found under
the same cover, usually medium-size to large rocks in cold, fast-
moving water, and in the same gravel riffles. Both species also
occurred together at locality (2). At locality (4), the Orconectes was
collected with C. chasmodactylus and C. robustus.

Procambarus (Ortmannicus) medialis Hobbs
Procambarus {Ortmannicus) pearsei (Creaser)
Procambarus {Ortmannicus) plumimanus Hobbs and Walton
These three species comprise a "disjunct" enclave at the northern
periphery of the range of the Planirostris Group of the genus
Procambarus, a fact that is of considerable importance in under-
standing the evolutionary history of the group. Hobbs (1975:15)
summarized their distributions: "The Neuse River basin appears to
mark the southern limit of the range of P. medialis, the northern
[Northeast] Cape Fear River basin marks the southwestern limit of
the range of P. plumimanus, and the range of P. pearsei encom-

North Carolina Crayfishes 109

passes most of the coastal plain lying in the Cape Fear River basin
(excluding the Northern [Northeast] Cape Fear) southward through
the Little Pee Dee River basin." The following records and discussions
support this statement, slightly expand and clarify the known ranges
of these species, add some information on their natural history, and
help explain a distributional anomaly for P. plumimanus.

Procambarus medialis

As mentioned by Cooper and Ashton (1985:10), although the
type locality of P. medialis is in the Tar-Pamlico basin in Halifax
County (Hobbs 1975:13), this and one other site near it in the drainage
of Deep Creek are the only places within this basin where the species
currently is known to occur. All other collection sites are within the
Neuse River basin. Eleven specimens were captured in wire minnow
traps at three localities in the Neuse basin during the N. lewisi study
(Braswell and Ashton 1985), and one of them was in the mainstem
Neuse River in Lenoir County. Identical traps were employed at over
180 stations in the Tar-Pamlico basin, from its headwaters in Person
County to central and southern Beaufort County, but yielded no
additional P. medialis. One of these stations was in Beech Swamp,
less than 5 air mi (8 air km) west of Deep Creek, and another was
in the Tar River near its confluence with Deep Creek in Edgecombe
County. There are no obvious ecological, edaphic, or physiographic
reasons, however, for this apparent severe restriction of the known
range of P. medialis in the Tar-Pamlico basin. During the N. lewisi
survey, little collecting was done in roadside ditches or isolated lentic
waters, and no digging of burrows was attempted. More thorough
sampling of these possibly preferred habitats might reveal a wider
distribution pattern for P. medialis in this river basin, although it
also inhabits larger, flowing waters in the Neuse basin.

The following additional localities are in the Neuse River basin:
Johnston Co.— {!) Black Crk at NC 50, 5.5 air mi (8.8 air km) NNE
Benson; 1 9 (NCSM C-144), 27 Jan 1979, trap, K. Everett; 1 9
(NCSM C-230), 30 Jan 1979, trap, KE; 1 9 (NCSM C-140), 10 Feb
1979, trap, KE; (2) Hannah Crk at US 701, 10 air mi (16 air km)
E Benson; 4 9 (NCSM C-1079), 22 Apr 1979, trap, P. S. Freed, 1 6
II, 1 9, 1 j 9 (NCSM C-1097), 23 Apr 1979, trap, PSF; (3) dry
roadside ditch, Truck Lane and East Rose St, Smithfield; 1 c? I, 1
6 II, 1 9 (NCSM C-2270), 17 Apr 1985, W. M. Palmer, ALB.
Lenoir Co.— (4) Neuse R at NC 55, 1.1 rd mi (1.8 rd km) E jet SR
1809, 5.2 air mi (8.3 air km) NE Kinston; 1 6 II (NCSM C-231), 29
Jan 1979, trap, PSF.

110 J. E. Cooper and A. L. Braswell

At locality (3), each of the three specimens was found under
a separate railroad crosstie along the dry ditch, near entrances to
burrows that could be seen to reach water. The ditch normally
contains water, but this area was in the midst of a prolonged drought
that had been broken only by some rainfall on the previous night.
Other ditches had some water. In captivity, the form I male, collected
in April, molted and died on 29 September 1985, but remained
form I.

Procambarus pearsei

Hobbs (1975:14) mapped 22 unspecified localities for P. pearsei
in the Cape Fear, Lumber, and Waccamaw basins in Bladen, Brunswick,
Columbus, Cumberland, Robeson, and Sampson counties. This same
map inadvertently indicated an additional locality for P. pearsei in
the Neuse River drainage of Johnston County. This should have been
for P. medialis; the position of the dot corresponds to a locality for
this species given by Hobbs (1975:13), ". . . 5.3 miles south of
Smithfield on U. S. Highway 706." The nearest headwater stream of
the Cape Fear basin, Mingo Swamp, rises at the extreme western edge
of Johnston County, about 2.5 air mi (4.0 air km) SSW of site (1)
above. Thus, the range of P. pearsei given by Hobbs (1989:70) as
"Johnston and Sampson counties, North Carolina, south to . . . .,"
currently requires the deletion of Johnston County.

The following localities add P. pearsei to the Cape Fear basin in
Bladen County, and extend the known range slightly west in the
Lumber River basin into Hoke and Scotland counties. CAPE FEAR
RIVER BASIN. Bladen Co.— (I) pond along SR 1327, 1.6 rd mi (2.6
rd km) NW jet SR 1325, 5.7 air mi (9.1 air km) W Ammon, in
vicinity Little Singletary and Horseshoe (Suggs Mill Pond) lakes; 1 6
I, 1 j 6 (NCSM C-304), 17 Mar 1979, P. S. Ashton, REA, Jr., ALB;
(2) Horseshoe Lake, 0.7 rd mi (1.1 rd km) NE SR 1327; 1 6 I, 8 j 6,
1 9, 6 j 9 (NCSM C-1602), 23 Feb 1980, PSA, REA, Jr, NCSM
Jr Curators. LUMBER RIVER BASIN. Hoke Co.— (3) Antioch Bay
off NC 211, 1.7 air mi (2.7 air km) SSE Antioch; 4 8 I, 3 9, parts of
9 (NCSM C-2240), 1 Feb 1985, RWL, DLS, DFL; (4) pools in
powerline cut Antioch Church Bay, E side NC 211, 0.6 rd mi (1.0 rd
km) SSE jet SR 1447, 2 air mi (3.2 air km) SSE Antioch; 1 6 II, 2
9 (NCSM C-2241), 23 Feb 1985, DLS, RWL, DFL. Scotland Co.—
(5) borrow pit near jet SR 1400 & 1413, ca. 3.5 air mi (5.6 air km)
NW Wagram; 1 S I, 1 j 6\ 2 j 9 (NCSM C-2011), 8 Oct 1974,
MRC, WMP, ALB, JEC.

North Carolina Crayfishes 111

The remains of a form I male and at least one other P. pearsei
(NCSM C-1277) were removed from the stomach of a redtailed hawk,
Buteo jamaicensis, found dead near Hallsboro, Columbus County, on
8 April 1983, and prepared as a museum specimen by Gilbert Grant.
Seven females carrying late-instar young (NCSM C-2242-2244, 2246-
2249) were found at a site in Robeson County on 23 February 1985
by DLS, RWL, and DFL. A female (NCSM C-1951) was found alive
on NC 211, about 3 air mi (4.8 air km) south of Bolton, Columbus
County, at night on 24 March 1975.

Procambarus plumimanus

Hobbs and Walton (1958:10-11) provided a locality for P.
plumimanus in Craven County and two in Duplin County, and Hobbs
(1975:14) mapped two localities in Duplin County, three in Craven
County, one in Carteret County, and one apparently on the Craven-
Carteret line. Hobbs (1989:70) added Jones County to the range of
the species, but gave no specifics since this was a general checklist.
The following localities add to our knowledge of the distribution
of this crayfish. NORTHEAST CAPE FEAR RIVER BASIN. Pender
Co. — (1) borrow pit "Back Island," Holly Shelter Game Lands, 0.3 mi
(0.5 km) N Lodge Rd; 1 6 II, 4 j 6,1 9, 1 j 9 (NCSM C-870), 4 j
8,5 j 9 (NCSM C-871), 3 Oct 1981, REA, Jr; (2) borrow pit Lodge
Rd, Holly Shelter Game Lands, ca. 2 air mi (3.2 air km) NW US 17,
5.8 air mi (9.3 air km) SW Edgecombe; 3 6 I, 2 j 6\ 1 9 (NCSM C-
872), 24 Jul 1981, REA, Jr; (3) dr of Harrisons Crk off NC 210, ca. 1
rd mi (1.6 rd km) N jet SR 1574, 4.2 air mi (6.7 air km) NW
Hampstead; 1 6 I, 1 j 8,2 9 (NCSM C-933), 13 Mar 1977, E.
Flowers, DLS. NEW (WHITE OAK) RIVER BASIN. Carteret Co.—
(4) borrow pit along SR 1125, 5.3 mi (8.5 km) NW Newport
(vicinity Northwest Prong Newport R); 1 9 (NCSM C-1942), 25
Mar 1975, MRC, JEC; (5) slough along Millis Rd (SR 1112
extended), 0.7 mi (1.1 km) W jet SR 1124, 5.5 air mi (8.8 air km)
WSW Newport (Jason Br of Southwest Prong Newport R); 2 6 \,
1 6 II, 1 j 6,1 9, 1 j 9 (NCSM C-2097), 25 Mar 1975, MRC,
ALB, JEC. Jones Co.— (6) drainage ditch along FSR 152 (Black
Swamp Rd), ca. 7 mi (11.2 km) ESE Maysville, 1 8 I (NCSM C-
294), 7 Apr 1979, DSL; (7) slough on Great Lake Rd (SR 1101
extended as FSR 126), ca. 10 air mi (16 air km) SE Maysville
(Hunter Crk dr); 2 6 I, 1 6 II, 1 j 6, 1 j 9 (NCSM C-2069), 26 Mar
1975, MRC, JEC. New Hanover Co. — (8) cypress pond at Carolina
Beach State Park; 1 6 I, 1 9 (NCSM C-850), 3 May 1980, PSA,

112 J. E. Cooper and A. L. Braswell

REA, Jr; this locality is on a narrow peninsula between the Cape Fear
River and Onslow Bay. Onslow Co. — (9) permanent ditch and pools
along US 17, 0.4 mi (0.6 km) N jet SR 1103, ca. 3.7 air mi (5.9 air
km) S Verona; 1 6 I, 1 6 II, 4 $ (NCSM C-70), 10 Aug 1976, M.
M. Browne.

Although the natural range of P. plumimanus appears to be
limited to the Northeast Cape Fear and New (White Oak) basins, and
this species did not appear at any of the N. lewisi survey stations
sampled in the Neuse basin (Cooper and Ashton 1985:10), two
localities for the species within the lower Neuse basin are known.
The type locality, "Roadside ditch 2.2 miles southeast of Havelock,
Craven County, North Carolina on Hwy. 70" (Hobbs and Walton 1958:
10) apparently lies close to East Prong Slocum Creek and not far
from the head of Hancock Creek, both north-flowing streams that
drain into the Neuse River on either side of the Cherry Point Naval
Reservation. Only swamps separate this area from nearby headwaters
of the Newport River not far to the south. The second locality, also in
the Neuse basin in Craven County, is a ditch along Catfish Lake Road
(SR 1100), 3.0 air mi (4.8 air km) southwest of the town of Croatan,
in the drainage of East Prong Brice Creek.

This apparent expansion of the range of P. plumimanus into the
lower Neuse basin possibly can be explained by the fact that in
Carteret, southern and eastern Craven, and southern Jones counties,
as in much of the poorly drained outer (tidewater) Coastal Plain,
contemporary drainage distinctions have become blurred. The drainage
divides, very low to begin with, have been breached by stream
channelization, a vast system of man-made drainage canals, and the
Intracoastal Waterway. Drainage canals along SR 1100 east and south-
east of Catfish Lake provide access from the White Oak River basin
to West Prong Brice Creek of the Neuse basin. Farther east, the
Harlowe (Clubfoot), Adams Creek, and other canals, link elements of
the Neuse River with the Newport and North rivers. Precisely what
roles the easternmost of these interbasin waterways might play in
crayfish distributions will not become clear until we have an under-
standing of the tolerance that various species display for the physico-
chemical features of their waters. Some crayfishes, notably P. a. acutus
and Fallicambarus (Creaserinus) fodiens (Cottle), and even C. diogenes,
are widely distributed in tidewater North Carolina. They often occur
in close proximity to saline estuaries and tidal creeks, and there is a
population of F. fodiens across Croatan Sound on Roanoke Island.
Interbasin connectors also are present in other coastal river basins
(see below).

North Carolina Crayfishes 113

THE HYDROLOGIC UNITS AND THEIR CRAYFISH FAUNAS

Seventeen major river basins generally are recognized within
North Carolina (Heath et al. 1975:152). All of them include smaller
hydrologic units, some of which are autonomous drainage systems
within the State. All but five of the major river basins lie within the
huge drainage of the Atlantic Ocean, and flow generally east and
south to empty into broad, saline estuaries and sounds of the ocean.
The exceptions are (1) the Hiwassee, Little Tennessee, French Broad,
and Watauga rivers, which drain north and west into the Tennessee
River, and (2) the New River, which drains north and west via the
Kanawha River into the Ohio River. These western North Carolina
waters ultimately flow into the Mississippi River drainage of the Gulf
of Mexico. In North Carolina, the west-flowing and east-flowing
montane headwaters of river systems are separated by the Appalach-
ian (Eastern Continental) Drainage Divide, represented in the State
by the Blue Ridge physiographic province.

The Waccamaw River and its tributaries, along with Lake
Waccamaw, are recognized as a hydrologically and faunistically
distinctive drainage unit by some biologists (see Bailey 1977:
269, 273; Shute et al. 1981:18-22), and are so treated here. Another
hydrologic unit, the Northeast Cape Fear River, probably also should
be considered an autonomous drainage system, because it does not
join the Cape Fear River until it reaches the estuary at Wilmington.
In addition, the Northeast Cape Fear is faunistically distinct, apparently
lacking over a dozen species of lowland freshwater fishes found
in the Coastal Plain portions of the Cape Fear itself (Rohde et
al. 1979:114-115: Menhinick 1991). The Northeast Cape Fear also
lacks two crayfish species found in the lower Cape Fear, but has one
species that is absent from the Cape Fear (see below). Historically as
well as faunistically, the Northeast Cape Fear appears to have more
in common with the New (White Oak) drainage system than with the
Cape Fear. During the Pleistocene, when sea level attained its
highest stand (as indicated by the Surry Scarp, which marks the marine
terrace formed by the Wicomico Sea), much of the southeastern
Coastal Plain of North Carolina was covered by the ocean (Rohde
et al. 1979:113, 116). This inundated area included today's New
(White Oak) River drainage, most of the Northeast Cape Fear, and
part of the lower Cape Fear. The early coastal Cape Fear and North-
east Cape Fear basins were at that time broadly separated, and they
are still separate systems today.

114 J. E. Cooper and A. L. Braswell

Tennessee-Ohio-Mississippi

Hiwassee River — The Hiwassee River system rises in the Blue
Ridge of northcentral Georgia, and in Clay and Cherokee counties,
North Carolina. It flows generally north and west across the southwest
corner of the State and into Tennessee, where it merges with the
Tennessee River. Two streams of this system head in the southwest
corner of Cherokee County and flow south into the Toccoa River in
Georgia, part of the Ocoee River system. The Ocoee then joins the
Hiwassee in Tennessee.

Cambarus hiwasseensis and C. parrishi are upper Hiwassee
endemics, occurring in both Georgia and North Carolina. The type
locality of C. hiwasseensis is a tributary to Peachtree Creek, 0.8 mi
(1.3 km) north of Peachtree School on US 64A, Cherokee County,
North Carolina (Hobbs 1981:260-261). Other species that occur in
the Hiwassee system in North Carolina but have broader ranges are C.
b. bartonii, C. nodosus, and C. acanthura. The latter two species
are known in North Carolina only from the Hiwassee. Hobbs (1981:
20, 22) reported C. latimanus in this system in Georgia, and said
(Hobbs 1981:263) that it was found with C. hiwasseensis and O.
erichsonianus at a locality on the Nottely River in North Carolina. In
addition, his range map for C. latimanus (Hobbs 1981:115) showed
Georgia localities west of Nottely Lake in Union County, and in
Fannin County just west of the North Carolina state line. Further-
more, Bouchard (1972:36) recorded the species "downstream in the
Ocoee (Tennessee) River drainage to the Little Frog Mountains in
Polk County, Tennessee." This area is in the Hiwassee basin just west
(downriver) of North Carolina. Thus, C. latimanus, which in
North Carolina is a species of the eastern Piedmont Plateau and the
Coastal Plain, also may occur in the Hiwassee basin of the Blue
Ridge in at least Cherokee County. It did not turn up, however, in
our Hiwassee field work, which included the Nottely River. Hobbs
and Peters (1977:27) cited a record for Cambarus (Hiaticambarus)
longirostris Faxon in the Hiwassee basin in Cherokee County, "Valley
River at Andrews . . . (Crawford 1961:244)," and Hobbs (1981:171-
172) found this species in the headwaters of the Nottely River in
Union County, Georgia, which is a considerable distance upriver from
North Carolina. The distribution maps in James (1966:12-13), however,
showed no localities for this species within the Hiwassee basin in
North Carolina. In July and September 1984, we failed to collect
this crayfish at any site within the system, including the Valley
River near Andrews. Nevertheless, C. longirostris could occur in the
Hiwassee basin in North Carolina. Cambarus carolinus also may occur

North Carolina Crayfishes 115

there, but specific records are lacking. There is a small, forest-green
Cambarus in Cherokee County that is as yet unidentified.

Little Tennessee River — The main artery of the Little Tennessee
River originates in the Blue Ridge of Rabun County, Georgia. Its
tributaries drain most of Macon County, and parts of Swain and
Graham counties, North Carolina. Its major eastern tributary, the
Tuckasegee River, heads in Jackson County and flows northwest into
Fontana Lake. Another eastern tributary, the Cullasaja River, begins
in southeastern Macon County and joins the Little Tennessee River
south of Franklin. A western tributary, the Cheoah River, heads in
southern Graham County and flows northwest through Santeetlah
Lake, to merge with the Little Tennessee River at Cheoah Dam near
the state line. A second major western tributary, the Nantahala
River, rises in western Macon County and along the eastern border of
Clay County, and flows primarily north into Fontana Lake. Beyond
this lake the Little Tennessee flows west into Tennessee, joining the
Tennessee River near Lenoir City.

One crayfish, C. georgiae, is endemic to the Little Tennessee,
where currently it is known from two sites along the main river in
Rabun County, Georgia, and several lower elevation sites in the Cullasaja
River watershed of North Carolina. More wide-ranging species of this
system are C. b. bartonii, C. longirostris, and C. asperimanus. Specific
localities for C. carolinus are reported here, and Hobbs and Peters
(1977:56) referred to a C. (J.) sp. in Jackson County (Tuckasegee
watershed), which may turn out to be C. carolinus or an undescribed
species. Orconectes spinosus is herein reported from the Cheoah
River watershed of this basin.

French Broad River — The uppermost tributaries of the French
Broad River rise on the western slopes of the Blue Ridge in Transylvania,
Henderson, and Buncombe counties. Its major western tributary, the
Pigeon River, heads in western Haywood County and is an autonomous
unit in North Carolina. However, it joins the French Broad near Newport,
Cocke County, Tennessee. The main eastern tributaries, the Cane,
Nolichucky, and Toe rivers, rise in Avery, Mitchell, and Yancey counties.
They, too, form an independent unit that flows northwest and enters
Tennessee as the Nolichucky. This river then turns west and merges
with the French Broad at the upper end of Douglas Lake. The French
Broad River initially flows northeast in North Carolina, then turns
northwest and enters Tennessee about 5.5 air mi (8.8 air km) downriver
from Hot Springs, Madison County. It joins the Holston River at
Knoxville to form the Tennessee River.

116 J. E. Cooper and A. L. Braswell

The endemic North Carolina crayfish, C. reburrus, is distributed
throughout the upper French Broad, but appears to be absent from the
Pigeon (and probably Nolichucky) watersheds, and from the main
French Broad system northwest of Asheville. The other species of
this river basin are C. b. bartonii, C. longirostris, C. asperimanus, C.
dubius, C. robustus (Hobbs and Peters 1977:30), and C. sp. C. The
type locality of C. asperimanus is Flat Creek, Montreat, Buncombe
County (Faxon 1914:391). We collected many specimens of what may
be an undescribed Puncticambarus in three streams in Madison County,
but for now these are assigned to C. sp. C. Bouchard (1978:36) reported
a specimen of C. latimanus from the French Broad in North Carolina,
extralimital to any part of the known range of the species, but considered
it probably an introduction.

Watauga River — The Watauga River rises in western Watauga
County west of Grandfather Mountain and flows north and west into
Tennessee. A southwestern tributary, the Elk River, heads in
northern Avery County. It is a minor independent subdrainage in
North Carolina, but joins the Watauga in Tennessee. The Watauga is
impounded just beyond North Carolina's borders,' but eventually
merges with the Holston River of the Tennessee River system.

The crayfish fauna of the Watauga River basin in North Carolina
is not well known, but consists of C. b. bartonii, C. longirostris, C.
asperimanus, C. dubius, and C. robustus.

New River — The two major western tributaries of New River —
North Fork New River and South Fork New River — head in Ashe and
Watauga counties, respectively. They flow northeast and merge at the
northern Ashe-Alleghany county line to form the New River just before
it enters Virginia. Eastern tributaries, Little River and Brush Creek,
rise in central and eastern Alleghany County, and flow northward
into Virginia to join the New River. The New River merges with the
Kanawha River near Charleston, West Virginia.

Cambarus chasmodactylus and probably Orconectes sp. B are
endemic to the New-Kanawha system, and in North Carolina they are
limited to, and close associates in, fast-flowing tributaries of the
New River. Records for these common species in the eastern tributaries
in Alleghany County, however, are sparse. The other members of the
New River crayfish fauna in North Carolina are C. b. bartonii, C.
asperimanus (Watauga County), C. dubius, and C. robustus.
Atlantic

Savannah River — The Savannah River begins as fast-flowing
streams in deep ravines on the eastern slopes of the Blue Ridge
in southeastern Macon, southern Jackson, and southwestern

North Carolina Crayfishes 117

Transylania counties. In North Carolina, its easternmost tributaries —
the Toxaway, Horsepasture, Thompson, and Whitewater rivers — flow
south as independent streams, entering Lake Jocassee and the Keowee
River system of the Savannah basin not far into South Carolina. West
of these rivers, the Chatooga River and several even more western
tributaries of that river head in Macon County and flow independently
south. The Chatooga River forms part of the upper boundary between
Rabun County, Georgia, and Oconee County, South Carolina. The
Tallulah River, which rises in the southeastern corner of Clay County
on the slopes of Standing Indian Mountin, flows south into Towns
and Rabun counties, Georgia, where it joins the Tugaloo River of
the upper Savannah system.

The crayfish fauna of the Savannah River basin in North Carolina
consists of the endemic C. reburrus (currently known in this system
only from the floodplain of the Horsepasture River in Jackson County),
C. b. bartonii, C. asperimanus, and Cambarus sp. (Hobbs and Peters
1977:38). The type locality of C. reburrus is in the Savannah basin
(see the preceding account for this species). Little is known of
the crayfish fauna of the Tallulah River in North Carolina, but
C. asperimanus and C. b. bartonii undoubtedly occur there.

Broad River — This system rises in the west as headwater streams
in southern and eastern Henderson, western Polk, and eastern
Buncombe counties, and in the north in southern McDowell and
northern Rutherford and Cleveland counties. A western tributary, the
North Pacolet River, heads in southern Polk and Henderson counties,
and flows independently into the Pacolet River of South Carolina.
The Pacolet and Broad rivers merge at the juncture of Cherokee,
Union, and Chester counties, South Carolina. Eastern tributaries
head in eastern Cleveland and western Lincoln and Gaston counties,
flow independently into South Carolina, then soon join the Broad
River at the line between Cherokee and York counties. The Broad
flows through the eastern foothills and western Piedmont of North
Carolina, across South Carolina, and merges with the Saluda River at
Columbia to form the Congaree River. The Congaree joins the Wateree
River (continuation of the Catawba River), and the Wateree flows
into the Santee system via Lake Marion.

The crayfish fauna of the Broad River basin in North Carolina
consists of C. b. bartonii (the eastern and southern limits of its
range here are unclear, but it is known from Cane Creek, McDowell
County, and Green River, Henderson County), C. sp. A (Hobbs and
Peters 1977:52), C. asperimanus (in southeastern McDowell and
northeastern Cleveland counties), and C. sp. C (which is probably

118 J. E. Cooper and A. L. Braswell

throughout the system). The ubiquitous P. a. acutus may turn up in
the eastern reaches of the basin, and it is not inconceivable that
exhaustive effort could reveal C. carolinus along the border with
South Carolina. The latter species occurs in the upper Tyger River
watershed of the Broad-Congaree basin, in Greenville and Pickens
counties, South Carolina (Hobbs and Bouchard 1973:60-61). A juvenile
male crayfish (NCSM C-2229) that strongly resembles Cambarus
(Puncticambarus) spicatus Hobbs was collected in the North Pacolet
River of the lower Broad basin in Polk County, but it would be
premature to add this species to the North Carolina fauna on the basis
of this specimen. The species is known with certainty only from a
tributary of the Broad River in Fairfield and Richland counties,
South Carolina (Hobbs 1989:27).

Catawba River — This heavily impounded system rises in North
Carolina, with its headwaters on the Blue Ridge escarpment in south-
eastern Avery, Caldwell, McDowell, and Burke counties. The streams
flow into and through the Piedmont Plateau. Tributaries that head in
southern Mecklenburg and western Union counties flow independently
south-southwest into South Carolina, where they join the Catawba
River in York County. The Catawba flows south into South Carolina,
where it enters the Wateree basin of the Santee drainage.

Cambarus b. bartonii and C. asperimanus appear to be limited
to the foothills and upper Piedmont sections of the Catawba River
basin. The former may occur as far south and east as Catawba County
at the Alexander County line (Hobbs and Peters 1977:45-46), unless
the population there is revealed to be C. sp. A. Cambarus asperimanus
has been found in streams in Burke, Catawba, and McDowell counties.
Cambarus dubius has been reported from Avery County in the
upper Catawba basin (Hobbs and Peters 1977:24). Cambarus sp.
A and C. sp. C may occur throughout the system, and C. sp. B
currently is known only from Catawba County. Cambarus reduncus is
known from Gaston, Mecklenburg, and Union counties. Procambarus
a. acutus has been collected in Lincoln and Gaston counties. Hobbs
and Peters (1977:45) recorded Cambarus (Hiaticambarus) longulus
from a locality in the Catawba basin in northern Caldwell County,
but there is evidence that the specimens from that site either came
from a tributary of the nearby Yadkin-Pee Dee River, or belong to
what appears to be an undescribed species under investigation by
JEC. James (1966:13) did not show C. longulus in the Catawba basin,
and Hobbs later (1989:20) wrote that it ranges "south to the Yadkin
basin in North Carolina . . . ."

North Carolina Crayfishes 119

Yadkin-Pee Dee River — The headwaters of the Yadkin-Pee
Dee basin rise in the eastern Blue Ridge of Virginia, in western
Surry, Wilkes, and southeastern Watauga counties, North Carolina,
and in the western Piedmont in southwestern Stokes and western
Forsyth counties, North Carolina. The main trunk of the Yadkin River
begins in Watauga County east of Blowing Rock, flows southeast to
northcentral Caldwell County, then makes an abrupt turn and flows
northeast to the juncture of Surry, Stokes, Yadkin, and Forsyth
counties. There it turns south and, at the Stanly-Montgomery county
line southeast of Badin Lake, joins the Uwharrie River, a major eastern
tributary that rises in northwestern Randolph County. This merger
forms the Pee Dee River. A major western tributary, the Rocky River,
heads in southern Iredell and northern Mecklenburg counties, flows
east, and joins the Pee Dee at the Anson County line below Lake
Tillery. Several creeks that drain southcentral Union County flow
south directly into the Lynches River, South Carolina, a major
western trunk of the Pee Dee River. Southeastern tributaries of the
Pee Dee drain southern Anson and Richmond counties, flowing
directly into South Carolina to join the Pee Dee River in Chesterfield
County. The Pee Dee flows across northern South Carolina and
merges with the Little Pee Dee River at the tip of Britton Neck. The
Pee Dee then continues into Winyah Bay.

In the uppermost sections of the Yadkin-Pee Dee basin occur
C. asperimanus, C. longulus, and C. dubius. The southern limits of
their distributions are unknown, but all three appear to be restricted
to higher elevations within this basin. Specimens tentatively
identified as C. b. bartonii have also been collected in the upper
reaches of this basin. Cambarus reduncus occurs in the Piedmont,
and C. sp. C apparently occurs throughout. Hobbs and Peters (1977:
18) reported C. sp. A from Mongtomery County, collected with C. sp.
C and P. a. acutus, inadvertently placing the locality ("creek, 4.6
mi . . . W Mt Gilead on SR 73") in the Catawba River basin. Holt
and Weigl (1979:24) described the ectocommensal branchiobdellid,
Xironodrilus bashaviae, from several creeks in Forsyth County, includ-
ing "Bashavia" (Barshavia = Bechewa) Creek, with C. b. bartonii its
host. This crayfish could turn out to be C. sp. A. Procambarus a.
acutus has been collected in Anson, Cabarrus, Davidson, Montgomery,
and Union counties. Fallicambarus fodiens and C. diogenes may occur
in that part of the basin draining the southeastern Piedmont and
Sandhills.

Lumber River — To the west this Coastal Plain system heads in
the Sandhills of southern Moore, eastern Montgomery, and Richmond,

120 J. E. Cooper and A. L. Braswell

Scotland, and Hoke counties. Some of the streams drain directly into
the Little Pee Dee River just across the South Carolina line. Eastern
tributaries rise in western Bladen and Columbus counties. The Lumber
River flows primarily east and south into South Carolina, where it
joins the Little Pee Dee River on the Marion-Horry county line south
of Nichols.

The crayfish fauna of the Lumber basin consists of C. diogenes,
C. sp. C, F. fodiens, P. a. acutus, Procambarus (Ortmannicus)
ancylus Hobbs, Procambarus {Ortmannicus) blandingii (Harlan), and
P. pearsei. A locality for the latter species, given as "CAPE FEAR
BASIN. Bladen County: (2) 3.7 mi (6 km) S Dublin on St Rte 41"
(Hobbs and Peters 1977:52), actually is in the Lumber River basin.
An unidentified Procambarus, currently represented in collections
by only juveniles and one female, has been found in the Lumber
River basin in Robeson and Scotland counties. When form I
males become available, this species may prove to be Procambarus
(Ortmannicus) lepidodactylus Hobbs, which is known from the system
in South Carolina, or an undescribed species of the Pictus Group
currently known only from the Waccamaw River basin (see below).

Waccamaw River — The northernmost headwaters of the
Waccamaw River drain southcentral Bladen and northern Columbus
counties, and its eastern tributaries drain western and northern
Brunswick County. Some of the northern tributaries enter Lake
Waccamaw, the largest of the natural Carolina Bay lakes, and the
Waccamaw River flows from the south shore of the lake. East of the
lake there is an extensive system of man-made drainage canals that
"connects Honey Island Swamp (Juniper Creek tributary) with Dans
Creek of the Cape Fear drainage and Big Creek which drains into
Lake Waccamaw, . . . ." (Shute et al. 1981:21). South of the lake,
the Waccamaw River forms part of the line between western
Brunswick and southeastern Columbus counties, then flows
southwest into South Carolina and enters Winyah Bay. The Shallotte,
Lockwood Folly, and Calabash rivers, which drain most of southern
Brunswick County, are here included in this system. The Calabash
River flows west into Little River in Horry County, South Carolina,
and is connected to the Waccamaw River through the Intracoastal
Waterway.

The crayfish fauna of the Waccamaw basin consists of C.
diogenes, C. latimanus (known from two localities in Brunswick
County), F. fodiens, P. a. acutus, P. ancylus, P. blandingii, and P.
pearsei. Specimens from this basin previously assigned to P.
lepidodactylus actually represent a new species being described by

North Carolina Crayfishes 121

JEC. Current evidence indicates that C. sp. C, which occurs in the
adjacent Lumber River basin, is absent from the Waccamaw basin.

Cape Fear River — The Cape Fear River, which begins in the
Piedmont Plateau and drains the largest watershed in the State, is
confined to North Carolina. Its major northern tributary, the
Haw River, heads in northwestern Guilford, southern Rockingham,
Orange, and Durham counties. A large southern tributary, the Deep
River, begins in southwestern Guilford and eastern Randolph counties,
flows southeast to northern Moore County, then turns east and north,
forming the northwestern boundary of Lee County. The Haw and
Deep rivers then merge in eastern Chatham County at the Lee County
line, forming the Cape Fear River. A central tributary, the Rocky
River, heads in northeastern Randolph County, flows southeast
through Chatham County, and joins the Deep River at the Lee County
line. A major central tributary in the Coastal Plain, the Black River,
drains Sampson County and the western part of Duplin County.
Another Coastal Plain tributary, the South River, divides Cumberland
and Bladen counties to the west from Sampson and Pender counties to
the east. The Black River merges with the South River in southern
Sampson County at the Bladen County line, and the South enters the
Cape Fear at the line between Pender and Bladen counties. The Cape
Fear then empties into the estuary at Wilmington, New Hanover
County.

Until recently, the endemic North Carolina crayfish, C. catagius,
was known only from its type locality, burrows in a lawn at East
Whittington Street in the southeastern section of Greensboro,
Guilford County (Hobbs and Perkins 1967:145), which is in the Haw
River subdrainage. Other colonies of this primary burrower have now
been discovered, and efforts are being made to more accurately
determine its range (Davis 1992:29). Cambarus reduncus is restricted
to the Piedmont and Fall Line zone, while C. latimanus, C. diogenes,
and C. sp. C are found in both the Piedmont and Coastal Plain. Procambarus
a. acutus appears to be common throughout the system, even into its
headwaters. Faxon's (1914:367) report of "Cambarus blandingii" from
Reedy Creek, near Greensboro, must have been based on P. a. acutus,
which D. G. Cooper collected from Lake Reidsville and a small lake
south of Reidsville, Rockingham County, in the Haw River subdrainage.
Fallicambarus fodiens apparently has made inroads into the Piedmont
of the Cape Fear basin; Hobbs and Peters (1977:46) recorded it from
the Rocky River in Chatham County. Procambarus ancylus is known
from White and Singletary lakes, Bladen County (Hobbs 1958:167),
and has been collected from a borrow pit near Colly Creek (NCSM

122 J. E. Cooper and A. L. Braswell

C-1949), which has a tributary stream entering it from Singletary
Lake. Procambarus pearsei is known from as far north in the system
as northern Sampson County, and might occur in extreme southwestern
Johnston County. Its type locality, a pond and ditch on NC 22 south
of Fayetteville, Cumberland County (Creaser 1934:4), is in this river
basin. Procambarus blandingii, which occurs in the Lumber and Waccamaw
basins, appears to be absent from the lower Cape Fear, but this needs
investigation.

The northern tributaries of the Northeast Cape Fear River originate
in the Coastal Plain in southern Wayne and northeastern Sampson
counties. The system drains almost all of Duplin, most of Pender,
western Onslow, and northern New Hanover counties. The Northeast
Cape Fear generally is considered an eastern trunk of the Cape Fear
River, but it flows into saline waters at Wilmington, independent of
the Cape Fear. As alluded to in the previous discussion of the Northeast
Cape Fear as a hydrologically and faunistically autonomous unit, there
are differences in the crayfish faunas of the two systems. Since the
Northeast Cape Fear is confined to the Coastal Plain, it is not surprising
that it does not harbor C. catagius or C. reduncus, both Piedmont
species. It does have P. plumimanus, however, which is absent from
the Cape Fear but present in the New (White Oak), and lacks P.
ancylus and P. pearsei, both of which occur in the Cape Fear, Lumber,
and Waccamaw. The rest of the crayfish fauna of the Northeast Cape
Fear is that of the coastal Cape Fear, i.e., C. latimanus, C. diogenes,
C. sp. C, F. fodiens, and P. a. acutus.

New {White Oak) River — The major hydrologic units of the New
(White Oak) drainage basin are the autonomous New, White Oak,
Newport, and North rivers and their tributaries. All are small and all
empty into saline coastal waters behind barrier islands in Onslow
Bay, and Bogue and Back sounds. The New River drains the center of
Onslow County, and extreme eastern Pender and New Hanover
counties. The White Oak drains eastern Onslow County and forms
part of the boundary between southeastern Jones and eastern Onslow
counties. Included in the White Oak unit are Catfish and Great lakes
of Croatan National Forest, southwestern Craven County. The Newport
and New rivers both drain southern Carteret County, and both, like
the White Oak, are artificially connected with the lower Neuse River
basin to the north (see the P. plumimanus account).

The crayfish fauna of the New (White Oak) basin consists of
C. diogenes, C. latimanus, F. fodiens, P. a. acutus, P. plumimanus,
and probably C. sp. C.

North Carolina Crayfishes 123

Neuse River — The Neuse is a second major drainage system
that heads and debouches in North Carolina (for a brief description
see Cooper and Ashton 1985:6-7). It includes Long, Little, and Ellis
lakes of Croatan National Forest, Craven County. Ellis Lake has a
direct connection with Southwest Prong Slocum Creek.

Cambarus reduncus is confined to the Piedmont Plateau and
Fall Line zone of this basin, where the southeasternmost known
localities are in central Wake County. Cambarus latimanus, C. diogenes,
C. sp. C, and P. a. acutus occur throughout the system. Procambarus
medialis, F. fodiens, and O. sp. A are essentially Coastal Plain species,
with F. fodiens and O. sp. A both edging into the Fall Line zone at
least as far west as southern Wake County. Two localities for P.
plumimanus, including the type locality, are known in the lower Neuse
basin. The listing by Harris (1903:56) of C. b. bartonii in the Neuse
was in error, and was probably based on Faxon's (1885:61) erroneous
report of the species from Kinston, Lenoir County, which also was
repeated by Ortmann (1931:131).

Tar-Pamlico River — Like the Cape Fear and Neuse rivers, the
Tar-Pamlico is confined to North Carolina (for a brief description see
Cooper and Ashton 1985:6-9). Lake Mattamuskeet, part of this system
in Hyde County, connects with the Alligator River of the Pasquotank
basin through drainage canals and the Intracoastal Waterway, which
also forms a direct connection between the Alligator and Pungo rivers.
Drainage canals also connect Pungo Lake of the Tar-Pamlico basin,
Washington and Hyde counties, with canals from New (Alligator)
Lake in the Pasquotank drainage of Hyde County.

The crayfish fauna of the Tar-Pamlico is the same as that of
the Neuse basin, except for the apparent expansion into the latter of
P. plumimanus. During the Pleistocene, these drainages "would have
been conjoined to form the Greater Pamlico River" (Lachner and
Jenkins 1971:62). In the Tar-Pamlico, however, P. medialis appears
to be limited to the immediate area of the type locality, a pool in
a roadside ditch, 0.6 mi (1 km) south of Scotland Neck, on US 258, in
southeastern Halifax County (Hobbs 1975:13). In the Neuse basin its
known range is more extensive, occupying an area from Johnston
County downriver to Lenoir and Pitt counties. In addition, the Tar-
Pamlico range of Orconectes sp. A extends from headwaters in
Granville County to eastern Pitt County, whereas in the Neuse, despite
intensive sampling, this species has not been found any farther
upriver than southern Wake County. The report by Faxon (1914:367)
of "Cambarus blandingii" from Lake Mattamuskeet was undoubtedly
based upon P. a. acutus.

124 J. E. Cooper and A. L. Braswell

Roanoke River — The Roanoke is another heavily impounded
river. Its major western tributary, the Dan River, begins in the Blue
Ridge in Virginia, and in northeastern Surry, Stokes, and Forsyth
counties, North Carolina. The Dan flows essentially east and north,
moving in and out of North Carolina in northern Rockingham and
Caswell counties, and acquiring tributaries in both. It joins the
Roanoke at Buggs Island Reservoir in Virginia. The Roanoke then
flows southeast into North Carolina, adding tributaries in northern
Granville and Vance counties and feeding into John H. Kerr Reservoir
and Lake Gaston. The river flows southeast from Lake Gaston, drains
southwestern Northampton, northern Halifax, and most of Bertie and
Martin counties, then empties into the western arm of Albemarle
Sound at Batchelor Bay.

Cambarus longulus and C. b. bartonii occur in the upper Dan
River watershed, where C. longulus is known from no farther east
than Rockingham County, and the eastern limits of the range of C. b.
bartonii are unknown. Cambarus sp. C apparently occurs throughout
the system. Cambarus diogenes and F. fodiens occur in the Coastal
Plain, but encroach on the Piedmont Plateau, where the western limits
of their ranges are uncertain. Procambarus a. acutus is found in all of
the Coastal Plain, and as far west in the Piedmont as eastern Caswell
County. Orconectes virginiensis is known from Bertie, Halifax, and
Martin counties.

Chowan River — The Chowan River basin is confined to the
northeastern Coastal Plain, rising at the confluence of the Nottoway
and Blackwater rivers just south of the Virginia state line. The southeast-
flowing Meherrin River heads in Virginia, enters North Carolina to
form part of the northern boundary between Northampton and Hertford
counties, then turns east across Hertford County to join the Chowan
at the Gates County line. The system drains northern and eastern
Northampton, Hertford, and most of southern and western Gates,
western Chowan, and northeastern Bertie counties, then empties
into the western arm of Albemarle Sound.

The crayfishes of the Chowan basin are C. diogenes, F. fodiens,
P. a. acutus, O. virginiensis, and possibly C. sp. C. The listing by
Harris (1903:53) of C. b. bartonii in the Chowan basin (in Virginia)
was in error, and was probably based on Faxon's (1885:61) erroneous
report of the species from Southampton and Lunenburg counties,
which also was repeated by Ortmann (1931:131)

Pasquotank River — North of Albemarle Sound, the streams
that form the northern part of this Coastal Plain system rise in the
southeastern tip of Virginia and in eastern Gates County, North
Carolina. This part of the basin includes the Pasquotank River that

North Carolina Crayfishes 125

drains northeastern Gates, northern Pasquotank, and the northern
half of Camden counties; the Perquimans River of Perquimans County;
the Little River that rises in the Dismal Swamp in central Pasquotank
County and forms part of the lower Pasquotank-Perquimans county
line; the North River that rises in the Dismal Swamp in Currituck and
Camden counties; the Northwest River, which begins in Norfolk
County, Virginia, and flows southeast into Tull Bay, Currituck County;
and the North Landing River, which begins in Princess Anne County,
Virginia, flows south into Currituck Sound, and connects with the
lower James River of Virginia via the Intracoastal Waterway. All
these rivers except the Northwest and North Landing flow into
Albemarle Sound.

South of Albemarle Sound the Pasquotank system includes the
north-flowing Alligator River, which drains parts of northern Hyde,
much of Tyrrell, and most of Dare counties; the Scuppernong River
that drains eastern Washington and northwestern Tyrrell counties;
and Phelps and New (Alligator) lakes. Both an immense system of
man-made drainage canals in the Dismal Swamp west of Phelps and
Pungo lakes, and the Intracoastal Waterway, link parts of the Pasquotank
basin with the Pungo River, Lake Mattamuskeet, and other parts of
the Tar-Pamlico basin in Beaufort and Hyde counties.

The crayfish fauna of the Pasquotank basin consists of C.
diogenes, F. fodiens, and P. a. acutus. Fallicambarus fodiens has
been found across Croatan Sound, near Manteo on Roanoke Island,
Dare County.

CRAYFISH DISTRIBUTIONS BY PHYSIOGRAPHIC PROVINCES

Cambarus is the dominant crayfish genus in North Carolina,
with 18 described species (not including C. " acuminatus") and at
least three known but undescribed species. Nine of these species are
limited to the Blue Ridge, three are limited to the Piedmont
Plateau, and two overlap the Piedmont and Coastal Plain (one of
these two also may be found in all three provinces). Distributions of
the taxa now subsumed under Cambarus sp. C, which includes
C. "acuminatus" will not be clarified until this complex has been
diagnosed. Suffice it to say that they occur in all three physiographic
provinces. The sole North Carolina species of Fallicambarus is
primarily Coastal Plain in distribution, but also has invaded the eastern
Piedmont Plateau. One of the two described Orconectes in North
Carolina occurs only in the Blue Ridge, as does an apparently
undescribed member of the genus. The other described Orconectes is
limited to the northeastern Coastal Plain, and a second undescribed

126

J. E. Cooper and A. L. Braswell

Table 2. Summary of crayfish distributions in major physiographic provinces
of North Carolina. X = present.

Species

Blue Ridge

Piedmont Plateau

Coastal Plain

Cambarus

(C.) b. bartonii

X

X

(C.) sp. A

?

X

(D.) catagius

X

(D.) latimanus

?

X

X

(D.) reduncus

X

(D.) sp. B

X

(//.) chasmodactylus

X

(H.) longirostris

X

(H.) longulus

X

X

(J.) asperimanus

X

X

(J.) carolinus

X

(J.) dubius

X

71

(J.) nodosus

X

(L.) diogenes

X

X

(P.) georgiae

X

(P.) hiwasseensis

X

(P.) parr is hi

X

(P.) reburrus

X

(P.) robustus

X

?

(P.) sp. C2

X

X

X

(T.) acanthura

X

Fallicambarus

(C.) fodiens

X

X

Orconectes

(C.) virginiensis

X

(P.) spinosus

X

(P.) sp. A

X

X

(P.) sp. B

X

Procambarus

(O.) a. acutus

X

X

(O.) ancylus

X

(O.) blandingii

X

(0.) lepidodactylus

?

(O.) medialis

X

(O.) pearsei

X

(O.) plumimanus

X

1 Known from east of the Blue Ridge in Avery, Surry, and Wilkes counties, but
eastern limits of range unknown.

2 Includes C. (P.) acuminatus (s. I.).

North Carolina Crayfishes 127

species of the genus is essentially Coastal Plain but occurs also at
the eastern edge of the Piedmont Plateau. Genus Procambarus, with
six, and probably seven, described native species, dominates the
Coastal Plain. Six of these species are limited to this province, but
the seventh also occurs deeply into the Piedmont. Table 2 is a summary
of species distributions by physiographic provinces.

ACKNOWLEDGMENTS— -We express our sincerest thanks to all
those collectors mentioned in this paper, without whose efforts we
would know considerably less about North Carolina crayfishes than
we do. Gary W. Woodyard provided the blue C. latimanus and helped
in other ways. Joseph F. Fitzpatrick, Jr. and Martha R. Cooper shared,
as usual, their knowledge and libraries. Dr. Fitzpatrick and an anonymous
reviewer provided invaluable comments on the manuscript. JEC is
also indebted to John E. Cooper, Jr., Martha R. Cooper, Gladys Leake,
Jesse Perry, Lynn Ferguson, John R. Holsinger, William J. Rishel,
Patricia S. and Ray E. Ashton, Jr., his co-author, and especially Don
Howard, without whose assistance this paper could never have been
completed.

Finally, we express our deepest gratitude to and for the late
Horton H. Hobbs, Jr., whose death was a great personal tragedy and
an incalculable loss to decapod research. His kindness and generosity,
his knowledge and insights, and his total integrity, shall never be
forgotten.

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132 J. E. Cooper and A. L. Braswell

ADDENDUM
After this paper went to press, V. Schneider and D. R. Lenat
collected the first Cambarus spicatus adults from North Carolina:
Cleveland Co., First Broad R at SR 1530, 6.1 air mi (9.8 air km)
WSW cntr Casar; 2 S II, 3 9 (NCSM C-2487), 20 Jun 1995.

Received 29 September 1994
Accepted 12 January 1995

133

NEW PUBLICATION

Reptiles of North Carolina by William M. Palmer and Alvin
L. Braswell was released by the University of North Carolina Press in
August 1995. It contains 428 pages, 76 color plates, 68 range maps,
170 tables, and 126 drawings by scientific illustrator Renaldo Kuhler.
The hardcover book is available from the publisher for $39.95 plus
shipping charges and N.C. sales tax (if applicable). Orders may be
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ERRATA

The authors of "Atlantic Ocean Occurrences of the Sea Lamprey,
Petromyzon marinus (Petromyzontiformes: Petromyzontidae), Parasitizing
Sandbar, Carcharhinus plumbeus, and Dusky, C. obscurus (Carcharhiniformes:
Carcharhinidae), Sharks off North and South Carolina" (Brimleyana
21:69-72) have requested that the following corrections be published:

Page 70: North Carolina paragraph, line 4, should have been
7.5-m-FL, not 3.

Page 71: Footnote1 to Table 1 should be host male, not female.

DATE OF MAILING

Brimleyana 21 was mailed on 25 April 1995.

134

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Adams, J. J. 1977. Food habits of the masked shrew, Sorex cinereus

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Adams, J. J. 1988. Animals in North Carolina folklore. Second

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Barnes, R. G. 1989. Northern flying squirrel. Pages 203-230 in

Mammals of the southeastern United States (J. J. Adams and J.

M. Smith, Jr., editors). Harper and Row, New York, New York.

Citing References in Text — Use parenthetical format, for example:
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STATE LIBRARY OF NORTH CAROLINA

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Publications of the North Carolina
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Endangered, Threatened, and Rare Fauna of North Carolina

Part I. A Re-evaluation of the Mammals

Mary K. Clark, editor, 52 pages, 1987, $5 postpaid

Part II. A Re-evaluation of the Marine and Estuarine Fishes

Steve W. Ross, Fred C. Rohde, and David G. Lindquist,
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Part III. A Re-evaluation of the Birds

David S. Lee and James F. Parnell, editors, 52 pages,
1990, $8 postpaid

Atlas of North American Freshwater Fishes

David S. Lee et al., 867 pages, 1980, $25 postpaid

Atlas of North American Freshwater Fishes, 1983 Supplement

D. S. Lee, S. P. Platania, and G. H. Burgess, 68 pages plus
looseleaf additions and corrections, 1983, $10 postpaid

A Distributional Survey of North Carolina Mammals

David S. Lee, John B. Funderburg, and Mary K. Clark,
70 pages, 1982, $5 postpaid

The Seaside Sparrow, Its Biology and Management

Thomas L. Quay et al., editors, 174 pages, 1983, $15 postpaid

Autumn Land-bird Migration on the Barrier Islands
of Northeastern North Carolina

Paul W. Sykes Jr., 50 pages, 1986, $5 postpaid

Potential Effects of Oil Spills on Seabirds and

Selected Other Oceanic Vertebrates Off the North Carolina Coast

David S. Lee and Mary C. Socci, 64 pages, 1989,
$8 postpaid

Bird Life of North Carolina's Shining Rock Wilderness

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95-001436

BRIMLEYANA NO. 22, JUNE 1995
CONTENTS

Aspects of the Feeding Ecology of the Little Grass Frog, Pseudacris ocularis
(Anura: Hylidae). Jeremy L. Marshall and Carlos D. Camp

Corrections of Records of Occurrence of Peromyscus polionotus (Wagner) and
P. gossypinus (LeConte) (Rodentia: Muridae) in the Blue Ridge Province
of Georgia. Joshua Laerm and James L. Boone

The Masked Shrew, Sorex cinereus (Insectivora: Soricidae), and the Red-backed Vole,
Clethrionomys gapperi (Rodentia: Muridae), in the Blue Ridge Province of
South Carolina. Joshua Laerm, Eric Brown, Michael A. Menzel, Amanda
Wotjalik, William Mark Ford, and Mary Strayer 15

Rediscovery of the Aquatic Gastropod Helisoma eucosmium (Bartsch, 1908),

(Basommatophora: Planorbidae). William F. Adams and Susan G. Brady 23

Effects of a Clearcut on the Herpetofauna of a South Carolina Bottomland Swamp.

Jospeh P. Phelps and Richard A. Lancia 31

First Record of the Water Shrew, Sorex palustris Richardson (Insectivora: Soricidae),
in Georgia with Comments on its Distribution and Status in the Southern
Appalachians. Joshua Laerm, Charles H. Wharton, and William Mark Ford 47

Florida Manatees, Trichechus manatus (Sirenia: Trichechidae), in North Carolina

1919-1994. Frank J. Schwartz 53

Recovery of the Cave Crayfish (Decapoda: Cambridae) Population in Peacock

Springs, Florida? W. J. Streever 61

Premolar Cementum and Noncementum Lengths As Potential Indicators of Age
for Beavers, Castor canadensis (Rodentia: Castoridae). Allan E. Houston
and Michael R. Pelton 67

Record Clutch Size for Chelydra serpentina (Testudines: Chelydridae) in Virginia.

Joseph C. Mitchell and Michael C. Odom 73

New Distributional Records for the Star-nosed Mole, Condylura cristata
(Insectivora: Talpidae), in North Carolina, with Comments on its
Occurrence in the Piedmont Region. Jeffrey C. Beane 77

Observations on North Carolina Crayfishes (Decapoda: Cambaridae).

John E. Cooper and Alvin L. Braswell 87

Miscellany 133