january 1989

number 15

EDITORIAL STAFF

Eloise F. Potter, Acting Editor

Eloise F. Potter, Managing Editor

Sheree Worrell, Production Manager

John B. Funderburg, Editor-in-Chief

Board

James W. Hardin

Department of Botany

N.C. State University

William M. Palmer

Curator of Lower Vertebrates

N. C. State Museum

David S. Lee

Curator of Birds

N.C. State Museum

Rowland M. Shelley

Curator of Invertebrates

N. C. State Museum

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

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

Contents will emphasize zoology of the southeastern United States, especially

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

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

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

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

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

invertebrate zoology, ichthyology, herpetology, ornithology, mammalogy, and

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

synthesizing reviews and papers of significant historical interest to southeastern

zoology will be included.

Suitability of manuscripts will be determined by the Editor and, where neces-

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

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

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

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

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

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

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

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

In citations please use the full name — Brimleyana.

North Carolina State Museum of Natural Sciences

North Carolina Department of Agriculture

James A. Graham, Commissioner

CODN BRIMD 7

ISSN 0193-4406

Occurrence of the Nine-banded Armadillo,

Dasypus novemcinctus (Mammalia: Edentata),

in South Carolina

John J. Mayer1

Savannah River Ecology Laboratory,

Drawer E, Aiken, South Carolina 29801

ABSTRACT. — The occurrence of the nine-banded armadillo, Dasypus

novemcinctus, in South Carolina has been poorly documented. The

recent discovery of a road-killed individual in Aiken County, capture

of a live individual in Barnwell County, and results of a questionnaire

sent statewide to wildlife personnel in South Carolina indicate an

increased frequency and concentration of records of the species in the

southern portion of the state. Results of the survey also revealed that

human importation of this mammal is still occurring in South Carolina.

Based on climatic limiting factors, the nine-banded armadillo should

be capable of expanding its range throughout most of South Carolina.

The presence of an established viable population of this species in

South Carolina remains uncertain.

Since the introduction of the nine-banded armadillo, Dasypus

novemcinctus Linnaeus, into Florida between 1915 and 1922 (Bailey

1924, Fitch et al. 1952), this mammal has expanded its range through or

into four states in the Southeast (Fitch et al. 1952, Humphrey 1974,

Hall 1981). However, the significance of its occurrence in South Carolina

is uncertain. Although sight records have been reported for this state

(three individuals by Golley 1966; one individual by Humphrey 1974;

ten individuals by Sanders 1978), these animals were assumed to be

escapees either from tourists’ automobiles traveling north from Florida

and Georgia or from circuses (Golley 1966; Sanders 1978; R. E.

Mancke, pers. comm.). No museum voucher specimens of this species

from South Carolina have been reported. In addition, undocumented

occurrences of the nine-banded armadillo in South Carolina have been

either stated or illustrated with maps by Hamilton and Whitaker (1979)

and Wetzel (1982).

On 2 August 1985, an adult male nine-banded armadillo roadkill

was found in Aiken County in the central portion of the Savannah

River Plant (SRP), a 77,000-ha federal nuclear facility that is closed to

public access. Therefore, it is unlikely that this animal was released by a

tourist traveling north. This specimen (ChM CM 1143) has been

Present address: Law Environmental, Inc., 112 Townpark Drive, Kennesaw,

Georgia 30144-5599.

Brimleyana No. 15:1-5, January 1989

1

2

John J. Mayer

deposited in the Charleston Museum. A subsequent check with other

biologists at this site resulted in six additional reports of roadkilled

armadillos in Aiken, Allendale, and Beaufort counties in South Carolina

during the spring and summer of 1985. The frequency of these sightings

suggests that this species is present in higher numbers in South Carolina

than had been suspected previously.

A second adult male nine-banded armadillo was discovered alive at

an elementary school in Williston, Barnwell County, on 26 June 1986

after becoming trapped in a drainage well. The animal was captured by

state wildlife personnel and later sent to the South Carolina State

Museum Commission.

A survey of the mammal collections in South Carolina resulted in

the location of another specimen and one additional sight record

(Charleston Museum files) from the state. The specimen (ChM CM1 142)

was taken alive in a barn in Bonneau, Berkeley County, in the mid-

1970s; it was later sent to the Charleston Museum where it was prepared

as a study skin. The additional sight record was from Piedmont,

Greenville County, in 1941.

In an effort to assess the recent population status and distribution

of this species in South Carolina, a questionnaire was sent to state

wildlife biologists, state wildlife law enforcement officers, superinten-

dents of the national forests in the state, and managers of the national

wildlife refuges in the state. The questionnaire requested details

concerning any recent sightings of this species in the respondent’s area

of South Carolina or any other sightings in the state known to the

respondent. It also asked whether or not the respondent believed that

the nine-banded armadillo was established in South Carolina. The

response to the 246 questionnaires was 57%.

Eleven respondents (8%) reported a total of 15 recent sightings of

the nine-banded armadillo in South Carolina, most of which were from

the southern portion of the state (Fig. 1). Most reports (73%) were of

roadkills along interstate highways and primary and secondary state

roads. Of the live sightings, two were seen along roadsides, one was

captured near a motel in Florence, Florence County, and one was

killed in a chicken coop by a farmer in Brunson, Hampton County.

Ninety-nine percent of the respondents did not believe that the nine-

banded armadillo was established in South Carolina at this time. Of the

two respondents who did believe this species to be established in the

state, only one reported any sightings. Two respondents stated that

they had encountered persons who either had been found in South

Carolina with armadillos that had been captured in Florida for release

in South Carolina or were in the process of capturing armadillos in

southern Georgia for transport to and release in South Carolina.

Nine-banded Armadillo in South Carolina

3

Fig. 1. Past and recent occurrence of the nine-banded armadillo, Dasypus

novemcinctus, in South Carolina. Solid dots indicate the localities of museum

specimens. Open circles indicate the 1980-to-present localities resulting from this

report. Solid triangles indicate the localities of the early (pre-1980) sightings in

the state resulting from this report, records at the Charleston Museum, and

records from Golley (1966), Humphrey (1974), and Sanders (1978). Limits of the

mean annual number of freeze-days are indicated by the numbered solid lines.

Routes of the Interstate Highway System in South Carolina are indicated by the

dotted lines.

Early records of the nine-banded armadillo in South Carolina were

most likely the result of escaped or released individuals that had been

imported into the state (Humphrey 1974, Sanders 1978). Most of these

records came from localities close to major highways (Fig. 1) that carry

tourist traffic north from Florida (Sanders 1978). Other records from

farther north along the Atlantic seaboard of the United States have

occurred in North Carolina, Washington, D.C., Delaware (Humphrey

1974, Lee et al. 1982), and Connecticut (UCONN 11249). The results of

the survey determined that introductions by humans are still occurring

in South Carolina. However, because of the recent increased frequency

and concentration of records of this species in the southern portion of

4

John J. Mayer

South Carolina adjacent to the Savannah River, it is possible that some

of these occurrences in the state represent natural range extensions.

Humphrey (1974) noted that the nine-banded armadillo has a strong

pioneering capability as indicated by the large number of extralimital

records in its distribution. The results of Humphrey’s (1974) study also

indicated that the distribution of the nine-banded armadillo had a lower

limit of about 380 mm annual precipitation and an approximate upper

limit of nine freeze-days per year (total number of days in a year during

which the maximum daily temperature does not exceed 0 degrees

Centigrade). Based on these data, Humphrey (1974) stated that the

range extension of the nine-banded armadillo could be expected to

reach at least the edge of the southern Appalachian piedmont in the

southeastern United States. Records of climatic data for South Carolina

from 1980 to 1985 (Anon. 1980-1985) indicate that the total annual

precipitation ranges from 733 to 1,517 mm and the average annual

number of freeze-days varies from 0 in southern South Carolina to 1 1 at

Caesar’s Head in the mountains (Fig. 1). Using Humphrey’s climatic

limiting factors, then, the nine-banded armadillo should be capable of

expanding its range throughout most of South Carolina. The only areas

that might be excluded would include the mountainous portions of the

extreme northwestern edge of the state.

In conclusion, reports of the nine-banded armadillo in South

Carolina are increasing at present, but because direct evidence for the

existence of an established viable population in South Carolina is

lacking, its status in the state remains uncertain.

ACKNOWLEDGMENTS. — I thank the many respondents to my

survey; Albert E. Sanders of the Charleston Museum (ChM) and

Robert E. Dubos of the University of Connecticut Museum of Natural

History (UCONN) for their helpful input; and I. Lehr Brisbin, Jr.,

Michael H. Smith, W. David Webster, Michael C. Kennedy, and James

M. Novak for critically reading earlier drafts of this manuscript. This

work was supported by Contract DE-AC09-76SROO8 19 between the

Institute of Ecology at the University of Georgia and the United States

Department of Energy.

LITERATURE CITED

Anonymous. 1980-1985. Climatological Data for South Carolina. Vol. 83-88.

NOAA National Climatic Data Center, Asheville, N.C.

Bailey, H. H. 1924. The armadillo in Florida and how it reached there. J.

Mammal. 5:264-265.

Nine-banded Armadillo in South Carolina

5

Fitch, Henry S., P. Goodrum, and C. Newman. 1952. The armadillo in the

southeastern United States. J. Mammal. 33:21-37.

Golley, Frank B. 1966. South Carolina Mammals. Charleston Mus. Press,

Charleston.

Hall, E. Raymond. 1981. The Mammals of North America. 2nd ed. John

Wiley and Sons, New York.

Hamilton, William J., Jr., and J. O. Whitaker, Jr. 1979. Mammals of the

Eastern United States. Cornell Univ. Press, Ithaca, N.Y.

Humphrey, Stephen R. 1974. Zoogeography of the nine-banded armadillo

( Dasypus novemcinctus) in the United States. BioScience 24:457-462.

Lee, David S., J. B. Funderburg, Jr., and M. K. Clark. 1982. A Distributional

Survey of North Carolina Mammals. N.C. Biol. Survey, N.C. State. Mus.

Nat. Hist., Raleigh.

Sanders, Albert E. 1978. Mammals [of the Coastal Zone of South Carolina]

Pages 296-308 in An Annotated Checklist of the Biota of the Coastal Zone

of South Carolina, R. G. Zingmark, editor. Univ. South Carolina Press,

Columbia.

Wetzel, Ralph M. 1982. Systematics, distribution, ecology, and conservation of

South American edentates. Pages 345-375 in Mammalian Biology in South

America, M. A. Mares and H. H.. Genoways, editors. Spec. Publ. Ser.,

Pymatuning Lab. Ecol. 6:1-539.

Accepted 26 September 1986

6

A DISTRIBUTIONAL SURVEY

OF NORTH CAROLINA MAMMALS

by

David S. Lee, John B. Funderburg, Jr., and Mary K. Clark

This book lists all the mammals of North Carolina and offers

species accounts and range maps for all of the non-marine species.

Introductory chapters describe the plant communities of the state as

they relate to mammal distribution and discuss local zoogeographic

patterns.

1982 72 pages Softbound ISBN 0-917134-04-4

Price: $5, postpaid. North Carolina residents add 5% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send to MAMMAL BOOK, N.C. State Museum of Natural Sciences,

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

Distribution and Seasonality of Branchiopod and

Malacostracan Crustaceans of the

Santee National Wildlife Refuge, South Carolina1

Charles K. Biernbaum

Grice Marine Biological Laboratory,

College of Charleston, 205 Fort Johnson,

Charleston, South Carolina 29412

ABSTRACT. — Distribution and seasonal changes in abundance of

branchiopod and malacostracan crustaceans were studied at the

approximately 6100-ha Santee National Wildlife Refuge, on the edge

of Lake Marion in the mid-coastal plain of South Carolina. A total of

42 species were collected, of which 19 are new records for the state.

Winter crustacean fauna is dominated by the cladocerans Simocephalus

serrulatus and Eurycercus ( Bulatifrons ) vernalis , the isopods Caecidotea

forbesi and Lirceus lineatus , the amphipods Hyalella azteea and

Crangonyx riehmondensis, mature individuals of the shrimp Pal-

aemonetes paludosus , and the crayfish Procambarus ( Ortmannicus )

blandingii. Spring is characterized by high water levels, population

pulses in several cladoceran species, occasional occurrences of the

anostracan Streptocephalus seali and conchostracan Eulimnadia

ventricosa , and more habitats for C. forbesi and P. ( O .) blandingii.

Summer crustacean fauna is dominated by H. azteea , Caecidotea

latieaudata , reduced numbers of C. forbesi, and immature specimens of

P. paludosus. Variations exist among species with respect to the types

of habitats inhabited (ditches, pools, impoundments, lake) and seasons

of abundance and reproductive activity. Most species have been

previously documented as being closely associated with aquatic

macrophytes, which reach high densities in several habitats.

Crustaceans are important components of practically all freshwater

ecosystems and many terrestrial ecosystems. Unfortunately, however,

little documented research has been done in South Carolina on the

distribution and ecology of nonmarine species, with the possible

exception of crayfish. Information about species occurrences, seasonal

changes in abundance, and seasonal reproductive characteristics is

especially valuable in areas that are managed for wildlife that may use

these crustaceans directly or indirectly as food. The Santee National

Wildlife Refuge, like many refuges, is very important as a winter

sanctuary for waterfowl, having more than 120,000 overwintering birds

annually (D. J. Voros, Santee National Wildlife Refuge, pers. comm.).

'Contribution Number 69 of the Grice Marine Biological Laboratory, College

of Charleston.

Brimleyana No. 15:7-30, January 1989

7

8

Charles K. Biernbaum

Waterfowl, though reduced in numbers, are abundant during other

times of the year as well. In addition to their trophic importance to

these birds, aquatic crustaceans are also important in food chains

leading to other animals, including the American alligator ( Alligator

mississippiensis ), an endangered species that is abundant within the

refuge boundaries.

Previous studies on freshwater and terrestrial crustaceans of South

Carolina are meager. There are many references to crayfish occurrences

in the state, one dating back more than 200 years (e.g., Bartram 1771;

Hobbs 1940, 1947, 1956a, b,c, 1958a, b, 1983; Hobbs and Carlson 1983;

Prins and Hobbs 1972; Hobbs III et al. 1976). Other taxa, however,

have not been as readily seen or surveyed. Important studies include a

survey of the Savannah River fauna (Patrick et al. 1966); several reports

of research involving a few crustacean species at the Savannah River

Plant, an approximately 80,000-ha federal production reactor and field

laboratory complex on the southwestern border of the state (e.g.,

Vigerstad and Tilly 1977; Thorp and Ammerman 1978; Cherry et al.

1979a, b; Giesy et al. 1980; Brown 1981; Dickson and Giesy 1981; Thorp

and Bergey 1981); some records of ostracod and copepod occurrences

(Hoff 1944; Ferguson 1952, 1954; Crawford 1957; Roache 1959); a

recent survey of zooplankton of an acidic (pH 4. 3-4. 5) cooling pond

(Mallin 1984); and documentation of the occurrence of two exotic

terrestrial amphipod species in the state (Biernbaum 1980). Except for

some early studies by a few biologists, some of which resulted in the

description of new species (Say 1818; Ellis 1940, 1941), very little work

has been done on nonmarine amphipods or isopods in the state. Fox

(1978) and Kelley (1978) summarized distributional information that is

known for these two groups in the coastal zone of South Carolina. No

information about the fauna or flora of the Santee National Wildlife

Refuge appears in the scientific literature. However, the United States

Department of the Interior has prepared pamphlets ( 1983:RF-42570-2,

42570-5, 42570-7; 1985:RF-42570-3) listing the species of fishes (based

on studies by P. Coleman), amphibians and reptiles (based on studies by

J. R. H arrison. III), and birds and mammals (based on many surveys)

occurring on refuge property.

STUDY AREA

Established in 1941, the Santee National Wildlife Refuge comprises

about 6100 ha in the mid-coastal plain of South Carolina (Fig. 1). The

four noncontiguous units of the refuge (Bluff, Dingle Pond, Pine Island,

and Cuddo) border Lake Marion, a reservoir created by the construction

of a hydroelectric dam on the Santee River. These units consist of

mixed pine-hardwood forests, croplands, marshes, ponds, and

impoundments.

Branchiopod and Malacostracan Crustaceans

9

Fig. 1. Map showing sampling areas, Santee National Wildlife Refuge, S.C.

The four units of the refuge are managed primarily for waterfowl.

Consequently, extensive complexes of water impoundments with

connecting ditches have been constructed on these sites. These impound-

ments vary in size, but a majority are 1 to 2 ha in size and quite shallow,

with most being virtually covered in summer with such rooted vegetation

as lilies or lotus. Other aquatic habitats occurring in the refuge are small

borrow pits and shallow swamps, some bordering the lake and others

found as isolated forest depressions.

There are significant seasonal variations in the amount of water

present in these aquatic habitats. The water level in Lake Marion is

reduced prior to spring in anticipation of increased runoff farther up the

Santee basin. As a result of such runoff, coupled with increased local

rainfall, water levels in the refuge are high in the spring, except for some

isolated habitats when spring rainfall is minimal. Usually, spring water

levels become so high that substantial areas of forest are also flooded, at

times up to 0.5 m in depth. Water levels then drop during summer and

fall, frequently resulting in the total drying-out of some impoundments,

ditches, and swamps.

At any one time, temperatures of aquatic habitats at the refuge are

highly variable because of differences in water depth and degree of

shading. The highest water temperature recorded during the study was

10

Charles K. Biernbaum

34 °C, but it is certain that temperatures occasionally exceed this in

small exposed pools or ditches on hot days. During the winter some

locations may rarely have a thin sheet of ice on the surface. Mid-June

pH measurements along the margin of Lake Marion ranged from 6.9 to

7.3, depending on location. Other aquatic habitats had pH values from

5.4 to 6.3.

Most aquatic habitats of the refuge, with the exception of swamps,

are characterized by having dense growths of a variety of aquatic plants.

Most such locations have large populations of floating plants, including

water lilies ( Nymphaea odorata ), American lotus ( Nelumbo luted), and

frequently water shield ( Brasenia schreberi ) and duck weed ( Lemna

spp.). Submergent plants that are very abundant include bladderwort

( Utricularia sp.), hornwort ( Ceratophyllum demersum), Brazilian water-

weed ( Elodea densa), water milfoil ( Myriophyllum sp.), Chara sp., and

a variety of filamentous green algae. Along the bordering shallows such

emergent plants as arrowhead ( Sagittaria latifolia ), water pennywort

( Hydrocotyle sp.), alligator weed ( Alternanthera philoxeroides ), smart-

weed ( Polygonum spp.), pickerel weed ( Pontederia cordata), and an

introduced false loosestrife ( Ludwigia uruquayensis) are usually abun-

dant. At the water’s edge a variety of sedges ( Cyperus spp.), rushes

(Juneus effusus , Scirpus spp.), and cattails ( Typha sp.) are frequently

found in dense stands. Many impoundments additionally have but-

tonbushes ( Cephalanthus occidentalis ) scattered throughout. Bald

cypress ( Taxodium distichum), water tupelo (Nyssa aquatica ), willows

( Salix spp.), and redbay ( Persea borbonia) are commonly found. Trees

usually dominating swamps include those listed above plus sweet gum

(Liquidambar styraciflua ), oaks ( Quercus spp.), and, in those swamps

and flooded woods not bordering the lake, some pines ( Pinus spp.).

METHODS

Sampling was done approximately every 4 to 8 weeks from January

1982 to November 1983. Occasional samples were taken during 1984.

Aquatic sampling was done primarily by use of a dipnet (0.9-mm mesh),

although some supplemental collections were made by plankton net

(100-^m mesh) at selected locations. Sampling was reduced in certain

areas during the cooler months to avoid disturbing overwintering

waterfowl. Specimens were collected at 105 refuge locations, over 75%

of which were sampled during at least two different months. Sixty-three

stations were sampled during at least two different seasons, 34 stations

during at least three seasons, and 16 for all four seasons.

Specimens were preserved in the field and returned to the laboratory

for identification and enumeration. Owing to the difficulty in identifying

females of Caecidotea to the species level, occurrences of species of this

Branchiopod and Malacostracan Crustaceans

11

isopod genus are based solely on the collection of males, plus those

females found at sites where males of only one species of Caecidotea

were collected. Individuals of the two dominant amphipod species,

Hyalella azteca and Crangonyx richmondensis richmondensis, were

identified as male, female, or juvenile. Because immature members of

these species resemble females, juveniles were defined as all individuals

shorter than the smallest recognizable male collected for each species. All

individuals of H. azteca were able to be sexed; however, many unsexed

C. r. richmondensis juveniles were collected. Voucher specimens of all

species have been deposited in the collections in the Division of

Crustacea of the United States National Museum of Natural History,

Washington, D.C.

RESULTS AND DISCUSSION

Branchiopods

Results of the crustacean survey are shown in Table 1. Two species

of non-cladoceran branchiopods were collected, the anostracan Strep-

tocephalus seali and the conchostracan Eulimnadia ventricosa. Both

species were found in one location only, two shallow pools adjoining a

shaded borrow pit (depression formed by soil excavation) in a forest.

They were absent in these pools in May 1982, present the following July

and September, and absent in December 1982. In July and September

S. seali was very abundant, with E. ventricosa less so. Also present were

large numbers of the cladoceran Simocephalus exspinosus and the

isopod Caecidotea forbesi. The pools, which contained water throughout

1982, were approximately 3 x 4 m and 4 x 4.5 m and were separated by

about 1 m; in July and September they were separated from the borrow

pit by a ridge approximately 0.5 m wide and 15 cm in elevation above

the water level. In July and September all four crustacean species were

abundant in the two pools, but absent from the borrow pit. There were

large numbers of small fish in the borrow pit, but none in the two pools.

By December 1982, water levels had risen as a result of rainfall so that

the pools and borrow pit were confluent. No anostracans or con-

chostracans could be found at that time. Spring and summer were drier

in 1983 than in 1982, and the pools were totally dry in May and August.

They contained water in the early summer of 1983, but no branchiopods

were found in this habitat during that year. Such sporadic occurrences

of non-cladoceran branchiopods is not uncommon (Pennak 1978).

Streptocephalus seali is the most widely distributed anostracan

species in North America, being found in pools and ponds from the

Canadian prairies south to Mexico and east to the Atlantic States

(Moore 1966, Fitzpatrick 1983). There is only one previous record of its

occurrence in South Carolina, and that is based on a single specimen

12

Charles K. Biernbaum

Table 1. Summary of the branchiopod and malacostracan crustaceans collected in the

Santee National Wildlife Refuge, S.C.

Class Branchiopoda

Order Anostraca

Family Streptocephalidae

Streptocephalus seali Ryder

Order Conchostraca

Family Limnadiidae

* Eulimnadia ventricosa Mattox

Order Cladocera

Family Sididae

Diaphanosoma brachyurum (Lieven)

*Latona setifera (O.F. Muller)

* Pseudosida bidentata Herrick

Sida crystallina (O.F. Muller)

Family Holopedidae

Holopedium amazonicum Stingelin

Family Daphnidae

Ceriodaphnia reticulata (Jurine)

*Daphnia laevis Birge

*Daphnia pulex Leydig

Scapholeberis kingi Sars

*Simocephalus exspinosus (Koch)

*Simocephalus serrulatus (Koch)

Family Bosminidae

Bosmina cf. B. longirostris { O.F. Muller)

Family Macrothricidae

Ilyocryptus spinifer Herrick

*Macrothrix rosea (Jurine)

Family Chydoridae

* A Iona cost at a Sars

* A Iona cf. A. guttata Sars

*Alona intermedia Sars

Alonella dadayi Birge

Alonella hamulata (Birge)

*Camptocercus cf. C. rectirostris Schodler

Chydorus cf. C. sphaericus (O.F. Muller)

*Eurycercus ( Bulatifrons ) vernalis Hann

*Monospilus sp.

Oxyurella brevicaudis Michael and Frey

* Pseudochydorus globosus (Baird)

Class Malacostraca

Order Isopoda

Family Asellidae

Caecidotea forbesi (Williams)

Habitat1 Occurrence2 Unit3

Branchiopod and Malacostracan Crustaceans

13

Table 1. Continued.

Habitat* Occurrence2 Unit3

t *Caecidotea laticaudata (Williams)

t *Caecidotea obtusa (Williams)

Lirceus lineatus (Say)

Family Armadillididae

Armadillidium vulgar e (Latreille)

Family Oniscidae

t * Porcellionides floria Garthwaite

and Sassaman

Family Trichoniscidae

* Miktoniscus halophilus Blake

Order Amphipoda (See also Addendum.)

Family Crangonyctidae

Crangonyx r. richmondensis Ellis

Crangonyx serratus (Embody)

Family Hyalellidae

Hyalella azteca (Saussure)

Order Decapoda

Family Cambaridae

Fallicambarus ( Creaserinus )

uhleri (Faxon)

Procambarus ( Ortmannicus )

blandingii (Harlan)

Procambarus ( Ortmannicus )

hirsutus Hobbs

Procambarus ( Scapulicambarus )

troglodytes (LeConte)

Family Palaemonidae

Palaemonetes paludosus (Gibbes)

D,I,L,S

I

I,L

T

1

5

1

B, C,DP

DP

C, DP,PI

B

T

4

B

D,I,L,S

I,L

D,I,L,S

1

4

B,C,DP,PI

DP

B,C,DP,PI

I

D,I,L,S

L

I

D,I,L

5

1

5

4

1

C

B,C,PI

B

PI

B,C,DP,PI

♦First record for South Carolina.

fRange extension.

'Habitat Codes:

Occurrence Codes:

3Unit Codes:

B: Bluff

C: Cuddo

DP: Dingle Pond

PI: Pine Island

D:Ditch

I: Impoundment, borrow pit

L: Lake Marion

S: Swamp; tree-shaded, shallow

T: Terrestrial

1: Commonly encountered and usually abundant when present.

2: Commonly encountered and usually not abundant, but

with incidences of large numbers.

3: Commonly encountered, but usually not abundant.

4: Not commonly encountered, but usually abundant when present.

5: Not commonly encountered and usually not abundant

when present.

14

Charles K. Biernbaum

(Dexter 1953). Eulimnadia ventricosa occurs in the Atlantic drainage

from Maryland to Georgia (Fitzpatrick 1983), but has not been

previously reported from South Carolina.

Of the 25 cladoceran species collected, only three were widely

distributed and, when present, usually abundant: Simocephalus serru-

latus, Eurycercus ( Bullatifrons ) vernalis, and Ilyocryptus spinifer. Of

these, S. serrulatus was the most commonly encountered species.

Simocephalus serrulatus had two pulses of increased numbers: May

through June and December, with June’s population increase being

especially pronounced. Population fluctuations of E. vernalis were

similar, with pulses occurring in June and from December through

January.

Four cladoceran species ( Simocephalus exspinosus, Ceriodaphnia

reticulata , Scapholeberis kingi , Macrothrix rosea ) were widely dis-

tributed in the refuge, but usually present in small numbers. There were

occasions, however, when each of these species was abundant.

Simocephalus exspinosus , for example, reached very high densities in

the borrow pit and adjacent pools referred to above from July through

January and in a swamp and flooded forest floor in January. It was

widely distributed at other locations and at other times, but in low

numbers. Ceriodaphnia reticulata , otherwise found in low numbers,

became very abundant in a ditch and impoundment in June 1983.

Scapholeberis kingi reached enormous numbers in a borrow pit in June

1983. There were fairly high numbers of Macrothrix rosea in a few

samples. Like the species mentioned above, Chydorus cf. C. sphaericus

and Diaphanosoma brachyurum were widely distributed, but never

collected in abundance. This is in contrast to the findings of Mallin

(1984), who found D. brachyurum to be one of only two major

cladoceran species (the other being Bosmina longirostris) in an acidic

South Carolina impoundment.

Three cladoceran species were very restricted in their distribution,

but usually abundant when present. In August 1982 Daphnia laevis

reached enormous numbers in a swamp adjoining Dingle Pond, a

habitat that was dry in August 1983. That species was found nowhere

else in the refuge. Elsewhere, Daphnia laevis has often been found in

temporary ponds (Brooks 1959). Sida crystallina was restricted to Lake

Marion and a few impoundments, being very abundant in the lake

during June 1983. Monospilus sp., although restricted to the lake, was

found there in large numbers.

Frey ( 1982a, b,c) urged caution when identifying cladoceran species

because several species, previously believed to be cosmopolitan, have

been found to be species complexes, e.g. Black’s (1980) report on

Bosmina longirostris and that by Frey (1980) on Chydorus sphaericus.

Branchiopod and Malacostracan Crustaceans

15

Also, there are several undescribed species of Camptocercus in the

United States (Fitzpatrick 1983). As pointed out by Hann (1982), to

distinguish Eurycercus vernalis from the closely related sibling species

E. longirostris, one must examine the anatomy of individuals of various

ages within a population; I did not do this. Nevertheless, the refuge is

within the presently recognized range of E. vernalis (North Carolina to

Louisiana) and a considerable distance from that of the sibling E.

longirostris (Indiana). Frey (1982a) has recently mentioned species in

the southern United States in particular when commenting on the

present systematic confusion within several cladoceran groups. He states

that the cladoceran fauna of the southern United States consists of

several undescribed species, many possibly constituting species pairs

with non-conspecific populations having the same name in the northern

part of the country.

With the above caveats in mind, I have determined that the 25

cladoceran species collected in the study include 14 not previously

reported from South Carolina (Table 1). All 14 species are found

virtually throughout North America except for E. vernalis (as described

above), Pseudosida bidentata (largely restricted to the southern states;

Fitzpatrick 1983), and Monospilus sp. The only species of Monospilus

previously reported from North America is M. dispar, which is known

only from the northern United States and Canada (Fitzpatrick 1983).

The species from the refuge differs from M. dispar primarily in the

shape of the labrum. Another possibly undescribed refuge species is

Alona cf. A. guttata , specimens of which differ slightly, but probably in

a taxonomically significant manner, from those of A. guttata Sars in

having a distal expansion of the postabdomen.

Isopods

There are three common aquatic isopod species ( Caecidotea forbesi ,

C. laticaudata , Lirceus lineatus) at the refuge and one very rare species

(C. obtusa)\ the rare species was found during both winters, but only in

Dingle Pond. Of these species, C. forbesi was the one most commonly

encountered throughout the refuge and throughout the year. Caecidotea

forbesi was most widely distributed from winter into early summer, with

occurrences decreasing during late summer and fall (Table 2). This

species was most abundant when the water was cool, either during the

winter at many sites or throughout the year at some shaded locations,

such as shallow forest swamps and shaded ditches. Although C. forbesi

was found in a wide variety of habitats, it was largely restricted to

shallow, shaded areas and was rarely found in impoundments. At

locations where the only species of Caecidotea collected was C. forbesi

(based on identification of males), brooding females were found in all

16

Charles K. Biernbaum

seasons. However, because females were not identified (taxonomically

important morphological characters were variable and overlapping in

these species), caution is urged when interpreting such reproductive

data.

Caecidotea laticaudata was most extensively distributed in the

summer (Table 2), when it was numerically most abundant. Although

found in various habitats, frequently co-occurring with C. forbesi, C.

laticaudata was largely restricted to the lake margin of the refuge units,

plus some immediately adjacent aquatic habitats such as impoundments

or swamps. Like C. forbesi , it was common in swamps; however, it did

not occur in such habitats in the interior of the units, as did C. forbesi.

Unlike C. forbesi , C. laticaudata was at times abundant in certain

impoundments next to Lake Marion. During the winter this species

occurred only in a few sunlit impoundments. Caecidotea laticaudata

occurred in shaded areas only in summer. At refuge sites where only C.

laticaudata was collected (based on identification of males), brooding

females were found in January and June. As mentioned above, one

must be cautious when using data based on unidentified females,

particularly because reproduction in January of a species like C.

laticaudata , which is rare in winter and abundant in summer, would be

unexpected.

Differences exist between refuge units with respect to occurrences

of Caecidotea species. These differences most likely result from inter-

unit variations in lake proximity and types of aquatic habitats present.

Prime examples are comparisons of the Bluff and Cuddo units. Bluff

unit is very narrow, with most of its area close to the lake; considerably

less than half of Cuddo unit is as close to the lake as is all of Bluff unit

(see Fig. 1). Bluff unit also has much more of its aquatic habitats

consisting of impoundments, rather than swamps or ditches, than does

Cuddo unit. Most likely due to these differences, C. laticaudata is the

dominant isopod in the Bluff unit and C. forbesi in the Cuddo unit.

As mentioned above, there is considerable seasonal variation in the

degree of flooding of aquatic habitats. In the dry summer and fall most

of the shallow habitats for C. forbesi dry out, greatly reducing the space

available for it and, as a result, its abundance. Parsons and Wharton

(1978) have reported a similar reduction in numbers of an unidentified

species of Asellus (= Caecidotea) when water levels dropped in summer

on a Georgia flood plain. However, because C. laticaudata is more

common in impoundments, seasonal dryness has far less effect on it.

Lirceus lineatus is widespread in the refuge from fall to early spring

(Table 2). During the winter it reaches its greatest abundance,

making up a substantial portion of the aquatic isopod fauna (Table 3),

but it virtually disappears in the summer. It was found in a wide variety

Branchiopod and Malacostracan Crustaceans

17

Table 2. Percentages by months of isopod taxa collected at Santee National

Wildlife Refuge, S.C. Stations included are only those where isopods

were found at least once. Data on species of Caecidotea are based

solely on the occurrence of males, unless only one species of Caecidotea

was collected at a particular locality. Data from 1982 and 1983 have

been combined.

Table 3. Percentages by months of all aquatic isopod specimens from Santee

National Wildlife Refuge, S.C., represented by Caecidotea spp. and

Lirceus lineatus. Data from 1982 and 1983 have been combined.

Caecidotea spp.

Dec-Jan 57.6

Mar-May 79.8

Jun-Jul 99.5

Aug-Sep 100.0

Nov 45.4

of habitats, but impoundments were more commonly occupied than

were shallow forest swamps or ditches. Brooding females were collected

only from December through March.

Oniscoid isopods were found only on and near the Indian Mound

Historic Site at Fort Watson on the edge of Bluff unit. This site is

outside the protected refuge area and is subject to significant human

disturbance. Repeated searches for terrestrial isopods elsewhere in the

refuge met with no success.

Collection of C. laticaudata , C. obtusa, Porcellionides floria , and

Miktoniscus halophilus at the refuge constitutes their first documented

occurrences in South Carolina. Caecidotea laticaudata has been reported

previously from Louisiana, Alabama, Mississippi, Kentucky, and Illinois

(Williams 1970, 1972; Fleming 1972). Caecidotea obtusa is also primarily

18

Charles K. Biernbaum

a southern species, having been reported from Georgia and Florida west

to Louisiana and southern Arkansas (Williams 1970, 1972; Fleming

1972). Caecidotea forbesi is widespread east of the Mississippi River,

with the exception of the Gulf coastal region (Williams 1970, 1972;

Fleming 1972). Williams (1970) has previously reported the occurrence

of C. forbesi in Anderson County, S.C. It is interesting that there

appear to be correlations between the geographical distribution of C.

laticaudata and A. forbesi and their respective seasonal occurrences at

the refuge (Table 2). The northerly occurring C. forbesi is most common

during cool seasons (March-May), whereas the southern species, C.

laticaudata , is most common in summer. Lirceus lineatus was originally

described by Say (1818) from Berkeley County, S.C. It is widespread

east of the Mississippi River from the Great Lakes through the Gulf

coastal region (Hubricht and Mackin 1949, Williams 1972). The oniscoid

Armadillidium vulgare is cosmopolitan. Kelley (1978) reported A.

vulgare from Aiken and Charleston counties, S.C. Porcellionides floria

has been recently described by Garthwaite and Sassaman (1985) from

the southern and western United States, Mexico, and the Bahamas. Its

presence at the refuge constitutes its northernmost documented occur-

rence in the eastern part of the country. Miktoniscus halophilus ranges

from Massachusetts to Georgia (Schultz 1975, 1976), but it has not been

previously reported from South Carolina.

Amphipods

Three amphipod species were collected in the refuge. Two species

(Hyalella azteca and Crangonyx richmondensis richmondensis ) were

common, whereas one species (C. serratus) was rare. Crangonyx serratus

was collected only in winter in one unit, Dingle Pond (Table 4). No

brooding females of this species were encountered. Fox (1978) reported

all three species from the state.

The most widespread and abundant amphipod was H. azteca ,

which reached high densities in practically all types of aquatic habitats.

Although common throughout the year, examination of its degree of

dispersion throughout the refuge (Table 5) and its abundance where it

was found (Table 6) indicates that populations in many habitats were

reduced during two periods. The first was a notable reduction in winter.

The second was in mid-to-late summer, when two important environ-

mental changes were evident. One was progressive drying out of many

habitats that were flooded during the spring peak in abundance of the

species. Another change in some locations having very dense growths of

aquatic vegetation was apparent deterioration in water quality, as

evidenced by abundant flocculent material in the water, the formation

of an organic film on the surface, and occasionally the odor of hydrogen

Branchiopod and Malacostracan Crustaceans

19

Table 4. Percentages by months of all amphipod specimens from Santee

National Wildlife Refuge, S.C., represented by each of the three

species collected: Hyalella azteca, Crangonyx r. richmondensis, and

Crangonyx serratus.

Table 5. Percentages by months of stations at Santee National Wildlife Refuge,

20

Charles K. Biernbaum

sulfide. Most such areas showed a dramatic reduction in numbers of

crustaceans, H. azteca included, while in nearby habitats that lacked

such apparently detrimental characteristics H. azteca was frequently

abundant. Brooding females of H. azteca were collected throughout the

year; however, reproduction was greatly reduced in fall and winter

(Table 7). Bousfield (1958, 1973) has previously reported that ovigerous

females of H. azteca occur from April to October, as reflected by studies

done in such northern locations as Ontario (Lindeman and Momot

1983), British Columbia (Hargrave 1970, Mathias 1971); Oregon (Strong

1972), Michigan (Cooper 1965), and New York (Embody 1912).

However, Strong (1972) reported that H. azteca reproduces all year in a

hot spring (12-40 °C) in Oregon.

Crangonyx r. richmondensis, equally varied as H. azteca in the

aquatic habitats occupied, was collected all year, but, in contrast to H.

azteca , was common only during winter (Tables 4 and 5); it was very

rare in summer. Mathias (1971) reported that Crangonyx is much more

tolerant of cold than is Hyalella , and that species of Crangonyx

frequently breed in winter and spring. No relationship was found

between the few summer occurrences of C. r. richmondensis and specific

habitats; it seems to become rare at all of its locations as winter passes

into summer. Reproduction in this species occurred at the refuge from

late fall through spring (Table 7), which is similar to Bousfield’s (1958)

report that ovigerous females of this subspecies occurred from December

to June in southern portions of its range.

Hyalella azteca occurs throughout North and Central America and

the Caribbean islands north to the tree line in Canada and Alaska in all

permanent fresh water that reaches a monthly mean summer temperature

of over 10 °C (Bousfield 1958). Cooper (1965) reported several sources

referring to the high degree of association between large populations of

this species and such aquatic plants as Chara, Elodea, and Myrio-

phyllum. Such an association probably accounts in large part for the

very high numbers of H. azteca frequently seen at the refuge, where

dense growths of aquatic plants are extremely common. The presence of

large numbers of waterfowl at the refuge provides a means of widespread

dissemination, as reported by Daborn (1976) and Swanson (1984) for

this species. Over Lake Marion in December 1982, J. Pinckney shot a

female wood duck (Aix sponsa) that had 10 amphipods in the breast

feathers. Three were retained and identified as H. azteca.

Crangonyx richmondensis was originally described from a site in

Berkeley County, S.C. (Ellis 1940). The subspecies C. r. richmondensis

occurs east of the Appalachians from Georgia north to Nova Scotia and

Newfoundland (Bousfield 1958, 1973). However, the distribution is

disjunct, with no records of occurrence between Massachusetts and

Branchiopod and Malacostracan Crustaceans

21

Table 6. Seasonal abundance of Hyalella azteca at sites where collected. Data

are expressed as percentage of such stations at which its occurrence

was common or uncommon. Data from 1982 and 1983 have been

combined.

Table 7. Sex ratios and percentages of females of Hyalella azteca and Crangonyx

22

Charles K. Biernbaum

Virginia. Holsinger (1972) has reported that the northern and southern

forms of this subspecies differ morphologically. In fact, the systematics

of the entire C. obliquis-richmondensis group is presently unclear

(Holsinger 1972).

Crangonyx serratus occurs in the coastal plain and piedmont areas

from Florida to Maryland (Holsinger 1972). Bousfield (1958) pointed

out that the first record of this species in the literature was Say’s (1818)

report of specimens from South Carolina, which Say identified as

“ Ampithoe dentata .” Holsinger (1972) mentioned that C. serratus

commonly co-occurs with C. r. richmondensis.

Decapods

Palaemonetes paludosus was found in large numbers in all types of

aquatic habitats, with the exception of those that are isolated and

occasionally dry out. It was most abundant from fall to early winter.

From midwinter through late spring there was a reduction in abundance,

with an increase from summer through late fall (Table 8). However,

some locations had abundant numbers at all seasons. Brooding females

of P. paludosus occurred from May through July. Postlarvae were

abundant by June, and immature individuals were dominant from then

through August. Average individual size increased notably through fall

and winter.

The type locality for P. paludosus is Charleston County, S.C.

(Gibbes 1850). It has been subsequently reported from the state by other

authors, including Hobbs et al. (1976). This species is common in fresh

waters of the coastal plain and lower piedmont east of the Appalachians

from New Jersey to Florida and thence west to Texas (Fitzpatrick

1983). It has also been introduced to several areas outside of its natural

range (Hobbs et al. 1976). Hobbs et al. (1976) report that the species is

most abundant where vegetation is dense, as is usually the condition in

Santee Refuge impoundments and ditches. The life history of the species

at the refuge agrees with reported studies: Ovigerous females occur in

spring; young appear in midsummer; young increase in size until they

reproduce the following spring, soon after which death occurs. Fleming

(1969) and Strenth (1976) have emphasized the importance of setal

characteristics of the appendix masculina when distinguishing among

different species of Palaemonetes. Both of these authors report that P.

paludosus has four apical setae on this structure. However, I found this

characteristic to be variable, with specimens collected having from four

to six apical setae. Significant variations in spine position on the telson

were also observed.

Of the four species of crayfish found, only one ( Procambarus

blandingii) was common. The other three species ( P . hirsutus , P.

Branchiopod and Malacostracan Crustaceans

23

Table 8. Seasonal abundance of Palaemonetes paludosus at stations where the

species was collected at least once during the study. Data are expressed

as percentage of stations at which the species was abundant or absent.

Data from 1982 and 1983 have been combined.

troglodytes , Fallicambarus uhleri ) were each found in only one location.

No brooding females of any species were collected.

Procambarus blandingii was commonly encountered throughout

the refuge in habitats ranging from vegetated margins of Lake Marion

to ditches, impoundments, and shaded swamps. Juveniles were found in

all seasons, but were particularly abundant in June. This species has

been reported from lentic and lotic sites in southern North Carolina and

from the Santee River to the Pee Dee River in South Carolina. Its type

locality is in Kershaw County, S.C. (Hobbs 1974).

Procambarus hirsutus was found only in one sample taken in

emergent vegetation along the edge of Lake Marion. Because immature

specimens are easily confused with those of P. blandingii , this species

may be more widespread in the refuge than this one sample indicates.

Procambarus hirsutus has been previously reported from streams in the

Edisto, Salkehatchie, and Savannah drainage systems in South Carolina.

Its type locality is Barnwell County, S.C. (Hobbs 1958a).

Procambarus troglodytes was found in one location that consisted

of a series of small unshaded puddles resulting from excavation work

less than 100 m from Lake Marion. Individuals were abundant in June

in these puddles, where they constructed large chimneys. Procambarus

blandingii occurred in some puddles and an impoundment within 30 m

of the puddles used by P. troglodytes ; however, neither species was

collected from a puddle inhabited by the other. Procambarus troglodytes

occurs in lentic and lotic habitats between the Altamaha River in

Georgia and the Pee Dee River in South Carolina (Hobbs 1974).

Fallicambarus uhleri was found in only one locality, a borrow pit,

and in small numbers. Procambarus blandingii occurred in much greater

24

Charles K. Biernbaum

numbers in this same borrow pit. Juveniles of F. uhleri were collected in

September and November. Fallicambarus uhleri occurs in lentic and

lotic habitats along the coastal plain from Maryland to South Carolina

(Hobbs 1974).

Horton H. Hobbs, Jr., stated (pers. comm.) that other crayfishes

are found near the refuge and may occur, perhaps rarely, in the refuge.

These include Cambarus ( Depressicambarus ) latimanus (Le Conte),

Cambarus ( Depressicambarus ) reflexus Hobbs, Cambarus ( Puncti -

cambarus ) acuminatus Faxon, and Procambarus ( Ortmannicus ) enplo-

sternum Hobbs. It is also possible that Procambarus ( Ortmannicus )

ancylus Hobbs and Cambarus {La cuni cambarus) diogenes diogenes

Girard may be encountered.

SUMMARY

Diversity of crustaceans within the Santee National Wildlife Refuge

is quite high, especially for the cladocerans. The high diversity of

cladocerans is due in part to the variety of aquatic habitats available,

from open lake to ditches and swamps, but probably is chiefly due to

the great amount of vegetation occurring in most of the impoundments,

in ditches, and along the margin of the lake. As suggested by Lemly and

Dimmick (1982) in a study of zooplankton in the littoral zone of some

North Carolina lakes, a large quantity of aquatic macrophytes provides

great habitat heterogeneity, resulting in increased species diversity in

vegetated areas (Brooks 1959). Of the 25 cladoceran species collected at

the refuge, at least 17 are known to be directly associated with vegetation

(Brooks 1959).

Crustaceans of the refuge go through significant seasonal fluctua-

tions in abundance and dominance. Such seasonal changes are correlated

with temperature changes as well as with such seasonal variations as the

amount of water in shallow aquatic habitats and the late-summer

deterioration of certain habitats owing to prolific growth of vegetation

and subsequent decomposition.

In winter the crustacean fauna of the refuge is dominated by the

cladocerans Simocephalus serrulatus, Eurycercus vernalis, and, to a

lesser extent, Ilyocryptus spinifer; the isopods Caecidotea forbesi and

Lirceus lineatus; the amphipods Hyalella azteca and Crangonyx r.

richmondensis ; mature Palaemonetes paludosus; and the crayfish

Procambarus blandingii. Early spring is characterized by reductions in

numbers of cladocerans, Lirceus lineatus , and Crangonyx r. rich-

mondensis, with an increase in numbers of Hyalella azteca. In late

spring the refuge has very high water levels, resulting in expanded

habitats for swamp- and ditch-dwelling Caecidotea forbesi and Pro-

cambarus blandingii. By June there are large numbers of juvenile

Branchiopod and Malacostracan Crustaceans

25

crayfish present. In addition, in June there are population pulses of

several cladoceran species, including Simocephalus serrulatus , Eurycer-

cus vernalis, Ceriodaphnia reticulata , Scapholeberis kingi, and Sida

crystallina. The occurrence of one or two population pulses during a

year is characteristic of many cladoceran species (Pennak 1978).

In summer, the cladoceran fauna, with the exception of a few

species, is again generally reduced in numbers. Peracarids that are

commonly encountered include Hyalella azteca, Caecidotea laticaudata,

and reduced numbers of Caecidotea forbesi. Dryness eliminates many

shallow-water habitats for Caecidotea forbesi and, in late summer, this

aridity, as well as the growth and decomposition of large quantities of

vegetation in certain locations, leads to reductions in numbers of

Hyalella azteca and other crustaceans at several sites. Populations of

Palaemonetes paludosus consist of large numbers of postlarvae and

juveniles in summer.

Late autumn is characterized by increasing numbers of Caecidotea

forbesi , Lirceus lineatus, and Crangonyx r. richmondensis , and

decreasing numbers of Caecidotea laticaudata and, to a moderate

extent, Hyalella azteca. Large numbers of Palaemonetes paludosus

mature during this period.

Two habitats on the refuge are quite distinctive and warrant further

investigation. Dingle Pond is unique in the refuge in that all seven

peracarid species occur there. The pond itself is in the latter stages of

hydrarch succession, characterized by a mixture of connected open

pools and swamps, with scattered tree- and bush-covered hummocks.

The second habitat is in Cuddo unit; it consists of a small, forest-

surrounded borrow pit encircled by a series of small, swampy pools. In

it I found Procambarus blandingii, Fallicambarus uhleri , Caecidotea

forbesi , Crangonyx r. richmondensis , Simocephalus exspinosus, and,

occasionally in summer, Streptocephalus seali and Eulimnadia ven-

tricosa. Streptocephalus seali and Eulimnadia ventricosa were restricted

to shallow pools bordering the borrow pit during low water when fishes

were abundant but limited to the borrow pit. That these two crustacean

species disappeared when the water rose, making the pools and borrow

pit confluent, suggests that predation by fishes may eliminate them. The

usual absence of non-cladoceran branchiopods from bodies of water

containing fishes is well known and has been thought to be the result of

predation on these large, easily seen, and almost totally defenseless

species (Pennak 1978).

ACKNOWLEDGEMENTS. — I thank the Managers (Paul Ferguson

and Glen Bond) and staff of the Santee National Wildlife Refuge for

permission to do the study and for their assistance. I also thank J. R.

26

Charles K. Biernbaum

Harrison, III, for suggesting the study and for assistance in sampling;

H. H. Hobbs, Jr., for assisting with crayfish identifications and providing

information on the occurrence of crayfish species in the area; N. E.

Strenth for examining specimens of Palaemonetes paludosus\ J.

Pinckney for amphipod specimens removed from a duck; and W. D.

Anderson, Jr., N. A. Chamberlain, and J. R. Harrison, III, for reviewing

the manuscript. Figure 1 was prepared by K. Swanson, and the

manuscript typed by C. Baldwin.

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Branchiopod and Malacostracan Crustaceans 27

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reproductive capacity of some fresh-water amphipods. Int. Rev. Gesamten

Hydrobiol. (Suppl.) 4:1-33.

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of Orangeburg County, South Carolina. Trans. Am. Microsc. Soc.

7 1(3):272-276.

Ferguson, Edward, Jr. 1954. A new ostracod of the genus Cypricerus from

South Carolina. Trans. Am. Microsc. Soc. 73(2): 189-192.

Fitzpatrick, Joseph F., Jr. 1983. How to Know the Freshwater Crustacea.

Wm. C. Brown Co., Dubuque, Iowa.

Fleming, Lawrence E. 1969. Use of male external genitalic details as taxonomic

characters in some species of Palaemonetes (Decapoda, Palaemonidae).

Proc. Biol. Soc. Wash. 82:443-452.

Fleming, Lawrence E. 1972. The evolution of the eastern North American

isopods of the genus Asellus (Crustacea: Asellidae). Int. J. Speleol.

4(3-4):22 1-256.

Fox, Richard S. 1978. Subclass Malacostraca: Superorder Peracarida: Order

Amphipoda. Pages 161-166 in An Annotated Checklist of the Biota of the

Coastal Zone of South Carolina, R. G. Zingmark, editor. Univ. South

Carolina Press, Columbia.

Frey, David G. 1980. On the plurality of Chydorus sphaericus (O. F. Muller)

(Cladocera, Chydoridae), and designation of a neotype from Sjaels0,

Denmark. Hydrobiologia 69(l-2):83-123.

Frey, David G. 1982a. Contrasting strategies of gamogenesis in northern and

southern populations of Cladocera. Ecology 63( 1 ): 223-24 1 .

Frey, David G. 1982b. The reticulated species of Chydorus (Cladocera,

Chydoridae): two new species with suggestions of convergence.

Hydrobiologia 93(3): 255-279.

Frey, David G. 1982c. Questions concerning cosmopolitanism in Cladocera.

Arch. Hydrobiol. 93:484-502.

Garthwaite, Ronald, and C. Sassaman. 1985. Porcellionides floria , new

species, from North America; provinciality in the cosmopolitan isopod

Porcellionides pruinosus (Brandt). J. Crust. Biol. 5(3):539-555.

Gibbes, L. R. 1850. On the carcinological collection of the cabinets of natural

history in the United States. With an enumeration of the species contained

therein and descriptions of new species. Proc. Am. Assoc. Adv. Sci.

3:165-201.

28

Charles K. Biernbaum

Giesy, John P., J. W. Bowling, and H. J. Kania. 1980. Cadmium and zinc

accumulation and elimination by freshwater crayfish. Arch. Environ.

Contam. Toxicol. 9:683-697.

Hann, Brenda J. 1982. Two new species of Eurycercus ( Bullatifrons ) from

eastern North America (Chvdoridae, Cladocera). Taxonomy, ontogeny,

and biology. Int. Rev. Gesamten Hydrobiol. 67(4):585-610.

Hargrave, Barry T. 1970. Distribution, growth, and seasonal abundance of

Hvalella azteca (Amphipoda) in relation to sediment microflora. J. Fish.

Res. Board Can. 27(4): 685-699.

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Mus. Leafl. 14:3-7.

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the genus Procambarus , with the description of a new species from South

Carolina. Fla. Entomol. 30(3):25-31.

Hobbs, Horton H., Jr. 1956a. A new crayfish of the genus Procambarus from

South Carolina (Decapoda, Astacidae). J. Wash. Acad. Sci. 46(4): 1 17-121.

Hobbs, Horton H., Jr. 1956b. A new crayfish of the genus Cambarus from

North Carolina and South Carolina (Decapoda, Astacidae). J. Elisha

Mitchell Sci. Soc. 72( 1 ):6 1 -67.

Hobbs, Horton H., Jr. 1956c. A new crayfish of the extraneus section of the

genus Cambarus with a key to the species of the section (Decapoda.

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Hobbs, Horton H., Jr. 1983. Distocambarus (Fitzcambarus) carlsoni . a new

subgenus and species of crayfish (Decapoda: Cambaridae) from South

Carolina. Proc. Biol. Soc. Wash. 96(3):429-439.

Hobbs, Horton H., Jr., and P. H. Carlson. 1983. Distocambarus (Decapoda:

Cambaridae) elevated to generic rank, with an account of D. crockeri new

species, from South Carolina. Proc. Biol. Soc. Wash. 96(3):420-428.

Hobbs, Horton H., Ill, J. H. Thorp, and G. E. Anderson. 1976. The

Freshwater Decapod Crustaceans (Palaemonidae. Cambaridae) of the

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Environ. Park Prog., Aiken.

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Branchiopod and Malacostracan Crustaceans

29

Kelley, B. J., Jr. 1978. Order Isopoda. Pages 167-170 in An Annotated

Checklist of the Biota of the Coastal Zone of South Carolina, R. G.

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amphipods Hyalella azteca and Crangonyx richmondensis occidentalis in

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of the fauna and flora of the Savannah River. Proc. Acad. Nat. Sci. Phila.

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Prins, Rudolph, and H. H. Hobbs, Jr. 1972. A new crayfish of the subgenus

Puncticambarus from the Savannah River drainage, with notes on

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Wash. 84(47):41 1-420.

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coastal sites in Georgia. Bull. Ga. Acad. Sci. 34:185-194.

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Name) and M. morganensis n. comb., reconsidered with notes on New

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Nat. 95( 1 ): 28-4 1 .

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freshwater species of Palaemonetes Heller of North America (Crustacea:

Decapoda). Smithson. Contrib. Zool. 228:1-27.

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1 00(2):47 1-474.

30

Charles K. Biernbaum

Thorp, James H., and E. A. Bergey. 1981. Field experiments on responses of a

freshwater, benthic macroinvertebrate community to vertebrate predators.

Ecology 62(2):365-375.

Vigerstad, T. J., and L. J. Tilly. 1977. Hyperthermal effluent effects on

heleoplanktonic Cladocera and the influence of submerged macrophytes.

Hydrobiologia 55( 1 ): 8 1 -85.

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Prot. Agcy., Washington, D.C.

Accepted 14 November 1986

ADDENDUM

In June 1986, two specimensbelonging to an undescribed amphipod

species in the genus Gammarus were collected from the sandy bottom

of Lake Marion adjoining Cuddo unit. This species belongs in the G.

fasciatus-tigrinus complex that, according to Fox (1978), includes two

or three undescribed species from South Carolina to Louisiana.

Taxonomic Analysis of the Coastal Marsh Raccoon

( Procyon lotor maritimus) in Maryland

Denise H. Clearwater, George A. Feldhamer,1

Raymond P. Morgan II, and Joseph A. Chapman2

Appalachian Environmental Laboratory,

Center for Environmental and Estuarine Studies,

University of Maryland, Frostburg State College Campus,

Frost burg, Maryland 21532

ABSTRACT. — Skulls of raccoons were collected from three

physiographic regions in Maryland. Based on multivariate analyses of

skull measurements, it is suggested that the coastal marsh raccoon,

Procyon lotor maritimus , be considered a synonym of P. 1. lotor.

Most taxonomic studies describing subspecies of raccoons ( Procyon

lotor) were based on qualitative cranial characteristics and pelage (see

Goldman 1950). Pelage is highly variable within geographic regions and

is a poor diagnostic character. The coastal marsh raccoon (P. 1.

maritimus) was described by Dozier (1948) on the basis of 34 specimens

(skins, skulls, or both) from marsh habitats on the Delmarva Peninsula.

According to Dozier (1948), skulls of P. 1. maritimus were smaller and

narrower than those of P. 1. lotor and had shorter postorbital processes.

He also stated that the two subspecies occupied different habitats on the

Delmarva Peninsula, with P. 1. maritimus in marshes and P. 1. lotor in

upland wooded areas. Neither the cranial nor the habitat differences

were quantified, however, and not all researchers accepted the coastal

marsh raccoon as a new subspecies. Paradiso (1969:145) felt that the

differences in P. 1. maritimus were “slightly marked” and within the

limits of individual variation of P. 1. lotor. Without commenting on the

validity of current taxonomy, Hall and Kelson (1959) and Hall (1982)

treated P. 1. maritimus as a valid subspecies. Our objective was to

determine if P. 1. maritimus was a valid taxon based on statistical

analyses of cranial characteristics.

MATERIALS AND METHODS

Skulls of male raccoons (n = 63) from the museum collections of

the Appalachian Environmental Laboratory and the National Museum

'Present address: Department of Zoology, Southern Illinois University,

Carbondale, Illinois 62901.

2Present address: Department of Fisheries and Wildlife, College of Natural

Resources, Utah State University, Logan, Utah 84322.

Brimleyana No. 15:31-36, January 1989

31

32

Clearwater, Feldhamer, Morgan, and Chapman

of Natural History were used in the analysis. Females were excluded

because of small, unequal sample sizes among groups. Adults only were

examined and distinguished from juveniles according to tooth wear

(Grau et al. 1970), closure of parietal sutures, and month of capture.

Raccoon skulls were grouped according to capture location in

Maryland into one of three physiographic regions (see Paradiso 1969):

(1) western, which included the Appalachian Plateau and Ridge and

Valley regions (n = 21); (2) central, which included the Piedmont region

(n = 23); and (3) eastern, which included the Coastal Plain region (n =

19). Skulls from the two initial groups represented P. 1. lot or. The third

group represented P. 1. maritimus, based on all captures being from the

type locality, Blackwater National Wildlife Refuge, Dorchester County,

adjacent Fishing Bay, or similar marsh habitats in five other counties on

Maryland’s eastern shore. Comparisons also were made with skulls of

P. l.fuscipes (n = 16) from Texas and Mexico.

Differences among groups were determined by analysis of variance

of 12 skull measurements (see Table 2), with Duncan’s Multiple Range

test, and by stepwise discriminant function analysis, after elimination of

correlated variables (r > 0.70). The selection criterion was the maximum

Mahalanobis distance. Analyses were conducted using programs of

SPSS (Nie et al. 1975).

Table 1. Actual and predicted group membership for male raccoon skulls from

western and central Maryland ( P . /. lotor), eastern shore of Maryland

( P . /. maritimus), and Texas ( P . l.fuscipes).

RESULTS AND DISCUSSION

In the discriminant function analysis of the three Maryland groups

and P. 1. fuscipes (Fig. 1), width of braincase had the highest standardized

coefficient (-0.849) in the first function, followed by basilar length

(0.830). The first function explained 69.0% of the variation among

groups. In the second function, width of rostrum (0.505) and width of

Table 2. Means ± one standard deviation, ranges (mm), and F values for 12 cranial characteristics from adult male raccoon

( Procyon lotor ) skulls from three regions of Maryland. * = P < 0.01.

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34

Clearwater, Feldhamer, Morgan, and Chapman

postorbital process (0.385) were the most important discriminating

variables and accounted for an additional 22.9% of among-group

variation. The overall percentage of individuals correctly classified was

only 63.3% (Table 1). Because all cases were used to derive the functions,

this percentage actually overestimated the accuracy of the functions

(Williams 1983). That the correct-classification rate was inflated is

further suggested by the calculation of Cohen’s kappa = 0.447 (Z = 4.96,

P < 0.0001), the chance-corrected percentage of agreement between

actual and predicted group memberships (Titus et al. 1984). Specimens

from central Maryland overlapped considerably with the western and

eastern groups. There also was considerable overlap among Maryland

raccoons and P. l.fuscipes, possibly because subadults may have been

included in the latter sample.

Significant differences (ANOVA F-test, P < 0.05) among the three

Maryland groups occurred in four skull variables. Duncan’s Multiple

Range test indicated the width of rostrum, width of braincase, and

width of postorbital processes were significantly smaller in the eastern

group; western and central groups were not different. The western

group had significantly shorter incisive foramina (Table 2). This supports

Dozier’s statement (1948) that the skull of P. /. maritimus is narrower

than in raccoons from western or central Maryland.

There is no ecological or geographic evidence (Vokes 1957) that

raccoons in Maryland were ever isolated. Dozier’s (1948) statements

that P. /. maritimus is restricted to marshland are not necessarily

supported by recent telemetry studies. Sherfy and Chapman (1983)

found that radiocollared raccoons at Blackwater National Wildlife

Refuge alternately used marsh and woodland as activity areas, although

sample size (n = 2) was very small. Thus, there probably is no ecological

barrier to isolate what Dozier termed maritimus from lotor. Raccoons

are highly mobile with observed morphological variation among

populations often explained by elevation, temperature, and habitat

factors (Kennedy and Lindsay 1984) as well as genetic considerations.

Although there is no clear consensus on subspecific criteria, we concur

with Paradiso’s (1969) opinion that the morphological differences among

Maryland raccoons do not justify designating those on the eastern shore

as a subspecies, and P. L maritimus Dozier should be considered a

synonym of P. /. lotor (Linnaeus).

ACKNOWLEDGMENTS. — We thank Kimberly Titus and Richard

Highton for critically reviewing an early draft of the manuscript. J.

Dunn, Appalachian Environmental Lab, and R. Fisher, National

Museum of Natural History, assisted in data collection. Computer time

Coastal Marsh Raccoon

35

Discriminant Function 1

Fig. 1. Ninety-five percent confidence ellipses around the group centroids for

raccoon skulls from (1) western and (2) central Maryland (P. L. lotor)\ (3) marsh

areas of Maryland’s eastern shore ( P . /. maritimus ); and (4) P. /. fuscipes from

Texas and Mexico. Function 1, from left to right, represents a gradient of

increasing skull length and decreasing width. Function 2, from bottom to top,

represents a gradient of increasing skull width.

was provided by the Computer Science Center of the University of

Maryland. This is Scientific Series No. 1752-AEL, Appalachian

Environmental Laboratory, University of Maryland.

LITERATURE CITED

Dozier, Herbert L. 1948. A new eastern marsh-inhabiting race of raccoon. J.

Mammal. 29:286-290.

Goldman, Edward A. 1950. Raccoons of North and Middle America. N.

Amer. Fauna No. 60.

Grau, Gerald A., G. C. Sanderson, and J. P. Rogers. 1970. Age determination

of raccoons. J. Wildl. Mgmt. 34:364-372.

Hall, E. Raymond. 1982. The Mammals of North America. 2 vol. John Wiley

and Sons, New York.

36

Clearwater, Feldhamer, Morgan, and Chapman

Hall, E. Raymond, and K. R. Kelson. 1959. The Mammals of North America.

2 vol. The Ronald Press Co., New York.

Kennedy, Michael L., and S. L. Lindsay. 1984. Morphological variation in the

raccoon, Procyon lotor, and its relationship to genic and environmental

variation. J. Mammal. 65:195-205.

Nie, Norman H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and D. H. Bent

(editors). 1975. Statistical Package for the Social Sciences. McGraw-Hill

Book Co., New York.

Paradiso, John L. 1969. Mammals of Maryland. N. Amer. Fauna No. 66.

Sherfy, Fred C., and J. A. Chapman. 1983. Seasonal home range and habitat

utilization of raccoons in Maryland. Carnivore 4:8-18.

Titus, Kimberly, J. A. Mosher, and B. K. Williams. 1984. Chance-corrected

classification for use in discriminant analysis: ecological applications. Amer.

Midi. Nat. 111:1-7.

Vokes, Harold E. 1957. Geography and Geology of Maryland. Md. Dep.

Geology, Mines, and Water Resour. Bull. No. 19.

Williams, Byron K. 1983. Some observations on the use of discriminant

analysis in ecology. Ecology 64:1283-1291.

Accepted 17 November 1986

Tolerance of Acidity in a Virginia Population of the

Spotted Salamander, Ambystoma maculatum

(Amphibia: Ambystomatidae)

Charles R. Blem and Leann B. Blem

Virginia Commonwealth University, Department of Biology,

Academic Division, Richmond, Virginia 23284

ABSTRACT. — We investigated apparent acid tolerance of egg masses

of the spotted salamander, Ambystoma maculatum , in the lower

piedmont and coastal plain of Virginia. All temporary ponds that we

tested in east-central Virginia were acidic (i.e., pH < 6.0), and the

majority (about 93%) had no ambystomatid egg masses in them during

the spring breeding season. However, a few ponds continued to

support successful spotted salamander populations, even though their

pH levels were below those that caused extensive mortality of eggs and

larvae in the laboratory. In laboratory tests, more than 50% of the eggs

removed from these ponds survived pH levels of 4.3 to 4.7, but no eggs

or larvae tolerated experimental exposure to a pH less than 4.0. Local

tolerance of low pH may result from natural selection of resistant

individuals through long exposure to the acidic, hoggish waters of the

coastal plain and lower piedmont. However, we hypothesize that

aluminum may be involved in apparent resistance to low pH. Some

coastal plain and piedmont soils are low in aluminum, and large

amounts of organic materials may bind and inactivate the aluminum

that is present. Lowered levels of dissolved aluminum may permit

survival of salamander embryos/ larvae at a low pH. This hypothesis is

supported by increased mortality of embryos in test containers to

which small (< 0.3 ppm) amounts of aluminum were added. Soils and

temporary ponds of the lower piedmont and coastal plain of Virginia

are naturally acidic and have little buffering capacity. Future

acidification of the environment as a result of acid precipitation is

likely in this region. Our tests indicate that slight decreases in pH of

temporary ponds may result in eradication of the spotted salamander

from this part of Virginia.

Tolerance of low pH by amphibians recently has received much

attention (for a review, see Pierce 1985). In general, most adult

amphibians are relatively tolerant of acidity or conditions associated

with low pH, but eggs and larvae may suffer extreme mortality under

mildly acidic conditions (Saber and Dunson 1978, Dunson and Connell

1982). Eggs and larvae of the spotted salamander, Ambystoma

maculatum , seem to be especially sensitive to low pH, and the species

may be decreasing in number at some locations because of acidification

of breeding ponds resulting from deposition of atmospheric acid (i.e.,

acid rain; see Pough 1976, Pough and Wilson 1977).

Brimleyana No. 15:37-45, January 1989

37

38

Charles R. Blem and Leann B. Blem

Pough (1976) and Pough and Wilson (1977) found that New York

populations of the spotted salamander suffered high mortality of larvae

at a pH lower than 6.0, and the species has declined or disappeared

from parts of the northeastern United States. However, Cook (1983),

working with spotted salamanders at sites within the Connecticut Valley

of Massachusetts, found that the percent of mortality of embryos /larvae

was small (i.e., usually less than 20%) in ponds of pH 4.2 to 6.0. He

concluded that survival of embryos /larvae in some populations of the

spotted salamander indicated embryonic acid tolerance, an idea echoed

by others (see Pierce 1985). The present study examines a similar

apparent tolerance of acidity in coastal plain and piedmont populations

of the spotted salamander in Virginia.

MATERIALS AND METHODS

Studies were conducted in the spring (February - April) of 1983

through 1986. Initially we surveyed temporary ponds in six counties

(Charles City, Chesterfield, Goochland, Hanover, Henrico, and

Powhatan) of the coastal plain and piedmont of east-central Virginia.

We specifically looked for egg masses, larvae, and spermatophores in all

168 temporary ponds we encountered.

In 64 of the ponds, we measured pH by means of an electronic,

portable meter (Digital Mini-pH meter) and confirmed these

determinations with two laboratory meters (Corning and Orion), Merck

colorpHast pH paper, or both. The pH of the remaining 104 ponds was

not measured, in the interest of time, because the ponds were near

others whose pH was known, or for both reasons. Although the point is

seldom mentioned in the literature, pH is difficult to measure in low

conductivity waters such as those encountered in temporary ponds. We

therefore were careful to duplicate most determinations with at least

two different meters, or with a meter and pH paper. Different techniques

never produced pH’s that differed by more than 0.2; duplicate

measurements were averaged in those instances where two methods

were used. Additionally, we confirmed most determinations by adding

small amounts of KC1 to aliquots of the solution to be tested before

using a pH meter. This increases the conductivity of the solution and

increases the accuracy and speed of measuring pH (F. Hawkridge, pers.

comm.).

Tests of pH-related mortality were performed on A. maculatum egg

masses that were removed from a temporary pond in Chesterfield

County, Virginia. This site is within the coastal plain and is acidic

(mean pH = 4.35 ± 0.30; calculated from 22 determinations made on

different days over a span of 4 years). One-liter aquaria filled with water

from the temporary pond were used for all tolerance tests as well as

Salamander Tolerance of Acidity

39

controls. All water was filtered through cheesecloth before eggs were

added. All tests were carried out in a controlled temperature (15°C)

cabinet at a photoperiod of 12L:12D. Oxygen levels in the aquaria

remained high (i.e., near saturation) through the tests and were monitored

periodically with an electronic oxygen meter.

Two egg masses were placed in each aquarium. Although we

measured masses having a volume greater than 200 cm3 and containing

more than 200 eggs, the volume of masses in our tests was 65.2 ± 10.2

cm3 (mean ± one standard error). These contained 57.9 ±0.8 eggs /mass.

Experimental groups included tests of survival of eggs in pond

water with acid added (either H2S04 or HNO3), or with aluminum

added. Aluminum used in all tests was in the form of A1 (S04)2. Acid

from a concentrated stock solution was added in tiny amounts until a

pH of 3.0 to 4.0 was obtained. The buffering capacity of individual egg

masses then caused acidity to change. This resulted in pH values of 3.1

to 4.0 (see Table 2). Aluminum sulfate was added in amounts sufficient

to produce 0.05, 0.10, or 0.23 ppm aluminum in the solution. Controls

were performed in exactly the same manner as experimental groups. In

addition, we tested survival of eggs in local spring water having a

slightly higher pH and little or no dissolved organic materials (pH =

4.65 ± 0.30; N = 5).

Egg masses were inspected daily, and embryos were classified

according to Harrison stage (Pough 1976; modification of Rugh 1962).

Mortality of salamander eggs may be a function of shrinkage of the

gelatinous matrix surrounding the eggs. Accordingly, the length and

width of each egg mass was measured with a plastic ruler at the

beginning of each test, and at 3, 6, 14, and 25 days. Volume of

individual masses was then computed from the prolate spheroid formula:

V = 0.523 AB2, where A is the length of the mass (cm), B is the width,

and V is the volume in cm . All free-swimming larvae were removed

within 48 hours of hatching, examined for morphological defects, and

released into a central holding tank. No long-term tests of mortality of

larvae were performed; references to larval survival in the present paper

are valid only for the 24 hours after hatching. Most surviving larvae

were returned to the breeding pond.

RESULTS

All temporary ponds (64) whose pH was known were acidic, and

the majority (63/64 = 98.4%) had pH values during the spring Ambystoma

breeding season that were below levels known to cause mortality (i.e.,

pH < 6.0; see Pough 1976, Pough and Wilson 1977, Cook 1983). A

small percentage (5/64 = 7.8%) had pH’s of less than 4.0 (Table 1). Of

the total of 168 ponds that were surveyed, only a few (12/ 168 = 7.1%)

40

Charles R. Blem and Leann B. Blem

had egg masses or larvae in them during the breeding season. However,

two ponds with pH’s less than 4.5 produced larvae, and one of these, a

large temporary pond in Chesterfield County, had more than 300 egg

masses in it during each of the five breeding seasons from 1982 through

1986.

Mortality of controls was highly variable among egg masses in the

laboratory, but in general was relatively low for water with such low pH

values (Table 2; see Pough 1976, Pough and Wilson 1977). Control

results for 1984 did not differ significantly (t = 1.6; p < 0.05) from those

of 1985, and neither of these groups was different from spring-water

tests (t = 1.3 and 0.4, respectively; p < 0.05). Mortality in control groups

occurred during early embryonic stages, that is, stages 1 through 9

(Rugh 1962). The only abnormalities noted were deformities of the

posterior trunk of hatched larvae and there were relatively few of these

(see Pough 1976).

Very small amounts of sulfuric or nitric acid decreased the pH of

test waters severely, indicating that water from these temporary ponds is

poorly buffered. Furthermore, even small decreases in pH resulted in

nearly total mortality of eggs/ larvae (Table 2). Mortality appeared to be

caused mainly by severe shrinkage of the total egg mass, as described by

Pough (1976). Shrinkage did not occur at pH levels of the natural ponds

(Fig. 1). In fact, egg masses increased in size as expected in newly

deposited eggs (Pough 1976). Acidity of pH less than 4.0 resulted in

shrinkage of egg masses, in some cases to less than one-third of the

original size (see Fig. 1). Shrinkage was roughly a function of pH, that

is, the lower the pH, the greater the amount of shrinkage. The most

severe cases resulted in egg masses less than one-fourth the expected

size.

Addition of aluminum sulfate to test aquaria resulted in higher

mortality, and the mortality rate increased rapidly with very small

increases in aluminum sulfate (Table 3). Slight increases of aluminum

(0.05 ppm) did not significantly increase mortality as compared with

pooled controls (t = 0.8), but 0. 10 and 0.23 ppm caused significant (p <

0.05) increases in mortality (t = 2.1 and 3.8, respectively).

DISCUSSION

The observation that one low-pH pond apparently has supported a

very large, successful population of spotted salamanders while few

others have done so, suggests that this population is acid-tolerant or

that the pond has one or more characteristics that reduce the deleterious

effect of low pH. Our control tests indicate that tolerance of acidity was

greater than that reported by Pough (1976) and was similar to that of an

acid-tolerant population investigated by Cook (1983). Field observations,

Salamander Tolerance of Acidity

41

Table 1. Acidity of 64 temporary ponds in the lower piedmont and coastal

plain of east-central Virginia. Sample size (N) represents individual

ponds having given pH values during early spring.

Table 2. Percent mortality of spotted salamander eggs exposed to different

aquatic media. Mortality rates are means ± one standard error;

ranges of mortality are in parentheses. N is the number of tests, two

egg masses per test. All egg masses were taken from a temporary pond

in Chesterfield County, Virginia.

42

Charles R. Blem and Leann B. Blem

as well as the great number of egg masses found in the Chesterfield

County pond, indicate that natural reproduction was successful there.

One hypothesis is that such success may represent evolution of acid

tolerance as a result of long exposure to low pH in this pond (e.g.,

Tome and Pough 1982, Cook 1983). However, there is another plausible

explanation. While low pH can have direct effects upon amphibian

larvae (Pough 1981, Freda and Dunson 1984), external cation

concentration may be the source of the damage (Hall and Likens 1981,

Freda and Dunson 1985; but see Dale et al. 1985). In most natural

waters, decreases in PH would increase the amount of free aluminum

(and other cations) available to aquatic organisms (see Hall and Likens

1981, Clark and Hall 1985). Decreasing pH may then cause mortality by

the increase in toxic aluminum, rather than by direct pH effects.

Aluminum has long been known to be detrimental to fish, particularly

their eggs and larvae (Freeman and Everhart 1981, Schofield 1980), and

it appears to have a similar effect on amphibian larvae (Clark and Hall

1985).

Our tests of aluminum toxicity (Table 3) support the hypothesis

that aluminum is harmful to spotted salamander eggs. It should be

noted that addition of Al (S04)2 decreased the pH of the aquaria media

slightly as a result of the addition of S04 ions (Table 3). The extent of

the extra acidity was only slight; this was not expected to affect egg

mortality because we found no significant correlation between mortality

and pH in the pH range of 4.2 to 4.8 (Table 2). Very small amounts of

aluminum increased mortality of larvae to the point that we estimate (by

extrapolation) that total mortality should occur at increases of 0.28 ppm

aluminum. This is similar to levels of toxicity for brook trout ( Salvelinus

fontinalis) eggs at similar pH’s (Hall and Likens 1981). This by no

means proves a causal relationship between aluminum concentration

and egg mortality, but does suggest that the hypothesis deserves further

testing.

Acidity of many temporary ponds in the coastal plain and eastern

piedmont of east-central Virginia is near the minimum tolerance levels

for spotted salamander embryos /larvae (see Pough 1976). Only one

larva survived any of our tests in which the pH was 4.0 or less, and

mortality in excess of 50% was observed in aquaria at pH’s of 4.3 to 4.4

or less. This indicates that only a slight further acidification of local

temporary ponds in east-central Virginia may result in total extirpation

of the species from the area.

Hatching success of spotted salamanders in acidic pools depends on

at least two factors. The first is the volume of the egg mass. However, in

the present study, there was no obvious relationship between mortality

rate and size of the egg mass. We excluded both very small and large,

Salamander Tolerance of Acidity

43

Fig. 1. Volume of spotted salamander egg masses as a percentage of the

original volume. The dotted line indicates no change.

irregular egg masses from the study because these may not be

representative of normal, single reproductive attempts. As a result, our

experiments were performed using egg masses that were rather uniform.

Although large egg masses may have a beneficial effect on larvae deep

within the mass by providing local buffering, we observed that mortality

was largely a matter of death of most eggs within single masses. Pough

(1976) noted that the buffering capacity of the gelatinous matrix of

ambystomatid egg masses was slight, but found that the few surviving

embryos in acid waters tended to be in the centers of masses. Evolution

of acid tolerance may take the form of natural selection of larger egg

masses, and this hypothesis deserves consideration. Second, hatching

success varies because temporary pools differ greatly in chemical

44

Charles R. Blem and Leann B. Blem

composition, depending upon the local soil composition and the nature

of the decomposing organic material in the water. For example, in our

study area, soil maps indicate a mosaic of soils that vary in acidity and

aluminum content (Hodges 1978). The local vegetation around breeding

ponds may be pines, hardwood forest, or old field. The acidity of

breeding pools may be increased by pine needles and/or oak leaves

through humic acids produced because of poor rates of decomposition,

but the products of other plants may not be so acidic (Smith 1986). The

interactions between pH, organic matter, and aluminum may be complex,

but in general it appears that high concentrations of organic compounds

may decrease available aluminum ions by binding with them (Pott et al.

1985).

In summary, the results of the present study indicate that we should

be cautious in attributing tolerance of acidity to populations of spotted

salamanders breeding in ponds of low pH. Even so, it appears that A.

maculatum in parts of Virginia has a dismal future because the slightest

increases in acidity may cause its extirpation.

ACKNOWLEDGMENTS. — We are indebted to the Department of

Biology, Virginia Commonwealth University, and particularly Martha

D. Berliner, for partial support of this study. We are grateful to Carolyn

Conway for the loan of equipment, to Fred Hawkridge for valuable

advice about water chemistry, and to Ralph Mendenhall for sharing

measurements of soil aluminum with us. Mark Zimmerman and members

of the 1984 physiological ecology class at Virginia Commonwealth

University helped make some measurements.

LITERATURE CITED

Clark, K. L., and R. J. Hall. 1985. Effects of elevated hydrogen ion and

aluminum concentrations on the survival of amphibian embryos and larvae.

Can. J. Zool. 63:116-123.

Cook, R. P. 1983. Effects of acid precipitation on embryonic mortality of

Ambystoma salamanders in the Connecticut Valley of Massachusetts. Biol.

Conser. 27:77-88.

Dale, J. M., B. Freedman, and J. Kerekes. 1985. Acidity and associated water

chemistry of amphibian habitats in Nova Scotia. Can. J. Zool. 63:97-105.

Dunson, W. A., and J. Connell. 1982. Specific inhibition of hatching in

amphibian embryos by low pH. J. Herp. 16:314-316.

Freda, J., and W. A. Dunson. 1984. Sodium balance of amphibian larvae

exposed to low environmental pH. Physiol. Zool. 57:435-443.

Freda, J., and W. A. Dunson. 1985. The influence of external cation

concentration on the hatching of amphibian embryos in water of low pH.

Can. J. Zool. 63:2649-2656.

Salamander Tolerance of Acidity

45

Freeman, R. A., and W. H. Everhart. 1971. Toxicity of aluminum hydroxide

complexes in neutral and basic media to rainbow trout. Trans. Amer. Fish.

Soc. 100:644-658.

Gosner, K. L., and I. H. Black. 1957. The effects of acidity on the development

and hatching of New Jersey frogs. Ecology 38:256-262.

Hall, R. J., and G. E. Likens. 1981. Chemical flux in an acid-stressed stream.

Nature 292:329-331.

Hodges, R. L. 1978. Soil survey of Chesterfield County, Virginia. U.S. Dept.

Agric., Washington, D.C.

Pierce, B. A. 1985. Acid tolerance in amphibians. BioScience 35:239-243.

Pough, F. H. 1976. Acid precipitation and embryonic mortality of spotted

salamanders, Ambystoma maculatum. Science 192:68-70.

Pough, F. H. 1981.' Mechanisms by which acid precipitation produces

embryonic death in aquatic vertebrates. U.S. Dept. Interior Office Res.

Tech., Res. Proj. Tech. Compl. Rept., 1-10.

Pough, F. H., and R. E. Wilson. 1977. Acid precipitation and reproductive

success of Ambystoma salamanders. Water, Air, Soil Poll. 7:307-316.

Pott, D. B., J. J. Alberts, and A. W. Elzerman. 1985. The influence of pH on

the binding capacity and conditional stability constants of aluminum and

naturally-occurring organic matter. Chem. Geol. 48:293-304.

Rugh, R. 1962. Experimental Embryology: A Manual of Techniques and

Procedures. 3rd Ed., Burgess Publ. Co., Minneapolis.

Saber, P. A., and W. A. Dunson. 1978. Toxicity of bog water to embryonic

and larval anuran amphibians. J. Exp. Biol. 204:133-142.

Schofield, C. L. 1980. Processes limiting fish populations in acidified lakes.

Pages 345-356 in Atmospheric sulfur deposition — environmental impact

and health effects, D. S. Shriner, C. R. Richmond, and S. E. Lindberg,

editors. Ann Arbor Sci., Ann Arbor, Michigan.

Smith, R. L. 1986. Elements of Ecology. Harper and Row, N.Y.

Tome, M. A., and F. H. Pough. 1982. Responses of amphibians to acid

precipitation. Acid Rain/ Fisheries, Amer. Fish. Soc., Bethesda, Md.,

245-254.

Accepted 5 May 1987

46

ATLAS OF NORTH AMERICAN FRESHWATER FISHES

by

D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins,

D. E. McAllister, J. R. Stauffer, Jr., and many collaborators

This very useful book provides accounts for all 777 species of fish

known to occur in fresh waters in the United States and Canada. Each

account gives a distribution map and illustration of the species, along

with information on systematics, distribution, habitat, abundance, size,

and general biology.

“. . . represents the most important contribution to freshwater fishes

of this continent since Jordan and Evermann’s ‘Fishes of North and

Middle America’ over 80 years ago.” — Southeastern Fishes Council

Proceedings.

1980 825 pages Index Softbound ISBN 0-917134-03-6

Price: $25, postpaid, North Carolina residents add 5% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send to FISH ATLAS, N.C. State Museum of Natural Sciences,

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

ATLAS OF NORTH AMERICAN FRESHWATER FISHES

1983 SUPPLEMENT

by

D. S. Lee, S. P. Platania, and G. H. Burgess

The 1983 supplement to the 1980 Atlas of North American

Freshwater Fishes treats the freshwater ichthyofauna of the Greater

Antilles. In addition to this bound supplement, there are 19 accounts,

mostly species not described in 1980, in looseleaf form to be added to

the 1980 volume. Illustrated by Renaldo Kuhler.

1983 67 pages Index Softbound ISBN 0917134-06-0

Price: $5, postpaid. North Carolina residents add 5% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send to FISH ATLAS, N.C. State Museum of Natural Sciences,

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

Population Structure and Biomass

of Sternotherus odoratus (Testudines: Kinosternidae)

in a Northern Alabama Lake

C. Kenneth Dodd, Jr.

National Ecology Research Center, U.S. Fish and Wildlife Service,

412 N.E. 16th Avenue, Room 250,

Gainesville, Florida 32601

ABSTRACT. — A population of the stinkpot, Sternotherus odoratus,

was sampled periodically during the summer of 1985 in a small lake in

northern Alabama. The population structure was similar to that

reported in other studies, although the sizes of turtles were intermediate

between populations located farther to the north and to the south. The

sex ratio was skewed toward females, but a relatively small sample size

hinders interpretation of the significance of this result. A population

density estimate of 148.5 turtles per hectare indicated a biomass of 10.6

kg/ ha. Mortality from drowning in traps was not influenced by the

sex, body mass, or size of the affected adult.

The stinkpot or common musk turtle, Sternotherus odoratus, is a

geographically widespread inhabitant of the eastern United States. This

omnivorous species frequents sluggish and still waters, particularly lakes

and ponds (Carr 1952, Ernst and Barbour 1972, Mount 1975). In

Alabama, stinkpots occur statewide (Mount 1975), and they inhabit

slow-moving or lentic waters, including large rivers such as the

Tennessee River.

In spite of the species’ ubiquitous nature, there have been relatively

few field studies of stinkpot ecology, and these for the most part have concentrated

on reproductive ecology (Tinkle 1961; Gibbons 1970a; Iverson 1977;

McPherson and Marion 1981, 1983; Gross 1982; Mitchell 1985a, 1985b).

Aspects of its life history and abundance have been reported for pond

populations in the north (Risley 1933, Cagle 1942, Wade and Gifford

1965, Ernst 1986), a large lake population in central Florida (Bancroft

et al. 1983), and a stream population in Oklahoma (Mahmoud 1969).

Although reproductive and intrapopulational morphological variation is

substantial, there is relatively little interpopulational morphological

variation (Reynolds and Seidel 1983). Little is known about geographic

variation in population density, or about population densities in different

habitat types within close proximity to one another.

During the course of a study of the ecology of the flattened musk

turtle, Sternotherus depressus, in the Warrior Basin of north-central

Alabama (Dodd et al. 1988). I had the opportunity to sample

Brimleyana No. 15:47-56, January 1989

47

48

C. Kenneth Dodd, Jr.

periodically a lake population of S. odoratus near one of the S.

depressus study sites. This paper provides data on the population

structure and biomass of stinkpots at that site, near the northwestern

boundary of the Warrior Basin.

STUDY AREA AND METHODS

The study was conducted on Brushy Creek Lake in the Bankhead

National Forest, Winston County, Alabama (T9S R7W S4N, Grayson

Quadrangle). Brushy Creek Lake was created when Brushy Creek was

impounded to serve recreational purposes. It has a surface area of 13.76

ha with a maximum depth of 3.7 m, although 9.36 ha (67.3%) is less

than 2.4 m. The study area (Fig. 1) comprised 0.9 ha on the north shore

of the lake, including 6.4% of the total lake surface area and

approximately 9.7% of the vegetated shallow-water habitat.

The primary submergent plants in Brushy Creek Lake were

pondweeds ( Potamogeton sp.) and bladderworts ( Utricularia sp.). Major

fish species included largemouth bass ( Micropterus salmoides), bluegill

(Lepomis macrochirus ), redear sunfish ( L . microlophus ), warmouth (L.

gulosus), crappie ( Pomoxis sp.), grass pickerel (Esox americanus), and

carp ( Cyprinus carpio). Other species of turtles seen or trapped were the

snapping turtle ( Chelydra serpentina ), eastern mud turtle ( Kinosternon

subrubrum), yellow-bellied slider ( Trachemys scripta ), and river cooter

( Pseudemys concinna). The major benthic components of the lake have

not been identified (G. Gaines, pers. comm.).

Trapping was conducted six times between 5 May and 5 September

1985, at approximately equal time intervals, using 2.54-cm mesh wire

funnel traps (Iverson 1979); sardines were used as bait. Cans were

partially opened or punctured and placed in the bottom of the trap. Ten

traps were set on each sampling date between 1700 and 2000 hours and

retrieved between 0800 and 1000 hours the next morning. The amount

of time traps were in the water was recorded to the nearest 0.5 hour.

Traps were set in shallow water with a heavy fringe of emergent and

submerged aquatic vegetation that provided abundant cover for stinkpots

and their prey.

The carapace length (CL) and plastron length (PL) were recorded

for each turtle to the nearest 0.1 mm using a dial caliper and standard

turtle measuring techniques. Mass was recorded to the nearest 0.5 g.

Turtles were considered adults if they measured greater than 65 mm CL

(McPherson and Marion 1981). Each turtle was assigned an identification

number (ID) by notching marginal scutes (Cagle 1939).

Pearson correlation coefficients were generated for CL versus PL,

and a linear regression line (y = a + bx) was fitted to the plotted points.

For body mass, the data were fitted to the general allometric equation y

Population Structure and Biomass of Stinkpot

49

SHORELINE

= FOOTPATH

SWAMPY OROUNO OR EMEROENT VEGETATION

^ BUSHES

SAND

INTERMITTENT STREAM

• WOODEN POST

■ METAL POST

LOQ

TRAP LOCATION

10 M

Fig. 1. Map of the study site, Brushy Creek Lake, Winston County, Alabama.

= axb, in the form log y = log a + b(log x), by the method of least

squares regression analysis. Other analyses used the Mest, X2 test of

independence, and 2-way analysis of variance (ANOVA) with adjust-

ments for unequal sample size. Statistical analyses were carried out

using the SAS program for microcomputers (SAS Institute Inc. 1985).

For all analyses, P < 0.05 was considered indicative of statistical

significance.

RESULTS AND DISCUSSION

Population Density. A total of 135 individual S. odoratus were

captured during 813 trap-hours. There were 26 recaptures for a total of

161 captures, or one turtle per 5.05 trap-hours. Unfortunately, 41

turtles, including five previously marked animals, drowned on 3 June

when water temperatures reached 32° C in the shallows. Such mortality

precluded a statistically rigorous estimate of the population size.

It is possible, however, to derive an estimate of the population

density of S. odoratus in Brushy Creek Lake if one assumes the

minimum number of turtles caught represents a minimum population

size within the study area and that immigration balances emigration. As

such, 135 stinkpots were caught in the study area, yielding an estimate

50

C. Kenneth Dodd, Jr.

of 148.5 turtles per hectare. Since Brushy Creek Lake has an area of

13.76 ha, there could be as many as 2,043 S. odoratus in the lake,

assuming an equal population density among areas. If adjustments are

made to confine the estimate to optimal habitat (67.3% of the lake

surface area based on depth profiles), a minimum population of 1,375

stinkpots in the lake would be indicated.

Published values of stinkpot density range from 8 to 700/ ha

(Mahmoud 1969, Iverson 1982, Congdon et al. 1986, Ernst 1986), and

Mitchell (pers. comm.) has found densities in two Virginia populations

at 188 and 194/ha. Given these figures, a density of 148 stinkpots per

hectare in Brushy Creek Lake seems reasonable.

Population Structure. Of the 135 stinkpots, 22 (16.3%) were juveniles

(< 65 mm CL); the smallest measured 39.3 mm CL. There was no

significant difference in CL between males and females (F = 1.30, P -

0.26), although males averaged slightly larger (M: x = 76.8 mm, range

66.6-90.5 mm, SD = 7.6; F: x = 75.0 mm, range 65.0-95.1 mm, SD =

5.5). The lack of differences in the CL of adult males and females was

not surprising since size dimorphism has not been reported for S.

odoratus except in the extreme southern portions of its range (Tinkle

1961, Bancroft et al. 1983).

Most turtles were in the 65- to 79.9-mm size classes, and none were

in the 50- to 54.9-mm size class (Fig. 2). Females outnumbered males in

all but the 85- to 89.9-mm size class. The relationship of carapace length

to plastron length was highly significant (Fig. 3) regardless of sex (Table

1).

The population structure of S. odoratus in Brushy Creek Lake was

nearly identical with populations in Pennsylvania (Ernst 1986) and

Florida (Bancroft et al. 1983). All three studies showed increasing

numbers of individuals in age classes up to intermediate CLs, followed

by a rapid decrease in the numbers of larger individuals. The main

differences are not in the structure of the populations per se, but in the

sizes of the animals. Pennsylvania stinkpots attain much larger CLs

than Florida turtles, and northern Alabama turtles are nearly intermediate

in this respect.

In some other studies (e.g., Wade and Gifford 1965), the population

structure is slightly different, showing what appears to be a more

gradual decline in numbers of larger turtles. The reasons for such

variation are unknown, although differences may be an artifact of

sampling technique. Unless data are available using comparable sampling

procedures, it may be premature to speculate on the underlying causes

of differences in population structure reported for distant populations.

Sex Ratio. Of the turtles larger than 65 mm CL, there were 30

males and 83 females, or one male per 2.8 females. This ratio is

significantly different from 1:1 ( X 2 = 12.68, P < 0.01). Proportionally

more females were caught in the May (6.25 females/ male) and August

PLASTRON (MM) t""1 T! NO. OF TURTLES

Population Structure and Biomass of Stinkpot

51

<50.0 50.0-54.9 55.0-59.9 60.0-64.9 65.0-69.9 70.0-74.9 75.0-799 80.0-849 85.0-89.9 >90.0

CARAPACE LENGTH (MM)

ig. 2. Population size structure of Sternotherus odoratus in Brushy Creek

ake, Winston County, Alabama.

CARAPACE (MM)

Fig. 3. Regression of carapace length versus plastron length for Brushy Creek

Lake Sternotherus odoratus. See Table 1 for values of r, a, and b. • = 1

observation; 0 = 2 observations; ★= 3 observations.

52

C. Kenneth Dodd, Jr.

samples (4.3 females/ male) as opposed to the other months (range 1.4

to 2.4 females/ male). Males were never caught in greater numbers than

females.

Several studies have reported sex ratios of 1:1 (Tinkle 1961,

Mahmoud 1969, Mitchell 1982, Ernst 1986) or favoring females (Risley

1933, Cagle 1942). Bancroft et al. (1983), with the largest sample size of

all studies to date, found that males outnumbered females (1.16:1) in a

Florida lake population. They attributed the skewed ratio to the tendency

of males to move greater distances than females, an explanation also

used by Dodd et al. (1988) to explain a similarly skewed ratio for S.

depressus in Sipsey Fork, Alabama. The tendency of males to move

more often and over greater distances than females may not explain sex

ratios in favor of females in Brushy Creek Lake, however.

It is possible that different sex ratios would result from more

extensive collections made at different times of the reproductive season

or from a sampling regime that placed traps over a larger portion of the

lake. In Alabama, K. R. Marion (pers. comm.) found that males were

caught more often in early spring and fall samples and that females

tended to be more common in late spring and early summer samples.

His overall results, however, still indicated a female-biased sex ratio. In

Virginia, J. C. Mitchell (pers. comm.) also found sex ratios in a stinkpot

population in an urban area that varied depending on season, although

in his study the overall sex ratio remained at 1:1.

Moreover, if there are differences in habitat selection between the

sexes, and if a sampling effort does not equally sample all potential

habitat, the resulting sex ratio may be biased and not indicative of the

overall sex ratio within the population. This may explain the female-

biased sex ratio of stinkpots in the present study. In any case, the

interpretation of studies reporting female-biased sex ratios in S. odoratus

is often difficult because of small sample sizes and differences in

trapping techniques between studies. Sex ratios based on relatively

small sample sizes should be interpreted with caution (Gibbons 1970b,

Bury 1979).

Mass. The average body mass for all stinkpots was 71.3 g (N =

108). Males averaged 78.9 g (range 46.0-130.0 g, SD = 22.6) and females

averaged 76.8 g (range 47.0-132.0 g, SD = 14.4). There was no significant

difference in body mass between adult males and females {t - -0.4819, P

- 0.63). Log(body mass) was significantly correlated with both log(cara-

pace length) and log(plastron length) (Fig. 4) regardless of sex. Power

function exponents ranged from 2.5 to approximately 2.9 (Table 1).

Only four studies have reported biomass data for S. odoratus in

lentic habitats. The estimates ranged from 1.2 to 41.7 kg/ ha, but mean

body mass varied considerably between studies (Wade and Gifford

1965; Iverson 1982; Congdon et al. 1986; J. C. Mitchell, pers. comm.).

Population Structure and Biomass of Stinkpot

53

A

Fig. 4. A. Regression of log-log transformed data of carapace length versus

body mass for Brushy Creek Lake Sternotherus odoratus. B. Regression of log-

log transformed data of plastron length versus body mass. See Table 1 for

values of r, a, and b. • = 1 observation; 0 = 2 observations; ★ = 3 observations.

54

C. Kenneth Dodd, Jr.

Table 1. Relationship between the dependent variables (Y) of plastron length

(PL) and body mass (W) and the independent variables (X) of carapace

length (CL) and plastron length for Brushy Lake Sternotherus

odoratus. For correlations involving body mass, the data were log-log

transformed. Units in g and mm. The statistical significance of

correlation coefficients is indicated (**, P< 0.01).

Assuming a density of 148.5 S. odoratus /ha in Brushy Creek Lake,

there was a minimum biomass of 10.6 kg/ ha.

Body mass may vary seasonally, especially among females (e.g.,

Branch 1984), and it may be among the most important variables in life

history studies (see Hedges 1985 and references therein). As such,

variation in body mass between populations of turtles may be of greater

importance than recognized to date. As Congdon et al. (1986) suggested,

factors such as habitat suitability, body size, and population age structure

may be more important in determining species-specific densities than

trophic position.

Mortality. On 3 June, 41 of 56 S. odoratus drowned in traps,

presumably because of high water temperature and low oxygen concen-

tration in the unshaded study site. These included 28 females, 4 males,

and 9 juveniles. There was no significant difference in the sex ratio of

drowned animals and the overall sex ratio of turtles that did not drown

(X2 = 3.35, P - 0.067) or between the carapace lengths of drowned (x =

75.2 mm, SD = 6.6) and non-drowned (x = 75.5 mm, SD = 5.9) adults (F

= 0.03; P - 0.87). Also, there was no relationship between the sex of the

turtles and their CL on their tendency to drown (F = 0.03; P = 0.85).

Although Ultsch et al. (1984) reported that S. odoratus could

survive submergence 5.2 days under anoxic conditions at 10° C, my

trap results suggest that stinkpots have considerably less tolerance to

Population Structure and Biomass of Stinkpot

55

low oxygen or anoxic conditions at high temperatures in the field.

Caution should be exercised in the placement of traps to prevent

drowning.

ACKNOWLEDGMENTS. — I thank Kevin M. Enge and James N.

Stuart for assistance with field data collection. Jose Gallo and Howard

I. Kochman provided valuable statistical advice. I thank Fred Cox, Ken

Marion, and Joseph Mitchell for their comments on an earlier draft of

the manuscript. Glen Gaines provided information on Brushy Creek

Lake. This study was conducted under contract No. 14-16-0009-84-1896

between the Office of Surface Mining, U.S. Department of the Interior,

and the U.S. Fish and Wildlife Service. Collecting was authorized under

scientific collecting permit No. 172 from the Alabama Department of

Natural Resources, and by the U.S. Forest Service, Bankhead National

Forest.

LITERATURE CITED

Bancroft, G. Thomas, J. Steve Godley, Dena T. Gross, N. Nan Rojas, Dareth

A. Sutphen, and Roy W. McDiarmid. 1983. The herpetofauna of Lake

Conway: Species accounts. Misc. Pap. A-83-5. Army Engineer Waterways

Experiment Station, Vicksburg, Miss.

Branch, W. R. 1984. Preliminary observations on the ecology of the angulate

tortoise ( Chersina angulata) in the eastern Cape Province, South Africa.

Amphibia-Reptilia 5:43-55.

Bury, R. Bruce. 1979. Population ecology of freshwater turtles. Pages 571-602

in Turtles, Perspectives and Research, M. Harless and H. Morlock, editors.

John Wiley and Sons, New York.

Cagle, Fred R. 1939. A system of marking turtles for future identification.

Copeia 1939(3): 170-172.

Cagle, Fred R. 1942. Turtle populations in southern Illinois. Copeia

1942(3): 155-162.

Carr, Archie. 1952. Handbook of Turtles. The Turtles of the United States,

Canada, and Baja California. Cornell Univ. Press, Ithaca, N.Y.

Congdon, Justin D., Judith L. Greene, and J. Whitfield Gibbons. 1986.

Biomass of freshwater turtles: a geographic comparison. Amer. Midi. Nat.

1 15(1): 165-173.

Dodd, C. Kenneth, Jr., Kevin M. Enge, and James N. Stuart. 1988. Aspects of

the biology of the flattened musk turtle, Sternotherus depressus, in northern

Alabama. Bull. Fla. St. Mus., Biol. Sci. 34(1): 1-64.

Ernst, Carl H. 1986. Ecology of the turtle, Sternotherus odoratus, in

southeastern Pennsylvania. J. Herpetol. 20(3):34 1-352.

Ernst, Carl H., and Roger W. Barbour. 1972. Turtles of the United States.

Univ. Presses of Kentucky, Lexington.

Gibbons, J. Whitfield. 1970a. Reproductive characteristics of a Florida

population of musk turtles ( Sternotherus odoratus). Herpetologica 26(2):268-270.

Gibbons, J. Whitfield. 1970b. Sex ratios in turtles. Res. Popul. Ecol.

12:252-254.

56

C. Kenneth Dodd, Jr.

Gross, Dena T. 1982. Reproductive biology of the stinkpot, Sternotherus

odoratus, in a central Florida lake system. Unpubl. Master’s thesis, Univ.

South Florida, Tampa.

Hedges, S. Blair. 1985. The influence of size and phylogeny on life history

variation in reptiles: a response to Stearns. Amer. Nat. 126(2):258-260.

Iverson, John B. 1977. Reproduction in freshwater and terrestrial turtles of

north Florida. Herpetologica 33(2):205-212.

Iverson, John B. 1979. Another inexpensive turtle trap. Herpetol. Rev.

10(2):55.

Iverson, John B. 1982. Biomass in turtle populations: a neglected subject.

Oecologia 55:69-76.

Mahmoud, I. Y. 1969. Comparative ecology of the kinosternid turtles of

Oklahoma. Southwest. Nat. 14( 1):3 1 -66.

McPherson, Roger J., and Ken R. Marion. 1981. The reproductive biology of

female Sternotherus odoratus in an Alabama population. J. Herpetol.

15(4):389-396.

McPherson, Roger J., and Ken R. Marion. 1983. Reproductive variation

between two populations of Sternotherus odoratus in the same geographic

area. J. Herpetol. 17(2): 18 1-184.

Mitchell, Joseph C. 1982. Population ecology and demography of the

freshwater turtles Chrysemys picta and Sternotherus odoratus. Unpubl.

Ph.D. dissert., Univ. Tennessee, Knoxville.

Mitchell, Joseph C. 1985a. Variation in the male reproductive cycle in a

population of stinkpot turtles, Sternotherus odoratus, from Virginia.

Copeia 1985( l):50-56.

Mitchell, Joseph C. 1985b. Female reproductive cycle and life history attributes

in a Virginia population of stinkpot turtles, Sternotherus odoratus. Copeia

1 985(4):94 1 -949.

Mount, Robert H. 1975. The Reptiles & Amphibians of Alabama. Agric.

Exper. Sta., Auburn Univ., Auburn.

Reynolds, Samuel L., and Michael E. Seidel. 1983. Morphological homogeneity

in the turtle Sternotherus odoratus (Kinosternidae) throughout its range. J.

Herpetol. 1 7(2): 1 13-120.

Risley, Paul L. 1933. Observations on the natural history of the common musk

turtle, Sternotherus odoratus (Latreille). Pap. Mich. Acad. Sci. Letters.

17:685-71 1.

SAS Institute Inc. 1985. SAS Introductory Guide for Personal Computers,

Version 6 Edition. SAS Institute Inc., Cary, N.C.

Tinkle, Donald W. 1961. Geographic variation in reproduction, size, sex ratio

and maturity of Sternotherus odoratus (Testudinata: Chelydridae). Ecology

42(l):68-76.

Ultsch, Gordon R., Christine V. Herbert, and Donald C. Jackson. 1984. The

comparative physiology of diving in North American freshwater turtles. I.

Submergence tolerance, gas exchange, and acid-base balance. Physiol.

Zool. 57(6):620-63 1.

Wade, Susan E., and Cameron E. Gifford. 1965. A preliminary study of the

turtle population of a northern Indiana lake. Proc. Indiana Acad. Sci.

74:371-374.

Accepted 3 June 1987

Distribution, Biology, and Conservation Status

of the Carolina Madtom, Noturus furiosus ,

an Endemic North Carolina Catfish

Brooks M. Burr, Bernard R. Kuhajda, Walter W. Dimmick,

and James M. Grady

Department of Zoology, Southern Illinois University,

Carbondale, Illinois 62901

ABSTRACT. — Noturus furiosus is endemic to the Tar and Neuse

river drainages, North Carolina, where it occurs in medium- to large-

size streams over sand, gravel, and detritus substrates. Because of its

endemicity and relatively limited distribution, N. furiosus became a

candidate for pre-listing studies by the Office of Endangered Species,

U.S. Fish and Wildlife Service. Recent survey work throughout the

Tar and Neuse rivers indicates that N. furiosus is reproducing and

undergoing recruitment at several localities. However, numerous

proposed projects and several recently constructed reservoirs pose

threats to the continued successful existence of this madtom.

Aspects of the general biology of N. furiosus were analyzed from

326 preserved specimens from both the Tar and Neuse river drainages.

Individuals live at least 4 years. The largest specimens seen were 101

mm SL (male) and 98 mm SL (female). Females mature at 2 to 3 years

and a mean SL of 75 mm. Mature oocytes, produced seasonally,

ranged from 79 to 298 (x = 126.3; N = 17) per female. Five nests, each

containing a clutch of embryos or larvae and guarded by a male, were

found in cans and bottles in pools or runs. All males guarding broods

were 3 to 4 years old and ranged in SL from 63 to 101 mm (x = 89.8).

Nests were observed in May at water temperatures of 20-25° C. Clutch

sizes ranged from 139 to 171+; embryos were spherical and light

yellow with yolk diameters averaging 3.2 mm. At about 1 day post-

hatching, larvae ranged from 9.1 to 10.0 mm TL; larval features were

similar to those described for other ictalurids. Stomachs of adults and

juveniles contained a variety of benthic organisms, but dipteran,

trichopteran, ephemeropteran, coleopteran, and odonate larvae or

nymphs composed more than 95% of the total food organisms

consumed.

The Carolina madtom, Noturus furiosus Jordan and Meek, is a

moderate-sized, boldly patterned catfish (Fig. 1) that is endemic to the

Tar and Neuse river drainages, North Carolina. It has remained poorly

known since its original description. Jordan and Meek (in Jordan 1889)

noted that “numerous specimens were taken” of N. furiosus (actually

fewer than 20), but did not comment further on its abundance. Bailey et

al. (1977) assigned N. furiosus a conservation status category of special

Brimleyana No. 15:57-86, January 1989

57

58

Burr, Kuhajda, Dimmick, and Grady

concern but made no further comment. Cooper and Braswell (1982)

stated: “Based on the very small numbers of specimens taken in recent

years, despite intensive sampling at many localities in both rivers, the

species seems to have experienced a serious decline.” They added, “Its

endemicity and apparent rarity make it vulnerable to extinction.”

Recently, Braswell and Cooper found the species to be relatively common

in October at two sites on the Tar River (Cooper and Ashton 1985).

Noturus furiosus was originally allied with N. miurus and N.

eleutherus (Jordan 1889). It is currently allocated to the subgenus

Rabida (Taylor 1969), which presently contains 14 other species (Taylor

1969, Douglas 1972, Etnier and Jenkins 1980). Taylor (1969) considered

N. furiosus to be a member of the furiosus species group including N.

placidus, N. stigmosus, and N. munitus. In an analysis of chromosomal

evolution of the genus Noturus , LeGrande (1981) did not include N.

furiosus. There has been some question regarding the taxonomic status

of N. furiosus because even Taylor (pers. comm.) has suggested that it

might be a geographic subspecific population of N. stigmosus. An

analysis of allozymes of all extant members of Noturus by James M.

Grady demonstrates that the species has several fixed alleles and can be

distinguished electrophoretically from other members of the furiosus

group.

The historical range of N. furiosus included varied habitats in two

physiographic provinces comprising all major tributary systems of the

Tar and Neuse rivers: Piedmont (Tar River and Neuse River [including

Eno River]) and Coastal Plain (Tar River [including Fishing Creek and

other minor streams] and Neuse River [including Little and Trent rivers,

Contentnea Creek, and other minor streams]). However, since 1963, N.

furiosus has been take at fewer than 12 localities. Use of ichthyocides,

electrofishing gear, and trapping in madtom habitat has revealed few

specimens of N. furiosus , although N. insignis and N. gyrinus are

encountered frequently.

Presented here is a report on the distribution, biology, and

conservation status of N. furiosus based on museum specimens and field

work through the summer of 1985. This paper is extracted from a report

to the Office of Endangered Species, U.S. Fish and Wildlife Service.

PROCEDURES

Collecting Methods. The field work accomplished from 1982 through

1984 was performed in most cases by two people with small-mesh, 10-

foot minnow-seines and dip nets. Both day and night collecting were

attempted at two sites. Locations where N. furiosus formerly had been

collected in relatively high numbers (5 to 10 individuals) were most

frequently visited. During May 1985, intensive collecting at nearly all

Carolina Madtom

59

Fig. 1. Noturus furiosus, 67 mm SL, Tar River, 5.5 airmiles NE Franklinton,

Franklin County, North Carolina, 17 October 1984 (NCSM 1 1089). Drawing by

Renaldo Kuhler.

previous sites of occurrence was performed using seines, 25-cm aquarium

nets, and snorkeling gear.

Field Surveys. A total of 66 visits to 16 sites within the Neuse River

drainage and 15 sites within the Tar River drainage were made with the

specific purpose of trying to locate extant populations of N. furiosus.

Little River, near Goldsboro, Wayne County, was visited almost monthly

during fall and spring, 1983 and 1984, to obtain baseline data on habitat

and life history. Unfortunately, extremely high water during several

months prevented field work.

Other information in this report has been obtained from museum

specimens in a number of research collections (see MATERIALS).

Geochronographic Figures. These figures, modified after Cashner

and Jenkins (1982), are maps showing locations, years, numbers of

collections, and results of collections in the Neuse and Tar rivers.

Locality data for all records are given under MATERIALS.

Reproductive Parameters. Specimens were sexed by examination of

gonads. GSI refers to the gonadosomatic index, calculated by total

weight of both ovaries or testes divided by eviscerated (= adjusted) body

weight X 1000. Weights of gonads and bodies were determined after

blotting, to the nearest 0.01 g, with a Mettler analytic balance.

Numbers of ova were determined by direct count from both ovaries.

Ova diameters were measured with dial calipers to the nearest 0.1 mm.

Ten randomly selected ova from each female were measured.

Nesting. Collecting sites were searched for potential nest sites (i.e.,

cans, bottles, mussel shells, boards, flat rocks, logs). Potential nest sites

were surrounded by a net, removed from the stream, and examined for

guardian adults, embryos, and larvae. If a nest was found, the parent(s)

60

Burr, Kuhajda, Dimmick, and Grady

and young were preserved for subsequent examination. In most cases,

mated pairs were returned to their nest sites if they were not guarding

embryos or larvae.

Aging. Age of some specimens in breeding condition was determined

by counting the number of annuli on cross sections of pectoral spines as

outlined by Clugston and Cooper (1960) and modified by Mayden and

Burr (1981). Many specimens were assigned to age classes based on

length-frequency analysis, although neither method of aging provided

satisfactory results. Lengths of juveniles and adults are expressed in mm

standard length (SL); lengths of larvae in mm total length (TL).

Food. Diet was determined from examination of the contents of

the stomach.

Substrate. Size ranges in centimeters of certain substrate types

mentioned herein are: sand, less than 0.3; pea or small gravel, 0. 3-3.0;

medium gravel, 3-5; large gravel, 5-8; small rubble, 8-15; medium

rubble, 15-22; large rubble, 23-30; and boulder, greater than 30.

MATERIALS

A virtually complete record of repository is given. Institutional

acronyms are: ANSP, Academy of Natural Sciences of Philadelphia;

DU, Duke University; CAS-SU, Stanford University (now at California

Academy of Sciences); NCSM, North Carolina State Museum of Natural

Sciences; SIUC, Southern Illinois Univeristy at Carbondale; UF,

University of Florida, Florida State Museum; UMMZ, University of

Michigan Museum of Zoology; UNCC, University of North Carolina at

Charlotte; and USNM, National Museum of Natural History. Numbers

of specimens (in parentheses) follow the catalog number or the acronym.

A total of 326 specimens (70 collections) were examined. Within

each drainage, collections are listed alphabetically by county. Collections

from the same site are listed in chronological order. An asterisk

preceding a locality denotes collections made by personnel of the North

Carolina Wildlife Resources Commission (NCWRC). To our knowledge,

these collections were not deposited in a recognized museum or university

collection, but are reported in Bayless and Smith (1962) and Smith and

Bayless (1964).

Neuse River System. CRAVEN COUNTY: NCSM 9939(1) Neuse

River near Streets Ferry, 8.7 airmi. NW New Bern, 28 January 1981.

DURHAM COUNTY: NCSM 1930(1) Eno River at SR 1004 bridge, 3

August 1961. GREENE COUNTY: *NCWRC(22) Contentnea Creek, 3

mi. S Scuffleton at Edwards Bridge, 6 September 1960. JOHNSTON

COUNTY: DU uncat. (3) Mill Creek, 1 mi. N Cox Mill, SR 1200 bridge,

8 June 1961; NCSM 13838(1) Little River at SR 1002 bridge, 1 mi. N

Carolina Madtom

61

Princeton, 19 May 1985; NCSM 8780(2) Little River at SR 1001 bridge,

4.25 airmi. SSW Kenly, 25-29 March 1979; NCSM 3420(5) Swift Creek,

3 mi. SW Smithfield, 18 July 1961; NCSM 13839(4) same site as

preceding, 19 May 1985; NCSM 1794(7) Little River, 1 mi. W Raines

Crossroads, 20 June 1961; DU uncat. (18) same site as preceding, 20 July

1961; NCSM 632(1) Middle Creek, at NC 210 bridge, 3 airmi. S

Smithfield, 20 July 1961; UNCC uncat. (?) Little River above SR 1001

bridge, 10.4 mi. E Smithfield, 16 May 1982. JONES COUNTY: NCSM

8245(5) Trent River at SR 1300 bridge, 4.75 airmi. NW Trenton, 25-26

September 1978; NCSM 13840(1) same site as preceding, 22 May 1985;

NCSM 8223(3) Trent River at SR 1129 bridge, 6 airmi. W Trenton, 25

September 1978; NCSM 8204(1) Trent River below NC 58 bridge, 4.75

airmi. NW center of Trenton, 25 September 1978; *NCWRC(10) Trent

River at NC 58 bridge, 3 mi. E Phillips Crossroads, 22 July 1960.

LENOIR COUNTY: SIUC 11730(22) Neuse River at NC 903 bridge,

5.5 mi. SSW LaGrange, 20 May 1985; NCSM 13841(1) Neuse River at

Kinston at Business NC 58-US 70 bridge, 22 May 1985; DU uncat. (5)

Neuse River, 5 mi. N Liddell, ca. 17 mi. W Kinston, 11 August 1960.

PITT COUNTY: NCSM 758(1) Little Contentnea Creek, at NC 102

bridge, 9 September 1960. WAKE COUNTY: USNM 48475(1) Crabtree

Creek, near Raleigh, July 1897; UMMZ 165885(1) Neuse River, near

Raleigh, 22 August 1897; NCSM 243(1), probably from Neuse River

near Raleigh, no date; USNM 67937(1) Neuse River, Raleigh, 27

August 1888; UMMZ 165884(1) Neuse River, 18 August 1902; CAS-SU

1380(2), USNM 39932(1), USNM 164109(2) Neuse River, at Millburnie,

near Raleigh, summer 1888. WAYNE COUNTY: NCSM 485(1), NCSM

486(1) Beaverdam Creek, 0.5 mi. upstream from jet. with Neuse River, 6

June 1961; NCSM 1242(1), NCSM 2715(2) Neuse River below Quaker

Neck Dam at Goldsboro, 7 June 1961; UF 31453(2) Neuse River, 2 mi.

downstream from H. F. Lee Plant, 9 August 1977; NCSM 2209(3) Little

River, 1 mi. W Goldsboro at NC 581 bridge, 13 June 1961; UF 23768(1)

same site as preceding, 25 April 1977; SIUC 5541(7) same site as

preceding, 22 May 1982; SIUC 8693(16) same site as preceding, 17

September 1983; SIUC 8960(3) same site as preceding, 14 October 1983;

SIUC 9754(1) same site as preceding, 30 April 1984; NCSM 13836(2),

NCSM 13837(2), SIUC 1 1685(1), SIUC 1 1777(1), same site as preceding,

19 May 1985; SIUC 11683(2) same site as preceding, 20 May 1985.

USNM 40572(1) Little River, near Goldsboro, 1888. WILSON COUNTY:

DU 851(9) Contentnea Creek, 3 mi. W Stantonsburg, 8 September

1960; *NCWRC(39) Contentnea Creek at NC 42 bridge, 4 July 1961.

Tar River System. EDGECOMBE COUNTY: ANSP 71335(1),

CAS-SU 3435(2), UMMZ (Indiana Univ. 7246)(1), UMMZ 167076(1),

USNM 20926(6) Tar River, near Tarboro, ca. 1877; NCSM 11087(1)

62

Burr, Kuhajda, Dimmick, and Grady

Tar River, 0.9 airmi. SE Tarboro, 3 October 1984; SIUC 11760(6) Tar

River at NC 44 bridge, N edge of Tarboro, 16 May 1985; NCSM

13832(2), SIUC 1 1776(1), SIUC 11679(8) same site as preceding, 17

May 1985; SIUC 1 1775(1) same site as preceding, 22 May 1985; NCSM

11090(11) Tar River, at NC 42 crossing, 4.9 airmi. E Pinetops, 30

October 1984; USNM 40398(1) Tar River, below Rocky Mount, 1888;

NCSM 13833(2), SIUC 1 1778(1) Tar River at SR 1252 bridge, 7.5 mi. E

Rocky Mount, 17 May 1985; USNM 191110(44), USNM 191071(15),

UMMZ 187094(8) Fishing Creek, below bridge at SR 1500, ca. 4.5 mi.

SW Lawrence, 19 September 1959; *NCWRC(37) same site as preceding,

12 August 1963; USNM 191099(8) Swift Creek, just above NC 97, W

Leggett, 19 September 1959; *NCWRC(16) Swift Creek at SR 1253

bridge, 1 mi. SSW Leggett, 9 July 1963; SIUC 4194(1) same site as

preceding, 21 May 1982; NCSM 13830(4), NCSM 13831(7) same site as

preceding, 16 May 1985; *NCWRC(9) Town Creek, 2 mi. upstream

from mouth, 10 June 1963. EDGECOMBE-H ALIFAX COUNTY:

*NCWRC(17) Fishing Creek at SR 1418 bridge near Enfield, 13 June

1963. EDGECOMBE-NASH COUNTY: USNM 191057(8), UMMZ

187097(4) Tar River, at railroad line, Rocky Mount, 19 September

1959; SIUC 1 1684(1) same site as preceding, 17 May 1985. FRANKLIN

COUNTY: *NCWRC(7) Tar River at SR 1003, W Louisburg, 26 June

1963; NCSM 11089(10) Tar River, upstream from bridge at SR 1003,

5.5 airmi. NE Franklinton, 17 October 1984; SIUC 11749(15) same site

as preceding, 15 May 1985; *NCWRC(17) Tar River at SR 1611 near

Bunn, 26 June 1963; *NCWRC(?) Sandy Creek at NC 58 bridge, 11

June 1963; NCSM 13834(1) Tar River at Louisburg, 18 May 1985.

FRANKLIN-VANCE COUNTY: NCSM 3077(1) Tar River, 4 mi. N

Franklinton at US 1 bridge, 7 July 1966; NCSM 13828(2) same site as

preceding, 15 May 1985; NCSM 13829(2) Tar River at SR 1203 bridge,

4.75 mi. NNW Franklinton, 15 May 1985. HALIFAX COUNTY:

*NCWRC(18) Little Fishing Creek at SR 1343 bridge near White Oak,

22 July 1963; *NCWRC(1) Beech Swamp at SR 1100 bridge, 13 June

1963. NASH COUNTY: *NCWRC(29) Swift Creek at SR 1003 bridge

near Red Oak, 10 July 1963; NCSM 13835(1) Tar River at NC 581

bridge, 0.5 mi. N Floods Chapel, 18 May 1985. NASH — HALIFAX

COUNTY: *NCWRC(18) Fishing Creek at SR 1506 bridge near Aventon,

12 June 1963. VANCE COUNTY: *NCWRC(20) Tar River at SR 1 101

bridge near Franklinton, 15 August 1963.

RESULTS

Habitat

All records of N. furiosus are from free-flowing streams. Permanent

occupation of lentic habitat is highly unlikely, and sampling in pools

Carolina Madtom

63

generally does not yield specimens. Gradients are moderate and stream

temperatures in summer are warm (diurnal maxima greater than 20° C).

Submersed macrophytes are generally absent; the Carolina madtom

does not appear to be strongly associated with emergent vegetation,

although small patches of vegetation may occasionally be present in

prime habitat. Water is generally clear to coffee-colored.

A large majority of records are from medium to large streams,

i.e., the Neuse and Tar rivers proper and the lower reaches of their

major tributaries. Most records from small streams are within a few

kilometers of the mouth and may represent forays of individuals, or

populations largely reliant upon recruitment from main rivers.

Stream sections inhabited have riffles, runs, and pools with N.

furiosus found during warm months in or near swift current; depth

usually 0.3 to 1.0 m. Young and juveniles tend to occupy shallower

riffles and runs in slower current than adults, although overlap is

typical. Habitat during cool months is unknown, but there is no reason

to suspect it would be strikingly different from that occupied during the

warm months. Common substrates in well-populated streams are leaf

litter, sand, gravel, and small rubble. The Carolina madtom occupies all

these substrates, but is most frequently taken over sand mixed with pea-

or medium-sized gravel and in leaf litter. In the lower Tar and Neuse

rivers, the species is frequently taken from debris piles in sandy areas.

The habitat of adult madtoms during the nesting season (May-July)

is in areas of moderate to slow flow where there is an abundance of cover

for nesting sites (e.g., beer and soda cans, bottles, mussel shells, flat

rocks, and stick piles). Because guardian males usually have empty

stomachs (Table 2), it is assumed they rarely leave a nest site to forage.

This may, in part, explain why collections made during the breeding

season often miss adult madtoms unless nesting sites are found or

ichthyocides are used.

Taylor (1969) noted that specimens taken from near the Fall Line

were collected in swift water about 1 m in depth over a gravel-rubble

bottom, whereas those taken on the Coastal Plain were in very shallow

water with little or no current over sand and small gravel. Our

observations of habitat agree closely with those of Taylor.

Distribution, Abundance, and Population Status

Noturus furiosus is known from the Neuse and Tar river drainages

of North Carolina (Fig. 2), occurring both above and below the Fall

Line. Extant populations of the fish are presently known with certainty

from 17 localities (Fig. 3-4). Generally, the species is rare or uncommon,

but this may relate to inadequate sampling of appropriate habitat and

lack of collections made after dusk. It has been taken with regularity in

Table 1. Characteristics of female Noturus furiosus in reproductive condition from the Tar and Neuse river drainages,

North Carolina.

64

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Carolina Madtom

67

Fig. 2. Distribution of Noturus furiosus, showing all known extant and

extirpated populations. Some dots cover more than one record locality. Solid

dots represent localities with extant voucher material; open circles represent

localities reported by Bayless and Smith (1962) and Smith and Bayless (1964)

for which voucher material is presumably not available. 1. Beech Swamp Creek,

2. Fishing Creek, 3. Swift Creek, 4. Swamp Creek, 5. Little Contentnea Creek,

6. Contentnea Creek, 7. Trent River, 8. Little River, 9. Beaverdam Creek, 10.

Mill Creek, 11. Middle Creek, 12. Swift Creek, 13. Crabtree Creek, 14. Eno

River.

only a few streams in recent years, the Little River system (Neuse

drainage), the lower Neuse River proper. Swift Creek (Neuse drainage),

the Tar River proper, and Swift Creek (Tar drainage).

Reviewed here are the status of madtom populations and their

distributions and habitats within tributary systems. Order of presentation

is from upstream to downstream, Neuse drainage first.

Neuse Drainage

Eno River system. — Only a single specimen of N. furiosus, captured

in 1961 (using rotenone), is known from the Eno River. Our collection

on the Eno River was made at night in May 1982 but did not reveal

additional specimens, although a typical variety of Piedmont fishes was

represented in our sample. Appropriate madtom habitat appeared scarce.

68

Burr, Kuhajda, Dimmick, and Grady

NEUSE RIVER

• 1985, 8 +

• 1984, 10

• 1983, 19

• 1982, 7

•1977, 1

o 1 985; 1 • 1 96 1 , 3

01983; 2 *1961, 39 *1888, 1 *1960, 22

Fig. 3. Geochronography of Noturus furiosus in the Neuse River drainage,

North Carolina. Map shows virtually all known records of capture. Solid dots

indicate capture sites, but not necessarily extant subpopulations. Offriver data

are: year of collection(s) preceded by (1) solid dot if N. furiosus taken, (2) open

circle if not taken, (3) asterisk if voucher material not available; following the

year are (4) number of specimens taken (this number is separated by a comma),

(5) the number of collections not yielding specimens (this number is separated

by a semicolon), (6) a “?” indicating number of specimens unknown, and (7) a

“+” sign indicating many individuals were returned to the stream. Numbers refer

to collection of juvenile and adult specimens only. A. Eno River, B. Crabtree

Creek, C. Swift Creek, D. Middle Creek, E. Mill Creek, F. Beaverdam Creek,

G. Little River, H. Contentnea Creek, I. Little Contentnea Creek, J. Trent

River.

with much of the stream bottom consisting of bedrock and large rubble.

Capture of N. furiosus above the Fall Line is rare, and the Eno River

record is at the extreme northwestern edge of the species’ range.

Neuse River near Raleigh. — This is the type locality of N. furiosus

where approximately seven specimens were taken in 1888, 1897, and

1902. Four daytime visits to the Neuse River near Raleigh in 1982, 1983,

and 1985 revealed large expanses of suitable habitat, but no madtoms

were collected. The changes in the river near this site include the

construction of a major reservoir (Falls Lake) upstream and pollution

Carolina Madtom

69

TAR RIVER

01985; 1 *1985; 1

*1963, 18 *1963, 29 *1963, 17

• 1959, 8

• 1985, 1

r

o

t — i — r

km

"I

50

Fig. 4. Geochronography of Noturus furiosus in the Tar River drainage, North

Carolina. Map shows virtually all known records of capture. Data format

explained in Figure 3. A. Beech Swamp Creek, B. Fishing Creek, C. Swift

Creek, D. Swamp Creek.

from the city of Raleigh. A large fish kill in July 1980 did not reveal any

Carolina madtoms (Alvin L. Braswell, pers. comm.). The temperature in

the Neuse River below Falls Lake dam was 18°C in mid-May 1985,

about 6 degrees cooler than surrounding streams, and perhaps too low

for madtom reproduction. Dead mussels abounded, but no other

invertebrates were seen. Few fishes were collected, and all but an eel,

Anguilla rostrata , were introduced sport fishes. It is unlikely that the

Carolina madtom occurs in the reservoir or near the polluted waters of

the city anymore.

Swift, Middle, Mill, and Beaverdam creeks. — A total of 1 1 specimens

were taken in 1961 from these four creeks. Upon re-collecting all these

creeks during the day in October 1983 and May 1985, we obtained five

specimens, all from Swift Creek. Suitable madtom habitat is present,

although limited, and the streams are relatively small, perhaps too small

to support large, viable populations of N. furiosus. Potential nest sites

in Swift Creek were occupied only by juveniles, although some nesting

probably occurs in the stream. Records from these streams may

70

Burr, Kuhajda, Dimmick, and Grady

represent forays of a few individuals from the nearby Neuse River

proper. We know of no substantial, long-term ecological perturbations

that have occurred in these streams since 1961.

Little River system. — Without question this stream harbors the

single largest extant population of the Carolina madtom known from

within the entire Neuse drainage. As many as 20 individuals have been

collected at any one time, and one of the largest series of males and

females in breeding condition was taken from the Little River in 1961.

Our several visits to Little River just west of Goldsboro from 1982

through 1985 revealed abundant madtom habitat, both for nonbreeding

and breeding individuals, and numerous mated pairs were found in cans

and bottles and returned to their nest sites. Substantial populations of

N. insignis and N. gyrinus are also known from the Little River. Except

for a small dam and impoundment, the Little River appears to be

relatively undisturbed.

Neuse River, Lenoir County. — In 1960, five specimens of N. furiosus

were collected in the Neuse River proper near the Lenoir County line.

We revisited this site in May 1985 and found N. furiosus to be the most

common benthic fish species. Water was low (<1 m) about 300 m

downstream of the bridge, and from two to eight specimens were

obtained from nearly every stick/ detritus pile found in moderate current

over sand. The fish was so common that many individuals were returned

(22 were preserved) to the river. No nesting sites were found here,

although a few scattered mussel shells were examined. A much smaller

population of N. furiosus is present on the lower Neuse River near

Kinston.

Contentnea Creek system. — Sampling activity in this stream has

been limited in recent years, but four collections using rotenone in 1960

and 1961 yielded a total of 71 N. furiosus. We resampled two of the

Contentnea Creek sites in May 1985, but found no N. furiosus. During

our visits, suitable madtom habitat was found to be limited. In fact, one

site was essentially a swamp rather than a flowing stream. We assume

the fish still occurs in the Contentnea system, but probably in limited

numbers owing to lack of adequate habitat.

Trent River system. — In 1960 and 1978, ten and nine specimens,

respectively, of N. furiosus were taken from three sites on the Trent

River. These were the first records of the species that far downstream in

the Neuse. Collection methods included ichthyocides, which are very

effective in securing madtoms. We revisited all three sites on the Trent

River in September 1983 and May 1985. Only one N. furiosus was

found even though adequate habitat was available. N. insignis was

common. About 75 potential nest sites were examined here, but they

were either empty or were being used by other fishes (e.g., N. insignis ,

A. rostrata) and the Neuse River waterdog, Necturus lewisi.

Carolina Madtom

71

Extreme lower Neuse River, Craven County. — Probably the most

surprising record of the Carolina madtom from the Neuse is an individual

that was trawled from the main river near New Bern in 2 to 3 m of

water over sand in 1981. Extremely heavy rain in December (the highest

in 15 years) preceded the capture of this fish in January; it had

apparently been swept downstream with the heavy flooding that followed

the rain. The collection should probably be considered a stray from

typical areas of occurrence upstream.

Tar Drainage

Tar River proper. — Both historically and recently the Tar River

proper has harbored the most substantial populations of N. furiosus

known anywhere. Six collections on the mainstream have yielded 45

specimens since 1877, and three large populations are known to be

extant. Searches by us, some U.S. Fish and Wildlife Service personnel,

and North Carolina State Museum employees in October 1984 and May

1985 revealed four new sites for the Carolina madtom in the Tar River.

In addition, several old sites were re-collected (e.g., Tarboro and Rocky

Mount). All sites had an abundance of madtom habitat, and numerous

additional specimens were observed but not preserved. Numerous mussel

shells, cans, bottles, jars, and other litter in the Tar River provide an

abundant variety of nesting sites and undoubtedly contribute to successful

reproduction in most summers. It should be noted that the recent

successful collections in the Tar River were made under ideal conditions

of extremely low, clear water and during peak recruitment or spawning

periods. The Tar River remains one of the finest large streams in North

Carolina and will probably continue to be the stronghold for the

species.

Swift and Fishing Creek systems.— Both of these streams offer

abundant habitat for N. furiosus and are physically similar to the Little

River near Goldsboro. The largest series ever taken of this species (67

individuals) was collected in 1959 from Fishing Creek during a combined

day and night sampling period. Re-collection of these sites between 1982

and 1985 yielded 14 specimens (all from Swift Creek), but sampling was

impossible in Fishing Creek because of extremely high water. Re-

collection of Fishing Creek under optimum conditions is especially

warranted because this stream may harbor one of the largest extant

populations of N. furiosus. Several large collections of N. furiosus were

made by NCWRC personnel in 1963 in Swift and Fishing creeks (Fig. 4)

after application of rotenone. To our knowledge, no voucher specimen

is extant for any of these collections, but the large number of specimens

taken (Fig. 4) is evidence that these two streams support, or did

support, large populations of N. furiosus.

72

Burr, Kuhajda, Dimmick, and Grady

Life History

Age, Maximum Size, and Weight-Length Regression. Specimens

(32-99 mm SL; N = 10) aged from pectoral spines were 1 to 4 years old.

Accurate readings of other pectoral-spine cross sections could not be

made. The largest specimen aged was 99 mm SL and 4 years old. The

largest male examined was 101 mm; the largest female 98 mm. Few

individuals reach 85 mm. The largest specimen recorded by Taylor

(1969) was 100 mm SL.

Length-frequency histograms of 248 specimens were difficult to

interpret, although several age classes appear to be represented (Fig. 5).

Populations seem skewed towards younger age classes as would be

expected in a healthy population experiencing normal recruitment. The

relatively large number of individuals collected in May under 40 mm SL

indicates that growth is slow in winter and early spring. In September

(N = 92), age 0 fish ranged in SL from 17 to 47 mm, age 1 from 49 to 67

mm, age 2 from 71 to 76 mm, and ages 3 to 4 from 79 to 86 mm. There

was considerable overlap in length of older age classes. Lack of adequate

sample sizes in most months precludes more meaningful comparisons of

age classes and sexual differences in growth.

Regression of body weight on length of specimens was similar for

both sexes. The relationship between adjusted body weight in grams

(W) and SL for males was Log W = -5.098 + 3.198 Log SL (r = 0.99; N =

56) and for females was Log W = -4.631 + 2.918 Log SL (r = 0.99; N =

77).

Age and Size at Maturity. Females of N. furiosus reached

reproductive maturity in a minimum of 2 years, although a vast ma-

jority of gravid females were 3-year-olds (x = 3.1 years; Table 1).

Other species of Noturus about the same size as N. furiosus reached

reproductive maturity in 1 to 2 years (e.g., N. exilis, Mayden and Burr

1981; N. miurus, Burr and Mayden 1982a; N. nocturnus , Burr and

Mayden 1982b). Mature females ranged in SL from 61 to 98 mm (x =

74.8; N = 16 for individuals 2 to 4 years old) and in adjusted body

weight from 4.82 to 17.85 g (x = 7.60; N = 16). No females older than 4

years or longer than 98 mm SL were seen.

Age at first spawning for males was not ascertained, although all

males found guarding nests or nest sites were 2 to 4 years old and longer

than 60 mm SL. Adjusted body weights of all males in breeding

condition ranged from 3.81 to 18.24 g (x= 8.46; N = 24).

Reproductive Condition in Males. In gross appearance, testes were

opaque white and fimbriate as in other ictalurids (see Sneed and

Clemens 1963; Mayden and Burr 1981). Weight of testes was positively

correlated with increasing body weight. For combined samples of

immature and mature males, linear regression of testes weight in

Carolina Madtom

73

OCTOBER

N= 20

O'

10-

juI

SEPTEMBER

N = 92

JUNE

N= 18

l_Lk

10

MAY

N = 92

STANDARD LENGTH (mm)

Fig. 5. Length-frequency distribution of Noturus furiosus in spring, summer,

and fall seasons. At least four age classes are represented.

milligrams (T) on adjusted body weight in grams (W) was T = -0.001 +

0.004W (r = 0.52; N = 17).

In 11 mature males collected from May to July, the GSI ranged

from 0.001 to 0.007 (x = 0.003). The largest relative testes weight

(equaling 0.7% adjusted body weight) was that of an 82-mm specimen

collected 8 June. The GSI of immature males was usually less than

0.002.

As in other species of the genus, there was no marked sexual

dimorphism outside the breeding season. Reproductively mature males

of N. furiosus had enlarged cephalic epaxial muscles, swollen lips, and

swollen genital papillae (Fig. 6). The male papilla was much like that

described for other species of Noturus in the subgenus Rabida (see Burr

and Mayden 1982a). The swollen heads of males are presumed to

function in nest guarding and perhaps in nest preparation.

Reproductive Condition in Females. Throughout the summer there

was little change in GSI for immature females (range = 0.001 to 0.008; x

= 0.005; N = 11). Lack of adequate numbers of specimens from the

remainder of the year prevented a comparison of GSI throughout fall,

winter, and spring. Ovarian growth in immature N. furiosus is

74

Burr, Kuhajda, Dimmick, and Grady

presumably similar to the variable, slight seasonal increase in other

species (Mayden and Burr 1981; Burr and Mayden 1982a, b).

GSI of mature females reached a peak in May (x GSI r 176) and

June (x GSI = 166) and began to taper off in July (x GSI = 105). Peak

GSI in May and June presumably corresponds to accelerated ovarian

recrudescence and oocyte vitellogenesis. Sixteen gravid females had a

mean adjusted body weight of 7.60 gm and a mean ovarian weight of

1.04 g (range = 0.34 to 2.34). The proportionally heaviest ovaries

(equaling 25.8% adjusted body weight) were those of a 70-mm female

collected from the Little River on 19 May (Table 1). GSI ratios from

gravid females of other medium-size to large madtoms collected May

through July were: 79 (N = 4) in N. nocturnus (Burr and Mayden

1982b), 107 (N = 7) in N.flavater (Burr and Mayden 1984), 145 (N = 12)

in N. miurus (Burr and Mayden 1982a), 149 (N = 33) in N. gyrinus

(Whiteside and Burr 1986), 175 (N = 3) in N. flavipinnis (Shute 1984),

and 21 1 (N = 11) in N. exilis (Mayden and Burr 1981).

Externally, gravid females appeared similar to immature specimens

except in having distended abdomens and swollen genital papillae

(Fig. 6).

Oocyte Diameter and Fecundity. As in congeners, ovaries of gravid

females contained two size classes of oocytes. Large, vitellogenic oocytes

were amber, ranged in diameter from 1.7 to 3.0 mm (x = 2.5; N = 130),

and were assumed to be the only oocytes spawned during one season.

Immature oocytes were small (0.2 to 1.8 mm; x = 0.9; N = 12) and

opaque white or yellow. In 17 gravid females there was a positive

correlation (r = 0.97) between mean oocyte diameter in mm (D) and

increasing GSI, with D = 1.46 + 0.006GSL

Vitellogenic oocytes in 17 gravid females ranged in number from 79

to 298 (x = 126.3). Asymmetry in oocyte number between right and left

ovaries occurred in several females, but the differences were not

statistically significant at the 0.05 level (Table 1). Skewed distributions

of oocytes have also been reported for N. exilis (Mayden and Burr

1981), N. flavus (Walsh and Burr 1985), and N. flavater (Burr and

Mayden 1984). As suggested by Walsh and Burr (1985), these differences

probably reflect individual variation since other gravid N. furiosus had

nearly equal numbers of oocytes between ovaries (Table 1).

The total number of mature oocytes in a female was positively

correlated with increasing SL, adjusted body weight, and age. For 17

gravid females, the regression of number of mature oocytes (F) on SL

was F = -248.60 + 5.0 1SL (r = 0.81), of the number of mature oocytes on

adjusted body weight in grams (W) was F = 16.37 + 14.47W (r = 0.90),

and of the number of mature oocytes on age in years (A) was

F = -140.30 + 87.04A (r = 0.77). The number of mature oocytes per gram

adjusted body weight ranged from 12.2 to 24.1 (x= 17.1; N = 16).

Carolina Madtom

75

Fig. 6. Genital papillae of Noturus furiosus. (A) 82-mm SL nonbreeding male,

19 September 1959, Tar River, Edgecombe-Nash counties (USNM 191057). (B)

79-mm SL breeding male, 20 May 1985, Little River, Wayne County (SIUC

1 1683). (C) 55-mm SL nonbreeding male, same as A. (D) 73-mm SL breeding

female, same as B. Left is anterior. 25X. Drawing by Karen L. Fiorino.

Sex Ratio. There was no significant deviation from a 1:1 sex ratio

in the total sample from North Carolina (126 males: 1 14 females), nor in

any of the monthly samples containing more than 10 individuals. In a

preliminary review, skewed sex ratios in other species of Noturus

(Mayden and Burr 1981) were thought to result from sampling bias,

particularly in older age classes.

Nesting. As judged from females with mature ova and males in

reproductive condition, the nesting season of N. furiosus extends from

about mid-May to the third week of July (Tables 1 and 2). The first

evidence of nesting N. furiosus was the discovery of a reproductively

76

Burr, Kuhajda, Dimmick, and Grady

mature pair in Little River, west of Goldsboro, in a 355-ml beer can on

22 May 1982; water temperature was 20° C in the shallow run where the

can was found. From 16 to 22 May 1985, in water ranging in

temperature from 22 to 25° C, we found two primary and two secondary

areas of nesting for N. furiosus. Both the Little River, west of Goldsboro,

and the Tar River in Tarboro contained substantial nesting populations

of the species. At the former site, in an area about 300 m in length,

seven solitary males in breeding condition were found in 355-ml cans

and bottles, or under shells (about 50 potential nesting sites were

examined); one female swollen with eggs was in a 355-ml can; three

pairs in reproductive condition were in cans; and one male with

embryos was in a beer can (Table 2). At Tarboro, in an area about 100

m in length, 11 solitary males in reproductive condition were found in

cans and bottles and under mussel shells (about 200 potential nest sites

were examined); two males, each with embryos, were found in glass

containers (Table 2). Subadults were numerous at both these sites, in

containers and other situations similar to those occupied by adults.

Secondary nesting areas for the species were found in the Tar River

in Rocky Mount and northwest of Heartsease. At both sites, we found

one breeding male with embryos or larvae in water ranging from 23 to

23.5° C. Only two other individuals, both juveniles, were found at each

of these sites. Potential nesting sites were uncommon at both areas.

Nests, pairs, and solitary adults in breeding condition were all

found in similar habitat; medium- to large-sized streams with pea- to

medium-sized gravel/ sand substrates and moderate-to-swift warm water

(22 to 25° C). Most nest sites were in runs above riffles or in pools with

current. Water depth varied from 26 to 120 cm (x = 66.4). All nests with

embryos or larvae were guarded by solitary males, 3 to 4 years old and

ranging in SL from 63 to 101 mm (x= 89.8). The stomachs of guardian

males were mostly empty or contained well-digested, unidentifiable

material.

Only three clutches captured were complete with 139, 147, and 171

embryos (x = 152). One clutch, containing more than 200 larvae and

guarded by the largest male, could not be captured (Table 2). A brood

of 16 larvae, considered incomplete, was captured with the guardian

male. Larvae were negatively phototaxic, and in one case dispersed

rapidly when nest cover was removed, precluding capture of the complete

brood.

Nest cans and bottles were generally free of substrate material, but

if present, it was tightly packed against the bottom of the container.

Nest containers or cover usually were partially buried in the substrate.

Nests were found as close as 5 m apart and ranged from a few

centimeters from the bank to the center of the stream.

Carolina Madtom

77

Cans or bottles with small openings are probably favored as nest

sites by large males because they are easy to protect. Head width in

eight reproductively mature males ranged from 15.1 to 29.0 mm (x =

22.4) and head depth ranged from 7.8 to 16.0 mm (x = 12.3). Pop-top

beer and soda can openings ranged from 18.5 to 28.0 mm in length (x =

23.7) and from 13.0 to 20.0 mm in width (x = 16.2). Thus, there is little

room to spare at the opening of a can once a male is inside. Guardian

males with their swollen heads generally face the opening of the

container, presumably blocking the entry of most potential predators. It

is not known whether a male enters a can with his head muscles and lips

already swollen or if these changes take place after he has selected his

nest site.

During May we found no instances of syntopic species of Noturus

nesting in areas with N. furiosus. Both N. gyrinus and N. insignis were

uncommon or not found in N. furiosus nesting habitat. Crayfishes,

juvenile madtoms, A. rostrata, and Necturus lexvisi may compete for

nest or hiding sites with N. furiosus. The waterdog and N. furiosus are

sympatric, and one nest of the former has been found under a flat rock

(Ashton and Braswell 1979). Although we did not find any N. furiosus

nests under rocks, the species may use such sites at least occasionally,

especially since close relatives of the species have been found to do so

(see Burr and Mayden 1982a, 1984). Like other ictalurid catfishes, N.

furiosus is a member of the speleophil reproductive guild as defined by

Balon (1975).

Clearly, N. furiosus has taken advantage of human litter as potential

nest sites, but the presence of several adults in reproductive condition

under mussel shells (mostly Elliptio complanata ) and bark indicates that

before the advent of modern man these were perhaps the only form of

adequate-sized cover available. Our survey work indicates that the

number of artificial nest sites (i.e., cans and bottles) far exceeds the

number of adequate-sized natural sites in the streams we intensively

sampled.

Larval Development. As in other species of Noturus, embryos

adhere to each other in a mass, but not to other surfaces. Chorion

diameters of 45 pre-hatchling embryos ranged from 3.8 to 4.4 mm (x =

3.9 mm). Yolk sacs of developing embryos were cream to light yellow,

similar to yolks of N. flavus (Walsh and Burr 1985), but in contrast to

the darker yellow or nearly orange yolks in N. albater (Mayden et al.

1980), N. exilis (Mayden and Burr 1981), N. miurus (Burr and Mayden

1982a), and N. nocturnus (Burr and Mayden 1982b). Yolk diameters

averaged 3.2 mm (range = 2.5 to 4.0; N = 45).

At about 1 day post-hatching, larvae ranged in TL from 9.1 to 10.0

mm (x = 9.5; N = 16), had well-developed maxillary and mandibular

78

Burr, Kuhajda, Dimmick, and Grady

barbels, rudimentary nasal barbels, and darkly pigmented retinae; small

pectoral and pelvic fin buds; and a continuous posterior fin fold

heightened in the regions of the anal and dorsal fins, and with

rudimentary caudal and anal rays (Fig. 7). A sprinkling of melanophores

occurred on the head and along the dorsal myomeres. Hatchlings of N.

furiosus exhibited tightly cohesive schooling behavior. Larvae in later

stages of development were not found. Early post-hatching larvae of N.

furiosus closely resembled N. nocturnus (Burr and Mayden 1982b) and

N. miurus (Burr and Mayden 1982a) in shape, pigmentation, and

overall developmental features. Early post-hatching larvae of other

species of Noturus are more heavily pigmented when compared with N.

furiosus (e.g., N. exilis, Mayden and Burr 1981; N. flav at er, Burr and

Mayden 1984).

The smallest juveniles known are 17 mm SL, and they have the

body shape and pigmentation pattern typical of adults.

Diet. A total of 200 stomachs of N. furiosus were examined; 88

were empty, 18 had unidentifiable disgested material, and 94 contained

some food. The large percentage of empty stomachs is probably a

sample bias resulting from many daytime collections. Madtoms feed

primarily during the evening with peaks at dawn and dusk (Mayden and

Burr 1981).

Dipteran, ephemeropteran, trichopteran, coleopteran, and odonate

larvae or nymphs compose more than 95% of the total food organisms

(Table 3). Dipteran larvae and ephemeropteran nymphs were the most

commonly eaten food source. Chironomid larvae accounted for 91% of

the dipterans eaten, with the remainder being culicid and simuliid

larvae. Ephemeropteran nymphs included members of the families

Baetiscidae, Heptageniidae, and Caenidae. Larvae of Hydropsychidae

were the predominant (69%) representatives of Trichoptera. Elmid

larvae made up 95% of the coleopterans, and almost all of the elmids

were members of the genus Stenelmis , with only one individual of

Dubiraphia represented. Odonates included nymphs of the genus

Hagenius, in the family Gomphidae, and one representative of

Coenagrionidae. Nematodes found in the stomachs were possibly

parasitic. Four individuals had large fish scales in their stomachs,

probably indicating benthic scavenging activities. Individual madtoms

ate a variety of organisms; one had representatives of six different taxa

in its stomach. Several individuals had sand grains and plant material

mixed in with their food, which probably was ingested incidentally.

Larger individuals ate larger food items (Table 3). The relatively

small-sized dipteran larvae decreased in the diet with increasing size of

the madtom. Percentages of larval dipterans in the diet of the three size

groups of madtoms listed in Table 3 varied from 35.5 to 26.8 to 0. In

Carolina Madtom

79

Fig. 7. Lateral view of 9.5-mm TL larva of Noturus furiosus (SIUC 1 1775).

Drawing by Karen L. Fiorino.

contrast, percentages of the larger-sized odonate nymphs in the diet

increased from 5.1 to 6.6 to 30.4 with increasing size of the madtom.

Comparisons of spring and summer diets are shown in Figure 8.

Elmid larvae were a significant food organism in the spring, but were

negligible in the summer samples. Simuliid larvae were present only in

summer collections, and the percentages of ephemeropteran nymphs

and trichopteran larvae increased in the madtom’s diet in summer. The

abundance of chironomid larvae and odonate nymphs in the diet of N.

furiosus was unaffected by the changing seasons.

It appears that N. furiosus , like other madtoms, is a nocturnal,

benthic insectivore. All madtoms studied to date are taste feeders and

are morphologically equipped for taste feeding with numerous gustatory

structures.

DISCUSSION

Historically, at least 24 distinct localities of occurrence of N.

furiosus have been recorded based on extant voucher material. Additional

literature records (Bayless and Smith 1962, Smith and Bayless 1964)

bring the total to 36. We have revisited all but ten of these sites since

1982. Nine sites in the Neuse River drainage and six in the Tar River

drainage have not yielded specimens. Several of these sites were too

flooded for adequate sampling and may harbor extant populations (e.g.,

Fishing Creek). In addition, we discovered seven new populations of N.

furiosus in the Tar and Neuse River drainages in 1984 and 1985,

indicating that successful and substantial reproduction has taken place

in recent years.

Prior to our field work in 1985, N. furiosus was considered to be a

rare species. This judgement was the general consensus of several

ichthyologists who had tried to collect the species in the 1970s and

80

Burr, Kuhajda, Dimmick, and Grady

Table 3. Stomach contents of Noturus furiosus from the Tar and Neuse river

drainages, North Carolina, by size class of madtom. Figures to the

left are percentages of diet derived from each food organism;

parenthetical figures are percent of stomachs in which food organism

occurred.

Carolina Madtom

81

Fig. 8. Composition of the diet of Noturus furiosus by spring and summer

seasons. Top figures are percentages of total number of food organisms

consumed; parenthetical figures are percentages of stomachs in which food

organisms occurred.

82

Burr, Kuhajda, Dimmick, and Grady

1980s. Further, the extensive trap data involved in collecting biological

information on Necturus lewisi (Braswell and Ashton 1985) did not

reveal a single specimen of the sympatric and syntopic Noturus furiosus.

Collecting conditions were ideal during our field work in May 1985 and

we concentrated our field efforts on locating only N. furiosus. These

factors probably contributed to our success. Because we found that N.

furiosus primarily inhabitats medium- to large-size streams, it is clear

that the species would be difficult to collect except when water levels are

low.

Although N. furiosus is relatively common at some sites, the species

appears to have experienced a decline and loss of habitat in other areas.

The greatest losses of N. furiosus habitat have occurred in the Neuse

drainage. Reservoir construction (Falls Lake), outflow of cold waters

below Falls Lake, and general pollution problems around Raleigh have

reduced habitat in the upper Neuse. A toxic chemical spill into the

Neuse River near Raleigh on 10 July 1980 caused a large fish kill, but

no N. furiosus were found. The Tar drainage seems to have experienced

fewer cases of severe habitat degradation. However, the Tar River from

below Rocky Mount to about 20 km downstream showed evidence of

extensive municipal and industrial effluents. We did not find N. furiosus

in that region, and Necturus lewisi is also absent there (Braswell and

Ashton 1985).

A number of federal and state projects are presently being completed

or are in the planning phase for both the Neuse and Tar drainages.

Most of these projects call for (1) removal of stream cover, (2) denuding

of stream banks, (3) dredging, (4) channelization, or (5) reservoir

construction at a number of localities (e.g., Fishing Creek, Tar River

near Tarboro, Contentnea Creek, Trent River) where N. furiosus is

known to occur. We predict that these activities will have a detrimental

effect on the quality of the habitat of N. furiosus, either by changing the

habitat altogether (e.g., reservoir construction), or by severely modifying

it (e.g., dredging).

Natural factors affecting the continued existence of N. furiosus

include a potential increase in predation owing to desiccation of streams

during drought. During late summer and fall most streams in the Neuse

and Tar drainages are reduced to low flow because of little rainfall. The

riffle habitat of the Carolina madtom is thus restricted in size, and the

species may be subject to increased predation by fish-eating birds and

snakes. Adults of other madtoms are eaten only rarely by piscivorous

fish, and in laboratory experiments they are usually eaten as a last

choice, probably because of their stout spines and the toxin-producing

glands associated with the spines (Case 1970). Predators we have

observed eating other species of adult Noturus on several occasions are

Carolina Madtom

83

water snakes of the genus Nerodia. However, most predation on N.

furiosus probably takes place during the larval stage. In other madtoms,

removal of guardian males from nest sites results in rapid loss of young

to fishes and crayfishes (Mayden et al. 1980, Mayden and Burr 1981).

Presently, N. furiosus is placed in the category of special concern

on North Carolina’s list of endangered and threatened animals (Bailey

et al. 1977); it is only being considered for listing by the Department of

the Interior. Nonetheless, fisheries biologists need to be aware of the

vulnerability of the species to sampling techniques. In the early 1960s,

numerous collections of N. furiosus were made from the Tar and Neuse

drainages using the ichthyocide rotenone (Bayless and Smith 1962,

Smith and Bayless 1964). Unfortunately, a majority of these rotenone

collections were made in June and July, during the breeding season of

N. furiosus. Because ichthyocides are extremely effective in killing

madtoms, we believe that the indiscriminate use of rotenone in North

Carolina stream surveys should be discouraged. Careful regulation and

monitoring may even be justified.

The general biology of N. furiosus as outlined here is similar to that

in previous reports of other species in the genus. There are, however,

two important aspects of reproduction in Noturus that remain unresolved.

Several authors (Menzel and Raney 1973, Mayden and Burr 1981,

Walsh and Burr 1985) have presented circumstantial evidence that

females in some species of Noturus may spawn with more than one male

in a breeding season. In several species, the mean number of vitellogenic

oocytes is about twice the mean number of embryos found in complete

broods. In this study, the mean number of vitellogenic oocytes (126.3)

was somewhat less than the mean number of embryos (152.0) from

complete broods of N. furiosus. Because of the positive correlation of

female body size with fecundity, we assume that polyandry does not

occur in this species. This assumption is supported by the results Blumer

(1985a) reported from field and laboratory experiments with a related

species, the brown bullhead, Ictalurus nebulosus.

One additional aspect of reproduction that remains unresolved in

Noturus is the contribution of females to parental care of embryos and

larvae. In all known studies of Noturus , including this one, only males

have been found guarding embryos or larvae. Because males generally

do not feed during the nesting and care-giving period they sustain a

greater cost of care giving (starvation and therefore reduced future

reproduction) than do males aided by their mates (Blumer 1985b).

Although a great deal has been learned about the natural history of

madtoms in the last five years, we still lack basic knowledge of

(1) spawning behavior, (2) nest construction (if any), (3) social behavior,

(4) contribution of females in parental care of embryos and larvae,

84

Burr, Kuhajda, Dimmick, and Grady

(5) number of clutches spawned per year by a single female, (6) behavior

and distribution of young after leaving the nest, (7) critical diurnal

habitats, (8) movements or migrations, (9) nocturnal behaviors, and

(10) winter habitat occurrence. Because none of this information is

known for N. furiosus, further research is needed to ensure our ability

to preserve and protect the species. Until propagation techniques have

been developed for madtoms, we recommend that spawning sites of N.

furiosus be protected and that collectors be discouraged from sampling

prime nesting areas.

ACKNOWLEDGMENTS.— We are grateful to Patti A. Burr,

Michelle J. Burr, Kevin S. Cummings, Michael A. Klutho, and Fred C.

Rohde for aid in collecting specimens. The following curators or staff

loaned specimens, provided laboratory space and locality information,

and extended numerous other courtesies: Barry Chernoff, Academy of

Natural Sciences at Philadelphia; John G. Lundburg, Duke University;

Alvin L. Braswell, John E. Cooper, David S. Lee, and William M.

Palmer, North Carolina State Museum of Natural Sciences, Raleigh;

Robert R. Miller and Douglas W. Nelson, University of Michigan

Museum of Zoology; and Susan Jewett, Wayne C. Starnes, and William

R. Taylor, National Museum of Natural History. Edward F. Menhinick,

University of North Carolina at Charlotte, provided us with locality

information for many records of N. furiosus. William Adams, U.S.

Army Corps of Engineers, Wilmington district, outlined major proposed

federal projects that might adversely affect the habitat of N. furiosus.

Frank J. Schwartz, University of North Carolina Institute of Marine

Sciences, Morehead City, was a generous and always helpful host to

BMB during his sabbatical leave (1983-1984). Karen Fiorino and Karen

Schmitt, Southern Illinois University at Carbondale, assisted in the

preparation of figures. Renaldo Kuhler, North Carolina State Museum

of Natural Sciences, prepared the illustration of Noturus furiosus.

This project was made possible through the efforts of Richard G.

Biggins and David S. Lee and was supported, in part, by a contract with

the U.S. Fish and Wildlife Service, the North Carolina State Museum

of Natural Sciences, and the SIUC Graduate School.

LITERATURE CITED

Ashton, Ray E., Jr. and A. L. Braswell. 1979. Nest and larvae of the Neuse

River waterdog, Necturus lewisi (Brimley) (Amphibia: Proteidae).

Brimleyana 1:15-22.

Bailey, Joseph R., and Committee. 1977. Freshwater fishes. Species list. Pages

278-280 in Endangered and Threatened Plants and Animals of North

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

N.C. State Mus., Raleigh.

Carolina Madtom

85

Balon, Eugene K. 1975. Reproductive guilds of fishes: a proposal and

definition. J. Fish. Res. Bd. Canada 32:821-864.

Bayless, Jack D., and W. B. Smith. 1962. Survey and classification of the

Neuse River and tributaries, North Carolina. Final Report, Federal Aid in

Fish Restoration, Job I-A, Project F-14-R. N.C. Wildl. Resour. Comm.,

Raleigh.

Blumer, Lawrence S. 1985a. Reproductive natural history of the brown

bullhead Ictalurus nebulosus in Michigan. Amer. Midi. Nat. 114:318-330.

Blumer, Lawrence S. 1985b. The significance of biparental care in the brown

bullhead, Ictalurus nebulosus. Environ. Biol. Fishes 12:231-236.

Braswell, Alvin L., and R. E. Ashton, Jr. 1985. Distribution, ecology, and

feeding habits of Necturus lewisi (Brimley). Brimleyana 10:13-35.

Burr, Brooks M., and R. L. Mayden. 1982a. Life history of the brindled

madtom, Noturus miurus, in Mill Creek, Illinois (Pisces: Ictaluridae).

Amer. Midi. Nat. 107:25-41.

Burr, Brooks M., and R. L. Mayden. 1982b. Life history of the freckled

madtom, Noturus nocturnus, in Mill Creek, Illinois (Pisces: Ictaluridae).

Occas. Pap. Mus. Nat. Hist. Univ. Kans. 98:1-15.

Burr, Brooks M., and R. L. Mayden. 1984. Reproductive biology of the

checkered madtom ( Noturus flavater) with observations on nesting in the

Ozark (N. albater ) and slender ( N . exilis ) madtoms (Siluriformes:

Ictaluridae). Amer. Midi. Nat. 112:408-414.

Case, Brian F. 1970. An ecological study of the tadpole madtom Noturus

gyrinus (Mitchill) with special reference to movements and population

flucuations. Unpubl. M.S. thesis, Univ. Manitoba, Winnipeg.

Cashner, Robert C., and R. E. Jenkins. 1982. Systematics of the Roanoke

bass, Ambloplites cavifrons. Copeia 1982:581-594.

Clugston, James P., and E. L. Cooper. 1960. Growth of the common eastern

madtom, Noturus insignis, in central Pennsylvania. Copeia 1960:9-16.

Cooper, John E., and R. E. Ashton, Jr. 1985. The Necturus lewisi study:

introduction, selected literature review, and comments on the hydrologic

units and their faunas. Brimleyana 10:1-12.

Cooper, John E., and A. L. Braswell. 1982. Noturus furiosus Jordan and

Meek, 1889. Carolina madtom. Account Red Data Book, Conservation

Monitoring Unit, International Union for Conservation of Nature and

Natural Resources, Cambridge, United Kingdom.

Douglas, Neil H. 1972. Noturus taylori, a new species of madtom (Pisces,

Ictaluridae) from the Caddo River, southeastern Arkansas. Copeia

1972:785-789.

Etnier, David A., and R. E. Jenkins. 1980. Noturus stanauli, a new madtom

catfish (Ictaluridae) from the Clinch and Duck Rivers, Tennessee. Bull.

Ala. Mus. Nat. Hist. 5:17-22.

Jordan, David S. 1889. Description of fourteen species of fresh-water fishes

collected by the U.S. Fish Commission in the summer of 1888. Proc. U.S.

Natl. Mus. 11:351-362.

LeGrande, William H. 1981. Chromosomal evolution in North American

catfishes (Siluriformes: Ictaluridae) with particular emphasis on the

madtoms, Noturus. Copeia 1981:33-52.

86

Burr, Kuhajda, Dimmick, and Grady

Mayden, Richard L., and B. M. Burr. 1981. Life history of the slender

madtom, Noturus exilis, in southern Illinois (Pisces: Ictaluridae). Occas.

Pap. Mus. Nat. Hist. Univ. Kans. 93:1-64.

Mayden, Richard L., B. M. Burr, and S. L. Dewey. 1980. Aspects of the life

history of the Ozark madtom, Noturus albater, in southeastern Missouri

(Pisces: Ictaluridae). Amer. Midi. Nat. 104:335-340.

Menzel, Bruce W., and E. C. Raney. 1973. Hybrid madtom catfish, Noturus

gyrinus X Noturus miurus, from Cayuga Lake, New York. Amer. Midi.

Nat. 90:165-176.

Shute, Peggy W. 1984. Ecology of the rare yellowfin madtom, Noturus

flavipinnis, in Citico Creek, Tennessee. Unpubl. M.S. thesis, Univ.

Tennessee, Knoxville.

Smith, William B., and J. D. Bayless. 1964. Survey and classification of the

Tar River and tributaries, North Carolina. Final Report, Federal Aid in

Fish Restoration, Job I-L, Project F-14 — R. N.C. Wildl. Resour. Comm.,

Raleigh.

Sneed, Kermit, E., and H. P. Clemens. 1963. The morphology of the testes and

accessory reproductive glands of the catfishes (Ictaluridae).

Copeia 1963:606-611.

Taylor, William R. 1969. A revision of the catfish genus Noturus Rafinesque

with an analysis of higher groups in the Ictaluridae. Bull. U.S. Natl. Mus.

282:1-315.

Walsh, Stephen J., and B. M. Burr. 1985. Biology of the stonecat, Noturus

flavus (Siluriformes: Ictaluridae), in central Illinois and Missouri streams,

and comparison with Great Lakes populations and congeners. Ohio J. Sci.

85:85-96.

Whiteside, Lisa A., and B. M. Burr. 1986. Aspects of the life history of the

tadpole madtom, Noturus gyrinus (Siluriformes: Ictaluridae), in southern

Illinois. Ohio J. Sci. 86:153-160.

Accepted 12 June 1987

Pelagic and Near-shore Plankton Communities

of a North Carolina Coastal Plain Reservoir

Michael A. Mallin

Department of Environmental Sciences and Engineering,

University of North Carolina at Chapel Hill,

Chapel Hill, North Carolina 27514

ABSTRACT.— The plankton community of Sutton Reservoir, an

estuarine-influenced cooling reservoir in the coastal plain of North

Carolina, was sampled monthly throughout 1985. Phytoplankton

densities were low to moderate and dominated by the Chlorophyceae

and Cryptophyceae, with chlorophyll a values indicating a mesotrophic

state. The zooplankton community was unusual compared with other

area reservoirs and reflected the estuarine influence of the lower Cape

Fear River and the reservoir’s geographic location. Crustacean

zooplankton were dominated by Diaptomus dorsalis, Eurytemora

affinis, Daphnia ambigua, and Bosmina coregonv, and the rotifers were

dominated by members of the Brachionidae. Zooplankton densities,

which exhibited a bimodal peak, were high relative to area reservoirs

and dominated by rotifers. Biomass was comparatively low and

dominated by crustaceans. Correlation analysis indicated a strong,

inverse relationship between crustacean zooplankton and phytoplank-

ton, and zooplankton grazing is suggested as the primary controlling

force in phytoplankton temporal population dynamics.

Sutton Reservoir, in New Hanover County, North Carolina, is one

of several water bodies located in the coastal plain of the southeastern

United States. These systems comprise both man-made reservoirs and

natural lakes (the Carolina bays). Most of these systems are very

important to wildlife, particularly migratory waterfowl; and bay lakes

often contain rare or endemic species (Sharitz and Gibbons 1982).

Sutton Reservoir is ecologically interesting in that it is subject to

both chemical and biological influence from the lower Cape Fear River

and displays characteristics of both fresh and brackish waters. The

objectives of this study were to describe the zooplankton community of

this unusual ecological system, compare the pelagic and near-shore

communities, discuss the relationship of the zooplankton community

with the reservoir’s chemical and physical characteristics and phyto-

plankton, and compare the Sutton Reservoir zooplankton with those of

other Southeastern systems.

Little has been reported in the literature regarding the plankton

communities of coastal-plain lakes and impoundments. Stoneburner

and Smock (1980) reported on the plankton community of an acid,

Brimleyana No. 15:87-101, January 1989

87

88

Michael A. Mallin

brownwater lake on a south Georgia barrier island. They found a sparse

phytoplankton community dominated by Chlamydomonas sp., Melosira

varians, and Peridinium pusillum. The zooplankton community

maintained high densities and was dominated by Diaptomus floridanus ,

Polyarthra vulgaris , Keratella cochlearis, and Daphnia ambigua. The

Great Dismal Swamp ecosystem in Virginia and northeastern North

Carolina was investigated by Anderson et al. (1977). They found that

the system was dominated by rotifers with high densities of Polyarthra

vulgaris and Conochiloides dossuarius. Dominant crustaceans were

Bosmina longirostris , Diaphanosoma leuchtenbergianum , Mesocyclops

edax, and Tropocy clops prasinus. Casterlin et al. (1984) studied the

algae of Lake Waccamaw, a large North Carolina bay lake, and found

increasing eutrophication occurring. The North Carolina Department of

Environmental Management has reported data concerning the mid-

summer algae and chlorophyll of several coastal-plain lakes (NCDEM

1984). The scarcity of plankton information about coastal-plain systems

leaves mainly inland water systems for comparisons.

SITE DESCRIPTION

Sutton Reservoir (Catfish Lake) is a 445-ha impoundment located

4.8 km northwest of Wilmington, N.C., adjacent to the Cape Fear

River. It was constructed in 1972 to provide cooling water for the L. V.

Sutton Steam Electric Plant, a 677-MWe closed-circulation, coal-fired

generating facility operated by Carolina Power & Light Company

(CP&L). The reservoir is U-shaped with a series of baffle dikes and has

a mean depth of about 2 m and a maximum depth of about 12 m (Fig.

1). It has a retention time of approximately 140 days, and the circulation

time around the reservoir is about 4 days. Power plant discharge has

caused midsummer reservoir water temperatures of 32 to 35 °C in

recent years (CP&L 1986).

The reservoir has no constant influent stream but receives intermit-

tent makeup water from the Cape Fear River, particularly during the

summer. The ionic composition of the water reflects the estuarine

influence (Table 1). The lake is well oxygenated, well mixed, and

circumneutral in pH (CP&L 1986). The fish community is dominated

primarily by typical Southeastern freshwater species such as largemouth

bass ( Micropterus salmoides ), gizzard shad ( Dorosoma cepedianum ),

bluegill ( Lepomis macrochirus ), and several other sunfish. Estuarine

influence is indicated by the presence of species such as mullet ( Muzil

cephalus), flounder ( Paralichthys spp.), and blue crabs ( Callinectes

sapidus ). Between 1972 and 1980 infestations of bladderwort ( Utricularia

vulgaris ) were common in Sutton Reservoir, with growth dense enough

in 1979 to cause a plant shutdown. Introduction of the redbelly tilapia

Reservoir Plankton Communities

89

Fig. 1. Map of Sutton Reservoir, New Hanover County, N.C., showing

sampling stations and baffle dikes.

( Tilapia zilli) in 1980 subsequently controlled the bladderwort (CP&L

1984b). In 1982 dense growths of pondweed ( Potamageton berchtoldii),

southern naiad ( Najas guadalupensis ), and coontail ( Ceratophyllum

demersum) supplanted the bladderwort and were not effectively

controlled by the tilapia. A dense blue-green algal bloom occurred

during the summer of that year as well (CP&L 1984a). The macrophytes

were controlled and nearly eliminated by the use of herbicides in

subsequent years (Schiller 1985). Increased densities of blue-green algae

still occur during summer, but nuisance blooms have not occurred since

the macrophytes were eliminated in 1983.

90

Michael A. Mallin

MATERIALS AND METHODS

Zooplankton samples were collected monthly from January 1985

through December 1985 at two locations using a 10.5-liter Juday-style

closing plankton trap fitted with a 75-jum mesh net. Traps have been

found to be more efficient than nets for zooplankton samples, especially

for smaller forms (Kankaala 1984). The 75-/im mesh is more efficient

than a larger size for capturing microzooplankton (Evans and Sell

1985). Station 6B was a midreservoir station over deep water (10 m),

and Station 6C was a near-shore station in about 1 m of water. Three

replicate samples were taken at each station at about the 0.5-m depth.

Samples were field preserved with formalin to 2% of volume.

In the laboratory the samples were mixed, and an aliquot containing

at least 100 organisms was removed and placed in a circular counting

chamber. Copepods, cladocerans, rotifers, and protozoans were counted

using a dissecting microscope and identified to the lowest practical

taxon using a compound microscope. Taxonomic keys included Pennak

(1953), Brooks (1957, 1959), Voigt (1951), Edmondson (1959), and

Wilson and Yeatman (1959). Biomass of zooplankton was determined

using literature biomass values for piedmont reservoir zooplankton

(Horton and Carter 1980). Biomass of species not listed was determined

by applying dry-weight regression equations (Dumont et al. 1975) to

specimens measured for length in the laboratory. A two-way analysis of

variance blocked on months was used to compare various loge (X + 1)

transformed zooplankton population density and biomass variables

between stations. A Type I error significance level of a - 0.05 was used

in the analyses. Densities are reported as number/ m3 and biomass as

mg/m3. Correlation analyses were run to detect linear relationships

between water temperature and selected zooplankton and plytoplankton

variables.

Phytoplankton and chlorophyll a were sampled monthly at a single

midwater station by combining whole water samples from the surface,

Secchi depth, and twice Secchi depth. Fifty ml of field-preserved sample

were sedimented in Utermohl settling chambers and examined for

phytoplankton taxonomic composition and density at 400X using an

inverted microscope. Chlorophyll a analysis was conducted spectro-

photometrically using the method described in Strickland and Parsons

(1972).

Monthly surface water samples were collected and analyzed for

nutrients and other chemical constituents by the CP&L Analytical

Chemistry Laboratory according to standard methods (USEPA 1979,

APHA 1981). Field measurements of water temperature, dissolved

oxygen, pH, and conductivity were also taken on a monthly basis

concurrent with the plankton samples.

Reservoir Plankton Communities

91

Table 1. Surface water characteristics of Sutton Reservoir during 1985.

Parameter

H20 temperature (°C)

Secchi depth (m)

pH

Conductivity (juS/cm)

P04 - P (mg / 1)

N02 + N03 - N (mg / 1)

Total alkalinity (mg/ 1 CaCo3)

Sodium (mg / 1)

Chloride (mg/ 1)

Chlorophyll a(pgl 1)

RESULTS AND DISCUSSION

Phytoplankton densities in Sutton Reservoir ranged from 480

units/ml in March to 7260 units/ml in June during 1985 (Fig. 2).

Densities and chlorophyll a values were moderate compared with other

lakes in the North Carolina coastal plain (NCDEM 1984). The

Chlorophyceae and Cryptophyceae were the two most important

phytoplankton classes, followed by the Bacillariophyceae and Cyano-

phyceae, respectively. Nuisance blue-green algal blooms observed during

1982 and 1983 were not manifested in 1985. Biomass as chlorophyll a

was greatest in June when densities of Cryptomonas ovata, Peridinium

spp., or larger diatom taxa such as Melosira were high (Fig. 2). Mean

chlorophyll a levels (5.7 pg/ 1) suggest a mesotrophic state for Sutton

Reservoir (Wetzel 1983).

Thirty-five zooplankton taxa from the Copepoda, Cladocera, and

Rotifera were identified from Sutton Reservoir samples collected during

1985 (Table 2). A great majority of the taxa were rotifers, with the

Brachionidae particularly well represented. Few of the crustaceans

were numerically dominant in the reservoir. Those which were dominant

included the copepods Diaptomus dorsalis and Eurytemora affinis and

the cladocerans Daphnia ambigua and Bosmina coregoni (Table 3).

Crustacean zooplankton densities displayed a bimodal peak in early

spring and again in fall and winter, with a minimum from May through

August. The rotifers generally maintained high densities from May

through the end of the year, with various species of Brachionus and

Keratella usually dominating. Other rotifers with high densities at

various times included Ascomorpha sp. and Conochiloides natans.

DENSITY (units/ml)

92 Michael A. Mallin

MONTH

PHYTOPLANKTON DENSITY

CHLOROPHYLL a

Fig. 2. Total phytoplankton densities and chlorophyll a values in Sutton

Reservoir during 1985.

Zooplankton densities (Table 4) were dominated by rotifers and

were high relative to other North Carolina reservoirs (Weiss et al. 1975,

DPC 1977, Mallin 1986). Zooplankton biomass values (Table 4)

indicated biomass dominance by copepods followed by cladocerans and

then rotifers. Zooplankton biomass in Sutton is low compared with

other North Carolina impoundments (Mallin 1986).

Temporal changes in dominance by organism density are illustrated

by the percent compostion over time of the total density by the major

taxa groups (Fig. 3). Rotifers comprised large percentages of the

densities in all seasons except spring. When the composition of biomass

in Sutton Reservoir is examined over time (Fig. 4), it is evident that

rotifers constitute a very small percentage of the zooplankton biomass,

which has two notable peaks each year. In spring the major contributors

to biomass were the copepod Eurytemora affinis followed by the

cladocerans Daphnia ambigua, Ceriodaphnia quadrangula, Diaphano-

soma brachyurum, Bosmina coregoni , and B. longirostris. In the fall,

CHLOROPHLLa (mq/I)

Reservoir Plankton Communities

93

Table 2. Zooplankton taxa identified from Sutton Reservoir during 1985.

Copepoda

Diaptomus dorsalis Marsh

Eurytemora affinis (Poppe)

Cyclops vernalis Fischer

Mesocyclops edax (Forbes)

Eucyclops agilis (Koch)

Cladocera

Daphnia ambigua Scourfield

Daphnia parvula Fordyce

Ceriodaphnia quadrangula (O. F. Muller)

Bosmina longirostris (O. F. Muller)

Bosmina coregoni Baird

Ilyocryptus spinifer Herrick

Alona sp.

Chydorus sphaericus (O. F. Muller)

Diaphanosoma brachyurum (Lievin)

Rotifera

Brachionus havanaensis (Rousselet)

Brachionus quadridentatus (Hermann)

Brachionus plicatilis (O. F. Muller)

Keratella americana (Carlin)

Keratella cochlearis (Gosse)

Keratella valga (Ehrenberg)

Keratella sp.

Platyias patulus (O. F. Muller)

Lecane sp.

Monostyla sp.

Trichocerca longiseta (Schrank)

Ascomorpha sp.

Asplanchna sp.

Synchaeta spp.

Polyarthra spp.

Filinia longiseta (Ehrenberg)

Pompholyx sulcata (Hudson)

Hexarthra sp.

Conochilus unicornis (Rousselet)

Conochiloides natans (Seligo)

Collotheca sp.

Table 3. Mean densities (no./m3) of important zooplankton taxa groups in Sutton Reservoir during 1985.

94

Michael A. Mallin

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Reservoir Plankton Communities

95

Table 4. Yearly mean densities (no./m3) and biomass (mg/m3) of major

zooplankton taxa groups at Sutton Reservoir during 1985.

c

a>

o

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a>

CL

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

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i I Rotifers

Fig. 3. Percent composition of total zooplankton density by major taxa groups

for Sutton Reservoir during 1985.

96

Michael A. Mallin

40

«

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

Month

■ Copepods

£v.v»l Cladocerans

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Fig. 4. Percent composition of total zooplankton biomass by major taxa groups

for Sutton Reservoir during 1985.

biomass was composed mainly of Diaptomus dorsalis , Daphnia ambigua,

B. coregoni, and Mesocy clops edax.

Comparisons were made to determine if there were differences

between the pelagic and near-shore zooplankton communities. The trap

sampling method was designed to capture typical planktonic species,

and therefore the common littoral taxa associated with the benthos or

littoral zone were not sampled. The results indicated that significantly

greater densities of copepods and rotifers were captured at Station 6B,

the pelagic location, than at 6C, the near-shore sampling station (Fig.

5). These differences were a result of greater densities of the copepods

Eurytemora affinis and Diaptomus dorsalis and the rotifers Brachionus

havanaensis, B. plicatilis, Keratella americana, K. cochlearis, and K.

valga in midwater as opposed to near shore.

The interesting ecological situation of Sutton Reservoir is illustrated

by its zooplankton taxonomic composition. The taxonomic composition

of the zooplankton is probably a result of the estuarine influence from

the makeup water and the reservoir’s coastal geographical location. The

two principal copepods, D. dorsalis and E. affinis , have not been

Organisms/m

Reservoir Plankton Communities

97

co

Month

Station 6B (pelagic)

G-EH3 Station 6C (nearshore)

Fig. 5. Total zooplankton densities by station for Sutton Reservoir during

1985.

described in the literature from North Carolina reservoirs. Diaptomus

dorsalis is typically found in states in the Gulf of Mexico region (Wilson

and Yeatman 1959). Eurytemora affinis is a variable marine form found

in North Carolina estuaries (Peters 1968), and its presence in Sutton

Reservoir is undoubtedly related to the estuarine influence of the lower

Cape Fear River. The cladocerans were mostly typical species of North

Carolina reservoirs with the exception of Ceriodaphnia quadr angularis .

Ceriodaphnia reticulata or C. lacustris are usually the members of that

genus reported from reservoir surveys of this area (Coker 1928, Weiss et

al. 1975, DPC 1977, Mallin 1986).

The taxa of rotifers found in Sutton Reservoir are normally present

in North Carolina reservoirs, but there were differences in the dominant

taxa. Members of the genus Brachionus have been noted in low

densities in several impoundments, but in Sutton Reservoir this genus

was dominant in the rotifer community. Some species of this genus

found in Sutton Reservoir, B. quadridentatus and B. plicatilis , are

abundant in salt or brackish water (Edmondson 1959). The appearance

of these two taxa coincided with an increase in chloride from 160 mg/ 1

98

Michael A. Mallin

in April to 260 mg/ 1 in July. The lack of taxonomic information in the

literature regarding other coastal-plain systems makes comparisons

difficult.

A strong phytoplankton-zooplankton interaction was suggested by

temporal population dynamics in Sutton Reservoir during 1985 (Fig. 2,

3, and 4). Based on the apparent inverse relationship shown on the

density figures, zooplankton grazing appeared to be a major factor

regulating the phytoplankton. To test this relationship, correlation

analyses were determined between important zooplankton and phyto-

plankton taxa groups (Table 5). Copepods, cladocerans, and selected

individual filter-feeding cladoceran taxa displayed significant linear

inverse relationships with total phytoplankton, Chlorophyceae, and

Cyanophyceae. These inverse relationships, along with the taxa of

phyto- and zooplankton present, suggest grazing was a controlling

factor in phytoplankton population dynamics. The chlorophyceae was

dominated by small, naked cells or flagellates such as Chlamydomonas,

Chlorella , and Selenastrum, which are considered easily accessible food

for filter-feeding crustaceans and are suppressed during grazing (Porter

1977, Vyhnalek 1983). The Chlorophyceae increased from 175 units/ml

in April to 995 units/ml in May, concomitant with a major crustacean

zooplankton decline (Table 2). The Cyanophyceae also showed a major

summer increase, but grazing is probably not a major controlling factor

for blue-greens, as members of this group are often difficult for

zooplankton to ingest and digest (Porter 1977). Increased water

temperatures (Tilman and Kiesling 1984, Lamberti and Resh 1985) or

nutrient availability (Tilman et al. 1986) are more probable Cyanophyte

controls in Sutton Reservoir.

SUMMARY

Sutton Reservoir is an estuarine-influenced coastal-plain system.

Phytoplankton densities in 1985 were low to moderate and dominated

by the Chlorophyceae and Cryptophyceae. A mean chlorophyll a value

of 5.7 fjig/ 1 suggests a mesotrophic state for this system.

The zooplankton taxonomic composition was unusual compared

with other North Carolina reservoirs and was probably a result of the

estuarine influence of makeup water from the lower Cape Fear River

and the reservoir’s geographic location. Dominant crustacean taxa were

Diaptomus dorsalis and Eurytemora affinis of the copepods and

Daphnia ambigua and Bosmina coregoni of the cladocerans. Rotifers

were dominated by various members of the Brachionidae. Densities

were high relative to other North Carolina impoundments and were

dominated by rotifers. Zooplankton biomass was comparatively low

and dominated by the crustaceans.

Reservoir Plankton Communities

99

Table 5. Results of correlation analyses between biomass of selected zoo-

plankton taxa, densities of selected phytoplankton taxa, and water

temperature, at Sutton Reservoir during 1985.

Total Water

* Results listed as correlation coefficients (r)/ probability (p)

Significantly greater densities of copepods and rotifers were

captured at a pelagic station as opposed to near shore. This was a result

of greater densities of E. affinis and D. dorsalis of the copepods and

Brachionus havanaensis, B. plicatilis, Keratella americana , K. cochlearis,

and K. valga at the pelagic station.

Correlation analyses indicated a strong, inverse relationship between

crustacean zooplankton (copepods and cladocerans) and phytoplankton.

Zooplankton grazing is suggested as a major controlling factor for

phytoplankton population dynamics with water temperature or nutrient

availability probably controlling the cyanophyceae.

ACKNOWLEDGMENTS.— I thank M. A. Pamperl and K. G.

Stone for phytoplankton data, M. M. Smart for chemical limnology

data and advice, and B. A. Carter and D. H. Schiller for sample

collection. The figures were produced by S. P. Price and the manuscript

was typed by members of the CP&L Harris Energy & Environmental

Center Word Processing Subunit.

100

Michael A. Mallin

LITERATURE CITED

Anderson, K. B., E. F. Benfield, and A. L. Buikema, Jr. 1977. Zooplankton of

a swampwater ecosystem. Hydrobiologia 55:177-185.

APHA. 1981. Standard Methods for the Examination of Water and

Wastewater. 15th ed. Amer. Pub. Health Assoc., Washington, D.C.

Brooks, J. L. 1957. Systematics of North American Daphnia. Mem. Conn.

Acad. Arts. Sci. 13:1-180.

Brooks, J. L. 1959. Cladocera. Pages 587-656 in Ward and Whipple’s

Freshwater Biology, 2nd ed., W. T. Edmondson, editor. John Wiley and

Sons, Inc., New York.

Casterlin, M. E., W. W. Reynolds, D. G. Lindquist, and C. G. Yarbrough.

1984. Algal and physicochemical indicators of eutrophication in a lake

harboring endemic species: Lake Waccamaw, North Carolina. J. Elisha

Mitchell Sci. Soc. 100:83-103.

Coker, R. E. 1928. Plankton collections in Lake James, North Carolina —

Copepods and Cladocera. J. Elisha Mitchell Sci. Soc. 41:228-258.

CP&L. 1984a. L. V. Sutton Steam Electric Plant Environmental Monitoring

Report 1982. Carolina Power & Light Co., New Hill, N.C.

CP&L. 1984b. L. V. Sutton Steam Electric Plant 1983 Annual Environmental

Monitoring Report. Carolina Power & Light Co., New Hill, N.C.

CP&L. 1986. L. V. Sutton Steam Electric Plant 1985 Annual Environmental

Monitoring Report. Carolina Power & Light Co., New Hill, N.C.

Dumont, H. J., I. Van de Velde, and S. Dumont. 1975. The dry-weight

estimate of biomass in a selection of Cladocera, Copepoda, and Rotifera

from the plankton, periphyton, and benthos of continental waters.

Oecologia 19:75-97.

DPC. 1977. Oconee Nuclear Station Environmental Summary Report 1971-

1976, Vol. 1. Duke Power Co. Steam Production Dept., Huntersville, N.C.

DPC. 1978. Baseline environmental summary report on the Yadkin River in

the vicinity of Perkins Nuclear Station. DUKEPWR/ 78-05. Duke Power

Co., Huntersville, N.C.

Edmondson, W. T. 1959. Rotifera. Pages 420-494 in Ward and VVhipple’s

Freshwater Biology, 2nd ed., W. T. Edmondson, editor. John Wiley and

Sons, Inc., New York.

Evans, M. S., and D. W. Sell. 1985. Mesh size and collection characteristics of

50-cm diameter conical plankton nets. Hydrobiologia 122:97-104.

Horton, W. T., and J. S. Carter. 1980. Estimation of zooplankton species body

weight in Lake Norman, North Carolina. Research Report EL/ 80/ 10.

Duke Power Co., Huntersville, N.C.

Kankaala, P. 1984. A quantitative comparison of two zooplankton sampling

methods, a plankton trap and a towed net, in the Baltic. Int. Revue ges.

Hydrobiol. 69:277-287.

Lamberti, G. A., and V. H. Resh. 1985. Distribution of benthic algae and

macroinvertebrates along a thermal stream gradient. Hydrobiologia

128:13-21.

Mallin, M. A. 1986. Zooplankton community comparisons among five

southeastern United States power plant reservoirs. J. Elisha Mitchell Sci.

Soc. 102:25-34.

Reservoir Plankton Communities

101

North Carolina Division of Environmental Management. 1984. Ambient Lakes

Monitoring Report 1983. N.C. Department of Natural Resources and

Community Development. Report No. 84-13.

Pennak, R. W. 1953. Freshwater Invertebrates of the United States. Ronald

Press, New York.

Peters, D. S. 1968. A study of relationships between zooplankton abundance

and selected environmental variables in the Pamlico River estuary of

eastern North Carolina. M.S. thesis. North Carolina State University,

Raleigh.

Porter, K. G. 1977. The plant-animal interface in freshwater ecosystems. Amer.

Scientist 65: 159-170.

Schiller, D. H. 1985. Aquatic vegetation control program for 1984. Carolina

Power & Light Co., New Hill, N.C.

Sharitz, R. R., and J. W. Gibbons. 1982. The ecology of southeastern shrub

bogs (pocosins) and Carolina bays: a community profile. FWS/OBS-82/04.

Fish and Wildl. Serv., U.S. Dept. Int., Atlanta, Ga.

Strickland, J. D. H., and T. R. Parsons. 1972. A Practical Handbook of

Seawater Analysis. Bull. Fish. Res. Bd. Canada, 167.

Stoneburner, D. L., and L. A. Smock. 1980. Plankton communities of an acid,

polymictic, brownwater lake. Hydrobiologia 69:131-137.

Tilman, D., and R. L. Kiesling. 1984. Freshwater algal ecology: taxonomic

trade-offs in the temperature dependence of nutrient competitive abilities.

Pages 314-319 in Current Perspectives in Microbial Ecology, Proceedings

of the 3rd International Symposium on Microbial Ecology, M. J. Klug and

C. A. Reddy, editors. Amer. Soc. Microbiol., Washington, D.C.

Tilman, D., and R. L. Kiesling, R. Sterner, S. S. Kilham, and F. A. Johnson. 1986.

Green, blue-green, and diatom algae: taxonomic differences in competitive

ability for phosphorus, silicon, and nitrogen. Arch. Hydrobiol. 106:473-485.

USEPA. 1979. Methods for the chemical analysis of water and wastes. U.S.

Environmental Protection Agency, EPA-600/ 4-79-020. Cincinnati, Ohio.

Voigt, M. 1957. Rotatoria, die radertiere Mittleleuropas. Vol. I and II.

Borntraeger, Berlin.

Vyhnalek, V. 1983. Effects of filter-feeding zooplankton on phytoplankton in

fish ponds. Int. Revue ges. Hydrobiol. 68:397-410.

Weiss, C. M., T. P. Anderson, P. H. Campbell, D. R. Lenat, J. H. More, S. L.

Pfaender, and T. G. Donnelly. 1975. An assessment of the environmental

stabilization of Belews Lake — Year IV and comparisons with Lake Hyco,

North Carolina, July 1973-June 1974. ESE Pub. No. 416. Dept, of Environ.

Sci. and Eng., School of Public Health, University of North Carolina,

Chapel Hill.

Wilson, M., and H. Yeatman. 1959. Free-living copepoda. Pages 735-861 in

Ward and Whipple’s Freshwater Biology, 2nd ed., W. T. Edmondson,

editor. John Wiley and Sons, Inc., New York.

Wetzel, R. G. 1983. Limnology. CRS College Publishing, Philadelphia, Pa.

Accepted 22 June 1987

102

FISHERMAN’S GUIDE

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by

Charles S. Manooch, III

Illustrated by Duane Raver, Jr.

Remarkable for its breadth of coverage, this book details the

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Technology, and Raver is nationally known for his paintings of wildlife.

“An excellent general reference book for scientific or non-scientific

audiences. ... It contains information not easily found in any other

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1984 376 pages Index Bibliography ISBN 0-917134-07-9

Price: $29.95, plus $1.25 for shipping. North Carolina residents add 5% sales

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Send to FISHERMAN’S GUIDE, N.C. State Museum of Natural Sciences,

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Reproductive Biology of the Brown Water Snake,

Nerodia taxispilota, in Central Georgia

Robert E. Herrington

Department of Biology,

Georgia Southwestern College,

Americus, Georgia 31709

ABSTRACT. — The brown watersnake, Nerodia taxispilota, is a large,

conspicuous, aquatic snake that occurs over much of the southeastern

Coastal Plain of the United States. At sexual maturity, females are

considerably larger in body length and mass than males. Males are

sexually mature at 2% years of age and approximately 58 cm SVL.

Females mature a year later at 85 to 90 cm SVL. Courtship occurs in

late April and early May, and may involve more than one male per

female. The young are born from late August to early September.

Litter size varied from 14 to 45 and was positively correlated with the

SVL of the female.

The natural history of many North American snakes remains

poorly known. Reasons for this include their relatively low population

densities, their secretive behaviors, and the seasonality of their activity

patterns. The brown water snake, Nerodia taxispilota (Holbrook),

occurs in the southeastern United States from Virginia to southern

Alabama (Conant 1975). The species is one of the largest members of

the genus and occurs in relatively high densities along many of the river-

swamps of central Georgia. These characteristics coupled with strong

arboreal basking tendencies make it an ideal subject for life history

studies. This report describes the growth and reproductive biology of N.

taxispilota in central Georgia.

MATERIALS AND METHODS

The study was conducted in two parts. From July 1976 through

July 1977, specimens (n = 59; 33 males, 26 females) were collected from

the Oconee, Ogeechee, and Flint river drainages. Snakes were sexed,

measured (snout-vent length = SVL, tail length = TL) to the nearest mm,

and weighed to the nearest 0.1 g; their reproductive tracts were examined

following dissection. Reproductive data recorded for males were the size

and wet weight of the testes (± 10 mg). A testis and ductus deferens

were fixed in a 10% formalin solution, embedded in paraffin, sectioned

at 6 to 8 (Lm), and stained with Harris’s hematoxylin and eosin (Luna

1968). Sections were examined microscopically (100-400X) for the

presence of spermatozoa and stage of spermatogenesis. Female re-

productive tracts were examined with a dissecting microscope; the

number and length of ovarian follicles, embryos, or both were recorded.

Brimleyana No. 15:103-1 10, January 1989

103

104

Robert E. Herrington

Between March 1977 and May 1981, I monitored a separate

population of N. taxispilota by mark-recapture techniques. The study

area was along a 900-m section of Commissioner Creek (Oconee River

tributary), 0.5 km N of Toomsboro, Wilkinson County, Georgia. At this

location, the creek is approximately 20 m wide and has a maximum

depth of 3 m. Numerous shallow oxbows and water-filled depressions

occur adjacent to the study area. The predominant woody vegetation

bordering the creek consists of tupelo ( Nyssa aquatica ), sweet gum

( Liquidambar styraciflua), bald cypress ( Taxodium distichum), and

alder (Alnus serrulatd).

I visited this site approximately four times per month from March

through October during 1977 and 1978. The frequency of visits was

reduced to monthly during 1979 and continued at this rate through May

1981. Using a canoe, I attempted to capture each specimen sighted.

Once captured, snakes were sexed, measured (SVL and TL), and individually

marked by clipping subcaudal scales (Blanchard and Finster 1933).

Adults were examined for evidence of recent courtship activities by

swabbing the interior of the cloaca (Fukada 1959). I palpated adult

females for enlarged ovarian follicles or embryos and checked for the

presence of cloacal plugs (Devine 1975). Most snakes were released near

the point of capture within 24 hours.

Eleven adult females captured outside the study area were

maintained in captivity for up to 6 months, and five of these produced

litters. Newborn snakes were measured and weighed as described above,

and released in the study area. Growth rates were determined from

recaptures and were calculated on the basis of an 8-month (240-day)

annual growth period, assuming no growth occurred during hibernation.

Where statistical treatment of data is provided in the text, the mean

value is followed by±l standard deviation. Statistical comparisons were

made using the student’s t test (Steel and Torrie 1980).

RESULTS AND DISCUSSION

Body size. The ratio of TL/ total length was significantly greater in

males (x = 0.257 ± 0.002, n = 50) than in females (x = 0.235 ± 0.001, n =

59, t = 6.2, p < 0.01). However, at maturity females are longer (SVL and

total length) and heavier than males (Fig. 1). I calculated allometric

equations separately for each sex in the form of y = ax*5, where y = body

weight (kg), x = SVL (m), and b = a derived constant. These equations

were 0.47X3-12 and 1.47X2-89, for males and females, respectively. These

compare favorably to the equation y = 0.50X313derived by Kaufman

and Gibbons (1975) for combined sexes of N. taxispilota from western

South Carolina. Semlitsch and Gibbons (1982) suggest that there is

strong selection in females for increased body size to allow production

Reproduction in the Brown Water Snake

105

SVL (cm)

Fig. 1. Relationship between SVL and total body weight for male (solid circles)

and female (x) Nerodia taxispilota from central Georgia.

of larger clutches, but selection for increased body size in males is weak

or absent. My data are consistent with this hypothesis, but other

hypotheses such as partitioning of food resources by prey size are

possible. The largest male and female examined from central Georgia

were 86 and 124 cm SVL, respectively, and six of 74 females (8.7%)

exceeded 100 cm SVL.

Growth. The age/ size-class structure of the mark-recapture

population was determined by plotting the SVL of all specimens collected

during April and May as a histogram (Fig. 2A) and transforming these

values onto probability paper as described by Harding (1949). Because

only mark-recapture data can positively age individual specimens, Figure

2B indicates the most likely age/ size-class assignments based on the

probability plot.

Five litters of N. taxispilota, totaling 83 individuals, were born in

captivity. Newborn snakes had a mean SVL of 23.3 ±0.1 cm and

weighed 10.9 ±0.1 g. Under natural conditions little growth occurred

before hibernation, and the following spring this group emerged from

hibernation with SVLs from 23 to 26 cm. One newborn snake marked

on 14 September 1976 had grown only 0.4 cm (SVL) when recaptured

on 16 March 1977.

106

Robert E. Herrington

Snakes in their first full season of activity were better represented

in spring samples because they became progressively more difficult to

locate as vegetation density increased from spring to summer. However,

during 1977, specimens from 23 to 33 cm SVL were collected between

March and August and are considered to represent first-year individuals.

Growth rates from recaptures (n = 4) averaged 9.6 ± 4.1 cm per season

(range 3.8 to 13.6 cm).

An additional cohort was apparent in the spring with SVLs ranging

from 39 to 53 cm. These snakes were approximately 19 months old and

beginning their second full season of activity. Growth appears to be

rapid during this period, but no recaptures were recorded for this size

group.

Specimens in their third full season of activity emerged with a

modal SVL of 62 cm. Recaptured males (n = 2) grew at a rate of 4.0 to

19.1 cm SVL per season (mean = 13.0 cm). Two females in this size class

were recaptured; one had increased only 0.2 cm after 6 weeks, whereas

the second had grown 3.8 cm SVL in 7 weeks (21.7 cm per season).

Males in the next larger size class (n = 6) increased in SVL from 2

to 4 cm per season (mean = 3.2 ± 1.8 cm), while females in the same

cohort (n = 3) maintained growth rates that averaged 12.2 ± 2.8 cm per

season. A reduction in female growth rates with the attainment of

sexual maturity is suggested by data on recaptures. Growth of three

measured in their fourth season of activity ranged from 8.8 to 13.6 cm

SVL per season (mean = 1 1.0 ± 2.4 cm). In the next larger size class (n =

4) growth was reduced to 3.2 to 6.4 cm SVL per season (mean = 5.7 + 2.0

cm).

Reproduction. In central Georgia, males are sexually mature when

they emerge from their third period of hibernation and are approximately

58 cm SVL. All males in this or larger size classes (n = 13) had

spermatozoa present in the ductus deferens irrespective of the month in

which they were collected. Spermatozoa were much more numerous in

late fall and spring than at other times of the year. This agrees with

observations of Mitchell and Zug (1984) for N. taxispilota in Virginia.

Two slightly smaller males (53 and 56 cm SVL) collected during

September had traces of sperm in their ductus deferens. Thus, size at

maturity of males from central Georgia is comparable to the 50.3-cm

size at maturity in Virginia (White et al. 1982).

Females are sexually mature between 85 and 90 cm SVL. The

largest immature female (92.5 cm SVL) was collected during August,

and the smallest mature female (86.0 cm SVL) gave birth in captivity.

The minimum size at maturity observed during this study (86 cm SVL)

is larger than that reported by White et al. (1982) for this species in

Virginia (72.5 cm SVL). During late summer a total of 12 females (70-80

NO. OF ANIMALS EST. AGE (MONTHS)

Reproduction in the Brown Water Snake

107

Fig. 2. Size-class structure for spring (April-May) Nerodia taxispilota

population: (A) Histogram of SVLs: males (solid bars) and juveniles (open bars)

above the line and females (solid bars) below the line. (B) Probable age-class

assignments for the same population, redrawn from a cumulative percent plot

on probability graph paper: juveniles = triangles, males = squares, and females =

open circles.

cm SVL) were either dissected or palpated in the field, and none

contained enlarged follicles or embryos. The larger size at maturity for

Georgia females may be the result of differences in growth rate or in the

age at maturity.

Macroscopic changes in the ovaries of N. taxispilota from central

Georgia are similar to those described by Betz (1963) for N. rhombifera,

by Bauman and Metter (1977) and Aldridge (1982) for N. sipedon, and

by White et al. (1982) for other populations of N. taxispilota. Ovarian

follicles gradually enlarge with increasing SVL in immature snakes.

Females in the second full season of activity had ovarian follicles less

than 3.0 mm in length. The following year follicles had increased to

between 5 and 9 mm in length. Females are sexually mature at the

beginning of their fourth full season of activity (approximately 43

months old). Vitellogenesis occurs rapidly, with follicle lengths often

exceeding 20 mm prior to ovulation. Ovulation apparently occurs from

108

Robert E. Herrington

late May to early June, but no adult females were available for

dissection during this period. However, ovulation did not begin earlier

than late May and was completed by early July. White et al. (1982)

found that ovulation occurred during late June in Virginia.

No observations have been reported concerning courtship in N.

taxisiplota. Between 24 April and 5 May 19797, four instances of

courtship activities were observed. In three of these, single females were

observed in close association with two to three males. All groups were

located on tree limbs that were 33 to 63 cm above the water surface.

Three of the four females were collected, and each had abundant sperm

in cloacal smears. Two of these females and three additional specimens

collected during May had in their cloacas gelatinous, semirigid structures

resembling sperm plugs (Devine 1975). These observations suggest that

multiple male courtship may be common in N. taxispilota. Multiple

male courtship was reported for N. sipedon by Mushinsky (1979).

Mating activities were centered between late April and early May.

No sperm were detected in cloacal smears of 18 mature females before

mid-April; however, between mid-April and mid-May, eight of ten

mature females contained sperm in cloacal smears. Six adult females

collected prior to 15 April and maintained in captivity failed to produce

young that year, presumably because mating had not occurred prior to

capture. No evidence of fall matings was observed in cloacal smears

taken from 29 adult females sampled during September and October.

Parturition occurred in captivity between 27 August and 9

September. Litter size was 16, 14, 26, 13, and 14, for females that were

86, 88, 94, 95, and 101 cm SVL, respectively. The earliest appearance of

a newborn specimen in the field was on 21 August 1977. Fecundity,

including the number of fetuses as well as the number of enlarged

ovarian follicies, ranged from 14 to 45 and was positively correlated

with the female SVL (r = 0.77), with SVL explaining 59% of the

variation. This agrees with the corresponding correlation coefficient of r

= 0.78 reported by Semlitsch and Gibbons (1978) for N. taxispilota from

western South Carolina.

Females apparently produce litters annually once sexual maturity is

reached. All females that were at least 92 cm SVL had either enlarged

ovarian follicles or embryos present, and at least four females were

known to have been gravid in consecutive years.

ACKNOWLEDGMENTS. — Part of this work was extracted from

a Master’s thesis submitted to the Department of Biological and

Environmental Sciences, Georgia College. Appreciation is extended to

J. D. Batson, E. R. Barman, and D. Staszak for guidance given during

Reproduction in the Brown Water Snake

109

the study. O. F. Anderson, C. Duke, D. Tucker, and R. L. Herrington

aided in field work. Earlier drafts of this manuscript were substantially

improved by comments from R. Mount, R. Wallace, J. Beneski, B.

Miller, and G. Zug. Special appreciation is extended to my wife Vicki

for preparing the histological material used in this study.

LITERATURE CITED

Aldridge, R. D. 1982. The ovarian cycle of the watersnake Nerodia sipedon,

and effects of hypophysectomy and gonadotrophin administration. Herpe-

tologica 38:71-79.

Bauman, M. A., and D. E. Metter. 1977. Reproductive cycle of the northern

watersnake Natrix s. sipedon (Reptilia, Serpentes, Colubridae). J. Herpetol.

11:51-59.

Betz, T. W. 1963. The gross ovarian morphology of the diamond-backed water

snake, Natrix rhombifera, during the reproductive cycle. Copeia

1963:692-697.

Blanchard, F. N., and E. B. Finster. 1933. A method of marking living snakes

for future recognition, with a discussion of some problems and results.

Ecology 14:334-347.

Conant, R. 1975. A Field Guide to the Reptiles and Amphibians of Eastern

and Central North America. 2nd ed. Houghton Mifflin Co., Boston.

Devine, M. C. 1975. Copulatory plugs in snakes: enforced chastity. Science

187:844-845.

Fukada, H. 1959. A method for detecting copulated female snakes.

Herpetologica 15:181-182.

Harding, J. P. 1949. The use of probability paper for the graphical analysis of

polymodal frequency distributions. J. Mar. Biol. Assn. UK 28:141-153.

Kaufman, G. A., and J. W. Gibbons. 1975. Weight-length relationships in

thirteen species of snakes in the southeastern United States. Herpetologica

31:31-37.

Luna, L. G. 1968. Manual of Histologic Staining Methods of the Armed

Forces Institute of Pathology. 3rd ed. McGraw Hill Book Co., New York.

Mitchell, J. C., and G. R. Zug. 1984. Spermatogenic cycle of Nerodia

taxispilota (Serpentes: Colubridae) in southcentral Virginia. Herpetologica

40:200-204.

Mushinsky, H. R. 1979. Mating behavior of the common water snake, Nerodia

s. sipedon, in eastern Pennsylvania. J. Herpetol. 13:127-129.

Semlitsch, R. D., and J. W. Gibbons. 1978. Reproductive allocation in the

brown water snake, Natrix taxispilota. Copeia 1978:721-723.

Semlitsch, R. D., and J. W. Gibbons. 1982. Body size dimorphism and sexual

selection in two species of water snakes. Copeia 1982:974-976.

Steel, R. G. D., and J. H. Torrie. 1980. Principles and Procedures of

Statistics — A Biometrical Approach. 2nd ed. McGraw Hill Book Co., New

York.

110

Robert E. Herrington

White, D. R., J. C. Mitchell, and W. S. Woolcott. 1982. Reproductive cycle

and embryonic development of Nerodia taxispilota (Serpentes: Colubridae)

at the northeastern edge of its range. Copeia 1982:646-652.

Accepted 17 July 1987

Movements of Land-based Birds

Off the Carolina Coast

David S. Lee and Kenneth O. Horner

North Carolina State Museum of Natural Sciences,

P.O. Box 27647, Raleigh, North Carolina 2761 1

ABSTRACT. — Although the occurrence of land-based birds at sea

during migration periods is well known, relatively little information is

available on the species composition of the flocks detected by radar.

This paper lists 96 species documented from the offshore waters of

North and South Carolina, offers evidence for offshore movements by

groups of birds other than nocturnal migrants, and suggests temporal

changes in flock composition.

It is well known that land-based birds regularly occur at sea during

migration periods, when flocks of birds or individual birds deliberately

or accidentally take oceanic routes. Various authors have demonstrated

(through personal observation and radar studies) that offshore movement

of nocturnal migrants occurs in the North Atlantic on a regular basis

(Scholander 1955; Drury and Keith 1962; Williams et al. 1977: Davis

1978; McClintock et al. 1978; Richardson 1978, 1980; Larkin et al. 1979;

Cherry et al. 1985; Williams 1985). Nisbet (1970) and others have

proposed that a long, over-water flight crossing the Atlantic directly to

South America is a normal and deliberate route for some species (e.g.,

Blackpoll Warbler). Although fall movements over the western North

Atlantic Ocean have been documented, there is relatively little in-

formation on the species composition of the migrant clouds detected.

Furthermore, individual records of land-based birds found at sea

generally have gone unrecorded and unreported. Here we identify some

of the offshore migrants, present evidence for offshore movements by

groups of birds other than nocturnal migrants, and suggest temporal

changes in flock composition.

Our sightings of land-based birds were for the most part recorded

incidental to studies of seabirds during the 10-year period from 1975 to

1986. They were made primarily 10 to 55 miles (16-88 km) off the North

Carolina coast between 30° and 35°N. Sightings were made without

optical aids, but identifications were often made with binoculars. It

should be emphasized that many land-based birds observed at sea could

not be specifically identified because of distance, boat movement,

atypical flight postures resulting from strong winds, and other adverse

conditions. We estimate that 55% of the land-based birds seen in flight

disappeared from the field of vision before they could be identified.

Brimleyana No. 15:1 1 1-121, January 1989

111

112

David S. Lee and Kenneth O. Horner

Therefore, we are unable to present data indicative of the actual

numbers of birds encountered or of the species’ relative abundance, and

comparison of tallies between trips is meaningless.

In the species list given below, observations are from off coastal

North Carolina unless otherwise indicated (SC = South Carolina). Most

records are from Lee’s 130 survey trips conducted in charter fishing

boats off Oregon Inlet, Dare County, N.C., but additional published

and unpublished records are included. Reports with incomplete data are

mentioned only when no other evidence is available to document the

species’ occurrence off the Carolina coast. Sightings for each species are

arranged by day and month. When more than a single bird was seen,

the number observed is presented in parentheses. Precise latitude and

longitude as determined by LORAN instruments are available for most

of Lee’s records, but these data have been omitted in the interest of

brevity. For selected species, distances from shore have been provided.

LAND-BASED BIRDS SEEN OFFSHORE

PODICIPEDIFORMES

Pied-billed Grebe ( Podilymbus podiceps): 19 July 1977 (NCSM

6167), 39 miles ESE Oregon Inlet; 10 August 1977, 30 miles E

Oregon Inlet.

Pelecaniformes

Brown Pelican ( Pelicanus occidentalis ): Summer 1976. Foraging

flocks approximately 50 miles SE Beaufort.

Double-crested Cormorant ( Phalacrocorax auritus ): 3 November

1979 (100 fathom contour).

ClCONIIFORMES

Great Blue Heron ( Ardea herodias ): 15 October 1979, 16

October 1979, 5 November 1979, 5 December 1985.

Snowy Egret ( Egretta thula ): 4 October 1980 (9), Chat 45:54.

Little Buie Heron ( Egretta caerulea ): 4 October 1980, Chat 45:54.

Cattle Egret ( Bubulcus ibis): 27 April 1968, Chat 33:102; 9

September 1979 (40); 7 October 1975 (SC).

Anseriformes

Northern Pintail ( Anas acuta): Fall 1983 or 1984 (flock).

Blue-winged Teal (Anas discors): 27 August 1979, 1 September

1979 (4), 8 September 1979.

Lesser Scaup ( Aythya affinis): 5 November 1979.

White-winged Scoter (Melanitta fusca): 6 September 1981 (6),

Chat 46:47.

Red-breasted Merganser (Mergus senator): 16 March 1984 (12+), 28

April 1983 (4), 19 May 1982 (3).

Land-based Birds Off Carolina Coast

113

Falconiformes

Osprey ( Pandion haliaetus ): 21 June 1985 (4), 30 September 1979.

Northern Harrier ( Circus cyaneus ): 4 October 1980, Chat 45:54.

Sharp-shinned Hawk ( Accipiter striatus ): 29 September 1979, 30

September 1979, 1 October 1979 (2), 12 October 1975.

American Kestrel ( Falco sparverius ): Fall, Kerlinger et al. 1983.

Merlin ( Falco columbarius ): 10 September 1979, 30 September

1979.

Peregrine Falcon ( Falco peregrinus ): 28 September 1979; 30

September 1979; 2 October 1979; 4 October 1980, Chat 45:54;

7 October 1985 (SC); 20 October 1982.

Gruiformes

Clapper Rail ( Rallus longirostris ): 1 September 1979.

King Rail ( Rallus elegans ): 9 September 1979, ca. 50 miles E Core

Banks.

Purple Gallinule ( Porphyrula martinica ): 19 August 1978.

American Coot ( Fulica americana ): 12 October 1975, 16 October

1979.

Charadriiformes

Black-bellied Plover ( Pluvialis squatarola ): 27 August 1979 (5).

Lesser Golden-Plover ( Pluvialis dominica ): 8 September 1979 (7),

15 October 1979.

Semipalmated Plover (Charadrius semipalmatus): 1 September 1979,

1 September 1984 (2).

Greater Yellowlegs ( Tringa melanoleuca ): 31 July 1984 (flock), 24

August 1979 (9).

Solitary Sandpiper ( Tringa solitaria ): 27 August 1979 (SC), 28

August 1979 (2), 1 September 1979 (2).

Willet (Catoptrophorus semipalmatus)'. 16 August 1984 (flock).

Spotted Sandpiper (Actitis macularia ): 1 September 1985 (SC).

Whimbrel (Numenius phaeopus): 19 May 1982.

Ruddy Turnstone ( Arenaria interpres ): 16 May 1979, 8 September

1979.

Red Knot ( Calidris canutus ): August 1984 (flock).

Sanderling ( Calidris alba): 22 May 1980, 7 September 1979 (4).

Semipalmated Sandpiper ( Calidris pusilla ): 1 September 1979, 1

September 1985 (SC).

Least Sandpiper ( Calidris minutilla ): 8 May 1980, 24 August 1979,

1 September 1979.

Stilt Sandpiper ( Calidris himantopus ): 24 August 1979.

Short-billed Dowitcher ( Limnodromus griseus ): 28 August 1979; 4

October 1980, Chat 45:54. [Dowitcher sp.: 24 August 1979

(10)].

114

David S. Lee and Kenneth O. Horner

COLUMBIFORMES

Mourning Dove (Zenaida macroura ): 2 October 1979, (2); 4

October 1980, Chat 45:54; 8 October 1978; 15 October 1979;

16 October 1979 (2); 4 November 1979 (2); 5 November 1979.

Apodiformes

Chimney Swift ( Chaetura pelagica ): 29 May 1980.

Cor AC 1 1 FORMES

Belted Kingfisher (Ceryle alcyon): 12 July 1980, 1 September 1979,

8 September 1979, 23 September 1985 (SC).

PlCIFORMES

Red-headed Woodpecker ( Melanerpes erythrocephalus ): 28 April

1983, 32 miles ESE Oregon Inlet.

Downy Woodpecker ( Picoides pubescens ): 6 August 1981, 30 miles

ESE Oregon Inlet.

Northern Flicker ( Colaptes auratus ): 2 October 1979, 12 October

1975, 15 October 1979.

Passeriformes

Acadian Flycatcher ( Epidonax virescens ): 31 August 1977, 1

September 1979.

Least Flycatcher ( Epidonax minimus ): September, Scholander 1955.

“Tropical” Kingbird (Tyr annus [melancholicus or couchii ]): 1

September 1985 (SC).

Tree Swallow ( Tachycineta bicolor ): 19 April 1980, 1 September

1979, 12 October 1975.

Bank Swallow ( Riparia riparia ) 30 August 1985.

Cliff Swallow ( Hirundo pyrrhonota ) 24 August 1979.

Barn Swallow ( Hirundo rustica ): 29 April 1980, 8 May 1980, 10

May 1978, 16 May 1979, 18 May 1977, 20 May 1977 (5), 22

May 1980, 5 August 1981, 7 August 1984, 9 August 1983, 9

August 1984, 10 August 1984 (2), 23 August 1979, 24 August

1979 (5), 26 August 1975, 27 August 1979 (3, SC), 29 August

1985.

American Crow ( Corvus brachyrhynchos ): 1 November 1979.

Red-breasted Nuthatch ( Sitta canadensis ): 9 September 1981 (2),

Chat 46:74.

Brown Creeper ( Certhia americana ): 15 October 1979.

House Wren ( Troglodytes aedon ): 8 October 1978, 12 October 1985

(3), 15 October 1979, 16 October 1979.

Golden-crowned Kinglet ( Regulus satrapa ): 12 October 1985, 4

November 1979.

Ruby-crowned Kinglet ( Regulus calendula ): 12 October 1985, 15

October 1979.

Land-based Birds Off Carolina Coast

115

Gray Catbird ( Dumetella carolinensis ): 7 Ocbober 1985 (SC), 15

October 1979 (2), 16 October 1979.

Cedar Waxwing ( Bombycilla cedrorum ): 15 October 1979 (4).

European Starling ( Sturnus vulgaris): 16 March 1984, 1 November

1979 (2), 4 November 1979 (2).

Orange-crowned Warbler ( Vermivora celata ): 1 September 1985

(SC).

Nashville Warbler ( Vermivora ruficapilla ): 24 September 1985.

Yellow Warbler ( Dendroica petechia ): 1 September 1979, 9 October

1987.

Cape May Warbler ( Dendroica tigrina ): 1 September 1979 (6); 7

September 1979; 8 September 1979 (8); 10 September 1981 (4),

Chat 46:74; 27 September 1979 (2); 15 October 1979; 16

October 1979 (2). The six birds on 1 September 1979 represent

early migrants.

Black-throated Blue Warbler (Dendroica caerulescens): 1 September

1979; 10 September 1981 (2), Chat 46:74; 9 October 1987.

September first is an extremely early migrant record.

Yellow-rumped Warbler ( Dendroica coronata ): 12 October 1985

(2), 15 October 1979 (8), 16 October 1979 (5, two different

boats), 25 October 1985 (2), 2 November 1979.

Black-throated Green Warbler ( Dendroica virens ): 10 September

1981 (4), Chat 46:74; 7 October 1985 (SC).

Yellow-throated Warbler ( Dendroica dominica ): 7 October 1985

(SC).

Prairie Warbler ( Dendroica discolor ): 2 September 1979.

Palm Warbler ( Dendroica palmarum ): 24 September 1985; 27

September 1979 (3); 28 September 1979; 2 October 1979; 4

October 1980, Chat 45:54; 7 October 1985 (5, SC); 15 October

1979 (3); 16 October 1979 (2).

Bay-breasted Warbler ( Dendroica castanea ): 9 October 1987, 15

October 1979, 16 October 1979 (2).

Blackpoll Warbler ( Dendroica striata ): 15 October 1979 (6), 16

October 1979 (2).

Black-and-white Warbler ( Mniotilta varia ): October, Scholander

1955.

American Redstart ( Setophaga ruticilla): 8 September 1979; 10

September 1979; 10 September 1981, Chat 46:47; 29 September

1979; 4 October 1980, Chat 45:54; 9 October 1987; 15 October

1979. Last date represents rather late migration record for

species.

Prothonotary Warbler ( Protonotaria citrea ): 1 September 1979, 7

September 1979.

116

David S. Lee and Kenneth O. Horner

Northern Waterthrush ( Seiurus noveboracensis ): 9 August 1984, 30

August 1985. 1 September 1979, 10 September 1979.

Kentucky Warbler (Oporornis formosus): 1 September 1985 (SC).

Mourning Warbler (Oporornis Philadelphia): 1 September 1979.

Common Yellowthroat ( Geothlypis trichas ): 1 September 1979, 1

September 1984. 7 September 1979, 8 September 1979, 7

October 1985 (2, SC), 15 October 1979.

Yellow-breasted Chat ( Icteria virens ): 7 September 1979.

Indigo Bunting ( Passerina cyanea ): 8 May 1980, 18 May 1977

(NCSM 6145).

Dickcissel ( Spiza americana ): October, Scholander 1955.

Rufous-sided Towhee ( Pipilo erythrophthalmus ): 23 October 1967,

Jensen and Livingstone 1969.

Chipping Sparrow ( Spizella passerina)-. 15 October 1979.

Tree Sparrow ( Spizella arborea ): 26 April 1985 (SC), Chat 50:

55-56.

Field Sparrow ( Spizella pusilla ): 15 October, 1979.

Savannah Sparrow {Passer cuius sandwichensis ): October, Scholander

1955.

Song Sparrow {Melospiza melodia ): 15 October 1979 (5), 4

November 1979 (2).

Swamp Sparrow (Melospiza georgiana): 12 October 1985.

White-throated Sparrow ( Zonotrichia albicollis ): 12 October 1985

(3), 15 October 1979 (3).

White-crowned Sparrow ( Zonotrichia leucophrys ): 12 October 1985,

15 October 1979 (3), 16 October 1979.

Dark-eyed Junco ( Junco hyemalis ): 12 October 1985 (5), 15 October

1979 (3), 4 November 1979 (2).

Lapland Longspur ( Calearius lapponicus ): 10 May 1978. Late date

for species in North Carolina; previously 9 May 1981 (Chat

45:1 10) was considered an extremely late date.

Bobolink ( Dolichonyx oryzivorus ): 9 September 1979, 27 September

1979.

Eastern Meadowlark ( Sturnella magna ): 23 October 1967, Jensen

and Livingston 1969.

Boat-tailed Grackle ( Quiscalus major): 2 June 1984.

Orchard Oriole ( Icterus spurius): 23 August 1979.

Northern Oriole ( Icterus parisorum): 1 September 1979; 8 September

1979; 9 September 1979; 10 September 1981, Chat 46:47; 23

September 1985 (SC); 1 October 1979.

DISCUSSION

Approximately 400 birds belonging to 12 orders and 97 species

(including 52 passerine species) were recorded. The absence from our

Land-based Birds Off Carolina Coast

117

records of several families of common East Coast migrants (e.g.,

thrushes and vireos) should be noted. Most of the birds seen were fall

migrants (90%), and more than 60% of all activity was recorded in

September and October. Only 1 1 species were recorded during the

spring migration period. Figure 1 illustrates the documented monthly

occurrence of land-based birds off the Carolinas.

Land-based migrants off the Carolina coast in August and early

September are predominantly non-passerine species, primarily shorebirds.

These non-passerine species dominate August migrants by 2: 1 over

passerine species. Passerine species become more numerous during

September. Most of these early records were of neotropical migrants

(Fig. 2). Furthermore, the 20 passerine species identified over the ocean

between 10 August and 10 September were largely different in geographic

origin from the 17 species seen between 6 October and 4 November.

Among the early passerine migrants were four parulids (Prairie, Protho-

notary, and Kentucky Warblers and Yellow-breasted Chat) that originate

from breeding grounds restricted primarily to southeastern North

America (i.e., the United States east of the Mississippi River). Their

median breeding range is about 34° N. These species, which apparently

fly from the northern part of their range southward over the Atlantic

Ocean, have been recorded as regular, though numerically few, migrants

to Bermuda (Drury and Keith 1962). Other early passerine migrants

that have a similar breeding range are the Acadian Flycatcher and the

Orchard Oriole. In addition, some of the other early-migrant parulids

(i.e.. Black-throated Blue, Black-throated Green, and Mourning War-

blers) are birds that occupy intermediate-latitude breeding ranges, and

individuals encountered in all likelihood originated from northern and

central Atlantic States.

These early-season species contrast with late-season parulid migrants

(i.e., Yellow-rumped, Bay-breasted, and Blackpoll Warblers) from

northeastern North America, which occur as late as 15 October or later

(Fig. 2). Each of these species comes from intermediate- to northern-

latitude breeding grounds (median breeding range ca. 49°N). Also

among the late-season encounters are short-distance migrants: Brown

Creeper, House Wren, Gray Catbird, Golden-crowned and Ruby-

crowned Kinglets, and Chipping, Field, Song, and White-throated

Sparrows. Only three species were recorded during both periods:

American Redstart and Common Yellowthroat, which have broad-

latitude breeding ranges, and the Cape May Warbler, which breeds at

the intermediate latitudes. Dates of encounters reported here are general-

ly within the periods reported for the respective species by Sykes (1986)

in his study of autumn land-bird migration along the Outer Banks of

North Carolina.

Many of the birds encountered, particularly small passerines, were

often near the point of exhaustion, and it was not uncommon to have

118

David S. Lee and Kenneth O. Horner

JAN FEB MAR APR MAY JUN JUL AUG SEPT OCT NOV DEC

MONTHS

Fig. 1. Monthly occurrence of passerine (dark bars) and land-based non-

passerine (light bars) birds off the Carolina coast.

them land on the boat. McClintock et al. (1978) found most of the

individuals landing on ships to be emaciated immatures. On several

occasions we watched warblers fall into the ocean or get caught in

waves. Gulls and jaegers have been seen eating exhausted song birds,

and the first Long-tailed jaeger ( Stercorarius longicaudus ) collected in

North Carolina had an Acadian Flycatcher in its stomach on 31 August

1977. Once we watched a Laughing Gull ( Larus atricilla ) capture an

exhausted Common Yellowthroat on the wing. It is possible that

weakened migrating song birds are an important food item for jaegers

at sea in fall. Although the mortality rate for offshore land-based birds

may be high, scavengers and predators certainly consume most of the

evidence quickly.

With the exception of the swallows and a few accidentals, we

consider the records of passerines reported here to represent fallout

from the offshore migrant clouds.

Our records were obtained on fair-weather days, weather conditions

under which radar has often revealed large numbers of birds passing

overhead. Radar studies indicate that migrants gain altitude as they

move out to sea, normally flying at heights that would keep them out of

view. Most passerines reported here were seen as they approached the

boat; their small size prohibited our seeing distant or high-flying birds.

Therefore, it is not possible to interpret the magnitude of offshore

migration based on chance encounters with what probably represent

middle latitude neotropical migrants

northern latitude neotropical migrants

short range migrants

Land-based Birds Off Carolina Coast

I I I I I I I I TT FT

Tf G >0

lo m rr

SI

i n n n i ii i i ii ii i

Tf O -0

snvnaiAiaNi jo H39i/vnN

Fig. 2. Occurrence of migrant passerine birds off the Carolina coast based on origin and destination of species.

120

David S. Lee and Kenneth O. Horner

exhausted individuals or those that were unable to keep up with the

migrating flocks.

Some of the hawks were observed at heights that made it difficult

to see them without the aid of binoculars. Because of this, we believe

that the offshore hawk movement is likely to be much more regular

than is suggested here. Kerlinger et al. (1983) discussed raptor migration

off the northeastern United States.

The offshore presence of sedentary species (e.g.. Downy Wood-

pecker) is difficult to interpret; certainly, some of these birds should be

regarded simply as accidentals. The occurence of a “Tropical” Kingbird

off the South Carolina coast on 1 September 1985 (Koeble, pers.

comm.) is noteworthy and at present inexplicable. A tropical storm that

was in the Gulf of Mexico at that time may in some way account for the

bird’s northward displacement. The Tropical Kingbird specimen taken

at Scarborough, Maine, early in the twentieth century is of the migra-

tory race Tyr annus melancholicus chloronotus (A.O.U. 1957).

Offshore movements of migrants are not limited to birds. Far

offshore we have seen butterflies [sulphurs (Pieridae), several species;

monarchs (Danaidae), Danus plexippus ], dragonflies [darners (Aesh-

nidae, Anax); skimmers ( Libellula sp.); the Globe Trotter, Pantala

flavescens , which is a cosmopolitan species], and bats. Red Bats,

Lasiurus borealis , were seen on 21 June 1985, 2 September 1984, 9

September 1979, and 4 November 1979.

ACKNOWLEDGMENTS. — We thank Wayne Irvin, Steve

Platania, and Mary Kay Clark for assistance during offshore trips and

Capt. Allen Foreman, Charles S. Manooch III, Richard Rowlett, Paul

DuMont, and Tim D. Koeble for sharing unpublished records of land-

based birds at sea. Rowlett’s records were particularly extensive and

useful. Janet M. Williams, Sidney A. Gauthreaux, Jr., and Robert

Dickerman reviewed a previous draft of the manuscript. Lee’s offshore

studies were financed in part by contract #92375-1 130-621-16, U.S. Fish

and Wildlife Service Laboratory, Slidell, Louisiana.

LITERATURE CITED

American Ornithologists’ Union. 1957. Check-list of North American Birds.

5th ed. Baltimore, Md.

Cherry, J. D., D. H. Doherty, and K. D. Powers. 1985. An offshore nocturnal

observation of migrating Blackpoll Warblers. Condor 87:548-549.

Davis, T. H. 1978. Pelagic birding trips to Cox’s Ledge from Montauk Point,

Long Island. Kingbird 28:131-149.

Drury, W. H., and J. A. Keith. 1962. Radar studies of songbird migration in

coastal New England. Ibis 104:449-489.

Land-based Birds Off Carolina Coast

121

Jensen, A. C., and R. Livingstone, Jr. 1969. Offshore records of land birds.

Kingbird 19:5-10.

Kerlinger, P., J. D. Cherry, and K. D. Powers. 1983. Records of migrant

hawks from the North Atlantic Ocean. Auk 100:488-490.

Larkin, R. P., D. R. Griffin, J. R. Torre-Bueno, and J. Teal. 1979. Radar

observations of bird migration over the western North Atlantic Ocean.

Behav. Ecol. Sociobiol. 4:225-264.

McClintock, C. P., T. C. Williams, and J. M. Teal. 1978. Autumnal bird

migration observed from ships in the western North Atlantic Ocean. Bird-

Banding 49:262-277.

Nisbet, I. C. T. 1970. Autumn migration of the Blackpoll Warbler: evidence

for long flight provided by regional survey. Bird-Banding 41:207-209.

Richardson, W. J. 1978. Reorientation of nocturnal landbird migrants over the

Atlantic Ocean near Nova Scotia in autumn. Auk 95:717-732.

Richardson, W. J. 1980. Autumn landbird migration over the western Atlantic

Ocean as evident from radar. Proc. XVII Int. Ornithol. Cong. ( 1 978):50 1 -

506.

Scholander, S. I. 1955. Landbirds over the western North Atlantic. Auk

72:225-239.

Sykes, Paul W., Jr. 1986. Autumn Land-bird Migration on the Barrier Islands

of Northeastern North Carolina. N.C. Biol. Survey, N.C. State Mus. Nat.

Sci., Raleigh.

Williams, T. C. 1985. Autumnal bird migration over the windward Caribbean

islands. Auk 102:163-167.

Williams, T. C., J. M. Williams, L. C. Ireland, and J. M. Teal. 1977.

Autumnal bird migration over the western North Atlantic Ocean. Amer.

Birds 31:251-261.

Accepted 27 August 1987

122

AUTUMN LAND-BIRD MIGRATION

ON THE BARRIER ISLANDS OF NORTHEASTERN

NORTH CAROLINA

by

Paul W. Sykes, Jr.

For three consecutive years Sykes investigated the autumn migration

of land birds in the Bodie Island and Pea Island area of coastal North

Carolina. During a 102-day period in 1965, he recorded 110,482

individual birds of 148 species. He was able to correlate major influxes

of migratory species with specific weather patterns. His data show

seasonal peaks of southward movement for the land-bird species that

pass along the North Carolina coast in large numbers. In addition,

Sykes recorded five species native to the western United States. Three of

these vagrants provided the first reports of Swainson’s Hawk, Sage

Thrasher, and Western Meadowlark for North Carolina.

1986 49 pages Softbound ISBN 0-917134-12-5

Price: $5 postpaid. North Carolina residents add 5% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: LAND-BIRD MIGRATION, N.C. State Museum of Natural

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

Distribution of the Southeastern Shrew,

Sorex longirostris Bachman, in Western Virginia

John F. Pagels

Department of Biology,

Virginia Commonwealth University, Richmond, Virginia 23284

AND

Charles O. Handley, Jr.

Division of Mammals,

U.S. National Museum of Natural History,

Smithsonian Institution, Washington, D. C. 20560

ABSTRACT. — A two-year study of the distribution and abundance

of shrews was conducted in Virginia. Pitfall traps (1508 in all) were

placed at 140 localities scattered around the state. Sorex longirostris

(73 specimens) was caught at 33 of the 107 localities within its potential

range (at 65% of Piedmont, 40% of Blue Ridge, 37% of Coastal Plain,

and 1 1% of Ridge and Valley localities). Other specimens were found

in museums and in bottles discarded along roadways. Altogether, 48 new

localities can be added to the compilation of Pagels et al. (1982) of

collecting sites of S. longirostris in Virginia. In the eastern lowlands S.

longirostris frequented all habitats sampled. In the mountains it was

caught mostly in fields and borders. Trapping results indicate that the

species reached the Valley of Virginia through several gaps in the Blue

Ridge, and that it reached southwestern Virginia through the Tennessee

Valley. Sorex longirostris was captured with five of the other eight

shrews known to occur in Virginia. It was not caught with S. cinereus,

and its distribution west of the Blue Ridge may be limited by the

presence of that species. The two exhibit contiguous allopatry, with S.

longirostris below 457 to 610 m (depending on latitude) and S. cinereus

above that elevation.

When we (Pagels et al. 1982) summarized the distribution of Sorex

longirostris in the Mid-Atlantic States, we questioned earlier reports of

a nearly statewide distribution in Virginia. We believed that this shrew

does in fact occur throughout the Coastal Plain and Piedmont lowlands

in Virginia (except on the Eastern Shore), but, although we reported it

west of the Blue Ridge in Page and Warren counties, we doubted that it

is widespread in the mountainous sections of the state. Specimens from

Giles County (Odum 1944) had been shown to have been misidentified

(French 1980a), and specimens from other montane counties,

Montgomery (Handley and Patton 1947) and Augusta (Bruce 1937),

Brimleyana No. 15:123-131, January 1989

123

124

John F. Pagels and Charles O. Handley, Jr.

apparently had been lost and could not be re-examined. Thus, the true

distribution of this shrew in western Virginia remained to be determined.

Following our earlier work on Sore: c longirostris, Pagels conducted

a more detailed field study of the distribution and ecology of shrews of

all species in Virginia. He collected S. longirostris at many localities

from which it had not been known and located previously collected

specimens that add significantly to our knowledge of the range of this

species in Virginia. Although we list and map all of the new localities, as

an update of our earlier work (Pagels et al. 1982), our present emphasis

is on the distribution of S. longirostris west of the Blue Ridge.

MATERIALS AND METHODS

We placed 1508 pitfall traps (16-oz. aluminum cans) at 140 localities,

mostly along highways, to form irregular transects in all five

physiographic regions of Virginia (Fig. 1). The cans, partly filled with

water early in the study but later with a formalin solution to help

preserve the specimens, were checked approximately bimonthly for 24

months between 1983 and 1985.

Four major cover types were sampled: old field, field-forest edge,

mixed forest, and hardwood forest. Within a given habitat, traps were

set irregularly in the best cover rather than at fixed intervals. Because

this study included all species of shrews, collecting effort sometimes was

directed toward a particular species. Thus, the intensity of sampling of

the various cover types was not the same in all regions.

Although we sampled 140 localities with pitfall traps, only 107 of

them (52 in the Coastal Plain and Piedmont, and 55 in the mountains)

were within the known range of Sorex longirostris. The other 33

localities were above 610 m, the upper limit of S. longirostris at this

latitude. The boreal habitats of these elevations are not likely to be

inhabited by this austral shrew. Thus, in Table 1 we listed these 33

localities as a separate division of the Ridge and Valley Province and

did not include them in the distributional totals.

COLLECTIONS

Altogether we found 115 specimens of Sorex longirostris at 48

localities in Virginia not listed in the compilation of Pagels et al. (1982).

The 48 new localities are plotted in Figure 1, and are grouped by

physiographic province in the following list of specimens (elevations in

parentheses were estimated by Pagels from topographic maps).

Because of our collecting technique, the specimens could only be

prepared as skulls or kept entire in 70% alcohol (skulls of some of those

Southeastern Shrew

125

Fig. 1. Collecting localities of Sorex l. longirostris in Virginia. Open circles

represent localities reported by Pagels et al. (1982). Solid circles represent

localities reported in this paper. Where localities are closely spaced, circles may

indicate more than one locality. Numerals indicate number of transects in each

county.

later were removed and cleaned). Most of the specimens are in the

Virginia Commonwealth University Mammal Collection: 73 taken in

pitfall traps, 8 found in bottles discarded along the roadsides (these

containers are effective deathtraps for shrews: Pagels and French

1987), and 5 contributed by Jack Cranford. The other 29 specimens are

in various museum collections: 1 in the U.S. National Museum of

Natural History (USNM), 2 in Lord Fairfax Community College

(LFCC), 9 in George Mason University (GMU), and 17 at Northern

Virginia Community College, Annandale Campus (NVCC-A).

COASTAL PLAIN. Essex Co.: 3.2 mi. NW Center Cross, 38 m, 1;

0.8 mi. NW Loretto, 31 m, 3; 2.4 mi. NW Loretto, 15 m, 4. Fairfax Co.:

Gunston Manor, on Mason Neck, 6.25 mi. SE Lorton, (9 m), 1 (GMU);

Isle of Wight Co.: 3.4 mi. SE Windsor, 23 m, 1. King George Co.: 0.5

mi. E Owens, 9 m, 6; 2.5 mi. S and 1 mi. E. Owens, 17 m, 2; 1.3 mi. N

Port Conway, 20 m, 4. Mathews Co.: 2.2 mi. NE Hudgins, 5 m, 3.

Prince George Co.: 1.2 mi. NE Disputanta, 23 m, 1; 2 mi. SSE

Hopewell, 37 m, 1. Prince William Co.: Woodbridge, Lake Ridge, (46

126

John F. Pagels and Charles O. Handley, Jr.

m), 12 (NVCC-A). Richmond Co.: 4 mi. S Warsaw, 5 m, 2; 5 mi. S

Warsaw, 5 m, 1. Southampton Co.: 8 mi. W Capron, 31 m, 3; 5.7 mi. W

Courtland, 21 m, 1; 5 mi. NE Sebrell, 24 m, 1. Sussex Co.: Warwick

Swamp, 1.4 mi. SE Sussex-Prince George Co. line, 23 m, 1.

Westmoreland Co.: 2.2 mi. SSW Colonial Beach, 6 m, 1; 1 mi. W Lerty,

47 m, 1. Total 50.

PIEDMONT. Amherst Co.: 0.1 mi. NW Amherst, 213 m, 2; Forks

of Buffalo, 280 m, 1. Campbell Co.: Brookneal, 125 m, 3. Charlotte Co.:

2.2 mi. S and 5 mi. W Charlotte, 146 m, 1; Cub Creek, 1 mi. W Phenix,

110 m, 1; 1.3 mi. W Phenix, 116 m, 2; Roanoke Creek, 6 mi. W

Keysville, 91 m, 6. Fairfax Co.: Dulles Airport, (91 m), 5 (GMU), 5

(NVCC-A); Fairfax (GMU Campus), 137 m, 1 (GMU); Fort Belvoir,

North Post, (31 m), 1 (GMU). Fauquier Co.: Conde (Rte. 737), 158 m, 1

(LFCC); 4 mi. NNE Marshall, 168 m, 6; 9.4 mi. NNE Marshall, 168 m,

2. Greene Co.: 0.6 mi. W Stanardsville, 183 m, 1. Pittsylvania Co.: 1 mi.

W Mount Airy, 165 m, 4. Prince William Co.: Haymarket, (101 m), 1

(GMU). Total 43.

BLUE RIDGE. Greene Co.: 4.7 mi. W Stanardsville, 411 m,

Rappahannock Co.: Chester Gap, 412 m, 4. Warren Co.: 1 mi. S Front

Royal, National Zoological Park Conservation Center, (305 m), 1

(LFCC); Linden, along branch of Manassas Run, 274 m, 2; 2.4 mi. W

Linden, beside Manassas Run, 213 m, 1. Total 9.

RIDGE AND VALLEY. Lee Co.: 2 mi. W Ewing, 415 m, 3; 3 mi.

W Jonesville, 421 m, 1; 1 mi. N Rose Hill, Poor Valley Branch, 433 m,

1. Montgomery Co.: VPI & SU, Blacksburg, 610 m, 5. Rockbridge Co.:

1.5 mi. NW Lexington, 305 m, 1; Vesuvius, 460 m, 1 (USNM). Scott

Co.: ca. 1 mi. E Hiltons, 442 m, 1. Total 13.

Associated Species

In this study, five of the other eight species of shrews known from

Virginia (Sorex fumeus, Sorex hoyi , Blarina brevicauda, Blarina

carolinensis, and Cryptotis parva) were captured at one or more sites

with S. longirostris. Because of their narrow habitat preferences and

limited distribution in Virginia, it is unlikely that Sorex dispar or Sorex

palustris will be found with S. longirostris. The masked shrew, Sorex

cinereus, has been captured with S. longirostris in west-central Indiana

(Rose 1980), but never in Virginia or elsewhere. In our area, S. cinereus

and S. longirostris exhibit contiguous allopatry. Sorex cinereus is

common at high elevations, and we found that its lowest elevational

limits (between 442 and 594 m) approximated the highest elevations at

which the southeastern shrew was collected (457 to 610 m). Mean

elevations of captures of the two species were 823 m versus 155 m.

Southeastern Shrew

127

Habitat

Habitats in which we found Sorex longirostris confirmed earlier

observations; the species occurs in a wide range of cover types (French

1980a, 1980b; Wolfe and Esher 1981; Pagels et al. 1982). Frequency of

captures of S. longirostris in various habitats in each of the physiographic

provinces of Virginia are given in Table 1. In the lowlands this shrew

was caught with similar frequency in all habitats. In the mountains,

however, it was caught more often in fields and field-forest edges than

in forest (14 versus 1).

Sorex longirostris was caught often in the Piedmont (at 65% of the

pitfall localities), Blue Ridge (40%), and Coastal Plain (37%), but

infrequently in the Ridge and Valley Province — at only 11% of the

localities. The rarity and limited distribution of S. longirostris in

western Virginia are emphasized by the very few captures there in spite

of extensive pitfall sampling (Table 1 and Fig. 2). Regardless of cover

type, the actual number of S. longirostris captured in pitfalls in the

Ridge and Valley Province, 7 vs. 66 in the other provinces, was

significantly less than the expected value adjusted for sampling effort

(X2 = 50.3, P< 0.01).

DISTRIBUTION IN WESTERN VIRGINIA

Our records extend the known range of S. longirostris in western

Virginia. Specimens from Scott and Lee counties bring the range of the

species to the edge of the Appalachian Plateau Province, where it

already is known to occur in Tennessee (French 1980b), Kentucky

(Caldwell and Bryan 1983), and West Virginia (French 1976). From the

localities in southwestern Virginia, the range of S. longirostris is

continuous southward along the Powell, Clinch, and Holston rivers into

the Tennessee Valley.

The specimens from Blacksburg confirm the occurrence of S.

longirostris there. This is close to the site where Handley collected a

specimen, subsequently lost, that he identified and published as Sorex

longirostris (Handley and Patton 1947). The external measurements of

this specimen, a male, total length 80 mm, tail vertebrae 32 mm, hind

foot 1 1 mm, ear 9 mm, and weight 3.0 g, are close to the mean for S.

longirostris.

We discovered that a specimen (USNM 521113) that had been

found floating dead in a swimming pool in Vesuvius, Rockbridge

County, in 1956 is a young S. longirostris , not Microsorex hoyi

winnemana Preble as we previously reported (Handley et al. 1980). This

establishes the occurrence of S. longirostris in the mid-Shenandoah

Valley. Later, we caught another specimen nearby, at Lexington.

128

John F. Pagels and Charles O. Handley, Jr.

In the Appalachians S. longirostris seems to be uncommon and to

be found only at relatively low elevations. Published records reveal a

general south-to-north gradient of the upper limits of its distribution:

North Carolina

Tennessee

Kentucky

West Virginia

Macon Co. 762 m

Sevier Co. 488 m

Knox Co. 335 m

Roane Co. ca. 305 m

(Gentry et al. 1968)

(Komarek and Komarek 1938)

(Caldwell and Bryan 1983)

(French 1976)

These localities are all on the west slope of the Appalachians, in the

Mississippi drainage. Our records from extreme southwestern Virginia,

with elevations of 427 to 457 m, fit into this gradient. Apparently a

different south-north gradient, displaced a little to the north, operates

on the east slope in Virginia:

We did very little sampling in the Blue Ridge, and most of the

capture sites there, all in the foothills, were near ones that we had

reported earlier (Pagels et al. 1982). Nevertheless, it must be assumed

that the Blue Ridge represents a formidable barrier to distribution of

Sorex longirostris. Gaps at the Roanoke and James rivers, Chester Gap

at the head of the Rappahannock River, Manassas Gap at the head of

Goose Creek, and the Potomac River Gap provide the only access for

this shrew to the lowlands behind the mountain range (Fig. 2).

We now have evidence that all of these passages except the Potomac

River Gap actually have been used by S. longirostris — the Roanoke

River Gap to Blacksburg, the Roanoke or James gaps (or both) to

Lexington and Vesuvius, and Chester Gap or Manassas Gap (or both)

to Front Royal and Luray. Specimens actually were taken in Chester

Gap and in Manassas Gap (at Linden).

Although there can be no doubt that Sorex longirostris has used

the gaps in the Blue Ridge to gain access to the Valley of Virginia, it is

not clear why its distribution in the Valley is so limited. In this area 45

of the pitfall localities were below 610 m, the highest elevation at which

S. longirostris has been found in Virginia (Fig. 2). Within the elevational

limits represented by the highest point where this shrew was found in

each basin, there were 23 pitfall localities. Sorex longirostris was caught

at only 5 (22%) of the 23 localities (2/8 of these were in fields, 3/13 in

ecotones, and 0/2 in forest). The species has reached the head of the

Roanoke River and crossed over the divide, barely into the extensive

Southeastern Shrew

129

Table 1. Number of sites sampled, frequency expressed as percentage of sites

where at least one Sorex longirostris was collected, and number of

individuals of S. longirostris captured in each habitat type and province.

New River lowlands, but seems to have made no further penetration.

Through the James or Roanoke gaps it has reached the head of the

Maury River watershed at Vesuvius, but has not been found at all in the

main valley of the James (including Cowpasture and Jackson rivers and

Craig and Catawba creeks). Similarly, it has been found a little way up

the South Fork of the Shenandoah River from Chester and Manassas

gaps, but it is unknown in the broad expanses of the Potomac and

lower Shenandoah valleys west of the Blue Ridge.

Can it be that the invasion of the Valley of Virginia by S.

longirostris is relatively recent, and that what we see are merely the

early stages of its occupation of habitat suitable for it but too low for S.

cinereusl Or, has S. cinereus , which undoubtedly occupied the whole

area during the Pleistocene, only recently withdrawn or begun to

130

John F. Pagels and Charles O. Handley, Jr.

Fig. 2. Distribution of Sorex longirostris west of the Blue Ridge in Virginia.

The heavy dashed line follows the Blue Ridge front. Arrows indicate low-

elevation gaps in the Blue Ridge and access from the Tennessee Valley. Shaded

areas represent potential S. longirostris habitat in western Virginia and adjacent

parts of Maryland and West Virginia: (1) Tennessee River watershed up to 457

m. (2) Roanoke River basin up to 610 m. (3) New River basin from the West

Virginia boundary up to 610 m (continuing down to the Ohio Valley in West

Virginia). (4) James River watershed up to 457 m. (5) Manassas Gap to

Shenandoah River basin up to 305 m. (6) Potomac River basin up to 305 m.

Solid circles represent pitfall trap localities where S. longirostris was caught;

open circles represent pitfall trap localities where S. longirostris was not found.

Solid triangles represent localities where S. longirostris has been taken by means

other than pitfall traps.

withdraw from the lower elevations of the Valley of Virginia, clearing

the way for S. longirostris ? Only more collecting can unveil the full

extent of details of the distribution of S. cinereus and S. longirostris and

its oddities in western Virginia.

ACKNOWLEDGMENTS.— We thank Jack Cranford of Virginia

Polytechnic Institute and State University, for donation of specimens.

Walter Bulmer of Northern Virginia Community College, Annandale

Campus; Carl Ernst of George Mason University; and Robert Simpson

of Lord Fairfax Community College allowed us to examine and report on

Southeastern Shrew

131

specimens under their care. We thank them for these privileges and

other kindnesses. Lauren Seymour compiled Figure 2. John E. Pagels

assisted in many of the collections, and Donald Young provided many

helpful comments on the habitat component of the study. John F.

Pagels’s efforts were supported by the Scholarly Leave Program of

Virginia Commonwealth University and the Non-game Wildlife and

Endangered Species Program of the Virginia Commission of Game and

Inland Fisheries. We are grateful to Darelyn Handley and Donald

Young for constructive criticism of the manuscript.

LITERATURE CITED

■=!

Bruce, James A. 1937. Sorex longirostris longirostris in Augusta County,

Virginia. J. Mammal. 18:513-514.

Caldwell, Ronald S., and H. Bryan. 1982. Notes on distribution and habitats

of Sorex and Microsorex (Insectivora: Soricidae) in Kentucky. Brimleyana

8:91-100.

French, Thomas W. 1976. The first record of the southeastern shrew, Sorex

longirostris , in West Virginia. J. W. Va. Acad. Sci. 48:120-122.

French, Thomas W. 1980a. Natural history of the southeastern shrew, Sorex

longirostris Bachman. Amer. Midi. Nat. 104:13-31.

French, Thomas W. 1980b. Sorex longirostris. Mammal. Species 143:1-3.

Gentry, John B., E. P. Odum, M. Mason, V. Nabholz, S. Marshall, and J. T.

McGinnis. 1968. Effect of altitude and forest manipulation on relative

abundance of small mammals. J. Mammal. 49:539-541.

Handley, Charles O., Jr., and C. P. Patton. 1947. Wild mammals of Virginia.

Comm. Game Inland Fisheries, Richmond.

Handley, Charles O., Jr., J. F. Pagels, and R. H. de Rageot. 1980. PYGMY

SHREW Microsorex hoyi winnemana Preble. Pages 545-547 in Endangered

and Threatened Plants and Animals of Virginia, D.W. Lindsey, editor.

Center for Environmental Studies, Va. Polytech. Inst, and State Univ.,

Blacksburg.

Komarek, Edwin V., and R. Komarek. 1938. Mammals of the Great Smoky

Mountains. Bull. Chicago Acad. Sci. 5:137-162.

Odum, Eugene P. 1944. Sorex longirostris at Mountain Lake, Virginia. J.

Mammal. 25:196.

Pagels, John F., C. S. Jones, and C. O. Handley, Jr. 1982. Northern limits of

the southeastern shrew, Sorex longirostris Bachman (Insectivora: Soricidae),

on the Atlantic coast of the United States. Brimleyana 8:51-59.

Pagels, John F., and T. W. French. (1987) Discarded bottles as a source of

small mammal distribution data. Amer. Midi. Nat. 1 18:217-219.

Rose, Robert K. 1980. The southeastern shrew, Sorex longirostris , in southern

Indiana. J. Mammal. 61:162-164.

Wolfe, James L., and R. J. Esher. 1981. Relative abundance of the southeastern

shrew. J. Mammal. 62:649-650.

Accepted 7 October 1987

132

DATE OF MAILING

Brimleyana No. 14 was mailed on 27 May 1988.

THE SEASIDE SPARROW,

ITS BIOLOGY AND MANAGEMENT

Edited by

Thomas L. Quay, John B. Funderburg, Jr., David S. Lee,

Eloise F. Potter, and Chandler S. Robbins

The proceedings of a symposium held at Raleigh, North Carolina,

in October 1981, this book presents the keynote address of F. Eugene

Hester, Deputy Director of the U.S. Fish and Wildlife Service, a

bibliography of publications on the Seaside Sparrow, and 16 major

papers on the species. Authors include Arthur W. Cooper, Oliver L.

Austin, Jr., Herbert W. Kale, II, William Post, Harold W. Werner,

Glen E. Woolfenden, Mary Victoria McDonald, Jon S. Greenlaw,

Michael F. Delany, James A. Mosher, Thomas L. Merriam, James A.

Kushlan, Oron L. Bass, Jr., Dale L. Taylor, Thomas A. Webber, and

George F. Gee. A full-color frontispiece by John Henry Dick illustrates

the nine races of the Seaside Sparrow, and a recording prepared by J.

W. Hardy supplements two papers on vocalizations.

“The Seaside Sparrow, with its extensive but exceedingly narrow

breeding range in the coastal salt marshes, is a fascinating species. All

the authors emphasize that the salt marsh habitat is at peril. . . . The

collection is well worth reading.” — George A. Hall, Wilson Bulletin.

1983 174 pages Softbound ISBN 0-917134-05-2

Price: $15, postpaid, North Carolina residents add 5% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send to SEASIDE SPARROW, N.C. State Museum of Natural Sciences,

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

133

MANUSCRIPT REVIEWERS

The editorial staff is grateful to the following biologists who

reviewed manuscripts for Brimleyana Numbers 11 through 15. The

acting editor especially appreciates the encouragement offered along

with the reviews.

Sydney Anderson, American Museum of Natural History

Thomas C. Barr, Jr., University of Kentucky

Ronald A. Brandon, Southern Illinois University at Carbondale

Alvin L. Braswell, North Carolina State Museum of Natural Sciences

Brooks M. Burr, Southern Illinois University at Carbondale

Charles C. Carpenter, The University of Oklahoma

Robert C. Cashner, The University of Oklahoma

Mary Kay Clark, North Carolina State Museum of Natural Sciences

John E. Cooper, Reidsville, North Carolina

Graham J. Davis, East Carolina University

Charles D. Duncan, University of Maine at Machias

Stephen R. Edwards, Museum of Natural History, University of Kansas

Carl Ernst, George Mason University

David A. Etnier, University of Tennessee

J. F. Fitzpatrick, Jr., University of South Alabama

Thomas W. French, The Nature Conservancy

Sidney A. Gauthreaux, Jr., Clemson University

Carter R. Gilbert, Florida State Museum, University of Florida

J. Christopher Haney, The University of Georgia

Stephen R. Humphrey, Florida State Museum, University of Florida

John B. Iverson, Earlham College

Robert E. Jenkins, Roanoke College

Ernest A. Lachner, Smithsonian Institution

David S. Lee, North Carolina State Museum of Natural Sciences

Robert W. Ling, Jr., Langley Air Force Base, Virginia

Charles Lytle, North Carolina State University

William C. McComb, University of Kentucky

Joseph C. Mitchell, University of Richmond

Robert H. Mount, Auburn University

Samuel C. Mozley, North Carolina State University

Jerry W. Nagel, East Tennessee State University

D. R. Oliver, Biosystematics Research Institute

William M. Palmer, North Carolina State Museum of Natural Sciences

Stewart B. Peck, Carleton University

George R. Pisani, The University of Kansas

Roger A. Powell, North Carolina State University

Richard R. Repasky, North Carolina State University

134

Robert K. Rose, Old Dominion University

Win Seyle, Savannah Science Museum, Savannah, Georgia

Rowland W. Shelley, North Carolina State Museum of Natural Sciences

Annelle R. Soponis, Florida Agricultural and Mechanical University

William R. Taylor, Smithsonian Institution

David K. Woodward, North Carolina State University

135

ENDANGERED, THREATENED, AND

RARE FAUNA OF NORTH CAROLINA

PART I.

A RE-EVALUATION OF THE MAMMALS

Edited by Mary Kay Clark

This book is a report prepared by a committee appointed in 1985

by the North Carolina State Museum of Natural Sciences to re-evaluate

the list of mammals presented in Endangered and Threatened Plants

and Animals of North Carolina (John E. Cooper, Sarah S. Robinson,

and John B. Funderburg, editors. N.C. State Mus. Nat. Hist., Raleigh,

1977), which is now out of print. Committee members were Mary Kay

Clark, David A. Adams, William F. Adams, Carl W. Betsill, John B.

Funderburg, Roger A. Powell, Wm. David Webster, and Peter D.

Weigh The report treats 21 species listed in the following status

categories: Endangered (5), Threatened (1), Vulnerable (6), and

Undetermined (9). Most species accounts discuss the animal’s physical

characteristics, range, habitat, life history and ecology, special sig-

nificance, and status (including the rationale for the evaluation and

recommendations for protection) and provide a range map and an

illustration of the animal’s external characters. Ruth Brunstetter and

Renaldo Kuhler illustrated the book. An introductory section contributed

by Ms. Clark discusses the changes in status that occurred in the decade

between 1975 and 1985. It also mentions efforts to protect marine

mammals and includes a checklist of the cetaceans known from North

Carolina.

1987 52 pages Softbound ISBN 0-917134-14-1

Price: $5 postpaid. North Carolina residents add 5% sales tax. Please make

checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: ETR MAMMALS, N.C. State Museum of Natural Sciences,

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

i

136

ENDANGERED, THREATENED, AND

RARE FAUNA OF NORTH CAROLINA

PART II.

A RE-EVALUATION OF THE MARINE AND

ESTUARINE FISHES

by

Steve W. Ross, Fred C. Rohde, and David G. Lindquist

This is the second in a series of reports by committees appointed in

1985 by the North Carolina State Museum of Natural Sciences to re-

evaluate the faunal lists presented in Endangered and Threatened Plants

and Animals of North Carolina (John E. Cooper, Sarah S. Robinson,

and John B. Funderburg, editors. N.C. State Mus. Nat. Hist., Raleigh,

1977), which is now out of print. The report on marine and estuarine

fishes by Ross, Rohde, and Lindquist treats one Endangered species, six

Vulnerable species, and four anadromous fishes that, while not formally

listed, are of some concern. Five species listed as being of Special

Concern in 1977 no longer warrant formal status. The publication

includes six original drawings by Renaldo Kuhler.

1988 20 pages Softbound ISBN 0-917134-17-6

Price: $3 postpaid. North Carolina residents add 5% sales tax. Please make checks

payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: ETR MARINE FISHES, N.C. State Museum of Natural Sciences,

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

FEB 23 1989

INFORMATION FOR CONTRIBUTORS

Submit original and two copies of manuscripts to Editor, Brimleyana, North Carolina

State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. In the case of

multiple authorship, indicate correspondent. Manuscripts submitted for publication in this

journal should not also be submitted elsewhere.

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belong. Example: Studies of the genus Hobbseus Fitzpatrick and Payne (Decapoda:

Cambaridae).

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second authors use initials followed by last name. Examples:

Woodall, W. Robert, Jr., and J. B. Wallace, 1972. The benthic fauna in four small

southern Appalachian streams. Amer. Midi. Nat. 88(2):393-407.

Crocker, Denton W., and D. W. Barr. 1968. Handbook of the Crayfishes of Ontario.

Univ. Ontario Press, Toronto.

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for proofreading.

BRIMLEYANA NO. 15, JANUARY 1989

CONTENTS

Occurrence of the Nine-banded Armadillo, Dasypus novemcinctus

(Mammalia: Edentata), in South Carolina. John J. Mayer 1

Distribution and Seasonality of Branchioped and Malacostracan

Crustaceans of the Santee National Wildlife Refuge, South Carolina.

Charles K. Biernbaum 7

Taxonomic Analysis of the Coastal Marsh Raccoon ( Procyon lotor

maritimus) in Maryland. Denise H. Clearwater , George A. Feldhamer,

Raymond P. Morgan II, and Joseph A. Chapman 31

Tolerance of Acidity in a Virginia Population of the Spotted Salamander,

Ambystoma maculatum (Amphibia: Ambystomatidae). Charles R.

Blem and Leann B. Blem f 37

Population Structure and Biomass of Sternotherus odoratus (Testudines:

Kinosternidae) in a Northern Alabama Lake. C. Kenneth Dodd, Jr. .. 47

Distribution, Biology, and Conservation Status of the Carolina Madtom,

Noturus furiosus, an Endemic North Carolina Catfish. Brooks M.

Burr, Bernard R. Kuhajda, Walter W. Dimmick, and James M.

Grady 57

Pelagic and Near-shore Plankton Communities of a North Carolina

Coastal Plain Reservoir. Michael A. Mallin 87

Reproductive Biology of the Brown Water Snake, Nerodia taxispilota, in

Central Georgia. Robert E. Herrington 103

Movements of Land-based Birds Off the Carolina Coast. David S. Lee

and Kenneth O. Horner Ill

Distribution of the Southeastern Shrew, Sorex longirostris Bachman, in

Western Virginia. John F. Pagels and Charles O. Handley, Jr 123

Miscellany 132

Manuscript Reviewers 133