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North Carolina State Library
Raleigh

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FEB 2 1981

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number 1

march 1979

EDITORIAL STAFF

John E. Cooper, Editor
Alexa C. Williams, Managing Editor
John B. Funderburg, Editor-in-Chief

Board

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

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

State Museum State Museum

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

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

State Museum State Museum

Martha R. Cooper, Associate Thomas L. Quay, Department

Curator (Crustaceans), N.C. of apology, N.C. State

State Museum University

James W. Hardin, Department Rowland M. Shelley, Chief

of Botany, N.C. State Curator of Invertebrates, N.C.

University State Museum

Brimleyana, the Journal of the North Carolina State Museum of Natural History,
will appear at irregular intervals in consecutively numbered issues. Con-
tents will emphasize zoology and general ecology of the southeastern Uni-
ted States, especially North Carolina and adjacent areas. Geographic
coverage will include AL, DE, FL, GA, KY, LA, MD, MS, NC, SC, TN,
VA and WV. Suitability of manuscripts will be determined by the
Editorial Board, and each paper adjudged suitable will be reviewed by ap-
propriate specialists. Final acceptability will be decided by the Editor.

Address all requests for information on purchase and exchange to the
Managing Editor, BRIMLEYANA, N.C. State Museum of Natural
History, P.O. Box 27647, Raleigh, NC 27611. Manuscripts and all other
correspondence should be addressed to the Editor at the same address.

In citations please use the full name — Brimleyana.

ISSN: BRIMD7 M tyfe

About the name of this journal . . .

The Brothers Brimley: North Carolina Naturalists

John E. Cooper

North Carolina State Museum of Natural History,
P.O. Box 27647, Raleigh, North Carolina 27611

During the middle years of the nineteenth century two sons were born
into a family of long time farmers living near Bedford in the midlands sec-
tion of England, northwest of London. One made his appearance at the
family home in the village of Willington, Bedfordshire, on March 7, 1861 ;
the other, a "seven months" baby, arrived unexpectedly at the home of
his maternal grandmother at Great Linford, Buckinghamshire, on
December 18, 1863. In their early youth, as the younger of them was to
write many years later, they "collected birds' eggs, caught small birds in
brick traps in the winter, went fishing, and meddled with living creatures
in general after the usual fashion of boys" and "had a reasonably good
working knowledge of the wild life around us." No one could then have
guessed, however, that these English farm lads would one day be
recognized as two of the most remarkable naturalists of their time in the
southeastern United States.

Their first step in this direction was initiated by misfortune and guided
by chance. Agriculture in England had experienced a series of poor
seasons in the 1870s, and by the end of the decade farm prices were at an
all time low. The family faced the sad reality that its only hope for new
beginnings lay in emigration to a distant land and made tentative plans
for a move to Australia or Canada. But before their plans became final,
an essentially accidental meeting with an official of the newly-formed
North Carolina Department of Agriculture, Immigration and Statistics
convinced them that America would be their Land of Opportunity.

Late on the night of December 31, 1880, Herbert Hutchinson Brimley,
nearly 20 years old, and Clement Samuel Brimley, barely 17, having
crossed the Atlantic by steamer to New York, arrived in Raleigh on the
heels of a blizzard. With them were their parents, Joseph and Harriet,
two sisters, and one of two living brothers. Their first stop was at a hotel
which, less than a year later, became the Agriculture Building, the very
place where they would ultimately labor for much of their lives. H.H.
Brimley 's remembrance of that time was written almost 50 years later:

"My first impression of Raleigh was that it was without question
the damndest place I had ever seen. Expecting to jump directly into

Brimleyana No. 1: 1-14. March 1979. 1

John E. Cooper

Clement Samuel Brimley
December 18, 1863 —July 23, 1946

the justly celebrated Sunny South, irrespective of time of the year
(December 31, 1880), I found a town with unpaved streets, ruts
hub-deep, frozen solid and covered with snow, and the temperature
down mighty close to zero. There were some board sidewalks, but
military tanks or caterpillar tractors would have been the only
suitable vehicles for negotiating those streets under the prevailing
conditions.

"The hotel, later used as the Agriculture Building, was not equip-
ped with running water, and that in the pitcher in the bedroom I
occupied was frozen solid. We had to pull up the carpets and use
them for blankets to keep from freezing to death the first night, no
artificial heat being provided in the rooms. Bedford, even at the

The Brothers Brimley

Herbert Hutchinson Brimley
March 7, 1861 — April 4, 1946

time known as an educational center, had paved streets and running
water in the houses. Raleigh depended on wells under the sidewalks
equipped with wooden pumps for its public water supply. Back-
yard and frontyard wells and pumps supplied some of the more
pretentious residences. In short, Bedford was a more or less finished
town, for its time. Raleigh conveyed the impression of being com-
paratively raw — the cows and hogs roamed the streets, giving it
something of a rural atmosphere. However, there was a restless,
pulsing air about the place and its people that impressed me."

The Brimleys soon found that their new land was not exactly an
agrarian paradise, and their attempts at farming the rocky Piedmont soil

4 John E. Cooper

near Method, between what is now west-central Raleigh and Cary, were
of limited success. H.H. turned to teaching in a one-room log schoolhouse
in House Creek Township near the site of present Meredith College, but
this venture also ended in failure, at least partly because his English ac-
cent and the southern drawl of his students proved incompatible. In his
words, "I will never know which of the three bodies were happier when I
resigned — the school committee, the students or myself. We just could
not understand each other."

Fortunately, although not endowed with much in the way of worldly
possessions, H.H. and C.S. brought with them two incredibly inquisitive
minds and an intense interest in nature developed in the hedgerows and
fields of the Ouse River valley. Immediately upon arriving in their adop-
ted land, and as time permitted between various unsuccessful attempts at
earning a living, they set about studying its wild creatures, especially the
birds. Sometime in 1882 or 1883 they came into possession of a 50-cent
book entitled Taxidermy Without a Teacher, and began dabbling in the art of
mounting and modeling animals. As their skills in this field developed, es-
pecially those of H.H. , a whole new enterprise enabling them to indulge
their love for the outdoors and natural history collecting opened before
them, and they drifted into a business under the name of "Brimley Bros.,
Collectors and Preparers." In 1942 H.H. wrote, "Following our arrival in
Raleigh in 1880 the main activity of my brother, C.S., and I in endeavor-
ing to keep the justly celebrated wolf from the not-too-securely fastened
door was a crude grade of custom taxidermy together with the collecting
of bird skins and eggs for wealthy men in the big cities, who vied with
each other over the comparative magnitude of their collections." As we
shall see, these unsure commercial beginnings were the foundations of
two outstanding careers in natural history.

At about the time the Brimleys were just beginning their "meddling
with living creatures," and probably well before either had ever heard of
North Carolina, a series of events was unfolding here, which, in
retrospect, almost seemed tied to them by some arcane cosmic threads.
They culminated in formation on March 12, 1877 of the N.C. Depart-
ment of Agriculture, Immigration and Statistics, and appointment of the
first commissioner of agriculture, Colonel Leonidas Lafayette Polk. Some
time before assuming this office Polk had urged the State Grange to es-
tablish a central headquarters containing a "Patron's State Musuem" for
the display of North Carolina's agricultural products. Shortly after
becoming commissioner he began such a museum himself in a room ad-
joining his office in the Briggs Building on Fayetteville Street in Raleigh,
which since 1875 had also housed the Geological Survey. Washington
Caruthers Kerr, state geologist, was maintaining a mineral collection

The Brothers Brimley 5

there, and often referred to it as the "geological museum" and even "state
museum." There were, in effect, two distinct "state museums" under one
roof, neither of them officially so designated. The mineral cabinet,
however, was a legal state collection, dating at least to the mid- 1850s
when the General Assembly required Ebenezer Emmons, Kerr's
predecessor, to establish such a display in the State Capitol. At any rate,
on February 20, 1879 the legislature placed the survey under the jurisdic-
tion of the Department of Agriculture and required the commissioner,
"by and with the consent and advice of the Board of Agriculture" to keep
a "museum or collection to illustrate the agricultural and other resources
and the natural history of the State." The geological and agriculatural
collections were thus combined under this broader mandate, and the
State Museum was founded.

In 1881 the Department of Agriculture purchased the old National
Hotel (site of the Brimleys' first Raleigh stop) to consolidate its offices.
The museum was assigned part of the space for displays and other collec-
tions, which included the remnants of exhibits made by the Board of
Agriculture at a number of expositions in this country and abroad. North
Carolina's participation in such expositions had proved quite advan-
tageous to a variety of economic enterprises, and the state's committment
to them continued. As the time for the 1884 State Centennial Exposition
at Raleigh approached, H.H. Brimley 's growing reputation as a taxider-
mist and preparator came to the attention of the board, which was plann-
ing and assembling exhibits on North Carolina's resources. Consequently
he was hired to mount a series of fishes for the display, under the direc-
tion of Stephen G. Worth, superintendent of fish and fisheries. Brimley
also made a collection of waterfowl from Currituck Sound for this exposi-
tion, as recounted in "Old Times on Currituck," published in North
Carolina Wildlife Conservation for March 1943.

The Brimley brothers' partnership became a going concern and gave
them marvelous excuse to collect and study the animals of North
Carolina, about which little was known at that time. Although they were
in the natural history business to earn a living, their intellectual interests
were not subordinated. 1884, in addition to being noteworthy as the year
of H.H. Brimley's first employment by the Board of Agriculture, was also
the year in which the brothers began publication of a series of notes on a
new and exciting fauna. Initially their contributions concerned birds, and
most appeared in the pages of Ornithologist and Oologist. These were only
the first efforts, however, in a long series of publications covering many
topics in a noteworthy array of journals, magazines, and newspapers.

In 1890 planning began for participation in the great World's Colum-
bian Exposition of 1893, and the State World's Fair Commission did not

6 J°hn E. Cooper

hesitate to select H.H. Brimley to collect and prepare native animals for
the North Carolina exhibits. He assembled and installed the Fish and
Fisheries Exhibit, which again included "aquatic birds," and remained
with it in Chicago throughout the exposition. The voluminous collections
gathered for the state's many displays, too excellent and valuable to dis-
card or relegate to storage, were directed by legislative action to join the
State Musuem upon their return to Raleigh. This further strained the
holding capacity of its none-too-capacious quarters, although the
available space had been nearly doubled by a minor addition in 1893, and
the collections remained rather jumbled and unusable. Among the pile of
materials were the disassembled bones of a Right whale known as
"Mayflower," and in 1894, not long after returning from Chicago, H.H.
Brimley was again hired by the board, this time for the singular task of
articulating and mounting the 46-foot skeleton for display. Since the
budget lacked money for such odd work he was hired under the job title
of "fertilizer inspector," as shown in the expenditure ledgers for that year.
The task took about three months, and this early piece of Brimley han-
diwork is still on display at the museum.

On April 15, 1895 the next logical step in the inexorable progression
was taken, and a new era signalled for the State Museum, when H.H.
Brimley was appointed its first full-time curator. He was also the sole em-
ployee and, as he wrote later, "I became expert with a feather duster and
pushed a wicked carpet sweeper! I had no funds beyond my princely
salary of $75 per month." His title remained curator until 1928, when it
was changed to director. H.H. continued to collect, prepare, ship, erect,
and occasionally to man the North Carolina exhibits at major fairs and
expositions. Each resulted in the addition of more and more collections to
the State Museum and the accumulation of a store of information on
North Carolina's natural history. An ever-growing fund of this informa-
tion was put into print in papers by the Brimleys. Not yet employed by
the state, C.S. mainly devoted his efforts to "Brimley Bros., Collectors
and Preparers." H.H. also continued to work in the business on a part-
time basis until 1907 or 1908, when he dropped out to devote his full
energies to the growing museum. By this time he had an assistant
curator, Tom Adickes, and a janitor, Bob Alston (who presumably
relieved the curator of his feather duster and carpet sweeper). The
museum now occupied the entire second floor of the Agriculture Building,
as well as another annex added in 1897.

In those early days the museum's holdings, as listed in the 1897 Hand
Book of the North Carolina State Museum, were diverse and even somewhat
cosmopolitan. Emphasis, of course, was on the geological and
agricultural resources of North Carolina, an immediate legacy of the

The Brothers Brimley 7

blending of survey collections with those assembled by Commissioner
Polk. By 1900, according to Curator Brimley 's report for that year, the
museum had accumulated "more than 100,000 specimens illustrative of
North Carolina . . . ." Although many of the materials obviously provided
data on the natural history of the state, there were at that time no collec-
tions maintained for purely scientific purposes. In this same report,
however, H.H. provided a portent of things to come, saying, "The
Zoological Department will be extended until the collections embrace
specimens of all the animal life native to the State." And, "Attention will
also be given to the flora of the State, and collections begun in that line.
This feature is at present unrepresented in the Museum." From the time
of this committment the museum moved steadily forward in most areas of
natural history. The old "National Hotel" was demolished in 1922 to
make way for a modern agriculture building, and the enlarged museum
reopened there in 1925.

H.H. Brimley continued to publish occasional zoological papers almost
until his death and received the singular honor in 1934 of being elected to
full membership in the American Ornithologists' Union, joining a select
assemblage of 150 of the nation's top students of birds. He was the only
North Carolinian to enjoy this privilege for quite a long time.
Nevertheless, his major talents were in areas other than technical
reporting. His reputation as a preparator of specimens and exhibits
climbed rapidly after he became curator of the State Museum. Among his
many accomplishments, assisted by various co-workers, were the model-
ing of a giant Ocean sunfish which weighed 1200 pounds and was seven
feet long and eight feet high; articulating and mounting the skeleton of a
54-foot Sperm whale estimated to weigh more than 50-tons in the flesh;
modeling a 17-foot Beaked whale and its fetus; modeling parts of a 35-
foot Whale shark, and whole modeling a 14-foot Basking shark and a
large Nurse shark; mounting and partially restoring the skeletal elements
of a mammoth; and mounting hundreds of other animals used in show-
case exhibits which he planned and prepared. He published a number of
papers on his methods and techniques and presented talks on the same
subjects at professional meetings. One such presentation, describing
methods of obtaining, preparing, and mounting whale skeletons, was
made at the 1929 meetings of the American Association of Museums and
published in The Museum Tears for November 1930. He was a lifelong
member of the AAS and seldom missed its annual meetings, often at-
tending at his own expense. He also belonged to the Museums' Associa-
tion of Great Britain.

Throughout his life, even into old age, H.H. was an obsessive out-
doorsman whose favorite activities included hunting and fishing, to which

8 John E. Cooper

he brought a scholar's attentions. Because of his deep involvement in
these pursuits he was an outspoken conservationist and advocate of
strong, well-enforced laws for the protection of game and other animals.
He characteristically led attempts to convince the legislature to end the
confusing array of local statutes which he saw as working to the disadvan-
tage of North Carolina's wildlife. It was not surprising, therefore, that he
and T. Gilbert Pearson, founder of the National Association of Audubon
Societies, were close, lifelong friends. The two met shortly after Brimley
became curator of the museum, when Pearson was just emerging as one
of America's pre-eminent ornithologists and conservationists. Judging
from their correspondence they took particular delight in treating each
other irreverently, and H.H. often sardonically addressed Pearson as
"My Dear Boy."

In the early 1900s Pearson and the Brimleys collaborated on a major
project, compiling data for and writing one of the first state bird books
ever produced in the south — Birds of North Carolina. The first edition of
this pioneer work, printed and ready for binding in 1913, was destroyed
by fire in the printer's plant and the whole edition was lost before official
publication. It finally appeared, with additional notes, in April 1919,
published by the N.C. Geological and Economic Survey and profusely il-
lustrated by Rex Brasher, Robert Bruce Horsfall, and Roger Tory Peter-
son. A second edition, published by the State Museum in 1942, quickly
sold out. The third and final edition, revised by David L. Wray of the
department's entomology division, and Harry T. Davis, the museum's
second director, was published by the museum in 1959.

H.H. Brimley retired as director of the State Museum in 1937. As his
replacement agriculture Commissioner W. Kerr Scott appointed Harry
T. Davis, who had joined the staff in July 1920 as assistant curator and
curator of geology. H.H. remained in the museum's employ as senior
curator of zoology and stayed active until his death. During this period he
did a great deal of work on the revised second edition of the bird book,
published several scientific papers, and made two of his finest fish mounts
(a 75-pound Channel bass and a 594-pound Blue marlin, then records for
the east).

At the turn of the twentieth century, while exciting developments were
occurring in the museum, a significant project was slowly and laboriously
unfolding in another division of the Department of Agriculture. It added
the vast realm of invertebrates, especially insects, to the fauna under
study within the department, and involved C.S. Brimley nearly two
decades before he became a state employee. Franklin Sherman, Jr. joined
the Division of Entomology, and became North Carolina's first state en-
tomologist in 1900. In addition to his duties in economic or applied

The Brothers Brimley 9

entomology, he began at once to collect and catalogue the insects and
other arthropods of the state. One of his major purposes was to have a
collection which would "give as complete an idea as possible of the insect
life of the State, showing the different stages of growth of each species,
and the food upon which they live. " By the time of Sherman's 1 902 report
the collection had grown to "30,000 specimens, which may be seen at any
time in the office of the Museum building."

Early in the century Sherman began publishing information ac-
cumulated in the project, and by the end of the first decade had produced
lists of North Carolina tiger beetles, tabanid flies, flat bugs, butterflies,
scorpionflies, dragonflies, and grasshoppers and their relatives. Many
ofthese papers were coauthored with C.S. Brimley while he was still a
private entrepreneur. C.S. met Sherman about the time he became state
entomologist, and Sherman, as C.S. later wrote, "revived my long flag-
ging interest in insects." Their association developed into a lasting
friendship, and in December 1919 C.S. entered the service of the Division
of Entomology to take charge of the "Insect Survey." By this time data
had accumulated on more than 5000 species known to occur in North
Carolina, and Sherman and his group had prepared several hundred
pages of manuscript "for what we hope may ultimately be a volume on
'The Insects of North Carolina,' in which every known species of insect in
the State shall be listed."

C.S. Brimley assiduously set about the task of expanding the collec-
tions and data files and enlarged their scope to include other members of
the state's fauna. "When Mr. Brimley came into the state service," wrote
Sherman in 1925, "he soon likewise card-catalogued the species of
Batrachians, Reptiles, and Mammals of the state which have been
recorded in papers by himself or others. With the Vertebrates thus on
record, and the Insects in full swing, it was merely an incident to include
the Invertebrates other than Insects." In addition to Sherman and
Brimley, other workers on the "Insect Survey" through 1925 were J. C.
Crawford, R.W. Leiby, C.L. Metcalf, Z.P. Metcalf, T.B. Mitchell, and
M.R. Smith.

In most respects C.S. was the more scientifically oriented of the
Brimley brothers and kept remarkably detailed records on southeastern
fauna and flora for more than 60 years. By the time he joined the depart-
ment he was already widely published in a number of journals, and his
amazing expertise, largely self-taught, extended to entomology, herp-
etology, mammalogy, ornithology, and botany. He was also quite
knowledgeable about many invertebrate groups other than insects. In
short, he was a virtuoso naturalist, whose "main interest for many years
zoologically," as he wrote in 1925, "has been to gain and disseminate

10 John E. Cooper

knowledge about the fauna of North Carolina, both vertebrates and inver-
tebrates, with especial regard to Herpetology and Entomology, an in-
terest very largely inspired and stimulated by Mr. Sherman."

C.S. Brimley's first publications were a number of ornithological notes
on which he appeared as junior author with his older brother, starting
with "Notes from middle North Carolina" in the October 1884 issue of
Ornithologist and Oologist. From then through 1894 he published another 70
brief solo notes, most of them in this same publication (which became
defunct in 1893), and a few in the Auk. He became active in herpetology
around 1890 and said that he was greatly helped by David Starr Jordan's
Manual of Vertebrates, which for a number of years was his "vertebrate 'Bi-
ble'." His first non-ornithological publication apparently was the 1895
"List of snakes observed at Raleigh, N.C.," in the American Naturalist.
During the next few years he published papers on amphibians, fishes, and
larval insects, more on reptiles and birds, lists of mammals of Raleigh and
of Bertie County, and a 32-page descriptive catalogue of the mammals of
the state. He also collaborated with Sherman on many of the insect lists.

As his notebooks on file in the State Museum archives show, C.S. kept
painstaking and meticulous records of all bird movements that occurred
in his vicinity. He published several summary papers on this subject, in-
cluding the 1917 "Thirty-two years of bird migration at Raleigh, North
Carolina," in the Auk. On December 1, 1930 he was awarded a certificate
from the Biological Survey, U.S. Department of Agriculture, in recogni-
tion of his 46 years of bird migration studies from 1885 to 1930. The
earlier records provided much of the migration data for Birds of North
Carolina.

In the course of his studies of the southeastern herpetofauna, C.S.
Brimley described several new species and subspecies. The first of these
were two salamanders named in 1912 — Plethodon metcalfi, after another
pioneer North Carolina scientist and co-worker Z.P. Metcalf, and the
subspecies Spelerpes (now Pseudotnton) ruber schencki, for C.A. Schenck,
director of the Biltmore Forest School. In 1924 C.S. recognized the en-
demic waterdog (an aquatic salamander) of the Neuse and Tar rivers as a
distinct subspecies, Necturus maculosus lewisi, naming it for Frank B. Lewis
who provided most of his specimens. A South Carolina salamander,
Plethodon clemsonae, followed in 1927, and two turtles in 1928 — Pseudemys
(now Chrysemys) vioscana from Louisiana, named for the naturalist Percy
Viosca, Jr., and the subspecies P. concinna elonae from a pond in Guilford
County not far from Elon College. However, only two of his new forms
have stood the test of time and further taxonomic studies. Necturus lewisi
was elevated to full species status in 1937 by Viosca, and Pseudotnton ruber
schencki is still recognized as a valid subspecies by some authorities

The Brothers Brimley

11

C. S. Brimley (left) and H. H. Brimley (right) in the basement prepara-
tion shop of the State Museum around 1944.

12 J°hn E. Cooper

although questioned by others. As would be expected C.S. also described
many insect species, primarily in the orders hymenoptera and diptera.
These included 13 psammocharid and 5 sphecid wasps, one each of
stratiomyid, cyrtid, conopid, sarcophagid and ortalid flies, and 4 asilid, 3
syrphid and 2 sciomyzid flies.

C.S. was a prolific writer who, before his career ended, published well
over 150 papers, notes and booklets on vertebrates, over 40 on inver-
tebrates, a 17-page paper on zoogeography, a partial bibliography of
North Carolina zoology, many popular natural history articles and ac-
counts, and a group of outlines for zoology lectures at the Biltmore
School. In addition to coauthoring the bird book with Pearson and his
brother, he wrote The Insects of North Carolina, which grew out of the early
manuscript begun by Sherman and others and included 35 years of
records on 9611 species. It was published in 1938 by the Department of
Agriculture. He also compiled the first supplement to this work,
published in 1942. A major summary of the amphibians and reptiles of
North Carolina, originally published as an annotated and illustrated
series in Carolina Tips from 1939 through 1943, was printed as a compila-
tion by Carolina Biological Supply Company in 1944. A similar collection
of North Carolina mammal accounts, written between 1944 and 1946, ap-
peared in 1946. Two installments toward a comprehensive series on fishes
of the state were published in the same outlet, but this project was in-
terrupted by C.S.'s death.

His publications also included a 20-year history of the North Carolina
Academy of Science. He and H.H. were founders of the Academy, and
C.S. was the only person without a college degree to ever serve as its
president. They were founders and life members, too, of the Raleigh
Natural History Club, and helped organize the Raleigh Bird Club and
the North Carolina Bird Club (now the two-state Carolina Bird Club).

In recognition of his outstanding contributions to the natural sciences,
on June 7, 1938 the University of North Carolina conferred on C.S.
Brimley the honorary degree of Doctor of Laws. This honor was all the
more remarkable in light of the fact that, while far from unlettered, both
the Brimley brothers were largely unschooled in any formal sense. H.H.
spent eight years in the Bedford County School at Elstow, excelling in
mathematics, football, and swimming, but left for a clerical job in
Howard's Iron Works before receiving a certificate. C.S. was educated in
the "common schools of Willington" until 1877, then attended the Bed-
ford County School through the close of the second term in July 1880.
"Attained the highest honors to be gained at that school," he wrote, "my
education on leaving being equivalent at least to completing a high grade
High School course or perhaps Freshman year in college." One writer,

The Brothers Brimley 13

W.T. Bost, later said that C.S. "had the sort of genius which made him a
great scientist without asking anything of the schools." Then, in reference
to his honorary doctorate, Bost added, "The University thinks it gave him
a degree; but in a larger sense he gave it one."

H.H. Brimley died at Rex Hospital on April 4, 1946, age 85, probably
the oldest active state employee in North Carolina's history. C.S. Brimley
died at his home a little over three months later, on July 23, 1946, while
dressing to go to his office in the Agriculture Building. Their productive
careers were dedicated to the Department of Agriculture; H.H. was
associated with the State Museum for over 60 years, 43 of them as direc-
tor, and C.S. with the Division of Entomology for at least 45 years, nearly
27 of them as an employee. Their service to the state of North Carolina
and its people, of course, cannot be measured in any such ordinary time
frame. They had a very profound influence on the scientific and
educational development of natural history in the southeast, particularly
in their chosen state, perhaps more than any other naturalists of their
time. The firm and enviable foundation which they laid was witnessed in a
letter of April 12, 1946 to C.S. from a friend in Charlottesville, Virginia.
"Both of you," he said, "have done a tremendous amount for the state in
stimulating it to go ahead with various things. Virginia has no such
museum, nor a collection of insects, nor such a bird book, or a catalogue of
its mammals — and a lot of other things are lacking because Virginia had
no Brimley brothers."

As a measure of the esteem in which these men were held by their
colleagues and other specialists who came later and appreciated their con-
tributions, a number of animals were named for them. These included the
fish Notropis brimleyi, the frog Pseudacns brimleyi, the salamander
Desmognathus brimieyorum, the millipeds Deltotana bnmleii and D. brimleardia,
and the hymenoptera Pedinaspis brimleyi, Halictus brimleyi, Colletes brimleyi,
and Ephuta pauxilla brimleyi.

Tributes to their influence on young naturalists, many of whom later
became outstanding scientists, are numerous. One of America's leading
ecologists, Eugene P. Odum (now Alumni Foundation Distinguished
Professor of Zoology and director of the Institute of Ecology at the Univer-
sity of Georgia), wrote in the preface to his 1949 compilation of H.H.'s
writings, T well remember my first visit with H.H. Brimley, when I was a
young high-school student. He took me under his wing and made me feel
at home immediately. The enthusiasm and sincerity with which he
worked and talked impressed me especially. In fact, H.H. Brimley and his
brother C.S. did more than anyone else to encourage me to develop my in-
terest in birds which later led me to go into teaching and research in
biology as a career."

A friend in England once wrote to H.H. Brimley: "But what a life you
have had, and what an interesting one; it was built for you and fitted your

14 J°hn E. Cooper

nature and inclinations; you must have reveled in it. It must have meant
any amount of strenuous effort; but after all, that is the gist of life, and
makes life worth living." These comments rather adequately summarized
the lives of both these gifted men. They came here as immigrant lads from
England and through diligence and dedication gave North Carolinians,
and indeed in a larger sense all Americans, the great legacy of a wealth of
knowledge about our natural heritage. This journal is dedicated to their
memory.

ACKNOWLEDGMENTS.—! would like to thank Alexa C. Williams,
John B. Funderburg, William M. Palmer, and Rowland M. Shelley, N.C.
State Museum, and our former employee Sarah S. Robinson, for
assistance in locating certain information; James F. Greene, Division of
Pesticides and Plant protection, N.C. Department of Agriculture, for
providing some of C.S. Brimley's notebooks; and Elaine H. Matthews,
Public Relations Division, same department, for providing specific docu-
ments. Alexa C. Williams, Martha R. Cooper, and David S. Lee, all of the
State Museum, made constructive criticisms of the manuscript, but any
remaining atrocities are mine.

SOURCES. — Quotations whose specific sources are not identified in the
text are from correspondence, unpublished manuscripts, and scripts and
sketches of various kinds contained in the archives of the North Carolina
State Museum of Natural History. The photographs are also from these
archives. Some information on H.H. Brimley came from brief biographical
sketches by Harry T. Davis (1946. J. Elisha Mitchell Sci. Soc. 62:128-129)
and C.S. Brimley (1946. Chat 10:42-43). Developments in the Department
of Agriculture were obtained from biennial reports of the Board of
Agriculture and the commissioner of agriculture published in Raleigh.
Franklin Sherman's comments anent the Insect Survey are from biennial
reports and from "Progress on State Insect Survey with comparative data
on other animal groups" (1925. J. Elisha Mitchell Sci. Soc. 47:129-134).
The comments by W.T. Bost are from the Greensboro Daily News, 27 July
1946. Information on Commissioner Polk mostly came from Stuart
Noblin's two-part article, "Leonidas Lafayette Polk and the North
Carolina Department of Agriculture" (1943. N.C. Historical Review
AT:103-121, 197-218). Some of H.H. Brimley's writings, including verse,
were compiled and annotated by E.P. Odum (1949. A North Carolina
Naturalist, H.H. Brimley. Univ. North Carolina Press, Chapel Hill). Ad-
ditional information on the Brimley brothers is contained in a history of
the Department of Agriculture's involvement in natural history studies
prepared by John B. Funderburg and me (in manuscript).

Accepted 19 December 1978

Nest and Larvae of the Neuse River Waterdog, Necturus
lewisi (Brimley) (Amphibia: Proteidae)

Ray E. Ashton, Jr.

and
Alvin L. Braswell

North Carolina State Museum of Natural History,
P.O. Box 27647, Raleigh, North Carolina 27611

ABSTRACT. — The first reported nest and hatchling larvae of Necturus
lewisi were found in Little River (Neuse River drainage), Wake County,
North Carolina, on 2 July 1978. Finding an adult male tagged for
behavioral studies with 60 Co wires led to their discovery. The nest was
under a flat, granite rock in 1.2 m of water at mid-river. Thirty-two
empty egg capsules, and three containing larvae which shortly emerged,
were attached to the underside of the rock. The male, found in a depres-
sion in the sand-gravel substrate directly beneath the eggs, was
apparently in attendance. Autopsy revealed that the male was in good
condition with an empty digestive tract. The presence of females of other
Necturus species in or near nests has been reported, but no males have
previously been verified in this situation. The hatchling larvae of N.
lewisi and N. maculosus are alike in color and pattern. However, post-
hatchling larvae of N. lewisi have a light mid-dorsum and dark sides,
while those of N. maculosus, as figured and described in the literature,
have a dark dorsum bordered on each side by a thin, light, dorsolateral
line. Necturus punctatus larvae are uniformly gray dorsally and lack strip-
ing. The distinctive post-hatchling larva of TV. lewisi lends added
credence to its current recognition as a full species.

INTRODUCTION

Necturus lewisi is a species of waterdog endemic to the Tar and Neuse
River drainages of North Carolina. Only one field study of this salaman-
der has been conducted (Fedak 1971). No accounts have been published
of its life history, and no descriptions of nesting, courtship and reproduc-
tion, or illustrations of larvae, have appeared. Adults and larvae are
described in Bishop (1926, 1943), Brimley (1924), Cahn and Shumway
(1926), Viosca (1937), and Hecht (1958), but the larva descriptions are
varied and contradictory. This paper describes the first discovered nest
and provides the first accurate descriptions and illustrations of TV. lewisi
hatchlings and older larvae.

MATERIALS AND METHODS

In November 1 977 we began a preliminary study of N. lewisi in the Lit-
tle River, a tributary of the Neuse River in northeastern Wake County.

Brimleyana No. 1: 15-22. March 1979. 15

16 Ray E. Ashton, Jr. and Alvin L. Braswell

One purpose was to develop methods of following movements, determin-
ing home range, and studying other behavior using radioactive tagging
and tracking techniques. Three adults (two females and a male) were
trapped in wire and plastic mesh minnow traps at that time, and each
was tagged with two 60 Co (35-50 mc) wires injected into the tail muscles
using methods described by Barbour et al. (1969).

We initially used a Thyac III Model 491 survey meter and scintillation
probe to locate animals, but monitoring proved difficult until a more sen-
sitive submersible Model 498 probe was obtained later in the study. The
tagged salamanders, which we had not located for three months, were
easily detected with this instrument. The nest was discovered in July 1978
while we were checking the location of the tagged male. Three larvae
were collected as they hatched from eggs, and four others were captured
by dip netting within 5 m of the nest site.

Larvae were preserved in 8 percent buffered formalin within one hour
after capture, and measurements, color observations and photographs
made within two hours after preservation. Measurements, made with an
ocular micrometer, are reported as snout-vent length (SV), measured
midventrally from tip of snout to vent pore; total length (TL); head
length (HL), from tip of snout to gular fold; head width (HW), at
posterior edge of eye socket; and tail width (TW), at widest point.

All specimens obtained in this study are deposited in the lower ver-
tebrate collections of the North Carolina State Museum of Natural
History (NCSM). Additional post-hatchling larvae and subadults from
various localities in both drainages were loaned by Duke University.

RESULTS

Nest. — On 2 July 1978 a nest and attending male (NCSM 19826;
147.6 mm SV) were found under a flat, granite rock (36 X 27 X 5.5 cm)
in 1.2 m of water near midstream, approximately 2 m from shore. The
underlying substrate was sand and fine gravel. Thirty-five egg capsules,
8-9 mm in diameter and each attached by a blunt stalk, were in an area of
about 60 cm2 on the underside of the rock. The rock's entire undersurface
was devoid of sessile and other invertebrates and debris. A depression in
the substrate, slightly larger than the nest area and about 40 mm deep,
apparently had been made and maintained by the male. The depression
narrowed and opened at the downstream edge of the rock, the upper edge
of which was embedded in the substrate. Water flow at the time was slow,
but this area is exposed to strong currents during flooding.

The male made no attempt to leave the nest until the rock was
removed. It was collected, preserved within two hours of capture, and
autopsied to verify sex and to determine general condition and any ob-

Necturus Nest and Larvae 1 7

vious effects of the radioactive tags. Prior to preservation the animal was
quite active and appeared healthy. Dissection revealed that the upper and
lower sections of the. digestive tract were empty, indicating that the
animal had not eaten recently. The liver appeared normal (compared
with other preserved specimens of similar size). The gall bladder was
filled with bile. Although no fat was evident the animal was not
emaciated. There was no indication of damage or irritation to muscle and
skin tissue from the tags.

Thirty-two of the 35 egg capsules found were empty, but three were in
the process of hatching and the larvae appeared within one hour.
Although we netted for approximately 5 m around the nest, only four ad-
ditional larvae were collected, all within one meter of the nest site. Large
rocks and gravel reduced netting efficiency.

Hatchlings (Fig. 1). — The mean total length of seven newly hatched
larvae is 22.8 mm (Table 1). The rounded head is totally unlike the
squared, elongate head of the adult and ranged from 3.5 to 3.7 mm (x =
3.5) long and 3.0 to 3.1 mm (x = 3.0) wide. The eyes are dark and appear
to be well formed. The forelegs are well developed, 2.0 to 2.2 mm long,
with the three outermost toes complete and the inner toe budlike. The
hind legs average 1.5 mm in length, are poorly developed, and are ad-
pressed to the lower tail fin. The tail is finned dorsally and ventrally, with
the dorsal fin slightly higher than the ventral and barely present at the
tail tip. Tail width in the series ranges from 2.9 to 3.2 mm (x = 3.0).
Because of the yolk sac the lower body is approximately 25 percent wider
than the upper body. The head and dorsum are rather uniform light
brown, and the color extends onto the area of the tail musculature. A
dark line extends from the nostril through the eye to the gills. Behind
each eye and above the dark line is a white, apparently rough, patch,
equal to or slightly smaller than the eye. The dorsal melanophores stop
abruptly on the side along a line level with the front and rear legs. Scat-
tered melanophores are present on the gills, upper surfaces of the legs,
lower jaw, and ventrolateral margins of the head. Melanophore concen-
trations are greatest in the area of the tail musculature, making the tail
slightly darker than the head and truck. The ventral tail fin is unpig-
mented, while the dorsal tail fin is clear on the margin with melanophores
increasing in number toward the musculature.

Post-hatchling larvae. — Color patterns were observed in 61 preserved N.
lewisi larvae from the Tar and Neuse drainages and in one live specimen
from the Neuse. Accurate description of some was difficult because of
varying degrees of fading. However, 48 percent of all specimens (N = 28)
between 21 and 41 mm SV have a distinct dark lateral stripe on each side,
and 37 percent have a light dorsum with the dark sides not well defined

Ray E. Ashton, Jr. and Alvin L. Braswell

m

■

mi

.*■>>■ -■

/„'-j

:-5:

i.<>-".

ff

be

Necturus Nest and Larvae

19

m»

Vj

s?/?

b&?

a

;■ ;/,*/

m

be
fa

20 Ray E. Ashton, Jr. and Alvin L. Braswell

(Fig. 2). These larvae also have a broad, light tan, dorsal stripe which
edges the costal grooves and extends from the snout onto the dorsum of
the tail and across its width. The dorsal region may have a scattering of
small, poorly defined, dark spots. The dark lateral stripe is continuous
from the nostrils through the eye and labial region to the tail where it
widens and includes all but the tail fringes. Irregular light blotches which
lack melanophores are scattered throughout the costal and tail region.
Most of the tail fin is lightly pigmented, but its edges lack pigmentation,
as does the tip of the tail. The underside is white or has a faint, reticulat-
ed pattern.

A larva, maintained alive for one year, measured approximately 30 mm
SV at capture and had a striped pattern which changed when it reached
45 mm SV. The lateral melanophores decreased in intensity while the
dorsal spots became darker and better defined. Sixty-three percent of the
preserved specimens (N = 33) between 45 and 72 mm SV show a similar
loss of striped pattern with a distinct increase in dorsal spotting. All
animals over 63 mm SV have an adult dorsal pattern.

DISCUSSION

The N. lewisi nest resembled reported stream nests of other Necturus
species in construction, location, and general conditions. The number of
eggs was greater than the 15 to 20 reported by Bishop (1943) for stream
nests of JV. maculosus, but within the range of 22 to 49 reported by Shoop
(1965) for N. m. louisianensis .

Although Bishop (1943) and others verified females near or attending
nests, few adults actually have been found in this situation. Most obser-
vers appear only to have surmised that nests are attended by females. Our

TABLE 1. Measurements (mm) of 7 newly hatched Necturus lewisi (NCSM 19827,
lot). SV = snout-vent length; TL = total length; TW = tail width; HL = head
length; HW = head width.

SV TL TW HL HW

15.3

22.8

3.0

3.5

3.0

15.5

23.6

3.2

3.7

3.0

16.3

23.0

2.9

*

*

15.5

23.1

3.0

3.5

3.1

15.0

22.2

3.0

3.5

3.0

15.9 .

23.4

2.0

3.5

3.1

14.9

21.8

3.1

3.6

3.0

x= 15.4 22.8 3.0 3.5 3.0

*damaged in capture

Necturus Nest and Larvae 21

deduction that the male present in the nest was attending it and not prey-
ing on hatchlings is based on the absence of larvae from the digestive tract
and the reluctance of the male to leave the nest. Broader studies which we
are now conducting on N. lewisi may provide evidence to corroborate this
observation.

Brimley (1924) stated, "Young specimens of the northern form (N.
maculosus) are said to be striped with black, but the smallest of ours that I
have seen (measuring only about 3 Vi inches in length) was spotted exact-
ly like larger specimens. In view of these facts it would seem fairly ap-
parent that the dwarf form occurring at Raleigh is a geographical variant
of subspecies of the true maculosus, and I propose for it the name Necturus
maculosus lewisi . . ."A similar description of the larva was given by Hecht
(1958). Viosca (1937) elevated N. m. lewisi to full species status and stated
that 55 mm larvae were dark on the sides and had a light dorsum marked
with faint spots. He also indicated that these spots develop into two or
three irregular rows as the larvae approach 90 mm, at which length all
striping disappears and they attain the adult spotted appearance. We
assume that Viosca 's measurements were total lengths, although this was
not indicated. If so, his observations compare favorably with ours of live
and preserved post-hatchling larvae, confirming that such larvae are quite
distinct from those of N. maculosus as illustrated by Bishop (1943), and N.
m. louisianensis as described by Shoop (1965).

The major difference is that N. maculosus post-hatchling larvae have a
dark dorsum bordered on each side by a thin, light, dorsolateral stripe.
However, newly hatched larvae of both species are indistinguishable from
each other. The larvae of Necturus punctatus, a species largely sympatric
with N. lewisi although occupying a broader range, are uniformly gray in
color with no evident striping (Brimley 1925, Bishop 1943). The dis-
tinctive post-hatchling larva of N. lewisi lends added credence to its
current recognition as a full species.

ACKNOWLEDGMENTS. — We wish to express our appreciation to
John C. Clamp, Patricia S. Ashton and John E. Cooper, North Carolina
State Museum, for their assistance in the field and their reviews of this
manuscript; Renaldo Kuhler, North Carolina State Musuem, for his skill
in illustrating the larvae; and Joseph R. Bailey, Duke University, for loan
of specimens. This study was partly supported by a grant from the
Carolina Conservationist program of the N.C. Wildlife Resources
Commission.

22 Ray E. Ashton, Jr. and Alvin L. Braswell

LITERATURE CITED

Barbour, Roger W., J.W. Hardin, J. P. Shafer, and M.J. Harvey. 1969. Home
range, movements, and activity of the dusky salamander, Desmognathus
juscus. Copeia 1 969(2) :273-297.

Bishop, S.C. 1926. Notes on the habits and development of the mudpuppy,
Necturus maculosus (Rafinesque). N.Y. State Mus. Bull. 265:5-60.

Bishop, Sherman C. 1943. Handbook of Salamanders. Comstock, Ithaca. 555 pp.

Brimley, C.S. 1924. The water dogs {Necturus) of North Carolina. J. Elisha
Mitchell Sci. Soc. 40(3-4) :1 66-1 68.

Cahn, A.R., and W. Shumway. 1926. Color variation in larvae of Necturus
maculosus. Copeia 1926(130) :4-8.

Fedak, Michael A. 1971. A comparative study of the life histories of Necturus lewisi
Brimley and Necturus punctatus Gibbes (Caudata: Proteidae) in North
Carolina. Master's thesis, Duke Univ., Durham. 103 pp.

Hecht, Max K. 1958. A synopsis of the mud puppies of eastern North America.
Proc. Staten Island Inst. Arts Sci. 27(l):l-38.

Shoop, C. Robert. 1965. Aspects of reproduction in Louisiana Necturus
populations. Am. Midi. Nat. 74:357-367.

Viosca, Percy, Jr. 1937. A tentative revision of the genus Necturus, with descrip-
tions of three new species from the southern Gulf drainage area. Copeia
1937(2):120-138.

Accepted 14 December 1978

A New Species of Xironodnlus Ellis 1918

from North Carolina (Clitellata:

Branchiobdellida)

Perry C. Holt

Department of Biology and Center for Systematics
Collections, Virginia Polytechnic Institute and
State University, Blacksburg, Virginia 24061

AND

Ann M. Weigl

Department of Biology, Wake Forest University,
Winston-Salem, North Carolina 27106

ABSTRACT. — Xironodnlus bashaviae, new species, is described from the
crayfish Cambarus bartonii collected at several sites in the upper drainage
of the Yadkin River, Forsyth County, North Carolina. It is the only
species of the genus known to occur east of the Blue Ridge Mountains.
The new species differs from other members of the genus primarily in
details of the reproductive system and dental formula.

None of the known species of the genus Xironodnlus Ellis 1918 has been
adequately described in published work. The discovery of a new species
outside the recorded range of the genus (Holt 1969) and the work of one of
us (W.) on the ultrastructure of this form affords us a propitious oppor-
tunity to present a description of this newly found species.

The literature pertaining to Xironodnlus is exceedingly sparse. Moore
(1894) recognized as Branchiobdella pulcherrima the first species now
assigned to the genus. Ellis (1919) formally erected the genus and more
fully described a species, X. formosus, which had been proposed in an
earlier paper (Ellis 1918). The effective date for both the erection of the
genus and the naming of its type-species (X. formosus) must derive from
this earlier work. There are other incidental references and some incom-
plete, but valid, species designations in the literature which will be cited
below, but the only detailed treatment of the genus is contained in Holt's
(1951) unpublished dissertation.

Our methods are those long used by Holt (1960 et seq.). Serial sections
were prepared by Weigl using material fixed in 70% ethanol and stained
with Harris' hematoxylin and eosin. The drawings (by H.) were done with
the aid of a camera lucida; all structures are illustrated with their anterior

Brimleyana No. 1: 23-29.March 1979. 23

24 Perry C. Holt and Ann M. Weigl

to the reader's right; cross-hatching indicates muscular and stippling,
primarily, glandular tissues. Measurements are approximations done with
an ocular micrometer and where averages are given the minimal and max-
imal measurements taken are enclosed in parentheses. The scanning elec-
tron micrograph was photographed by Weigl.

Genus Xironodrilus Ellis 1918

Type-species: — Xironodrilus formosus Ellis 1918, by subsequent designation
(Ellis 1919).

Diagnosis: — Body depressed widening gradually from segment I to seg-
ment VII; deferent ducts entering ental end of spermiducal gland;
prostate absent; bursa spherical; penis protrusible (modified from Hobbs
et al. 1967).

Remarks: — Members of the genus are distinguished from those of
Xironogiton Ellis 1919, and Ankyrodnlus Holt 1965, the only other American
branchiobdellids with a flattened ("depressed") body form, by the ental
entry of the vasa deferentia into the spermiducal gland. The species of
Xironodrilus are known from the Blue Ridge Province of the Appalachians
and the Interior Low Plateaus into the glaciated regions of Michigan
(Holt 1969): the species described herein is the only one known from the
Piedmont of the Appalachians east of the Blue Ridge Mountains.

Xironodrilus bashaviae, new species
Figures 1 and 2

Type- specimens: — Holotype, USNM 53641 taken on Cambarus bartonn from
Hunter's Creek, 0.8 km from the intersection of State Routes 1463 and
1446, Forsyth County, North Carolina, by Ann M. Weigl, 9 September
1975 (PCH 3376; AW 21 A); three paratypes (PCH 3376) in the VPI &
SU Center for Systematics Collections; three paratypes (AW 21 C) in the
collections of Ann Weigl.

Diagnosis: — Medium large branchiobdellids (average length about 3.5
mm); dental formula 4/4, lateral teeth of each jaw longer than median
ones; bursa subspherical; ejaculatory duct long, thick, curving laterodor-
sad from bursa; spermiducal gland large, with prominent lumen; sper-
mathecal duct long, bulb long, subcylindrical.

Etymology: — For Bashavia Creek, of which Hunter's Creek is a tributary.

Description: — Specimens of Xironodrilus bashaviae are large, but not among
the largest, branchiobdellids, averaging about 3.5 mm in length (pre-

New Species Branchiobdellid

25

Fig. 1. Structural details of Xironodnlus bashaviae new species. A, ventral view of
holotype; B, upper jaw; C, lower jaw of paratype; D, reproductive systems in ven-
tral view. Abbreviations: b, bursa; ed, ejaculatory duct; sb, spermathecal bulb; sd,
spermathecal ectal duct; sf, sperm funnel; sg, spermiducal gland; ve, vas efferens.

served material). The holotype (selected for the clarity with which internal
structures may be seen) has the following dimensions: total length, 2.8 mm;
greatest diameter, 0.6 mm; head length, 0.6 mm; head diameter, 0.4 mm;
diameter, segment I, 0.3 mm; diameter, sucker, 0.4 mm. The average dimen-
sions, with minima and maxima given in parentheses, of the holotype and
four specimens selected at random from the type-series are as follows: total

26

Perry C. Holt and Ann M. Weigl

- •••

Fig. 2. Scanning electron micrograph of a portion of the body of X. bashaviae.
Note ciliary tufts.

length, 3.4 mm (3.4-4.2 mm); greatest diameter, 0.8 mm (0.6-1. 2mm); head
length, 0.7 mm (0.6-0.8 mm); head diameter, 0.5 mm (0.4-0.6 mm); diameter,
segment I, 0.4 mm (0.3-0.5 mm); diameter, sucker, 0.5 mm (0.4-0.5 mm). In
the holotype and one paratype, segment V was greater in diameter than
segment VII. The latter, which bears the ovaries and developing eggs, is
normally greater in diameter among all members of the order. In these
two specimens there are no large eggs.

Scanning and transmission electron microscopy of X. bashaviae have
revealed the presence on the outer epithelium of numerous bristle-like
structures, each composed of a tuft of cilia. The tufts are most abundant in
the region of the mouth, but are also found on other parts of the head as
well as on the body segments (Fig. 2). They are not visible by light
microscopy, though "sensory" hairs have been detected in the mouth
region of other branchiobdellids (Franzen 1963:370; Moore 1895:499;
Holt, unpub. observ.).

The animals widen uniformly from segment I to their greatest diameter
in segments VI and VII and from there become narrower towards the
sucker and are uniformly, but not excessively, flattened, except for the
terete head. The posterior annulus of each segment is only slightly less in

New Species Branchiobdellid 27

diameter than the anterior one, conferring a generally smooth appearance
to the outline of the body.

There is a pronounced external sulcus or constriction of the head im-
mediately posterior to the position of the jaws. No other external evidence
of the segmentation of the head is apparent. Both upper and lower lips
bear very shallow and narrow emarginations. Oral papillae are not detec-
table in our specimens. There is one internal pharyngeal sulcus, deep, and
closely compressed, delimiting the posterior 1/3 of the head region from
the anterior 2/3.

The jaws are subequal in size, their width about 1/8 the greatest
diameter of the head. They consist of subquadrangular plates carrying
prominent teeth-bearing ridges. The dental formula is 4/4 and the lateral
teeth are longer than the median ones and often noticeably diverge
laterad. The teeth may be blunted, particularly the median ones,
presumably by wear. The jaws are brown; the teeth colorless.

The sperm funnels are narrow; not conspicuously set off from the vasa
efferentia by constrictions. The vasa deferentia are short and thick, enter-
ing the ental borders of the spermiducal gland at widely separated por-
tions of its ventral surface.

The spermiducal gland is short and thick and roughly subspherical. Its
ventral (ental) border lies just dorsal and to the side of the bursa; its dorsal
border extends to about the mid-portion of its segment. Its most dis-
tinctive feature, aside from shape, is the capaciousness of its lumen
(obscure in most branchiobdellids) which is expanded so that it appears
as a thin-walled sac filled with a clear fluid. Moreover, again unusual,
there are small amounts of spermatozoa clustered in the central portion.
The wall of the spermiducal gland is, however, composed of the usual ele-
ments (Holt 1949) of a peritoneal investiture, a thin muscular covering
and a lining of glandular epithelium.

The ejaculatory duct is a long and prominent tube composed of the
usual layers of muscle. The bursa is small and subspherical, in diameter
about 3/4 that of the segment (VI) in which it lies. There is a short ectal
(outlet), narrowed portion and the penis is a simple, protrusible, muscular
cone, exserted, one presumes, by the eversion of the bursa.

The spermatheca has a relatively long ectal duct and the cylindrical
bulb bends dorsad between the gut and the body wall. The curvature of
the organ precludes, in the absence of tedious and essentially impossible
procedures, any just estimate of its length.

Variation: — Beyond the usual differences in size and those produced by
differing degrees of contraction at death, there are few variations of note.
The relative lengths of the teeth seem to differ, but the lateral teeth are

28 Perry C. Holt and Ann M. Weigl

always at least as long as the median ones and usually longer. There is a
possibility that the dental formula may vary slightly, but it is constant for
all the specimens examined in which it could be determined.

Affinities'. — In the true sense of the word, until there is a monographic
revision of the genus, the affinities of X. bashaviae cannot be determined.
Xironodrilus formosus has a small spermiducal gland without the expanded
lumen of X bashaviae and lacks an ejaculatory duct. The dental formulas
also differ: that of X. formosus varies from 4/3 to 6/5 (Holt 1951).
Xironodrilus pulcherrimus (Moore 1894) has a dental formula of 3/3 with the
middle tooth shorter than the lateral ones and its spermiducal gland is
slender and lacks the expanded lumen of X. bashaviae. In addition,
specimens of X '. pulcherrimus (Moore 1894; Holt 1951) are somewhat larger
than any of those of X. bashaviae we have measured. Xironodrilus appalachius
Goodnight 1943 also has a dental formula of 3/3 with the middle tooth
longer than the lateral ones and a reproductive system similar to that of X.
pulcherrimus. Specimens of X. appalachius are of approximately the same size
as those of X. bashaviae (Holt 1951). Xironodrilus dentatus Goodnight 1940 is
characterized by a dental formula of 4/5, 5/5 or 5/4 and is recorded from
Oklahoma and Missouri. The jaw shape and other features of this species
also differ from X. bashaviae (Holt, unpub. data).

Host: — Cambarus bartonu (Fabricius 1798).

Distribution: — The type-locality, two sites in Bashavia Creek and one in
Fries Creek, are all part of the upper drainage of the Yadkin River in For-
syth County, North Carolina.

Material examined: — The type-series and 38 other specimens mounted en-
tire and portions of several serially sectioned animals.

LITERATURE CITED

Ellis, Max M. 1918. Branchiobdellid worms (Annelida) from Michigan crayfish.

Trans. Am. Microsc. Soc. 37(1):49-51.
1919. Branchiobdellid worms in the collections of the United States

National Museum, with descriptions of new genera and species. Proc. U.S.

Natl Mus. 55:241-265, pis. 10-13.
Franzen, A. 1963. Notes on the histology of Xironogiton instabilia (Moore, 1893)

(Family Branchiobdellidae) with special reference to the muscle cells. Zool.

Bidr. Upps. 35:369-384, pi. I-IV.
Goodnight, Clarence J. 1940. The Branchiobdellidae of North American

crayfishes. 111. Biol. Monogr. 77(3): 1-75.
1943. Report on a collection of branchiobdellids. J. Parasitol. 29:100-

102.

New Species Branchiobdellid 29

Hobbs, Horton H., Jr., P.C. Holt, and M. Walton. 1967. The crayfishes and their
epizootic ostracod and branchiobdellid associates of the Mountain Lake,
Virginia, region. Proc. U.S. Natl. Mus. 723:1-84.

Holt, Perry C. 1949. A comparative study of the reproductive systems of Xironogiton
instabilius instabilius (Moore) and Cambanncola philadelphica (Leidy) (An-
nelida, Oligochaeta, Branchiobdellidae). J. Morphol. #4(3):535-572.

1951. The genera Xironodnlus and Pterodnlus in North America with

notes on other North American genera of the family Branchiobdellidae
(Annelida, Oligochaeta). Ph.D. dissert., Univ. Virginia.

1960. The genus Ceratodnlus Hall (Branchiobdellidae, Oligochaeta),

with the description of a new species. Va. J. Sci. (N.S.) 77(2):53-77.

1965. On Ankyrodrilus , a new genus of branchiobdellid worms (An-
nelida). Va. J. Sci. (N.S.) 76(1)9-21.

1969. The relationships of the branchiobdellid fauna of the southern

Appalachians, pp. 191-219 in Holt, Perry C. (ed.). The distributional

history of the biota of the southern Appalachians, Part I: Invertebrates.

Res. Div. Monogr. 1, Va. Polytech. Inst., Blacksburg. 295 pp.
Moore, J. Percy. 1894. On some leechlike parasites of American crayfish. Proc.

Acad. Nat. Sci. Phila. 759:419-429, pi. XII.
1 895. The anatomy of Bdellodnlus illuminatus, an American discodrilid. J.

Morphol 7tf(2):497-540, pis. XXVIII-XXXII.

Accepted 14 November 1798

A New Cryptic Species of Salamander of

the Genus Plethodon from the

Southeastern United States

(Amphibia: Plethodontidae)

Richard Highton

Department of £00 logy, University of
Maryland, College Park, Maryland 20742

ABSTRACT. — A new species of woodland salamander, Plethodon
websteri, is described. It is a member of the P. welleri group of eastern
small plethodons and is morphologically very similar to P. dorsalis, but
differs from it at over 80% of 26 genetic loci as determined by elec-
trophoresis. This represents an extreme case of genetic divergence
without accompanying morphological change. The geographic range of
P. websteri includes east central Alabama and west central Georgia with
disjunct populations in Clarke County, Alabama; Winston County,
Mississippi; and McCormick County, South Carolina. The ranges of P.
dorsalis and P. websteri are largely allopatric, but the two have been found
sympatrically at one locality in Jefferson County, Alabama.

An electrophoretic study of genetic variation in salamanders of the
Plethodon welleri group (Larson and Highton 1978) showed that an un-
described species occurs in Mississippi, Alabama, Georgia and South
Carolina. Morphologically it is so similar to Plethodon dorsalis Cope that
there are no known standard taxonomic characters that distinguish the
two species. Yet genetically they are so different that they share less than
20% of their alleles at the 26 genetic loci evaluated electrophoretically.
Thus they represent a most extraordinary example of evolutionary
genetic divergence without accompanying morphological change.

There is no doubt of the specific distinctness of the two species. They
differ genetically from each other more than do some genera, for example
Notophthalmus and Tancha (Ayala 1975). They have been taken sym-
patrically in Jefferson County, Alabama, and no electrophoretic hybrids
were detected. The absence of diagnostic morphological characters re-
quires that the species be diagnosed exclusively on the basis of elec-
trophoretically detectable differences in the mobility of protein molecules.
This makes it difficult to identify living or preserved salamanders. For-
tunately, the ranges of the two species appear to be largely allopatric so
that most individuals may be identified on the basis of geographic
provenance. Moreover, in and near the zone of contact between the two

Brimleyana No. 1: 31-36. March 1979. 31

32 Richard Highton

species, there is character displacement in color morph frequency, so that
even in the zone of sympatry all available specimens that have been ex-
amined electrophoretically may be correctly allocated to species on the
basis of color morph.

I name the new species for my friend, the late T. Preston Webster, who
first called my attention to the remarkable amount of electrophoretic
divergence found in southern populations of Plethodon dorsalis representing
the new species.

Plethodon websteri, new species
Diagnosis: — An eastern small Plethodon of the P. welleri group (Highton
1962) that has virtually the same range of variation as P. dorsalis for all
known morphological taxonomic characters, but that differs from P. dor-
salis at most genetic loci evaluated electrophoretically. All samples of the
two species are completely separable (they do not share a single common
allele) at 14 of 26 genetic loci (Larson and Highton 1978): fumarase,
glutamic oxaloacetic transaminase- 1, indophenol oxidase-1, indophenol
oxidase-2, heart lactate dehydrogenase, muscle lactate dehydrogenase,
leucine aminopeptidase, malate dehydrogenase- 1, malate dehydrogenase-
2, peptidase-2, transferrin, and general proteins B, C and D. Most pop-
ulations of the two species also are distinct at 6 additional loci: esterase,
isocitrate dehydrogenase- 1, isocitrate dehydrogenase-2, phosphoglu-
comutase, phosphoglucose isomerase and general protein A. Good diag-
nostic loci are fumarase, indophenol oxidase-1, indophenol oxidase-2,
heart lactate dehydrogenase, malate dehydrogenase-2, and protein C (the
polypeptides of P. websteri migrate cathodally to those of P. dorsalis) and
leucine aminopeptidase, malate dehydrogenase- 1, peptidase-2 and
protein B (the polypeptides of P. websteri migrate anodally to those of P.
dorsalis). Plethodon websteri differs from P. welleri in the same morphologi-
cal ways as does P. dorsalis (it has modal number of 19 trunk vertebrae
compared to 17 in P. welleri; its belly is heavily mottled with red, white
and black chromatophores compared to the black belly with small white
spotting of P. welleri; and there is color pattern dimorphism in P. websteri:
a red or yellow striped dorsal pattern morph and a dark brown unstriped
morph in P. websteri compared to an unstriped brown dorsal color pattern
heavily mottled with brassy flecking in P. welleri). The electrophoretic
data and their genetic analysis are presented in Larson and Highton
(1978).

Holotype: — NMNH 204814, an adult male collected 0.6 km east, 0.9 km
south of Howelton, Etowah County, Alabama, on 7 January 1976 by
Scott Bunting, Richard Highton, Mark Kielek and Allan Larson.

New Species Plethodon

33

®

Fig. 1. Electrophoretic localities of P. dorsalis (hollow circles) and P. websten
(solid circles) in the southeastern United States. Approximate limits of the ranges
of the two species are indicated. The locality in Jefferson County, Alabama where
the two species are sympatric is indicated by a half solid circle.

Paratypes: — NMNH 204815-34, topotypes.

Other material: — Living individuals have been examined from all localities
listed in table 1 and preserved specimens from most localities will be
depositied in the collection of the National Museum of Natural History

(NMNH).

Description of Holotype: — Before preservation, the length from the tip of
the snout to the anterior angle of the vent was 39 mm, to the posterior
angle of the vent 41 mm, and the total length 78 mm. There are 18 costal
grooves (equivalent to 19 trunk vertebrae) and the vomerine teeth number
6 on each side. A red dorsal stripe with irregular edges was present in life;
it is wider and brighter red on the tail than on the dorsum. The legs and
sides were brown with abundant yellow and red spots and brassy flecks.

34

Richard Highton

Table 1. Localities and number of P. dorsalis and P. websteh identified electrophoretically.

No

th Latitude

West Longitude

Spe

aes State

County

A'

°

"

o

"

P. dot

salts Alabama

Blount

1

34

09

03

86

27

18

"

Cullman

1

34

10

22

86

53

31

"

DeKalb

3

34

23

38

85

37

38

"

Fayette

7

33

45

43

87

45

05

"

Jefferson

7

33

43

33

86

49

15

"

Jefferson

4

33

46

55

86

49

10

"

Lawrence

48

34

18

25

87

20

10

"

Tuscaloosa

2

33

26

38

87

29

47

"

Walker

7

33

42

20

87

23

22

"

Winston

3

34

03

30

87

20

45

Arkansas

Independence

23

35

52

30

91

46

26

"

Pope

16

35

38

28

93

04

03

, „

Stone

45

35

59

05

92

16

02

Georgia

Dade

16

34

52

02

85

31

58

Illinois

Pope

61

37

22

54

88

40

20

"

Union

46

37

32

43

89

26

14

Indiana

Crawford

15

38

16

35

86

32

10

"

Parke

61

39

53

14

87

11

20

Kentucky

Franklin

55

38

11

24

84

52

53

> )>

McCreary

2

36

52

15

84

21

55

Mississippi

Tishomingo

15

34

36

23

88

10

32

Missouri

Taney

4

36

40

14

93

18

37

Oklahoma

Adair

21

35

50

13

94

39

20

, >>

Cherokee

1

35

58

03

94

48

55

> >j

Sequoyah

34

35

34

47

94

31

20

"

"

2

35

37

47

94

34

50

Tennessee

Bledsoe

32

35

38

32

85

19

55

'

Blount

9

35

38

20

83

44

51

'

"

36

35

39

56

83

47

04

> "

Montgomery

34

36

31

00

87

30

35

'

Moore

7

35

20

55

86

20

30

' "

Washington

6

36

10

32

82

31

17

Virginia

Scott

3

36

38

05

82

26

52

' "

"

15

36

37

50

82

35

22

P. we

bsieri Alabama

Blount

17

34

05

12

86

20

57

' "

"

7

34

08

03

86

23

09

'

Clarke

66

31

32

55

87

55

48

'

Cleburne

37

33

29

15

85

47

28

> >)

Etowah1

64

34

02

51

86

10

38

'

"

51

34

04

06

86

18

43

> 11

Jefferson

4

33

43

33

86

49

15

5 11

Lee

36

32

36

17

85

17

57

,

Shelby

9

33

21

37

86

28

38

, ii

"

1

33

22

03

86

39

49

Georgia

Cobb

28

33

58

34

84

34

56

>' i)

Upson

91

32

47

38

84

15

30

Mississippi

Winston

30

33

09

10

89

02

50

South Carolina

McCormick

81

33

41

20

82

09

15

'

"

8

33

43

48

82

11

02

'Type locality of P. websteri.

New Species Plethodon 35

The chin and belly had red, white, and black pigment in the following
proportions: chin 50:40:10, belly 30:60:10. It is a mature male with a
rounded mental gland as in P. dorsalu and P. welleri (Highton 1962:fig.
2D).

Distribution: — P. websten is known from east central Alabama and west
central Georgia. Apparently disjunct populations occur in Clarke
County, Alabama, Winston County, Mississippi, and McCormick
County, South Carolina (figure 1).

Variation in P. websteri: — The modal number of trunk vertebrae in all
known populations of P. dorsalu is 19. This is also true for P. websteri with
the exception of the two samples from McCormick County, South
Carolina. Both have slightly more individuals with 20 trunk vertebrae
than they do with 19 (mean = 19.6 in the more southern sample and 19.8
in the more northern sample).

As in P. dorsalis, most samples of P. websteri include salamanders of both
the red striped and unstriped morphs, and individuals intermediate be-
tween the two. Because of the difficulty of classifying intermediate in-
dividuals, it is impossible to objectively quantify the data on the frequen-
cies of the color morphs. This is in contrast to several other species of the
genus in which few or no intermediates between the two color morphs oc-
cur. In spite of the difficulty in classifying a few individuals, most P.
websteri from the immediate vicinity of the zone of contact between P.
websteri and P. dorsalis in Blount and Etowah Counties, Alabama are of
the red striped morph, while P. dorsalis from nearby Lawrence, Cullman
and Blount Counties are of the unstriped morph. At the locality where
the two species were taken sympatrically in Jefferson County, Alabama, 8
animals of each morph were collected. Of the 1 1 that were examined elec-
trophoretically, all 4 P. websteri are of the striped morph and all 7 P. dor-
salis are of the unstriped morph. No intermediates are present and no
genetic hybrids were found. This same kind of character displacement in
color morph frequencies has been reported for contact zones of several
other pairs of species of eastern small Plethodon (Highton 1962, 1972).

Remarks: — A photograph of a living individual from Lee County,
Alabama appears in Mount (1975:133) and is presumably P. websteri
since the locality is within the range of the species.

Although the holotype of P. websteri has not been subjected to elec-
trophoretic analysis, there is little doubt that it belongs to this species.
Sixty-four other salamanders from the type locality have been studied
electrophoretically and all are P. websteri.

Additional studies of the zone of contact and/or overlap of the ranges of
the two species in Alabama are needed to determine the nature of their

36 Richard Highton

geographic and ecological interactions as well as the details of the color
morph character displacement in that area.

No morphological differences in standard taxonomic characters used to
distinguish species group taxa in the genus Plethodon have been detected in
this pair of sibling species, but this does not preclude the possibility that a
multivariate morphometric analysis might detect differential characters.
Such a study would be especially interesting because of the very large
amount of genetic divergence between P. dorsalis and P. websten.

ACKNOWLEDGMENTS. — I wish to thank all those persons who helped
with the field work, including several who provided information on
localities, as well as those who aided in the laboratory work, especially
Allan Larson and the late T. Preston Webster. The work was supported
by the National Science Foundation (grants DEB 76-10583 and GB-
37320).

LITERATURE CITED

Ayala, Francisco J. 1975. Genetic differentiation during the speciation process.

Evol. Biol. 5:1-78.
Highton, Richard. 1962. Revision of North American salamanders of the genus

Plethodon. Bull. Fla. State Mus. Biol. Sci. 6:235-367.
1972. Distributional interactions among eastern North American

salamanders of the genus Plethodon. pp. 139-188 in Holt, P.C. (ed.). The

distributional history of the biota of the southern Appalachians, Part III:

Vertebrates. Res. Div. Monogr. 4, Va. Polytech. Inst. State Univ.,

Blacksburg. 306 pp.
Larson, Allan, and R. Highton. 1978. Geographic protein variation and

divergence in the salamanders of the Plethodon wellen group (Amphibia:

Plethodontidae). Syst. Zool. 27:431-448.
Mount, Robert H. 1975. The reptiles and amphibians of Alabama. Auburn Univ.

Agric. Exp. Stn, Auburn. 347 pp.

Accepted 27 November 1978

The Female Reproductive Cycle in North Florida
Kinosternon bauni (Testudines: Kinosternidae)

John B. Iverson1

Florida State Museum and Department of apology,
University of Florida, Gainesville, Florida 32611

ABSTRACT. — Female striped mud turtles are generally active
throughout the year, with maximum activity in March and October and
minimum in February and July. Females mature at an age of 5 to 6
years and a plastron length of 70 to 75 mm.

Vitellogenesis is continuous from July through the following May.
Ovulation and oviposition occur from at least September to June. At
least three clutches are produced annually. Egg size averages 28 X 17
mm and is not correlated with female size or clutch size. The mean
length of laboratory incubation is 119 days and hatchlings average 19.2
mm PL. Clutch size ranges from one to five (usually two or three) and is
positively correlated with female size. Each clutch comprises about 8
percent of total body weight.

INTRODUCTION

Few reproductive studies exist for mud turtles (genus Kinosternon) other
than Kinosternon subrubrum (Mahmoud and Klicka 1972; Gibbons 1975;
Iverson, 1979) and Kinosternon flavescens (Mahmoud and Klicka 1972;
Christiansen and Dunham 1972). Studies by Sexton (1960) of K. scor-
pioides and by Moll and Legler (1971) of K. leucostomum comprise the only
other detailed studies. Most Kinosternon reproductive information is anec-
dotal.

The present knowledge of reproduction in K. bauni is based almost en-
tirely on Einem's (1956) and Lardie's (1975) observations in central
Florida. The purpose of this report is to provide more complete informa-
tion on the female reproductive cycle of striped mud turtles, Kinosternon
bauni from northern Florida. This study, with others now in progress by
this author, should soon permit an analysis of reproductive strategies
within the genus.

MATERIALS AND METHODS

Female turtles were collected whenever possible in Alachua, Levy, and
Marion counties in north Florida (usually within 50 km of Gainesville)

Present address: Department of Biology and Joseph Moore Museum, Earlham
College, Richmond, IN 47374.

Brimleyana No. 1: 37-46. March 1979. 37

38 John B. Iverson

from January 1972 through December 1976. Due to the proclivity of
adults and hatchlings to terrestrial activity in north Florida (Carr 1952),
this species is more frequently encountered on land than any other local
turtle. Most turtles were collected as they attempted to cross roads. Many
of these were dead on the road, but remained intact enough to provide
suitable specimens for measurement and dissection. Turtles were also ob-
tained by trapping, seining, and diving. In addition, specimens from
north Florida in the Florida State Museum (University of Florida) collec-
tion were included in the samples.

Ovarian follicles, oviducal eggs, and corpora lutea were examined and
measured in each specimen. All straight-line measurements were taken
with dial calipers to the nearest 0.1 mm. Carapace length (CL) and
plastron length (PL) were also recorded. Measurements from preserved
turtles did not significantly differ from the data collected from fresh
specimens and were included in the analysis.

Several clutches of shelled oviducal eggs were removed and incubated
to hatching at 27° to 29 °C. Data from eggs in natural nests were also
recorded. Most turtles were deposited in the Florida State Museum, Uni-
versity of Florida, but a representative series was retained by the author.
Means are followed by ± one standard deviation. All measurements are
in mm.

SEASONAL ACTIVITY

Striped mud turtles were collected throughout the year, both on land
and in water. Individuals may be found active on all but a few of the
coldest winter days. This species thus exhibits the longest annual activity
cycle of any previously studied kinosternid in the United States
(Christiansen and Dunham 1972).

Annual terrestrial activity seems to be bimodal. Striped mud turtles are
most frequently encountered (especially on land) in March when spring
rains are filling the ponds and marshes, and in October as water levels
are dropping. Few turtles were collected during late summer when water
levels and temperatures are maximal, or in mid-winter (February) when
temperatures are minimal. Wygoda (1976) found a similar annual ac-
tivity pattern in K. bauni inhabiting seasonally flooded hardwood swamps
in central Florida.

Since annual activity is bimodal, two (or more) epidermal scute annuli
are often produced by an individual turtle during any one year. Winter
annuli are apparently always produced and are usually more distinct
than those produced in the summer. This variation in annuli production
often makes aging turtles, based on scute annuli, difficult.

Kinosternon Reproductive Cycle 39

AGE, GROWTH, AND SIZE AT MATURITY

Females possessing ovarian follicles exceeding 7 mm diameter (or
oviducal eggs or corpora lutea) during the months of July through May
were considered mature. The three smallest mature females had plastral
lengths of 69.4, 70.2 and 72.7. The four largest females judged to be im-
mature measured 66.6, 67.9, 74.8, and 79.6 PL. Maturity therefore occurs
at sizes between 70 and 75 PL (75 to 80 CL). I believe the 79.6 PL im-
mature was reproductively anomalous since nine other females with
plastron lengths between 75 and 80 mm were mature. The smallest
mature female dissected by Einem (1956) was 81.2 mm CL and the
largest immature female was 71.1 mm CL.

Age at maturity was estimated by counts of clearly visible winter (or
primary) abdominal scute annuli as described by Sexton (1959). The
oldest immature females bore six clear primary annuli (74.8 and 79.6
PL); the youngest mature female had only five (73.1 PL), indicating the
usual age at maturity to be five or six years.

Plastron lengths calculated from the abdominal scute annuli lengths of
19 juvenile and young adults in the manner of Ernst et al. (1973) in-
dicated turtles in their first winter averaged 17.9 PL (Range = 14.4-20.6,
N = 19). Those in their second winter averaged 32.7 PL (25.5-39.0, N =
19); in their third winter, 45.7 (34.3-56.8, N = 14); fourth, 57.8 (49.3-
64.3, N = 11); fifth, 68.9 (62.8-76.9, N = 9); sixth, 74.3 (69.1-82.0, N =
8); and in their seventh, 75.1 (71.1-83.8, N = 7). If turtles are maturing at
plastral lengths of 70-75, these data support the contention that maturity
occurs during the fifth to sixth years of age.

Average size of 101 adult females previously examined from throughout
Florida (Iverson 1978b) was 86.2 PL (91.6 CL) [52 males averaged 73.1
PL, 83.7 CL]. The largest female measured 105.1 PL and 114.7 CL. The
PL-body weight (in gm) regression, based on six females (69.1-90.0 mm
PL, and 64.5-143.1 gm), is Wt - 3.36 PL - 154.84 (r = 0.92; p<0.01).
From this regression the average female (86.2 PL) weighs 135.1 gm.

FEMALE REPRODUCTIVE CYCLE

The ovarian cycle is nearly continuous (Table 1), with only a short
summer quiescent period (coincident with the summer reduction in ac-
tivity). Ovulation occurs from late August or early September to early
June. Based on excavated nests and the presence of oviducal eggs, females
apparently nest from September through June. Females continue to yolk
follicles to replace those ovulated during all but the last of this period.
Follicular enlargement is curtailed only from late May through June.
During the remainder of the year enlarged follicles (> 7 mm) are

40 Jonn B. Iverson

typically found in the female reproductive tract along with oviducal eggs
and/or corpora lutea.

Ova were removed from oviducal eggs to determine their approximate
size at ovulation. Twenty-two excised yolk masses averaged 16. 24*1. 06
(Range = 14.5-18.0) in diameter. Maximum diameter of an ovarian folli-
cle was 17.5. Only four females had ovarian follicles exceeding 16 mm in
diameter.

The ovaries of one dissected female (98.8 PL), collected 22 April 1972,
were anomalous in that her ovaries bore no corpora lutea or follicles > 4
mm in diameter.

Of 50 females with oviducal eggs and corpora lutea, 16 exhibited
evidence of transuterine migration of ova. Net migration was away from
the tract with the larger ovary in 11 (68.8%) of the cases, and probably
served to equalize reproductive tract volumes.

The length of time that eggs were retained in the oviducts is not known.
No females with oviducal eggs had the corresponding corpora lutea in
any state of regression; all appeared fresh (maximum corpora lutea
diameter is 6-7 mm). A physiological mechanism such as that suggested
by Moll and Legler (1971) may allow female K. baurii to retain their eggs
until suitable nesting sites and conditions can be found, without
possibility of subsequent ovulation.

Table 1 . Percentage of mature female Kinosternon baurii bearing enlarged follicles
(> 7.0 mm), oviducal eggs, and corpora lutea for each month.

% with

% with

Sample

Enlarged

Oviducal

% with

Size

Follicles

Eggs

Corpora lutea

9

100

77.8

100a

2

100

50

50

25

100

56

92

9

88.9

44.4

55. 5a

15

73.3

53.3

73.3

7

57.1

28.6

42. 9a

1

100

o

100

4

100

25

25

4

100

100

100

9

100

88.9

100a

3

100

100

100

7

100

100

100

Month

January . .
February .
March . . .

April

May

June

July

August . . .
September
October . .
November
December

includes one female with two sets of corpora lutea.

Kinosternon Reproductive Cycle

41

5-

4-

M 3

CO

3 2
U

23

24

25

26 27 2(

EGG LENGTH

29

30

Fig. 1. Relationship between clutch size and egg size (mean length for clutch in mm) in
Kinosternon baurii. Each symbol represents at least one record. Least squares regression is y =
-0.08X + 4.83; r = -0.16, p>0.5 (N = 49).

110

100-

O

z

^ 90-

z
o

80-

70

V . •

22

23

24

25

26 27 28 29

EGG LENGTH

30

31

32

Fig. 2. Relationship between body size (PL in mm) and egg size (mean length for clutch
in mm) in Kinosternon baurii. Each symbol represents at least one record. Least squares
regression is y = 0.36X + 76.67; r = 0.093, p>0.05 (N = 38).

42

John B. Iverson

Fig. 3. Typical mid-incubation crack in egg of Kinostemon baurii after 103 days incubation.
Oviducal egg was removed from female 26 September 1976, crack first appeared 74 days
later. Egg hatched after 118 days incubation. X marks top of egg.

Natural nests with eggs in various stages of incubation were found on
the following dates: 8 March, 25 April, 7 May, and 20 May 1972; and 19
May (2), and 25 May (2) 1973. Eggs were found buried under 1-3 cm of
sand (5 cases) or under moist leaf litter 1 cm or less in depth. Two captive
turtles oviposited on 8 February and 17 February 1972, the former in
sand and the latter in the water of its tank. Carr (1952) reported finding
natural nests from April to June in sand and in piles of dead hyacinths.
Captive nestings were observed on 7 March, 9 June, and 19 November by
Lardie (1973), and on 4 June by Nicol (1970).

EGGS, INCUBATION, AND HATCHING

The eggs of Kinostemon baurii have been described by Einem (1956) and
present observations do not differ from his. Mean egg size for 114 eggs I
examined was 27.55 (longest diameter; Range = 22.8-32.8) by 16.63
(shortest diameter; Range = 13.6-19.3). Shells of 2 eggs (28.6X16.7;
28.6X16.8) were 0.23-0.28 mm thick at their ends (10 measurements) and
0.34-0.40 thick along the perimeter of their minor axes (10 measure-
ments). Eight eggs (X length 26.6) averaged 4.45 gm each (3.22-5.26).

Kinostemon Reproductive Cycle

43

22-

•

21-

•

X 20-

t—

O

z

^ 19-

•

•
•y^

•

y^ •

•

§ 18-

\—

CO

<

^ 17-

y/ •

•

16-

yS «

•

15

22

23

24

25 26 27 28

EGG LENGTH

29

30

Fig. 4. Relationship between hatchling size (PL in mm) and egg length (mm) in Kinoster-
non baurii. Least squares regression is y = 0.611 X + 1.846; r = 0.813, p<0.01 (N = 14).

4-

£ 3
to

X

a

3 2

u

1-

•• • •

70

80 90

PLASTRON LENGTH

100

no

Fig. 5. Relationship between clutch size (based on counts of corpora lutea) and body size
in Kinostemon baurii. Each symbol represents at least one record. Least squares regression is y
= 0.048 X -1.630; r = 0.442, p<0.01 (N = 60).

44 John B. Iverson

Seventeen oviducal eggs from central Florida K. baura measured by
Einem (1956) averaged 28.0 (Range = 25.0-31.8) by 16.6 mm (15.8-
17.2). Lardie (1975) recorded a single central Florida egg of 28X16, and
Nicol (1970) measured six south Florida eggs which averaged 23.75 (21-
27) by 14.0 (12.5-16). Egg length was not significantly correlated with
clutch size (Figure 1) or female PL (Figure 2). No seasonal trends in egg
size were identifiable.

Fourteen clutches of oviducal eggs were incubated. Three clutches were
opened and preserved after 88, 109, and 114 days, respectively. The
remainder were incubated to hatching. At least some of the eggs of all
clutches exhibited mid-incubation cracking of the egg shell (Figure 3) as
described for Kinosternon bauru by Einem (1956) and for Sternotherus minor
by Iverson (1978a). Not all eggs in any one clutch developed these cracks.
Crack formation occurred after an average incubation period of
78.8±16.7 days (Range = 58-113; N = 9 eggs).

Mean incubation time to hatching for 11 clutches was 11 8.8 ±11.3 days
(Range = 97-143). Eggs from any single clutch hatched within 24 hours
of each other, except for one clutch in which 9 days separated the
hatching of the 2 eggs. Five eggs incubated by Einem (1956) hatched after
96 to 129 days. Nicol (1970) hatched 4 eggs from the same clutch after 91,
93, 102, and 107 days, respectively. Three eggs from a clutch laid 9 June
1969 were hatched by Lardie (1975) after 117-119 days. Clutches hatched
in my laboratory in January (1), February (1), March (3), April (1), June
(1), August (2), and September (1). The hatching process was as
described by Einem (1956).

Twenty-eight captive hatchlings averaged 19.2 PL (Range = 15.45-
22.0) and 22.5 CL (Range = 17.55-25.0). Plastron length of hatchlings
was significantly positively correlated (r = 0.813) with egg size (Figure
4). Hatchling-sized turtles were collected in the field on 5 January, 27
January (8), 28 January (5), 4 March (4), 9 March (2), 7 August, 15 Sep-
tember, and 7 December. Most of these had incompletely or very recently
closed umbilical openings. Average size of 13 of these neonates was 19.6
PL (Range = 16.55-22.0) and 22.4 CL (Range = 20.4-25.0). Hatchlings
from three eggs incubated by Lardie (1975) each measured 19 PL (22
CL); five hatched by Einem (1954) measured 18.1-20.3 PL (20.5-25.0
CL); and four hatched by Nicol (1970) averaged 17.5 CL (15.0-18.5).

CLUTCH SIZE

Clutch size in Kinosternon bauru ranged from one to five, two or three be-
ing the usual complement. Mean clutch size was insignificantly different
whether estimated by counts of enlarged follicles over 10 mm diameter
(2.69±0.85; N = 48), oviducal eggs (2.60±0.96; N = 50), or corpora

Kinosternon Reproductive Cycle 45

lutea (2.50±0.87; N = 82). Twelve clutches examined by Einem (1956)
and Lardie (1975) averaged 2.33 (Range = 1-3). Nicol's (1970) record of
a 6-egg clutch probably represents the maximum for the species. As
Einem's (1956) data suggest, clutch size is positively correlated (p <
0.01) with plastron length and increases an average of one egg for each
20.8 mm increase in PL (Figure 5). No seasonal trends in clutch size were
identifiable.

ANNUAL REPRODUCTIVE POTENTIAL
The ovaries of four females bore two distinct sets of corpora lutea.
Clutch sizes of these turtles were 3 (larger set of corpora lutea) and 2 (6
October; 95.4 PL), 2 and 2 (5 January; 101.3 PL), 1 and 1 (20 April; 77.8
PL), and 5 and 4 (20 June; 95.4 PL). All but the last of these also had a
set of pre-ovulatory follicles > 10 mm diameter. The ovaries of the first
female suggest that following the first clutch of the reproductive season
(late August to early September ?), she might possibly nest again in Oc-
tober. If her indicated inter-nest period of about 2 months were main-
tained, 6 clutches could be produced annually. This relatively long inter-
nest interval presumably allows for nearly complete luteal regression bet-
ween ovulations and may explain why only 4 of 84 females had more than
one set of identifiable corpora lutea.

Mature females certainly produce at least three clutches each year.
Einem (1956) also suspected that three clutches might be produced an-
nually, and Lardie (1975) reported a captive central Florida female (115
mm CL) which produced three clutches of three eggs each on 7 March, 9
June, and 16 November 1969. If each clutch averages 2.5, average annual
reproductive potential is at least 7.5. Field studies will be necessary to
determine the actual number of annual clutches.

Reproductive effort per clutch was estimated by the ratio of mean
clutch weight (mean egg weight X mean clutch size) to mean total female
weight. Clutches in Kinosternon baurn average 8.23 per cent (4.45X2.50X
100/135.1) of female weight. Unfortunately, similar estimates are
available for no other kinosternid turtle.

ACKNOWLEDGMENTS. — I thank Dr. Walter Auffenberg for making
the Florida State Museum facilities available to me. Thanks are also due
Diderot Gicca, Howard Kochman, Peter Meylan, and Brick Rainey for
able field assistance. Numerous other people diligently retrieved
specimens killed while crossing roads. Dale R. Jackson offered valuable
comments on an early draft of the manuscript. My wife, Sheila, typed the
manuscript.

46 John B. Iverson

LITERATURE CITED

Carr, Archie F. 1952 Handbook of turtles: the turtles of the United States,

Canada, and Baja California. Cornell Univ. Press, Ithaca. 542 pp.
Christiansen, James L., and A.E. Dunham. 1972. Reproduction of the yellow

mud turtles (Kinosternon flavescens flavescens) in New Mexico. Herpetologica

28 (2): 130- 137.
Einem, G.E. 1956. Certain aspects of the natural history of the mud turtle,

Kinosternon bauni. Copeia 1956(3): 186- 188.
Ernst, Carl H., R.W. Barbour, E.M. Ernst and J.R. Butler. 1973. Growth of the

mud turtle, Kinosternon subrubrum, in Florida. Herpetologica 29(3):247-250.
Gibbons, J. Whitfield. 1975. Reproductive strategies of the mud turtle {Kinosternon

subrubrum). Paper presented at annual ASIH meeting, Williamsburg,

Virginia.
Iverson, John B. 1978a. Reproductive cycle of female loggerhead musk turtles

(Sternotherus minor minor) in Florida. Herpetologica J4(l):33-39.
1978b. Variation in striped mud turtles, Kinosternon bauni (Reptilia,

Testudines, Kinosternidae). J. Herpetol. 72(2) : 135-142.
1979. Reproduction and growth of the mud turtle, Kinosternon subrubrum,

in Arkansas. J. Herpetol. 73(1):105-111.
Lardie, Richard L. 1975. Observations on reproduction in Kinosternon. J. Herpetol.

9(2):257-260.
Mahmoud, I.Y., and J. Klicka. 1972. Seasonal gonadal changes in kinosternid

turtles. J. Herpetol. 6(3/4 ): 183-1 89.
Moll, Edward O., and J.M. Legler. 1971. The life history of a neotropical slider

turtle, Pseudemys scnpta (Schoepff) in Panama. Bull. Los Ang. Co. Mus.

Nat. Hist. Sci. 77:1-102.
Nicol, R. 1970. Striped mud turtle has tiny offspring. Tortuga Gazette 6(2) :4.
Sexton, Owen J. 1959. A method of estimating the age of painted turtles for use in

demographic studies. Ecology 40:716-718.
1960. Notas sobre la reproduccion de una tortuga Venezolana, la

Kinosternon scorpwides. Mem. Soc. Cienc. Nat. La Salle 20(57):189-197.
Wygoda, Mark. L. 1976. Terrestrial activity of the striped mud turtle, Kinosternon

bauni, in a seasonal hardwood swamp forest. Master's thesis. Univ. South

Florida, Tampa. 61 pp.

Accepted 25 June 1978

Fishes of the Gauley River, West Virginia

Charles H. Hocutt

Appalachian Environmental Laboratory,
University of Maryland, Frostburg, Maryland 21532

Robert F. Denoncourt

Department of Biology, York College of
Pennsylvania, York, Pennsylvania 17405

AND

Jay R. Stauffer, Jr.

Appalachian Environmental Laboratory,
University of Maryland, Frostburg, Maryland 21532

ABSTRACT. — The confluence of Gauley River and New River forms
the Kanawha River a short distance above Kanawha Falls, West
Virginia. A survey of fishes of Gauley River in 1976 yielded 50 species,
25 of which were not previously reported in the literature. Six species
{Lampetra aepyptera, Moxostoma erythrurum, Ictalurus natalis, Ictalurus
nebulosus, Noturus flavus, and Percina caprodes) established new distribution
records above Kanawha Falls, which is generally recognized as a major
barrier to fish dispersal. Additional verified records increased the total
known ichthyofauna to 58 species. These data suggest that fishes which
successfully negotiated Kanawha Falls may have found the Gauley
River a less strenuous route than the New River for upstream dispersal.
Gauley River fauna also may have been influenced by stream captures
with Greenbrier and Elk rivers.

INTRODUCTION

Gauley River rises in Webster and Pocahontas counties, West Virginia,
and flows west-southwest to Gauley Bridge where it joins New River to
form Kanawha River (Fig. 1). The main-channel Gauley is 168 km long,
occupies a drainage basin of 3497 km2 and has an average gradient of 6.1
m/km (Reed 1974). Its headwaters are characterized by broad, meander-
ing, low gradient streams draining the Plateau; the lower section is well
known for long rapids, cataracts, large boulders and a deep, narrow V-
shaped valley bordered by sandstone cliffs (Reger 1920).

Gauley River apparently retains the old channel it developed on a
peneplain during the Cretaceous (Reger 1921), as evidenced by
numerous ancient meanders representative of an old base-level bed (Hen-
nen 1919). The length of the river, 168 km, as compared to the airline dis-
tance, 95 km (Reed 1974), is indicative of the amount of meandering.

Brimleyana No. 1: 47-80. March 1979. 47

48 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Subsequent uplift of the area during the late Tertiary (Reger 1920)
revived the parent stream and caused rapid cutting which resulted in for-
mation of a great gorge, often incised 150 m or more into the Plateau. The
presence of a V-shaped lower valley, rather than U-shaped, indicates that
the river has not progressed far into its erosional cycle. Many tributaries
approach base-level maturation in their heads as well, but have
significantly increased gradients in their lower sections indicative of
Plateau uplift and stream rejuvenation (Reger 1920). For instance,
Meadow River, a major southern tributary, drops over 207 m in the last
18.5 km with an average gradient of 11.4 m/km (Reger 1921).

The only previous systematic survey of the fishes of the Gauley River
was that of Addair (1944), who reported on 24 species. C.L. Hubbs, E.C.
Raney, and F.J. Schwartz made occasional collections in the drainage,
but did not publish their data. Reed's (1974) discussion of the fishery of a
portion of the drainage was limited primarily to game species. Jenkins et
al. (1972) reported some Gauley River records, but did not discuss the
fauna in detail. Ross and Perkins (1959) and Ross (1959) discussed fishes
of the New (upper Kanawha) River, but data presented by Jenkins et al.
(1972) are more recent.

Zoogeographically, the Gauley River ichthyofauna is identified with
the New River drainage (Jenkins et al. 1972), i.e., that portion of the
Kanawha River drainage above the 7.3 m high Kanawha Falls (Denon-
court et al. 1975), which has been considered as a major barrier affecting
upstream dispersal of fishes (Jenkins et al. 1972; Lachner and Jenkins
1971). Endemism is reportedly high in the upper Kanawha (New/ Gau-
ley) River drainage (Hocutt et al. 1978), with the following fishes occur-
ring nowhere else: Nocomis platyrhynchus , Notropis scabriceps, Phenacobius
teretulus, Etheostoma kanawhae and E. osburni. Coitus carolinae ssp. (Robins
1954), long thought to be a New River endemic, is also known from one
spring in Jefferson County, Tennessee (Etnier, pers. comm). Exoglossum
laurae and Percina oxyrhyncha, species associated with the unique New River
fauna, have wider distributions than once thought (Jenkins et al. 1972;
Hocutt et al. 1978; Hocutt, in press). The form previously recognized as
Percina maculata in the upper Kanawha drainage is another endemic
species (E. Beckham, pers. comm.).

Various localities within the Gauley River system have been proposed
by the Corps of Engineers, Huntington, W. Va., District, as potential
sites for location of hydroelectric facilities. Among these sites is a location
on the main-channel Gauley River at Swiss that would inundate much of
the lower gorge. Collison Creek and Muddlety Creek are among the alter-
nate sites. The U.S. Fish and Wildlife Service, aware of our efforts to sur-
vey the streams of West Virginia (Hambrick et al. 1973; Denoncourt et al.

Gauley River Fishes 49

1975; Stauffer et al. 1975; Denoncourt et al. 1977; Stauffer et al. 1977;
Hocutt et al. 1978; Stauffer et al. 1977; and Hocutt et al. 1977), contract-
ed this investigation to serve as a basis for their position regarding the
proposed Corps' projects on Gauley River.

METHODS AND MATERIALS

Gauley River is rather isolated and offers several distinctive problems
in sampling for fishes. Preliminary planning indicated a need to use a
variety of collecting gear. Streams were sampled primarily with 1.5 X 3.0
m nylon seines with 3.2 mm mesh or with a pulsated DC electrofishing
unit. In more open waters and pools a 1.5 X 7.6 m nylon seine with 3.2
mm mesh was often employed. The electrofishing unit was used ex-
clusively in lower stream sections characterized by large rubble, boulders
and high gradient. Four localities in the drainage were sampled with
emulsified rotenone using techniques recommended by Hocutt et al.
(1973). Trotlines and gill nets were fished overnight in some of the larger
pools of Gauley River; these techniques were ineffective due to water
clarity and depauperate fauna, so the data were combined with seine data
for the particular localities.

A total of 52 stations was sampled in the system (Table 1, Fig. 1), with
the expressed purpose of obtaining a representative qualitative sample
(Hocutt et al. 1974) at each station. Stations were sampled a single time,
except for Station 30 which was sampled by seine and by electrofishing;
for the purposes of this report, data obtained in these two collections were
combined. Table 2 lists each fish species collected by station. Data are
organized for discussion by main-channel and its tributaries. Subse-
quently, an annotated list of species collected in this survey and by Ad-
dair (1944) and Reed (1974) is presented.

All specimens were preserved in a 10 percent formalin solution, unless
collected by rotenone. Rotenone collections were preserved in 20 percent
formalin (Hocutt et al. 1973). All collections were catalogued into the
Fish Museum, Appalachian Environmental Laboratory (AEL 142-194,
226), and stored in 40 percent isopropanol.

Museum records of Gauley River specimens were verified where possi-
ble. Museums housing collections from the system include: Cornell Uni-
versity (CU); Ohio State University (OSU); University of North Caro-
lina (UNC); U. S. National Museum (USNM); University of Michigan,
Museum of Zoology (UMMZ), where Addair's (1944) collections are
catalogued; and Virginia Polytechnic Institute and State University
(VPISU).

Jenkins et al. (1972) defined drainages, systems, and basins, and their
classification is followed here. Their suggestion that use of these terms be
standardized has merit, but certain ambiguities remain. Critical com-
ment is reserved for discussion elsewhere.

50 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Fig. 1 . Map of Gauley River drainage, West Virginia, with fish sampling localities noted.

Gauley River Fishes 51

Table 1. Fish sampling localities on the Gauley River, West Virginia. Ap-
palachian Environmental Laboratory (AEL) catalogue number, date of collection
and time of collection are in parentheses.

Station No. Locality Description

1 Headwaters of Gauley River, mouth of Big Run, Webster Co. (AEL 165;
6/9/76; 1100).

2 Gauley River, old tipple at Jerrysville, Webster Co., WV (AEL 167;
6/9/76; 1245).

3 Gauley River, 3.2 km s on Route 20 from junction Co. Rd. 42, Bolair,
Webster Co. (AEL 180; 6/9/76; 1445).

4 Gauley River at Gauley Mills, Webster Co. (AEL 179; 6/9/76; 1630).

5 Confluence of Gauley and Cherry rivers, Route 20 bridge, Nicholas Co.
(AEL 181; 6/11/76; 1130).

6 Gauley River, at mouth of Meadow River, Carnifex Ferry, Fayette Co.
(AEL 187; 7/6/76; 0900).

7 Gauley River, secondary road paralleling Peter's Creek, then downstream
along Gauley for ca. 8.0 km, Nicholas Co. (AEL 188; 8/2/76; 1000).

8 Gauley River, mouth of Laurel Creek, Swiss, Nicholas Co. (AEL 185;
8/26/76; 1300).

9 Gauley River, first riffles above mouth, midway between Vanetta and
Gamoca, Nicholas Co. (AEL 147; 8/9/76; 1430).

10 Williams River, Williams River Road, Williams River Campground, 4.8
km from Handley, Pocahontas Co. (AEL 186; 6/11/76; 0900).

11 Williams River, Scenic Route 150 bridge, Pocahontas Co. (AEL 182;
6/10/76; 0930).

12 Williams River bridge at Dyer, Webster Co. (AEL 165; 6/8/76; 1900).

13 Confluence North and South Forks of Cranberry River, Pocahontas Co.
(AEL 163; 6/10/76; 1230).

14 Cranberry River at Little Bee Run, 0.96 km upstream of bridge at Big
Rock Campgrounds, Nicholas Co. (AEL 166; 6/8/76; 1750).

15 Summit Lake, just off Route 39, Pocahontas Co. (AEL 161; 6/10/76;
2030).

16 North Fork of Cherry River, north bend of picnic area near road to Sum-
mit Lake, Greenbrier Co. (AEL 169; 6/8/76; 1030).

17 South Fork Cherry River, 5.3 km upstream of bridge across North Fork of
Cherry, e of Richmond, Greenbrier Co. (AEL 183; 6/8/76; 1200).

18 Laurel Creek, at confluence McMillion Creek, Namo Chapel, Greenbrier
Co. (AEL 175; 6/8/76; 1430).

52 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Table 1. (Com.)

Station No. Locality Description

19 Laurel Creek, 0.48 km s of Route 39, Fenwick, Nicholas Co. (AEL 193;
6/8/76; 1345).

20 Cherry River, Route 20 bridge, Holcomb, Nicholas Co. (AEL 178;
6/10/76; 1530).

21 Panther Creek, Route 39/20 bridge, east of Nettie, Nicholas County (AEL
176; 6/11/76; 1400).

22 Deer Creek at Deepwell, Nicholas Co. (AEL 191; 6/11/76; 1220).

23 Hominy Creek at Hominy Falls, Nicholas Co. (AEL 184; 6/11/76; 1530).

24 Hominy Creek, ford 6.4 air km e of Mt. Nebo, Nicholas Co. (AEL 142;
7/7/76; 1515).

25 Collison Creek, Nicholas Co. (AEL 157; 7/7/76, 1100).

26 Meadow River, co. rd. bridge just w of Grassy Meadows, Greenbrier Co.
(AEL 150; 7/8/76; 1400).

27 Meadow River, 0.27 km w of Station 26, w of Grassy Meadows, Green-
brier Co. (AEL 149; 7/8/76; 1330).

28 Meadow River, co. rd. bridge nw of Meadow Bluff, Greenbrier Co. (AEL
151; 7/8/76; 1400).

29 Meadow River, Route 60 bridge, n of McRoss, Greenbrier Co. (AEL 153;
7/8/76; 1630).

30 Meadow River, Russelville, WV (Babcock Railroad Junction), Fayette Co.
(AEL 159; 7/10/76; 1545) (AEL 192; 8/25/76; 1300).

31 Meadow River at mouth on Anglins Creek, Nicholas Co. (AEL 155;
7/7/76; 1300).

32 Meadow River, below new Route 19 bridge, Fayette Co. (AEL 173;
7/6/76; 1000).

33 Mouth of Meadow River, Fayette Co. (AEL 152; 7/5/76; 1500).

34 Little Clear Creek, Raders Run Railway Crossing, on Co. Rd. 1, Green-
brier Co. (AEL 174; 6/11/76; 1800).

35 Little Clear Creek, Route 60 bridge, Shawyer's Crossing, Greenbrier Co.
(AEL 156; 7/10/76; 1300).

36 Big Clear Creek at Anjean, Greenbrier Co. (AEL 177; 6/11/76; 1650).

37 Big Clear Creek, Route 60 bridge, Rupert, Greenbrier Co. (AEL 172;
7/10/76; 1215).

38 Little Sewell Creek, co. rd. bridge, just downstream of confluence Boggs
Creek, Greenbrier Co. (AEL 170; 7/8/76; 1500).

39 Sewell Creek at Lilly Park, Greenbrier Co. (AEL 158; 7/8/76; 1030).

Gauley River Fishes 53

Table 1. (Cont.

Station No. Locality Description

40 Sewell Creek, Route 60 bridge, Rainelle, Greenbrier Co. (AEL 144;
7/8/76; 1115).

41 Dogwood Creek, Saturday Rd. Bridge, Fayette Co. (AEL 162; 7/10/76;
1010).

42 Mouth of Rich Creek at Jodie, Fayette Co. (AEL 160; 7/11/76; 1040).

43 Big Beaver Creek, Route 41 bridge, Craigsville, Nicholas Co. (AEL 189;
8/24/76; 1420).

44 Brushy Fork, Route 43 bridge, Muddlety, Nicholas Co. (AEL 168;
7/11/76; 1700).

45 Muddlety Creek below confluence Brushy Fork and McMillion Creek, s of
Muddlety, Nicholas Co. (AEL 194; 7/11/76; 1745).

46 Lower Muddlety Creek at end of private drive off Route 39, Rev. Mycott
property, Nicholas Co. (AEL 190; 8/25/76; 1100).

47 Confluence of Peter's Creek and Buck Garden Creek, Gilboa, Nicholas Co.
(AEL 143; 7/11/76; 1500).

48 Peter's Creek, Summersville Dam rd. bridge, Drennen, W. Va. (AEL 171;
7/11/76; 1415).

49 Peter's Creek, along co. rd., 3.7 km s of Lockwood, above major water
falls, Nicholas Co. (AEL 145; 7/11/76; 1310).

50 Twentymile Creek, at confluence Ash Fork, Nicholas Co. (AEL: 7/9/76;
1830).

51 Bell's Creek, first bridge (residential) above confluence Twentymile Creek,
Fayette Co. (AEL 148; 7/9/76; 1700).

52 Confluence of Twentymile Creek and Bells Creek, Nicholas-Fayette Co.
line (AEL 154; 7/9/76; 1930).

RESULTS
Sub-drainages

Main-channel
That portion of the Gauley River above the mouth of Cherry River is
approximately 67 km in length and has an average gradient of 4.5 m/km;
the major tributaries are Williams, Cranberry and Cherry rivers, in
descending order (Fig. 1). From the confluence of Cherry River to the
Route 39 bridge (near the head of Summersville Reservoir), Gauley River

54 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

drops 104 m in 24 km (gradient, 4.2 m/km), and begins the descent into
its gorge. Summersville Dam is constructed across the gorge at or near
the mouths of Battle Run and McKee Creek, and has a normal pool
elevation of 503 m to above the Route 39 bridge approximately 22.5 km
upstream. In the vicinity of the dam the gorge is cut nearly 152 m below
the level of the Plateau. Hominy and Big Beaver creeks are the major
tributaries between Summersville Dam and Cherry River.

Progressing downstream 9.7 km from the Summersville Dam to Car-
nifex Ferry at the mouth of Meadow River, Gauley River descends an ad-
ditional 65 m at an average gradient of 6.7 m/km. Meadow River, the
only significant tributary, has a large drainage basin, near 932 km2, as
compared to 1932 km2 in the Gauley River basin above the confluence.
Collison Creek also enters this section of the river with an average
gradient of 28.4 m/km over its 10 km length. From Carnifex Ferry to
Swiss the Gauley River continues its run through the gorge and is charac-
terized by torrential water, boulder and bedrock substrate, and an
average gradient over 3.8 m/km for the 30.5 km distance. The river then
approaches base-level with an average gradient of 1.1 to 1.3 m/km, and
with long riffle and pool habitats alternating over the last 14.5 km to its
confluence with New River. These physical characteristics and associated
stresses influence distribution of fishes throughout the drainage. In this
survey, 31 species were collected in the main-channel Gauley River, Sta-
tions 1 through 9 (Table 2).

Williams River
Williams River heads against Dry Mountain, Pocahontas County, at
an elevation of 1210 m and discharges into Gauley River near Co wen at
an elevation of 689 m. Length of the stream is 54 km and average gradient
is 11.5 m/km. The drainage basin of Williams River, 337 km2, is larger
than Gauley River above their confluence. A total of 23 species was
collected in the Williams River subdrainage, Stations 10 through 12
(Tables 1 and 2; Fig. 1).

Cranberry River
Cranberry River rises in Pocahontas County at approximately 1402 m
in elevation, flows north, then southwest to enter Gauley River at Cran-
berry Station at 585 m. Length of the river is 51.5 km and gradient is 15.9
m/km. The basin is 181 km2 in size. Associated with the river is the Cran-
berry Back Country and Wilderness Study Area, a 14690 hectare tract
that is regulated by the U.S. Forest Service and has been promoted for in-
clusion as a Wilderness Area. Nine species were collected from two sta-
tions (13, 14) on Cranberry River (Tables 1 and 2; Fig. 1). Generally,

Gauley River Fishes 55

Cranberry River has a depauperate fauna which probably resulted from a
past history of intensive logging and mining, as well as naturally low pH
waters draining Cranberry Glades.

Cherry River
Cherry River, a major southern tributary to Gauley River, rises at 1341
m in Greenbrier County and flows generally in a northeast direction for
43.4 km to its mouth at Curtin. The drainage basin is 445 km2 in size and
gradient averages 17.4 m/km. Cherry River is rather industrialized in its
lower section and domestic sewage also adds to the degradation of water
quality. Major tributaries to Cherry River are the North Fork, South
Fork and Laurel Creek. A small impoundment, Summit Lake, is located
off Route 39, and discharges into the North Fork. Six stations (15-20)
were located in the Cherry River drainage (Tables 1 and 2; Fig. 1). Sum-
mit Lake (Station 15) is a well used recreation area stocked with Lepomis
macrochirus , Micropterus salmoides and salmonids.

Panther Creek
Panther Creek is a small southern tributary to Gauley River east of
Nettie. It is approximately 15.4 km long with an average gradient of 23.3
m/km. One collection was made on Panther Creek at Station 21 yielding
only 4 species (Tables 1 and 2; Fig. 1).

Hominy Creek
Hominy Creek is a principal southern tributary to Gauley River, with
its source near 1097 m in elevation at Grassy Knob, Greenbrier County.
Its length is approximately 35.1 km with an average gradient of 18 m/km.
The drainage basin is about 272 km2. A vertical drop of 6.1 m occurs at
Hominy Falls. A total of 15 species was collected from the system (Sta-
tions 22-24) (Tables 1 and 2; Fig. 1).

Collison Creek
This stream is a small tributary to Gauley River below Summersville
Dam. It is being considered as a possible site for impoundment by the
Corps. Total length is 9.5 km and gradient is 28.8 m/km. The drainage
basin is 24.9 km2. One collection (Station 25) on Collison Creek yielded 7
species (Tables 1 and 2; Fig. 1).

Meadow River
Meadow River, the major tributary to Gauley Rvier, rises in eastern
Summers County at approximately 1202 m and flows north to northwest
along the Fayette-Greenbrier and Fayette-Nicholas county lines to its

56 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

mouth at Carnifex Ferry. Its total length is 80.5 km and the drainage
basin is 932 km2. Gradient averages 10.4 m/km and increases from head
to mouth. Meadow River headwaters are along the broad Appalachian
Plateau, but in the last 17.7 km it enters its own scenic gorge to Carnifex
Ferry. Sixteen stations were located in the Meadow River system: main-
channel (Stations 26-33), Little Clear Creek (34-35), Big Clear Creek (36-
37), Sewell Creek (38-40), and Dogwood Creek (41). A total of 28 species
was collected in the system (Tables 1 and 2; Fig. 1).

Rich Creek
This is a small tributary to Gauley River at Jodie, W. Va. Total length
is near 12.2 km and average gradient is 45 m/km. The stream was sur-
veyed at its mouth (Station 42), particularly in a pool area adjacent to
but not a part of Gauley River. A total of 15 species was collected (Tables
1 and 2; Fig. 1).

Big Beaver Creek
This stream rises at 792 m in Webster County and is approximately
27.4 km in length. The gradient is 10.2 m/km and the drainage basin is
101 km2. Seven species were collected at Station 43 (Tables 1 and 2; Fig.
1).

Muddlety Creek
Muddlety Creek, about 32 km in length, rises at an elevation of 731 m
and enters Gauley River at Route 39 bridge. The drainage basin is 172
km2 and gradient is 8.0 m/km. The stream had been rechanneled along
much of the section below Muddlety, and road construction was present
adjacent to the stream. Coal washings were abundant in the substrate.
Thirteen species were collected from Stations 44-46 (Tables 1 and 2; Fig.
1). In general, the fauna was depauperate for the above reasons. This
stream is presently being considered for impoundment by the Corps.

Peters Creek
Peters Creek originates north of Summersville and flows southwest for
28.1 km to its confluence with Gauley River. Gradient averages 10.8
m/km and the basin is approximately 135 km2. A substantial waterfall of
about 10-12 m exists 1.6 km above its mouth. Eleven species were collec-
ted from this subdrainage at Stations 47-49 (Tables 1 and 2; Fig. 1).

Twentymile Creek
Twentymile Creek is the largest northern tributary to Gauley River. It
occupies a drainage basin of about 272 km2, and has a total length of 43.1

Gauley River Fishes 57

km. Gradient averages 10.2 m/km. Its principal tributary is Bells Creek.
Twentymile Creek was relatively productive with 21 species collected at
three localities (Stations 50-52) (Tables 1 and 2; Fig. 1). This was
probably due to a combination of factors, including its close relationship
to Gauley River below the gorge.

Annotated List of Species

The discussion of species collected in this survey is supplemented by
other collections cited in the literature and verified museum records.
Species are presented in phylogenetic order (Bailey et al. 1970). Con-
sidering the minimal information available on the Gauley River
ichthyofauna, this discussion should prove beneficial for future com-
parisons.

Petromyzontidae

Lampetra aepyptera. — One specimen of the least brook lamprey (AEL
181), a non-parasitic species, was collected in Gauley River at the mouth
of Cherry River. The specimen was an ammocoete taken over a detritus
bank from an eddy at the lower tip of a mid-channel island. Continued
seining and bank kicking did not produce additional specimens.

This record is the first report of L. aepyptera above Kanawha Falls. Its
presence in other West Virginia drainages was documented by Addair
(1944), Jenkins et al. (1972) and Stauffer, Denoncourt and Hocutt (ms.).
The specimen was taken above Summersville Reservoir, which infers an
established population prior to dam construction. Hocutt (1975) and
Stauffer (1975) did not report it from the upper New River system,
Virginia though they made extensive collections throughout the drainage.

Anguillidae
Anguilla rostrata. — Addair's (1944) collections of the American eel from
Greenbrier River (one collection) and New River (two collections) were
the only documented reports of this species in the upper Kanawha River
drainage. A single specimen of A. rostrata (ca. 1 m TL), reported in this
survey from Gauley River at the mouth of Meadow River below Sum-
mersville Dam (AEL 187), was found dead on the bank with an angler's
hook and line attached.

Salmonidae
Salmo gairdnen. — Rainbow trout are routinely stocked by the W. Va.
Department of Natural Resources.

Salmo trutta. — Brown trout are also commonly stocked in West Virginia.

58 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Salvelinus fontinalis. — The Brook trout is indigenous to much of the Ap-
palachian Plateau and natural populations are to be found in pristine
habitats common to Gauley River headwaters.

Cyprinidae

Campostoma anomalum. — The Stoneroller is common throughout the up-
per Kanawha River drainage. Addair (1944) collected it from eight of 19
sampling localities on Gauley River. This survey reports it from 27 of 52
stations. Verified museum records are CU 32399; UMMZ 95280, 95293,
108166, 108176, 165698, 165705, 165754.

Clinostomus funduloides . — Addair (1944) reported the Rosyside dace
from two Kanawha River localities, one of which was on Meadow River
near Russelville (Station 30, Fig. 1). Hocutt et al. (1978) reported it from
three localities in the Greenbrier drainage. It was found in this survey at
five stations: Cherry River (Stations 17 and 20) and Meadow River (Sta-
tions 38, 39, 40). Its distribution appears limited to higher elevations and
colder waters. Another record is from Cherry River (UNC 3315).

Encymba buccata. — Addair (1944) collected the Silverjaw minnow in six
of 87 collections above Kanawha Falls, including three localities in the
Gauley drainage: Twentymile Creek, Peters Creek and Williams River. It
was common at ten of the 20 localities where we collected it. Factors af-
fecting its distribution (Wallace 1972; 1973) appear to be favorable under
present conditions. Previous museum records are UMMZ 95294; USNM
(RVM-260).

Exoglossum laurae. — The Tonguetied minnow was collected by Addair
(1944) at three Williams River localities. It was not abundant in this sur-
vey, with only eight specimens collected from headwater stations of
Williams River (Station 11, AEL 182), Cranberry River (13, AEL 163)
and Cherry River (17, AEL 183; 18, AEL 175).

Nocomis platyrhynchus . — The Bigmouth chub is endemic to the upper
Kanawha River drainage above Kanawha Falls (Lachner and Jenkins
1971). The sibling species, jV. micropogon, occurs commonly in all other
drainages of the greater Ohio River, including the lower Kanawha River,
and Atlantic slope heading in West Virginia. Many diagnostic charac-
teristics of the two species are similar, with overlap occurring between
morphometric and meristic ranges; it is distinguished from TV.
platyrhynchus chiefly by tuberculation patterns. Despite six new dis-
tributional records for fishes above Kanawha Falls, it was necessary to
consider all micropogon-group chubs collected in Gauley River as JV.
platyrhynchus for the reasons cited.

Addair (1944) collected N. platyrhynchus from five localities. This survey
found it at 23 stations where it was often abundant. The use of elec-

Gauley River Fishes 59

trofishing gear and rotenone provided advantages of capture which Ad-
dair did not enjoy. Chubs were no doubt present throughout the drainage
at the time of his survey, but they are difficult to collect by seine due to
their habitat preference for rapid runs with large rubble and boulder sub-
strate. Other records are UNC 6083, CU 28867; UMMZ 95281, 108168,
165699, 165706, 165755.

Notemigonus crysoleucas. — Collection of the Golden shiner from Gauley
River at Carnifex Ferry (Station 6) represents the first record of this
species from the Gauley drainage. It is widely used by fishermen, and its
occurrence is attributed to bait bucket introduction.

Notropis albeolus. — Previous efforts by Addair (1944) and others did not
yield representatives of the Notropis subgenus Luxilus from Gauley River.
Hocutt et al. (1978) noted that Gilbert (1964) and R.D. Ross (VPISU
2429) took N. albeolus from Greenbrier River, and Hocutt (1974) and
Stauffer (1975) found it in New River, Virginia. We collected the White
shiner from nine localities where its presence was rare to abundant. The
reason for its absence in previous collections is speculative.

Notropis chrysocephalus. — The Striped shiner is also a member of the sub-
genus Luxilus (Gilbert 1964). Gilbert (1964) considered N. chrysocephalus to
be introduced to the upper Kanawha drainage, but common in other
Ohio River drainages. We collected it at five localities, but it was abun-
dant only at the mouth of Cherry River (Station 5). It was apparently
collected by Schwartz from Gauley River east of Bolair (UNC 685) and
Williams River (UNC 6087), but these records are not confirmed.

Notropis photogems. — Addair (1944) reported the Silver shiner from one
Peter's Creek locality and two Meadow River stations. We found it at ten
stations, distributed from the headwaters to our most downstream station
on Gauley River. The only other record of the Silver shiner from the
Gauley system is UMMZ 95283.

Notropis rubellus. — The Rosyface shiner was one of the most abundant
and widely distributed minnows collected in this survey, taken from 25
stations. Addair (1944) collected it from ten of 19 stations on Gauley
River. Other records are UNC 6086, UMMZ 95285, 108170, 165700,
165707, 165756.

Notropis scabnceps. — The New River shiner is endemic to the upper
Kanawha River drainage. We collected a single specimen from Williams
River (Station 12; AEL 164). Although never abundant, Addair (1944)
found it more widely distributed, collecting it from seven localities: Peters
Creek (1 station), Muddlety Creek (1), Cherry River (2) and Williams
River (3). This species deserves recognition by the W.Va. Department of
Natural Resources as a threatened or endangered species. Its ecological
requirements are not known, but its distribution appears to coincide with

60 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

high elevation, cold water streams. We found it at one of 52 localities in
the Gauley River (Station 12, AEL 182) and Hocutt et al. (1978) found it
at 5 of 32 localities on Greenbrier River. It was also previously collected
from Peters Creek (UMMZ 95295), Muddlety Creek (UMMZ 108172),
Panther Creek (UMMZ 1081178) and Williams River [USNM-(RVM-
260)] in the Gauley system.

Notropis spilopterus. — The Spotfin shiner was the only species collected
by Addair (1944) not found in this investigation. He collected it from Big
Creek (a lower tributary to Gauley River) and two localities on Meadow
River, and concluded it avoids high upland streams. It is also known from
Gauley River, Route 41 bridge (CU 32391, 32391) and at the confluence
of Gauley River and Twentymile Creek (UMMZ 95284).

Notropis stramineus. — Addair (1944) found the Sand shiner at Big Creek
and two Meadow River localities. We collected it at six localities (Sta-
tions 9, 48-52) in the lower Gauley River system.

Notropis telescopus. — Collections of the Telescope shiner from Stations 8
(15 specimens; AEL 185) and 50 (1; AEL 146) represent the first records
of this species from Gauley River. Gilbert (1969) considered N. telescopus
as introduced to New River and Hambrick et al. (1973) first reported it
from West Virginia. Hocutt et al. (1978) subsequently found it in Green-
brier River. Additionally, we have recent records from tributaries to
Kanawha River below Kanawha Falls.

Notropis volucellus. — Addair (1944) found the Mimic shiner common in
New and Kanawha rivers, but collected it at only two Gauley River
localities. We collected it from six localities (Stations 4, 5, 6, 8, 9 and 52).
Its distribution in the main-channel Gauley indicates an ability to
tolerate, if not a preference for, large water. Evidence presented by
Hocutt et al. (1978) supports this hypothesis. Other records from the
Gauley system are CU 32532; UMMZ 95282, 108171.

Phenacobius teretulus. — The Kanawha minnow is endemic to the system
above Kanawha Falls, but is rarely taken in West Virginia. Hocutt et al.
(1978) reported three specimens from Greenbrier River. Hambrick et al.
(1975) reviewed life history aspects of P. teretulus, noting only three collec-
tions of the species in West Virginia, all predating 1940. We collected
only two specimens from upper Laurel Creek (Cherry River system; Sta-
tion 18, AEL 175) during this study. Hocutt et al. (1978) recommended
its consideration as an endangered species; these data support that con-
clusion.

Pimephales notatus. — Surprisingly, Addair (1944) collected the Bluntnose
minnow from only one locality in the Gauley system, Twenty-mile Creek.
We collected it at 32 stations, making it one of the more widely dis-
tributed species encountered. There is a distinct preference of this species

Gauley River Fishes 61

for sandy, silt-laden pool or eddy habitats. Other records include: CU
32531; UMMZ 95286; USNM (RVM-260).

Pimephales promelas. — This study yielded 14 specimens of the Fathead
minnow from upper Williams River (Station 10, AEL 186). They were
collected in a standing back water area clogged with filamentous algae,
and represent the first records of the species from the system. Williams
River is noted for its trout fishing and the occurrence of this minnow is at-
tributed to bait bucket introduction.

Rhimchthys atratulus. — The Blacknose dace was a widely encountered
species in this study, found at 23 stations. Addair (1944) collected it from
11 localities. It is known from other collections: UMMZ 95297, 108173,
108177, 108183, 131827, 165701; USNM 196458.

Rhinichthys cataractae. — The Longnose dace was not reported from
Gauley River by Addair (1944). We found it at 18 localities where it was
locally abundant. Other records are from: Williams River [OSU (LM-76-
7), collected concurrent with this survey; UMMZ 165708]; Cranberry
River (UMMZ 165702); Peters Creek (UMMZ 95296); and Gauley
River (CU 32398).

Semotilus atromaculatus . — The Creek chub is a headwater species which
Addair collected at nine localities; we sampled it from 37 stations. Other
records are OSU (LM-76-7); UMMZ 95292, 108167, 165703, 165709,
165757; USNM (FJS 710).

Ictaluridae

Ictalurus natalis. — One specimen of Yellow bullhead (AEL 150) from
upper Meadow River is the first record of this species from the Gauley,
and also represents the first record of the species above Kanawha Falls
(Jenkins et al. 1972). It was collected from a small pool in the
headwaters, bordered by pasture. The specimen may be representative of
an indigenous population, or introduced to the system. Addair (1944)
collected two specimens from a tributary to lower Kanawha River.

Ictalurus nebulosus. — The collection of a Brown bullhead (AEL 174) from
upper Little Clear Creek is the first confirmed record of the species above
Kanawha Falls (Jenkins et al. 1972; Hocutt 1974; Stauffer 1975). It was
collected with electroshocking gear in a pool averaging 1.4 m in depth, 9.1
m wide and 30 m long. Other bullheads were observed swimming at the
surface, but not collected.

Noturus flavus . — Eight specimens (AEL 185) of the Stonecat collected by
rotenone from the Gauley River at Swiss, West Virginia (Tables 1 and 2:
Fig. 1), represent the first reported record of this species from the entire
Kanawha River drainage above or below Kanawha Falls. Station 8 was
characterized by a long riffle/run (90 m) habitat that opened into a pool

62 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

and was channelized on the right side of an island at the mouth of Laurel
Creek. A collection (CU 32540) of the Stonecat from Gauley River, Route
41 bridge, in 1951 has been confirmed (E. Brothers, pers. comm.), which
infers that N. flavus is native to the Gauley drainage.

Pylodictus olivaris. — Flathead catfish collected at the mouth of Meadow
River (Station 33) by rotenone are the first specimens from the Gauley
system reported in the literature. It was known from previous surveys
above and below Kanawha Falls (Addair 1944; Jenkins et al. 1972;
Hocutt 1974; Stauffer 1975), and is routinely taken by anglers (pers.
comm.) from Gauley River. Specimens previously collected from Gauley
River are housed at Cornell (CU 32397, 32535).

Centrarchidae

Ambloplites rupestris. — Addair (1944) did not collect the Rockbass from
the Gauley system, but Reed (1974) reported it. We collected it from 22
stations and it was rarely abundant. Other records include: CU 32400,
32534; and UMMZ 9529, 165760.

Lepomis cyanellus. — Jenkins et al. (1972) considered the Green sunfish as
probably introduced above Kanawha Falls. Our records of it from eight
stations are the first reported from the Gauley. It was not abundant,
although widely distributed, and was collected almost exclusively from
small to moderate size streams with pool habitats.

Leopmis gibbosus. — We collected one adult specimen, a probable in-
troduction, from lower Big Clear Creek (Station 37, AEL 172). The site
was characterized by a long base-level pool, and banks which were
strongly undercut. Another museum record from Gauley is CU 32539.

Lepomis macrochirus . — Addair (1944) collected the Bluegill from a lower
Gauley tributary, Big Creek. We found it at four stations (6, 15, 28 and
43).

Micropterus dolomieui. — Smallmouth bass were collected by Addair
(1944) from five stations and by Reed (1974) from various localities. It
was widely distributed in this survey, being collected at 22 localities.
Other records are CU 32392; and UMMZ 95289, 95291, 165713, 165761.

Micropterus punctulatus. — The species was not collected by Addair (1944)
from Gauley River, although he found it at several localities in the
Kanawha drainage where it is native (Jenkins et al. 1972). It was collec-
ted by Hocutt (1974), Stauffer (1975) and Hocutt et al. (1978) from the
New system. We found it at five stations in this survey.

Micropterus salmoides. — We collected the Largemouth bass from three
stations. It is regarded as introduced.

Gauley River Fishes 63

Percidae

Etheostoma blenmoides. — Greenside darters were collected from upper
Meadow River (2 stations), Cranberry River (1) and Williams River (1)
by Addair (1944). Other collections are OSU (LM-76-7); CU 25393,
32396, 32536; UMMZ 95279, 165710, 165758. We collected it from 26
stations and it was locally abundant.

Etheostoma caeruleum. — The first record of the Rainbow darter above
Kanawha Falls was reported in Hocutt et al. (1973) from New River just
above the mouth of East River. Subsequently, it was collected from East
River by Hambrick et al. (1973) and Stauffer et al. (1975), and from New
River (Stauffer 1975). A total of 272 specimens was taken in this survey
from six stations on the lower Gauley (AEL 146, 147, 148, 154, 160, 185).
No other records are known from above the Falls. Two dams, several
cataracts and the New River gorge separate this population from the
small one described by Hocutt et al. (1973) in the vicinity of East River.
Masnik, Hocutt, and Stauffer (ms) made over 200 collections in the upper
New River system, West Virginia, Virginia and North Carolina, 1971-
1975, and no other populations of E. caeruleum were located. Thus, the
East River population appears to be a relict, virtually isolated from other
populations.

Eheostoma flabellare. — Addair (1944) collected the Fantail darter from 14
of 19 localities sampled. We also found it widely distributed. Museum
records are: OSU (LM-76-7); UMMZ 95278, 108174, 108181, 108182,
131826, 165704, 165711, 165759.

Etheostoma nigrum. — Addair (1944) was the first to report the Johnny
darter above Kanawha Falls, collecting it at two localities: New River
above the confluence with Gauley River; and Glade Creek, tributary to
New River. He felt that its absence in Gauley River was related to rapid
currents and lack of suitable substrate for food organisms. We found it at
13 localities distributed from the headwaters to the mouth. Its distribu-
tion and absence above Summersville Dam indicate that it is indigenous
to the system. The only record located in addition to Addair (1944) was
from Gauley River, Route 41 bridge south of Summersville, West
Virginia (CU 32537).

Etheostoma osburm. — The Finescale saddled darter is endemic to the
Kanawha River drainage above Kanawha Falls. Jenkins et al. (1972)
reported it from Elk River below the Falls; however, the validity of these
data is doubted (R.E. Jenkins, pers. comm.). Our recent efforts in Elk
River have yielded only E. variatum, a sibling species. Addair (1944) found
E. osburm in upper Gauley River (1 station), Williams River (1) and
Cherry River (2). We found it at ten localities (Tables 1 and 2; Fig 1), be-
ing more predominant in the headwaters than the lower drainage. Other

64 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

collections are OSU (LM 76-7); CU 25394; UMMZ 165712. Collections
catalogued from Gauley River as E. kanawhae (UNC 7006) and E. vanatum
(UNC 6704) are regarded as E. osburm; we are not aware of verified
museum records of E. kanawhae from West Virginia.

Percina caprodes. — Specimens of the Logperch collected from Stations 8
(AEL 185) and 9 (AEL 147) represent the first reported records of this
species above Kanawha Falls. Addair (1944) collected it below the Falls,
and concluded that it was one of the darters unable to make it over that
physical barrier into the upper Kanawha drainage. Our specimens were
collected by rotenone (Station 8) and electrofishing (9) from extensive rif-
fle/run habitats characterized by moderate to large rubble substrate. Our
experience indicates that the Logperch is most often collected just below
a "lip" or dropoff where the riffle becomes a run. Other specimens of P.
caprodes are known from upper New River, Grayson County, Virginia
(R.E. Jenkins, pers. comm.), but not reported in the literature.

Percina cf. maculata. — The endemic relative of the Blackside darter oc-
curs above Kanawha Falls in the Kanawha River drainage (Jenkins et al.
1972; Hocutt 1974; Stauffer 1975; Hocutt et al. 1978) and is currently be-
ing described by E. Beckham. Addair (1944) collected it from one station
on upper Williams River. We collected it from six localities chiefly in cold
upland streams (Tables 1 and 2; Fig. 1). Past experience indicates that it
is widely distributed throughout the New River system, but rarely abun-
dant. The only other museum record from Gauley River is CU 28866.

Percina oxyrhyncha. — The Sharp nose darter has long been identified with
the ichthyofauna of the upper Kanawha River drainage; however, its dis-
tribution is wider than once expected (Jenkins et al. 1972; Hocutt and
Hambrick 1973; Denoncourt et al 1977; Hocutt et al. 1978; Thompson
1977; Hocutt, in press). Further, it was once considered by the U.S.
Department of Interior for posting as a threatened or endangered species.
Recent collecting in the upper New River (Hocutt et al. 1973; Hocutt
1974; Stauffer 1975) has shown that it is locally abundant. Hocutt et al.
(1973) collected 33 specimens from New River at Lurick, Virginia.
Specimens from eight Gauley River stations are the first reported from
the system (Table 1 ; Fig. 1). At the mouth of Meadow River (Station 33),
54 specimens were collected with rotenone. As suggested by Denoncourt
et al. (1977), the preference of adult P. oxyrhyncha for habitats with a large
rubble and boulder substrate probably accounts for the paucity of known
specimens before our work in the 1970's. The only other records of it from
the Gauley system are CU 28868 and 32538 from Route 41 bridge south
of Summersville.

Gauley River Fishes 65

Stizostedion vitreum. — Reed (1974) first discussed Walleye in the Gauley
River system. A rotenone sample from the mouth of Meadow River
produced one specimen in this survey!

Cottidae

Cottus bairdi. — Prior to this survey, records of Cottus were not known
from Gauley River. Three specimens of C. bairdi were collected from
Williams River, Station 11 (AEL 182). They were distinguished by
characters recommended by Robins (1954). One other collection exists
(OSU-LM 76-7), made concurrently with this survey from Williams
River.

Cottus carolinae ssp. — C. carolinae were collected from habitats with
moderate to large rubble in Williams River (Stations 10, 11) and Peters
Creek (47, 48).

Other Species
Other species are known to occur in the Gauley River system. Perca
flavescens, Lepomis auntus, Pomoxis annularis, and Pomoxis mgromaculatus, oc-
cur in Summersville Reservoir (C. Clower, pers. comm.), where they
presumably have been introduced. Reed (1974) reported stocking of Esox
masquinongy in Meadow River. Ictalurus punctatus (CU 32393, 32533) was
taken from Gauley River, Route 31 bridge, 4.8 km south of Summersville,
prior to constuction of Summersville Dam. Subsequent to our survey, the
W. Va. Department of Natural Resources collected specimens of Phoxinus
oreas 36 km upstream of the mouth of Williams River (R.L. Miles, pers.
comm.).

Expected Species
Hocutt et al. (1978) treated the fishes of Greenbrier River and included
information on species known from the main-channel New River between
Claytor Lake, Virginia, and Kanawha Falls, West Virginia. It is possible
that some of these species may potentially occur in Gauley River, e.g.,
Hybopsis dissimilis and Labidesthes sicculus. Claytor Lake, Bluestone Reser-
voir and the Union Carbide impoundment at Hawks Nest may be
barriers for downstream dispersal of upper New River fishes (Ross 1959;
Ross and Perkins 1959; Hambrick et al. 1973; Jenkins et al. 1972; Hocutt
1974; Hocutt et al. 1978; and Stauffer 1975). While the complete effec-
tiveness of Kanawha Falls as a barrier to the upstream dispersal of fishes
from the lower Kanawha River is debatable (Hocutt, in press), it no
doubt is a limiting factor for many species.

66

Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Table 2. Number of each species collected from Gauley River, 1976, by station.
(*new literature records to the Gauley River ichthyofauna; **first literature
records for species above Kanawha Falls).

Species 1

**Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdneri

*S. trutta

Salvelinus fontinalis

Campostoma anomalum

Clinostomus funduloides

Ericymba buccata

Exoglossum laurae

Nocomis platyrhynchus

*Notemigonus crysoleucas

*Notropis albeolus

*N. chrysocephalus

N. photogems

N. rubellus

N. scabriceps

N. stramineus

*N. telescopus

N. volucellus

Phenacobius teretulus

Pimephales notatus

*P. promelas

Rhimchthys atratulus 21

*R. cataractae

Semotilus atromaculatus 5

Catostomus commersom

Hypentelium nigricans

**Moxostoma erythrurum

**Ictalurus natalis

**/. nebulosus

**Noturus flavus

*Pylodictus olivans

Ambloplites rupestns

*Lepomis cyanellus

*L. gibbosus

L. macrochirus

Micropterus dolomieui

* M. punctulatus

*M. salmoides

Etheostoma blennioides

*E. caeruleum

E.flabellare ' 17

*E. nigrum

E. osburni 1

* *Percina caprodes

P. maculata

*P, oxyrhyncha

Stizostedion vitreum

*Cottus bairdi

*C. carolinae

Total Species 4

Total Specimens 44

Stations
3 4

8

35
13

11
2

7
106

10

1
1

57

333

383

4

86

21

974

22

195

40

494

24

2

3

2

1

12

3

3

18

3

5

10

1

18

10

6

2

1

25

30

11

1

3

4
15

2

1
3
3

162

3

6

11

13

14

17

60

189

668

574

1723

Gauley River Fishes

67

Table 2. (Cont.)

Species 7

**Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdnen

*S. trutta

Salvelinus fontinalis

Campostoma anomalum 2

Clinostomus funduloides

Ericymba buccata

Exoglossum laurae

JVocomis platyrhynchus 39

*Notemigonus crysoleucas

*Notropis albeolus

*N. chrysocephalus

N. photogenis

TV. rubellus 1

N. scabriceps

N. stramineus

*N. telescopus

N. volucellus

Phenacobius teretulus

Pimephales notatus 2

*P. promelas

Rhinichthys atratulus

*R. cataractae 27

Semotilus atromaculatus

Catostomus commersoni

Hypentelium nigricans 14

**Moxostoma erythrurum

**Ictalurus natalis

**/. nebulosus

**Noturus flavus

*Pylodictus olivans

Ambloplites rupestns

*Lepomis cyanellus

*L. gibbosus

L. macrochirus

Micropterus dolomieui 1

*M . punctulatus

*M . salmoides

Etheostoma blennioides 24

*E. caeruleum

E. flabellare

*E. nigrum 39

E. osburni

* *Percina caprodes

P. maculata

*P. oxyrhyncha 4

Stizostedion vitreum

*Cottus bairdi

*C. carolinae

Total Species 10

Total Specimens 153

Stations
9 10

11

12

9

10
32

2

2
5

67

18

7

58

36

1

25
08

50
243

180

1

15

5

93

10

16

1

14

31

38

9

91

4

1

1

40

8

5

118

3

10

36

2

1

30

2

4

1

1

2

2

17

33

16

9

35

6

3

33

54

13

2

1

9

3

25

4

10

10

3

3
10

20
481

22
808

12
279

19
180

11
68

68 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Table 2. (Cont.)

Stations
Species 13 14 15 16 17 18

* *Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdnen 1 1 1 4

*S. trutta

Salvelinus fontinalis 2 3

Campostoma anomalum 38 30

Clinostomus funduloides 6

Encymba buccata

Exoglossum laurae 1 1 1

Nocomis platyrhynchus 22 10

*Notemigonus crysoleucas

*Notropis albeolus

*N. chrysocephalus

N. photogenis 2

N. rubellus

N. scabnceps

N. stramineus

*N. telescopus

N. volucellus

Phenacobius teretulus 2

Pimephales notatus

*P. promelas

Rhinichthys atratulus 19 20 89

*/?. cataractae 4 9 2

Semotilus atromaculatus 1 1 1 21

Catostomus commersoni 1 9

Hypentelium nigricans 3

**Moxostoma erythrurum

**Ictalurus natalis

**/. nebulosus

**Noturusflavus

*Pylodictus olivaris

Amblophtes rupestris

*Lepomis cyanellus

*L. gibbosus

L. macrochirus 5

Micropterus dolomieui 2

*M. punctulatus

*M. salmoides 2

Etheostoma blennwides

*E. caeruleum

E.flabellare 15 2 6 11

*E. nigrum 2

E. osburni 7 7

**Percina caprodes

P. maculata 4

*P. oxyrhyncha

Stizostedion vitreum

*Cottus bairdi

*C. carolinae

Total Species 8 3 3 2 12 12

Total Specimens 44 4 8 21 101 189

Gauley River Fishes

69

Table 2. (Cont.;

Species

19

20

Stations
21 22

23

241

**Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdnen

*S. trutta 3

Salvelinus fontinalis 2

Campostoma anomalum 1 47 1 3

Clinostomusfunduloid.es 1

Encymba buccata

Exoglossum laurae

Nocomis platyrhynchus 14 41

*Notemigonus crysoleucas

*Notropis albeolus

*N. chrysocephalus 21 8

N. photogems 1

N.rubellus 81 10

N. scabnceps

N. stramineus

*N. telescopus

TV. volucellus

Phenacobius teretulus

Pimephales notatus 1

*P. promelas

Rhimchthys atratulus 98 32 2

*R. cataractae 3 16

Semotilus atromaculatus 5 4 4 10

Catostomus commersoni 3 2 4

Hypentelium nigricans 16 3 9

**Moxostoma erythrurum 41

* *Ictalurus natalis

**/. nebulosus

**Noturus flavus

*Pylodictus olivaris

Ambloplites rupestns 5 4

*Lepomis cyanellus

*L. gibbosus

L. macrochirus

Micropterus dolomieui 4

*M. punctulatus

*M. salmoides

Etheostoma blennioides 8 4 2

*E. caeruleum

E.flabellare 12 10 21 1 2

*E. nigrum 11 2

E. osburni 5

**Perana caprodes

P. maculata 4 1

*P. oxyrhyncha

Stizostedion vitreum

*Cottus bairdi

*C. carolinae

Total Species 1 15 4 10 3 13

Total Specimens 1 230 115 77 7 112

1**Moxostoma erythrurum (AEL 226), four specimens collected in Summersville Reservoir at mouth of
Hominy Creek by gill net in conjunction with this survey are not included in total number of species and
specimens at Station 24.

70 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Table 2. (Cont.)

Stations
Species 25 26 27 28 29 30

**Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdneri

*S. trutta

Salvelinus fontinalis

Campostoma anomalum 5 20

Clinostomus funduloides

Ericymba buccata 4

Exoglossum laurae

Nocomis platyrhynchus 86

*Notemigonus crysoleucas

*Notropis albeolus

*N. chrysocephalus

N. photogenis

N.rubellus 21 14 146

TV. scabriceps

N. stramineus

*N. tehscopus

N. volucellus

Phenacobius teretulus

Pimephales notatus 24 101 45 375 7 15

*P. promelas

Rhimchthys atratulus 54 2

*R. cataractae 1

Semotilus atromaculatus 7 2 50 6

Catostomus commersoni 10 2 5 3 1

Hypentelium nigricans 4

**Moxostoma erythrurum

**Ictalurus natalis 1

**/. nebulosus

**Noturusflavus

*Pylodictus olivaris '.

Ambloplites rupestris 2 2 1 24

*Lepomis cyanellus 4 6 1

*L. gibbosus

L. macrochirus 3

Micropterus dolomieui 19

*M. punctulatus 1 5

*M. salmoides

Etheostoma blenmoides 1 7 3

*E. caeruleum

E.flabellare 4 1

*E. nigrum 15

E. osburni 5

**Percina caprodes

P. maculata

*P. oxyrhyncha 8

Stizostedion vitreum

*Cottus bairdi

*C. carolinae

Total Species 7 8 4 9 6 1

Total Specimens 104 120 101 417 35 341

Gauley River Fishes

71

Table 2. (Cont.)

Stations
Species 31 32 33 34 35 36

**Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdnen 15 10

*S. trutta 1

Salvelinus fontinalis 1

Campostoma anomalum 2 7 2 1

Clinostomusfunduloid.es

Ericymba buccata 4 1 3 2

Exoglossum laurae

Nocomis platyrhynchus 9 251 188 5 2

*Notemigonus crysoleucas

*Notropis albeolus 9

*N. chrysocephalus

N. photogenis

N.rubellus 110 20 14 16 39

N. scabriceps

N. stramineus

*N. telescopus

N. volucellus

Phenacobius teretulus

Pimephales notatus 4 11 6 85 150

*P. promelas

Rhinichthys atratulus 1

*R. cataractae

Semotilus atromaculatus 24

Catostomus commersoni 5

Hypentelium nigricans 2

**Moxostoma erythrurum

**Ictalurus natalis

**/. nebulosus 1

**Nolurus flavus

*Pylodictus olivaris 5

Ambloplites rupestns 1 8 28 3 1

*Lepomis cyanellus 7 11

*L. gibbosus

L. macrochirus

Micropterus dolomieui 1 72 41

*M . punctulatus

*M. salmoides

Etheostoma blenmoides 1 3 14

*E. caeruleum

E.flabellare 5 10 1 1

*E. nigrum 1 7

E. osburni

**Percina caprodes

P. maculata

*P. oxyrhyncha 24 54

Stizostedion vitreum 1

*Cottus bairdi

*C. carolinae

Total Species 15 12 16 13 11 5

Total Specimens 171 530 478 178 223 42

8

24

54

99

2

1
1

19

6
1

6

77

16

5

1

17

7

225

83

17

3

5

12

18

40

1

15

1

7
26

72 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Table 2. (Cont.)

Stations
Species 37 38 39 40 41 42

**Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdneri

*S. trutta

Salvelinus fontinalis

Campostoma anomalum

Clinostomusjunduloid.es

Ericymba buccata 33

Exoglossum laurae

Nocomis platyrhynchus

*Notemigonus crysoleucas

*Notropis albeolus

*N. chrysocephalus

N. photogenis

N. rubellus 7

TV. scabriceps

N. stramineus

*N. telescopus

N. volucellus

Phenacobius teretulus

Pimephales notatus 1 09

*P. promelas

Rhinichthys atratulus 2

*R. cataractae

Semotilus atromacuiatus 15

Catostomus commersoni 7

Hypentelium nigricans 1

**Moxostoma erythrurum

**Ictalurus natalis

**/. nebulosus

**Noturus flavus

*Pylodictus olivaris

Ambloplites rupestris 2 3

*Lepomis cyanellus 1

*L. gibbosus 1

L. macrochirus

Micropterus dolomieui 1 2 3

*M. punctulatus

*M. salmoides

Etheostoma blennioides 1 2 5

*E. caeruleum 1 24

E.flabellare 1 37

*E. nigrum 21

E. osburni

* *Percina caprodes

P. maculata

*P. oxyrhyncha

Stizostedion vitreum

*Cottus bairdi

*C. carolinae

Total Species 10

Total Specimens 178

4

166

33

1

8

2

1

22

11

23

32

29

211

5

9

9

3

526

1

32

11

6

11

3

15

130

106

245

33

1291

Gauley River Fishes

73

Table 2. (Cont.)

Species 43 44

* *Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdneri

*S. trutta

Salvelinus fontinalis

Campostoma anomalum

Clinostomus funduloides

Ericymba buccata 67 14

Exoglossum laurae

Nocomis platyrhynchus

*Notemigonus crysoleucas 2

*Notropis albeolus

*N. chrysocephalus

N. photogenis

TV. rubellus

N. scabriceps

TV. stramineus

*N. telescopus

N. volucellus

Phenacobius teretulus

Pimephales notatus 100 21

*P. promelas

Rhimchthys atratulus 1

*R. cataractae

Semotilus atromaculatus 69 10

Catostomus commersoni 19 1

Hypentelium nigricans 1

**Moxostoma erythrurum

* *Ictalurus natalis

**/. nebulosus

**Noturus flavus

*Pylodictus olivaris

Ambloplites rupestris

*Lepomis cyanellus 1 2

*L. gibbosus

L. macrochirus 4

Micropterus dolomieui

*M. punctulatus 1

*M. salmoides 1 1

Etheostoma blennioides 1

*E. caeruleum

E.flabellare 1

*E. nigrum

E. osburni

**Percina caprodes

P. maculata

*P. oxyrhyncha

Stizostedion vitreum

*Cottus bairdi

*C. carolinae

Total Species 7 8 4

Total Specimens 96 183 38

Stations
45 46

47

48

2
24

45
1

52

8
2

12

4
26

8
138

74 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Table 2. (Cont.)

Stations
Species 49 50 51 52 Total

**Lampetra aepyptera

*Anguilla rostrata

*Salmo gairdneri

*S. trutta

Salvelinus fontinalis

Campostoma anomalum 7

Clinostomus funduloides

Ericymba buccata 33

Exoglossum laurae

Nocomis platyrhynchus

*Notemigonus crysoleucas

*Notropis albeolus

*N. chrysocephalus

N. photogenis

N.rubellus 102

JV. scabriceps

JV. stramineus 23

*JV. telescopus

JV. volucellus •

Phenacobius teretulus

Pimephales notatus 75

*P. promelas

Rhinichthys atratulus

*R. cataractae

Semotilus atromaculatus 9

Catostomus commersoni 18

Hypentelium nigricans 89

**Moxostoma erythrurum

**Ictalurus natalis

**/. nebulosus

**Noturus flavus

*Pylodictus olivaris

Ambloplites rupestris

*Lepomis cyanellus

*L. gibbosus

L. macrochirus

Micropterus dolomieui 22

* M . pundulatus

*M . salmoides

Etheostoma blennioides

*E. caeruleum

E.flabellare

*E. nigrum

E. osburni

**Percina caprodes

P. maculata

*P. oxyrhyncha

Stizostedion vitreum

*Cottus bairdi

*C. carolinae

Total Species 9 15 15 16 49

Total Specimens 378 380 325 84 12,518

24

33

5

92

55

15

4

107

5

13

15
42

4
92

7

5
1

4

13
1

43

9

9

16
14

1
20

1

1

1
7

5

2

2
1

2

3

1

8

91

5

2

4

1
2

Gauley River Fishes 75

ZOOGEOGRAPHY

This survey established many distributional records. The remoteness of
the drainage and its inaccessibility, particularly in the lower two-thirds of
the main-stem and its tributaries, has undoubtedly been a deterrent to
sampling. The upper New River is generally more accessible. Perhaps for
this reason, students of West Virginia fishes have tended to avoid the
Gauley in their efforts.

The confluence of Gauley and New rivers forms Kanawha River above
Kanawha Falls, long considered to be a barrier to upstream dispersal of
fishes. All the endemic forms of the upper Kanawha (New) drainage oc-
cur in the Gauley, exluding Etheostoma kanawhae which is not known from
West Virginia. However, the presence of six species found in Gauley
River in this survey and not previously reported from the upper Kanawha
drainage infers a relationship of the Gauley with the lower Kanawha
drainage. The new distributional records indicate that either Kanawha
Falls was not as effective a barrier as supposed, or conditions were once
more favorable than at present for the dispersal of fishes above the Falls
into Gauley River. In either case, it is assumed that other physical
barriers were effective in limiting dispersal of fishes into the upper New
River system. The zoogeographic relationships summarized below are
discussed in detail by Hocutt (in press).

The wide distribution of Etheostoma nigrum in the Gauley drainage sup-
ports a hypothesis that more favorable conditions once existed for fishes
to negotiate Kanawha Falls. It is more often found in slack water habitats
over sand and detritus, and avoids strong currents. Thus, we reason that
it is not a recent migrant over Kanawha Falls. Cole (1971) supports this,
postulating that the present E. nigrum distribution resembles that of pre-
Pleistocene times, and that populations east of the Appalachians are a
consequence of relatively recent stream capture.

Subsequent to the Tertiary uplift of the Appalachians, the Teays River
had a cutting advantage due to its volume and gradient. The evolving
(New River) gorge with its multitude of rapids and cataracts probably
served as an effective barrier. Kanawha, Wylie, Bull and Sandstone Falls
are remnants of that rejuvenated period (Hocutt et al. 1978). Prior to re-
cent impoundment, Kanawha Falls was the first of the series of natural
barriers that served as a faunal filter (Hocutt in press).

The Pleistocene (Neff et al. 1970; Hocutt et al. 1978) impoundment of
Teays River would have inundated Kanawha Falls if indeed the Falls ex-
isted at that time. Gauley River, smaller in drainage and volume of water,
cut its gorge more slowly than the Teays and offered a route of dispersal
for fishes (Hocutt, in press). Many species present today serve as relict
populations to that time. The large numbers of Etheostoma caeruleum

76 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

support the contention that conditions once facilitated passage of fish up-
stream of the Falls area. Lampetra aepyptera, Moxostoma erythrurum, Ictalurus
natalis, I. nebulosus, and Noturus flavus are native to the greater Ohio River
drainage (Jenkins et al. 1972), thus their presence in the Gauley system
may be explained by a Pleistocene impoundment. The capture of an im-
mature L. aepyptera above Summersville Dam suggests a population pre-
dating construction; the presence of N. flavus in a 1951 collection (CU
32540) is similar evidence for an established population.

Gilbert (1969) regarded Notropis telescopus as introduced to the New
River system, with all previously known records being very recent and
from only a small part of the upper New River, Virginia (Jenkins et al.
1972). Subsequently, it was collected by Hocutt et al. (1978) from Green-
brier River, West Virginia, in this survey, and from Kanawha River
below Kanawha Falls (Hocutt and Stauffer, ms). These data suggest
three alternatives for its distributional status: (1) it is indeed an in-
troduced species to the upper New system that has rapidly extended its
range downstream by negotiating Bluestone Reservoir and the Union
Carbide impoundment at Hawks Nest; (2) it is an introduced species to
both the upper (New) and lower Kanawha drainages; or (3) it is a native
species that is rarely encountered in the Kanawha drainage, its presence
probably related to stream capture with the Tennessee drainage (Ross
and Carico 1963).

Notropis scabriceps and Phenacobius teretulus are identified as part of the
unique faunal assemblage of New River. Their almost exclusive distribu-
tion in the upper Gauley River system could be related to stream piracy
with Greenbrier River (Wright 1934; Hocutt, in press), or to present-day
ecological factors limiting their distribution. Wright (1934) felt there was
evidence that the East Fork of Greenbrier River and Knapp Creek at one
time continued a westerly flow beyond the present Greenbrier. The East
Fork would have joined Shavers Fork in the vicinity of Cheat Bridge.
Knapp Creek's westernly continuation is in approximate alignment with
Stony Creek (reversed) into Laurel Creek of Williams River (Gauley
system). Wright (1934) stated that little geological evidence existed to
support this contention in light of the fact that erosion of the Greenbrier
Valley limestone erased traces of stream diversion. Biological evidence
(Hocutt et al. 1977, 1978; Hocutt, in press) supports Wright's (1934)
hypothesis. Additionally, if his contention were correct, dispersal of upper
New River fauna (e.g., N. scabriceps, P. teretulus, Cottus spp.) into the
Gauley system via Greenbrier River would have been possible.

Once fauna entered Gauley River via Williams River from the Green-
brier, distribution would be related to downstream and lateral dispersal.
Little Beaver Creek may once have flowed into Muddlety Creek via

Gauley River Fishes 77

Harris Fork of the latter (Reger 1921). Peters Creek apparently captured
Muddlety Creek drainage via Arbuckle Branch, and could have once
flowed through the present valleys of Otter and Little Elk creeks to a con-
fluence with Gauley River at Swiss, 12. 9 km downstream of its present
mouth (Reger 1921). Back Fork of McMillion Creek may have once
flowed into Persinger Creek. Similarly, fishes may have dispersed up-
stream through this series of captures.

Elk River of the lower Kanawha drainage has captured part of the
Gauley system (Campbell 1896) and continues to encroach on Gauley
River waters. Only 4.8 km separate their main-channels near Webster
Springs, and capture by the Elk appears imminent (Reger 1920); the Elk
valley is nearly 244 m lower than the Gauley valley at Webster Springs.
Also, Anthony and Laurel creeks, tributaries to Birch River of the Elk
system, are presently encroaching on Beaver and Muddlety creeks of
Gauley River (Reger 1921). There may have been faunal interchange
during the Elk River capture of Gauley River drainage (Campbell 1896).
Noturus miurus (UNC 7629; identifications verified) is recorded from
Williams River (Gauley system), but is suspected to represent a mistaken
locality. No specimens were collected by us from Williams River after
repeated sampling, but the species is known from nearby Elk River
(Taylor 1969).

In summary, these data support a hypothesis that Kanawha Falls was
once more navigable to fishes than at present (Hocutt, in press).
Ichthyofauna once above the Falls area could migrate up either the New
River gorge and its series of montane stresses, or up Gauley River, a
rigorous but less stressful route. Biological evidence supports Wright's
(1934) contention of piracy between the Greenbrier and Gauley rivers,
with fauna having dispersed into each drainage from the other. Facts may
be masked by introductions of various species (e.g., Hocutt and Ham-
brick 1973) into the Gauley system, and by extensive logging and mining
operations in the basin during the past 80 years.

ACKNOWLEDGMENTS. — We particularly wish to express our
gratitude to D. Harris, C. Clower and W. Tolan of the Ecological Services
Division, U.S. Fish and Wildlife Service, Elkins, West Virginia. The spirit
in which this study was conducted is a tribute to their cooperation. Our
appreciation is further extended to various persons of the West Virginia
Department of Natural Resources (W. Va. DNR) who assisted whenever
possible and directed the rotenone samplings. Also, we are appreciative of
the Handley and Elkins facilities which the W. Va. DNR allowed us to use
during the course of this investigation. Fred C. Rohde, University of North

78 Charles H. Hocutt, Robert F. Denoncourt, Jay R. Stauffer, Jr.

Carolina, Morehead City, confirmed the identification of Lampetra aepyp-
tera. Drs. David A. Etnier, University of Tennesee, and Robert E.Jenkins,
Virginia Commonwealth University, offered critical comment for improve-
ment of the manuscript.

LITERATURE CITED

Addair, J. 1944. The fishes of the Kanawha River system in West Virginia and
some factors which influence their distribution. Ph.D. dissert., Ohio State
Univ., Columbus. 225 pp.

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description and distribution of the sharpnose darter, Percina oxyrhyncha.
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Accepted 27 June 1978

Historical Review of the Carolina Parakeet
in the Carolinas

Daniel McKinley

Department of Biological Sciences,

State University of New York at Albany,

Albany, New York 12222

ABSTRACT. — Parrots or parakeets appear in many lists of Carolina
birds recorded by such early voyagers, explorers and promoters as
Thomas Hariot (1588), William Hilton (1664), Thomas Ashe (1682),
Samuel Wilson (1682), John Lawson (1709), Mark Catesby (1731) and
others. These references, when authentic, can be safely assigned to the
now extinct Carolina parakeet, Conuropsis carolinensis . The species was so
named by Linnaeus (1758) from a drawing of a specimen taken in South
Carolina by Catesby. Despite association of the region with the bird's
name, a long history constitutes the bulk of evidence on the species in
the Carolinas. There are no specimens, exceedingly few precise claims
by ornithologists, and no specific references to eggs, migratory move-
ments or young. Little can be found to validate North Carolina's claim
to parakeets after about 1770 (William Bartram). For South Carolina,
matters are more complex: widely spaced but fairly persistent records
bring the bird's history there down to about the end of the Civil War,
with a final, no doubt storm-tossed, bird accidentally occurring about
1885. There was a flurry of alleged sightings in the decade of the 1930s,
but the birds either disappeared without documentation or were not
there in the first place.

INTRODUCTION

In an account of the Carolina parakeet, Conuropsis carolinensis
(Linnaeus), history and biology must mix, for the species is extinct. That
notably handsome bird, so often remarked by early travelers, thus joins
the passenger pigeon and other vanished species in a group about which
we (as a civilization) know pretty much all that we shall know. Veteran
ornithologists know about it, of course, and sometimes allege a good
many things that a careful historian of the species learns are not true.
But, ask a concerned American citizen to name ten exterminated species
of animals and the Carolina parakeet will probably not be among them.

Except for its name, the parrot of Carolina was not uniquely as-
sociated with the Carolinas. It was widely, if somewhat erratically, dis-
tributed in the eastern United States, being found from Florida to Texas
and well up the Arkansas, Missouri, Ohio and Mississippi river valleys.
This report, emphasizing evidence on distribution in North and South

Brimleyana No. 1: 81-98. March 1979. 81

82 Daniel McKinley

Carolina, extends my geographical history of the parakeet (McKinley
1960, 1964, 1965, 1976, 1977a, b, c, 1978a, b, c). Some quotations from
the literature may seem unnecessarily long, but they impart to modern
readers the enormous impact made by the new land upon early ob-
servers, and leave to critical minds the final evaluation of these pioneer
statements.

Early and late, there is chaff among the grain. Since "Carolina" al-
ready has an open-ended quality about it, for example, it may be well to
sandwich in here a quaint and innocent early allusion to the parakeet in
the New World, although largely extrapolated from the cartographic. Ex-
cept for Hariot's Roanoke Island report for North Carolina, it is also the
earliest attribution of the species to the area of Carolina. Sanson d 'Ab-
beville, writing about 1653, said nothing about parakeets in Florida or
Virginia, but in regard to a vague region in between that he called (in
translation) "the Appalachians," he wrote: "In this region there are par-
rots, pigeons, turtle doves, eagles, ducks, magpies, sparrows and many
other types of birds." A map accompanying his account shows an inland
area denominated "Apalatchy Monts" which runs more or less east-west
between a tremendously northerly-swollen Florida and a Virginia that is
sort of hunched up against the Atlantic Ocean (1959:48).

PARAKEETS IN NORTH CAROLINA: THE REPORTS

For a state with few substantial records of the parakeet, North Carolina
has a history of the species that is resplendent in its antiquity. Thomas
Hariot (1588) reported that the ill-starred little colony on Roanoke Island
had parrots. In what was certainly America's first example of science for
science's sake, he wrote: "There are also Parats, Faulcons & Marlin
haukes, which although with us they bee not used for meate, yet for other
causes I thought good to mention" (in Quinn 1955:359). Unfortunately,
John White, dedicated planter of the colony (and grandfather of Virginia
Dare, born there), did not figure the parakeet among the lovely illustra-
tions of natural history subjects that he left to a careless posterity (Hulton
1965).

Thus, Dare County has an early claim to parakeets. The next report af-
firmed that the species was found in southeastern parts of the state. Cap-
tain William Hilton of the West Indian island of Barbados carefully sur-
veyed the coast of the Carolinas in autumn 1663. His explorations of Cape
Fear River may have taken him to the vicinity of Fayetteville, Cumber-
land County, with noteworthy descriptive results. In early November, he
described the return down-river toward the sea (1664): "So we returned
. . . viewing the Land on both sides the River, and found as good tracts of
land, dry, well wooded, pleasant and delightful as we have seen any where

Carolina Parakeet in Carolinas 83

in the world, with great burthen of Grasse on it, and in some places very
high, the woods stor'd with abundance of Deer and Turkies every where;
we never going on shoar, but saw of each also Partridges great store,
Cranes abundance, Conies, which we saw in several places; we heard
several Wolves howling in the woods, and saw where they had torn a Deer
in pieces. Also in the River we saw great store of Ducks, Teile, Widgeon,
and in the woods great flocks of Parrakeeto's; the Timber that the woods
afford for the most part consisting of Oaks of four or five sorts, all differing
in leaves, but all bearing Akorns very good." To certify that the expe-
dition was not bent entirely upon esthetic and scientific ends, he
enumerated the game taken: "In that time as our business called us up
and down the River and Branches ... we kill 'ci of wild-fowl, four Swans,
ten Geese, twenty nine Cranes, ten Turkies, forty Duck and Mallard,
three dozen of Parrakeeto's, and six or seven dozen of other small Fowls,
as Curlues and Plovers, etc." (Salley 1911:46, 53). The decimation of a
continent was underway.

But, such early writers served their own days only, if any at all, and
seem to have been soon forgotten. More seminal, however, was the work of
John Lawson, loyal adopted son of North Carolina. I think it probably sig-
nificant that he left no evidence of having seen parakeets in the arduous
journey from Charleston, South Carolina, to Pamlico Sound, North Caro-
lina, in the period of late 1700 to late February 1701. That trip took his
party up the Santee River and its tributary, the Wateree, to the vicinity of
Union County, North Carolina. From there they went northward deep
into central parts of the latter state, thence eastward to coastal "Pampti-
cough" (Pamlico), a distance of some 550 miles (see Lefler's comments,
Lawson 1967).

Lawson's 1709 account of the aboriginal and natural history of the
colony of North Carolina has extensive accounts of many kinds of animals
and plants, including parakeets. It is full of information — and misin-
formation — that must have cost him much conversation and cor-
respondence. "Parrakeetos are of a green Colour, and Orange-Colour'd
half way their Head, " he wrote in part. "Of these and the Allegators, there
is found none to the Northward of this Province" (1967:146-147).

Colonel William Byrd of Westover, ever in search of an outlet for his
restless energies (and of a source of income) had something to say about
parakeets in North Carolina. He had little use for the scurvy inhabitants of
that state, but mentioned as an extenuating circumstance in their failure
to plant orchards that "paraqueets" frequently raided fruit trees in
autumn (1929:77-78). Or so he said. I suspect the passage was put in for
literary effect; his secret diary, from which he later wrote up the public ac-
count, does not mention the parakeets at all {op. at.).

84 Daniel McKinley

When Benjamin Franklin's printing partner, Hugh Meredith, grew
tired of the bonds of city life, he dissolved the partnership (July 1730) and
apparently went to live among Welch kinsmen that summer and fall near
Cape Fear in Brunswick and New Hanover counties. He spent the winter
at the mouth of Black River, South Carolina, "near 100 miles West of
Brunswick." His letters to Franklin describing Cape Fear, published in
April and May 1731, were presumably written that winter or early spring
and, of course, refer to the previous summer. The Cape Fear region,
Meredith reported, had no chestnut, but other Pennsylvania trees were
present, plus "Cypress, Laurel, Bay red and yellow, Live Oak and Swamp
Oak, all Evergreen except Cypress; with several Sorts whose Names I
know not. Pheasants and Heath-hens here are none, but all other Fowl
common with you are. Parraquets in Summer, and greater Plenty of
Turkeys than ever I saw in Pennsylvania. Here are Foxes, Wolves, Wild-
cats, Possums, Raccoons, and Panthers always, and Bears sometimes in
great plenty; also plenty of Deer, but Beavers here are none, nor any
Ground-Squirrels, tho' plenty of Gray and Flying Squirrels; Alligators are
very numerous here but not very mischievous; however, on their Account
Swimming is less practis'd here than in the Northern Provinces" (1922:26-
27). It is unfortunately not clear whether Meredith meant to say that
parakeets were absent in winter. If we infer this, it must be realized that it
would have been hearsay information in his case.

John Brickell, who practiced medicine in Edenton for many years, took
part of his material straight from Lawson and added bits of his own.
Whether to trust him at all is a question at times. For example, he ob-
served in 1737, in regard to what he called "Black small-Crows" (by
which he apparently meant blackbirds of some sort that were enemies of
corn), that they "Build their Nest in hollow Trees as the Parakeetoes do"
(1911:179, 181). This is something that our native blackbirds do not do
and which is largely a matter of popular conviction with the parakeet, for
nobody ever got around to observing it conclusively. (A source of ad-
ditional Brickell claims is the Virginia traveler, the Rev. John Clayton, as
shown by Simpson and Simpson [1977], although Clayton contributed
nothing on the parakeet. The Simpsons reproduce [page 4] a plate that
appeared in the 1737 edition of Brickell where a "Parekeetoe" is among
the fairly recognizable denizens of Carolina.)

Aside from William Bartram's rather circumstantial contribution of
1791, the story of the parakeet in North Carolina very nearly ends with
Lawson and Brickell. When Bartram wrote that the "parrot of Carolina,
or Parrakeet" was among "natives of Carolina and Florida, where they
breed and continue the year round," North Carolina, although not ex-
pressly said, was probably meant. He had spent a good deal of the time

Carolina Parakeet in Carolinas 85

from 1761 to 1765 and 1770 to 1772 at the plantation of his uncle William
who lived near present-day Council, Bladen County, in the Cape Fear
River country. He wondered that parakeets did not appear in his native
Pennsylvania, since they could, he thought, easily fly from North
Carolina, "where they are very numerous" (1958:182, 190-191).

In a search of literature lasting nearly 20 years I have found no records
of parakeets in the inland central and western three-quarters of North
Carolina. Negative reports are never of much value singly, although I have
cited Lawson's account above. I am also impressed by such a diary as that
of Lieutenant Reeves, an intelligent Revolutionary War soldier of the
Pennsylvania Line (1897). He crossed the entire central North Carolina
region from north to south in the spring of 1782, alert to all natural
phenomena, but saw no parakeets until he was within South Carolina.

NORTH CAROLINA: A CRITIQUE OF RETROSPECTS

Considering enormous geographic differences between North and South
Carolina, it is unfortunate that even ornithologists have so often lumped
them as "the Carolinas." It is certainly doubtful if a statement that,
"They apparently were common in the Carolinas up to 1850, or perhaps
1860, but must have disappeared from there soon after that" (Bent
1940:3) ought to stand as any sort of North Carolina record. H. H.
Brimley wrote that the Austro-riparian or Louisianian Life Zone, charac-
terized by the alligator, marsh rabbit, big eared bat and chuck- will's-
widow, "formerly . . . received added brilliance in North Carolina by the
presence of the gaudy and noisy Carolina Parroquet," but he offered no
significant evidence of its occurrence (1896:66). C. S. Brimley had been
asked by old people "what was the bird that used to roam over the state
before the Civil War and eat cocklebur," but this is not satisfactory proof
of its existence or time of disappearance (MS. note, N. C. State Mu-
seum).

The first state-wide bird list described the species as among those gone
from North Carolina due to "changes in their environment," an instance
of misplaced precision if there ever was one. No doubt optimistically, it
was thought that the species might still be looked for as an accidental visi-
tor in southeastern parts of the state (Atkinson 1887:50, 65).

Being totally unhelpful, both Hasbrouck (1891:374) and Smithwick
(1 897 :21 2) cited Catesby as the only previous authority for the parakeet in
North Carolina, despite the fact that Catesby did not refer to that state
(Wayne 1917:3). Pearson et al. also mentioned "Catesby 's record in
1731" (1919:184), adding to its geographic ambiguity the fact that the
date was the rather belated year of publication, not the time when he

86 Daniel McKinley

would have seen them there anyway. They later added William Byrd's so-
called record (1942:192). They chose to ignore, no doubt wisely, a manu-
script note dated 10 January 1925 that C. S. Brimley had filed in the State
Museum: "Mr. John Handy Ford of Wilmington told Mr. J. C. Crawford
recently that he took the eggs of this species some ten years ago in the
swamps near Wilmington."

THE PARAKEET IN SOUTH CAROLINA: EARLY YEARS
References that call the parakeet a "formerly abundant permanent resi-
dent" (Bent 1940:3) and "common in the Carolinas up to 1850" (Sprunt
and Chamberlain 1949:292) are rather devoid of substantiating details.
Elliott Coues, in the first critical list of South Carolina birds, wrote that
the species "appears to have been in former times a common bird: but its
occurrence has not been noted of late years" (1869:119). But that, except
for the negative second part, is also empty. Even Wayne's ambitious orni-
thology of the state provided an account that was obscure and lacking in
details (1910:10). Bent mentioned the range of the parakeet as formerly
including the Pine Barrens and Edding Island (1940:10), but both these
attempts to particularize distributions require qualification, as will be
shown.

The story of the parakeet in South Carolina begins modestly enough.
"T. A., Gent." (supposedly Thomas Ashe, "Gentleman," a ship's clerk —
and not a man named "Gent") included a list of birds in a promotional
letter written to a friend, as was done by many early English explorers and
exploiters. He described in glowing terms the region of "Charlestown,"
where he lived, probably about 1680-1682: "Birds the Country yields of
differing kinds and Colours: For Prey, the Pelican, Hawk, and Eagle, etc.
For Pleasure, the red, copped and blew Bird, which wantonly imitates the
various Notes and Sounds of such Birds and Beasts which it hears, where-
fore, by way of Allusion, it's call'd the mocking Bird; for which pleasing
Property it's there esteem 'd a Rarity. Duck, Mallard, Widgeon, Teal,
Curlew, Plover, Partridge, the Flesh of which is equally as good, tho'
smaller than ours in England. Pigeons and Parakeittoes. In Winter huge
Flights of wild Turkies, oftentimes weighing from twenty, thirty, to forty
pound . . . They have a Bird I believe the least in the whole Creation,
named the Humming Bird; in bigness the Wren being much Superior . . .
they continue between the Tropiques the whole year round . . . but I am
informed, that in the more Northern parts of America they sleep the whole
Winter" (Salley 1911:151-152). Although the worthy clerk perhaps got
painted buntings ("red, copped and blew") confused with mockingbirds
and his typesetter put "In Winter" with turkeys instead of parakeets, he
ought not to be ridiculed for believing that various small birds hiber-

Carolina Parakeet in Carolinas 87

nated; many scientists of his day agreed with him. The year 1682, one
judges, was a good one for wild turkeys, and the frontier mentality has fer-
vently abided by forty-pound turkeys ever since.

Samuel Wilson, who was probably never in the colonies, seconded the
sterling hopes of economic opportunities promoted by T. A., Gent, (and
may even have purloined his bird list from that, or a common, source).
Putting a Pounds-Shillings-Pence sign on everything that he could, he
listed trees, other plants, fruits, mammals and birds found in the Charles-
ton area : "Here are also in the woods great plenty of wilde Turkeys, Par-
tridges, something smaller than those of England, but more de[l]icate,
Turtle Doves, Paraquetos, and Pidgeons: On the grass planes the whis-
tling Plover and Cranes and divers sorts of Birds unknowne in England."
He also listed a number of waterfowl (1682; Salley 1911:170-171).

John Lawson, as I have already noted, did not mention parakeets in his
long trip inland from Charleston in the winter of 1700-1701, suggestive
negative evidence that the "Carolina" part of the bird's name was never
more than a formality. It remained for his near-contemporary, visiting
naturalist Mark Catesby, to put South Carolina's claim to the parakeet
firmly on record (1731 :1 1) and, incidentally, to bring it to attention of the
scientific world. The latter event came to official fruition through the rest-
less genius of the great Karl von Linne, known to the Latin-mongering
elite of that time as Carolus Linnaeus. Linnaeus merely cited the species'
homeland as "Carolina" and duly provided it with an enduring specific
scientific name that says, in Latin, the same thing (1758:97). He said very
little more, for he had not a specimen but only Catesby 's plate and ac-
count from which to elaborate upon his legitimate binominal for the
species. That Catesby referred to South Carolina only needs to be re-
peated; he did not mean to include North Carolina (Wayne 1917:3; con-
sult also Catesby's places of residence in America: Frick and Stearns
1961). As to the status of the species, Catesby was sketchy and it is not
clear whether he meant to imply that the species left Carolina twice a year,
in winter and again, as the French naturalist Buffon put it, "in the love
season," to reappear later in the season of harvest (Buffon 1792-1793:235-
237). (Buffon was often misled by what he called the voice of reason — ac-
tually, his own preconceptions — and not only had no qualms about de-
moting New World forms to poor relations of Old World species, but also
held firmly to his decision that parrots only bred in the tropics: hence,
the Carolina parrot by simple calculation was but a migrant out of the
French tropical colony of Guiana.)

Thomas Pennant, who, like Buffon, had not been in Carolina, wrote at
first that "a few are found as far north as Carolina." He later amended that
view to include Virginia, but considered it mainly a migratory bird even in

88 Daniel McKinley

Carolina (1773:6; 1792, 1:282). Pennant's contemporary, John Latham,
who became a universal genius in ornithology by taking uncritically from
all previous authors, threw his net widely: "This bird inhabits Guiana,
migrating into Carolina and Virginia in Autumn." He leaves one a little
staggered by citing only Catesby for this monstrous combination (1781-
1785:227).

In the midst of all this copying from each other, it is a pleasure to record
observations of someone who actually saw a parakeet in "Carolina." The
alert Lieutenant Enos Reeves marched southward into South Carolina in
April 1782 and left a letter date-lined Congaree, Richland County, South
Carolina, 20 April. In it he related intimate details of the back country.
The countryside had changed dramatically after Charlotte had been
reached and Rowan County, North Carolina, was crossed. His group ap-
proached McCord's Ferry on the Congaree River: "Here is the first
place that I have come across the Palmetto tree or rather species of it
called the Palmetto Royal and Parrots or rather Parroquetts, and I am
told, that Alligators are to be found in this River" (1897:475-476).

William Bartram in 1791 offered first of all what everyone already knew
or at least said often enough: parakeets "are natives of Carolina and
Florida, where they breed and continue the year round." The "year
round" part may refer to Florida alone and, more critical in this case, it is
uncertain whether he meant to include South Carolina in his generaliza-
tion, for his longest stays in Carolina were in North Carolina (1958:182,
190-191). In fact, although he seems to have had substantial personal
knowledge of parakeets in North Carolina, this whole statement may sim-
ply be a bow to the Catesby tradition.

John Davis, an itinerant English tutor, spent autumn and early winter
1799 on the plantation of Thomas Drayton, apparently at Coosawhat-
chie, Jasper County. He went with his pupil on hunting forays: "we
fired in vollies at the flocks of doves that frequent the corn fields; some-
times we discharged our pieces at the wild geese, whose empty cackling
betrayed them; and once we brought down some paroquets that were di-
recting their course over our heads to Georgia" (1909:91).

Robert Mills, an early historian of the state, included the "perroquet"
in his statewide list, which included about 93 species; in county by county
enumerations of birds that frequently repetitively included passenger
pigeons, the only county specifically listed as having the "parroquet," as
he spelled it the second time, was Beaufort County (1826:101,378).

Garrulous John James Audubon must also be cited. His genius for
burying good observation amidst verbiage and glittering generality again
asserted itself. In Volume II of his Ornithological Biography he told of keep-
ing a couple of young black vultures in a coop in the yard, giving them "a

Carolina Parakeet in Carolinas 89

great number of Red-headed Woodpeckers and Parokeets, birds then
easy to procure, as they were feeding daily on the mulberry trees in the
immediate neighbourhood of my orphans" (1834:35). The area no doubt
was Charleston, at the home of the Reverend John Bachman. An Audu-
bon letter of 24 December 1833 mentions that vultures eat freshly killed
birds and that this was the second experiment of this sort, being a repeti-
tion of what he, Audubon, had performed before (Herrick 1917, 2:55).
Audubon had been at Bachman's place several times, but various cir-
cumstances indicate that the above events must have occurred during the
visit that ended with his leaving Charleston in early June 1832, after he
had spent the spring there. Both young black vultures and ripe mul-
berries would have been available in South Carolina before his de-
parture. I am confident that this is a good South Carolina record of the
parakeet and, despite its involved nature, perhaps more trustworthy than
some claims, where literary effect or promotional advantage may be sus-
pected.

Audubon's record is in fact the kind of first-hand, unstudied and spon-
taneous evidence so sadly lacking in the history of the parakeet (even in
Audubon's own formal accounts). Memories are notoriously unreliable
and yet end up being much of what little we have. Memories also be-
come encrusted with information from sources quite removed from the
original observations, if any. Samuel Scoville, Jr., an amateur or-
nithologist, visited South Carolina in May 1937. Conversation turned to
parakeets (then alleged to be abroad in the state, as will be described
later), and a landscape artist in Charleston related that his grandfather,
born in the late 1830s, "used to tell of running out into his mother's gar-
den in Charleston, when he was a little boy, to scare away the paroquets
from the orange trees. Every year, too, he would ride over to Virginia
Springs on his pony, while the rest of the family went in the family car-
riage, and on the way he would frequently see 'conures' — the Carolinian
name for paroquets" (1940:560). The story may be true in the main; its
date of around the 1840s seems acceptable, but I doubt seriously if
Americans of that time would have called parakeets by the later pet-store
name of "conure."

Scoville 's date of around 1840 agrees with the memories of George
Twiggs. He "was greatly interested in birds and . . . spent his boyhood on
plantations in Aiken (South Carolina) and Richmond (Georgia) coun-
ties, and . . . died at the age of eighty in 1930." He "never observed this
species . . . but ... his father told him that they were not uncommon in
Aiken County during his young manhood, about 1840" (Murphey
1937:24).

These memories place the species as present more or less into the

90 Daniel McKinley

1840s. Plantation owner J. Motte Alston declared, however, that it had
disappeared from the Santee River area (probably Georgetown County)
"before my day" — he was born in 1821. His grandmother recalled large
flocks of them, probably around 1780 (1953: 13). This dramatically un-
even pattern of distribution, with vague allegations of previous abun-
dance, seems to typify the parakeet over much, perhaps most, of its range.
Any popular conjuration of it as a rival of the passenger pigeon's millions
must be rejected out of hand.

Broome's attribution of breeding, year-round status to the parakeet in
South Carolina was probably uncritical (1837:65). That in the hack
historical work by Simms certainly was (1843:13). Ramsay's history con-
sidered the "perroquet" as permanent resident (1809, 2:185), as un-
informative a remark as that of Professor Gibbes (1848:vi) who prepared
a list that has been widely cited but which seems to have come straight
out of Audubon's check-list of American birds (1839:189), even pre-
serving Audubon's generic howler of Centurus for the parakeet.

A more reliable sounding record, on the other hand, has come down
from Albert Twiggs, who had a long continued interest in natural history.
As a 17-year old soldier "in the Confederate Army attempting unsuc-
cessfully to stem Sherman's march from Savannah to Beaufort and
Charleston ... he had seen a number of flocks of paroquets on the Com-
bahee River and in the pine woods between Yemassee and the coast, on
numerous occasions" (Murphey 1937:24). The time of this observation
can be calculated as late autumn 1864 and the place extreme southern
South Carolina. It seems to be one of the last observations upon the
species as a probably continuous resident in South Carolina, all other re-
ports being at widely spaced intervals.

One of these later, perhaps accidental, appearances has been described
for me by Jay Shuler (letter 1961). Dr. Eddie McClellan, an intelligent
and interested observer, had recalled that a parakeet appeared after a big
storm in 1885 and was killed with a slingshot in McClellansville,
Charleston County. Since the species still existed in considerable num-
bers in parts of Florida at that time, such an occurrence is quite possible.
But, it is also clear that there were no contemporary reports of parakeets
in the South Carolina area. Witness Walter Hoxie's suggestion that
"Parrot Ridge" on Edding Island, near Frogmore, Beaufort County, was
"a name which designates many localities hereabouts and was doubtless
bestowed by the early settlers when the gaudy Parrakeets flocked in this
region" (1886). Hoxie was a talented and experienced ornithologist and
he certainly had no first-hand knowledge of parakeets in South Carolina.
His passing comment, more etymology than ornithology to begin with,
was a poor reed for Bent to have leaned on in naming Edding Island a

Carolina Parakeet in Carolinas 91

former locality for the species in the state. No one seems to have docu-
mented Hoxie's claim that "many localities hereabouts" have the term
"parrot" or "parakeet" in their names, and I cannot even precisely place
his "Parrot Ridge."

Leverett M. Loomis, a careful student of ornithological history, turned
up no surprises in his history of certain South Carolina birds, but per-
haps was somewhat wide of the mark to place "the time of the dis-
appearance of the Paroquet from our local fauna" as about 1826 (1886).
But, his caution was commendable alongside the error of Hasbrouck
(1891:374) who indicated that Waldo Irving Burnett listed the parakeet
as present in the "pine barrens" in 1851. That unfortunate misfiring was
heard round the world. Wayne (1910), Ridgway (1916), Pearson et al.
(1919) and, as mentioned, Bent all allude to parakeets in the pine
barrens, if not Burnett by name. Burnett's paper "On the fauna of the
Pine Barrens of upper South Carolina" did indeed appear, as every-
body's bibliography says, in 1851. It is a list of species observed, "with a
few words on the 'conformability of individuals of the Fauna to each
other' — whatever they may be," as Elliott Coues put it (1878:637).
However, there is no reference to the Carolina parakeet in it.

The general failure of observers to leave definite records might justly be
called Footnote Number One to the tragedy of the parakeet in South
Carolina. Footnote Number Two came later. Paul M. Rea remarked edi-
torially in 1919: "Tradition says that many years ago nearly a dozen
Carolina Parrakeets were destroyed because they were not in sufficiently
good condition to be exhibited. Some of these specimens undoubtedly
lived in South Carolina. The Parrakeet is now almost extinct and it is not
known that a single specimen from this state is in existence" (1919:7).
"Tradition says" was no doubt just a polite way to avoid naming names
and exposing someone to ridicule for the rashness of his action.

SOUTH CAROLINA: THE PRESENT CENTURY

Footnote Number Three to the parakeet in South Carolina may or may
not have been a tragedy, for it may be that parakeets were not involved.
Many real or alleged sightings of the Carolina parakeet have come in over
the years. Nearly all such claims from the 1930s and 1940s, interestingly
enough, were from South Carolina. It is from there, of course, that there
came what, from sheer bulk of documentation, must remain the
Gran'daddy of all "rediscoveries" of the parakeet. First, let me review
summarily the body of published matter.

It all began when George M. Melamphy, working on a wild turkey pro-
ject in the Santee Swamp, Georgetown County, talked to Alexander
Sprunt, Jr., in 1933-1934, and reported several times seeing parakeets and

92 Daniel McKinley

ivory-billed woodpeckers. The sighting of the latter rare species was
finally fully substantiated, although the good news came to naught and
the species could not be saved in that area. Sprunt and Chamberlain, in
their study of South Carolina birds (1949: 292-293), later summarized the
situation and the two authors fell neatly into two positions in the parlia-
mentary arena, with Sprunt stoutly arguing for the Ayes.

In spite of, as will be seen, some equivocation by Robert P. Allen at the
time (evident from Audubon Society records), his considered opinion in
1949 was that he had not seen parakeets (op. at.: 294). Roger Tory Peter-
son also later confidently rejected the whole claim (1948:204, 207).

The contemporary published record is skimpy. John H. Baker, presi-
dent of Sprunt and Allen's parent organization, the National Audubon
Society, reported in Bird-Lore (1938) that there were no observations suf-
ficiently definite to be considered scientific, although investigations would
continue. The official pronouncement of the influential American Or-
nithologists' Union Protection Committee (1939) was negative. On the
other hand, Samuel Scoville, Jr., an amateur bird-watcher, visited the
Santee with some other people in May 1937 and managed to catch sight
of a swiftly flying bird that appeared to him to be green in color
(1940:564).

Although convinced that he had seen a parakeet, it was personal con-
viction alone for Scoville. Time magazine put it much more forcefully in a
sensationalist note in 1941 : "The Carolina parakeet . . . last reported seen
in 1904 and long thought extinct, is not. Last week an official of the
National Audubon Society confessed that a Charleston ornithologist has
been watching parakeets in the Santee River swamps for five years." Ac-
tually, not many Audubon officials would have wanted their names as-
sociated with such a "confession" by that time. Revisionary hindsight, as
has been shown above, even further eroded this ebullient pronounce-
ment.

The ghost of the Santee parakeets, however, has not remained laid.
George Laycock, field editor of Audubon (the modern name of Bird-Lore),
has blown new life into the old drama of Santee Swamp, and thinks a
negative conclusion less than scientifically proved (1969). No uncritical
sensationalist, Laycock had just proved that even in zoos where the public
record ought to have been straight from the beginning the parakeet lived
three years longer than all the official textbooks said. The orthodox had
their dates thoroughly mixed up, with misinformation from several
sources congealing into the received version. As an example of how in-
correct details accrue to an already dubious but popular conclusion, one
report even had it that the region where the birds were allegedly sighted
"has since been destroyed by a power project" (Greenway 1967:322) —

Carolina Parakeet in Carolinas 93

which is not quite the case, since the dams and reservoirs are con-
siderably upstream from the area in question.

In discounting the discounters, Laycock convinced himself that there
was some fire amidst all the smoke of the decade of the 1930s. Through
the kindness of Les Line, editor of Audubon, I have read the very con-
siderable amount of smoke generated in those faraway times. In the
following review, I plan to quote from the Audubon Society archives only
when some commentary upon the earlier published record or the good ac-
count by Laycock seems called for. Any apparent brusqueness is in the
interest of brevity. The decision is still open and I leave the reader to his
instincts.

George Melamphy, dismissed acidly by world famous ornithologist
Ludlow Griscom as "not a bird student," did have some knowledge of
wild turkeys and I can see no particular reason for him to mislead any-
one in regard to other birds. Besides, he did apparently correctly alert
Sprunt and others to the presence of ivory-billed woodpeckers. The
preliminary report by Sprunt on 10 April 1935, relating definite but un-
dated Melamphy sightings, described the region in question, some 25
miles above the delta of the Santee River, as "a tract of unbroken wilder-
ness and absolutely virgin timber." His enthusiasm was probably justi-
fied, but Audubon official Lester L. Walsh on 24 December 1937 (after
the chilling Griscom episode, to be recounted shortly) was more re-
strained in his analysis: "Lest any misapprehension exist relative to the
extent of virgin timber let me say that most of the cypress and gum in the
region gives indications of having been cut at one time or another." There
were, however, unlumbered patches of small extent and some places
judged adequate habitat for parakeets.

A "Cracker" (Sprunt's term) named W. F. "Red" Welch, who took on
the very part-time job of warden for a section of the Santee tract over
which the Audubon Society was able to gain slight control, also reported
seeing a parakeet, but he may have been shoring up his job. He sub-
mitted a couple of feathers which looked interesting enough to Allen that
he sent them to Alexander Wetmore of the U. S. National Museum for
identification. Wetmore reported them to be meadowlark feathers.

Another local man, Warren J. Shokes (described as a man of "simple
honesty" by Sprunt but who struck Griscom as "quite capable of bare
fabrication") became official warden on 1 February 1936. He reported
seeing a parakeet, with adult coloring, on 17 February. By the end of
December he had reported five sightings of the parakeet, and on
Christmas Day his son, Hollie, saw what Sprunt recorded as "a beautiful
adult Carolina Paroquet." Hollie thought the bird had a rather darker
band around the base of the neck than was shown in the picture given

94 Daniel McKinley

him. By then, in their report for the period 26 November to 12 December
1936, both Sprunt and Allen had pretty much put their stamp of ap-
proval upon the notion that there were parakeets present. Despite Allen's
later recantation, consider this: "From the details of these and previous
observations and the established nature of the evening flyway we have no
hesitation in identifying these birds as Carolina Paroquets (Conuropsis c.
carolinensis)." It is perhaps not surprising that, in the face of such
persnickity taxonomic overkill, talented and experienced Griscom should
have emphasized that "neither gentleman had had any previous ex-
perience with wild parrots at any time or place."

Things slowed down in 1937. There was a February report of a para-
keet from the elder Shokes but nothing else until 1 1 September when one
was sighted. Shokes, with what seems to me a suspicious haggling over ir-
relevant — or spurious — details, insisted that there was some "'speckl-
ing'" around the shoulders of the latter bird, although agreeing other-
wise that it was adult.

This time of lull was fated to coincide with the visit of top brass. Gris-
com and others descended in the period 7 to 16 December, during a
stretch of bad weather. Griscom obviously was in a no-nonsense mood
which, as a dean of American field ornithologists, he had some right to
be. He pronounced it "most improbable that these birds were Carolina
Paroquets; that they were more likely to be Parrots of other species that
had escaped from captivity or been released." (It might have been
questioned whether they were parrots at all.) Anyway, Griscom was con-
cerned for the good name of the Society, should all that leak to the press,
and he also wanted to keep investigations alive on the slim chance that
something might turn up.

Some of Griscom's opinions of various people involved have been cited.
It ought to be said that he considered the younger Shokes, Hollie, "a
thoroughly honest and attractive fellow," although pretty largely lacking
in critical capacities to make ornithological decisions. To exemplify his
estimation Griscom pointedly noted that Hollie 's sighting of what Sprunt
had accepted as "a beautiful adult Carolina Paroquet" had, under grill-
ing, become "a strange looking bird unlike anything he had ever seen be-
fore; that it was generally 'bluish in color with a yellow topknot,' and was
apparently catching insects on the bank of the creek." Griscom sug-
gested that "flight-lines" would not be held to by the birds over any very
long period of time; that parrots were not usually late in going to roost
(he might have pointed out that mourning doves, so like parakeets in size
and perhaps even in pattern of flight, frequently careen about quite late);
and that, even though he could be wrong about the Carolina parakeet,
which he had never seen, parrots usually called and chattered loudly

Carolina Parakeet in Carolinas 95

when flying to feeding and roosting stations.

Evidently Hollie had failed to produce any birds on another visit by
Sprunt in March 1938 (correspondence is missing). He did, allegedly
with some reluctance, report to Sprunt in mid-November that he had
seen parakeets in early June: two adults and what was presumably a
young one "being taught to fly." The adults were reported to have raised
quite a chatter. Sprunt thought this significant, for Hollie was "rather
deaf, and . . . the noise made by the birds must have been considerable
for him to hear it." Hollie had seen a lone adult — "One of those same
birds" — on the first of September. He had reported neither incident to
Sprunt spontaneously, not wanting to "stir up things again."' By that
time, however, the Shokeses were not in Audubon hire, the Santee
Sanctuary had been discontinued, and Hollie was anxious to have em-
ployment.

What it all adds up to is difficult to calculate, but I find it hard to share
Laycock's conclusion with enthusiasm. On the other hand, in the years
since 1940 various reports of surviving parakeets have come to Sprunt and
others. Nothing worthwhile ever evolved from any of them. It must be
said, however, that nobody investigates them wholeheartedly — such re-
ports are now filed (or referred) and forgotten. It is as if the hot potatoes
and burned fingers of one generation deter those who come later from
taking a chance.

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3:408.

Carolina Parakeet in Carolinas 97

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72:274-287.
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98 Daniel McKinley

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Univ. S. C. Press, Columbia; Charleston Mus. Contrib. 77. xx + 585 pp.

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don. See: Salley, 1911, pp. 161-176.

Accepted 27 November 1978

Estimates of Fish Populations in Two Northeastern
North Carolina Swamp Streams

William H. Tarplee, Jr.1

Department of ^oology, North Carolina State University,
Raleigh, North Carolina 27607

ABSTRACT. — Fish populations were sampled in two northeastern
North Carolina swamp streams, Duke and Hoggard Mill Creek, from
May through August 1972. Water conditions permitted partitioning the
streams into 0.4 km sections with nets, and estimating the abundances
of 30 species in randomly chosen areas using the Petersen method. Some
biomass estimates were also made. Population estimates in Duke
Swamp varied from 6630 to 33,734 fish per surface hectare. In Hoggard
Mill Creek estimated numbers ranged from 17,656 to 103,891 fish per
surface hectare. Large variations were found in numbers of fish among
sections of stream, but these variations were not uniform from species to
species. Biomass estimates ranged from 195 kg to 1607 kg offish per sur-
face hectare for the two streams.

INTRODUCTION

Swamp streams are generally defined as streams located in lowland
areas which flood periodically, inundating the vegetated flood plain for
extended periods during the year. These areas are usually characterized
by the presence of bald cypress, Taxodium distichum, and tupelo gum, Nyssa
aquatica. The plant communities in these systems were described in detail
by Wells (1928), Beaven and Oosting (1939), and Hall and Penfound
(1943). Although swamp stream ecosystems are known to have diverse
communities (Viosca 1928, Wharton 1970), few studies have been con-
ducted regarding the abundance of their components.

Fish populations in two swamp streams were examined in this study
from May through August 1972, in an effort to determine their composi-
tion and magnitude. Attempts were made to estimate the numerical
abundance, and in some cases the biomass, of each species.

MATERIALS AND METHODS
The Study Area

Duke Swamp and Hoggard Mill Creek are swamp streams in North-
eastern North Carolina (Fig. 1). Timber has been logged along both

Present address: Carolina Power and Light Company, P. O. Box 1551, Raleigh,
NC 27602.

Brimleyana No. 1: 99-112. March 1979. 99

100

William H. Tarplee, Jr.

TO WINDSOR'

Fig. 1. Maps of Duke Swamp (upper left) and Hoggard Mill Creek (lower right)
study areas, showing sampling sections.

streams but most areas have returned to the gum-cypress dominant forest
type. In a few recently logged areas dense stands of aquatic vegetation
and dense shrub layer have developed.

Approximately 13.7 stream km of Duke Swamp, a tributary to Lassiter
Swamp and the Chowan River in eastern Gates County, were designated
the Duke Swamp study area (Fig. 1). Hoggard Mill Creek, tributary to

Fish Population Estimates 101

Table 1. Fishes collected in Duke Swamp and Hoggard Mill Creek, May 1972
through August 1972 (P = Present). Names from Bailey et al. (1970).

Hoggard

Duke Mill

Species Swamp Creek

Bowfin, Amia calva P P

American eel, Angmlla rostrata P P

Eastern mudminnow, Umbra pygmaea P P

Redfin pickerel, Esox amencanus amencanus P P

Chain pickerel, Esox niger P P

Golden shiner, Notemigonus crysoleucas P P

Ironcolor shiner, Notropis chalybaeus P

Unidentified shiner, Notropis sp P

Creek chubsucker, Enmyzon oblongus P P

Yellow bullhead, Ictalurus natalis P P

Brown bullhead, Ictalurus nebulosus P P

Tadpole madtom, Noturus gyrinus P P

Swampfish, Chologaster cornuta P P

Pirate perch, Aphredoderus sayanus P P

Lined topminnow, Fundulus lineolatus P

Mosquitofish, Gambusia affims P

Mud sunfish, Acantharchus pomotis P P

Flier, Centrarchus macropterus P P

Banded pigmy sunfish, Elassoma zonatum P

Black banded sunfish, Enneacanthus chaetodon P

Bluespotted sunfish, Enneacanthus glonosus P P

Banded sunfish, Enneacanthus obesus P

Redbreast sunfish, Lepomis auritus P

Pumpkinseed, Lepomis gibbosus P P

Warmouth, Lepomis gulosus P P

Bluegill, Lepomis macrochirus P P

Largemouth bass, Micropterus salmoides P P

Black crappie, Pomoxis nigromaculatus P P

Swamp darter, Etheo stoma jusijorme P

Sawcheek darter, Etheostoma sernferum P

Yellow perch, Perca flavescens P

26 25

the Cashie River near its confluence with the Roanoke River, is located in
Bertie County. The Hoggard Mill Creek study area extended approx-
imately 9.3 stream km downstream from the Bertie County SR 1301
bridge on Cucklemaker Swamp and the SR 1001 bridge on Flat Swamp
(Fig. 1). The main stream channels of both streams were measured and
divided into 0.4 kilometer study sections.

Both streams varied considerably in size within the study area. Main
stream channels were generally 2 to 8 m wide and a few cm to 6 m deep.

102

William H. Tarplee, Jr.

Table 2. Population estimates of fishes collected from Duke Swamp, May through
August 1972. N = number of each species per surface hectare; CL =
80% confidence limits; B = biomass in kg per surface hectare; %N =
percent of total number; %B = percent of total biomass; * = data not
available or < 0.1; and °o = infinite.

Species

Bowfin

American eel

Eastern mudminnow

Redfin pickerel

Chain pickerel

Golden shiner

Creek chubsucker . .
Yellow bullhead ....
Brown bullhead ....
Tadpole madtom . . .

Pirate perch

Mud sunfish

Flier

Blackbanded sunfish
Bluespotted sunfish .

Banded sunfish

Pumpkinseed

Warmouth

Bluegill ..•

Largemouth bass . . .

Black crappie

Swamp darter

Sawcheek darter ....

TOTALS

Section 1

N

22

. 121

7

. 1871

40

12

CL

(62-co)
(5-oo)

(1240-4549)

(*-*)

(*-*)

(255-1003)

(541-18688)

(72-oo)

(640-2031)
(54- co)

(546- oo)

(72-co)

(126-oo)

(*-*)
(42- off)
(106-oo)

(*-*)
(*-*)
(*-*)
(12-co)

B

8.3

1.7

.1

65.5

2.6

.1

13.1

35.6

4.8

6.5

4.5

41.0
*

1.6

*

7.1

2.6

*

*
*
*

%N

.3

1.8

.1

28.2

.6

.2

5.5

14.9

1.5

16.3
1.6

17.7
2.1
3.9

.0
1.2
3.3
.2
.2
.0
.3

%B

4.3

.9

.1

33.6

1.3

.1

. 363

991

99

0

. 1082

6.7
18.2

2.5

3.3

109

2.3

. 1174

21.0

136

. 259

0

.8

2

79

3.6

217

1.3

10
10

2

22

195.1

The variation in volume of stream flow through a year had considerable
influence on stream width due to the nearly uniform topography of the
watersheds. Rises in stream level of approximately 1 m caused width in-
creases of up to 0.8 km.

Population Estimates

Four study sections were randomly selected in each stream for fish pop-
ulation sampling. Sampling was conducted during summer low water
periods using a combination of capture methods in an attempt to reduce
bias from gear selectivity. Sections were blocked off with nets, and cylin-
drical poultry-wire traps, gill nets, a seine and a backpack Smith Root
Type V electrofishing unit were used for collecting fish.

Fish Population Estimates

103

Table 2. (Continued)

Section 5

Species N

Bowfin 25

American eel 546

Eastern mudminnow ... 1300

Redfin pickerel 7050

Chain pickerel 0

Golden shiner 670

Creek chubsucker 11764

Yellow bullhead 4700

Brown bullhead 502

Tadpole madtom 0

Pirate perch 5916

Mud sunfish 7

Flier 546

Blackbanded sunfish .... 7

Bluespotted sunfish 242

Banded sunfish 0

Pumpkinseed 148

Warmouth 0

Bluegill 247

Largemouth bass 7

Black crappie 0

Swamp darter 0

Sawcheek darter 57

TOTALS 33734

CL

B

%N

%B

(*-*)

9.2

.1

1.0

(309-10381)

7.6

1.6

.8

(813-4653)

11.7

3.9

1.2

(4653-17238)

246.6

20.9

25.8

(321-oo)

6.1

2.0

.6

(6217-224102)

423.2

34.9

44.2

(2150-oa)

169.0

13.9

17.7

(250-co)

24.5

1.5

2.6

(4465-9449)

35.4

17.5

3.7

(*-*)

*

.0

(358-3010)

19.1

1.6

2.0

(*-*)

*

.0

(119-0O)

1.5

.7

.2

(91-co)

*

.4

(141-co)

2.9

.7

.3

(*-*)

*

.0

(40-oo)

.1

.2

.0

956.9

The Petersen single census mark-recapture method (Ricker 1958) was
used in estimating populations. Sampling was divided into a marking
period and a censusing period. A week generally was required for sampl-
ing each study section and at least one night separated the two periods.
During the marking period all fish captured in good condition and large
enough to mark (generally > 75 mm) were fin clipped and released. Fish
too small to mark were counted and this count was considered a minimal
population estimate. All fish (of the size marked) captured during census-
ing were inspected for marks. The same sampling effort was employed
during the marking period and the censusing period.

An indication of fish biomass was obtained from the product of the
numerical population estimates and the mean weight of a sample of fish
(by species) collected from the study streams using the same collection
techniques (Tarplee 1975). In cases where the number of individuals con-
sidered in obtaining the mean weight was small, data from 1973 rotenone
samples (Pardue et al. 1975) were also used.

104

William H. Tarplee, Jr.

Table 2. (Continued)

Species

Bowfin

American eel

Eastern mudminnow

Redfin pickerel

Chain pickerel

Golden shiner

Creek chubsucker . .
Yellow bullhead ....
Brown bullhead ....
Tadpole madtom . . .

Pirate perch

Mud sunfish

Flier

Blackbanded sunfish
Bluespotted sunfish .

Banded sunfish

Pumpkinseed

Warmouth

Bluegill

Largemouth bass 0

Black crappie 0

Swamp darter 0

Sawcheek darter 12

TOTALS 10389

Section 6

N

CL

B

%N

%B

12

82

929

. 3039

27
237

(*-*)

(40-od)

(425-oo)

(2078-6133)

(12-oo)
(lll-oo)

(1048-1969)

(484-2046)

(109-3403)

(786-4623)

(20-OO)

(1018-36934)

4.6

1.1

8.3

106.3

1.8

1.2

50.0

26.1

8.7

*

7.7

1.5

67.1

.1

.8

8.9

29.3

.3

2.3

13.4

7.0

1.7

.0

12.4

.3

18.5

1.6
.4

2.6

36.5

.6

.4

. 1391

726

178

2

. 1285

17.2
9.0
3.0

2.6

35

.5

. 1920

23.1

0

131

0

(64-oo)

(30-co)
(15-00)

(163-oo)

.8

*

2.2
3.7

1.3

.5

.2

2.9

.3

54

25

.8

. 304

1.3

(*-*)

291.1

RESULTS AND DISCUSSION

The 27 species of fish collected from Duke Swamp and 24 species
collected from Hoggard Mill Creek were typical of lowland blackwater
streams (Table 1). Differences in the species lists can be largely attribut-
ed to those smaller species less susceptible to capture, and those represent-
ed by low numbers of individuals which may actually occur in both
streams. Several additional species were reported from these areas by Par-
due et al. (1975).

Population estimates of markable-size fish in the sampled sections of
Duke Swamp and Hoggard Mill Creek are presented on a per hectare
basis in Table 2 and 3. The area of water considered was only that con-
tained within the main stream channel and did not reflect increases in sur-
face area which occurred when water levels rose above the main channel
banks.

The Petersen method is a single census technique, so confidence limits

Fish Population Estimates

105

Table 2. (Continued)

Species N

Bowfin 74

American eel 334

Eastern mudminnow . . . 801

Redfin pickerel 5960

Chain pickerel 0

Golden shiner 667

Creek chubsucker 79

Yellow bullhead 3121

Brown bullhead 1586

Tadpole madtom 0

Pirate perch 4159

Mud sunfish 35

Flier 1952

Blackbanded sunfish .... 0

Bluespotted sunfish 326

Banded sunfish 1001

Pumpkinseed 17

Warmouth 49

Bluegill 0

Largemouth bass 0

Black crappie 0

Swamp darter 0

Sawcheek darter 0

TOTALS 20161

Section 34

CL

B

%N

%B

(54-®)

27.6

.4

5.0

(208-oc)

4.7

1.7

.9

(408-od)

7.2

4.0

1.3

(5083-7440)

208.5

29.6

38.1

(566-927)

6.1

3.3

1.1

(59-420)

2.8

.4

.5

(1433-O0)

112.2

15.5

20.5

(882-30500)

77.7

7.9

14.2

(3425-5513)

24.9

20.6

4.5

(*-*)

1.5

.2

.3

(1762-2271)

68.3

9.7

12.5

(158-oo)

1.9

1.6

.3

(549-18898)

*

5.0

(*-*)

*

.1

(37-oo)

4.4

.2

.8

547.8

were estimated by considering R (recaptures) as a binomial and using
tables of confidence limits of binomial proportions (Mainland et al. 1956).
The 80 percent level of confidence was selected for use in examining these
estimates, as variability is often high in field studies of fish and wildlife
populations and an 80 percent confidence level is often adequate for
ecological evaluation and management.

There was considerable variation in numbers of fish between the sam-
pled sections of Duke Swamp, with estimates ranging from 6630 fish
weighing 195.1 kg per surface hectare to 33,734 fish weighing 956.9 kg
(Table 2). Redfin pickerel, Yellow bullhead, Pirate perch and American
eel, the most abundant fishes collected (pooled estimates), made up 69.0
percent of the estimated total. Large differences existed among species in
the variation in population estimates between study sections. For exam-
ple, Creek chubsucker estimates varied considerably between sections
while Redfin pickerel were abundant in all sections.

106 William H. Tarplee, Jr.

Table 2. (Continued)

Totals (Based on Pooled Data)

Species N CL B %N %B

Bowfin 17

American eel 1433

Eastern mudminnow ... 1065

Redfin pickerel 2933

Chain pickerel 1 33

Golden shiner 185

Creek chubsucker 381

Yellow bullhead 2310

Brown bullhead 1035

Tadpole madtom 2

Pirate perch 2110

Mud sunfish 168

Flier 554

Blackbanded sunfish .... 15

Bluespotted sunfish 49

Banded sunfish 232

Pumpkinseed 17

Warmouth 20

Bluegill 49

Largemouth bass 5

Black crappie 2

Swamp darter 2

Sawcheek darter 12

(*-*)

6.2

.1

1.8

(326-11937)

20.1

11.3

5.9

(494-2911)

9.6

8.4

2.8

(2671-3487)

102.7

23.0

30.4

(40-1334)

8.5

1.0

2.5

(143-250)

1.7

1.5

.5

(346-423)

13.7

3.0

4.1

(1574-5866)

83.2

18.1

24.6

(376-3571)

50.7

8.1

15.0

(*-*)

.0

.0

.0

(1762-2553)

12.7

16.6

3.8

(43-1542)

6.9

1.3

2.0

(484-640)

19.4

4.4

5.7

(•-•)

*

.1

(*-*)

.3

.4

.1

(59-2009)

*

1.8

(*-*)

*

.1

(*-*)

1.7

.2

.5

(*-*)

.6

.4

.2

(*_*)

*

.0

(*-*)

*

.0

(*-*)

*

.0

(*-*)

.0

.1

TOTALS 12729 338.0

Pirate perch, American eel, Yellow bullhead and Redfin pickerel were
the most numerous species in Hoggard Mill Creek, making up 79.2 per-
cent (pooled estimate) of the total (Table 3). Total estimates ranged from
17,656 to 103,891 fish per surface hectare and biomass estimates ranged
from 218.8 kg to 1606.9 kg per surface hectare among the four sections
sampled. The Pirate perch was the only species abundant in all sections.
As in Duke Swamp there was considerable variation in the number of in-
dividuals of several species between study sections. Section 2, the section
with the greatest fish density, primarily contained juvenile American eels,
while Section 21 had a density of only 74 eels per surface hectare.

In Duke Swamp 70 percent of the estimated total fish biomass was
made up of Redfin pickerel, Yellow bullhead and Brown bullhead. Most
of the additional 30 percent was composed of American eel, Flier, Pirate
Perch and Creek chubsucker. Six species — Yellow bullhead, Redfin
pickerel, American eel, Pirate perch, Flier and Mud sunfish — composed

Fish Population Estimates 107

Table 3. Population estimates of fishes collected from Hoggard Mill Creek, May
through August 1972. N = number of each species per surface hectare;
CL = 80% confidence limits; B = biomass in kg per surface hectare; %N
= percent of total number; %B = percent of total biomass; * = data not
available or < 0.1; and °° = infinite.

Species N

Bowfin 10

American eel 2780

Eastern mudminnow ... 22

Redfin pickerel 526

Chain pickerel 2

Golden shiner 32

Creek chubsucker 190

Yellow bullhead 818

Brown bullhead 2

Tadpole madtom 1426

Pirate perch 12792

Mud sunfish 314

Flier 128

Bluespotted sunfish 944

Redbreast sunfish 2

Pumpkinseed 7

Warmouth 146

Bluegill 5

Black crappie 5

Sawcheek darter 934

Yellow perch 0

Section 19

CL

B

%N

%B

(7-oo)

3.7

.0

1.7

(1619-15590)

38.9

13.2

17.8

(10- oo)

.2

.1

.1

(314-2936)

18.4

2.5

8.4

(*-*)

.1

.0

.0

(20-oo)

.2

.2

.1

(131-670)

6.8

.9

3.1

(430-15750)

29.4

3.9

13.4

(*-*)

.0

.0

.0

(66O-00)

7.2

6.8

3.3

(9953-19217)

76.6

60.7

35.0

(151-oo)

12.9

1.5

5.9

(67-co)

4.5

.6

2.1

(442-oo)

5.6

4.5

2.6

(*-*)

*

.0

(*-*)

*

.0

(106-oo)

13.0

.7

5.9

(*-*)

.1

.0

.0

(*_*)

*

.0

(423-oo)

1.2

4.4

.5

.0

.0

.0

TOTALS 21083 218.8

92 percent of the estimated fish biomass in Hoggard Mill Creek. This
comparison indicates that most of the biomass in both these streams is
made up of species desired by fisherman.

The great variation in population estimates among the sections and be-
tween the streams sampled is probably primarily due to habitat difference.
Some variation, however, may be due to changes in sampling gear ef-
ficiency in different stream types. The larger numbers of Pirate perch and
American eel in Hoggard Mill creek may be related to the greater amount
of aquatic vegetation in that stream. Section 2 of Hoggard Mill Creek con-
tained dense beds of aquatic vegetation and a soft, muddy substrate. Den-
sities of juvenile American eel and Pirate perch were much greater there
than in any other sampling area in either stream.

In the larger sections of streams studied (Section 1 in Duke Swamp and
part of Section 19 in Hoggard Mill Creek) some of the sampling gear, such

108 William H. Tarplee, Jr.

Table 3. (Continued)

Section 8

Species N CL B %N %B

Bowfin 0

American eel 1836

Eastern mudminnow . . . 996

Redfin pickerel 7144

Chain pickerel 5

Golden shiner 133

Creek chubsucker 363

Yellow bullhead 4223

Brown bullhead 0

Tadpole madtom 1 166

Pirate perch 9298

Mudsunfish 1866

Flier 2291

Bluespotted sunfish 2098

Redbreast sunfish 5

Pumpkinseed 0

Warmouth 57

Bluegill 0

Black crappie 0

Sawcheek darter 44

Yellow perch 22

.0

.0

.0

(1181-6541)

25.2

5.8

3.8

(492-oo)

9.0

3.2

1.4

(6061-9343)

249.9

22.6

37.6

(*-*)

.3

.0

.0

(*-*)

1.0

.4

.2

(232-2009)

13.1

1.2

2.0

(2644-15068)

126.9

13.4

19.1

.0

.0

.0

(549-oo)

5.8

3.7

.9

(7309-14435)

55.7

29.5

8.4

(1223-5256)

78.9

5.9

11.9

(1554-5542)

79.8

7.3

12.0

(951-oo)

12.6

6.7

1.9

(*-*)

*

.0
.0

(•-*)

5.0

.2

.8

.0

.0
.0

.0

(27-co)

.1

.1

.0

(*-*)

.9

.1

.1

TOTALS 31547 664.2

as the backpack electrofishing unit, became less effective and resulted in
capture of fewer individuals and wider confidence limits.

This variation in fish population with habitat distribution appears to be
a characteristic of these swamp systems. In this study we attempted to es-
timate the fish populations in the study area by using randomly chosen,
relatively large sampling areas. The results provide a list of species found
in these swamp streams, estimates of their abundance, and, perhaps most
important, an indication of the variation found among areas within
streams as well as between streams.

The number of fish collected that were too small to mark includes both
species with a small adult size and juvenile fishes. These fish are included
in the area species list (Table 1) and their numbers were reported by Tar-
plee (1975). Largest differences occurring between the two streams were
in Swampfish and Ironcolor shiner, which were much more abundant in
Hoggard Mill Creek than in Duke Swamp.

The estimates of fish biomass in both creeks are slightly higher than

Fish Population Estimates

109

Table 3. (Continued)

Section 2

Species

Bowfin

American eel

Eastern mudminnow

Redfin pickerel

Chain pickerel

Golden shiner

Creek chubsucker . .
Yellow bullhead ....
Brown bullhead ....
Tadpole madtom . . .

Pirate perch

Mud sunfish

Flier

Bluespotted sunfish .
Redbreast sunfish . .

Pumpkinseed

Warmouth

Bluegill

Black crappie

Sawcheek darter ....
Yellow perch

TOTALS 103891

N

CL

B

%N

%B

. 37265

(23754-106347)

.0
35.9

32.4

. 18453

(8209-oo)

165.8

17.8

10.3

. 11080

(6578-62012)

387.5

10.7

24.1

0

.0

.0

.0

30

(17-00)

.2

.0

0

.0

. 3877

(2323-21663)

139.1

3.7

8.7

0

.0

.0

.0

173

(109-3274)

.9

.2

.1

. 23951

(17880-37189)

143.5

23.1

8.9

536

(324-10232)

22.0

.5

1.4

2192

(1171-41752)

188.8

2.1

11.7

6304

(2891-oo)

37.8

6.1

2.4

0

.0

0

.0

0

.0

.0

.0

0

.0

.0

.0

0

.0

30

(17-oo)

.0

.0

.0

0

.0

1606.9

values reported for Louisiana backwater areas and rivers by Lantz (1970a,
b), and within the range of values reported for the same Louisiana
backwater area by Lambou (1959). These biomass estimates are also
slightly higher on the average than fish populations reported from North
Carolina swamp streams by Bayless and Smith (1963) and Tarplee et al.
(1971), although most values in this study were within the range reported
in the literature.

ACKNOWLEDGMENTS. — This work was supported by the N. C. Wild-
life Resources Commission, the Water Resources Research Institute of the
University of North Carolina, U. S. Division of Ecological Services, and
the N. C. Cooperative Fishery Unit. Dr. Melvin T. Huish provided guid-
ance and assistance in all phases of the study. This paper is from a thesis
submitted in partial fulfillment of the requirements for the degree of
Master of Science at North Carolina State University.

110

Table 3. (Continued)

William H. Tarplee, Jr.

Species N

Bowfin 0

American eel 74

Eastern mudminnow . . . 3902

Redfin pickerel 5864

Chain pickerel 0

Golden shiner 57

Creek chubsucker 30

Yellow bullhead 1035

Brown bullhead 0

Tadpole madtom 0

Pirate perch 6370

Mud sunfish 74

Flier 74

Bluespotted sunfish 166

Redbreast sunfish 0

Pumpkinseed 0

Warmouth 5

Bluegill 0

Black crappie 0

Sawcheek darter 5

Yellow perch 0

TOTALS 17656

Section 21

CL

B

%N

%B

(44-co)

1.0

.4

.3

(3158-5389)

35.1

22.1

10.8

(4979-7386)

205.1

33.2

63.1

.0

.0

.0

(*-*)

.4

.3

.1

(•-•)

1.1

.2

.3

(591-06)

37.2

5.9

11.4

.0

.0

.0

.0

.0

.0

(4675-8634)

38.1

36.1

11.7

(59-1404)

3.0

.4

.9

(40-co)

2.6

.4

.8

(119-co)

1.0

.9
.0
.0

.3

(•-•)

.4

.0

.1

.0

.0

.0

(*-*)

.0

.0

.0

.0

.0

325.0

LITERATURE CITED

Bailey, Reeve M., J. E. Fitch, E. S. Herald, E. A. Lachner, C. C. Lindsey, C. R.

Robins, and W. B. Scott. 1970. A list of common and scientific names of

fishes from the United States and Canada. 3rd edition. Am. Fish. Soc. Spec.

Publ. 6. 149 pp.
Bayless, Jack, and W. B. Smith. 1965. The effects of channelization upon the fish

populations of lotic waters in eastern North Carolina. N. C. Wildl. Resources

Comm., Raleigh. 14 pp.
Beaven, George F., and H. J. Oosting. 1939. Pocomoke Swamp: a study of a

cypress swamp on the eastern shore of Maryland. Bull. Torrey Bot. Club

66:367-389.
Hall, Thomas F., and W. T. Penfound. 1943. Cypress-gum communities in the

Blue Girth Swamp near Selma, Alabama. Ecology 24:208-217.
Lambou, Victor W. 1959. Fish populations of blackwater lakes in Louisiana.

Trans. Am. Fish. Soc. 88:7-15.
Lantz, Kenneth E. 1970a. An ecological survey of factors affecting fish production

in a Louisiana backwater area and river. La. Wildl. Fish. Comm. Fish. Bull.

5. 60 pp.

Fish Population Estimates 1 1 1

Table 3. (Continued)

Totals (Based on Pooled Data)

Species

Bowfin

American eel

Eastern mudminnow

Redfin pickerel

Chain pickerel

Golden shiner

Creek chubsucker . .
Yellow bullhead ....
Brown bullhead ....
Tadpole madtom . . .

Pirate perch

Mud sunfish

Flier

Bluespotted sunfish .
Redbreast sunfish . .

Pumpkinseed

Warmouth

Bluegill

Black crappie

Sawcheek darter ....
Yellow perch

TOTALS 36507 616.1

1970b. An ecological survey of factors affecting fish production in a

Louisiana natural lake and river. La. Wildl. Fish. Comm. Fish. Bull. 6. 92

pp.
Mainland, Donald, L., L. Herrera, and M. I. Sutcliffe. 1956. Tables for use with

binomial samples. Dep. Med. Stat., N. Y. Univ. College Med., New York. 79

pp.
Pardue, Garland B., M. T. Huish, and H. R. Perry, Jr. 1975. Ecological studies of

two swamp watersheds in northeastern North Carolina — a prechanneli-

zation study. Water Resour. Res. Inst. Univ. N. C. Rep. No. 105, Raleigh.

455 pp.
Ricker, William E. 1958. Handbook for computations for biological statistics of

fish populations. Bull. 119. Fish. Res. Board Can. 300 pp.
Tarplee, William H., Jr., D. E. Louder, and A.J. Weber. 1971. Evaluation of the

effects of channelization on fish populations in North Carolina's coastal plain

streams. Proc. 25th Annu. Conf. Southeast. Assoc. Game Fish Comm.:431-

446.
Tarplee, William H., Jr. 1975. Studies of the fish populations in two eastern

North Carolina swamp streams. Master's thesis. North Carolina State Univ.,

Raleigh. 66 pp.

N

CL

B

%N

%B

2

(*-*)

1.3

.0

.2

. 8569

(5409-1 3S38)

120.0

23.5

19.5

. 1940

(1483-2607)

17.5

5.3

2.8

. 3973

(3482-4571)

139.1

10.9

22.6

2

(*-*)

.1

.0

.0

133

(59-492)

1.2

.4

.2

. 222

(136-408)

8.0

.6

1.3

. 4265

(2340-761)

153.5

11.7

24.9

2

(•-*)

.1

.0

.0

. 2797

(623-2238)

14.0

7.7

2.3

. 12071

(10316-14201)

72.4

33.1

11.8

907

(521-1658)

37.1

2.5

6.0

. 1332

(825-2656)

46.6

3.6

7.6

173

(*-*)

1.0

.5

.2

2

(*-*)

*

.0

5

(2-44)

*

.0

47

(32-89)

4.1

.1

.7

2

(*-*)

.0

.0

.0

2

(*-*)

*

.0

59

(*-*)

.1

.2

.0

2

(*-*)

.1

.1

.0

1 1 2 William H. Tarplee, Jr.

Viosca, Percy, Jr. 1928. Louisiana wetlands and the value of their wildlife and
fishery resource. Ecology 9:216-229.

Wells, B. W. 1928. Plant communities of the coastal plain of North Carolina and
their successional relations. Ecology 9:230-242.

Wharton, Charles H. 1970. The southern river swamp — a multiple use environ-
ment. Bur. Business Econ. Res. Sch. Business Admin., Georgia State Univ.,
Atlanta. 45 pp.

Accepted 9 January 1979

Some Snake Food Records from the Carolinas

E. E. Brown

Box 343, Davidson, North Carolina 28036

ABSTRACT. — Some 690 food items were recovered from digestive
tracts of 479 snakes of 32 species from North and South Carolina. The
most extensive series were from the colubrids Regina septemvittata, which
contained only crayfish; Virginia striatula, which fed exclusively on
earthworms; Coluber constrictor, whose diet was varied; Opheodrys aestiuus,
which contained mostly lepidoptera larvae, plus orthoptera and
arachnids; Elaphe obsoleta, which consumed small mammals, young
birds, and birds' eggs; and the crotalid, Agkistrodon contortnx, which con-
tained mostly small mammals and lepidoptera larvae. Comments made
by other authors anent secondary ingestion and lengthy retention of in-
digestible residues are questioned.

INTRODUCTION

Studies of the food habits of snakes from the Carolinas, or closely adja-
cent areas, include those of Uhler et al. (1939), Hamilton and Pollack
(1955, 1956), Savage (1967), and Garton and Dimmick (1969). Similar
studies in other areas are those of Raney and Roecker (1947), Allen and
Swindell (1948), Clark (1949), Carpenter (1952), Barbour (1956), Brown
(1958), Klimstra (1959a,b), Bush (1959), Fitch (1960, 1963a,b), Burkett
(1966) and Wharton (1969). Wright and Bishop (1915) included some
food notes in their Okefenokee work.

This paper provides information on 690 food items from 479 snakes of
32 species, all from North and South Carolina. For better or worse, all
common and scientific names used here for reptiles and amphibians are
those provided in Collins et al. (1978).

MATERIALS AND METHODS

A great many of the snakes were road-kills, but many also were secured
alive. The presence of food in live specimens could usually be detected by
palpation, and it could then be removed by manual manipulation. Dead
specimens required dissection. Specimens with empty stomachs are not
reported. Emphasis has purposely been placed upon the taking of certain
food items rather than upon the volume of such items, especially when
many small items of relatively similar size were involved.

Brimleyana No. 1: 113-124. March 1979.

113

114 E. E. Brown

RESULTS

Nerodia erythrogaster erythrogaster, Redbelly Water Snake.

Five stomachs of this species contained 12 food items: 10 amphibians
(6 undetermined tadpoles, 4 Bufo terrestris); and 2 fish (a Micropterus
salmoides and a "sunfish").

Snakes involved were from Columbus Co., NC, and Horry and Sumter
cos., SC.

Nerodia sipedon sipedon, Northern Water Snake.

Specimens were mostly from a number of sandy-bottomed, channelized
streams of the North Carolina Piedmont and several mountain streams in
the northwestern corner of the state. Food habits in lacustrine habitats
might differ.

Forty-five items were recorded from 30 stomachs: 17 anurans (6 Bufo w.
fowleri, 3 Hyla chrysoscelis, 3 Scaphiopus holbrooki, 2 Rana catesbeiana, 1 Hyla
cruafer, 1 Rana spenocephala, and 1 undetermined); 17 salamanders (5 un-
determined larvae, 4 Desmognathus fuscus, 2 larvae of Pseudotnton sp., 1
Eurycea bislineata wilderae, 1 Plethodon glutinosus, 4 undetermined); and 11
fish (5 Semotilus atromaculatus ; 1 Etheostoma sp., 1 Ictalurus sp., 1 Hypentelium
nigricans, 1 "sunfish," 2 undetermined).

I was surprised at the prominence of amphibians (75.6 percent of all
items), and initially felt that this might correlate with the limited fish life
in the channelized streams. However, 15 of the 19 food items from the un-
channelized mountain streams were amphibians. Two of the larger
snakes had eaten Rana catesbeiana with displacement volumes of 110 and
135 ml. Three of the largest meals taken represented 26, 40 and 56 per-
cent of the weight of the respective snakes.

Specimens were from Alleghany, Ashe, Cabarrus, Caldwell, Lincoln,
Mecklenburg, and Watauga cos., NC.

Nerodia fasciata fasaata, Banded Water Snake.

Twelve stomachs of this Coastal Plain form contained 20 food items: 12
frogs and toads (6 Bufo terrestris, 3 Hyla gratwsa, 1 Hyla chrysoscelis, 2 un-
determined); 4 salamanders (2 Desmognathus auriculatus, 1 Necturus
punctatus, 1 undetermined); and 4 fish (3 Fundulus sp., 1 undetermined).
This species has been observed eating dead anurans on the highway on a
rainy June night in Brunswick County, North Carolina.

Snakes examined were from Brunswick, Carteret, Craven and Colum-
bus cos., NC, and Chesterfield and Sumter cos., SC.

Regina septemvittata, Queen Snake.

Raney and Roecker (1947), reporting on 45 stomachs from Erie
County, New York, found crayfish in 44 and a dragonfly nymph in each

Snake Food Habits 115

of 2. Wood (1949) reported the contents of 6 Ohio stomachs and 43 dis-
gorgings as all crayfish, except for a single catfish. He further proposed
that crayfish, either dead or alive, are probably taken in the soft-shelled
condition following molting. My material supports these observations.
Thirty-two stomachs contained remains of 35 crayfish, and fecal samples
from 31 additional snakes contained gastroliths from 49 crayfish. The
crayfish found in many stomachs appeared to have been soft-shelled, but,
in a few instances, I could not be certain of this. The fecal gastroliths
presumably were from crayfish at, or extremely close to, the molting
state.

The majority of crayfish were small to medium in size, but several were
relatively large. One first-year snake had eaten a very small crayfish, 20
mm total length (TL). However, a 32 cm TL snake, probably in its
second year, contained a crayfish of 50 mm TL. No more than two
crayfish were found in any stomach. Three of 31 fecal samples contained
gastroliths from three crayfish. Of 24 crayfish whose orientation was
noted, 15 had been swallowed head-first, 9 tail-first.

Specimens were from Alexander, Alleghany, Ashe, Buncombe, Cataw-
ba, Guilford, Iredell, Mecklenburg, Transylvania, Watauga and Wilkes
cos., NC.

Seminatrix pygaea paludis, Carolina Swamp Snake.

Only four specimens, all from Brunswick County, North Carolina, con-
tained food. Two had earthworm remains; a third had a very small fish
(not over 20 mm TL), partly digested and unidentified; and the fourth
contained fragments of a small, unidentified arthropod.

Storena dekayi, Brown Snake.

Eight stomachs, from Columbus, Mecklenburg and Rowan counties,
North Carolina, contained 5 earthworms and 4 small slugs.

Storena occipitomaculata occipitomaculata, Northern Redbelly Snake.

Two stomachs, from Cabarrus and Columbus counties, North
Carolina, yielded three very small (0.1 ml volume) slugs.

Thamnophis sirtalis sirtalis, Eastern Garter Snake.

Fourteen stomachs contained 16 food items: 8 salamanders (including
Pseudotriton sp., Plethodon glutinosus, Ambystoma opacum, Eurcyea bislineata, 3
undetermined); 6 anurans (1 Bufo queracus, 1 Bufo w. fowlen, 1 Hyla
chrysocelis, 1 Rana palustris, 2 undetermined); and 2 earthworms (found in
2 of 5 snakes from the mountains). One snake was observed eating small,
dead frogs on a highway in Brunswick County, North Carolina, on a
rainy June night.

116 E. E. Brown

Snakes examined were from Brunswick, Columbus, Davidson, Jackson,
Mecklenburg and Watauga cos., NC, and Hampton Co., SC.

Thamnophis sauritus sauntus, Eastern Ribbon Snake.

Thirteen stomachs yielded 14 relatively small amphibians: 11 anurans
(3 Acris gryllus, 2 Pseudacris tnseriata fenarum, 1 Pseudacris brimleyi, 2 Hyla
femoralis, 1 Hyla sp., 1 just-metamorphosed Rana catesbeiana, 1 Bufo sp.);
and 3 salamanders (2 Desmognathus fuscus, 1 Eurycea bislineata).

Snakes examined were from Ashe, Brunswick, Cabarrus, Jones and
Mecklenburg cos., NC and Colleton and Lee cos., SC.

Virginia striatula, Rough Earth Snake.

Forty specimens contained only remains of earthworms. In 15 of these
specimens the evidence consisted of dark, earthy material, plus setae in
the intestine.

Snakes were from Brunswick, Columbus, Duplin, Guilford, Mecklen-
burg and Rowan cos., NC.

Virginia valeriae valenae, Eastern Earth Snake.

Two of seven specimens examined contained only earthworm remains.
The snakes were from Mecklenburg Co., NC, and Kershaw Co., SC.

Heterodon platyrhinos, Eastern Hognose Snake.

Toads were the only food I found in this snake. Five stomachs, all from
Mecklenburg County, North Carolina, contained 6 Bufo woodhousei fowleri.
One snake, 40 cm TL and weighing 26 gm, had just swallowed a toad
weighing 29 gm, a truly gigantic meal.

Diadophis punctatus, Ringneck Snake.

These snakes feed largely upon small, slender-bodied prey, difficult to
detect by palpation. Twelve stomachs contained 12 food items: 9
salamanders (3 Plethodon jordani "metcalfi,\ 1 P. anereus, 1 P. glutinosus, 1
Eurycea quadridigitata, 3 undetermined); and 3 earthworms. All stomachs
(8) from the mountain region contained salamanders; the earthworms
were in 3 of 4 Coastal Plain snakes. One mountain specimen contained a
small lepidoptera larva (phalaenid = "noctuid") along with a salaman-
der. The salamander might have disgorged the insect larva, but I could
not be certain of this.

Specimens were from Alexander, Avery, Caldwell, Columbus, Duplin,
Macon, and Watauga cos., NC, and Dorchester Co., SC.

Carphophis amoenus amoenus, Eastern Worm Snake.

Seven stomachs yielded only earthworms; 16 others had dark, earthy
material and earthworm setae in the intestines.

Snake Food Habits 1 1 7

Snakes examined were from Caldwell, Columbus, Duplin, Mecklen-
burg and Stanly cos., NC.

Farancia erytrogramma er tyro gramma, Rainbow Snake.

In late April in Sumter County, South Carolina, a snake (86 cm TL)
was found at night laboriously attempting to swallow a mammoth tad-
pole of Rana heckscheri which it had dragged some 3 m up the bank of a
pond.

Coluber constrictor constrictor, Northern Black Racer.

The food of this snake is quite diverse. A series of 86 items from 53
stomachs included reptiles, mammals, amphibians, arthropods, birds and
a small snail, in that order of frequency and apparently all taken indepen-
dently. There was great variation in size of animal taken, from a fairly
large vole or snake to a small lepidoptera larva or lycosid spider. Reptiles
and mammals ranked almost equally in bulk and, with amphibians, com-
prised 80 percent of the food items and 95.4 percent of volume. The 38
reptiles included: 22 lizards (9 Scincella lateralis, 5 Eumeces sp., 2 Ophisaurus
sp., 2 Anolis carolinensis, 1 Sceloporus undulatus, 1 Cnemidophorus sexlineatus, 2
undetermined); 15 snakes (5 Carphophis amoenus, 3 Opheodrys aestwus, 2
Nerodia fasciata, 1 Coluber constrictor, 1 Tantilla coronata, 1 Virginia stnatula, 1
undetermined, and a fragment of shed skin); and 1 young Kinosternon
subrubrum. The 15 mammals included: 3 Microtus pinetorum (adult and
young), 1 Microtus pennsylvanicus (young), 4 Peromyscus leucopus (mostly
young), 1 Sigmodon hispidus, 1 Mus musculus and 5 undetermined. The 16
amphibians included: 6 Acns gryllus, 1 Rana sphenocephala, 1 Rana virgatipes,
2 just-metamorphosed Rana catesbeiana, 2 Hyla chrysoscelis, 2 undetermined
frogs and 2 Desmognathus fuscus. While arthropods made up 15 percent of
the items, their bulk was very limited (1.3 percent of volume). They in-
cluded 7 lepidoptera larvae, 3 lycosid spiders, 2 moths and 1 diptera
larva. Remains of a small, unidentified bird occured in each of three
stomachs (2.9 percent of volume). At least two of these were nestlings.

Sixty-four percent of the stomachs contained a single food item, but
one contained seven small frogs (mostly Acns), another had two young
mice and two Desmognathus fuscus, and a third, collected on an April morn-
ing, had three very fresh Carphophis amoenus. On a morning in early May, a
110 cm TL male racer was found vigorously swallowing a 50 cm TL male
of its own species. The hatchling eastern mud turtle found in a September
specimen was still alive, and had been in the snake's stomach for at least
two hours.

Snakes examined were from Brunswick, Cabarrus, Chatham, Colum-
bus, Guilford, Iredell, Lincoln, Mecklenburg, Montgomery, Moore,

118 E.E.Brown

Richmond, Rowan, Rutherford and Watauga cos., NC, and Abbeville,
Barnwell, Berkeley, Charleston, Dorchester, Horry, Kershaw, Lancaster,
Marlboro and Sumter cos., SC.

Masticophis flagellum flagellum. Eastern Coachwhip.

Sixteen food items were recovered from 12 stomachs: 8 reptiles (6
Cnemidophorus sexlineatus, 1 Eumeces sp., 1 35 mm CL Chrysemys concinna); 5
mammals (1 Peromyscus sp. 4 undetermined); and 3 arthropods (2 cicada
nymphs and 1 large Amblyomma tuberculatum, a tick). Both cicada nymphs
were in a small snake from Charleston County, South Carolina. The tick,
in the stomach of a larger snake, was 27 mm long and engorged with
blood. Cooney and Hays (1972) regard this tick as host specific in the
adult and nymphal stages to the tortoise, Gopherus polyphemus, and they
believe it to be the largest tick in the United States, possibly the largest
known. (A number of non-engorged ticks were collected from a large tor-
toise that was intercepted nearby.) The snake involved was found in a
shallow Gopher tortoise burrow in Jasper County, South Carolina, along
with specimens of Bufo terrestns. However, neither this snake nor another
found in a similar situation had taken a toad.

Snakes examined were from Cumberland and Hoke cos., NC, and
Charleston, Chester, Chesterfield, Edgefield, Jasper, Kershaw, Lexing-
ton, McCormick and Sumter cos., SC.

Opheodrys aestivus, Rough Green Snake.

This snake is partial to insect and arachnid prey. Fifty-nine stomachs
contained 125 food items, mostly of three major groupings: lepidoptera
(59.2 percent of items: 2 moths, 72 larvae); orthoptera (17.6 percent of
items: 15 grasshoppers, 4 field and tree crickets, 2 mantids, 1 undeter-
mined); and arachnids (13.6 percent of items: 12 spiders, 1 spider egg co-
coon, 4 phalangids). One tiger beetle, 5 undetermined insects and 4 small
snails were also present. Lepidoptera occurred almost twice as frequently
as both the other major groups combined and larvae of phalaenids,
hesperiids, goemetrids, nymphalids and sphingids were recognized. One
was only 10 mm in length. Recognizable spiders included lycosids,
araneids and attids. Stomachs contained from one to eight food items.

These snakes were collected from April to October, with 59 percent
taken during June, July and August. They were from Alexander,
Brunswick, Chatham, Columbus, Craven, Guilford, Iredell, Mecklen-
burg, Richmond and Rowan cos., NC, and Berkeley, Colleton, Dillon,
Horry and Sumter cos., SC.

Elaphe guttata guttata, Corn Snake

Ten food-laden specimens of this largely Coastal Plain form were en-
countered. They yielded 10 food items: 9 mammals (3 Microtus

Snake Food Habits 119

pennsylvamcus, 2 Peromyscus sp., 1 Oryzomys palustris, 3 undetermined
remains); and 1 unidentified bird.

Snakes examined were from Brunswick and Columbus cos., NC, and
Colleton, Dorchester, Horry and Sumter cos., SC.

Elaphe obsoleta obsoleta, Black Rat Snake.

Thirty-nine stomachs yielded 51 food items, including four sets of eggs
counted as single items. This snake appears to feed mainly on small
mammals and young birds. Mammals comprised 59 percent of the food
items (64 percent of volume) in this sample, and birds 37 percent of the
items (34 percent of volume). Half of the 30 mammals were Microtus sp.
(9) and Peromyscus sp. (6). Others were: 2 young Rattus norvegicus (one of
27 cm TL, the other smaller), 2 Tamias striatus, 2 Condylura cristata, 2
Sigmodon hispidus, 1 Mus musculus, 1 Glaucomys volans, 1 Blanna brevicauda
carolinensis, 1 Sciurus carolinensis (tail only), 1 young Sylvilagus flondanus and
2 undetermined.

One snake contained remains of two hen eggs, another had two
Carolina Wren eggs, and one from the mountains had swallowed ten Ruf-
fed Grouse eggs. Recognizable young birds from other snakes included:
American Robin, Cardinal, Yellow-breasted Chat, three Blue Jays and a
hatchling Bobwhite. In nine other cases bird remains found were not
identifiable.

One specimen had eaten a group of six adherent snake eggs, whose
size, shape, number and other features suggested the possibility of
Lampropeltis getulus. A first-year snake weighing 12 gm contained a
Scelopours undulatus.

Six snakes had taken unusually large meals that ranged from 24 to 46
percent (average 36 percent) of the weight of the snake.

Snakes examined were from Anson, Ashe, Burke, Cabarrus, Caldwell,
Iredell, McDowell, Mecklenburg, Montgomery, Stanly, Stokes, Watauga
and Wilkes cos. NC, and Aiken Co., SC.

Elaphe obsoleta quadrwittata, Yellow Rat Snake.

Food was found in two specimens from Berkeley and Horry counties,
South Carolina: a young rabbit weighing 140 gm; and fragments of
several pale blue eggs, possibly those of an Eastern Bluebird.

Pituophis melanoleucus melanoleucus, Northern Pine Snake.

A single, medium-sized specimen, road-killed in late June in Richmond
County, North Carolina, contained four Bobwhite eggs.

Lampropeltis getulus getulus, Eastern Kingsnake.

Eight stomachs yielded eight food items: 5 reptiles (1 Ophisaurus
attenuatus, 2 Opheodrys aestivus, 1 Diadophis punctatus, 1 large Lampropeltis

120 E. E. Brown

calligaster rhombomaculata) ; 2 turtle eggs, from two specimens collected in
late May and mid-June; and a decaying post-nestling Mockingbird
known to have been eaten as carrion.

Snakes examined were from Carteret, Columbus, Mecklenburg and
Surry cos., NC, and Colleton, Orangeburg and Williamsburg cos., SC.

Lampropeltis triangulum elapsoides, Scarlet Kingsnake.

Two snakes, from Berkeley and Sumter counties, South Carolina, con-
tained three specimens of the lizard, Scincella lateralis.

Lampropeltis calligaster rhombomaculata, Mole Kingsnake.

Six stomachs yielded 12 food items: 11 small mammals (1 adult
Microtus pinetorum, 10 young Microtus pennsylvamcus, 1 undetermined); and
1 snake (Opheodrys aestwus). One snake had taken three young meadow
voles which, from their sizes, were clearly from two different litters. The
largest meal, five young meadow voles, had been taken by an egg-laden
female snake in early May.

Snakes were from Mecklenburg and Rowan cos., NC, and Spartan-
burg Co., SC.

Cemophora coccinea copei, Northern Scarlet Snake.

Some early literature statements regarding food of Cemophora are rather
vague. Dickson (1948) first noted this snake's feeding on reptile eggs, and
contributions by Neill (1951), Minton and Bechtel (1958) and Palmer
and Tregembo (1970) also focused on this topic.

Most Cemophora that I examined did not contain food. A specimen from
Moore County, North Carolina, taken in late July, contained remains of
three small snake eggs (possibly Diadophis). An individual from
Brunswick County, collected in late May, had swallowed two small eggs,
apparently of a snake. A specimen collected in mid-June in Sumter
County, South Carolina, contained a collapsed egg "shell" 8 mm long,
evidently of Scincella lateralis. It was far down in the intestine and appar-
ently was going to pass through the gut undigested. This snake has been
cornered in a sunken cistern and partly swallowed, tail first, by a large
Bufo terrestns. The intestines of several snakes contained one or more small
masses of sandy or earthy material, but these masses did not include
earthworm setae. The stomachs of one or two of my early snakes con-
tained pale, yellow, fluid material. When I later saw Dickson's (1948)
note, I realized this might have been ingested egg material.

Tantilla coronata, Southeastern Crowned Snake.

Five snakes contained remains of five small centipedes and two small
beetle larvae of cucujoid type.

Snake Food Habits 1 2 1

Snakes examined were from Alexander and Cabarrus cos., NC, and
Richland Co., SC.

Micrurus fulvius Julviuf, Eastern Coral Snake.

The intestine of a single specimen for Aiken County, South Carolina,
contained smooth scales of a snake.

Agkistrodon contortnx, Copperhead.

A series of 35 stomachs yielded 62 items, chiefly small mammals and
larvae of lepidoptera. Mammal remains ranked first in volume (59.2 per-
cent) but second as percentage of total food items (37.1). Among the
remains of 23 mammals, those identifiable were: 5 Peromyscus sp., 4
Microtus pennsylvamcus, 1 Microtus pinetorum, 1 Zjapus hudsomus, 1
Reithrodontomys humulis and 1 Blarina brevicauda carolinensis. Insects ranked
second in volume (24.9 percent) and first in percentage of food items
(56.5). They included 33 larvae of lepidoptera, 1 cicada nymph and 1
adult dragonfly. The relative prominence of these larvae is due partly to
one large copperhead's having taken 20 specimens of Amsota senatona
(Citheroniidae). This caterpillar occurred also in four other stomachs.
Another snake had eaten two large saturniid moth larvae. Eight of the
nine specimens that contained larvae were collected in September or Oc-
tober. The presence of a fresh adult dragonfly in one stomach was sur-
prising. In the caudal end of the same stomach was mouse-colored fur of
some small mammal that apparently had been eaten much earlier. James
D. Brown has given me verbal permission to report that each of three
snakes whose remains he examined in northern Craven County, North
Carolina, in September, 1975, contained a large milliped, apparently of
" Spirobolus" type (Narceus).

Three of my specimens contained reptiles (6.9/4.8 percent): 1
Carphophis amoenus, 1 Diadophis punctatus and 1 Ophisaurus sp. Both snakes
had been eaten by copperheads in their first year of life. Remains of a
young bird, possibly a Rufous-sided Towhee or Blue Grosbeak, were
found in another stomach.

Two copperheads had taken especially sizable meals, 36 and 62 percent
of their weights. In only five cases out of the 35 did stomachs contain
more than one food item.

Snakes examined were from Anson, Brunswick, Burke, Caldwell, Co-
lumbus, Mecklenburg and Robeson cos., NC, and Colleton, Dillon and
Horry cos., SC.

Agkistrodon piscworus piscivorus, Eastern Cottonmouth.

Three specimens, from Horry and Berkeley counties, South Carolina,
contained food. A 395 mm TL snake had a small frog of the genus Rana,

122 E.E.Brown

and a 600 mm TL specimen contained a shrew, Cryptotis parva. The in-
testine of a 380 mm TL specimen contained mammal fur and smooth
snake scales.

Crotalus homdus, Timber Rattlesnake.

Three specimens from Alleghany and Buncombe counties, North
Carolina, had eaten four mammals: 2 Microtus pennsylvamcus, 1 Tamias
striatus and 1 Ochrotomys nuttalli.

Two young snakes, of what may or may not be recognized as the sub-
species C. h. atricaudatus, contained a young Peromyscus sp. and a Microtus
pennsylvamcus. A larger snake, 140 cm TL, had eaten a Sciurus carolinensis .
These last snakes were from Brunswick and Burke cos., NC, and Ches-
terfield Co., SC.

DISCUSSION

Neill and Allen (1956) commented rather vigorously and dogmatically
concerning "secondarily ingested food items" in snake stomachs. Their
warning was valid, but their paper, which considerably overstated the
case, is still being quoted uncritically. It is evident that some early
workers, e.g. Surface (1906), made no effort to distinguish between
primary, and possible secondary, items. No doubt there have been oc-
casional oversights and misinterpretations since those times. However, to
imply that a copperhead or black racer (among others) may not take
arthropods at times seems extremely questionable, if not actually absurd.
Savage (1967) recorded 29 cicadas in 42 copperheads from the Great
Smokies region, and his study apparently did not include a season of
periodical cicada emergence. The finding of centipedes in southern
Sistrurus by Hamilton and Pollack (1955) probably should have suggested
to the rest of us that we know too little about that snake. Although it in-
volves a totally different small viper, a figure in Copeia (1967, p. 224) is
thought provoking in this connection.

Heavier skeletal parts of some beetles are, of course, highly resistant to
digestion by snakes. However, softer materials (e.g. moths; caterpillars;
cicada nymphs; abdominal regions of spiders and of mantids and other
orthoptera; small centipedes) appear to pose no appreciable problem.

Neill and Allen (1956) also referred to lengthy retention of indigestible
food residue in the colon of such large snakes as Python and Eunectes. I may
not be qualified to comment on this, since my chief experience has been
with small, eastern forms, especially Nerodia. However, as I showed in a
later paper on water snake food (Brown 1958), such retention occurs
when a snake is without food for an extended period. A meal, or
sometimes even a massive drink of water, may stimulate it to pass such

Snake Food Habits 123

previous residue. A specimen that was feeding regularly and frequently
would pass all residue through without appreciable delay. This feature,
using a "marker" meal, was found to be useful in roughly gauging "total
digestive time" at average summer temperatures. The later ingestion of a
"chaser" meal helps to ensure that the marker meal does not dally along
the way. The observer, of course, must experiment a bit to see whether
the marker material is all being voided at one time.

ACKNOWLEDGMENTS. — I am indebted to many persons for the con-
tribution of specimens, especially the late J. Oscar Gant and Rob B.
Knox, and also to Ruben T. Mcintosh, Tom Daggy, James D. Brown,
James Gant, C. K. Bartell and John Lammers. Tom Daggy was kind
enough to identify many of the insects. Some of the field trips on which
specimens were collected were partially supported by Davidson College
Faculty Research Funds.

LITERATURE CITED

Allen, E. Ross and D. Swindell. 1948. Cottonmouth moccasin in Florida. Her-
petologica 4 (First Suppl.):l-16.

Barbour, Roger W. 1956. A study of the cottonmouth, Ancistrodon piscivorus
leucostoma Troost, in Kentucky. Trans. Ky. Acad. Sci. 77(1 ) :33-4 1 .

Bothner, Richard C. 1974. Some observations on the feeding habits of the cotton-
mouth in southeastern Georgia. J. Herpetol. #(3):257-258.

Brown, E. E. 1958. Feeding habits of the northern water snake, Natnx sipedon
sipedon Linnaeus. Zoologica (NY) 43:55-71.

Bush, Francis M. 1959. Foods of some Kentucky herptiles. Herpetologica
75(2):73-77.

Burkett, Ray D. 1966. Natural history of cottonmouth moccasin, Agkistrodon
piscivorus (Reptilia). Univ. Kans. Publ. Mus. Nat. Hist. /7(9):435-491.

Carpenter, Charles C. 1952. Comparative ecology of the common garter snake
{Thamnophis s. sirtalis), the ribbon snake {Thamnophis s. sauntus), and Butler's
garter snake (Thamnophis butleri) in mixed populations. Ecol. Monogr. 22:235-
258.

Clark, Robert F. 1949. Snakes of the hill parishes of Louisiana. J. Tenn. Acad.
Sci. 24(4):244-261.

Collins, Joseph T., J. E. Huheey, J. L. Knight and H. M. Smith. 1978. Standard
common and current scientific names for North American amphibians and
reptiles. Soc. Study Amphib. Reptiles Herpetol. Circ. No. 7. 36 pp.

Cooney, Joseph C. and K. L. Hays. 1972. The ticks of Alabama (Ixodidae:
Acarina). Ala. Agric. Exp. Stn. Bull. 426 (Auburn Univ.), 40 pp.

and 1972. Bionomics of the gopher tortoise tick, Amblyomma tuber-
culatum Marx. J. Med. Entomol. 9(3) :239-245.

Dickson, John D., III. 1948. Observations on the feeding habits of the scarlet
snake. Copeia 1 948(3) :216-217.

Fitch, Henry S. 1960. Autecology of the copperhead. Univ. Kans. Publ. Mus. Nat.
Hist. 7J(4):85-288.

124 E.E.Brown

1963a. Natural history of the racer Coluber constrictor. Univ. Kans. Publ.

Mus. Nat. Hist. 75(8) :351-468.

1963b. Natural history of the black rat snake {Elaphe o. obsoleta) in

Kansas. Copeia 1 963(4) :649-658.
Garton, John S. and R. W. Dimmick. 1969. Food habits of the copperhead in

middle Tennessee. J. Tenn. Acad. Sci. 44(4) :1 13-1 17.
Hamilton, W. J., Jr. and J. A. Pollack. 1955. The food of some crotalid snakes

from Fort Benning, Georgia. Nat. Hist. Misc. 140:1-4.
and 1956. The food of some colubrid snakes from Fort Benning,

Georgia. Ecology 37(3):519-526.
Klimstra, W. D. 1959a. Food of the racer, Coluber constrictor, in southern Illinois.

Copeia 1 959(3) :2 10-2 14.
1959b. Food habits of the cottonmouth in southern Illinois. Nat. Hist.

Misc. 168 AS.
Minton, Sherman A., Jr. and H. B. Bechtel. 1958. Another Indiana record of

Cemophora coccinea and a note on egg-eating. Copeia 1 958(1 ):47.
Neill, Wilfred T. 1951. Notes on the natural history of certain North American

snakes. Publ. Res. Div. Ross Allen's Reptile Inst. /(5):47-60.
and E. R. Allen. 1956. Secondarily ingested food items in snakes. Her-

petologica 72(3): 172- 174.
Palmer, William M. and G. Tregembo. 1970. Notes on the natural history of the

scarlet snake Cemophora coccinea copei Jan in North Carolina. Herpetologica

26(3):300-302.
Raney, Edward C. and R. M. Roecker. 1947. Food and growth of two species of

water snakes from western New York. Copeia 1 947(3) : 1 7 1- 1 74.
Savage, Thomas. 1967. The diet of rattlesnakes and copperheads in the Great

Smoky Mountains National Park. Copeia 1967(1 ):226-227.
Surface, H. A. 1906. The serpents of Pennsylvania. Zool. Bull. Pa. Dept. Agric.

IV: 113-208.
Uhler, F. M., C. Cottam and T. E. Clarke. 1939. Food of snakes of the George

Washington National Forest, Virginia. Trans. N. Amer. Wildl. Conf. 4:605-

622.
Wharton, Charles H. 1969. The cottonmouth moccasin on Sea Horse Key,

Florida. Bull. Fla. State Mus. Biol. Sci. 74(3):227-272.
Wood, John T. 1949. Observations on JVatnx septemvittata (Say) in southwestern

Ohio. Am. Midi. Nat. 42(3):744-750.
Wright, Albert H. and S. C. Bishop. 1915. A biological reconnaissance of the

Okefinokee Swamp in Georgia. II. Snakes. Proc. Acad. Nat. Sci. Phila.

67:139-192.

Accepted 4 January 1979

The Status of the Outer Banks Kingsnake, Lampropeltis
getulus sticticeps (Reptilia: Serpentes: Colubridae)

Richard M. Blaney

Department of Biology, West Virginia State
College, Institute, West Virginia 25112

ABSTRACT. — The Outer Banks Kingsnake, described as Lampropeltis
getulus sticticeps by Barbour and Engels (1942), is analyzed. Aspects of
color pattern, proportions, and scutellation characteristic of L.g. floridana
and L.g. getulus are found in this population, with no characteristics
unique enough to warrant the status of subspecies. Instead, it should be
recognized as a probably relictual, intergrade population.

The taxonomic status of kingsnakes on the Outer Banks of North
Carolina has been a source of controversy since the description of
Lampropeltis getulus sticticeps by Barbour and Engels (1942), and its subse-
quent relegation to the status of "problematical" by Wright and Wright
(1957). During the course of my work on L. getulus (Blaney 1971, 1977), I
examined a series of specimens from the Outer Banks that, when com-
pared with other material from throughout the range of the species,
showed the population to be highly variable and to exhibit characteristics
of both L.g. getulus and L.g. floridana. I therefore concluded that it should
not receive nomenclatural recognition. These conclusions, data and
photographs were provided to J.D. Lazell in 1971 for consideration at his
request. However, on the basis of several new specimens, Lazell and
Musick (1973) reported that L.g. sticticeps should be regarded a valid sub-
species. The purpose of this paper is to reexamine the data and reconsider
the interpretation of the kingsnake inhabiting the intra-Capes zone.

That this Outer Banks population is unusual is not at all in question.
The unique environment of the area, admirably described by Lazell and
Musick (1973), surely contributes to the maintenance of its integrity by
isolation. What is in question is the level of differentiation of the popula-
tion relative to the species as a whole, and the causes of its distinctiveness.

Lazell and Musick (1973) described the basic pattern of L.g. sticticeps as
"exactly like that of the nominate L.g. getulus: bold white or yellow
(usually cream) transverse bars cross the dorsum and are connected
laterally," except that in L.g. sticticeps light spotted scales occur in the
dark interspaces above scale row eight. They further stated that the spot-
ted patterns of L.g. sticticeps and L.g. floridana differ in that "in L.g.
floridana the light dorsals in the dark interspaces result primarily from
progressive lightening of each dorsal scale basally" while the spots are

Brimleyana No. 1: 125-128. March 1979. 125

126 Richard M. Blaney

"well-centered" in L.g. sticticeps. All specimens that I examined from the
Outer Banks have the basal spotting characteristic of L.g. flondana
(Blaney 1977). Even the dorsal view of the head of the Outer Banks
specimen illustrated by Lazell and Musick (1973) shows this type of
spotting.

Lazell and Musick (1973) stated that "the most striking feature of L. g.
sticticeps is the Pituophis-\ike head shape." They admitted difficulty in
quantifying this observation, and suggested that it is the rostral shape
and proportion that imparts this appearance. The rostral shape does dif-
fer from that found on most individuals of L.g. getulus from adjacent pop-
ulations, but this is a highly variable character. It is, however, the same
as in L. g. flondana. Lazell and Musick (1973) dismissed this similarity as
"coincidental, because the head shapes are not similar." The similarity in
rostral shape cannot arbitrarily be dismissed; head shapes are, indeed,
similar. Head shape varies individually in the Outer Banks population, in
L.g. getulus, and in L.g. flondana (Fig. 1). The comparative head
photographs provided by Lazell and Musick (1973) represent the ex-
tremes of this variation. They also stated that L.g. sticticeps "is propor-
tionately stouter and heavier than is nominate getulus, and this reflected in
average scale counts: dorsal rows and ventrals." Neither of these charac-
teristics reflects stoutness, a relative term at best. Stout specimens can be
found in L.g. getulus and L.g. flondana at the extremes of variation of both
characteristics (Blaney 1977). However, the number of dorsal scale rows
is significant. Lampropeltis g. getulus have a maximum of 21 dorsal scale
rows throughout its range; L.g. flondana has 23 dorsal scale rows (Blaney
1977). Another indication of L.g. flondana influence on the Outer Banks
population is that at least two specimens have more than 21 dorsal scale
rows.

Lazel and Musick (1973) presented three lines of evidence that point
away from L.g. flondana influence in L.g. sticticeps. First, L.g. sticticeps has
fewer transverse light bars. I have shown geographic variation in the
number of dorsal bands (Blaney 1977). The Outer Banks population
averages 25.3 dorsal bands compared with 21.5 and 22.6 on the adjacent
mainland, and 23.8 to the north on the Virginia coast. Southern Florida
L.g. flondana average about 54; L.g. getulus in northern Florida have the
lowest average of 18, and 32 is the high in the northern extremes of the
range. Intergradc populations of L.g. getulus and L.g. flondana in central
Florida average in the 40s, but a disjunct L.g. flondana population in ex-
treme northeastern Florida averages 25. Clearly, the number of dorsal
bands does not exclude the possible influence of L.g. flondana.

Their second argument is that "L.g. sticticeps strongly diverges from
floridana and nominate getulus in ventral counts. . ." However, analysis of

Outer Banks Kingsnake Status

127

A

?# .♦* / >* •§

Fig. 1. Variation in head shape of adult Lampropeltis getulus. TOP ROW: Outer
Banks Kingsnakes, North Carolina — A. Dare Co., Hatteras (Museum Com-
parative Zoology 128104); B. Carteret Co., Core Banks, ca 24 km n Cape Lookout
(North Carolina State Museum 12450); C. Dare Co., Hatteras Island, Buxton
(NCSM 10912); D. Carteret Co., Core Banks, 24 km n Cape Lookout (NCSM
12449). MIDDLE ROW: Lampropeltis getulus getulus, Florida— E. Gulf Co., 16 km
n Apalachicola (R.M. Blaney coll. 4997); F. Gulf Co., 8 km e Wewahitchka
(RMB 4998); G. Duval Co., 3.7 km s Ponte Verde (RMB 4937); //.Jefferson Co.,
Wacissa (RMB 4578). BOTTOM ROW: Lampropeltis getulus flondana, Florida—/.
Dade Co. (RMB 4588); J. Collier Co., Immolakee (RMB 4773); K. Dade Co.,
Homestead (RMB 4888); L. Palm Beach Co., Belle Glade (RMB 4624).

128 Richard M. Blaney

geographic variation in number of ventral scutes in L. getulus (Blaney
1977) shows that subspecies definitions relying on such counts are un-
reliable. Ecological influences on these counts must be considered. For
example, kingsnakes inhabiting the pinewoods of southeastern Louisiana
average 212 ventrals, while those inhabiting lowlands in the same
geograhic area average 206 ventrals. Such considerations would account
for the lower average number of ventrals (203) in the Outer Banks pop-
ulation than in mainland populations to the east (210) and north (209),
and certainly should not be compared with the average (215) of
kingsnakes from southern Florida. Ventral count comparisons are useful
only for delimiting populations, not subspecies.

Thirdly, Lazell and Musick (1973) dismissed the biogeographic con-
cept (Blaney 1971, 1977) of dynamic populations in their statement, "It
would be remarkable if a southern form extended hundred [sic] of km
north of its present range during a presumably colder climatic regime."
Such northward expansion, however, might be possible along the exposed
continental shelf where any climate changes would be moderated by the
ocean. Nonetheless, it was not my contention that southern forms
migrated north and outcompeted northern forms to inhabit the exposed
continental shelf. Rather, I gave evidence (Blaney 1971, 1977) that L.g.
getulus was derived from primitive L.g. floridana stocks and suggested that
the nominate form differentiated in the northern part of its range to its
present characteristics. In certain isolated areas, such as the Outer Banks
of North Carolina, and Sapelo Island, Georgia, some of the genetic traits
of the parental form (L.g. floridana) persist.

In summary, no characteristics exist in the Outer Banks population
described as L.g. sticticeps that make it unique enough for taxonomic
recognition. The characteristics present are only those of L.g. getulus and
L.g. floridana, and therefore it should be recognized as a probably relic-
tual, intermediate (= intergrade) population.

LITERATURE CITED

Barbour, T., and W.L. Engels. 1942. Two interesting new snakes. Proc. New

England Zool. Club 7:101-104.
Blaney, R.M. 1971. Systematics of the Common Kingsnake, Lampropeltis getulus

(Linnaeus). Ph.D. dissert., La. State Univ., Baton Rouge.
1977. Systematics of the Common Kingsnake, Lampropeltis getulus

(Linnaeus). Tulane Stud. Zool. Bot. 79(3 & 4):47-103.
Lazell, J.D., Jr., and J. A. Musick. 1973. The kingsnake, Lampropeltis getulus

sticticeps, and the ecology of the Outer Banks of North Carolina. Copeia

1973(3) :497-503.
Wright, A.H., and A. A. Wright. 1957. Handbook of snakes of the United States

and Canada. Vol 1. Comstock, Ithaca. 564 pp.

Accepted 22 November 1978

Notes on the Natural History of the Terrestrial Leech,

Haemopis septagon Sawyer and Shelley

(Gnathobdella: Hirudinidae)

Rowland M. Shelley and Alvin L. Braswell

North Carolina State Museum of Natural History,

P.O. Box 27647, Raleigh, North Carolina 27611

and

David L. Stephan

Department of Entomology, North Carolina State University,

Raleigh, North Carolina 27607

ABSTRACT. — The terrestrial leech, Haemopis septagon Sawyer and
Shelley, inhabits moist floodplains near water sources in North Carolina
and appears to be widespread in the Coastal Plain and eastern Pied-
mont of the state. Large earthworms seem to be the primary food source.
Some individuals display light yellowish stripes along the lateral margin,
and one reddish juvenile has been collected, possibly reflecting a
recessive genotype for color. Immature specimens have been encoun-
tered in April and are characterized by a flattened female gonopore and
reduced body size. Anatomical, ecological, and geographical similarities
with H. terrestns (Forbes) suggest a close phylogenetic relationship for
the two land leeches of North America.

The terrestrial leech, Haemopis septagon Sawyer and Shelley, inhabits the
Coastal Plain and eastern Piedmont Plateau provinces of the Carolinas
and Virginia (Sawyer and Shelley 1976). The only other land leech in the
United States, Haemopis terrestns (Forbes), occurs in the Mississippi and
Ohio River valleys from the southern Great Lakes south to Louisiana and
eastward along the Gulf Coast to Gainesville, Florida (Sawyer 1972,
Sawyer and Shelley 1976). The latter species occurs in damp soil under
rocks and logs and feeds on large earthworms (Forbes 1890, Sawyer
1972). Little is known about its life history. Sawyer and Shelley (1976)
described the anatomy of H. septagon, but because the number of
specimens was limited, and most were preserved without habitat data, no
ecological or reproductive information was provided.

In the past few years we have collected a number of specimens of H.
septagon in North Carolina, especially from the upper Neuse River Basin
of Wake County, and present the following observations on its habitat,
life history, and color variation to supplement the popularized account by
Shelley (1977). Six individuals were immature, as revealed by overall size
and the condition of the female gonopore.

Brimleyana No. 1: 129-133. March 1979. 129

130 Rowland M. Shelley, Alvin L. Braswell, David Stephan

7"*-

• ■r^L£s.-.y,".~.:.*rr

.:*£^iL-

ii^.W?- •

*-*" v^i^' Ji-."-i*?.'\.

-. jT-: ^ &. V ■' *
•a' ■ -•*& - -*>

1

apn-iM^

■■:.^!W2j;.r,.'

..;,»*..-

:^'-- ■

,-*i£^

I imm

fl

Fig. 1. Dorsal view of adult Haemopis septagon, specimen from 0.5 km nw Falls,
Wake Co., North Carolina.

Terrestrial Leech Natural History 131

As described by Sawyer and Shelley (1976), the dorsal pigment pattern
of H. septagon (Fig. 1) is typically dark olive-green with a faint,
longitudinal middorsal stripe and numerous scattered black flecks; the
venter is lighter olive-green without flecks. Variation in this pattern has
been noted among the individuals reported below. Juveniles and some
adults also display yellowish marginal stripes, which are most con-
spicuous on the leech from Duplin County (NCSM PI 77) (see locality
data below). Here the pattern resembles that exhibited by H. terrestris
(Sawyer 1972, Fig. 13F). A reddish immature leech was collected in a
sample containing five additional fully pigmented juveniles of the same
size and, presumably, the same age. This individual is uniformly red on
both surfaces, possibly because of blood in the underlying musculature,
and the eyes, middorsal stripe, and flecks are faintly visible. Its color is
similar to that of an albino specimen of Dina absolom Johansson (Erpob-
dellidae) from a cave in Yugoslavia (Johansson 1913). Other reports of
albino leeches are of white specimens, however, and include an uniden-
tified species of Philaemon (Haemadipsidae) from a cave in New Guinea
(Ewers 1974), and Erpobdella punctata punctata (Leidy) (Erpobdellidae)
from a lake in southern Michigan (Sawyer 1970). It is noteworthy that
the juvenile H. septagon was collected with five fully pigmented juveniles of
apparently the same age. Thus, its reduced pigmentation may be reflec-
tive of a recessive genotype for color.

The six juveniles mentioned above and two adults were collected on 19
April 1976 in northern Wake County. The imm^tures were generally
smaller than the adults in physical dimensions, and the female gonopore
was flattened and contiguous with the ventral surface instead of elevated
and nipple-shaped. The gonopores conformed to the species description
in location, the male being 24 annuli posterior to the oral sucker and 6 Vi-
7 annuli anterior to the female opening. Elevation of the female gonopore
is apparently achieved later in development and is not characteristic of
young individuals.

The habitat of H. septagon is similar to that mentioned earlier for H.
terrestris. The leech is usually encountered on moist floodplains in
deciduous forests; the areas immediately surrounding floodplain ponds
and backwaters of rivers and creeks seem to be preferred. Most in-
dividuals are found under or inside partially buried, rotting logs, and
some have been taken from wet leaf litter. Several specimens were collec-
ted from seepage areas on hardwood slopes. All of the leeches have been
found close to but not in standing water, in areas also inhabited by large
earthworms. During preservation some specimens regurgitated portions
of earthworms, which appear to be their primary food.

The following new localities, all in North Carolina, are reported for H.

132 Rowland M. Shelley, Alvin L. Braswell, David Stephan

HAEMOPIS TERRESTRIS
HAEMOPIS SEPTAGON

Fig. 2. Distribution of the terrestrial leeches of the United States, taken in part
from Sawyer (1972) and Sawyer and Shelley (1976). Each circle represents a
single collecting locality except for those sites occurring close together, which are
represented by one symbol. Open circles are based on literature records believed
to be valid.

septagon; coupled with those listed by Sawyer and Shelley (1976) they in-
dicate that the species is widespread in the Coastal Plain of the state. The
leech is common at the type locality in spring and summer. All specimens
referred to are deposited in the invertebrate collections of the North
Carolina State Museum of Natural History (NCSM).

Person Co.— 12.8 km ne Roxboro, off co. rd. 1518 near Mayo Cr.
(Roanoke dr.), 1 adult, 5 April 1977, A.L. Braswell (NCSM P223).
Granville Co. — 4.8 km se Creedmoor, along co. rd. 1721, 1.9 km e jet. co.
rd. 1722, under log near trib. Robertson Cr. (Neuse dr.), 1 adult, 27 April
1977, J.E. Cooper and R.E. Ashton (NCSM P227). Durham Co.— 9.9 km
sw Durham, along NC hwy 54, 0.2 km e jet. NC hwy. 751, floodplain
Third Fork Cr. (Cape Fear dr.), 1 adult, 11 July 1975, R.M. Shelley and
J.C. Clamp (NCSM PI 26). This site is about 1.6 km e of the type locality.
Wake Co. (all Neuse dr.)— along co. rd. 1918, 0.3 km se jet. co. rd. 1909,

Terrestrial Leech Natural History 133

under log in seepage area near small trib. Newlight Cr., 1 adult, 18
February 1976, A.L. Braswell and N. Murdock (NCSM P175); 5.6 km n
Cary, along US hwy 1-40, 0.5 km nwjct. co. rd. 1795, floodplain Crabtree
Cr., 2 adults, 28 March 1976, A.L. Braswell and N. Murdock (NCSM
P178); near Neuse R. at NC hwy. 98, 1 adult, 19 April 1976, A.L.
Braswell and D.L. Stephan (NCSM P181); 0.5 km nw Falls, under wet
leaves in seepage area, about 15 cm from water and 46 m from Neuse R.,
1 adult, 18 February 1976, A.L. Braswell and N. Murdock (NCSM
P176); 7.7 km sw Wake Forest, along co. rd. 2000 near Neuse R., 2
adults, 6 immatures (one reddish in color), 19 April 1976, A.L. Braswell
and D.L. Stephan (NCSM PI 80). Johnson Co.— Smithfield, in lawn or
plant bed at 831 Ward St. (Neuse dr.), 1 adult, 10 April 1977, M. Dublin
(NCSM P224). Duplin Co.— 13.6 km ne Kenansville, Goshen Swamp
(Cape Fear dr.) near NC hwy. 11, 1 adult, 6 March 1976, B.S. Martof
and J.H. Reynolds (NCSM P177). Pasquotank Co.— Elizabeth City, crawl-
ing on ground near service station (Pasquotank dr.), 1 adult, 8 March
1975, R. Mann (NCSM P186).

The similarities between H. septagon and H. terrestns are noteworthy. In
addition to terrestrial habits they have comparable color patterns, with a
dark base color and darker middorsal and lighter lateral stripes. These
markings are better defined in H. terrestns than in H. septagon. External
differences include the configuration of the adult female gonopore
(elevated in H. septagon, flattened in H. terrestris) and the annular separa-
tion of the gonopores {6V2-I annuli in H. septagon, 5-51/2 annuli in H.
terrestris). The Carolina species seems to be more dependent upon
moisture and is rarely found more than a few meters from a water source.
According to Sawyer (1972), H. terrestris is usually found "well away from
the water" and thus seems to be better adapted to terrestrial life.
Anatomical, ecological, and geographical (Fig. 2) comparisons suggest
that the two may be sister species only one step removed from a common
ancestor. Conceivably, either could have been the stock from which the
other was derived, existing relatively unchanged since the time of
divergence. They may prove to occur sympatrically in Georgia and
southern South Carolina, and future efforts should be directed toward
determining the nature of the terrestrial leech populations of these areas.

ACKNOWLEDGMENTS. — We are grateful to Roy T. Sawyer for his
constructive comments on an early draft of the manuscript. We also
thank the persons cited in the text for assistance in securing specimens of
H. septagon. Figure 1 was prepared by Renaldo G. Kuhler, North Carolina
State Museum scientific illustrator.

134 Rowland M. Shelley, Alvin L. Braswell, David Stephan

LITERATURE CITED
Ewers, W.H. 1974. Trypanosoma aunawa sp. n. from an insectivorous bat,

Miniopterus tristris, in New Guinea, which may be transmitted by a leech. J.

Parasitol. 60:172-178.
Forbes, S.A. 1890. An American terrestrial leech. Am. Nat. 24:646-649.
Johansson, L. 1913. Uber eine neue von Dr. K. Absolon in der Herzegowina

entdeckte Hohlenbewohnende Herpobdellide. Zool. Anz. 42:77-80.
Sawyer, Roy T. 1970. Observations on the natural history and behavior of

Erpobdella punctata (Leidy) (Annelida: Hirudinea). Am. Midi. Nat. 83:65-

80.
1972. North American freshwater leeches, exclusive of the Piscicolidae,

with a key to all species. 111. Biol. Monogr. No. 46; 155 pp.

and Rowland M. Shelley. 1976. New records and species of leeches

(Annelida: Hirudinea) from North and South Carolina. J. Nat. Hist.
10:65-97.
Shelley, Rowland M. 1977. North Carolina's terrestrial leech. Wildl. N.C.
47(9):24.

Accepted 8 November 1978

New Records of the Salamanders

Ambystoma talpoideum (Holbrook) and

Hemidactylium scutatum (Schlegel) in

North Carolina (Amphibia: Ambystomatidae

and Plethodontidae)

Alvin L. Braswell and Nora A. Murdock1

North Carolina State Museum of Natural History,
P.O. Box 27647, Raleigh, North Carolina 27611

ABSTRACT. — Disjunct populations of Ambystoma talpoideum, a salaman-
der which primarily occurs in the southeastern Coastal Plain and the
Mississippi Valley, were found in Union and Surry counties in the Pied-
mont. They are North Carolina's first verified records from east of the
eastern Continental Divide. Hemidactytium scutatum, formerly known in
North Carolina only from a few localities in the Piedmont and Moun-
tains, was found at three localities in the Coastal Plain of the state.
These are the first records for the species from this physiographic
province south of Virginia. Several other sites heretofore considered dis-
junct now seem to be part of a more or less continuous range.

Knowledge of a species' distributional boundaries is a prerequisite for un-
derstanding the environmental factors that govern its range. Ambystoma
talpoideum and Hemidactylium scutatum have ranges that are imperfectly
known. Both species apparently have disjunct populations that suggest
past ranges different from those that now exist (Smith 1957, Conant 1960,
Dowling 1956). Shoop (1964) and Neill (1963), respectively, provided
reviews of the A. talpoideum and H. scutatum literature. Ambystoma talpoidum
is considered a southern species with northern disjunct populations,
whereas H. scutatum is thought to be a northern species with southern dis-
juncts. This paper summarizes previously reported localities in North
Carolina, adds new records, and provides significant range extensions for
both species.

Ambystoma talpoideum

Ambystoma talpoideum, the Mole Salamander, is chiefly an animal of the
southeastern Coastal Plain and the Mississippi Valley. In the northern

Present address: U.S. Fish and Wildlife Service, Division of Wildlife Ser-
vices, 1801 Assembly Street, Columbia, SC 29201.

Brimleyana No. 1: 135-139. March 1979. 135

136 Alvin L. Braswell and Nora A. Murdock

portion of its range it is known from several disjunct localities (Shoop
1964, Conant 1975). In North Carolina, A. talpoideum has been reported
only from the southwestern Mountains: Cherokee Co. — Huheey and Stupka
(1967) [= 19.3 km w Murphy; Field Museum of Natural History =
FMNH, 10 specimens]; Henderson Co. — (Fowler and Dunn 1917;
Academy of Natural Sciences of Philadelphia, 1 specimen). Bishop (1943)
mentioned its occurrence in Transylvania County, but we have been un-
able to locate voucher specimens.

Additional North Carolina A. talpoideum localities, with numbers of
specimens following museum designations, are: Buncombe Co. — 37.0 km ne
Brevard (Charleston Museum = ChM, 30), 11.3 km ssw Asheville
(University of Michigan Museum of Zoology = UMMZ, 1 gilled), 12.1
km sw Asheville (North Carolina State Museum of Natural History =
NCSM, 16), 3.6 km nnw town of Avery Creek (NCSM, 6 + 40 gilled);
Cherokee Co.— 8.0 km w Murphy (ChM, 11), 6.4 km w Murphy (ChM, 6),
9.3 km sw Murphy (University of Kansas = KU, 1), 8.0 km sw Murphy
(NCSM, 14 gilled); Macon Co. —22.5 km wsw Franklin (NCSM, 6 + 185
gilled); Surry Co. —Pilot Mountain State Park (NCSM, 5 + 11 gilled);
Union Co. —26.2 km ne Monroe (NCSM, 10 + 131 gilled). The Bun-
combe County records probably are from the same locality ("Sandy Bot-
tom"), reported in various ways by different collectors. Investigation of
the Cherokee County records revealed that the probable locality for all
specimens previously (Huheey and Stupka 1967) and herein reported,
with the possible exception of the KU specimen, is 8.0 km southwest of
Murphy. All North Carolina localities are shown in Fig. 1.

The specimens from Union and Surry counties represent the first North
Carolina records east of the Eastern Continental Divide. Both localities
are in the Yadkin- Pee Dee River drainage of the Piedmont. Ambystoma
talpoideum is not known from this drainage in South Carolina. The Surry
County locality, only 31 km south of the Virginia border, is the most
northeastern record for the species. It is about 250 km north of the
nearest South Carolina locality near Columbia (Shoop 1964), and 212 km
northeast of the nearest mountain locality near Asheville, Buncombe
County.

The North Carolina A. talpoideum localities apparently are disjunct and
represent relict populations from a time that favored a more northern dis-
tribution. Smith (1957) attributed disjunct northern populations of A.
talpoideum in the Prairie Penninsula region to the post-Wisconsin
"Climatic Optimum" and the following "Xerothermic" period. The fac-
tors that Smith discussed may explain the North Carolina populations.
However, the climate of the post-Wisconsin period and its effects on the
southeastern United States are far from fully understood (Wright 1976).

New Salamander Records

137

Fig. 1. Distribution of (A) Ambystoma talpoideum and (B) Hemidactylium scutatum in
North Carolina. A single symbol may represent several clustered localities.
The open circle on (A) indicates a county record without precise locality
data.

Hemidactylium scutatum

Hemidactylium scutatum, the Four-toed Salamander, has a somewhat con-
tinuous range in the northeastern states and is relatively common there.
Conant (1975) considered it absent from the Atlantic Coastal Plain south
of Virginia, and it apparently is rare in the southeastern states. The
species was formerly known in North Carolina only from a few localities
in the Piedmont and Mountains: Cherokee Co. — near Andrews (Huheey
and Stupka 1967; FMNH, 2); Durham Co.— Duke Forest (Gray 1941;
UMMZ, 1); Graham Co.— near Topton (Huheey and Stupka 1967;
UMMZ, 1); Wake Co.— 8 mi. [12.9 km] nw Raleigh (Hurst 1964; NCSM,
1). Recent discoveries indicate that it is more widely distributed.

Additional North Carolina localities for H. scutatum are: Bladen Co. —
10.5 km ese Kelly (NCSM, 3); Buncombe Co. — 3.6 km nnw town of Avery
Creek (NCSM, 3 + 1 larva); Chatham Co.— -2.2 km ssw Wilsonville

138 Alvin L. Braswell and Nora A. Murdock

(NCSM, 7 + 42 larvae); Cherokee Co.— 8.0 km sw Murphy (NCSM, 14
larvae), 2.0 km ese Andrews (KU, 10); Cumberland Co. — near Linden
(NCSM, 1); Duplin Co.— A. 2 km w Wallace (NCSM, 2); Henderson Co.—
2.4 km w Etowah (NCSM, 1); Macon Co. — 22.5 km wsw Franklin
(NCSM, 9 + 7 larvae), 16.9 km sw Franklin (NCSM, 1); Orange Co.— 9.1
km nnw Chapel Hill (Duke University, 9 + larvae; NCSM, 2); Person
Co.— 3.2 km se Bethel Hill (NCSM, 2), 12.9 km ne Roxboro (NCSM, 2);
Polk Co. — near Tryon (National Museum of Natural History = USNM,
1); Rockingham Co.— 12.1 km s Reidsville (NCSM, 1); Surry Co.— Pilot
Mountain State Park (NCSM, 4); Wake Co.— 14 additional localities
(NCSM, 105). Most specimens are females that were collected with their
eggs. All localities are shown in Fig. 1.

The Bladen and Duplin County localities are 144 km south-southeast
and 126 km southeast respectively, of the nearest Piedmont locality in
Wake County. Excluding an old and questionable record from
Charleston, South Carolina (Neill 1963), the localities in Bladen, Duplin,
and Cumberland counties are the first from the Atlantic Coastal Plain
south of Virginia.

More field work is needed before any definite conclusions can be made
about disjunct or relict populations of H. scutatum. Since Neill (1963),
several authors have provided additional records of H. scutatum in the
south: Mount (1975, Alabama); Dundee (1968, Mississippi); Fugler and
Folkerts (1967, Florida); Carter (1968, Oklahoma); and Watkins (1969,
Missouri). These records are reflected in Conant's range map (1975).
Although somewhat restricted in its habitat requirements, this secretive
salamander appears more widely and uniformly distributed than
previously thought. Several sites heretofore considered disjunct now seem
to be part of a more or less continuous range.

ACKNOWLEDGMENTS. — We thank Bernard S. Martof, N.C. State
University, for his criticisms of the manuscript, and William M. Palmer,
N.C. State Museum, for his help with the manuscript and for providing
information on major museum holdings. Drs. R.C. Bruce and J.R. Bailey
contributed much to our current knowledge of H. scutatum in the state. S.
Alford, J.W. Braswell, Jr., J. Brellenthin, J. Clayton, E.E. Flowers, J.
Gillespie, D. Lee, D. Lockwood, G. Mortoro, J.C. Nicholls, Jr., J.H.
Reynolds, F. Scott, R.C. Shoop, D.L. Stephan, and R. Yates helped in
various ways. The Highlands Biological Station, Highlands, was a base of
operations for some of the field work.

New Salamander Reocrds 1 39

LITERATURE CITED

Bishop, Sherman C. 1943. Handbook of Salamanders. Comstock, Ithaca. 555 pp.
Carter, William A. 1968. First record of Hemidactylium scutatum in Oklahoma. Her-

petologica 24(4) ^328.
Conant, Roger. 1960. The queen snake, Natrix septemvittata, in the Interior

Highlands of Arkansas and Missouri, with comments upon similar dis-
junct distributions. Proc. Acad. Nat. Sci. Phila. 7 72(2):25-40.
1975. A Field Guide to Reptiles and Amphibians of Eastern and Central

North America. Houghton Mifflin, Boston. 429 pp.
Dowling, Herndon G. 1956. Geographic relations of Ozarkian amphibians and

reptiles. Southwest. Nat. 7(4): 174-1 89.
Dundee, Harold A. 1968. First record of the four-toed salamander, Hemidactylium

scutatum, in Mississippi, with comments on its disjunct distribution in

Arkansas and Louisiana. J. Herpetol. 7(1-4) : 101-1 03.
Fowler, H.W., and E.R. Dunn. 1917. Notes on salamanders. Proc. Acad. Nat.

Sci. Phila. 69:7-28.
Fugler, Charles M., and G.W. Folkerts. 1967. A second record of Hemidactylium

from Florida. Herpetologica 23(1 ):60.
Gray, I.E. 1941. Amphibians and reptiles of Duke Forest and vicinity. Am. Midi.

Nat. 25(3):652-658.
Huheey, James E., and Arthur Stupka. 1967. Amphibians and Reptiles of Great

Smoky Mountains National Park. Univ. Tennessee Press, Knoxville, 98

pp.
Hurst, George A. 1964. Notes on Hemidactylium scutatum and Hyla squirella in Wake

County, North Carolina. Bull. Phila. Herpetol. Soc. 72(1-4):19.
Mount, Robert H. 1975. The Reptiles and Amphibians of Alabama. Auburn

Univ., Agric. Exp. Stn., Auburn. 347 pp.
Neill, Wilfred T 1963. Hemidactylium scutatum. CAT AMER AMPHIB REPT: 2.1-

2.2.
Shoop, Robert C. 1964. Ambystoma talpoideum. CAT AMER AMPHIB REPT: 8.1-

8.2.
Smith, Philip W. 1957. An analysis of post- Wisconsin biogeography of the Prairie

Peninsula region based on distributional phenomena among terrestrial ver-
tebrate populations. Ecology 35:205-218.
Watkins, Larry C. 1969. A third record of the four-toed salamander, Hemidactylium

scutatum, in Missouri. Trans. Kans. Acad. Sci. 72(2):264-265.
Wright, H.E., Jr. 1976. The dynamic nature of Holocene vegetation: A problem

in paleoclimatology, biogeography, and stratigraphic nomenclature. Quat.

Res. (N.Y.) 6:581-596.

Accepted 16 November 1978

An Albino Sculpin from a Cave in the New River Drainage of
West Virginia (Pisces: Cottidae)

James D. Williams

Office of Endangered Species,
U. S. Fish and Wildlife Service, Washington, D. C. 20240

AND

William M. Howell

Department of Biology, Samford University,
Birmingham, Alabama 35209

ABSTRACT. — An albino Banded sculpin, Cottus carolinae, with atypical
morphological features, is reported from Buckeye Creek Cave, Green-
brier County, West Virginia. This is the first report of albinism in the
genus. One atypical feature of the specimen is presence of a frenum, a
structure unknown in any member of the family Cottidae. Other
atypical features include enlargement of the cephalic canal pores and fu-
sion of the postmandibular pores, and dorsal spine and pelvic fin ray
counts that are unusual for a member of the C. carolinae species group.
The atypical features are described and compared with those of nor-
mally pigmented C. carolinae, which also occur in the cave.

INTRODUCTION

A male albino Banded sculpin (Fig. 1), 67 mm standard length (SL),
was collected in Buckeye Creek Cave by J. A. Stellmack on 3 September
1967. We are reporting this specimen as the first record of albinism in the
genus Cottus, family Cottidae. It is somewhat surprising that this condi-
tion has not previously been reported for sculpins considering their fre-
quent occurrence in subterranean waters. The specimen is of particular
interest since it has several atypical morphological features in addition to
the albinistic condition.

Buckeye Creek Cave is located approximately 4 km southwest of Ren-
ick, Greenbrier County, West Virginia. The entrance to the cave is in a
shallow limestone sink in a hillside alongside a pasture. A small stream 1
to 2 m wide which flows into the cave appears to be a tributuary of Spring
creek, Greenbrier River drainage. This drainage is developed on lime-
stone and has many subterranean streams. The albino sculpin was col-
lected in the aphotic zone of the cave, approximately 200 m from the en-
trance. It was sighted and taken in an open area in clear water approx-
imately 15 cm deep over a gravel bottom. At the time of capture the body

Brimleyana No. 1: 141-146. March 1979. 141

142

James D. Williams and William M. Howell

Fig. 1. Albino Banded sculpin, Cottus carolinae, (67 mm SL) from Buckeye Creek
Cave, Greenbrier County, West Virginia, collected 3 September 1967.

was white to cream colored, without any dark pigment. Eye color was not
noted at that time (J. A. Stellmack, pers. comm.).

On 7 August 1970 we visited Buckeye Creek Cave to search for ad-
ditional specimens of Cottus. No albinos were seen, but 48 normally
pigmented Cottus carolinae, ranging from 18 to 91 mm SL, were collected
inside and outside the cave. We found specimens to a distance of 400 m
inside the cave, which was as far as we explored.

RESULTS AND DISCUSSION

Other atypical features present in the Buckeye Creek Cave albino in-
clude: (1) a frenum; (2) enlargement of the cephalic canal pores and fu-
sion of the postmandibular pores; and (3) dorsal spine and pelvic fin ray
counts that are atypical for a member of the Cottus carolinae species group
(Table 1). The presence of a frenum in the albino sculpin is most un-
usual. This condition is not known to occur in any other member of the
genus Cottus or in the family Cottidae.

Enlargement of the cephalic canal pores and fusion of the postman-
dibular pores may be associated with the albinistic condition, but are
more likely results of conditions in the cave environment. McAllister
(1968) reported enlarged pores on sculpins from poorly lighted environ-
ments, such as deep lakes and muddy water, compared with sculpins in
well lighted environments. Fusion of the two postmandibular pores into a
single large pore is a condition that rarely occurs in surface dwelling
sculpins. Seven of the 20 specimens (29-91 mm SL) of the 48 normally-
pigmented specimens taken inside Buckeye Creek Cave and outside
around the entrance had the postmandibular pores fused into a single
pore.

Albino Sculpin from Cave

143

Table 1. Frequency distribution of numbers of fin rays and lateral line pore
counts for Cottus bairdi and C. carolinae from the New River drainage
(Robins 1954); a recent collection of C. carolinae from Buckeye Creek
Cave (New River drainage); and an albino C. carolinae from Buckeye
Creek Cave.

Dorsal

spines

6

7

8

N

X

Cottus bairdi

New River drainage

Cottus carolinae

New River drainage
Buckeye Creek Cave
Albino

1

70

27
28

64

69

2

134

96

30

1

7.48

7.72
7.07
6.00

Dorsal

rays

15

16

17

18

N

X

Cottus bairdi

New River drainage

Cottus carolinae

New River drainage
Buckeye Creek Cave
Albino

11
3

68

46

25
1

55

50

2

2
1

136

97

30

1

16.35

16.54
15.97
16.00

Dorsal spines

; plus rays

22

23

24

25

26

N

X

Cottus bairdi

New River drainage

Cottus carolinae

New River drainage
Buckeye Creek Cave
Albino

7

3
1

32

12
23

65

36

4

27
42

1

131

91

39

1

23.86

24.35
23.03
22.00

Anal rays

11

12

13

14

N

X

Cottus bairdi

New River drainage

Cottus carolinae

New River drainage
Buckeye Creek Cave
Albino

6

1

2

75

36

13

1

52

59
14

2
1

135

97

29

1

12.37

12.62
12.41
12.00

144 James D. Williams and William M. Howell

Table 1 (Continued)

Pectoral rays (left and right fins)

13

14

15

16 17 18

N

X

Cottus bairdi

New River drainage

Cottus carolinae

New River drainage
Buckeye Creek Cave
Albino

11

96

185

3

1

20

47 129 17
35 25
1

312

196
60

2

14.69

16.88
16.42
15.50

Pelvic rays (left and right fins)

3

4 N

X

Cottus bairdi

New River drainage

Cottus carolinae

New River drainage
Buckeye Creek Cave
Albino

37
2

235 272

182 182
60 60

2

3.86

4.00
4.00
3.00

Lateral line pores

18

19

20

21 22 23

24

25 26

Cottus bairdi

New River drainage

Cottus carolinae

New River drainage
Buckeye Creek Cave
Albino

1

6

8

10 18 25

14

2
2

11 5

6 7
3 5
1

Lateral line pores

27

28

29

30 31 32

33

N X

Cottus bairdi
New River drainage

Cottus carolinae

New River drainage
Buckeye Creek Cave
Albino

21
5

38
1

8

2 2 8

3

100 22.68

97 28.04
16
1 25.00

Albino Sculpin from Cave 145

Two atypical fin ray counts were observed in the albino. The low dor-
sal spine count (6) is unusual for species of the Cottus bairdi or C. carolinae
complex. Three pelvic rays in the albino are atypical for the C. carolinae
complex, but is not unusual for, and can be characteristic of, some species
of the C. bairdi complex. All fin rays except caudal rays were unbranched,
which is the usual condition in the genus. Relyea and Sutton (1973)
reported a cave population of Yellow bullheads, Ictalurus natalis, in which
some individuals had no pelvic fins while others had a deformed caudal
fin and a deformed, reduced or absent adipose fin. These bullheads were
somewhat depigmented but not albinistic.

The preopercular armature of the albino sculpin consists of one
moderate-to-large upturned spine with two smaller spines below it. The
dentition is well developed, with the palatine patch almost touching the
vomerine patch. In length the palatine patch is about equal to the width
of the vomerine patch. The preopercular armature and dentition are
typical for C. carolinae.

Body proportions of the albino fall within the range of variation of both
C. carolinae and C. bairdi as reported by Robins (1954). Body proportions,
expressed as thousandths of standard length, are: head length/346; eye
length/82; lateral line length/845; body depth/224; caudal peduncle
depth/82; caudal fin length/228; pelvic fin length/ 187; pectoral fin
length/262.

There have been numerous reports of albinism in fishes (see Dawson
1964, 1966, 1971), most of which reported the albinistic condition but did
not note any unusual or atypical morphological features associated with
albinism. Bridges and Limbach (1972) demonstrated through breeding
tests that albinism in Rainbow trout, Salmo gairdneri, is a simple auto-
somal recessive character. They found no significant pleiotropic effect of
the mutant gene.

Speciation can be expected to be greater in the folded limestones of the
Appalachians than in the flat-bedded limestones of, say, the Interior Low
Plateaus because dispersal routes are more likely to be disrupted and
populations isolated. Examples of several invertebrates which evolved in
isolated cave systems in the Greenbrier Valley of West Virginia were
mentioned by Culver et al. (1974). Besharse and Holsinger (1977)
described a new species of subterranean salamander from a cave in this
valley. We recognize that this albino may represent a "stray" from a true
cave-adapted population which is specifically distinct. However, the atyp-
ical features of the specimen, i.e., presence of a frenum, unusual dorsal
and pelvic fin ray counts, and enlarged cephalic canal pores, may be en-
vironmentally induced. We feel that the assignment of a specific name is

146 James D. Williams and William M. Howell

not appropriate at this time, and hope that this report will stimulate spe-
leologists and ichthyologists to search for additional albino Cottus in the
caves of this area.

ACKNOWLEDGMENTS. — We wish to express our gratitude to the
following individuals for their assistance with the preparation of this
paper. John E. Cooper, North Carolina State Museum of Natural His-
tory, and C. Richard Robins, University of Miami, reviewed and made
comments on the manuscript. J. A. Stellmack, Pennsylvania State Uni-
versity, collected and donated the specimen to John Cooper and provided
information relative to its capture. James F. McKinney photographed the
specimen. Field work was supported by a Samford University Faculty
research grant to W. M. Howell.

LITERATURE CITED

Besharse, Joseph C, and J. R. Holsinger, 1977. Gynnophilus subterraneus, a new

troglobitic salamander from southern West Virginia. Copeia 1977 (4):624-

634.
Bridges, W. R., and B. Limbach. 1972. Inheritance of albinism in rainbow trout.

Heredity 63(3): 152- 153.
Culver, David, J. R. Holsinger, and R. Baroody. 1972. Toward a predictive cave

biogeography: the Greenbrier Valley as a case study. Evolution 27(4) :689-

695.
Dawson, C. E. 1964. A bibliography of anomalies of fishes. Gulf Res. Rep.

7(6):308-399.
1966. A bibliography of anomalies of fishes, supplement 1. Gulf Res. Rep.

2(2): 169- 176.
.. 1971. A bibliography of anomalies of fishes, supplement 2. Gulf Res. Rep.

3(2) :2 15-239.
McAllister, Don E. 1968. Mandibular pore pattern in the sculpin family Cottidae.

Natl. Mus. Can. Bull. 223(6):58-69.
Relyea, Kenneth and B. Sutton. 1973. Cave dwelling yellow bullheads in Florida.

Fla. Sci 30(l):31-34.
Robins, C. Richard. 1954. A taxanomic revision of the Cottus bairdi and Cottus

carolinae species groups in eastern North America. (Pisces, Cottidae). Ph.D.

Dissert., Cornell Univ., Ithaca. 248 pp.

Accepted 10 January 1979

Occurrence of the Milliped Pachydesmus

crassicutis incursus Chamberlin in the

Kings Mountain Region of North Carolina

and the Coastal Plain of South Carolina

(Polydesmida: Xystodesmidae)

Rowland M. Shelley

North Carolina State Museum of Natural History,
P.O. Box 27647, Raleigh, North Carolina 2761 71

AND

Marianne Filka

Department of £00 logy, North Carolina State
University, Raleigh, North Carolina 27607

ABSTRACT. — The milliped Pachydesmus crassicutis incursus Chamberlin
is reported from specific localities in the Kings Mountain region of
North Carolina, the northeastern limit of the genus, and the Piedmont
and southern Coastal Plain of South Carolina. Males are generally
larger than females, and individuals of both sexes are generally darker in
color and larger in the Kings Mountain region than in the southern part
of the range. Gonopodal variation is evident in the apical portion of the
secondary tibiotarsus, especially in the subterminal process, which may
be blunt, pointed, or absent. The secondary tibiotarsus is longer than
the primary branch in individuals from southern South Carolina, but
the two structures are subequal in specimens from the Kings Mountain
region.

Millipeds of the genus Pachydesmus are the largest polydesmoids in
North America. They occur in the southeastern states from South
Carolina and Tennessee to eastern Texas, but tend to be secretive and are
generally encountered less frequently than other xystodesmids. Hoffman
(1958) recognized two species: Pachydesmus clarus (Chamberlin) and P.
crassicutis (Wood), the latter represented by eight subspecies. The Mis-
sissippi River divides their ranges, with the former occurring on the
western side and the latter on the eastern side. The easternmost represen-
tative, P. crassicutis incursus Chamberlin, inhabits the foothills of Georgia
and South Carolina adjacent to the southeastern edge of the Blue Ridge
Mountains, and is known definitely from Tallulah Falls, Habersham
County, Georgia; and Easley, Pickens County, and Taylors, Greenville

Adjunct Assistant Professor of Zoology, North Carolina State University.

Brimleyana No. 1: 147-153. March 1979. 147

148 Rowland M. Shelley and Marianne Filka

County, South Carolina. The sole North Carolina record is a male from
an unspecified locality in the mountains, which Hoffman surmised was
probably either near Tryon, Polk County, or "in Macon County between
Franklin and Clayton, Ga."

In the summers of 1975 and 1976 we discovered P. c. incursus in moist
seepage areas in the Kings Mountain region of North Carolina
(Cleveland and Gaston counties), just across the border from South
Carolina, and can now confirm its occurrence in the state. This area
might be the one referred to previously, as it consists of several peaks over
300 m, two of which, the Pinnacle of Kings Mountain and Crowders
Mountain, rise to 520 m and 495 m, respectively. However, the region is
located in the south-central Piedmont Plateau Province, about 88.5 km
east of the Blue Ridge Front, and lies within the range postulated by
Hoffman for the subspecies. Nevertheless, the prior North Carolina
record could conceivably be from the Kings Mountain region, since to our
knowledge Pachydesmus has never been authentically collected from the
Appalachian Mountain portion of the state.

Wray (1967) reported "Pachydesmus retrorsus" from "probably valley of
French Broad River," western North Carolina. The specimen that he
cited, however, was the questionable one from North Carolina that
Hoffman had earlier (1958) identified as P. c. incursus. Thus, P. retrorsus
does not occur in North Carolina and is hereby deleted from the fauna of
the state. This error could have been avoided by simply reviewing
Hoffman's paper, which also reduced P. retrorsus to a subspecies of P.
crassicutis.

In the summers and springs of 1976-1977 the senior author collected P.
c. incursus from 16 localities in the South Carolina Piedmont and two in
the lowland region of the Coastal Plain. Of the Coastal Plain specimens,
those from Barnwell County were found along the edge of a swamp in
association with Sigmona latior hoffmam Shelley, and the female from
Jasper County was found under moist leaves in a live oak thicket. In the
Piedmont, the milliped was encountered primarily in hardwood localities
near water sources but was also taken from pine areas with thick vines
and undergrowth. One female from Newberry County was discovered un-
der pine-bark mulch on a walkway at Molly's Rock Picnic Area, Sumter
National Forest, in a primarily pine forest. These collections indicate that
the subspecies is common in the Piedmont and Coastal Plain of South
Carolina and can probably be collected from similar habitats south of the
Savannah River in Georgia.

The known North Carolina and new South Carolina localities for P. c.
incursus are listed below. Except for two samples in the Florida State
Collection of Arthropods (FSCA) and one in the American Museum of

New Milliped Records 149

Natural History (AMNH), all specimens are deposited in the inver-
tebrate collection of the North Carolina State Museum of Natural History
(NCSM).

NORTH CAROLINA: Cleveland Co.— 6.6 km sw Kings Mtn. (town),
along co. rd. 2245 at crossing of Dixon Branch Cr., 0.3 km nwjct. co. rd.
2283, 2 c3\ 39 (1 juv.), 16 August 1975, R.M. Shelley and J.C. Clamp
(NCSM A537); 9.3 km s Kings Mtn. (town), along co. rd. 2245, 0.2 km n
jet. co. rd. 2288, 3 9, 16 August 1975, R.M. Shelley and J.C. Clamp
(NCSM A541); 9.1 km sw Kings Mtn. (town), along co. rd. 2283, 1.3 km
ne jet. NC hwy. 216, ltf, 8 July 1976, M. Filka and W.W. Thomson
(NCSM A1060); and 4.8 km s Kings Mtn. (town), along co. rd. 2289, 1.0
km w jet. NC hwy. 161, 2 tf, 18 October 1976, M. Filka and G. Wicker
(NCSM A2239). Gaston Co.— 8.5 km sw Gastonia, along co. rd. 1122 at
Crowder Cr., 1.4 km w jet. co. rd. 1131, 2 tf, 2 9, 16 August 1975, R.M.
Shelley and J.C. Clamp (NCSM A547); 6.4 km sw Gastonia, along co.
rd. 1126, 0.8 km s jet. co. rd. 1113, 1 9, R.M. Shelley and J.C. Clamp
(NCSM A549); 7.7 km sw Gastonia, along co. rd. 1131, 0.2 km nwjct. co.
rd. 1133, 1 9, M. Filka and W.W. Thomson (NCSM A1091); and 6.8 km
w Gastonia, along co. rd. 1106, 2.4 km ejet. co. rd. 1236, 1 ^, 16 October

1976, M. Filka and G. Wicker (NCSM A2255).

SOUTH CAROLINA: York Co.— Kings Mtn. State Park, tf, 29, 20
August 1976, R.M. Shelley (NCSM A1361). Chester Co.— 21.9 km w
Chester, Woods Ferry Recreation Center, Sumter National Forest, 6<3\ 4
August 1976, R.M. Shelley (NCSM A1363); and 9, 1 May 1977, R.M.
Shelley (NCSM A1504). Union Co.— 11.7 km s Union, SC hwy. 16 at
Tyger R., 9, 5 August 1976, R.M. Shelley (NCSM A1364); and 10.3 km
nw Union, SC hwy. 279 at Fair Forest Cr., 29, 2 May 1977, R.M. Shelley
(NCSM A1509). Newberry Co.— 14.9 km ne Newberry, Molly's Rock Pic-
nic Area, Sumter National Forest, 9> 5 August 1976, R.M. Shelley
(NCSM A1365); 8.5 km s Whitmire, SC hwy. 32 at Indian Cr., 9, 2 May

1977, R.M. Shelley (NCSM A1520); 18.8 km ne Newberry, SC hwy. 81
at Enoree R., 3^, 29, 5 August 1976, R.M. Shelley (NCSM A1366); and
19.3 km sw Newberry, jet. SC hwys. 72 and 211, $, 6 August 1976, R.M.
Shelley (NCSM A1367). Saluda Co.— 8.5 km ne Saluda, along SC hwy.
39, 1.4 km n jet. SC hwy. 450, 9, 4 May 1977, R.M. Shelley (NCSM
A1527). Greenwood Co. — 19.1 km s Greenwood, along SC hwy. 48, 0.2 km
w jet. SC hwy. 63, tf, 9 August 1976, R.M. Shelley (NCSM A1371); and
19.3 km se Greenwood, unnumbered rd. at Cuffytown Cr. along edge of
Sumter National Forest, 9, 9 August 1976, R.M. Shelley (NCSM
A1372). Oconee Co. — 9.1 km se Oakway, SC hwy. 66 at Beaverdam Cr., tf,
10 June 1978, R.M. Shelley and W.B. Jones (NCSM A2062). McCormick
Co. — 6.6 km se Plum Branch, SC hwy. 21 at Stephen's Cr., 9, 5 May

150 Rowland M. Shelley and Marianne Filka

Fig. 1. Dorsal view of 9 Pachydesmus crassicutis incursus from 9.3 km s Kings
Mountain (town), Cleveland County, North Carolina. Total length 63.8 mm.

1977, R.M. Shelley (NCSM A1535); Baker Creek State Park, 3tf, 9, 8
August 1976, R.M- Shelley (NCSM A1369); Hickory Knob State park,
9, 9 August 1976, R.M- Shelley (NCSM A1370); and 7.4 km n Mt. Car-
mel, SC hwy. 46 at Little R., tf, 9, 5 May 1977, R.M. Shelley (NCSM
A1542). Laurens Co. — 15.8 km n Laurens, SC hwy 97 at Beaver Dam Cr.,
9, 9 May 1977, R.M. Shelley (NCSM A1567). Orangeburg Co.—
Orangeburg, 4tf, 2 juvs., 20 July 1961, D. Dowling (FSCA), and 4(J, 9, 3
August 1961, D. Dowling (FSCA). Barnwell Co.— Barnwell State Park, 5
tf, 7 August 1976, R.M. Shelley (NCSM A1368). Jasper Co.— 12.9 km s
Hardeeville, along US hwy 17A, 0.6 km w SC hwy. 170A, 9, 2 July 1977,
R.M. Shelley (NCSM A1601); and Ridgeland, 9, 6 April 1975, D. Brody
(AMNH).

Only three males of P. crassicutis incursus were examined by Hoffman for
his revision, and all had been preserved since early this century. We
therefore submit the following observations on color, body proportions,
and gonopod variation to supplement his description.

In life most individuals are dark, chocolate brown dorsally with light
yellow paranota (Fig. 1); the venter is a little lighter in color than the
paranota. Specimens from Barnwell and Greenwood counties, South
Carolina, are light, brownish-gray dorsally with cream-colored paranota.

The body dimensions of males and females collected in the summer of
1976 are shown in Table I. Males are generally longer and wider than
females and have a higher W/L ratio. Individuals of both sexes are larger
in the Kings Mountain region and become smaller in a south-south-
eastward direction. This correlates with the paling of body color and
probably reflects more favorable environmental conditions in the Kings
Mountain region.

New Milliped Records

151

Figs. 2-3. Left male gonopod of Pachydesmus crassicutus incursus from 9.3 km s
Kings Mountain (town), Cleveland County, North Carolina. Drawn x50. 2,
medial view. 3, lateral view.

Figs. 4-9. Apical variation of secondary tibiotarsus of Pachydesmus crassicutis in-
cursus. Specimens from Woods Ferry Recreation Center, Chester County, South
Carolina. Medial views, drawn xlOO.

152

Rowland M. Shelley and Marianne Filka

Fig. 10. Distribution of Pachydesmus crassicutis in southeastern United States. •
P. c. incursus; A = P. c. denticulatus; and ■ = P. c. retrorsus.

Table 1. Body dimensions of Pachydesmus crassicutis incursus
(all measurements are averages and in mm.)

<s

County

No. inds.

W L W/L%

No. inds.

W L

W/L

Cleveland, NC

3

12.0 62.8 19.2

5

11.2 69.0

17.8

Gaston, NC

2

13.2 66.9 19.7

3

11.9 65.5

18.1

York, SC

1

12.6 61.7 20.4

2

11.2 57.7

19.4

Chester, SC

6

12.3 63.0 19.5

Union, SC

1

11.8 63.6

18.6

Newberry, SC

4

10.9 57.2 19.1

2

11.4 61.7

18.5

Greenwood, SC

1

10.8 56.5 19.1

1

9.9 54.7

18.1

McCormick, SC

3

11.5 60.9 18.8

1

11.2 54.8

20.4

Barnwell, SC

4

10.2 52.6 19.3

New Milliped Records 1 53

On the gonopods (Figs. 2, 3), the subapical process of the secondary
tibiotarsus is reduced from the configuration depicted by Hoffman (Figs.
7c and 8c, pp. 201 and 203), so that it appears as a small tooth on the
main stem of the structure. It may be blunt or pointed (Figs. 4.-9) and is
absent from one male from Chester County (Fig. 9). Similarly, the apical
portion of the secondary tibiotarsus varies and may be blunt or pointed
and curved distad or ventrad (Figs. 4-9). The primary and secondary
tibiotarsi are subequal in length in specimens from North Carolina and
York, Chester, and Newberry counties, South Carolina, but the latter is
slightly longer in specimens from the southern part of the range. Den-
ticulations along the anterior edge of the primary tibiotarsus vary and are
essentially absent from some gonopods. The coxal apophysis and the
remainder of the gonopod are as described by Hoffman (1958).

This increased knowledge of variation of P. c. incursus does not affect the
status of P. c. denticulatus Chamberlin, which occurs in the Piedmont
region of north-central Georgia. The two subspecies are distinguished by
several features, most notably the configuration of the coxal apophysis,
which is tri-lobed in P. c. incursus and smoothly rounded with the distal
margin entire in P. c. denticulatus.

The range of P. crassicutis incursus can thus be expanded to include the
Kings Mountain region of North Carolina and the Piedmont and Coastal
Plain of southern South Carolina (Fig. 10). Efforts to find the diplopod in
North Carolina adjacent to the Kings Mountain region have been unsuc-
cessful, and this area therefore represents the known northeastern range
limit of the genus.

ACKNOWLEDGMENTS.— We thank John C. Clamp and William W.
Thomson for assistance in collecting Pachydesmus from North Carolina;
Howard V. Weems, Jr., for access to material in the FSCA; and Norman
I. Platnick, for access to that in the AMNH. Specimens from Kings
Mountain, Barnwell, Baker Creek, and Hickory Knob State Parks, South
Carolina, were collected with permission of the S. C. Department of
Parks, Recreation, and Tourism, Division of State Parks. The photograph
is courtesy of Curtis Wooten, N. C. Wildlife Resources Commission.

LITERATURE CITED

Hoffman, Richard L. 1958. Revision of the milliped genus Pachydesmus
(Polydesmida: Xystodesmidae). Proc. U. S. Natl. Mus. 108: 181-218.

Wray, David L. 1967. Insects of North Carolina, Third Supplement. N. C. Depart.
Agric. Div. Entomol., Raleigh 181 pp.

Accepted 8 November 1978

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CONTENTS

The Brothers Brimley: North Carolina Naturalists. John E. Cooper . . 1

Nest and Larvae of the Neuse River Waterdog, Necturus lewisi
(Brimley) (Amphibia: Proteidae). Ray E. Ashton, Jr. and
Alvin L. Braswell 15

A New Species of Xironodrilus Ellis 1918 from North Carolina

(Clitellata: Branchiobdellida). Perry C. Holt and Ann M. Weigl 23

A New Cryptic Species of Salamander of the Genus Plethodon from
the Southeastern United States (Amphibia: Plethodontidae).
Richard Highton 31

The Female Reproductive Cycle in North Florida Kinosternon bauni
(Testudines: Kinosternidae). John B. her son 37

Fishes of the Gauley River, West Virginia. Charles H. Hocutt,
Robert F. Denoncourt and Jay R. Stauffer, Jr 47

Historical Review of the Carolina Parakeet in the Carolinas.
Daniel McKinley 81

Estimates of Fish Populations in Two Northeastern

North Carolina Swamp Streams. William H. Tarplee, Jr 99

Some Snake Food Records from the Carolinas. E. E. Brown 113

The Status of the Outer Banks Kingsnake, Lampropeltis getulus
sticticeps (Reptilia: Serpentes: Colubridae). Richard M. Blaney 125

Notes on the Natural History of the Terrestrial Leech, Haemopis
septagon Sawyer and Shelley (Gnathobdella: Hirudinidae).
Rowland M. Shelley, Alvin L. Braswell and David L. Stephan 129

New Records of the Salamanders Ambystoma talpoideum (Holbrook)
and Hemidactylium scutatum (Schlegel) in North Carolina
(Amphibia: Ambystomatidae and Plethodontidae).
Alvin L. Braswell and Nora M. Murdoch 135

An Albino Sculpin from a Cave in the New River Drainage of
West Virginia (Pisces: Cottidae).
James D. Williams and William M. Howell 141

Occurrence of the Milliped Pachydesmus crassicutis incur sus

(Chamberlin) in the Kings Mountain Region of North Carolina
and the Coastal Plain of South Carolina (Polydesmida:
Xystodesmidae). Rowland M. Shelley and Marianne Filka 147