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North Carolina State Library
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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 £00logy, 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
Speaes 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. websieri 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
> >) Etowah 1 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. Iverson 1
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.
5
a
15 73.3 53.3 73.3
7 57.1 28.6 42.
9
a
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-
zo
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
Xa3
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 Cowen 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-clu