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A New Nonparasitic Species of Lamprey,
Genus Lethenteron (Petromyzonidae),
from Eastern Tributaries of the Gulf of Mexico, U.S.A.
National Museum of Natural Sciences
Publications in Zoology, No. 12
Published by the
National Museums of Canada
Staff editor
Frances Smith
Crown copyrights reserved
National Museum of Natural Sciences
National Museums of Canada
Ottawa, Canada
Fourth quarter 1975
Information Canada
Catalogue No. NM95-10/12
P0987654321
Y798765
Printed in Canada
ISSN 0068-8037
Musée national des Sciences naturelles
Publications de Zoologie, n° 12
Publié par les
Musées nationaux du Canada
Droits réservés au nom de la Couronne
Musée national des Sciences naturelles
Musées nationaux du Canada
Ottawa, Canada
Quatrième trimestre 1975
Information Canada
N° de catalogue NM95-10/12
T0987654321
A798765
Imprimé au Canada
A New Nonparasitic Species of Lamprey,
Genus Lethenteron (Petromyzonidae),
from Eastern Tributaries of the Gulf of Mexico, U.S.A.
V. D. Viadykov,
Edward Kott,
and S. Pharand-Coad
Acknowledgements
The present study was made possible through the generous help
of many persons and institutions.
The study specimens were kindly loaned by the following
persons: Dr. Branley A. Branson and Mr. Bruce H. Bauer, Depart-
ment of Biological Sciences, Eastern Kentucky University; Drs.
Warren V. Brigham and Larry M. Page, Section of Faunistic Surveys
and Insect Identification, Illinois Natural History Survey; Drs.
Hubert Boschung and Thomas S. Jandebeur, Department of
Biology, University of Alabama; Dr. Frank J. Bulow, Department
of Biology, Tennessee Technological University; Dr. George L.
Harp, Arkansas State University; Dr. John S. Ramsey, Department
of Fisheries and Allied Aquacultures, Auburn University; and Dr.
Martin L. Wiley, Natural Resources Institute, University of Maryland.
Many measurements of specimens were made by Mrs. Jacqueline
Lanteigne-Courchéne, a student, and photographs were prepared
by Mr. G. Ben-Tchavtchavadze, both of the Department of Biology,
University of Ottawa.
Dr. Carl L. Hubbs, Scripps’ Institution of Oceanography, Dr. Don
E. McAllister and Mr. C.G. Gruchy, National Museum of Natural
Sciences, Ottawa, carefully read the manuscript and offered many
valuable suggestions. This study was supported by Grant No.
A-1736 from the National Research Council of Canada.
To all these persons and institutions the authors extend their
most sincere thanks.
Summary
À new nonparasitic lamprey, Lethenteron meridionale, from streams
of the Tennessee, the Alabama, and the Tombigbee river systems,
is described and illustrated. The holotype (NMC 74-249) is depos-
ited in the National Museum of Natural Sciences, Ottawa, Canada.
The species is distinguishable from Lethenteron lamottenii, its
nearest relative, by fewer myomeres (50-58), dentition, and pig-
mentation. By its row of posterial teeth L. meridionale is easily
separable from Lampetra aepyptera, which occurs in the same
watersheds. The description is based on an examination of 79
transformed individuals and two ammocoetes.
Résumé
Cet article décrit le Lethenteron meridionale, une espèce nouvelle
de lamproie non parasite des ruisseaux tributaires des rivières
Tennessee, Alabama et Tombigbee. L’holotype (NMC 74-249) se
trouve au Musée national des Sciences naturelles, a Ottawa,
Canada. Cette nouvelle espèce diffère du Lethenteron lamottenii,
par sa dentition, sa pigmentation et son nombre inférieur de
myomères (de 50 à 58). Sa rangée de dents postérieures la dis-
tingue aussi de la Lampetra aepyptera qu’on rencontre dans les
mêmes réseaux hydrographiques. La description se fonde sur une
étude de 79 individus métamorphosés et de deux ammocètes.
Biographical Notes
V. D. Viadykov
Vadim D. Vladykov was born in Russia and studied at Charles
University in Prague, where he specialized in zoology and anthro-
pology, receiving his Ph.D. in 1925. In 1930 Dr. Viadykov came to
Canada, and from 1930 to 1936 was employed by the Biological
Board of Canada (now the Fisheries Research Board of Canada)
as a fisheries biologist. In 1936 he became a Canadian citizen.
For the next two years he worked as a fisheries biologist for the
State of Maryland and as professor of ichthyology at the University
of Maryland. In 1938 he returned to Canada and joined the Biology
Department of the University of Montreal as professor of ichthyol-
ogy, a post he held until 1942. From 1943 to 1958 he was director
of the Biological Laboratory of the Quebec Department of Fisheries.
In 1958 he joined the staff of the University of Ottawa as professor
of biology, a position he held until his retirement in 1973, at which
time he was appointed professor emeritus. In the same year he
accepted the honorary position of research associate at the
National Museum of Natural Sciences. Dr. Vladykov was elected
a fellow of the Royal Society of Canada in 1963.
Dr. Vladykov is credited with more than 185 publications, and
his specialties are the biology of the American eel and the taxon-
omy of salmonids, sturgeons, and, especially, Holarctic lampreys.
He has described several new species of lampreys from Europe
and North America. Among Dr. Viadykov’s contributions to science,
the studies of particular significance are those dealing with the
description of characters by which larval lampreys (ammocoetes)
of nonparasitic species and those of the parasitic sea lamprey
(Petromyzon marinus) can be distinguished. This work now enables
biologists to separate larvae of nonparasitic lampreys from those
of parasitic lampreys, a task formerly impossible. For Canada and
the United States this is a great financial and ecological saving,
because treatment of streams and brooks is now restricted to
those which contain larvae of the destructive sea lampreys. In
recognition of his study of lampreys, a new species from central
Europe was described under the name Eudontomyzon vladykovi
Oliva & Zanandrea, 1959.
Edward Kott
Edward Kott obtained his academic training from the University of
Toronto. He received his B.A. in 1960 and his Ph.D. in 1964. From
1963 to 1965 he was a lecturer in biology at Lakehead University,
Thunder Bay, Ontario. In 1965 Dr. Kott joined the Fisheries Re-
search Board of Canada, and from 1965 to 1967 worked at the
Marine Ecology Laboratory of the Bedford Institute of Oceanogra-
phy, Dartmouth, Nova Scotia. He is now associate professor of
biology at Wilfrid Laurier University, Waterloo, Ontario. His research
has been in the ecology of small mammals, community studies,
and the ecology and systematics of lampreys. At present Dr. Kott
is working in cooperation with Dr. V.D. Vladykov on the system-
atics of North American lampreys.
Biographical Notes
S. Pharand-Coad
Sylvie Pharand-Coad was born in Hull, Quebec, in 1950. She gra-
duated from the University of Ottawa with an Honours B.Sc. in
Zoology in 1972 and a B.Ed. in 1973. For four years (1971-74),
Mrs. Pharand-Coad worked as a research assistant to Dr. V. D.
Vladykov on the systematics of Holarctic lampreys. Her research
interests are centred on the systematics and ecology of lampreys,
amphibians, and reptiles. At present she is a high-school science
teacher with the Ottawa Board of Education.
Introduction
The senior author (Vladykov 1973) recently
described a nonparasitic lamprey, Lampetra
pacifica, from the western United States. To
compare critically this western species with
the nonparasitic Lampetra aepyptera, from
eastern and southeastern United States, he
wrote to W.V. Brigham, who had just (1973)
described spawning of Lampetra aepyptera
in Tennessee. Thanks to Dr. Brigham’s co-
operation, we were able to obtain specimens
of L. aepyptera from several institutions in
the following states: Alabama, Arkansas,
Georgia, Kentucky, Maryland, Ohio, and
Tennessee.
Upon examining the specimens received,
we found two species of nonparasitic lam-
preys, one of which is Lampetra aepyptera
(Abbott 1860), the other, which we here treat
as a new species, referred to the genus
Lethenteron Creaser and Hubbs 1922, of the
family Petromyzonidae.*
The aim of the present article is to des-
cribe this new species and compare it with
two other nonparasitic species, Lampetra
aepyptera and Lethenteron lamottenii** (Le
Sueur 1827).
*Alternate spellings of the family are found in the
literature (Viadykov 1974).
**The name /amottenii is now, commonly without
valid reason, spelled as /amottei (American Fish-
eries Society, Committee on Names 1970).
Methods and Materials
All measurements and counts were typically
made on the left side of the specimen, as
established by Vladykov and Follett (1965).
Measurements
All measurements are expressed as percent-
ages of the total length of the specimen;
the disc length (d) is also expressed as a
percentage of the branchial length (B,—B,).
The following abbreviations are used, as in
Vladykov and Follett (1965).
a-C Tail length, the distance from the
posterior edge of the cloacal slit to
the end of the caudal fin.
Trunk length, the distance from the
posterior edge of the last (seventh)
branchial opening to the anterior
edge of the cloacal slit.
Branchial length, the distance from
the anterior edge of the first bran-
chial opening to the posterior edge
of the last (seventh) branchial open-
ing.
d Disc length, longitudinal diameter,
with the oral fimbriae included, mea-
sured with the disc closed.
Prebranchial length, the distance
from the anterior edge of the disc
(in transformed individuals) or of the
upper lip (in ammocoetes) to the
anterior edge of the first branchial
opening.
Second dorsal fin maximum height,
measured along the highest fin ray.
0 Eye length, the horizontal diameter
of the eye.
TL Total length, the distance from the
anteriormost oral fimbria to the end
of the caudal fin.
B.-a
Bi-B;
hD,
In addition, the number of oral papillae,
the number of anterior oral fimbriae con-
tained within a section equal to the width of
the base of the supraoral lamina, and the
number of velar tentacles were counted.
In identifying ammocoetes, the number of
trunk myomeres and body proportions were
used. Pattern and intensity of pigmentation
on head and tail were also noted (Vladykov
1950).
Methods and Materials
Material Examined
The material of the new species of Lethen-
teron consists of 79 transformed individuals
from southern United States: 65 from Ten-
nessee, 11 from Alabama, and 3 from Geor-
gia; and 2 ammocoetes: one each from
Tennessee and Alabama.
The comparative material of Lampetra
aepyptera consists of 53 transformed indi-
viduals (28 from Alabama, 3 from Arkansas,
5 from Kentucky, 6 from Maryland, 3 from
Ohio, and 8 from Tennessee) and 13 ammo-
coetes (10 from Alabama and 3 from Ken-
tucky). Data for each collection are given in
the Appendix.
10
Description of New Species
Lethenteron meridionale sp. nov.
Gulf Brook Lamprey
Figures 1-6
Etymology
The species name meridionale is derived
from the Latin meridies for midday, or south,
and refers to the southern distribution. It is
used in the neuter form to agree with
Lethenteron.
Holotype
An adult male 104 mm in total length from
Blue Springs Creek, a tributary of the Ten-
nessee River, near Hillsboro, Coffee County;
3 February 1968; J.S. Jandebeur and J.D.
Williams; UAIC 2830, now re-catalogued as
NMC 74-249. It bears the plastic tag W667.
This new nonparasitic species belongs to
the genus Lethenteron Creaser and Hubbs
1922. This genus is well characterized by
the arrangement of its teeth (Vladykov and
Follett 1967).
Diagnosis
The new species clearly differs from its
nearest relative, Lethenteron lamottenii (Le
Sueur), also nonparasitic, by its low number
of trunk myomeres in transformed speci-
mens, varying between 50 and 58 (average
54), whereas in L. lamottenii the myomeres
range from 64 to 74 (average 68). Other
differences are summarized in Table 12.
From another nonparasitic species, Lam-
petra aepyptera (Abbott) living in the same
watersheds, Lethenteron meridionale is
readily distinguishable by possession of
posterials,* varying in number from 10 to 21
(average 15.4). Posterials are completely
lacking in aepyptera, as in all other species
of Lampetra. Other differences between
meridionale and aepyptera are given in
Tables 7 and 11.
Characters of Holotype
The holotype, a male 104 mm in total length
when received.
Measurements of the holotype — referable
to maturity stage 3 (Vladykov and Follett
1965) — each expressed as a percentage of
* It is regrettable that Scott and Crossman (1973)
omitted posterials on their schematic drawings of
the disc of L. lamottenii, in spite of the fact that
these teeth are always present in all species of
Lethenteron.
total length, are: prebranchial length, 10.6;
trunk length, 46.6; tail length, 30.8; maximum
height of the second dorsal fin, 5.3; disc
length, 6.3; eye length, 1.9. Disc length as a
percentage of branchial length is 59.1. Trunk
myomeres (Figures 1 and 2), 52.
The already cornified teeth are small. On
each side of the disc there are three inner
lateral teeth, all unicuspid. On the anterior
field there are about 34 teeth, all practically
the same size; on the posterior field there
is a single row of 18 small posterials. The
supraoral lamina is provided with one cusp
at each end; the infraoral lamina has about
12 weakly developed cusps; the marginals
are in two rows. The three lingual laminae
are so poorly developed that the cusps are
uncountable (Figure 3).
Colour of the specimen preserved in 4-5%
formalin is dark greyish brown on the flanks
and back, and whitish along the ventral
surface.
Although the holotype is in the early stage
of maturity, the dorsal fins are already high
and close together, and the genital papilla
is well-developed. There is an extensive,
dark pigmentation of vertically elongated
spots on the dorsal fins. A dark blotch sur-
rounds the posterior section of the noto-
chord; towards the edges of the caudal fin,
the pigment becomes less pronounced
(Figure 1).
Characters of Paratypes
Collection data for paratypes are given in
the Appendix.
Transformed Specimens
Total length (Tables 1 and 2)
The 55 males examined range in total length
from 96 to 136 mm (average 115.8 mm); the
24 females range from 96 to 141 mm
(average 116.7 mm).
Body proportions (Tables 1 and 2)
For the 55 males the average measurements
as percentages of total length are: prebran-
chial length, 11.7 (range 10.2-12.9); branchial
length, 10.2 (range 9.1-11.4); trunk length,
48.0 (range 43.6-52.7); tail length 30.2 (27.3—
33.9); maximum height of second dorsal fin,
4.5 (3.0-5.7); eye length, 1.8 (range 1.5-2.1);
disc length, 5.2 (4.2-6.5). Disc length as a
percentage of branchial length is 51.5 (range
41.7-66.7).
11
Description of New Species
For the 24 females the average measure-
ments are: prebranchial length, 10.7 (range
7.8-13.0); branchial length, 10.0 (range 9.3-
11.4); trunk length, 51.4 (49.2-53.9); tail
length 28.0 (26.0-30.3); maximum height of
second dorsal fin, 3.8 (range 2.5-5.1); eye
length, 1.7 (range 1.4-2.1); disc length, 4.8
(range 2.8-5.7). Disc length as a percentage
of branchial length is 47.5 (range 27.6-57.1).
Trunk myomeres (Table 5)
The number of trunk myomeres in the trans-
formed specimens of meridionale varies
from 50 to 58 (average 53.8).
Dentition (Tables 7-9)
The teeth are small and often not fully corn-
ified; hence some counts of them are un-
certain. The teeth on the anterior field are
uniformly small, averaging 16.1 (range 5-34)
in number. The single row of posterials is
typical for all species of the genus Lethen-
teron.
The posterials range in number from 10 to
21 (average 15.4), but are often weakly
developed and difficult to count. The supra-
oral lamina has two blunt cusps, one at each
end; as an exception, one specimen, Tag
W533, from Alabama, has a strongly devel-
oped median cusp on the supraoral lamina.
The rounded and weakly developed cusps
on the infraoral lamina average 9.9, ranging
from 6 to 13 in number.
The typical number of inner laterals in the
genus Lethenteron is six, three on each side
of the disc. Infrequently, fewer than six
(rarely aS many as eight) cornified inner
laterals develop in meridionale. In three spec-
imens (Table 8) all inner laterals are lacking.
The average number of inner laterals is 5.5
(range 0-8). Although in the genus Lethen-
teron all inner laterals are typically bicuspid,
in L. meridionale they are usually unicuspid.
The marginals tend to form two rows
around the disc.
The transverse and longitudinal lingual
laminae are weakly developed and the cusps
are uncountable.
Anenlarged median cusp on the transverse
lamina is typical of other species of Leth-
enteron (Figure 4), but does not develop in
L. meridionale.
Other characters
Velar tentacles were counted in two speci-
mens of meridionale where only three ten-
1124
tacles were found: one median with a single
lateral on either side.
The average number of oral papillae
around the disc in 55 specimens of L. meri-
dionale is 14.2 (range 8-25). The number of
anterior fimbriae within the width of the
base of the supraoral lamina is somewhat
variable. In 10 specimens from Tennessee
the average is 17.2 (range 13-22) and in 5
from Alabama the average is 11.0 (range
9-15). The difference between the Tennessee
and Alabama specimens appears to be
related to the width of the supraoral lamina,
which averages 2.6 mm in Tennessee speci-
mens and 1.9 mm in those from Alabama.
There are dark chromatophores in the
buccal cavity but none extend onto the
tongue.
Colouration (Table 10)
The sides and back of specimens fixed in
4-5% formalin are greyish brown and the
lower surface is whitish.
Dark pigmentation on the second dorsal
fin, if developed, consists of a series of
vertically elongated narrow dark spots. There
is a distinct black blotch (Figures 1 and 2) on
the caudal fin. The intensity of dark pigmen-
tation varies with the geographical region; in
specimens from the Tennessee-Mississippi
drainage pigmentation is very pronounced,
in contrast with those from the Tombigbee
and Alabama rivers, where it is weakly deve-
loped.
Sexual dimorphism
The external appearance of males and fe-
males of Lethenteron meridionale is similar
to that of L. lamottenii as described by
Vladykov (1949).
There are sexual differences in body pro-
portions (Tables 1, 2, and 4). Disc length (d),
prebranchial length (d-B,) and tail length
(a-C) are greater in males; females have a
relatively longer trunk (B,-a).
All specimens were in prespawning condi-
tion. One female (Tag W556, from Alabama),
maturity stage 3, 122 mm in length, collected
on 5 December 1963, had 2,154 eggs aver-
aging 0.63 mm in diameter.
Ammocoetes
Only two ammocoetes were available, one
from Tennessee (Tag W677), 100 mm long,
and the other from Alabama (Tag W552),
142 mm. Body proportions expressed as
percentages of total length were as follows:
d-B,, 7.0-8.5; B,-B,, 11.2-12.5; B,-a, 50.5-
53.5; a-C, 28.5. The myomeres number 54
and 55.
As in the transformed specimens, the
ammocoete from Tennessee is more heavily
pigmented than the one from Alabama. The
limited number of specimens does not permit
a critical description of the ammocoetes of
Lethenteron meridionale.
Comparison of Lethenteron meridionale with
L. lamottenii
In the Tennessee River tributaries from which
the holotype and topotypes of L. meridionale
were obtained, one male and one female of
L. lamottenii (Tags W679 and W680) were
collected, in Putnam County, Tennessee.
These Tennessee specimens of the two
nonparasitic species are compared in Table
12. The principal differences are found in
the number and degree of development of
different types of teeth (Figures 2 and 4). The
number of myomeres in meridionale is 50-
58, whereas each of the two specimens of
lamottenii from Tennessee has 68. Smith-
Vaniz (1968) indicated 63-70 for /amottenii
from the Tennessee River system in Alabama,
and Trautman (1957) reported 63-73 from
Ohio. Several authors, Valdykov (1949),
Manion and Purvis (1970), and Kott (1974),
observed a similar number in material from
the St. Lawrence River and Great Lakes
areas.
Another striking difference between me-
ridionale and lamottenii is in dark pigmen-
tation of the second dorsal fin. In meridionale
there are often several vertical narrow spots,
whereas in lamottenii individual spots are
absent, but a dark-brown tint made up of
minute chromatophores is noticeable over
the fin membrane (Figures 1 and 5).
Lethenteron meridionale is asmall species,
attaining less than 150 mm in total length,
whereas some individuals of L. /amottenii
have been reported to reach a “gigantic”
size, up to 300 mm (Manion and Purvis 1970),
but ordinarily reach 217 mm (Kott 1974).
13
Comparison of Lethenteron meridionale with
Lampetra aepyptera
In exterior appearance Lethenteron meri-
dionale and Lampetra aepyptera are similar
(Figure 2). For this reason all specimens
that we now identify as meridionale were
originally determined as Lampetra aepyptera
in collections of the University of Alabama.
Similarity between meridionale and aepyp-
tera was observed in number of trunk myo-
meres (Tables 5 and 6), in dark pigmentation
of fins, and in size. However, a close exam-
ination of body proportions (Tables 1-4) and
dentition (Tables 7-9) revealed differences.
See Table 10 for degree of pigmentation.
Lethenteron meridionale and Lampetra
aepyptera are compared in detail in Table
11. The principal differences between them
are in numbers and degree of development
of teeth on the anterior and posterior fields.
The number of anterials average 16 (range
7-34) in meridionale and 8 (range 2-21) in
aepyptera. The posterials are always present
in all species of Lethenteron. In meridionale
the number of posterials averages 15.9 (range
10-21). In all species of Lampetra, including
aepyptera, the posterials are absent (Figures
3 and 6). Even in histological sections,
Seversmith (1953) found no evidence of
posterials in aepyptera.
The number of inner laterals averages 5.5
(range 0-8) in meridionale and 4.6 (range
1-6) in aepyptera. In our material three
specimens of meridionale are devoid of inner
laterals. A similar loss in aepyptera was
reported by Raney (1952).
Two counts suggest that the fecundity of
meridionale may be significantly higher than
that of aepyptera: a 122-mm female of
meridionale (Tag W556, from Alabama) con-
tained 2,154 eggs, whereas a 118-mm speci-
men of aepyptera (Tag W537, also from
Alabama) collected on 2 February 1963,
contained 1,547.
Lethenteron meridionale and Lampetra
aepyptera are placed in separate genera on
the basis of the presence or absence of
posterial teeth. In spite of their allocation to
different genera, the two species are similar
in myomere number, size, pigmentation, and
reduction in number of teeth. One can
speculate that these similarities may be due
to parallelism, in which case assignment to
different genera is warranted. Or, aepyptera
may have lost its posterials and may share
a recent common ancestor with meridionale.
Lamprey collections from other Gulf states
would hopefully cast light on this problem.
14
Geographical Distribution of Lethenteron
meridionale
This species has been collected so far only
in freshwater streams in the eastern drainage
of the Gulf of Mexico, that is, in Tennessee,
Alabama, and Georgia. In Tennessee it has
been collected in several tributaries of the
Tennessee River system. It is significant to
note that in streams of Putnam County, Ten-
nessee, lamottenii and aepyptera are present
in addition to meridionale. Several samples
of meridionale were obtained from the Mobile
Basin to which the Alabama and Tombigbee
rivers belong. From the Alabama River sys-
tem meridionale has been collected in Geor-
gia and Alabama. Specimens have also
been obtained from the Tombigbee River in
Alabama.
The distribution of meridionale corres-
ponds to that of 16 other freshwater species
of Alabama, which Smith-Vaniz (1968) con-
cluded probably originated in the Tennessee
River system, whence they invaded the
headwaters of the Alabama River system
and Tombigbee River.
Figure 1
Enlarged photographs of the head and tail regions
of Lethenteron meridionale, holotype, Tag W667,
&, total length 104 mm, from Tennessee.
15
Figure 2
Enlarged photographs of two nonparasitic lam-
preys. Upper: Lethenteron meridionale, Tag W667,
holotype, &, total length 104 mm, from Tennessee.
Lower: Lampetra aepyptera, Tag W548, & , total
length 99 mm, from Alabama.
16
Figure 3
Enlarged photograph of the disc of Lethenteron
meridionale, holotype, Tag W667, o”, total length
104 mm, from Tennessee.
17
Figure 4
Enlarged photograph of the disc of Lethenteron
lamottenii, Tag W681, o, total length 162 mm,
from Gatineau River, Quebec, collected on 17
May 1958, by M. Corbeil and A. Lutz.
18
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uO p9}99/[09 ‘2eqenD “JaAly neesunedy wos ‘Www
291 uiBue] 1eJ0} ‘ © ‘LgoM Gey ‘ueyowe, uole}
-U9y197 JO UolBei jez ay} jo yde1Boyoud pebiejuy
G oinbi4
19
Figure 6
Enlarged photograph of the disc of Lampetra
aepyptera, Tag W572, o’, total length 93 mm,
from Alabama.
Note: The pinning of the disc for photography
caused a certain distortion on the left side of the
photograph. Hence, three small supplementary
marginal teeth, which could be mistaken for
posterial teeth, are brought closer to the infraoral
lamina.
20
Table 1
Body proportions (as percentages of total length) of 55 males of Lethenteron meridionale
from Tennessee, Alabama, and Georgia
TL d-B: B;-B; B--a a-C hD, 0 d d
Tag No. (mm) TL TL TL TL TL HE TL B;-B;
Tennessee
W673 96 12.5 9.9 45.8 31.8 5.2 2.1 5.2 52.6
W634 97 12.4 10.3 45.9 31.4 5.2 2.1 6.2 60.0
W618 98 12.8 9.7 48.5 29.1 4.6 2.0 5.1 52.6
W670 98 12.8 9.7 45.4 32.1 5.1 2.0 5.6 57.9
W633 101 11.4 9.9 48.5 29.7 5.0 2.0 5.4 55.0
W667* 104 12.1 10.6 46.6 30.8 5.3 1.9 6.3 59.1
W615 105 12.9 11.4 44.3 31.0 4.8 1.9 6.2 54.2
W631 105 12.4 10.5 46.7 30.5 4.8 1.9 6.3 59.1
W645 105 11.9 10.0 48.6 29.5 5.2 1.9 5.7 57.1
W635 105 12.4 11.0 46.2 30.5 4.8 1.9 5.7 57.1
W632 108 12.5 9.7 46.3 31.5 5.6 1.9 6.5 66.7
W643 109 12.4 10.6 47.7 29.4 5.0 1.8 5.5 52.2
W668 109 11.0 10.6 49.1 29.4 4.6 1.8 5.5 52.2
W629 110 12.3 10.5 46.4 30.9 5.5 1.8 5.0 47.8
W639 110 12.7 10.0 47.3 30.0 5.5 1.8 5.5 54.5
W637 111 10.8 10.8 47.7 30.6 5.4 1.8 5.4 50.5
W644 111 12.2 10.8 45.0 32.0 5.4 1.8 5.4 50.5
W613 112 12.1 10.7 45.5 31.7 5.4 1.8 5.4 50.0
W660 112 12.9 10.7 45.5 30.8 4.9 1.8 5.8 54.2
W671 112 121 9.8 47.3 30.8 4.9 1.8 5.8 59.1
W648 113 12.4 9.7 46.9 31.0 5.3 1.8 5.3 54.5
W661 113 11.9 11.1 46.0 31.0 4.4 1.8 6.2 56.0
W666 113 11.5 9.7 49.6 30.1 49 1.8 5.8 59.1
W657 114 11.8 10.5 47.4 30.3 4.8 1.8 5.7 54.2
W674 114 11.8 9.6 47.8 30.7 4.8 1.8 5.7 59.1
W646 115 11.7 9.1 47.8 31.3 5.2 1:7 5.7 61.9
W659 115 12.2 9.6 48.3 30.0 4.8 17 6.1 63.6
W658 115 11.3 10.0 48.3 30.4 5.2 17 5.2 52.2
W612 116 11.2 9.9 47.0 31.5 3.4 07 4.7 47.8
W619 116 12.1 9.9 45.7 32.3 5.6 17 5.2 52.2
W630 116 12.5 10.3 48.7 28.4 5.2 17 5.6 54.2
W638 117 12.0 10.3 43.6 34.2 5.1 1.7 5.6 54.2
W649 117 12.0 10.3 45.3 32.5 4.7 17 4.7 45.8
W665 117 12.0 9.8 48.3 29.9 4.7 47 5.6 56.5
W642 119 11.8 10.1 46.2 31.9 5.0 17 5.5 54.2
W647 119 11.3 10.1 46.2 31.9 5.0 17 5.5 54.2
W672 120 12.1 10.0 46.7 31.3 5.0 1.7 5.4 54.2
W636 122 12.3 tia 45.9 30.3 4.9 1.6 5.3 48.1
W616 124 121 10.5 47.6 29.8 5.2 2.0 6.5 61.5
W640 124 11.7 10.9 45.6 31.9 4.4 1.6 5.2 48.1
Table 1
age d-B: B,-B; Ba a-C hD: 0 d d
Tag No. (mm) Wt ae TL ae TL WL TL B:-B;
W656 124 Aer 9.3 45.2 33.9 5.6 1.6 5.6 60.7
W641 126 11.9 10.3 47.6 30.1 4.4 1.6 5.6 53.8
W655 126 11.5 9.5 47.2 31.7 5.2 1.6 5.2 54.2
W676 132 10.2 9.8 52,7 27.3 3.0 1.5 4.5 46.2
W677 136 11.0 9.9 50.4 28.7 3.7 1.5 4.8 48.1
Mean 112.8 12.0 10.2 47.0 31.0 5.0 1.8 5.5 54.7
Alabama
W533 116 11.2 9.5 50.7 30.2 3.9 124 4.3 45.5
W581 116 12.1 10.3 50.0 27.6 - 1.8 5.2 50.0
W583 116 1122 9.5 50.0 29.3 - 17 6.0 63.6
W582 117 11.5 9.8 49.6 29.1 - 2.1 5.6 56.5
W555 118 11.4 10.2 50.0 28.4 3.0 ln 4.2 41.7
W554 122 11.1 10.2 50.8 27.9 3.7 1.6 4.9 48.0
W558 131 10.3 9.2 51.9 28.6 3.1 1.5 4.2 45.8
Mean 119.4 11.3 9.8 50.4 28.7 3.4 etl 4.9 50.2
Georgia
W596 103 12.1 lee, 45.1 31.2 4.9 1.9 5.0 47.8
W598 110 11.8 10.5 49.5 28.6 4.5 1.8 52 50.5
W597 123 11.8 10.2 45.5 32.5 SA 1.6 DA 56.0
Mean 112.0 11.9 10.6 46.7 30.8 5.0 1.8 5.2 50.5
Combined
Mean 114.7 ul 10.2 48.0 30.2 4.5 1.8 5.2 515
*Holotype
22
Table 2
Body proportions (as percentages of total length) of 24 females of Lethenteron meridionale
from Tennessee and Alabama
TL
Tag No. (mm)
W650 96
W620 99
W621 100
W662 105
W626 107
W627 108
W622 109
W623 109
W669 109
W625 dti
W652 111
W628 113
W651 114
W624 115
W653 116
W663 116
W664 116
W654 118
W617 119
W675 141
Mean 111.6
W553 119
W543 121
W556 122
W557 125
Mean 121.8
Combined
Mean 116.7
d-B, Bi-B Ba a-C hD; 0
TL TL ie TE TL TE.
Tennessee
13.0 10.4 50.5 26.0 4.7 2.1
12.1 10.6 50.5 26.8 5.1 2.0
12.0 10.0 51.0 27.0 5.0 2.0
11.4 11.4 50.0 27.1 4.3 1.9
10.7 10.3 52.3 26.6 4.2 1.9
10.6 10.2 52.8 26.4 4.6 1.9
11.0 9.6 50.9 28.4 4.6 1.8
ies) 10.6 50.0 28.0 4.6 176
11.5 10.6 51.8 26.1 4.7 1.8
10.8 9.5 52.3 27.5 4.5 1.8
wales) 9.9 BATA 27.6 4.7 127
11.1 9.3 51.8 27.9 4.4 1.8
10.5 10.1 51.8 28.1 4.4 1.8
10.9 9.6 51.3 28.3 4.3 17
10.8 9.9 51.7 27.6 4.7 let
11.2 112. 49.6 28.0 4.7 5 IEA
10.3 9.5 51.7 28.4 3.9 127
11.4 11.4 50.0 27.1 4.3 1.9
10.1 10.1 49.6 30.3 3.4 ei,
7.8 10.3 53.9 28.0 2.5 1.4
11.0 10.2 Sis 27.6 4.4 1.8
Alabama
10.5 9.7 51:3 28.6 2.9 1.7
9.1 9.9 537 PATES) 2.9 atl
10.7 10.2 49.2 29.9 3.3 1.6
10.8 9.6 52.0 27.6 3.2 1.6
10.3 9.8 51.5 28.3 Oat 1%
10.7 10.0 51.4 28.0 3.8 174
B;-B;
Table 3
Body proportions (as percentages of total length) of 23 males and 12 females of Lampetra
aepyptera from Tennessee and Alabama
Tag No.
5072 (VDV)
5073 (VDV)
5075 (VDV)
5071 (VDV)
5074 (VDV)
5076 (VDV)
Mean
W517
W548
W549
W524
W539
W540
W530
W550
W528
W526
W531
W551
W578
W579
W546
W522
W580
Mean
Combined
Mean
5070 (VDV)
5069 (VDV)
Mean
we
(mm)
121
134
135
142
142
145
136.5
91
99
99
101
107
109
110
111
112
112
115
118
119
122
123
124
128
110.1
117.0
140
151
145.5
95
97
99
102
103
d-B;
TL
11.6
11-9
11.9
11.6
12.0
10.3
11.4
10.9
11.2
Uist,
11.3
12.6
10.8
9.2
B,-B,
TL
10.1
10.2
B--a
wit
Males
Tennessee
50.4
50.4
48.5
48.9
50.0
49.7
49.7
Alabama
48.4
47.5
48.5
46.5
48.1
48.6
49.1
50.5
47.8
49.6
50.0
48.7
48.3
48.4
49.2
47.2
47.7
48.6
49.2
Females
Tennessee
53.2
54.3
a-C
ho:
ME
6.0
5.5
ND = a a oS
ON
œ o
_
[Ce]
B,-B;
Table 3
TL d-B, B.-B; B-a a-G hD; We d
Tag No. (mm) ie TL Tite TL TL TL TE
W523 106 11.3 9.9 51.4 27.4 4.7 1.9 4.7
W541 107 10.7 10.3 50.9 28.0 33 1.4 5.1
W542 111 10.4 10.4 50.9 28.4 3.6 1.8 4.5
W547 112 10.3 10.3 50.4 29.0 3.6 1.8 4.9
W537 118 9.3 9.3 50.8 30.5 3.0 1.7 3.8
Mean 105.4 10.7 10.0 50.8 28.6 4.0 1.8 4.9
Combined
Mean 112.1 11.0 10.2 52.3 28.4 4.9 1.8 5.1
25
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Table 8
Number of inner lateral teeth, combined for both sides of disc, in Lethenteron meridionale
and Lampetra aepyptera from Tennessee, Alabama, and Georgia;* sexes combined.
No. of Number of inner lateral teeth
Species specimens 0 1 2 3 4 5 6 7 8 Mean
L. meridionale 77 3 2 2 = 3 2 61 2 2 5.5
9.92% 2:6 2:6 ~ 3.9 2.6 79.2 2.6 2:6 1000
L. aepyptera 31 - 2 2 4 3 8 12 - - 4.6
- Gist = 76:5) NC PCR - 100.0
* From Georgia we have three specimens of L. meridionale and none of L. aepyptera.
**Figures in this line are percentages.
Table 9
Frequency of occurrence of cusps on inner lateral teeth in Lethenteron meridionale and
Lampetra aepyptera from Tennessee, Alabama, and Georgia;* sexes combined.
L. meridionale L. aepyptera
Inner lateral No. of Mean Mean
teeth cusps % no. of cusps % no. of cusps
Anterior 1 91.7 1.1 63.9 1.4
2 8.3 36.1
Middle 1 73.5 1.3 33.3 12
2 26.5 61.1
3 - 5.6
Posterior 1 84.8 dee 85.1 sl
2 1912 14.9
Supplementary 1 100.0 1.0 - -
posterior
*From Georgia we have three specimens of L. meridionale and none of L. aepyptera.
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Table 11
Detailed comparison of Lethenteron meridionale with Lampetra aepyptera, based on Tables
1-9. Numbers in italics refer to means, and ranges (in parentheses), for each character.
Character
L. meridionale
L. aepyptera
Similarities
Supraoral lamina
Marginals
All lingual laminae
Number of myomeres
Number of velar tentacles
Buccal cavity
Tongue
Dark pigmentation of second
dorsal and caudal fins
Body proportions, except length
of disc and height of second
dorsal fin
Total length (mm),
sexes combined
Differences
Posterial teeth
Anterial teeth (type)
Anterial teeth (number)
Total number of inner
lateral teeth
Middle lateral tooth
Infraoral lamina and number
of cusps
Number of oral papillae
Length of disc as percentage
of TL
Height of second dorsal fin
as percentage of TL
Number of eggs
and their diameter
Length of female
with two cusps,
one at each end
in a double row
typically with obsolete cusps
54.0
(50-58)
3
pigmented
nonpigmented
quite similar
similar
96-141
present
small, uniform,
but numerous
16.1
(7-34)
5.5
(0-8)
typically with one cusp
cusps degenerate
9.9
(6-13)
14.2
(8-25)
male: 5.2
(4.2-6.5)
female: 4.8
(2.8-5.7)
male: 4.5
(3.0-5.7)
female: 3.8
(2.5-5.1)
2,154 (0.6 mm)
122 mm
with two cusps,
one at each end
in a double row
typically with obsolete cusps
54.5
(50-59)
typically 3
pigmented
nonpigmented
quite similar
similar
91-151
absent
small, uniform, but few
8.1
(2-21)
4.6
(1-6)
typically with 2 cusps
cusps degenerate
15.8
(10-24)
male: 5.8
(4.3-6.7)
female: 5.1
(3.8-5.6)
male: 5.0
(3.3-7.0)
female: 4.9
(3.0-6.0)
1,543 (0.8 mm)
118 mm
Table 12
Comparison of two nonparasitic species of Lethenteron from Tennessee. Numbersinitalics
refer to means, and ranges (in parentheses), for each character.
Character
Anterial teeth
Number of posterial teeth
Total number of inner lateral
teeth
cusps on inner lateral teeth
Infraoral lamina and
number of cusps
Marginal teeth
All lingual laminae
Transverse lingual lamina
Number of myomeres
Number of velar tentacles
Buccal cavity
Dark pigmentation of second
dorsal fin
Total length (mm)
L. meridionale
L. lamottenii
uniform in size
15.9
(10-21)
515
(0-8)
all typically
with one cusp
cusps degenerate
10.1
(6-13)
in two rows
with cusps typically
obsolete
middle cusp
not enlarged
53.8
(50-58)
3
pigmented
several vertical
narrow spots
96-141
of two sizes: large near
supraoral lamina and smaller
towards marginals
21.5
(20-23)
6.0
(6)
all bicuspid
cusps prominent
9.5
(9-10)
in one row
with cusps well developed
middle cusp enlarged
68.0
(68)
7
nonpigmented
dark-brown tint without
distinct spots
153-162
Appendix
Complete data for the collections of Lethenteron
meridionale and Lampetra aepyptera used in this
study appear in the following sequence:
ES
Tag number. Small white plastic tags (15 mm
long by 5 mm wide) of two series were used.
Series a: On one side of the tag is the letter W,
followed by a number; on the other side
“Canada” is written.
Series b: On one side there is only a number,
and on the other the letters VDV, the initials
of the senior author, appear.
Number of specimens
Sex (in transformed specimens)
Total length
Localities in each river system (presented al-
phabetically)
Date of collection
Collector
Repository institution abbreviated as follows:
ASUMZ Arkansas State University Museum of
Zoology, State University, Arkansas
AU Auburn University, Auburn, Alabama
O1 B © D
OND
CBL Chesapeake Biological Laboratory,
Solomons, Maryland
EKU Eastern Kentucky University, Museum
of Fishes, Richmond, Kentucky
INHS Illinois Natural History Survey,
Urbana, Illinois
NMC National Museum of Natural Sciences,
National Museums of Canada, Ottawa
TTU Tennessee Technological University,
Cookeville, Tennessee
UAIC University of Alabama Ichthyological
Collection, University, Alabama
9 Catalogue number of repository institution
Lethenteron meridionale
Holotype and Topotypic Paratypes
W662-66 and 668 (392, 105-16 mm; 3%, 109-
17 mm); collected with holotype, W667 (now
NMC 74-249); Blue Springs Creek, a tributary of
the Tennessee River, near Hillsboro, Coffee
County; 3 February 1968; J.S. Jandebeur and J.D.
Williams; UAIC 2830.
Other Paratypes
Tennessee: Tennessee River system
W669-74 (one®, 109 mm; 5, 96-120 mm);
Bean’s Creek, 2 miles southeast of Hillsboro at
Stephenson, Coffee County; 4 February 1968;
J.S. Jandebeur and J.D. Williams; UAIC 2837.
W620-49 (92 , 99-115 mm; 219, 97-126 mm);
Bradley Creek at Hillsboro, Coffee County; 3
February 1968; J.S. Jandebeur and J.D. Williams;
UAIC 2831.
W650-61 (59, 96-118 mm; 7c”, 112-26 mm);
unnamed spring, tributary of Bradley Creek at
Hillsboro, Coffee County; 3 February 1968; J.S.
Jandebeur and J.D. Williams; UAIC 2832.
34
W612 (ones, 116 mm); Elk River at Elk Head,
Grundy County; 3 February 1968; J.S. Jandebeur
and J.D. Williams; UAIC 2835.
W617-19 (one®, 119 mm; 207, 98-116 mm);
Henly Creek, 2.1 miles southeast of Pelham,
Grundy County; 3 February 1968; J.S. Jandebeur
and J.D. Williams; UAIC 2834.
W613 (ones, 112 mm); Hurricane Creek, at
Awalt, Franklin County; 28 March 1968; J.S.
Jandebeur and J.D. Williams; UAIC 2866.
W675-77 (one®, 141 mm; 207, 132-36 mm),
W978 (one ammocoete, 142 mm); Spring Creek,
Putnam County; date and collector unknown; TTU.
W615-16 (25, 105-24 mm); unnamed creek,
2 miles southwest of Winchester Spring, Franklin
County; 28 March 1968; J.S. Jandebeur and J.D.
Williams; UAIC 2864.
Alabama: Tombigbee River system
W553-58 (32, 119-25 mm; 307, 118-31 mm),
W552 (one ammocoete, 100 mm); spring tributary
of Binnion Creek, Samantha, Tuscaloosa County;
5 December 1963; J.D. Williams and W.M. Howell;
UAIC 1091.
W543 (oneQ, 121 mm); Blue Creek about 1%
miles north of Vina, Frankline County; 24 January
1966; B. Wall and H.T. Boschung; UAIC 1837.
W533 (ones, 116 mm); Coldwater Spring, 5.7
miles west of Oxford, Calhoun County; 31 August
1966; J.D. Williams, J.G. Armstrong, and B.R.
Wall; UAIC 2349.
W581-83 (30, 116-17 mm); first creek south
of Tuscaloosa on Alabama highway 69, Tusca-
loosa County; 15 March 1963; A.B. Stapp and J.
Barnes; UAIC 991.
Georgia: Alabama River system
W598 (one, 110 mm); Blue Spring, 0.55 miles
north of Varnell, Whitfield County; 4 March 1966;
R.D. Caldwell and W.M. Howell; UAIC 1868.
W596 (ones, 113 mm); spring west of U.S.
highway 41, 6.8 miles south of Dalton, Whitfield
County; 4 March 1966; R.D. Caldwell and W.M.
Howell; UAIC 1873.
W597 (oneo’, 123 mm); spring pond run-off,
Y% mile north of Varnell, Whitfield County; 4
March 1966; R.D. Caldwell and W.M. Howell;
UAIC 1871.
Lampetra aepyptera
Alabama: Alabama, Tennessee, and
Tombigbee River systems
W526 (one, 112 mm); Beaver Creek, 4.4 miles
northwest of Aliceville, Pickens County; 12 March
1966; R.D. Caldwell and H.T. Boschung; UAIC
1892.
W902 (one ammocoete, 80 mm); Big Reedy
Creek, tributary to Alabama River, 1.6 miles north
of Choctaw Bluff, Clarke County; 8 April 1972;
J.R. Ramsey and W.L. Shelton; AU 6268.
W547-51 (one®, 112 mm; 4, 99-118 mm);
Big Sandy Spring, Tuscaloosa County; 8 March
Appendix
1964; R.D. Caldwell and J.C. Hall; UAIC 1225.
W541-42 (29, 107-11 mm); tributary to Car-
roll’s Creek, 5 miles north of Tuscaloosa, Tus-
caloosa County; 15 March 1963; J.D. Williams
and W.M. Howell; UAIC 1837.
W539 (ones, 107 mm); Cave Spring, 1% miles
east of junction of Morgan County; 5 March 1967;
J.G. Armstrong and J.D. Williams; UAIC 1982.
W578-80 (37, 119-28 mm); Gurley Creek, on
U.S. route 79 near Blout-Jefferson County line,
Jefferson County; 1 March 1970; Lee Barclay and
Vickie Barclay; UAIC 3342.
W517-19 (22, 97-103 mm, onec, 91 mm);
branch of Hurricane Creek at Alberta City, Tus-
caloosa County; 22 March 1956; W. Herndon and
J. Zambernard; UAIC 796.
W572 (one? , 93 mm); Hurricane Creek, Tus-
caloosa, Tuscaloosa County; 23 March 1956; L.B.
Cooper; UAIC 711.
W523-24 (oneQ, 106 mm; onec’, 101 mm);
unnamed spring, Lauderdale County; 23 March
1967; J. Armstrong and T. Jandebeur; UAIC 1991.
W529 (one2, 99 mm); Little Cypress Creek,
%> mile west of Zip City, Lauderdale County; 23
March 1967; B.R. Wall, H. Harima, and J.F.
Thompson; UAIC 2509.
W531-32 (one?, 102 mm; one, 115 mm);
tributary of Middle Cypress Creek at Bethel Berry
Church, Lauderdale County; 29 January 1969;
Wall, Harima, and Mettee; UAIC 3240.
W522 (ones, 124 mm); New River below U.S.
highway 278 at Natural Bridge, Marion County;
30 March 1968; Caldwell, Wall and Barclay; UAIC
2885.
W528 (onec’, 112 mm); New River below U.S.
highway 278 at Natural Bridge, Marion County;
3 April 1965; J.D. Williams, D. Caldwell, and W.M.
Howell; UAIC 1593.
W537 (one®, 118 mm); Polebridge Creek, 11
miles north of Tuscaloosa, Tuscaloosa County;
2 February 1963; J.D.W. and W.M. Howell; UAIC
1161.
W544 (one2, 95 mm); Schultz Creek, 4 miles
north of Centerville, Bibb County; 29 March 1966;
W.M. Howell; UAIC 1903.
W896-900 (5 ammocoetes, 66-125 mm); Seven
Springs, tributary to Choccolocco Creek, 1.3 miles
north-northeast of Choccolocco, Calhoun County;
11 September 1971; J.E. McCaleb and L.L. Ram-
sey; AU 4819.
W530 (onec’, 110 mm); tributary of Sipsey
River, 2 miles south of Fayette, Tuscaloosa
County; 25 February 1966; R.D. Caldwell and Bill
Shamblen; UAIC 1863.
W540 (one, 109 mm); South Sandy Creek,
Talladega National Forest, Bibb County; 1 April
1962; T.H. Walker, R.B. Phillips, and B. Hepner;
UAIC 882.
W546 (ones, 123 mm); South Sandy Creek,
Tuscaloosa County; 17 January 1964; R.D. Cald-
well and W.M. Howell; UAIC 1112.
W892-95 (4 ammocoetes, 98-132 mm); Town
Creek, tributary to Cotaco Creek near Somerville,
Morgan County; 12 July 1966; G.R. Hooper; UAIC
1848 and UAIC 1873.
Arkansas: Mississippi River system
W742 (one, 110 mm); Piney Creek, Izard County;
17 February 1973; W.J. and R.S. Matthews and
G.L. Harp; ASUMZ 1878.
W740-41 (29, 110-12 mm); Saddle Mill Pond
branch, South Folk River, Fulton County; 27 Jan-
uary 1973; G.L. Harp; ASUMZ.
Kentucky: upper Mississippi River system
W2316 (oneo’, 150 mm); Clear Creek, 2 miles
southeast of Disputanta, Rockcastle County; 20
March 1966; Steve Stacy; EKU 168.
W607 (onec’, 143 mm); creek, 6 miles south of
Fairdale, Bullitt County; 7 April 1968; R.T. Schaaf;
INHS.
W609-10 (2c, 121-36 mm); creek one mile
west of Means, Montgomery County; 12 April
1968; P.W. and D.M. Smith; INHS.
W747-49 (3 ammocoetes, 62-67 mm); north
fork of Rough River, Breckinridge County; 18
April 1970; Branson and Class; EKU 421.
W608 (one? , 140 mm); Walters Creek, 4 miles
north of Magnolia, Larue County; 28 March 1964;
M.E. Braasch and P.W. Smith; INHS.
Maryland: Chesapeake Bay basin
W1104-08 (one®, 92 mm; 4, 100-12 mm);trib-
utary of Battle Creek, 5 miles southwest of
Prince Frederick, Calvert County; 4 April 1964;
Brooke Kaine; CBL 2266.
W1101 (one®, 128 mm); below Uriebille Pond,
Kent County; 14 November 1958; J. Longwell;
CBL 1969.
Ohio: Ohio River system
W588-89 (one® , 113 mm; onec’, 135 mm); Crane
Hollow, Ohio River, Hocking County; 20 April
1967; Andrew White; UAIC 2846.
W587 (oneco’, 123 mm); Laurelville; 17 April
1969; Andrew White; UAIC 4504 .
Tennessee: Tennessee River system
5069-76 (VDV), (22, 140-51 mm; 69, 121-45
mm); Falling Water River, 7%2 miles southeast of
Cookeville, Putnam County; date and collector
unknown; TTU.
35
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36
Viadykov, V.D.
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