1838.
_Tyrse argus_ Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 48,
1844.
_Aspidonectes nuchalis_ Agassiz, Contr. Nat. Hist. United States,
1(pt. 2):406, 1857.
_?G[ymnopus] olivaceus_ Wied-Neuwied, Nova Acta Acad.
Leopold.-Carol., 32:55, pl. 5, 1865.
_Type._--Lectotype, Museum d'Histoire Naturelle, Paris, No. 8808;
large stuffed female obtained by C. A. Lesueur from the Wabash River,
New Harmony, Posey County, Indiana (Pl. 52).
_Range._--Northeastern United States and extreme southeastern Canada
in tributaries flowing into the Mississippi River from the east, and
the St. Lawrence River drainage; extreme southern Quebec and Ontario,
Canada, east through southern Great Lakes region to Wisconsin, and
south through New York, western Pennsylvania and Illinois to Tennessee
and western Virginia (see map, Fig. 19).
_Diagnosis._--Juvenal pattern of large, thick-bordered black ocelli,
often 9-10 millimeters in diameter in center of carapace on adult
males, and 2-3 millimeters in diameter on hatchlings (mean OD/PL,
Michigan, .066); only one dark marginal line separating pale rim of
carapace from dorsal ground color.
_Description._--Plastral length of smallest hatchling, 2.7 centimeters
(UMMZ 89950, INHS 3143); of largest male, 14.5 centimeters (UMMZ
72512); of largest female, 31.0 centimeters (UMMZ 40866).
Carapace olive, having large ocelli in center but smaller ocelli or
spots at sides; ocelli often interrupted; pale rim of carapace not
four or five times wider posteriorly than laterally, separated from
darker ground color of carapace by one dark marginal line; large
females often having remnants of ocelli at sides of carapace on
mottled and blotched background; pattern on snout of pale,
dark-bordered stripes that unite forming acute angle in front of eyes;
well-defined dark markings in subocular and postlabial region; pattern
contrasting with ground color on side of head; postlabial stripe
interrupted, diffuse; pale postocular stripe having blackish borders
interrupted, not uniting with postlabial stripe; dorsal surface of
soft parts of body having contrasting pattern, largest blackish marks
on hind limbs; elongate tail of adult males having pale dorsolateral
bands with well-defined lower blackish borders; underparts whitish,
often having blackish marks, except in center of plastral area; dark
marks on webbing of limbs, palms and soles; dark streaks often
coincident with digits; small conical tubercles on anterior edge of
carapace on adult males; conical or equilateral tubercles on anterior
edge of carapace of large females; accessory knoblike tubercles in
nuchal region and in middle of carapace posteriorly on large females.
Ontogenetic variation in PL/HW, mean PL/HW of specimens having
plastral lengths 7.0 centimeters or less, 4.09, and exceeding 7.0
centimeters, 5.50; ontogenetic variation in CL/CW, mean CL/CW of
specimens having plastral lengths 8.5 centimeters or less, 1.12, and
exceeding 8.5 centimeters, 1.21; mean CL/PCW, 2.02; mean HW/SL, 1.30
(including subspecies _hartwegi_); mean CL/PL, 1.39.
_Variation._--Variant individuals include: UMMZ 72512, an adult male,
having some ocelli seven millimeters in diameter that are almost solid
spots; UMMZ 89659 having postocular and postlabial stripes connected
on right side of head; UMMZ 95615, 52948, 54402 having inner dark
borders of pale stripes on snout represented by short dashes and dots
(a ragged line connecting anterior margins of orbits on 54402); UMMZ
52948, 89659 having interrupted, black marginal lines on carapace with
ends of some segments oriented inward and overlapping portion of
adjacent segments; UMMZ 81699, female having plastral length of 19.0
centimeters, lacking conspicuous tubercles on anterior edge of
carapace; UI 2403, CNHM 92204 having extensive dark mottling and
marbling on throat and neck, undersurface of limbs and posterior
portion of carapace.
_Comparisons._--_T. s. spinifer_ can be distinguished from all other
subspecies of _T. spinifer_ by the presence of large black ocelli
(diameter 9-10 mm. on adult males, 2-3 mm. on hatchlings) in
combination with only one dark marginal line. _T. s. spinifer_
resembles _asper_ in having ocelli or dots on the carapace but differs
from _asper_ in having only one dark marginal line and larger ocelli.
_T. s. spinifer_ differs from _hartwegi_ only in the large size of the
ocelli. _T. s. spinifer_ resembles _hartwegi_ and _asper_ but differs
from _pallidus_, _guadalupensis_ and _emoryi_ in having blackish spots
and ocelli on the carapace and lacking whitish dots. _T. s. spinifer_
resembles _hartwegi_, _asper_, and _pallidus_ and differs from
_guadalupensis_ and _emoryi_ in having conical or knoblike tubercles
on the anterior edge of the carapace on large females.
_T. s. spinifer_ differs from the subspecies _asper_, _guadalupensis_
and _emoryi_ in having a relatively narrower head, and from _emoryi_
in having a relatively wider carapace. _T. s. spinifer_ resembles
_hartwegi_ and _asper_ but differs from the other subspecies in having
the carapace widest at a plane approximately one-half way back on the
carapace. The subspecies _spinifer_ and _hartwegi_ have longer snouts
than _pallidus_, _guadalupensis_, and _emoryi_. _T. s. spinifer_
differs from _asper_ but resembles all the other subspecies in having
a relatively longer plastron.
_Remarks._--Lesueur's description of _Trionyx spiniferus_
(1827:258-261, Pl. 6) seems to be based mostly, if not entirely, on a
large female (length of carapace, 13 inches), which was "Le plus grand
des individus observes ..." (_op. cit._:258); an accompanying
illustration depicting the dorsal surface of the bony carapace is
unusual in lacking neurals (Pl. 6, E). Duméril and Bibron (1835:481)
mentioned eight or nine additional specimens that Lesueur sent to the
Museum of Natural History in Paris. Dr. Jean Guibé informed me under
letter dated September 24, 1959, that a larger stuffed female, bearing
catalog number 8808 is regarded as the holotype, and that there are
seven additional specimens (1949, 4143, 8807, 8809-12) in the museum
at Paris. All turtles were obtained by Lesueur from the Wabash River.
To my knowledge no specimen that was available to Lesueur has been
specifically designated as a type. Because the description seems to be
based on one specimen, undoubtedly No. 8808, this specimen has been
regarded as the holotype. However, Lesueur referred to several
specimens and did not mention a type in the original description;
consequently I prefer to regard No. 8808 as a lectotype.
Lesueur also described _Trionyx ocellatus_ (_op. cit._:261-263) as a
variety of _T. spiniferus_ having ocelli, or parts thereof, on the
carapace and mentioned three specimens. The total number of specimens
that were available to Lesueur is unknown. One young alcoholic
specimen having ocelli is in the British Museum (Natural History)
(Gray, 1855:69). The same letter from Dr. Guibé stated that a specimen
in the Museum of Natural History, Paris, No. 6957, having a carapace
17 centimeters in length, conforms to the characters of _ocellatus_ as
mentioned by Lesueur, and was obtained from the Wabash River by
Lesueur. Two of the specimens mentioned by Lesueur (_loc. cit._) are
stated to be females. No. 6957 is an adult male and clearly shows the
juvenal pattern; it is regarded as the lectotype of _T. ocellatus_
Lesueur, a name-combination, which is a synonym, based on a secondary
sexual difference in pattern.
Rafinesque (1832:64) described a soft-shelled turtle from "the River
Hudson between the falls of Hadley, Glen and Baker, and further up to
the source" as _Apalone hudsonica_. The most outstanding
characteristic was the presence of five claws on the digits of each
limb. Rafinesque's recording of this characteristic was perhaps
influenced by the illustration of a softshell in Bartram's _Travels_
that showed each limb with five, clawed digits. Perhaps this was the
basis for Boulenger (1889:245, footnote) regarding _Apalone_ as
"mythical." The large, yellowish, black-bordered spots, one behind and
one in front of the eye presumably represent segments of the
postocular stripe and the stripe on the snout; Rafinesque described
the carapace as "entire ... the margin is yellowish unspotted, then
comes a circular black line ..." and having "many round spots
occulated and clouded by having a brown margin, with grey dots
within." Except for five claws, the description is applicable to a
softshell and referable to _T. s. spinifer_. To my knowledge, the only
other records of the occurrence of soft-shelled turtles in the Hudson
river drainage are those of Eights (_in_ Bishop, 1923:120, Mohawk
River at Cohoes), and DeKay (1842:7, Mohawk River and Hudson River
near Albany); presumably these records are the basis for the comments
of Holbrook (_in_ Bishop, _loc. cit._), and symbolized as an isolated
locality by Conant (1958:318, map 35). The type locality of _Apalone
hudsonica_ is herein restricted to the Hudson River, near Baker's
Falls, Saratoga County, New York.
Gray (1844:48) proposed the name _Tyrse argus_ for a specimen reported
to have come from Sierra Leone, West Africa; later (1855:68), he
referred the species to the genus _Trionyx_. After comparison with a
specimen of _T. spiniferus_ Lesueur, Gray (1864:89) was "doubtful
whether there must not have been some confusion about the habitat of
the specimen [which formed the basis of the description of _Tyrse
argus_], and whether it is not more probably a North American
species." The same author (1869:222; 1870:109) listed _Tyrse argus_ as
a synonym of _Callinia spinifera_ (= _Trionyx spiniferus_ Lesueur).
Agassiz (_op. cit._:406-07) described _Aspidonectes nuchalis_ on the
basis of three adults from the Cumberland River and a number of young
from the headwaters of the Tennessee River. Boulenger (1889:245,
footnote 2) suggested that the status of _A. nuchalis_ required
further investigation. The species was not generally recognized after
the turn of the century. Barbour and Loveridge (1929:226) listed MCZ
1908 (one of the juveniles) and 1623-25 as cotypes. Stejneger
(1944:52) showed that _nuchalis_ was not distinguishable from _T. s.
spinifer_, and (_op. cit._:49) listed MCZ 1623-25 as cotypes. Schmidt
(1953:110) restricted the type locality to the Cumberland River, near
Nashville, Tennessee.
Agassiz (_loc. cit._) mentioned that _nuchalis_ "differs strikingly
from Asp. spinifer in the much more elongated form of the male, and in
the great development of the marginal spines and of the tubercles upon
the carapace, ... But the most prominent specific character consists
in the marked depressions on either side of the blunt median keel, and
also in the triangular dilation of that keel behind the front margin
of the carapace." These characters seem to be of no taxonomic worth. I
have seen three syntypes (MCZ 1623-25) that undoubtedly correspond to
the three adult specimens mentioned by Agassiz. All are females,
measuring 19.5, 22.0, and 19.0 centimeters, respectively, in plastral
length, and lack a contrasting mottled pattern on the carapace; the
juvenal pattern is obscured, except for blackish spots at the edge of
the carapace on MCZ 1625, and parts of an ocellus on MCZ 1624. The
dorsal surfaces of the limbs are boldly marked. MCZ 1623, showing the
diagnostic feature mentioned by Agassiz, is photographed by Stejneger
(_op. cit._:Pls. 14, 15), and may be regarded as the lectotype of
_Aspidonectes nuchalis_ Agassiz. MCZ 1908 is one of the young syntypes
mentioned by Agassiz, and is referable to _spinifer_. The juvenal
pattern consists of spots and ocelli; the plastron measures 3.1
centimeters in length, and the carapace 4.2 centimeters.
Wied-Neuwied (1865:55-57, Pl. 5) described the species _?G_[_ymnopus_]
_olivaceus_, but was uncertain whether his interpretation was based on
a species, a variety or a secondary sexual difference. Wied-Neuwied
mentioned that Lesueur had already named this soft-shelled turtle as
_Trionyx ocellatus_, and agreed with Lesueur that those turtles having
occulated spots on the carapace were distinguishable from _T.
spiniferus_ and _T. muticus_. But because Duméril and Bibron in their
_Erpétologie Général_ failed to recognize _T. ocellatus_, Wied-Neuwied
felt obliged to bring it to the attention of his American colleagues
and he renamed it. Wied-Neuwied also stated, in the context of a
synonym, "Beschreibung einer Reise in Nord-America Bd. I., pag. 140."
This comment presumably refers to his earlier description of _T.
annulifer_ (1838:140); seemingly Wied-Neuwied considered _T.
annulifer_ and _G. olivacea_ as conspecific, although there is no
mention of _annulifer_ in the text proper. Stejneger (_op. cit._:49)
designated the type locality of _T. annulifer_ as the Ohio River at
Pittsburgh, Pennsylvania, and of _Gymnopus olivacea_ as New Harmony,
Wabash River, Illinois (_lapsus_ for Indiana).
_Trionyx spiniferus_ was questionably considered distinct from _T.
ferox_ by Lesueur who listed "Testudo ferox Gm. Tortue de Pennant?"
and "Trionyx georgicus Geoffr.?" as synonyms. Subsequently, most
authors considered _T. spiniferus_ synonymous with _T. ferox_ until
Agassiz (1857) pointed out differences between the two species.
The average size of the ocelli on the carapace of the subspecies
_spinifer_ decreases westward toward the Mississippi River; ocelli of
different sizes occur on different individuals from the same state and
presumably from the same population. For example, INHS 2281, plastron
9.9 centimeters in length, from Effingham County, Illinois, has some
ocelli eight millimeters in diameter, whereas a larger male from the
same locality, UI 1322, plastron 11.6 centimeters in length, has the
largest ocelli only five millimeters in diameter. For convenience, all
softshells having locality data from states east of the Mississippi
River are referred to _spinifer_, recognizing that intergradation
occurs with _hartwegi_ over a broad area paralleling the Mississippi
River. The type locality of _spinifer_ is in an area where most
turtles do not have the larger ocelli (diameter of seven to ten mm. on
adult males); however, some individuals from the Wabash River (UMMZ
63523, adult male, plastron 11.5 cm. in length, ocelli diameter seven
mm.) agree with more "typical" _spinifer_ to the east. Intergradation
with _asper_ possibly occurs in that part of the Tennessee River in
eastern Tennessee as exemplified by UMMZ 59198.
Published reports indicate that _T. s. spinifer_ is not abundant in
some of the northeasterly parts of its geographic range. Adams and
Clark (1958:10) wrote that few softshells at Long Point on the
Canadian side of Lake Erie are "ever collected and the area's game
keepers report ... (none) ... seen in recent years. They also tell of
recurrent severe stormy winters in which the muddy bottom of the
marshland was repeatedly churned up and frozen. Such climatic
conditions could easily destroy a large part of the _Trionyx_
population overwintering in the mud bottom." Wright (1919:8) reported
that softshells are "rarely seen" in bays on the New York side of Lake
Ontario, and Babcock (1938:53) wrote that _spinifer_ "is not common in
Lake Champlain."
_T. s. spinifer_ probably extended its geographic range into the
Hudson River drainage of New York _via_ the Erie Canal (connected
Buffalo and Albany) after its completion in the early 1800's (DeKay,
1842:7). Now, the New York Barge Canal (essentially the Erie Canal,
but with minor changes in course and the addition of several spurs)
provides an avenue for dispersal of _spinifer_ to the Hudson River
drainage, Lake Ontario and intervening waterways in New York (Mertens,
1928:199). Netting (1944:86-87), however, suggested that _spinifer_
occupied Lake Champlain, the Finger Lakes, Mohawk River and upper
Hudson in the late stages of the formation of the Great Lakes.
A publication not seen by me is that of Mansueti and Wallace (1960).
Its title suggests that _Trionyx_ occurs in Maryland.
The unsuccessful introduction of _T. s. spinifer_ in the Delaware
drainage in New Jersey has been discussed by Fowler (1907:213), who
wrote that they were found as early as the late 1860's and were
introduced when young presumably to stock aquaria. Records of
occurrence include Cooper's Creek, Camden County (Stone, 1906:168);
Woodbury, Gloucester County (Cope, 1894:889); and Paulins Kill at
Hainesburg, Warren County (Johnson, 1894:889).
Surface (1908:122) believed that soft-shelled turtles "have doubtless
been introduced into the eastern part of Pennsylvania through the
canal from the Western and Central part of New York," and Roddy (_in_
Neill, 1951:21) suggested that the species may be found in the
Susquehanna River. Babcock (1919:420) mentioned a young specimen of
_spinifer_ in the collection of the Boston Society of Natural History
that was obtained "in White River, Vermont," a tributary of the
Connecticut River of the Atlantic Coast drainage; seemingly this
record has not been accepted and the species is not established. To my
knowledge, populations of _T. s. spinifer_ do not occur in rivers of
the Atlantic Coast drainage, except probably the Hudson-Mohawk
drainage.
Stockwell (1878:401) wrote that _spinifer_ was found "as high as
Athabasca." Presumably Stockwell referred to Lake Athabaska in
northern Alberta and Saskatchewan, Canada, a region where soft-shelled
turtles are unknown; see also the comments by Stejneger (1944:52).
_Specimens examined._--Total 250 as follows: ALABAMA: _Morgan_: UMMZ
99578, "near" Decatur.
ILLINOIS: _Adams_: INHS 2150, Quincy. _Bond_: INHS 8345, Greenville.
_Carroll_: CNHM 42116, Ordnance School Proving Ground. _Cass_: INHS
2151, Beardstown. _Champaign_: INHS 2273, 2311, 2413, 3142, "near"
Seymour; INHS 4229, Champaign; INHS 6163, Sidney. _Christian_: INHS
1560, Pana. _Coles_: INHS 1968-69, 2 mi. W Charleston. _Cumberland_:
INHS 2282, Greenup. _De Witt_: INHS 7674, Farmer City. _Effingham_: UI
1322, 2281, 19365, "near" Effingham. _Fulton_: INHS 5531, 2 mi. NE
Bluff City, Schyler County; UI 23449, Liverpool; UI 24611, Spoon
River, 18 mi. NW Canton. _Hancock_: USNM 53522, 59277, "near"
Hamilton. _Iroquois_: INHS 6869-70, 2.5 mi. N Crescent City.
_Jackson_: TU 1369 (12), Elkville. _Kane_: CNHM 42400, Aurora.
_Kankakee_: CNHM 324, Momence. _Kendall_: UI 2411, Plano. _Logan_:
INHS 7171-72, 6 mi. N Lincoln. _Madison_: USNM 60571. _Macoupin_: UI
2401-02, Beaver Dam Lake. _Mason_: CNHM 346, 470, INHS 1122, 1559,
5756-58, UI 42, 2404, Havana, Lake Chautauqua. _Mercer_: CNHM 3220,
New Boston. _Morgan_: CNHM 2067 (2), 3290, 3303-04, 3306, INHS 2152,
2154, 5132-37, USNM 54747, Meredosia. _Moultrie_: INHS 8989, 2 mi. NW
Lovington. _Peoria_: UI 2406-10, Peoria. _Pope_: INHS 5505, Lake
Glendale. _Putnam_: UMMZ 81604-14, 5 mi. N Henry, Marshall County.
_Schuyler_: UI 2405, "near" Ripley, Brown County. _Scott_: INHS 2149,
2153, Naples. _Union_: CNHM 18623, 6 mi. SW Jonesboro. _Vermilion_:
INHS 3142, Muncie; INHS (1 untagged); UI 1970, 3209, Danville; UI
2403, 1.5 mi. E Oakwood; UI 16265, Kickapoo State Park. _Wabash_: USNM
12061, Mt. Carmel. _Winnebago_: INHS 7185, Kishwaukee Forest Preserve;
INHS 7294, 1/2 mi. S Shirland. _County unknown_: USNM 7661.
INDIANA: _Bartholomew_: UMMZ 61060, 10 mi. W Columbus. _Carroll_: USNM
42905-06, Burlington. _Clark_: UMMZ 110599, 14-mile Creek, 3 mi. NW
Charleston. _Decatur_: UMMZ 55416, 3 mi. S Westport. _Elkhart_: UMMZ
105598, Elkhart River, south of Goshen. _Gibson_: UMMZ 89744, Foot's
Pond. _Johnson_: UMMZ 108062, 2 mi. S Trafalgar. _Knox_: USNM 22711,
Vincennes. _Kosciusko_: AMNH 8379, UMMZ 84287 (5), Winona Lake; UMMZ
110235, Wawasee Lake. _Lake_: CNHM 11019, 11021-24, Crown Point.
_Marion_: UMMZ 103393, Ravenswood; UMMZ 110236, 1 mi. N Lawrence.
_Marshall_: CNHM 39299; USNM 33495, Yellow River north of Burr Oak;
USNM 33496-501, 35404, 42583-84, Lake Maxinkuckee. _Wells_: UMMZ
63523, Wabash River, Bluffton. _County unknown_ (Lagrange or
Marshall): USNM 50670, Twin Lakes.
KENTUCKY: _Casey_: UMMZ 112252, trib. of Green River, south of
Yosemite. _Green_: UMMZ 116718, Little Barren River, 1.5 mi. E Monroe,
Hart County. _Rockcastle_: UMMZ 98767, Rockcastle River, 5 mi. above
Livingston.
MICHIGAN: _Allegan_: UMMZ 42112, Kalamazoo River. _Barry_: UMMZ 53874,
Thornapple River, 3 mi. NW Hastings. _Bay_: UMMZ 74670. _Branch_: UMMZ
95615, 1 mi. S Kinderhook; UMMZ 70748, Hog Creek. _Calhoun_: UMMZ
89950 (3); UMMZ 79133, near Battle Creek. _Cass_: UMMZ 40866-67,
53005, Diamond Lake; UMMZ 40868, 52948, Long Lake. _Jackson_: UMMZ
72494. _Kalamazoo_: UMMZ 42130, 80534, Kalamazoo; UMMZ 90506, Gull
Lake; UMMZ 92599, Kellogg Bird Sanctuary. _Lenawee_: UMMZ 72457,
Devil's Lake; UMMZ 74662, Wolf Lake Park. _Livingston_: UMMZ 54401,
76190, Portage Lake. _Monroe_: UMMZ 44604-06, USNM 51213, "near"
Monroe. _Newaygo_: UMMZ 63469. _Oakland_: UMMZ 64363, Hay's Creek;
UMMZ 96539, Clinton River. _Ottawa_: UMMZ 81699. _St. Joseph_: UMMZ
38876, 38889, "near" White Pigeon; UMMZ 96537, Corey Lake. _Van
Buren_: UMMZ 90003, Wolf River, west of Kalamazoo, Kalamazoo County.
_Washtenaw_: SM 2035, 2038, 2105, UMMZ 39847, 96538, "near" Ann Arbor;
UMMZ 35765, 35769, 74518 (2), Portage Lake; UMMZ 54402-03, Little
Lake; UMMZ 89659, Huron River, Dexter; UMMZ 110583-85. _County
unknown_ (Washtenaw or Livingston): UMMZ 54400, Huron River near
Portage Lake.
MISSISSIPPI: _Adams_: MCZ 46615, UMMZ 76446, "near" Natchez; MCZ
46621, 46633, USNM 01084, 01086, Washington. _Coahoma_: AMNH 5289,
5285-86, Moon Lake. _Lafayette_: MCZ 37173, Oxford; USNM 7650,
Abbeville? (reported from Abbeville, South Carolina by Pickens,
1927:113; see discussion by Stejneger, 1944:50, and my comments on
page 509 beyond). _LeFlore_: USNM 73668-69, Greenwood. _Madison_: USNM
95192, Big Black River. _Washington_: USNM 115980, Deer Creek.
_Yazoo_: UMMZ 86669, Panther Creek west of Yazoo City; UMMZ 83304,
Yazoo City.
NEW YORK: _Monroe_: CNHM 92001-02, Genesee River, Rochester. _Wayne_:
AMNH 69931, CNHM 92004, Sodus Bay.
OHIO: _Athens_: UMMZ 111793, east branch Shade Creek. _Franklin_: USNM
26290. _Lucas_: USNM 51214, Toledo. _Pike_: UMMZ 99309, Morgan's Fork,
Sunfish Creek. _Warren_: AMNH 4763, Little Miami River, 3 mi. below
Morrow. _County unknown_: USNM 21128-29, Cuyahoga River.
TENNESSEE: _Benton_: UMMZ 113036, Eagle Creek, 1/2 mi. E Holliday.
_Bradley_: UMMZ 59197, west branch of Chestnee Creek, 7 mi. E
Cleveland. _Claiborne_: USNM 86677, 5 mi. SE Cumberland Gap, Powell
River. _Davidson_: MCZ 1623-25, Cumberland River near Nashville
(restricted locality); USNM 7165-67, Nashville. _Decatur_: KU 3000,
Perryville. _Hamilton_: USNM 131861, Chattanooga. _Monroe_: TU 16058,
Little Tennessee River, 10 mi. N Madisonville. _Obion_: UMMZ 53199,
USNM 102911, Reelfoot Lake. _Overton_: UMMZ 69561 (2), Wirmingham.
_Sevier_: TU 16132, UMMZ 86735, USNM 86681-82, near Sevierville; UMMZ
86734, Walden Creek "near" Gatlinburg. _County unknown_: MCZ 1908,
headwaters of Tennessee River.
VIRGINIA: _Smythe_: USNM 101386, Holston River, Seven Mile Ford.
WEST VIRGINIA: _McDowell_: USNM 33767, Dry Fork, Perryville (county
questionable, perhaps Randolph County).
WISCONSIN: _Chippewa_: CNHM 8223, Lake Wissota, mouth of Yellow River,
Anson Twp. _Polk_: UMMZ 72511-12, St. Croix River "near" Never's Dam.
_County unknown_: CNHM 15971, Eau Claire River.